https://cpw.cvlcollections.org/files/original/28eabfc4589cf192f6162bfb1422a33e.pdf 359344a47c785eb13fe4f17d0b1512c6 PDF Text Text 1 JOB PROGRESS REPORT State of _ _C~O~L~O~RADO=~--------Project No. ~S~E=---~3~-~2=----------- Endan2ered Wildljfe Tnvestjgatiao Work Plan No. - - ~ l = - - - - - - - - -- Job No. Job Title Colorado Sguawfish Propagation Study . Period Covered: July 1, 1978 to June 30, 1979 Personnel: Gary Barta, Walter Graul, Charles Haynes, Robin Knox, David Langlois, Tom Lytle, Clee Sealing, Ken Stead and John Torres. ABSTRACT A Colorado squawfish hatchery feasibility study was initiated a.t the Rifle Falls State Fish Hatchery near Rifle, Colorado. An engineering plan was prepared to accomodate research and production of 100,000 young fish annually . The hatchery plan includes: a hatching house to research spawn taking techniques, egg incubation, and fry rearing; two half- acre ponds and one quarter-acre pond for rear ing fingerlings and overwintering brood fish, respectively; four concrete raceways for additional fingerling rearing space;. a pollution abatement pond to restore water quality; and, a quarantine raceway to hold wild brood fish. ��3 COLORADO SQUAWFISH David Langlois, PROPAGATION STUDY Tom Lytle, and Charles Haynes PROGRAM NARRATIVE OBJECTIVES To design Colorado squawfish propagation facilities for construction at the Rifle Falls fish hatchery capable of producing 100,000 juvenile Colorado squawfish yearly; and, to begin construction of such facilities by June 30, 1979. INTRODUCTION With the knowledge that Colorado squawfish can be successfully propagated in a hatchery (Toney 1979), the State of Colorado decided to investigate and refine production technology. Spawn-taking techniques, handling of brood fish, and rearing methods for progeny are some of the research unknowns which must be answered to maximize production of fingerling squawfish on an annual basis. It was also decided that a facility devoted totally to Colorado squawfish production and research would enhance the chances of large-scale success. The Rifle Falls State Fish Hatchery became a candidate site for a feasibility study on the following criteria: 1) it appeared to have land and water available to develop the facility; 2) it is adjacent to the historical habitat of the Colorado squawfish, i.e., the Colorado River and, 3) the existing hatchery and regional staff had shown an interest in the culture of endangered species. While hatchery propagation alone will not result in the declassification of the species, such a source of young fish will aid the recovery effort in several ways. Fish of a uniform size and age are needed for bioassay work, including salinity, pesticide, and heavy metal toxicity studies. This knowledge would supplement (not replace) habitat requirement studies using wild fish. Taxonomic work with larval squawfish would facilitate field identification of wild fish, making it easier to monitor annual reproduction. Life history studies could also be conducted using hatchery fish both in the hatchery environment and in historic habitat. Squawfish are needed to investigate competition and predation with other fishes. migration, growth, and survival. Such uses of hatchery - reared squawfish demonstrate tbe value of the hatchery product on the squawfish recovery effort. SYNOPSIS OF PROGRESS On August 2, 1978 personnel from the Division of Wildlife and the U.S. Fish and Wildlife Service met at the Rifle Falls hatchery to tour the unit, evaluate potential construction sites, and discuss the operation of the squawfish facility. Three possible sites were discussed and it was decided that the Division Engineering Section should design a construction plan. The regional office made the engineering request. As a result, an on-site inspection was completed by the engineering section and three alternative hatchery plans were prepared for review during the winter months. �4 On March 5, 1979 the Division of Wildlife and the U.S. Fish and Wildlife Service met again to review the three engineering designs and select the best one for further work. The details of the hatching house and pond layout were also reviewed. Final approval is pending. The hatching house hopefully will provide an environment conducive to survival and spawning of fry and fingerlings. It includes several features requiring advanced fish culture and engineering technology. As water enters the hatching house from the settling pond or the existing trout hatchery source, it will be passed through an ultraviolet filter'to kill potential pathogens. A water pump will push water through the system at a flow up to 225 gallons per minute. A water heater will be used to increase water temperature to approximately 70 to 750 F (This will be a research item) because the incoming water will seldom be warmer than 550 F. By recirculating the heated water at a rate of 90 percent recirculated and 10 percent fresh water, it will be possible to significantly reduce heating costs. The energy source will either be electrical, propane or solar powered but in any case will have a back-up power source. Particulate material will be removed when the water passes through a settling basin. Additionally, the water will pass through a denitrifying filter to reduce toxic ammonia waste products. Aerators will be used to restore dissolved oxygen to saturation and eliminate possible gas bubble disease caused by supersaturated nitrogen. Holding facilities include two indoor raceways for brood stock spawning and six small troughs to incubate eggs and rear fry. A small storage area is also included. Production raceways and ponds, the quarantine facility, and the broodstock overwintering pond will be located outdoors. In the raceways , it will be possible to rear fingerling squawfish and conduct diet, growth and survival experiments. Similar experiments can be conducted in the ponds for comparison with raceway data. The quarantine facility will be used to isolate wild brood fish as they are captured and delivered to the hatchery. Disease inspection will be made in this raceway. The brood stock overwintering pond will be used to produce forage for the brood fish, hold the brood fish during non-spawning months, and research brood stock maintenance techniques. In July of 1979., regional personnel visited Hatchery in Arizona to tour their squawfish techniques. the Willow Beach National Fish facility and discuss propagation The letting of bids and contracting of construction work were not accomplished by June 30, 1979. Additional review of the hatchery design was requested by regional and research personnel. Also the Colorado Legislature requested a reconsideration of the plan to propagate squawfish at the Rifle Falls unit. When, and if, these concerns are resolved, construction will begin. �5 LITERATURE CITED Toney, D.P. 1974. Observations on the propagation and rearing of two endangered fish species in a hatchery environment. Proc. West. Assoc. State Game and Fish Comm. 54: 252-259. Prepared by: Dave Langlois Biologist Final Edit by: Nongame �6 JOB PROGRESS REPORT State of COLORADO Endangered --~~~~~---------------- Project No. SE-3-2 Work Plan No. Job Title: 1 II ----------------------- Nesting Performance of Peregrine Falcons in Colorado Period Covered: Personnel: Job No. Wildlife Investigation July 1, 1978 - June 30, 1979 J. Enderson, Colorado College; D. Berger, B. Busby, and B. Pendleton, G. Craig, Colorado Division of Wildlife, J. Hogan and S. Petersurg ABSTRACT In 1979, the number of pairs of breeding peregrines declined although an additional site was occupied. The reduced population makes establishment of trends difficult, but the population appears to be continuing to decline. Reoccupancy of one site and presence of yearling birds at two other sites offer hope that trend may reverse soon. Although augmentation efforts obscure natural reproduction, estimates of probable natural reproduction are provided. Pesticide residue of whole eggs collected in 1978 are provided and shell thickness data for 1978 eggs and some hatched 1979 eggs are also presented. Potential prey species were collected at four eyrie sites but residue analysis is not yet available. �7 NESTING PERFORMANCE OF PEREGRINE FALCONS IN COLORADO Gerald R. Craig P. N. OBJECTIVE The objective of this study is to annually monitor the breeding numbers and reproduction of Colorado peregrine falcons in an effort to document further population declines as well as eventually record the population's response to various recovery efforts which are implemented. Additionally, health of the population may be monitored indirectly by analyzing pesticide residue levels in the falcon's eggs and principal prey they feed upon. Information obtained from these investigations will be made available through annual reports to the Rocky Mountain/Southwest Peregrine Falcon Recovery Team as well as cooperating agencies to aid in evaluation of various recovery efforts. SEGMENT OBJECTIVES 1. Annually document the number of breeding their productivity in Colorado. pairs of nesting peregrines 2. Annually survey potential habitats to locate previously peregrines and document their reproductive success. 3. Annually document egg shell thinning and 'pesticide residues nesting peregrines in Colorado. 4. Collect samples of principal avian prey species utilized by nesting peregrines and analyze the samples for pesticide residues. 5. Compile data and submit reports and the Rocky Mountain/Southwest in evaluating recovery efforts. and unknown nesting in eggs of to appropriate state and federal personnel Peregrine Falcon Recovery Team for use Note - These objectives correspond to tasks 1113.,1114., 212., and 221. in the approved American Peregrine Falcon Recovery Plan (Rocky Mountain/Southwest Population). PROCEDURES lao Visit all nest sites throughout Colorado which have been occupied within the past three years and observe them from a distance with spotting scopes and binoculars to establish the presence of breeding adults. All occupied sites will be revisited periodically throughout the nesting season to document reproductive success. lb. Prior to, or immediately after hatching of the eggs, nest sites will be visited and egg shell fragments and addled eggs will be collected for pesticide analysis. Egg shells will be measured for a thickness index according to standardized methods. Egg contents will be shipped to the Fish and Wildlife Research Laboratory at Patuxent for analysis. �8 1c. Successful nests will be visited prior to fledging of the young and they will be banded and color marked. These sites will be kept under surveillance to determine actual fledging success. 2. Favorable habitats will be surveyed for potential nesting sites. The surveys will be conducted from the ground, but where necessary a helicopter may be used in remote regions. When new pairs are located they will be surveyed as outlined in la, 1b and 1c. 3. Ten to twenty individuals of each of 5 prinicpal avian prey species will be collected from representative hunting areas utilized by breeding peregrines. This will be done at two sites annually. The individuals of each species will be combined into a single sample for pesticide residue analysis. The analysis will be accomplished by the Fish and Wildlife Service Laboratory at Patuxent or an independent laboratory recognized by the Patuxent Laboratory. 4. Compile data and submit reports and the Rocky Mountain/Southwest to appropriate state and federal personnel Peregrine Falcon Recovery Team. METHODS AND MATERIALS Nesting Investigations. Methods for nesting in the Procedures enumerated above. investigations have been described Pesticide Analysis of Prey. Adult individuals of potential prey species within 8.3 km (5 mi) of active eyries were collected. The species were selected on the basis of high prevalence, and both migratory, non-migratory, herbivorous and insectivorous forms are included. Specimens were taken by shotgun and stored frozen. After thawing, the birds were weighed, plucked, and the feet, beak and large and small intestine removed. The remainder was finely chopped, weighed and a sample taken for pooling. Species pooled samples were frozen in acetone-washed foil and submitted for analysis. Analytical prodecures followed by Raltech Scientific Services, WARF Institute, Inc.,) at Madison, Wisconsin were as follows: Inc. (formerly Each sample is al.lowed to thaw in the refrigerator overnight. A 10 gram aliquot is weighed into a 250 ml beaker and mixed with a spatual with 150 grams of sodium sulfate. At the same time an aliquot is weighed into a preweighed 100 ml beaker for moisture determination and placed in a 400C oven for 2 weeks. After 2 weeks, weigh the beaker and dry sample back and calculate percent of moisture. The 10 gram portion is allowed to dry overnight and is transferred to a 43 x 123 Whatman extraction thimble and plugged with glass wool. Place in a large soxhlet extraction aparatus and extracted with 50:50 Ethyl Ether: Petroleum Ether for 8 hours. Sample is removed and taken down just to dryness on a steam bath. Sample is brought to 25 ml with 25% Toluene in Ethyl Acetate. A 5 ml aliquot is transferred to Gel Permeation apparatus. The settings are 28 mins discard, 14 mins collect and 2 mins wash. The sample is taken down on a flash evaporator just to dryness and made to 10 ml for injection. �9 Injection: Hewlett Packard 5710A with Ni 63 Electron capture detector, with allta injector and hooked to a Hewlett Packard integration computor model 3352C. Column: 1.5% OV-17 x 1.95% QF-1 on 80/100 G.C.Q. Column Temperature: 2000C; Detector Temperature: 3000C; DDT Retention time of approximately 9.8 mins; Injection Temperature: 2500C; Carrier: 95% argon; 5% Methane Flow 31 ml/min. .. Lipid Determination: From the 25 ml volumetric containing the sample a 5 ml aliquot is pipetted into a preweighed 2 dram vial. These are placed into a 400C oven for 2 days, desiccated, weighed and percent lipid calculated. Pesticide Analysis of Peregrine Eggs. Infertile and unhatched eggs which are encountered in the wild, as well as those wild eggs which are removed for incubation in captivity and subsequently fail to hatch were analyzed for pesticide residues. Pesticide analysis of eggs was accomplished by the U.S. Fish and Wildlife Service Research Center at Patuxent. In an effort to maintain uniformity in results, Patuxent should continue to perform the analysis. Samples were prepared for shipment by placing the egg contents in foil wrapped glass jars which had been rinsed twice in laboratory grade acetone. The samples were then frozen and shipped in dry ice to the laboratory. Analysis techniques are the same as those described in Cromartie et a!. 1975. Pesticide Monitoring J. 9:11-14. Since the eggs have undergone dehydration during incubation, residue values were corrected to fresh weight values assuming 85% moisture content. RESULTS AND DISCUSSION Productivity of Nesting Peregrines In 1979, .8 sites were occupied by pairs of falcons of which two pairs comprised yearling members. Site 6 was occupied by an adult male and yearling female for the second consecutive year and experienced no reproduction. The adult male at site 7 was replaced by a yearling male during the courtship period, so this pair was also unsuccessful in producing eggs. In addition lone adult males were present at 3 other sites (sites 27,31 and 32). A lone adult male was observed briefly in early April at site 26, but was not sighted on subsequent visits. A pair of prairie falcons which had occupied adjacent cliffs in previous years relocated their eyrie to the peregrines' nest cliff and reared young. A single adult male was observed near the eyrie ledge at site 31 late in the breeding season. This site was occupied by a pair in 1978, but the female did not return. Site 32 was frequented by a lone adult male for the second year after its discovery in 1978. During this breeding season, a lone adult male at site 5 succeeded in attracting an adult female for the early portion of the season but she disappeared shortly before egg laying. The first reoccupancy of an abandoned nest cliff was observed at site 5 which was occupied by a lone adult male in 1977 and vacant in 1978. Unfortunately, the female also disappeared at initiation of egg laying. Unseasonable late spring storms may have been a factor in reducing the number of successful pairs in 1979. The females at sites 5 and 28 appeared gravid �lO • and ready to lay but disappeared during or immediately following a spring snow storm which lasted several days. The cause of their disappearance could not be determined. One egg in a clutch of three at site 9 was eight days late hatching (under artificial conditions) which suggests that the female may have suspended laying activities or resorbed an egg and initiated full term incubation prior to completion of the clutch. At site 27, one egg of the second clutch had a piece of hardened mud affixed to one side indicating unusual moisture conditions at the nest. This egg successfully hatched under artificial conditions but would probably have died since the adult would not have been able to turn it properly. In all, only four pairs of peregrines produced clutches of eggs and all succeeded in rearing and fledging young. Due to augmentation activities which were undertaken at all four sites, it is difficult to determine actual productivity since the pairs were not permitted to reproduce naturally. Table 1 summarized the assumed natural reproduction had the pairs not received manipulation of clutches or augmentation of broods. The various manipulative activities are described in further detail in the report for Work Plan II, Job 3 of this publication. Table 2 summarizes actual reproduction of the pairs after augmentation. Had the pairs been permitted to reproduce naturally, they would have averaged 1.25 young fledged per pair which is generally considered the threshold value necessary to sustain the population. Manipulation activities increased the fledging success to 1.50 young per pair. This is lower than the 1978 values primarily due to the presence of 2 nonbreeding pairs and two immature pairs (Le. a pair consisting of at least one yearling). When only laying pairs are considered (seven pairs in 1978 and four pairs in 1979), the assumed natural reproduction remained constant at 2.5 young per laying pair while the actual reproduction increased from 2.29 to 3.0 per laying pair. The increased fledging success partially offset the 43 percent reductions of laying pairs in 1979. Location of New Nesting Pairs Beginning in 1965, exploration of potential nesting habitats was intensified in an effort to locate previously undiscovered breeding pairs of falcons. To date, it is estimated that approximately 75 percent of the state has been surveyed which has resulted in the addition of 10 new nest sites (sites 24 through 33 in Table 3). From 1975 to 1978, 8 sites were located through an investment of approximately 150 man days of search effort (19 man days per site). Approximately 260 man days of search were expended in locating 2 new nesting sites in 1978 and 1979 (130 man days per site). The increased effort with reduced results would support the conclusion that the majority of the nest sites have been located and it is unlikely that a significant breeding population awaits discovery. Population Trends Efforts to locate new nest sites tend to obscure actual population trends. Since only occupied nests can be located, inflated values result in the percent of occupied sites calculations (Table 4). In order to obtain a more representative estimate of population trends, only those sites known prior to 1975 should be considered (as indicated above. in 1975 efforts �11 were undertaken to locate new nesting pairs). In 1979, only 5 of the 23 (22%) "historic" (pre 1975) sites were occupied. This represents an increase of one site from 1978 when only 17 percent were occupied. This recent upward trend is attributed to the reoccupancy of site 5 which was occupied by a lone adult male in 1977 and vacant in 1978. The presence of two yearlings in pair bonds at sites 6 and 7 draw mixed conclusions. The optomistic conclusion is that sufficient recruitment is occuring to maintain nesting pairs. The pessimistic response is the population does not contain sufficient members of adults to displace yearlings at breeding sites. Although it is impossible to quantify with the small remaining size, it is apparent the population is still declining. The presence of yearlings andreoccupancyof a vacant site offers some hope that the trend may reverse. Table 1. Site No. Assumed natural production~1 Site Received Manipulation of managed and unmanaged No. of Eggs Which Would Have Been Produced No. of Young Which Would Have Hatched sites. No. of Young Which Would Have Fledged 6 No 0 0 0 fJ_! No 0 0 0 7]:/ No 0 0 0 9 Yes 3 2 2 14 Yes 4 4 2 27 Yes 3 3 4 28 No 0 0 0 30 Yes 4 3 3 14 12 10 Totals 3.50 eggs per laying pair; 3.0 young hatched per laying pair; 3.0 young per brood; 1.43 young fledged for all pairs; 1.67 young fledged for each adult pair; 2.5 young fledged per laying pair; 2.5 young fledged per successful pair. II Natural reproduction not been manipulated. is estimated for managed ~I Pair consisted of an adult and a yearling. sites as though they had �12 Table 2. Actual Productivity Site Received Manipulation Site No. of managed No. Eggs Produced and unmanaged No. Eggs Hatched sites. No. Young Fledged Pair Returned to Pair 5 No 0 0 0 0 r}_1 No 0 0 0 0 7.11 No 0 0 0 0 9 Yes 3 3 3(3 c.p.)!:_1 3 14 Yes 7 6 4(1 c.P.) 21/ 27 Yes 6 4 4 (/+ c.P. ) 4 28 No 0 0 0 0 30 Yes 8 2 4 3 24 15 15 12 Totals 6.0 eggs per laying pair; 3.75 young hatched per laying pair; 3.75 young per brood; 1.70 young fledged for all pairs; 2.0 young fledged per adult pair; 3.0 young fledged per laying pair; 3.0 young fledged per successful pair • .11 Pair consisted of an adult and a yearling. II 11 Where indicated by (C.P.) captive produced young augmented production. The 3 young fledged prematurely, 1 died of injuries sustained and another disappeared, probable victim of owl predation. in the fall �Table 3. Site Occupancy of Colorado peregrine falcon eyries, 1964-1979. Pre1964 1 2 3 4 5 6 7 8 + 9 + + + 64 65 66 67 P P P P P A A A M P 68 69 70 71 72 73 74 P P M P P P P P P P P P P P P P P P P P P P P P P P A P P P P P P P P P V 10 11 12 13 14 15 16 17 18 19 2Q 21 22 23 24 25 26 27 28 + + + + + + + + + 76 77 78 79 M V P M V M V V V V V V V V V V V V V v V V V p!i._/ P P P v V P P P P P p-!/ P P P p!!_/ p!i._/ P V V V pE_/ V V P P A V F P P V V V V V V P p]j V V V V V V V V V V V p!!._/ p p P P V V V V A M V V V A V F P V V A V V v P V p 1/ v v v v v v V V V V V V V V V V V V V V V V V V V V V V A P V A V V V V V V V V A + 75 v V V V V V V V V V V V V V V V V V V V V V V V V V V V V V p p p p p p p p p V V V V V p V V V V V V V v p:i./ p V M M P ---------------------------------------------------------------------------------------------------------------- ,_. w �Table 3. Site Occupancy Pre1964 of Colorado 65 66 peregrine 68 67 falcon eyries, 69 70 71 1964-1979 72 (continued). 73 74 75 76 29 30 31 77 78 79 P P P P A p!!j M M P 32 M 33 P = Pair, M = Male, F = *These are neighboring site 10 to site 11. Female, A sites = = Lone Adult, V (approximately Lone adult male observed 2/ Adult male replaced 1/ An adult female was found dead in the vicinity !!/ Pair consisted ~/ Adult !l_/ Pair consisted by a yearling season, attracted male midway of an adult male and yearling female replaced by immature Blank Spaces 1 mile apart) and this could represent . !/ throughout Site was Vacant, through No Data Available a shift of one pair from incubation. of the eyrie, could not confirm presence through breeding male. I-' .:::- female in June. female. female midway of an adult female and yearling yearling = P season. of a pair. �15 Table 4. Occupancy and productivity Year of Colorado peregrine eyries, 1972-1979. 1972 1973 1974 1975 1976 1977 1978 1979 Eyries visited 15 23 24 26 27 31 32 33 Occupied 11 12 9 8 8 12 11 12 Adult pairs 8 11 7 6 5 11 7 6 Immature pairs!/ o o o o 2 o 2 2 Lone adults 3 1 2 1 1 1 2 3 1 5 2 4 6 5 4 2/0 11/0 5/0 6/4 11/5 16/11 12/8 0.2 1.6 0.8 1.2 1.0 2.0 2.0 2.0 2.2 2.5 1.5 1.8 3.0 3.0 eyries Successful pairs Total young fledged No. of young augmented Young/adult pair2/ Young/successful pair Percent of sites occupiedl/ 73% 52% 38% 31% 30% 39% 34% 36% Percent of sites w/ adult pairs 53% 49% 29% 27% 26% 35% 22% 18% 1/ At least one member of the pair was in juvenile 2/ 1.25 young fledged per pair is considered 3/ 80%-90% of the eyrie sites should normally year. Egg Shell Thinning plumage. normal reproduction. be occupied and Pesticide in any particular Residues Peregrine eggs failing to hatch in 1978 await analysis for chlorinated pesticide and PCB residues. Table 5 summarizes egg shell thicknesses for all eggs collected in Colorado from 1973 through 1978. Several eggs are awaiting measurement and so are not included in the list at this time. Several conclusions may be drawn from the collection of measurements: 1) Variation of shell thickness is as great between eggs in the same clutch as it is between clutches from different sites. Shell thickness varied by .03 mm or greater between eggs in the same clutch in 7 our of 12 cases. Extreme variations greater than .05 mm was experienced in two cases. 2) There is often considerable variation in shell thickness from the same site in different years. Average shell thickness of clutches varied by .02 mm or more at 3 sites (sites 7, 9 and 15) on succeeding years. 3) Although the sample is not large, as a group, the 1978 eggs 'do not appear to have the extreme downward variation observed in earlier samples. �16 Table 5. Eggshell condition of Colorado peregrines. Eyrie Year 1 2 1973 1973 1974 " " " 5 " 6 " " " " " " " 1975 1971 " 1975 " " " " " " 1976 " " " " " " " " " 1977 " " " " " " " " 7 " " " " " Ii " " " " 9 H " .325 .305 .287 .282 .259 .259 .259 .284 .294 .300 .246 .292 .295 .259 .259 .3021./ .305 1./ .272 2) " " " " Thickness (mm)1./ W/Membrane W/O Membrane " " " " " 1973 1974 1976 " " 1977 " " 1978 " " 1977 " " " 1978 " " " Remarks 1.62 1.52 1.43 1.39 .319 .299 Hatched 1.49 .231 .278 .288 .244 .203 .236 .216 Hatched .185 .203 .226 ]../ .229 1'/ .196 1/ .216 .267 .287 .279 .269 .300 1./ .274 1./ .269 1./ .279 1) .254 .272 .249 .300 .302 .269 .274 .244 .318 .333 .351 .307 2./ .323 1/ .292 1/ .272 .274 .282 Ratcliff's Index 1.1 .191 .203 .201 .203 1.1 .1931/ .248 1.32 1.32 1.43 1.35 1.48 1.40 1.LI2 1.43 1.33 Broken Hatched Hatched Broken Small Embryo 27 day embryo Hatched Hatched Died at hatch Large embryo Large embryo Hatched Hatched 27 day embryo No development Hatched Hatched 27 day embryo 1. LI5 .188 .244 .216 .191 .239 1./ .251 2) .229 1/ 1.42 1.41 1.36 1.68 1.72 1.61 1.65 1.72 1.62 1.50 1.49 1.53 Addled Addled Hatched Hatched Hatched Hatched No development No development No development Hatched Hatched No development Hatched Hatched Hatched " ------------------------------------------------------------------------------ �17 Table 5. Eggshell condition of Colorado peregrines (continued). Eyrie Year " " 1979 " II 11 " 11 " 14 " " " " " 1974 1975 1975 " 1974 1975 1977 " " " 1978 " " " " " " " " " " " " " " " " " " 1979 " " " " " 25 1977 " " " " " " " " Ii " 26 1977 " " " " 27 1978 " " " " " " 1979 .285 .328 .338 .307 .297 .305 .297 .284 .305 .203 .292 .310 .284 .289 1.48 .262 .264 .241 .244 .234 .252 .218 .226 .201 .211 .284 .322 2/ .312 2/ 2-/ .287 .320 .320 .305 .305 .315 .269 .267 .302 .267 .307 .267 .244 .246 .259 .307 .323 Ratcliff's Index .206 .201 .229 .246 1.64 1.61 1.61 1.55 1.50 1.49 1.55 1.60 1.36 1.47 1.55 1.48 2/ 2/ 2/ .224 .244 .196 .211 2/ 2/ .318 .196 .185 .231 .231 .208 .244 1.41 1.34 1.58 1.42 1.57 1.35 1.43 1.50 1.46 1.59 1.61 1.50 " 30 1978 " " " " " " " " " " Thickness (nun) 1/ W/Membrane W/O Membrane " " .312 .307 .292 .292 .312 2/ .292 2/ .323 2/ .312 .293 .246 .244 .229 .234 1.73 1.55 1.43 1.54 1.57 1.37 1.64 Remarks Hatched Hatched .Hatched Hatched Hatched Spoiled Spoiled Spoiled Hatched Hatched Hatched No Development No Development No Development Hatched Hatched Hatched Hatched Hatched Hatched Hatched 26 day Embryo 26 day embryo No development No development Cracked Cracked 24 Day embryo Hatched Cracked Hatched Hatched Hatched Hatched Hatched Large Embryo No development Large Embryo No development No development No development Hatched Hatched 1979 " .249 " " -------------------------------------------------------------------------------- �18 Table 5. Eggshell condition of Colorado peregrines (continued). Thickness (mm) 11 WIMembrane w/o Membrane Eyrie Year Means 1979 1978 1977 1976 1975 1974 1973 1940 11 Pre- .320 .307 .280 .283 .273 .280 .295 .395 (n=15) (n=24) (n=26) (n=9) (n=8) (n=7) (n=3) .248 .226 .209 .211 .235 Ratcliff's Index Remarks (n=13) (n=16) (n=17) (n=9) (n=7) .289 (n=3) II Measurements taken around waist of egg. 1/ From second clutch of eggs. 11 Eggshells from Alberta, Saskatchewan, Montana; Anderson and Hickey, 1972. In Procceedings of XVth Int. Ornith. Congr. �Table 6. Chemical Residues in Colorado peregrLne eggs, 1978 (Values given are in parts per million) Compound 7:1,12:./7:1,2 15:1,3 15:2,1 15:2,2 29:1,3 30:1,3 30:2,1 30:2,2 30:2,3 29.07 40.77 30.17 8.35 20.16 17.01 19.35 18.85 DDE 7.89 7.66 DDD -- -- 0.15 0.11 0.84 DDT 0.18 -- 0.51 0.46 0.38 -- 0.15 0.10 0.11 0.11 Dieldrin 0.24 0.22 0.11 0.15 0.18 0.19 0.14 0.10 0.10 0.11 Heptachlor epoxide 0.29 0.30 0.63 0.69 0.59 0.38 0.71 -- 0.64 0.64 Oxychlordane - - -- 0.l3 0.15 0.13 0.10 0.20 0.18 0.19 0.18 I-' \.0 cis-Chlordane -- -- 0.11 0.13 B-BHC -- - - 0.16 0.18 0.16 0.12 0.17 0.19 Est. PCB 0.14 0.08 2.11 2.69 1.93 0.55 0.88 0.73 trans-Nonachlor cis-Nonachlor Endrin Est. Toxaphene HCB = None detected }j Corrected to 85% moisture content ]j example: 7:1,1 means Site 7, egg 1 of first clutch .. 0.18 0.19 0.89 0.90 �20 Egg Shell Thinning and Pestide Residues Peregrine eggs obtained from wild sites in 1979 under the augmentation program which failed to hatch have been submitted for chlorinated pesticide and PCB residue analysis and the results have not yet been received. Since eggs must be submitted intact, shell thickness measurements could not be taken and the data will be provided in a subsequent report. The shell thickness data for the 1978 eggs as well as those eggs which hatched in 1979 are provided in Table 5. While the data has not yet been subjected to statistical scrutiny, it would appear that there may beatrend to slightly thicker eggs in the past two years. Pesticide residue data for the 1979 eggs are not yet available from the laboratory and will be provided in a subsequent report. The data from the eggs analyzed in 1978 are provided in Table 6. Chlorinated Hvdrocarbon Residue Analysis in Prey Species Seven adult individuals each of potential avian prey species were collected within 5 miles of four peregrine eyries (sites 1, 6, 27, and 28). Four species were collected at site 1, five species at site 6, ten species at site 27 and five species at site 28. The samples have been prepared and submitted for residue analysis but the results have not been forthcoming in time for this publication and will be provided in a subsequent report. Prepared by: C.R. Gerald R. Crai Sr. Wildlife Biologist �21 JOB PROGRESS REPORT State of COLORADO ----~~~~--------------------- Project No. SE-3-2 ----~------------------------ Work Plan No. Job Title: II Endangered Wildlife Investigations Job No. 2 Physical and Biological Analysis of Colorado Peregrine Nesting Habitat Period Covered: Personnel: July 1, 1978 - June 30, 1979 J.E. Enderson, Colorado College; M. Berman, B. Busby, G. Craig, M. McWhorter, J. Patz, B. Pendleton and J. Rucks, Colorado Division of Wildlife· ABSTRACT The physical features for all historic and currently occupied Colorado peregrine eyries have been recorded and are presently being analyzed. Prey transects were conducted at 6 sites, of which 4 sites were repeated in an effort to compare information from different years. Results are being compiled and analyzed. �22 PHYSICAL AND BIOLOGICAL ANALYSIS OF COLORADO James H. Enderson PEREGRINE NESTING HABITAT and Gerald R. Craig P. N. OBJECTIVE The primary objectives of this study are to examine physical and biological features of peregrine eyrie sites in an effort to eventually delineate specific factors which favor occupancy by peregrines. This information will aid in delineating those habitats which should be protected or enhanced for future occupancy by the falcons. SEGMENT OBJECTIVES 1. Establish physical and biological parameters which are uniform at all eyrie sites currently or historically occupied by peregrine falcons. 2. Delineate those human activities by nesting peregrine falcons. 3. When no alternative method exists, adults at specific eyrie sites may be trapped after their young have hatched and radio packages will be affixed to them to monitor their movements anQ hunting ranges. 4. Assemble the data and prepare a report of the results for use in designating new and potential eyrie sites by wildlife agencies and land managers. Note - These objectives correspond American Peregrine Falcon Recovery METHODS The investigation conformed and disturbances which are tolerated to tasks 1111. and 1112. of the approved Plan (Rocky Mountain/Southwest Population) AND MATERIALS to the following broad procedures: Procedures. 1a. Annually, eight known historic and currently occupied peregrine eyrie sites in Colorado will be visited and the following physical aspects will be recorded: topography. geology, elevation, snow depth and precipitation, mean temperature, soil, presence and distance to water, and cliff characteristics. In addition, a series of photographs will be taken of each site. Each year, eight new sites will be surveyed until all known sites have been studied. lb. The vegetative types of the habitat within a distance of 15 miles of eight nesting cliffs will be cataloged whenever possible, appropriate Forest Service vegetative maps will be utilized and meadows and other �23 potential hunting areas will be located on maps. 1c. Through use of standardized census techniques, avian prey diversity and abundance will be calculated for the months of April, May, June and July. This will be done for at least two sites annually. Censuses will be run once monthly in two localities for each habitat type represented. 2. Human activities,land use practices and audio and visual disturbances will be noted at each site and within an area of 15 miles .of each site. If breeding pairs are present, they should be observed to note their reactions to potential disturbances. 3. At one or two sites annually, adult peregrines may be trapped and equipped with radio packages. This will be accomplished after the young have hatched and will extend for a period of approximately 3 weeks. The adults will be monitored as they embark upon hunting forays and locations will be triangulated primarily from the ground. Periodically, hunting adults will be tracked using a fixed-wing-aircraft to follow the radio signals. All locations of radio marked falcons will be recorded on appropriate topographic maps of the region. 4. Compile data forms and photographs taken during 1, 2 and 3 into notebooks. Prepared and submit reports of the results to appropriate state personnel, federal agencies and the Rocky MOlintain/Southwest Peregrine Falcon Recovery Team. Physical Analysis. Physical analysis of nesting cliffs and the immediate vicinity was accomplished by visiting the site and making a visual inspection of the area. Photographs were taken of the nest cliff and panoramas were compiled of the surrounding area from vantage points at the top of the cliffs. Vegetative information was also recorded during the visits and was further augmented from timber maps of the region. Elevational information was obtained from topographic maps of the region. Prey Abundance. Line-transect. counts of bird populations we re taken in June and July at the seven eyries under study. Transects 805 m (0.5 mi) long were established within 8 km of each eyrie to quanify bird populations within the hunting range of resident peregrines. Transects were usually set in the most prevalent plant communities in each region and in the nearby riparian habitat where possible. Bird counts were made between 0630 and 1000 hrs. by an observer walking the transect in 45-60 minutes, and then returning after a 15 minute pause. Species and numbers of individuals were recorded for all seen or heard, and unidentified birds were classed in three size groups. Individuals of each species for the "round trip" on the transect were divided by two to yield the average number of individuals seen on the 805 m �24 transect on a given day. These daily means were in turn averaged to give the mean number of individuals of each species for each transect in the study period. Each species was classified in one of three subjective categories of vulneraD,ility to peregrine predation, based on the relative amount of time each species was seen to fly in the open at least 15 m from the ground or other cover. "A" category includes forms often seen away from cover, e.g. Clark's nutcrackers; "B" includes species often seen in cover but given to occasional long flights in the open, e.g. robins and f LLcke rs ; "c" category includes species that seem infrequently subject to peregrine attack, e.g. rufous sided towhees. Large species such as buteos, ravens, and large waterfowl were given a "not applicable" (N.A.) rating. RESULTS AND DISCUSSION In 1979, physical data were collected for all 32 peregrine falcon eyrie sites in Colorado. The data include physical descriptions of the nest site and vegetational communities in the vicinity, as well as photographs. This information is currently being analyzed and a final report is in progress. Line transects were conducted to record avian prey species composition at the following eyries: 1 (2 Site Site 5 (2 Site 6 (2 Site 7 (2 Site 9 (4 Site 12 (2 Site 14 (2 and abundance transects) transects) transects) transects) transects) transects) transects) Transects which had been established in 1977 were again run at sites 5,7 and 14. Transects at site 1 which were conducted in 1978 repeated in 1979. The purpose of repeating a sample of the transects was to provide comparison between years. The transect information is being compiled and is not available for publication in this report. Details will be provided in the 1980 progress report. Prepared by: ogist �25 JOB PROGRESS State of REPORT COLORADO --~~~~~--------------- Project No. SE-3-2 Endangered ~~~~----------------- Work Plan No. II Job Title: Reintroduction Period Covered: Personnel: Job No. and Augmentation Wildlife Investigations 3 of Peregrine Falcon Production July 1, 1978 - June 30, 1979 J. Enderson, Colorado College; J. Hogan and S. Petersburg, National Park Service; T. Reed, S. Warner and C. Button, Bureau of Land Management; W. Heinrich and W. Burnham, The Peregrine Foundation; D. Berger; B. Busby, G. Craig, R. Gaines, R. Meese, B. Pendleton and W. Russell, Colorado Division of Wildlife. ABSTRACT A total of 25 young peregrines were released into the wild in Colorado through various techniques. Twelve young were fledged as a result of manipulation and augmentation efforts at wild eyrie sites. Ten young were released through hacking. One young released in 1978 returned to a hack site and fed with:the young released in 1979. Three young were released through a cross fostering experiment at a prairie falcon nest. �26 REINTRODUCTION AND AUGMENTATION OF PEREGRINE FALCON PRODUCTION Gerald R. Craig P. N. OBJECTIVE The objective of this study is to augment natural to sustain the wild peregrine population. reproduction in an effort SEGMENT OBJECTIVES 1. Augment natural production of peregrines by various techniques. 2. Monitor the results of the efforts, compile data and submit reports to appropriate state and federal agencies and the Rocky Mountain/ Southwest Peregrine Falcon Recovery Team. Note - These objectives correspond to jobs 222., 3133., 321., 322., 331., 332., and 3211. in the approved American Peregrine Falcon Recovery Plan (Rocky Mountain/Southwest Population). METHODS AND MATERIALS Augmentation efforts will be undertaken according to the following methods: 1a. Breeding pairs of peregrines will be observed to determine dates of initiation of egg laying. Within a week to 10 days after completion of the full clutch of eggs, the eyrie will be visited and all the eggs removed and artificially incubated. Approximately two weeks after removal of the eggs, the pair will recycle and lay a second clutch. The second clutch mayor may not be replaced with dummy eggs which the adults will be permitted to incubate. Dummy eggs should be substituted in situations where there may be concern about the adults' ability to incubate the eggs without breaking them. After acsuitable period (if dummy eggs were substituted), the site will be revisited and the dummy eggs will be replaced with chicks from the eggs which were incubated and hatched in captivity. If the adults were permitted to hatch their own eggs, they will be permitted to continue to rear and fledge them. Since there will be a 28 day difference in age between the young produced from the first clutch and those produced from the second clutch, the young from the first clutch will be placed in other wild eyries containing similarly aged broods. Several representatives may also be retained for captive propagation purposes if similarly aged broods cannot be located. lb. As in 1a., initiation eyrie site dummy eggs breeding pairs will be kept under surveillance to determine of egg laying. Shortly after completion of the clutch, the will be visited and all the eggs removed and replaced with which the adults will be permitted to incubate. Since the �27 wild eggs usually are thin-shelled they will be artificially incubated to avoid their being accidentally crushed by the adult. After several weeks, the site will be revisited and young peregrines will be exchanged for the artificial eggs. Up to four young may be placed at each site to assure the maximum number of young are fledged. 1c. Captive produced young may be released at unoccupied or potential sites without the benefit of protection or care from adults through the technique of Ithacking". Young falcons of three to four weeks of age will be placed on a suitable ledge at a potential reintroducton cliff site. They will then be cared for and fed by human attendants until they are flying and capable of feeding themselve. In this manner, the young falcons will return to the site at which they were reared and hopefully breed. This approach requires constant attendance and observation in order to protect the vulnerable young and insure they have sufficient food while they are in the eyrie. Because of this. the first two techniques will receive priority attention. If there are no additional adult breeding pairs as required by approaches 1 and 2, then young will be placed into the wild using this technique. RESULTS AND DISCUSSION Data gathered during the period from March 1, 1977 through July 1, 1977 had been reported in Federal Aid P-R Progress Report. January 1979. As of July 1, 1977 the investigation was shifted to the Endangered Wildlife Investigations and the 1978 breeding season results are presented here. Recycling Efforts Pairs at sites 14, 27 and 30 were kept under observation early in the breeding season and recycled to produce second clutches of eggs. Although the pair at site 9 were kept under surveillance from initiation of courtship, they qpparently did not initiate egg laying until late in the season, so the decision was made not to induce them to recycle. It is possible that this pair may have laid a clutch of eggs early in the season and abandoned them, but this could not be confirmed. It is interesting to note that the pair at site 9 laid their clutch during the same period as the three recycled pairs. Following is a synopsis of events occurring at the three sites whic.h were recycled: Site 14 The clutch of 4 eggs was completed April 25. Usually au egg is laid every other day, which puts the date of deposit of the first egg at April 19. Four days after completion of the clutch (April 29), the eyrie ledge was visited and all the eggs were removed and transported to Fort Collins. Colorado for artificial incubation. Three of the four eggs eventu:el1y hatched and survived. Orie egg died early in development. Approximately 20 days after removal of their eggs, the pair relocated to the same nest ledge they had used �28 on a pinnacle in 1978 and initiated egg laying. The site was visited on May 27 and the clutch of three eggs were removed and replaced with plastic replicas which the adults accepted and incubated. The second clutch of eggs was also transported in a portable incubator to Fort Collins. All three of the eggs in the second clutch were fertile and hatched. One young died after hatching, but the other survived. On June 19 the climbers returned and replaced the plastic eggs with 4 peregrine nestlings which the adults immediately accepted and fed. Approximately 10 days prior to fledging, two young fell from the nest ledge and a third fell to a lower ledge the next day. One young died of injuries sustained in the fall. A second young made its way up the talus slope below the eyrie to a ledge on the face but could not be relocated when the cliff was visited. It is assumed that it was killed and carried off by a predator, probably a great horned owl. The adults continued to feed the remaining two young despite the fact that they were on separate portions of the cliff. It is puzzling that the young fell from the ledge since four young successfully fledged from the same location in 1978. The falls appeared to happen when one of the young failed to negotiate a short gap in the ledge and another young fell along with a rock it was perched upon. It is probable that these accidents would not have occurred had the pair been permitted to remain at the location they selected for their first clutch. Site 27 The pair completed their first clutch of three eggs on or about April 20. Climbers visited the eyrie ledge on April 22 and removed the three egg clutch. The eggs were transported to Fort Collins where one hatched and survived. All eggs were fertile but two died early in development, possibly as a result of faulty incubation in transit. The pair relocated approximately 200 yards down the cliff and completed their second clutch between May 10 and 13. The nest ledge was visited on May 20 and the three egg clutch was removed and replaced with plastic replicas, which the adults accepted. All three eggs in the second clutch were fertile and hatched. One chick died after hatching and the other two survived. The site was rev~sited on June 6 and the dummy eggs were replaced with 4 peregrine nestlings which were subsequently reared by the adults. All four young successfully fledged from the site. It is interesting to note that when the second clutch was removed a hardened piece of mud was attached to the side of one egg. A week previously, a late spring storm caused intermittent snow and rain for several days. Undoubtedly, the moisture caused some of the nest substrate to adhere to the egg. Because of the mud, the egg could not have hatched since the adult could not turn it properly. Secondly, one egg hatched 4 days earlier than the other two indicating that the female was forced to initiate incubation upon deposition of the first egg rather than delaying until the clutch was complete. This was probably done to keep the egg from chilling during the storm. Since the egg hatched 4 days early, it is likely the remaining eggs would have not have hatched since incubation normally terminates within 48 hrs after the hatch. �29 Site 30 At the time the nest ledge was visited on May 4, it was estimated that the pair had been incubating their clutch of 4 eggs for a week to ten days. Three of the eggs were normal, but the fourth was one quarter the size of the other eggs. This egg typt cal.Lycalled a "poult egg - was not fertile. Such undersized eggs are typically laid when birds initiate their egg laying careers, but have been produced even by seasoned breeders in captive situations. This appears to be the first time such an egghas been encountered with wild nesting peregrines. All eggs were removed and transported to Fort Collins for incubation as previously described. The three normal sized eggs were all fertile but died early in development, possibly as a result of faulty incubation while in transit. The pair moved approximately ~_mile down the escarpment and completed their second clutch between May 22 and 25. The site was visited May 26 and the four eggs were replaced with plastic replicas which the adults accepted. The second clutch of eggs were treated in the same manner as the first. All four eggs were fertile and two young were hatched and survived. On June 20, four nestling peregrine chicks were placed on the nest ledge and the dummy eggs were removed. At the time of fledging, only three young could be counted in various locations along the cliff despite two days of observation. Thus it is assumed that one young died prior to that time. Augmentation Efforts Only one wild eyrie (site 9) was augmented without the recycling effort. In reality, augmentation is identical to recycling, except the pair is not induced to produce a second clutch. When the eggs are first encountered, the eggs are simply replaced with plastic replicas which are subsequently exchanged for young peregrines. This procedure permits the wild eggs to receive more gentle treatment under controlled conditions, incubation parameters of temperature and humidity can be adjusted to the requirements of individual eggs, and finally the optimum member of young can be returned to the pair to be reared. Generally, this approach is taken instead of recycling when one or a combination of the following occurs: 1. Sufficient information is not available to determine the exact age of the eggs. It is unwise to attempt to recycle pairs which have-been incubating eggs in excess of 10 days. As more time is invested in incubation, the greater the likelihood the adults will abandon a renesting attempt. 2. Recycling would delay the nesting effort sufficiently into the summer to reduce the chances of successful f Ledg i.ng of the young. Extreme summer temperatures can stress the young at critical periods and prey may not be as abundant later in the season. 3. The pair may relocate which is inaccessable augmentation. to an inferior nest site or select a site and thereby curtail egg replacement or brood Site 9 fulfilled each of these conditions and the decision was made not to recvcle them. Further, this pair had not demonstrated their reproductive �30 capability. When the site was augmented in 1977, the two young peregrines which were placed in the nest failed to fledge. In 1978, the pair abandoned the dummy eggs about one week after the exchange. Therefore, it was decided that young prairie falcons should replace the dummy eggs as soon as possible after the wild eggs had been exchang~d for plastic eggs. Events occurred in the following sequence: Activities throughout April indicates that the pair may have selected an eyrie and partially completed a clutch of eggs, but aborted. -, May 5, the pair were discovered incubating 3 eggs. May 20, the 3 wild eggs were replaced with two 10 day old prairie falcon chicks, thus eliminating the intermediate step of replacing the eggs with plastic replicas. June 6, the site was revisited and the prairie falcon nestlings were replaced with three 21 day old peregrine chicks. Time was critical since the prairie falcons were well developed and approaching the flight stage. The adult peregrines immediately accepted and reared the younger peregrine falcon nestlings. All three young fledged successfully. Three young were also produced from the wild eggs. It is unlikely that one of the young would have hatched if the eggs remained with the wild pair since it hatched eight days later than the other two. The adults would have ceased incubating shortly after the hatch of the two eggs. The delayed hatching time of this egg is attributed to a late spring snowstorm which apparently caused the pair to temporarily cease laying activities and initiate full time incubation in order to protect the first two eggs from chilling. It is a matter of conjecture whether or not another egg was reabsorbed or laid elsewhere in the interval after the second egg was laid and before the last egg was deposited. Results of the recycling and augmemtation efforts are surrnnarized in Table 1 and 2. Table 1 summarizes the assumed productivity of the four sites had they not received manipulation. Even though the act of manipulation obscures the natural productivity inferences can be made to estimate the probable natural reproduction had manipulation not occurred. Generally, the estimated natural production is optimistic since it is not likely the wild pairs would have been as successful at hatching the thin shelled eggs as the Peregrine Fund. As can be seen from Table 1, natural production of the four sites would have been 14 eggs which would have yielded 12 young and fledged 10. Actual production due to manipulation yielded 24 eggs, hatched 15 young and fledged 12. The number of young per pair was increased from 2.5 to 3.75. However overall fledging success was increased a lesser amount from 2.5 to 3.0 primarily due to loss of young from natural causes after augmentation. Portable Incubator Problems It is essential to have a portable incubator which is reliable and keeps eggs at a constant temperature in transit from the wild nests to the incubation facilities. Such a system has undergone gradual metamorphosis from a styrofoam ice. chest with a light bulb in 1976 to a sophisticated solid state unit utilizing a water bath in 1979. Unfortunately the only true test of the machinery seems to occur during the field season with wild peregrine eggs. After the first clutches from sites 27, 14 and 30 had been transported, hatch ability was reduced And there "(vasan indication that �31 Table 1. Assumed natural production of manipulated sites Site No. No. of Eggs Which Would Have Been Produced No. of Young Which Would Have Hatched No. of Young Which Would Have Fledged 9 3 2 2 14 4 4 2 27 3 3 3 30 4 3 3 14 12 10 Totals 3.50 eggs per pair; 4.0 young hatched per pair; 2.5 young fledged per pair. Table 2. Site No. Actual Productivity of manipulated sites No. of Eggs Produced No. of Young Hatched No. Young Returned Per Pair No. of Young Fledged 9 3 3 3 3 14 7 6 4 2 27 8 2 4 3 30 6 4 4 4 24 15 15 12 Totals 6.0 eggs per pair; 3.75 young hatched per pair; 3.75 young returned per pair; 3.0 young fledged per pair. �32 six of the eggs had probably died at the time of transport. Two of three eggs from site 27 died, one of the four from site 14 failed, and all of the eggs in the first clutch from site 30 succumbed. Even though the eggs are held a relatively short time in the portable incubator, adverse incubation temperatures during that period are quite critical. The eggs from site 30 were held the longest in transit (an estimated 12 hours). Subsequent rechecking of the incubator established that where the eggs contacted the water heater, the temperature was 98.50 F. the temperature gradient dropped to 88.50 F. on the opposite site pf the egg. The incubator was modified so the eggs were suspended between two water heaters and the embryo mortality declined significantly. Only two eggs of the 13 remaining eggs hatched. The two eggs which failed were part of a four egg clutch resulting from the recycle of site 30. Again, the eggs were held for a prolonged period (approx. 24 hrs.) in the incubator and further adjustment may be necessary. Reintroduction Efforts Two locations were selected for hack activities. One was in the vicinity of site 18 and was activated for the second year. In 1978, four young were successfully released from the hack box. In 1979 five young were successfully released from the site. During that period, a yearling male from the previous year returned to the hack box to receive food along with the five recently released young. The second hack site was established adj acent to site 25 and seven young were originally placed in the hack box. Two weeks after all young were on the wing, two females were retrapped and will be held in captivity until adults, then released with adult males at potential nesting cliffs in an effort to establish breeding adults at sites to shortstop juvenile and yearling mortality. Five young were eventually fledged from the hack site near site 25. Cross Fostering Efforts Although it had not originally been planned, an opportunity arose to replace broods of prairie falcons with young peregrines at two eyries near site 11. Both prairie falcon eyries were exceptionally late and one had a brood of small young in early July and the other was incubating two eggs. Since additional young were available for placement in the wild, the decision was made to place 3 young peregrines in each of the two sites. Unfortunately, one site failed for unknown causes, but two of the young peregrines were saved and only one perished. The second cross foster site experienced a black fly outbreak and two young died as a result of a blood parasite. The eyrie was dusted with pyretharon and the two young remaining from the first site were placed with the lone youngster at the second site. All fledged successfully. Time will tell if this release method is effective. Prepared by: G. nior Wildlife Biologist �33 JOB PROGRESS State of __ ~C~O=LO~RA~D~O~ Project No. _ SE-3-2 Work Plan No. REPORT Endangered II Job No. Falcon Habitat Protection Wildlife Investigations 4 Job Title: Peregrine Activities in Colorado Personnel: H. Browning, G. Craig, R. Meese, and P. Waters, Colorado Division of Wildlife; D. Cook, T. LaMay and C. Rackham, u.S. Forest Service, J. Randall, u.S. Fish and Wildlife Service. ABSTRACT The U.S.Forest Service and Colorado Division of Wildlife entered into a Memorandum of Understanding to manage and protect an occupied peregrine falcon nest in southwestern Colorado. Activities involve surveillance, a Special Closure to prevent trespass, and supervision of tours of Indian ruins adjacent to the site. �34 PEREGRINE FALCON HABITAT PROTECTION IN COLORADO ACTIVITIES Gerald R. Craig P. N. OBJECTIVE The objective of this study is to document man-caused peregrine nests and implement procedures to eliminate disturbances at the activities. SEGMENT OBJECTIVES 1. Reduce or eliminate sites. human disturbances occurring at specific eyrie 2. Protect important feeding areas or eyrie sites located on private land through cooperative agreement, lease, purchase or exchange of estate. 3. Prepare annual reports of the results of the endeavors. Note - These objectives correspond American Peregrine Falcon Recovery to tasks 1222. and 123. in the approved Plan (Rocky Mountain/Southwest Populations). PROCEDURES 1. Two observers will be stationed at eyrie sites and will keep them under constant surveillance from initiation of egg laying until after the young have fledged. The observer will be situated in such a position that his presence will not disturb the falcons and yet he will be capable of viewing the vicinity and note any intruders. The observer should be present at the site during the daylight hours. If possible, the observer should have access to a packset radio to communicate with Division of Wildlife field personnel should assistance be required. This work will also be coordinated with the Special Agent-in-Charge at the Denver Federal Center. In addition to maintaining surveillance, the observers will be expected to keep field notes of various activities of the breeding peregrines. 2a. When important feeding areas of peregrines are located on private lands, an effort will be made to contact the landowner and negotiate to assure that the area remains suitable as a hunting area for peregrines. In general, the landowner will be encouraged to continue to provide the habitat types preferred by key prey species, continue to plant crops which support the prey or undertake other activities which continue to benefit the peregrine's prey. No funding request for this activity is included at this point since it is hoped ·that negotiations may be accomplished without disbursement of funds to the landowner. �35 2b. One currently active peregrine eyrie site is situated upon private land. The landowner has been contacted and has been made aware that the peregrines reside upon his land. He has expressed an interest in assuring the existence of the site and should be approached to enter into a cooperative agreement to continue to protect the site. Should he contemplate disposing of the land, a clause in the agreement should provide that the Division be provided the first option to acquire the land. Possible lease of the land by the Division should also be investigated. 2c. Where no other alternative protective method exists, land acquisition will be undertaken to purchase the actual nesting cliff and those feeding areas which must be maintained to insure the integrity of the area for occupancy by peregrines. When such actions are warranted, this job will be amended to request funding for land purchase. 3. Prepare annual reports of the activities accomplished under this job and provide them to the appropriate state and federal personnel and the Rocky Mountain/Southwestern Peregrine Falcon Recovery Team. MATERIALS AND METHODS Two observers were employed jointly by the U.S. Forest Service and the Colorado Division of Wildlife to keep an active peregrine eyrie under surveillance from April 20, through September 15, 1978. The site was kept under surveillance during daylight hours from several vantage points which provided views of the surrounding area and nesting cliffs. The observation points were located at a sufficient distance from the nest site to avoid disturbance of the falcons. Binoculars and spotting scopes were frequently used to scan the vicinity for intruders, although it was soon discovered that the falcons were the best watchdogs and would vocalize and dive at intruders in the vicinity of nesting cliff. The observers were also equipped with radios to contact local Wildlife Conservation Officers or Forest Service field personnel in case assistance was required. In addition to protecting the falcons from direct human harassment, the observers recorded the falcons' behavior throughout the breeding season. Particular effort was made to record any response the falcons made to man-related disturbances. RESULTS AND DISCUSSION The peregrine nest site near Chimney Rock in the San Juan National Forest poses several unique problems. The first is that a pair of peregrine falcons have occupied the site since prior to 1943. Second, during a period of 9 years, the falcons were not present and the Forest Service began development of Indian ruins adjacent to the site. Plans were underway to continue development and expand access to permit tours of the ruins when the falcons returned to their historic nest in 1973. As a result of their presence, the �• 36 J Forest Service curtailed further development of the tours because of potential disturbance to the falcons. This caused much concern and focused public attention on the falcons. In 1976, the Forest Service and Division of Wildlife decided it would be prudent to station two observers to df.scoucage trespass and offer some protection to the falcons while management plans were formulated and appropriate land closures implemented. In March of 1977, the Forest Service and Division of Wildlife consumated a Memorandum of Understanding concerning management of the Chimney Rock Site. The agencies agree to meet annually to discuss and app rove activities proposed in the vicinity which might have impact upon the falcons or their hunting areas. In keeping with the spirit of the agreement, the Forest Service enacted a Special Closure within approximately one-half mile radius of the site on November 17, 1978. The closure limits access to the area by unauthorized persons from February 1 through September 30 of each year. Provision was made to permit tours of the ruins which were immediately outside the closure so long as the tours were supervised by the Forest Service and occurred at predetermined times. As agreed upon at the coordination meeting, experimental tours were conducted twice weekly in June to ascertain the response of the falcons to such congregations. Generally, tour sizes were smaller than in 1978, probably due to shift in responsibility from the Durango Chamber of Commerce to administration by the Forest Service. The Forest Service limited tour sizes to less than 35 persons and restricted the length of the visit to the upper ruins to an hour. At no time did the falcons show any reaction to these tours. This is probably due to a combination of factors: 1. The falcons occupied an eyrie on the pinnacle furthest from the upper ruins. The eyrie was partially buffered from view of the ruins by an intervening pinnacle which provided more visual security to the falcons. 2. Tours were better organized and controlled. The most severe reaction to a tour occurred July 14, 1978 when a large group of children visited the upper ruins. Undoubtedly, the noise and activity caused the falcons to react. 3. Tours were generally spread through the week rather than one day after another. Thus, the falcons probably did not become sensitized to their occurance. 4. Visits to the upper ruins (which are most visible and closest to the falcons) were limited to an hour and occurred at the end of the tour. In this way, the most disturbing activity took place at the last. 5. The tours were conducted late in the morning around 10:00-11:00 AM which was the period that the falcons were usually away from the nest hunting. After August 1, the frequency of the tours was increased with no adverse response from the falcons. to three per week Trespass problems did increase in 1979 with approximately eight cases of illegal entry occurring. None of the entry seemed to be associated �37 with the falcons, people were either hiking in the vicinity or going up to the ruins. As a result, it is evident that the closure signs need to be placed in more conspicuous locations near entry points. Presently, the closure notices are posted at the one-half mile boundary which is up to three-quarters the distance from the entry points. At that stage most people are reluctant to turn around. This year, the U.S. Fish and Wildlife Service assigned agents to provide surveillance from mid June through mid July. This effort seemed redundant and was not properly coordinated. In the future the effort must be coordinated to maximize manpower use. Special Agent Jack Randall did provide valuable assistance by arranging his schedule to coincide with periods the surveillance crew was absent, thus keeping the site under 24 hour observation. His dedication bore fruit when he observed the young peregrines fall from the nest ledge. Although little could be done, this solved what would have been a mystery in the disappearance of two young. Publicity about this site makes it the best known peregrine eyrie in Colorado. This popularity is bound to increase the number of trespass problems as well as the possibility of illegal removal. Surveillance activities will have to continue and must be coordinated among the responsible law enforcement agencies to make most efficient use of manpower. On August 2, and Wildlife and plan for existence of Prepared by: 1979 a Biological Opinion was obtained from the U.s. Fish Service which agreed that the Forest Service's management the site would "contribute to the conservation and continued the species." c. Gerald R. r Wildlife Biologist �38 JOB FINAL REPORT State of COLORADO --~~~==~----------- Project No. SE-3-2 -------------------- Work Plan No.II Endangered Birds: Job No. 5 ---------------------------------- Job Title __~D~e~t~e~rm~i=n=a=t=1=·o=n=_o~f_=th~e_=P~o~t~e=n~t~i~a=l~f~o~r~E=s=t=a=b=I=1=·s=h~1=·n~g_ _ Population Period Covered: Personnel: of Greater Sandhill 1 July 1978 through Cranes in Southwestern Colorado 30 June 1979 W. Graul, C. Fox, H. Geduldig, G. Seville, J. Frothingham. ABSTRACT From 7 September 1978 through 6 June 1979 aerial and terrestrial investigagat ions were conducted to locate habitats suitable for a potential reintroduction project to establish a second Colorado population of Greater Sandhill Cranes (Grus canadensis tabida) on historicgl nesting grounds. From 36 potential sites located, 4 were rated to have at least moderate potential, 1 exhibiting high potential, for nesting reintroduction. Plans for obtaining, housing, and feeding of captive cranes are discussed. A 10-year strategy for the reintroduction is presented along with projected expenses. Project Objectives(s): Determine whether it is feasible to establish a nesting population Sandhill Cranes in historical habitat in southwestern Colorado. Segment Objective of Greater (s): 1. Verify the presence of suitable crane nesting habitat in southwestern Colorado and map the location of this habitat. 2. Determine the feasibility of establishing a new crane nesting population based on existing knowledge, and if feasible, develop the procedure to be followed. �39 ACKNOWLEDGEMENTS This report is largely the synthesis of information gathered from numerous governmental and pri~ate agency personnel. I wish to express my gratitude for input from George Archibald and Chris LaRue of the International Crane Foundation (ICF) , David Ruff and John Curran of the United States Forest Service (USFS), Gene Patton and Bill Wilson of the Arapaho National Wildlife Refuge (NWR), Melvin Nail of the Monte Vista NWR, Phil Bailey and Bruce Baker of the Bureau of Land Management (BLM) , ·Ernest House of the Ute Mountain Ute Tribe, T.E. Taylor of the Bureau of Indian Affairs (BIA) , Mr. Hanson and Charles Holcomb of the Soil Conservation Corps (SCS) , and Jim Houston, Tom Henry, Tom Sherill, Richard Fentzlaff, Wayne Russell, Joe Frothingham, and Gordon Sevelle of the Colorado Division of Wildlife (CDOW), Special gratitude is extended to my supervisor, Walter Graul, Nongame Wildlife Research Leader of the CDOW for both his guidance in field investigations and attention to the numerous details necessary for the completion of this report. �40 INTRODUCTION This project was undertaken to determine the feasibility of establishing a breeding population of Greater Sandhill Cranes (Grus canadensis tabida) on historical nesting grounds in southwestern Colorado. Historically~ the Greater Sandhill Crane nested in those regions of Colorado west of the Continental Divide, from the San Juan Mountains north to the Wyoming State Line, in the mountain parks up to 9500 ft (2896m) (Bailey and Ni.edrach 1965). With the settlement of these regions in the early 1900's, the Colorado nesting crane populations were reduced solely to those few birds remaining in northern Colorado (Routt County, north of Hayden and Steamboat Springs). With subsequent protection of these few remaining birds, the Routt County population had slowly increased to a 1977 population estimate of about 150 breeding cranes. Although this population has experienced a steady increase in numbers, and a slight expansion of its breeding range, its survival may be in jeopardy from an increase in mining, timber, and recreational activities (Bieniasz 1978). While mitigation of these activities and protection of these cranes must continue, the establishment of a second, additional nesting population on its historical breeding grounds in Colorado would contribute in upgrading this subspecies' current legal status from "Endangered" (Colorado Wildlife Commission 1973) to one of various levels of security above "Threatened". Since (1) young and/or adult Greater Sandhill Cranes can be obtained readily from other states (e.g., Idaho), (2) captive male Sandhill Cranes have successfully attracted, nested, and reared young with wild migratory females (Longley 1970 and Konrad 1975), and (3) there appears to be a strong attraction for mated Sandhill Cranes and their young to return during subsequent spring migrations to their most recent nesting site (Longley 1970), the establishment of an additional nesting population seemed worthy of further investigations. This report documents the location of potentially suitable Greater Sandhill Crane nesting habitat and describes a methodology feasible for reintroduction of a nesting population in this habitat. All cost estimates are as of 1 July 1979. �41 LITERATURE REVIEW Nesting Habitat: Taylor (1975) found that Sandhill Cranes on the Hiawatha National Forest in Michigan prefer to nest in bogs and marshes with upland openings within several kilometers for feeding. The cranes nest near water. They prefer sparse ground cover, and will use pole size timber stands with a sparse understory for nesting. Clearcuts of aspen and jack pine and prescribed burns will maintain this sort of habitat. Beaver ponds will remain active if aspen stands are allowed to flourish near streams. Taylor determined that isolation of nest sites is of prime importance to cranes. Drewien (1973) maintained that cranes at, the Greys Lake National Wildlife Refuge in Idaho prefer to nest in 1) west meadows or marsh edges less than forty-five meters from shore, 2) islands, 3) dry upland meadows, 4) marshes greater than forty-five meters from shore, or 5) upland dikes. He found that vegetation, and in some pastures cow manure is used. Cranes will also nest on abandoned muskrat lodges. Drewien observed that the nest site, roost site, escape cover, wate r , and feeding meadows are located in contiguous area. He also stated that suitable nesting habitat is avoided if it is close to a road, unless it is screened by tall vegetation. Littlefield and Ryder (1968) surveyed nests on the Malheur National Wildlife Refuge in Oregon and found that all nests were located in or near free water except for two which were located in a dry meadow two hundred yards from water. Three nests were among willows while 118 were in open meadows and marshes. Nests were constructed mainly of the previous years' bulrush and burreed. They observed that water, nesting cover, and a feeding meadow were essential components of all territories. Nests were deserted after human disturbance or water level fluctuations. Walkinshaw (1949) noted that in general Sandhill Cranes usually nested in a large marsh that was open with shallow water and that contained dense stands of grass sedge, phragmites, rushes, and leather leaf. The nest site was located in open or semi-open habitat where sedges and rushes or other tall growth hide the crane while on the nest or when walking in the immediate area. In Idaho, he found that no shrubs were right around the marsh, but on one side there was a row of willows behind which was a nest. In Michigan the nest sites varied from being in shrubs in the marsh to being 500 meters away from the marsh. Most nest were found in water, some on open islands. Water depth averaged 17.52 cm. Walkinshaw stated that cranes don't like human or animal intrusions and prefer remote regions isolated by brush, forests, mountains, or buttes. Blake (1974) studied the Greater Sandhill Cranes in Colorado. She found that here cranes nest in willow swamps along streams, often on old beaver dams. In California Park the willows are bordered by sage; in other places they are bordered by grassy areas. Blake found that cranes require minimal disturbance, feeding meadows, nest cover, and water. Nests are constructed in sage, on beaver dams or muskrat lodges, or in willows around beaver ponds. Cranes will not nest where they are highly visible to humans. In 1975, Blake studied the Colorado cranes as they arrived at �42 their staging areas and move slowly up to their breeding grounds. The cranes arrived at their staging area April 1, and began daily movements up the willow drainages following snowmelt. Due to a late snowfall nesting was delayed for two weeks,_ and started 20 May. Blake watched 35 pairs, and noted that most nested by the willows, while four nests were found on beaver dams, one on a lodge, one in sage, and one on an irrigation ditch. Cranes in Colorado were again studied in Routt County in 1976 and 1977 by Bieniasz (1978). She located 40 breeding pairs during her two-year study. They nested along small willow-bordered streams in sage brush parks. Cranes need adequate water adjacent to the nest, nest cover, and nearby feeding sites. Bieniasz outlined critical sandhill crane habitat as staging areas used in spring and fall migration, and potential nesting and loaf areas free from human disturbance, below 9500 ft. elevation and within one quarter mile (0.4 km. ) of any willow-lined drainage that carries water through June. Bieniasz located 19 nests in California park during 1976 and 1977. One. nest was used both years. Breeding pairs nested on small drainages, usually one pair to a drainage. On larger streams a few pairs were found. Nest sites are often separated by ridges along the creeks. Nests were composed of willow twigs from leftover beaver cuttings and grass. Six nests were located where water was present only during May, June, and July. However they were all within one meter of slow moving water and were surrounded by dense cover, generally willows. Bieniasz determined that fishermen, sheep, and cattle were the major disturbances to the crane population. Sheep that were driven through the nest areas could cause nest desertion. Cattle would feed together with cranes, and seemed to.have less effect on them that the sheep. Fishermen wading through the willows often flushed cranes from their nests. Nesting History in Colorado: The Greater Sandhill Crane was a regular migrant in Colorado, particu~ larly in the western counties (Bailey and Niedrach 1965). Records of early naturalists and explorers indicate that sandhill cranes were abundant as a breeding species in the western mountain parks. In 1876 Carter found three nests at the mouth of the Blue River and one along the Colorado (Grand) River in 1877. Drew (1881) recorded the Sandhill Crane at Animas Peak in San Juan County. Warren (1904) found a nest on a mud bank of a small lake in northwestern Gunnison 'County. He describes the location as a plateau of approximately 8000 feet, between Ragged Mountain and Muddy Creek. The area contained ponds and lakes from 60 feet to 100 yards in diameter. He found the nest on June 5, 1903, 20 feet from the shore of a small lake. The nest was made of marsh grass. Peters (1925) cites the breeding areas of "Megalornis canadensis Mexicanus" to include Loveland, Middle Park, Gunnison, and San Juan Counties. Reintroduction of Cranes: As no reintroductions of cranes have thus far been accomplished, all possible methods must be investigated. Konrad (1975) has surveyed previous reintroduction attempts that have been made in Japan and the United States. �43 Based on his study he has described four methods when reintroducing captive cranes into the wild. first step is to select suitable habitat. It is area that has surrounding habitat for dispersing that areas selected should be on preserves which development. that may be successful He determined that the important to choose an young cranes. He stated will not be subjected to The first method of reintroduction that he suggests involves transplanting via the use of foster parents, based on the work of Rod Drewien at the Greys Lake National Wildlife Refuge in Idaho, where Greater Sandhill Cranes are being used as foster parents for Whooping Cranes. This method has sh~Nll promise, however, a suitable foster parent must be present in the chosen transplant site. Where no foster species is available, Konrad determined that pairs of captive-raised cranes may be wing-clipped and placed in a large enclosure in suitable habitat. The pen would contain a shelter and the cranes would be provided with food. After four weeks of acclimatization the cranes could be allowed to roam. They could establish a nesting territory the first summer and hopefully nest during the following spring. This method may not be feasible with migratory birds, however, because they would not have learned the migration route. Konrad suggested capturing migratory birds in the fall and holding them over the winter. They could be placed in pens in spring, and after a period of acclimatization they could be allowed to roam in the area. A winter feeding station may suffice instead of housing. He stated that clipped wing feathers could be pulled after the birds have nested, which will permit them to fly. Birds can be expected to return to the area where they first nested successfully. Another method he described would be to place captive, wing-clipped pairs in open-topped enclosures. They would not be released, and would hopefully raise young. The young would be allowed to fly, and would be able to disperse into the surrounding areas to establish territories and nests as wild birds. When using captive pairs it is important to determine the sex of each bird, as homosexual pairs form in captivity (Archibald 1974). Feather pulp analysis, chromosomal examination, and Unison calling (three calls of the female to everyone of the male) can be used to judge the sex of the individuals in a pair (Konrad 1975). The fourth method considered by Konrad is to place captive two-year old cranes in open-topped enclosures in areas located under the migration route. The cranes could then attract mates from the wild flocks. When a pair bond is formed, the pair would be allowed freedom of movement in the area. The captive bird could be kept wing-clipped until the pair nested, after which the clipped feathers could be removed so the captive may recover flight ability. The wild bird will presumably show the captive bird the migration route, however, until the birds have nested successfully the captive will have to be housed during the winter. The wild bird may either migrate in th.e fall, or remain withi.ts mate in winter. �44 There are several reports of captive cranes attracting wild mates. Longley (1970) wrote of a young crane confiscated from a farmer. The crane was wing-clipped and allowed to roam over thirty acres at the Carlos Avery Wildlife Area in Minnesota. A female crane flying overhead joined the captive in early spring and they remained together until the female migrated in fall. She returned to her mate in spring, and they nested. They were unsuccessful the first summer, but raised two young the following year. The young followed the female on migration. The next summer the pair produced one chick, and two more were reared the year after. Thus, in three years the pair produced five young. Five cranes were seen flying over the pair, and these may have been their young returning to the area. A migrating female Greater Sandhill Crane paired with a captive male at the International Crane Foundation in Wisconsin in March 1975 (Konrad 1975). They were allowed to roam until June. Then the male's primaries were clipped and he was placed in an enclosure with a shelter and fed pellets. The female roam~d free. They regarded the enclosure as their territory and would return to it after foraging outside. Hyde (in Konrad 1975) reported that his captive female and a wild male Greater Sandhill Crane paired and reared young. In Japan, Takahashi (in Konrad, 1975) captured five wild male Manchurian Cranes, wing-clipped them, and placed them in large breeding enclosures. Two died, but three succeeded in attracting and mating with wild females. Takahashi hand-reared the first clutch and allowed the cranes to raise the second clutch. The females were not wing-clipped and the young could fly with them. Three young raised in this way have joined wild flocks. Reintroductions of Other Species of Birds: The Canada Goose (Branta canadensis maxima), the Trumpeter Swan (Olor buccinator), and the Nene Goose (Branta sandvicensis) have been successfully reintroduced in many areas. The techniques used in these transplants may be helpful for crane reintroduction projects. Konrad (1975) cited the basic procedures for transplanting the Canada Goose to include the use of immature, wing-clipped birds, places in holding pens for acclimatization, then released in small groups to act as decoys to attract wild geese. Like the Sandhill Cranes, Canada Geese have a strong tendency to return to natal areas, and to areas where they first nested. Szymczak (1973) describes a Canada Goose release project on the eastern slop of Colorado, north of Boulder. Goslings or eggs were used, the eggs incubated by White Rock hens. The goslings were wild-trapped at 7 - 8 weeks of age, using a drive trap and crates for transport. The birds were held at the Wildlife Research Station in Fort Collins and were integrated with the hand-reared birds. At 9 - 10 weeks they were released. A continuous supply of grain was provided, using self-feeders. Feeding continued for six years, then was discontinued except during hunting season. An Air Aqua unit was used to keep the lake water open during the winter. Hunting, fishing, and trapping were prohibited for twelve years. The population increased from 500 in 1963 to 1300 in 1967. Trumpeter Swans were established at the Malheur National Wildlife Refuge in Oregon (Konrad 1975). The birds were wing-clipped and held in a semi- �45 captive conditions. They were held in pens over the winter, and were allowed to fly the following summer. A fe~v mated pairs of Nene Geese were placed in a 1 hectare open-topped enclosure in suitable habitat. The adults were wing-clipped but the young could fly. The young dispersed and established breeding territories. Rearing Captive Cranes: Methods of rearing cranes were discussed by Archibald (1974) of the International Crane Foundation. The diet used there for immature cranes is a conbination of Turkey Starter crumbles plus several vitamins and minerals (Table 5). Chicks are fed this diet from 36 hours of age to 30 days. From 30 days to two years of age they are fed Turkey Grower pellets (22% protein). At two years they are fed a 16% protein diet, except during breeding season when they are given a 36% protein diet. In order to encourage young cranes to imprint on one another, Archibald places a transparent screen between adjacent chicks in the brooder. At three weeks, cranes are placed in a 3 x 1.2 m enclosure, using heat lamps until the birds are fully feathered. Young cranes are highly susceptible to leg injury for the first 60 days of age. Archibald found that inter-chick aggression subsides at 40 days, and chicks can be placed together in outdoor enclosures containing soil, low grass, water, food, and a shelter. The birds must be carefully watched however, as fighting may still occur. Archibald stated that wings must be clipped after each moult. Usually the flight feathers of one wing are clipped. Breeding adults are held in breeding enclosures measuring 18.3 x 12.2 m, 18.3 x 18.3 m, or 24.4 x 18.3 m at the Foundation, depending upon the size of the species. Two inch (5 cm.) cyclone fencing is used, buried 30 cm in gravel. The enclosures are located away from ponds and marshes to minimize chances of disease. A small fountain and a shade tree are located in each enclosure, and grass is not mowed. Fences separating pairs are covered with vines or canvas to reduce contact and aggression between the pairs. Partitions of ~" plywood may also be used. Archibald found that breeding pairs must have privacy. Dried grass is placed in the breeding enclosure for nest construction. He determined that vocalizing cranes nearby act as a stimulus to breeding pairs. He state that personnel from the Patuxent Wildlife Reasearch Center in Maryland found that breeding pairs of Greater Sandhill Cranes are reproductively.sensitive to photo-period, and that extended daylight is beneficial to use from February until laying discontinues. A two-month-old"chick may be placed in an adjacent enclosure in visual contact with a breeding pair. Archibald found that once accepted, the chick and adults form a family structure that lasts until the chick is removed from the pair the following spring. He determined that the association with the chick increases the pair's reproductive drive. An attempt was made to raise Greater Sandhill Cranes in captivity at the Fish Springs National Wildlife Refuge in Utah (Kraft 1975). The plan was to rear cranes at the facility and reintroduce them to m~Fshes in the refuge area. The refuge personnel first attempted to incubate eggs but found that �46 too discouraging. They then obtained chicks from the Greys Lake National Wildlife Refuge in Idaho. The chicks were three to four days old and at this age pecking order fighting was a major problem. They found that birds seven to ten days old did not fight, and recommended that for future projects only birds five days or older be collected to avoid losses due to fighting. Kraft kept seven young birds in a garage and fenced backyard, and provided them with cardboard boxes for shelters. At 5 - 6 weeks they moved the birds to a one-acre enclosure, and provided a shelter, although it was not used. Running water is necessary in winter to prevent frozen toes. Kraft recommended that enclosures be placed away from tall vegetation where predators can hide. He found that cranes must have their wings clipped at least three times a year beginning in July. It is advisable to clip one wing to cause an unbalanced flight. He used a walk through drive trap to capture birds for wing-clipping. Kraft fed chicks pablum with water four times a day and crushed hard boiled eggs, including shells, once a day. At one week he introduced puppy chow, and gave eggs until the birds were four weeks of age. At eight weeks, the cranes were almost adult size, and were fed Purina dog chow. At twelve weeks grains (corn and millet) replaced dog chow. Kraft noted that the adult cranes ate mostly wild foods found in their enclosures, and only supplemented their diets with the grains. He found that cranes engage in territorial fighting in a pen, and because of this no more than four adults should be kept in a 6-Acre pen. Walkinshaw (1949) found ~hat captive crane chicks can be fed egg whites and dicalcium phosphate. At three weeks they will consume a variety of insects, earthworms, baby food, fruit, popped corn, meat, and cottage cheese. At one month crickets, katydids, .and grasshoppers were added to the diet. He described the consumption of approximately 150 - 200 insects every morning and evening. At three months cranes will eat corn ears in addition to their insect diet. As it may be necessary to trap wild cranes for the project, techniques of trapping were investigated. Wheeler and Lewis (1972) trapped cranes on the Platte River, and found that the use of "landing strips" of mowed and raked hay or alfalfa contributed greatly to their success. Also, taxidermy decoys were helpful in attracting cranes to the area. The nets used were 30 x 60 feet. They set 1 - 6 nets at each site and used three rockets. The nets were placed in parallel rows and directed at each other. The average catch was eight cranes per net firing. �47 METHODS Location of Potential Reintrodution Sites Fall Study: Initial work on verification of the presence of suitable crane nesting habitat in southwestern Colorado was undertaken by Caryn Fox. In order to establish guidelines for optimum nesting habitat, she studied p'resent Ly used nesting grounds in Routt Co. for 2 weeks, beginning 7 September 1978. She examined nesting sites described by Kathy Bieniasz (1977) in California Park, Slater Park, and around Hahns Peak. Photographs were taken in the California Park willows and streams where neasting activities occurred. In order to locate potential reintroduction sites, from 23 September 1978 through 20 October 1978 she made 2 aerial surveys (in Cessna-185) covering most of the historical Greater Sandhill Crane nesting range in southwestern Colorado, and then contacted U.S. Forest Service (USFS), Bureau of Land Management (BLM) , National Park Service (USPS), Colorado Division of Wildlife (CDOW) personnel to obtain site-specific information on dates of spring snow melt, availability of water, type and extent of land use, dates of recreational use, relationship to the path of migrating cranes, and availability of housing for captive cranes. With 4-wheel drive truck, and on foot, she visited 35 potential sites to determine area, height of willows, number and size of ponds, presence of beaver lodges, stream width, types of surroundings, presence of nearby drainages and marshes (for potential crane population expansion), proximity of roads, trails, buildings, and signs of grazing and recreational pressure, Photographs were taken at each site. Subsequently, each site was assigned a rating based upon habitat quality (from'guidelines established from prior study of Routt Co. Nesting grounds), human and animal impact, ownership of land (e.g., private, government), and usability. Spring Study From 16 April 1979 through 6 June 1979 I continued verification of the presence of suitable cranes nesting habitat. Due to requirements that the potential reintroduction sites be under currently used Greater Sandhill Crane spring migration routes and in historical nesting areas, serious consideration during this facet of the study was given only to those sites that are in the Gunnison Region. Initially, I concentrated my efforts on the 5 Gunnison Region Group I sites (for explanation of Group I and II status see RESULTS AND DISCUSSION). After consulting with District Wildlife Manager (DWM) Tom Henry, I added 2 Gunnison Region Group II sites (#17, Union Park, and #19, Texas Creek), and one additional site, Slaughterhouse Gulch, not previously examined. These 8 potential sites were evaluated for reintroduction through consideration of the following factors: 1) suitability snow accumulation and distribution along willow-lined drainages in comparison with similar habitats in the known nesting areas in California Park during the pre-nesting and early nesting period of Greater Sandhill Cranes (mid-April through May), �48 2) potential methods of access by CDOW personnel (for construction and maintenance of breeding pens and care for captive cranes) during use of the site, 3) availability, and distance from the site, of potentiallyavailable housing during the period the site would be in use, 4) land use restrictions and requirements affecting" the reintroduction and related activities, and 5) any other factors affecting reintroduction on activities remaining to be investigated. Four aerial surveys (in Cessna-185) were completed during pre-nesting and early nesting periods, primarily to evaluate snow accumulations. During the same period, attempts were made to reach the 8 sites in 4-wheel drive truck and on foot, primarily to evaluate site access by road. In early June 1 conducted field examinations of the 3 most promising sites to pin-point potentially-suitable habitat locations "for the erection of breeding pens. Personal contacts with USFS, BLM, Bureau of Indian Affairs (BIA), Soil Conservation Service (SCS), and CDOW personnel, as well as with private interests, were used to gain information relevant to all the above factors. Reintroduction Methodology Fall stl'dy Initial work on reintroduction logistics was undertaken by Caryn Fox. She also prepared the LITERATURE REVIEW. In addition to information gathered from this review, she received advice from personal communication with personnel representing the International Crane Foundation (rCF), the National Audubon Society (NAS), and the Denver Zoological Park. Spring study Finalization of reintroduction logistics was completed from mid-April through June of 1979. It was necessary to design and determine location of both holding and breeding pens, care for captives, methods for presenting captives to the reintroduction sites, and determining cost estimates. I relied principally on information from personal contacts with ICF and CDOW personnel, as well as, with commercial suppliers in the Fort Collins and Denver areas. Facilities at the Arapaho and Monte Vista National Wildlife Refuges (NWR) and the CDOW's Fort Collins Research Station (FCFS) were visited and personnel contacted to evaluate potential use for holding captive cranes. �49 RESULTS AND DISCUSSION Reintroduction Site Location Spring snow accumulation For the Steamboat Springs Region (currently-used crane nesting grounds), 15-year average 1 April snow depths on the 8500 ft (2590m) Hahns Peak SCS Snow Course and the 8650 ft (2640 m) Elk River #2 SCS Snow Course are, respectively 41 in (104 em) and 52 in (132 cm) , For the Gunnison Region (potential crane nesting grounds), 44-year 1 April average snow depth on the 9000 ft (2740m) Crested Butte SCS Snow Course is 50 in (127cm) • . In 1979, however, in the Steamboat Springs Region, 1 April snow depths were 53 in (135cm) and 66 in (168 cm), or 37% and 52% above the 15-year averages for the Hahns Peak and Elk River #2 SCS Snow Courses, respectively. In the Gunnison Region, snow depth for 1 April was 58.5 in (149 cm), or 17% above the 44-year average for the Crested Butte SCS Snow Course. If we assume that years of greater-than-average snow accumulation do not render the currently used Steamboat Springs nesting sites unsuitable for nesting Greater Sandhill Cranes, then the potential reintroduction sites in the Gunnison Region should not be considered unsuitable due to their frequent 1 April heavy snow accumulations. It is probable, however, that nesting phenology would be retarded during years of heavy spring snow accumulations and/or late snow melt. Therefore, in this reintroduction program, it may be necessary in some years to retard the presentation of captive cranes to the reintroduction sites from 2-4 weeks. Site analysis Fall study Caryn Fox located, and investigated on the grounds, 35 potential reintroduction sites, principally in the Gunnison, San Luis Valley, and North Park Reg:ions. Eleven potential sites were assigned "Group I " status: "excellent habitat with little or no human impact." Eight potential sites were assigned "Group II" status: "excellent habitat but may have heavy fishing pressure, cattle grazing, or heavy 4-wheel drive use'." For the "Group II" sites she determined that " •.. these areas, while not usable themselves, may be indicative of similar habitat in more remote and inaccessible locations." Sixteen potential sites were assigned "Group III" status: " •••unless they have one or two outstanding features, such as being in specially protected locations, they should not be considered for use as transplant sites." The following summary charts, Tables 1,2, and 3, prepared by Caryn Fox, list the 35 sites according to their "Group" designation and location. The charts contain information on the habitat, size, setting, snow melt, and human and livestock impact for each site, as well as proximity to the migration path and capacity for population expansion. Problems are listed for "Group II" and "Group III" sites to indicate reasons for their lower ratings. �TABLE 1: GROUP I NESTING AREAS GUNNISON REGION If 2 Title Castle Mtn. Wilderness Spring Ck. Res. 3 Brush Creek Willows 5mi x~mi 4 Slate River Willows 8mix~i 5 Blue Mesa Rd. Willows '-2mi x 100' 1mi x 200' 1 Habitat Type Willows, wet meadows Willows Size 2mi x'-2mi 3mi x 500' Setting Snow Melt Timber, mead, Late May, Mtn. park June Timber, sage Apr. - May Mtn. gulch Pasture, sage Apr. - May Rolling hills Timber, sage May Mtn. streams Sage April Rolling hills Impact Cattle, hikers July-Sept. Hikers, horses July-Sept. Cattle, Fishing .June.-Sept. Tourist, mines June-Sept. Cattle Migration Expansion Path Area Occasional Yes Occasional Yes Occasional Yes Occasional Yes Staging 5 mi. north Yes V1 0 6 7 8 MONTE VISTA REGION Monte Vista Nat'l Wildlife Refuge Blanca Wildlife Habitat Area Rio Grande Wildlife Area NORTH PARK REGION Arapaho Nat' 1 Wildlife Refuge 10 Moose Release Area 11 Southwestern North Park 9 Marsh ,wet meadows Marsh, wet meadows Wet mead,mar Willows 5000 Acres 470 Acres 2000 Acres Vli 11ows 10mi x 200' Willows 3mi x '-2mi Willows 2mi x 200' Meadow,sage Valley Sage Valley Grain.fields Valley Sage Valley Timber,sage Mtn. gulches Sage Rollin hills Feb. - Mar. Feb. - Mar. Feb. - Mar. April April May Hunting Oct. Fishing, hunt. July-Oct. None Migrant rest stop Roosting area Feeding rest stop Fishing, hunt. Aug. - Oct. Fishing, resid. July-SeEt. Cattle . June-Sept. Nesting 6 mi. west Nesting 20 mi west Nesting area Yes Some Yes Yes Yes Yes �TABLE: 2 GROUP II NESTING AREAS GUNNISON REGION ff Title Habitat Type 12 Lottis Creek Size 13 Ohio Creek Marshes, beaver ponds Willows 3mi x ~i 14 Rustler's Gulch Willows 3/4mi x ~i 1mi x 100' ~mi x ~i 15 Gothic Area Willows 2mi x ~i 16 Dustin Gulch Willows 2mi x "Gni 17 Union Park Willows 2mi x I;mi 18 Willow Creek Willows 1mi x ~mi 19 Texas Creek Willows smi x "Gni Setting Sage, Timber Mtn. Park Pasture, sage Valley Sage,timber Mtn. pa_rk_ Sage, timber Gulch Sage, timber Mtn. Park Sage, grasses Mtn. park Pasture,timb. Mtn. Gulch Sage, timber Mtn. gulch Snow Melt May April Apr. - May Apr. - May Apr. - May May May May Impact Fish ,hunt,cattle July-Oct. Hiking, cattle June-Oct. Camping ,fish July-Oct. Fishing,cattle July-Oct. Fishing, hiking Cattle, 4WD July-Oc_t.__ __ Cattle, resid. June-Oct. Fishing, 4WD June-Oct. Migration Path Yes Problems Uranium drilling Occasional Moly mine tailings pod Occasional Heavy ~ recreation Occasional Heavy recreation Stage Heavy 10 mi south recreation Stage Heavy in area grazing Stage Heavy in area recreation Stage Heavy in area recreation n_ �TABLE: 3 GROUP III NESTING AREAS GUNNISON REGION fI 22 Title Roaring Judy Fish Hatchery Route 149 Cebolla Game Area Tomichi Creek 23 Cebolla Creek Willows 2mi x ~i 24 Cimmarron Ck. Rd. Willows 5mi x 100' 25 Cochetopa Creek Willows 1mi x ~i 26 Quartz Creek Willows 2mi x 50' 27 Saguache Park Willows 2mi x ~i 28 Razor Creek Willows 3mi x 1/8mi 29 Willows 2mi x ~mi Willows 8mi x Willows Imi x ~i Willcrws 3mi x 100' 33 34 Ohio Pass, Kebler Pass, Coal Ck. Rd. Lake City Cutoff, Vulcan Cottonwood Pass road Rainbow Lake Rd. Dry Creek Soap Creek Rd. Cement Creek Willows Willows lmi x 20' 2mi x 1/8mi 35 MONTROSE REGION Hart's Basin Willows Imi x ~i 20 21 30 31 32 Habitat Type Willows Size Yzmix 50' Willows 2mi x 50' Willows 5mi x 50' Setting __ Snow_l1elt__. Itnpact Pasture April Fishing ,cattle Valley July-Oct. Sage Mar.- Apr. Cattle,recrea. Valley Apr.-Oct. Pasture,sage Mar.-Apr. Cattle Rolling hills April-Oct. Sage,timber April-May Fishing ,hiking Steep gulch Pasture Mar.-Apr. Fishing,cattle Valley _May-Oct. Pasture Mar.-Apr. Cattle Valley May-Oct. Pasture,sage April Cattle, resid. Valley April-Oct. Timber, mead, May Fishing ,cattle Mtn. park June-Oct. Pasture,sage April Cattle,resid. Valley April-Oct. Sage,timber May Mines,recrea. Ptn. park _J_\l_ne-Oct. Sage April Tourists,cattle Narrow gulch May-Oct. Timber May Cattle,recrea. Narrow gulch June-Oct. Sage Mar.-Apr. Cattle,recrea, Rolling hills May-Oct. Sage - gulch April Recrea,May-Oct. Timber, gulch April Recrea,May-Oct. Migration Path Occasional Yes Stage on creek Yes Stage in area Stage on creek Yes Yes Stage in area Occasional Reservoir Orchards Mar.-Apr. Recreation April-Oct. Heavy recreation Grazing Roads Too narrow Few willows Heavy V1 grazing N Few willows }1any cattle Few willows Many cattle Yes Moly mine Many People Too close to road Few willows Near road Many people Yes Yes Many people Many people Yes Many People u sot Problems Too small Occasional Yes �53 Spring study #1 Castle Creek Wilderness: Aerial observation on 26 April and 16 May 1979 revealed snow accumulation along the willows on Castle Creek to be approximately 50% greater than snow accumulation about the willow-lined drainages of California Park for the same time period. DWM Cliff Coghill claimed that snow frequently covers most of this site into July, potentially prohibiting access to both nestbuilding materials and a natural food source. Access to this site is provided by USFS Road 728, also called the Castle Mountain Wilderness Streams Road, a private road currently controlled by the Castle Mountain Irrigation Road and Recreation Association. This road runs: west from the Ohio Creek Road (USFS Road 780) to the eastern edge of both the West Elk Wilderness Area and the Potential reintroduction site. There appears to be little difficulty, currently, with our use of this road, although the USFS is contemplating litigation to gain greater control over its use, and with the current RARE II proposals for extending the "Wilderness Area" designation further east, motorized travel eventually may be prohibited. As of 26 April 1979, a snow bank created by Gunnison County snow plowing operations on the Ohio Creek Road prevented access to the private road. As of 14 May 1979, I was able to drive a 4-wheel drive truck about 1 mi (1.6 km) from the Ohio Creek Road before snow accumulation prevented further driving, with over 5 mi (8 km) of road remaining to the potential reintroduction site. It appears that access plans would require the use of snow machine, snow shoes, or skis to the wilderness boundary, and snow shoes and/or skis within the Wilderness Area. Landing of a helicopter within the West Elk Wilderness Area is prohibited. The Castle Creek Cow Camp, operated under USFS permit by the Ochs Brothers, would be potentially available for housing CDOW personnel during most of the time period in question. It is located approximately 0.5 mi (0.8 km) east of the Wilderness Area boundary on the Castle Mountain Wilderness Streams Road. A CDOW-owned 9-ft (2.7-m) camper placed just east of the Wilderness Area boundary before winter conditions inhibit access on the private road would provide a viable alternative to use of the Castle Creek Cow Camp. Although David Ruff (Land and Recreation Manager, Taylor Ranger District, Gunnison National Forest) assured me of the USFS's desire to aid us in the reintroduction activities, it is unlikely that the CDOW could obtain permission to construct any breeding pens that would both be compatible with reintroduction needs (relatively permanent and predator proof) and not in violation of Wilderness Area statutes. Cattle are authorized on this site annually, beginning 16 June + 14 days (pees comm, John Curran, District Ranger, Taylor Ranger District, Gunnison National Forest). Human impact at this site is negligible prior to the normal crane hatching of young. �54 Although this area possesses excellent Greater Sandhill Crane nesting habitat and relatively light animal and human activity, the duration of snow cover into early summer and the conflict between our activities and wilderness area statutes give this site low potential as a reintroduction site. #2 Spring Creek Reservoir: Aerial observation on 26 April and 16 May 1979 revealed snow accumulation along the willows on Spring Creek and Flag Creek to be approximately 50% greater than snow accumulation about the willow-lined drainages of California Park for the same time period, potentially prohibiting access to both nest-building materials and a natural food source. Access to this site is provided by USFS Road 744, running north about 15 mi (24 km) along Spring Creek from the Taylor Canyon Road (USFS Road 742). As of 26 April 1979, USFS Road 744 was plowed clear for vehicular traffic approximately 2.5 mi (4.0 km) from USFS Road 742, leaving almost 13 mi (21 km) of impassable road to the potential reintroduction site. On 14 May 1979, I was able to drive approximately 3.4 mi (5.4 km) on USFS Road 744 from USFS Road 742, still almost 12 mi (19 km) short of the potential reintroduction site. Although this site could be reached by snow machine over those portions inaccessible to 4-wheel drive truck, DWM Tom Henry held the opinion that avalanche danger along Spring Creek during pre-nesting and early nesting periods might inhibit safe access to the site. Approach to this site by helicopter might be the only safe, feasible method during this period during years of heavy snow accumulation. Although the Boston Cattle Camp is located only about 4 mi (6 km) south of this potential reintroduction site, the permittee refused to make this facility available to CDOW'reintroduction personnel. The nearest available housing is located at 2 resorts, approximately 2 mi (3 km) north of the Taylor Canyon Road along Spring Creek (and approximately 13 mi (21 km) from the site). Their posted rates for cabins are about $135/wk. Since we would be using housing at least partly during the "off-season", they would substantially reduce their rates (pers comm., resort owner-mangers) for CDOW personnel needs. A CDOW-owned 9-ft (2.7-m) camper placed near this potential reintroduction site before winter conditions inhibit access along USFS Road 744 would provide a viable alternative to use of the resort housing. David Ruff assured me of the USFS's wish to expedite our reintroduction activities at this site. Upon our submission of a "letter of intent" (outlining our reasons for choice of this particular site, the time period in question, etc) and our submission of the "Special Use Application and Report" (Reference FSM 2712), the USFS would be ready to enter into a cooperative transplant agreement with the CDOW for reintroduction activities at this site. Cattle are authorized on this site annually, beginning 1 June + 14 days (pers comm., John Curran). Human impact at this site is negligible prior to the normal crane hatching of young. �55 Although this area possesses excellent Greater Sandhill Crane nesting habitat and relatively light human and animal impact, the duration of snow cover well into the nesting period gives this site low potential as a reintroduction area. #3 Brush Creek: Aerial observation on 26 April and 16 May 1979 revealed snow accumulation along the willows on Brush Creek to be approximately 25% less than snow accumulation about the willow-lined drainages of California Park for the same time period. During the 16 May 1979 flight, several large (50-ft (15-m) diameter) openings were observed along the lower West Branch and East Branch of Brush Creek, potentially allowing access to both nestbuilding materials and a natural food source. Access to this site is provided by USFS Road 738 running northeast about 7 mi (11 km) along the East River and Brush Creek to the site from CO 135. As of both 26 April and 14 May 1979, USFS Road 738 had been plowed for approximately 3.5 mi (5.6 km), the remaining approximately 3.5 mi (5.6 km) to the potential reintroduction site having too much snow for passage of a 4-wheel drive truck. This last portion of road to the site should be easily passable in a snow machine. Steve Rieshl may have cabins available for CDOW personnel, located on lower Brush Creek in SE ~, S-28, T.13 S., R.85 W., along the western edge of the potential site. A CDOW-owned 9-ft (2.7-m) camper placed near this potential reintroduction site before winter conditions inhibit access along USFS Road 738 would provide a viable alternative to use of the privately-owned cabins. David Ruff assured me of the USFS's wish to expedite activities at this site. the reintroduction Cattle are authorized on this site annually, beginning 6 July + 14 days. The potential for increased cross-country activity and a planned summer camp at the potential reintroduction site may conflict with the crane reintroduction requirements. It is lik~ly that some use of snow machines for the transport of CDOW personnel and equipment during the pre-nesting period will be necessary. This area possesses excellent Greater Sandhill Crane nesting habitat, currently light spring-time human and animal impact, and less spring-time snow accumulation than similar habitats in California Park. However, this site is given only moderate potential as a reintroduction site because of the probable increased human activity. #4 Slate River: Aerial observation on 26 April and 16 May 1979 revealed snow accumulation along the willows on the Slate River to be approximately 10% greater than snow accumulation about the willow-lined drainages of California Park for the same time period, potentially prohibiting use by cranes. �56 Access to this site is provided by USFS Road 734, running northwest about 5 mi (8 km) along the Slate River to the site from CO 135. As of 26 April 1979, USFS Road 734 was plowed clear for approximately 2.5 mi (4.0 km) from CO 135; and, as of 14 May 1979, this road had been plowed beyond the potential reintroduction site, but was still not readily passable by 4-wheel drive truck. This last portion of road should be passable in a snow machine. There is no available housing closer to this site than what is connnerciallyavailable in Crested Butte. A CDOW-owned 9-ft (2.7-m) camper placed near this potential reintroduction site before winter conditions inhibit access along USFS Road 734 would provide a viable alternative. David Ruff assured me of the USFS's wish to expedite the reintroduction activities at this site. Cattle are authorized on this site annually, beginning 1 June ± 14 days. This site receives heavy use from skiers, hikers, and fishermen. In addition, there is a high probability of increased mining activities in the vicinity of this site. Thus, although this area possesses favorable Greater Sandhill Crane nesting habitat, it appears that recreational and mining activities will conflict with the crane reintroduction. In addition, the duration of snow cover well into the nesting period gives this site low potential as a reintroduction site. #5 Blue Mesa Road: Aerial observation on 26 April 1979 and ground observation on 15 May 1979 revealed virtually no snow remaining about the willo~.,salong Willow and Pine Creeks. Access to this site is provided by the Blue Mesa Road, running south about 6 mi (10 km) along Pine Creek Mesa between Pine and Willow Creeks to the site from US 50, west of Lake Fork Campground on Blue Mesa Reservoir. As of 26 April 1979, the Blue Mesa Road was in the process of being plowed to clear the few drifts remaining between US 50 and the~ite, all snow having disappeared from the area by 15 May 1979. It is possible that a snow machine would be necessary for access during the pre-nesting and early nesting periods in years of unusually late and/or heavy snow accumulation. The only housing for CDOW personnel near the site is owned by the Ute Mountain Ute Tribe, who have not responded to my inquiries regarding use of their facilities. The Sapinero Trading Post, located about 10 mi (16 km) from the site at Sapinero on US 50 does have primitive cabins available for about $150/mo. A CDOW-owned camper placed near the potential reintroduction site would provide a viable alternative to use of either the Ute or Sapinero facilities for housing. Phil Bailey (Realty Specialist, BLM, Montrose) have assured me of the BLM's readiness to enter into a cooperative transplant agreement with the CDOW for reintroduction activities on those portions of the site on BLM property. �57 They would be willing to take care of any required environmental assessments. Ernest House (Brunot Area Agreement Hunting Commission and Councilman, Ute Mountain Ute Tribe) has granted permission to the CDOW for reintroduction activities on those portions of the site on Ute tribal lands. It is likely that portions of the site will change ownership between the BLM and the Ute Mountain Ute Tribe, the areas to be affected still in question. At present, most of the suitable habitat is on portions controlled by the Utes. Cattle are authorized on the BLM lands, beginning between 10 and 20 May. Only limited areas along Willow and Pine Creeks possess suitable nesting habitat, the expanses of willows along the drainages are quite limited in area and length. Consequently, the potential for growth of the reintroduced population might be severely limited. In addition, homesites are being developed along the south bank of Blue Mesa Reservoir and eventually may extend to close proximity with the few suitable nesting habitats. While snow accumulation appears to be minimal during spring migration, the shortcomings of limited habitat and uncertainty of future land ownership and use give this site only moderate potential as a reintroduction site. #17 Union Park: Aerial observation on 16 May 1979 revealed snow accumulation in the willow-lined marsh in Union Park to cover approximately 50% of the site. Access to the site is provided by USFS Road 752, running south about 4.0 ffii(6.4 km) from Taylor Park, or by USFS Road 758, running east about 3.5 mi (5.6 km) from the Lottis Creek Campground on the Taylor Canyon Road. As of 14 May 1979, neither of these roads was passable by 4-wheel drive truck. Snow machine or helicopter should provide access to this potential reintroduction site with little difficulty. The closest available housing for reintroduction personnel is at the Taylor Park Trading Post on the southeast edge of Taylor Park Reservoir. Their rates are $12/day. A CDOW-owned camper placed near this potential reintroduction site before winter conditions prevent access along USFS Road 752 (or 758) would provide a viable alternative to use of the commercially-availab~e housing at Taylor Park. David Ruff assured me of the USFS's wish to expedite activities at this site. the reintroduction Cattle are authorized on this site annually beginning 1 July 14 + days. Interspersed with the USFS property in Union Park are numerous small private land holdings which receive high recreational use. Also, the University of Minnesota operates a summer geology camp inUnion Park. A dump for uranium tailings is being proposed (pers comm, David Ruff). Although this area possesses excellent Greater Sandhill Crane nesting habitat, it appears that recreational and potential mining activities may conflict with CDOW reintroduction activities. A snow machine would be essential for access, at least during the pre-nesting and early nesting periods. Consequently, this site is given low potential as a reintroduction site. �58 #19 Texas Creek: Aerial observation on 16 May 1979 revealed snow accumulation in the willows along Texas Creek to cover approximately 10% of the site. Access to this site is provided by the Texas Creek Road, a 4-wheel drive road running east about 2.5 mi (4.0 km) along Texas Creek from the northeast edge of Taylor Park Reservoir. As of 14 May 1979, USFS Road 742 was not passable by 4-wheel drive truck from approximately 0.25 mi (0.4 km) south of the west end of the Texas Creek Road, and the Texas Creek Road was impassable. A snow machine should provide access to this potential reintroduction site with little difficulty. The nearest available housing for CDaW reintroduction personnel is at the Taylor Park Trading Post. A CDaW-owned camper placed near this potential reintroduction site would. provide a viable alternative to the commercially-available housing at Taylor Park. David Ruff assured me of the USFS's wish to expedite activities at this site. the reintroduction Cattle are authorized on this site biennially beginning 11 July + 14 days. While it is likely that use of snow machine during the pre-nesting period will be necessary, the area possesses excellent Greater Sandhill Crane nesting habitat and receives relatively little human and livestock use before mid-July. This site is given high potential as a reintroduction site. Slaughterhouse Aerial the willows Gulch: observation on 16 May 1979 revealed snow accumulation about along Slaughterhouse Gulch to cover approximately 40% of the site. Access to the site is provided first by heading south on USFS 765 from Tin Cup for about 0.5 (0.8 km), then heading west about 1 (1.6 km) along Slaughterhouse Creek. As of 14 May 1979, .USFS Road was plowed as far as Tin Cup (it had been all winter), but passage beyond Tin Cup in 4-wheel drive truck was not possible due to snow accumulation. A snow machine would be needed to provide access to Road mi 765 this site. The nearest available housing for CDaW reintroduction personnel is at the Taylor Park Trading Post. A camper placed near this potential reintroduction site before winter conditions prevent access along the roads from Tin Cup would provide a viable alternative to the commercially-available housing at Taylor Park. David Ruff assured me of the USFS's wish to expedite activities at this site. the reintroduction Cattle are authorized on this site annually, beginning 1 July days, reintroduced population may be limited by the lack of nearby. ± 14 �Table 4. Summary of 8 Potential Reintroduction Sites Site Spring Snow Accumulation 1) Accessibility2) Nearest Housing Land Use ReStrictions Human &/or Potential for Animal Impact3) Reintroduction 111 Castle Creek Wilderness 50% greater snow machine, snowshoes, &/or skis probably required Castle Creek Cow Camp, 0.5 mi (0.8 km) east of site no motorized travel, no permanent structures negligible low 112 Spring Creek Reservoir 50% greater snow machine (avalanche danger) &/or helicopter probably required Char-B and Spring Creek Resorts, $135 /wk, 13 mi (21 km) from site negligible negligible low 113 Brush Creek 25% less snow machine probably required cabins adjacent to site, or Crested Butte negligible moderate light recreational use now, will probably become heavy 114 Slate River 10% greater snow machine probably required Crested Butte negligible low heavy recreational use, potential mining 115 Blue Mesa Road much less, virtually no snow cover snow machine may be required during years of unusually late melt Sapinero negligible on BLM lands, must file reports to Utes if on Ute lands. light 1117 Union Park much less snow machine required Taylor Park negligible low heavy recreational use, potential mining 1119 Texas Creek much less snow machine may be re(luired Taylor Park neglibible somewhat less snow machine probably req.uired Taylor Park negligible Slaughterhouse Gulch I I moderate4) light high light moderateS) \JI \.0 �60 Table 4 (cont.). Summary of 8 Potential Reintroduction Sites. Notes: 1) 2) 3) 4) 5) in comparison with known nesting sites in northwest Colorado during April and May during April, May, and June this site has limited suitable nesting habitat this site lacks other suitable nesting habitat in vicinity for potential population expansion �61 The streams at higher elevations are often snow covered in April, and those at lower elevations receive high recreational pressure. It is likely that snow machines will be required for access during the pre-nesting and early nesting periods. This area possesses excellent Greater Sandhill Crane nesting habitat, little use before mid-July, and lesser snow accumulation than similar habitat in California Park. The limitations imposed by its small area and recreational use in adjacent habitats give this site only moderate potential as a reintroduction site. Reintroduction sites with moderate-high potential The Brush Creek (#3), Blue Mesa (#5), and Slaughterhouse Creek sites possess characteristics giving them moderate potential for Greater Sandhill Crane reintroductions. During the next few years, the Brush Creek site .Ls likely to experience human impacts that will conflict with the needs of nesting cranes. The Blue Mesa site has only limited suitable habitat, and potential intrusion of housing developments during the next few years is also likely to conflict withthe needs of nesting cranes. Slaughterhouse Creek in itself appears quite suitable, but future natural expansion of any reintroduced nesting population would probably be severely limited by the lack of other nearby suitable habitats. Considering the factors for reintroduction site suitability (see METHODS), the Texas Creek (1119) site holds the greatest potential for use in establishing a new nesting population of Greater Sandhill Cranes in Colorado. Captives While capturing male cranes during their third fall (when just over 2 years old) and placing them in breeding pens their third spring might appear to simplify the inherent difficulties in caring for captives and eliminate the first 2 years of the reintroduction, problems frequently encountered with caring for wild adults in captivity, their concomitant failure in achieving readiness for breeding, and the lack of opportunity for their obtaining philopatric ties to the reintroduction site (during their second spring) make the capture of 5-to lO-week old chicks a more feasible alternative (per comm. , George Archibald). The implication that care of cranes from chick to breeding adults will simplify matters, however, is not realistic. Their maintenance will necessitate, at least initially, long periods of observation and care. Housing and feeding requirements must be met (see Holding pens for captive cranes and Feed for captives). Approximately 15 pre-flight chicks should be captured on the nesting grounds (e.g. Grays Lake. Idaho) by running down 5- to lO-week old chicks. After leg banding, the chicks will be transported in cardboard boxes by truck to the nearest airfield, by airplane to Fort Collins, Colorado, and by truck to the FCRS. Blood samples will be taken from each banded chick and sent to the ICF for individual sex determination. �62 In the northwest part of the FCRS there are 21 pens potentially available for housing these cranes, including 20 small enclosures, measuring 9.0 ft (2.7 m) by 12.0 ft (3.7 m) by 6.5 ft (2.0 m) high, and a single large enclosure, measuring 72.0 ft (21.9 m) by 30.0 ft (9.1 m) by 6.5 ft (2.0 m) high. Initially, the young cranes will be housed together in the large enclosure. Ten plywood shelters, measuring 4.0 ft (1.2 m) 8 ft (2.4 m) by 4 ft (1.2 m) high will be constructed to provide protection from the elements. Water will be provided from 10 galvanized steel feed pans (placed over poultry heater bases to prevent icing in winter). Purina Game Bird Flight Conditioner will be provided, ad libitum, from an additional 10 galvanized steel feed pans until the chicks are 16 weeks old. During these first 16 weeks the young cranes will be carefully observed, those individuals displaying overt aggressive behavior or those lacking sufficient assertion to obtain feed will be isolated in the smaller pens. It is possible that those cranes failing to feed may initially have to be force fed by CDOW personnel (per comm., George Archibald). From the age of 16 weeks the juvenile cranes will be fed Purina Game Bird Maintenance Chow, ad libitum. In early April of their second year,S male captive cranes will be selected from the FCRS for use at the reintroduction site. At this point, the fencing between 10 of the smaller pens adjacent to the large pen at the FCRS will be dismantled, creating a second large pen measuring 45.0 ft (13.7 m) by 30.0 ft (9.1 m) by 6.5 (2.0 m) high. A visual barrier of 54-in (137-cm) canvass fabric will be erected between the two large pens. Those captives remaining at the FCRS will be separated into the 2 large pens, by sex, to prevent pair-bond formation amongst the captives. Observation of the captives must be continued and isolation of overtly aggressive individuals to the ten remaining small pens to the north may be necessary. Purina Game Bird Maintenance Chow will continue to be provided, ad libitum. During the third fall, with the return of the 5 captives from the reintroduction site to the FCRS, segregation by sex will be discontinued (pair bonds unlikely to form during the fall and winter) until the first days of the following March. During the remaining years of the study, a schedule of observation and care for at least 2 hours per day, isolation of overtly aggressive individuals, the provision of Purina Game Bird Maintenance Chow, ad libitum, and separation of the sexes during the nesting season will be followed. Since the potential for mortality amongst captives is always present, it is probable that some of the males not previously used at the reintroduction site may now have to be recruited as replacements. If mortality amongst captives reaches a degree such that 5 males are no longer available at the FCRS at the 'commencement of the nesting season, female captives may have to be recruited for use at the reintroduction sites. Hopefully, they would be successful in attracting wild, migrating males. Although there are more records in the crane literature of captive males attracting and nesting with wild females than of captive females attracting and nesting with wild males, Hyde (in Konrad 1975) does report a captive female attracting and nesting with a wild male crane. At the onset of the final nesting season of this reintroducion project, any captives remaining at the FCRS may be used for other studies, contributed to zoological parks, or released at the spring migration stop at the Monte Vista NWR. �63 Food for Captive Cranes There are several complex diets described in the literature for the feeding of captive cranes (Table 5). It is clear that nutritional needs vary with crane maturity. As an alternative to preparing our own diets, the Ralston Purina Company (Cherkerboard Square, St. Louis, MO) offers a variety of prepared feed mixes readily available locally from Northern Colorado Feeders Supply, Inc. (359 Linden St., Ft. Collins, CO). In the interests of availability, reliability, and simplicity, their use is recommended. Purina Game Bird Flight Conditioner is recommended for cranes under 16 weeks old. Purina Game Bird Maintenance Chow is recommended afterwards, except that Purina Game Bird Breeder Layena should be fed during the breeding season to those cranes designated for breeding activities. Quantities and costs of feed for this project are estimated in Table 6. �64 Table 5. Compilation of Recommended Immature Cranes Taken From 36 hrs. to 1 month: Archibald 100g turkey starter crmbls. (1974) 2sg Magnesium sulfate lsg Zinc carbonate 250mg Folic acid sOmg Biotin 300mg Choline CI. sOOmg Pyridoxine 109 Niacin 50g Vitamin Bl2 2000g Corn meal 1 month to 2 years: turkey grower pellets (22% protein) 36 hrs. to 1 week: Pablum & water Crushed hd. bId. egg. (l/bird inc. shells) 1 week: Moistened Friskies puppy chow Phase out Pablum 4 weeks: Discontinue h.b. eggs 6 weeks: Dry puppy chow 8 weeks: Purina dog chow 12 weeks: Corn & millet Schmitt 36 hrs. to 6 weeks: (pers. comm.) Game bird chow Dicalcium phosphate Raw greens Kraft (1975) 1) Diets for Captive-Reared Mature Cranes 2 years +: Non-breeding: Turkey breeder pellets (16% protein) Breeding (Feb-end Laying) 36kg/907kg turkey str. Pre-Mix 519kg Soybean meal 9kg Animal fat 36kg Pex poultry Pre-Mix 138kg Meat & bone meal 91kg 17%alfalfa meal 36kg Pical phosphate 11 kg Salt 68kg Limestone (=36% rotein) U weeks +: Wild foods Corn & millet 6 weeks +: !z;Duck chow 3/4 Turkey chow Trout chow Raw greens Crickets Mealworms Cranes. Cost 1) 16% protein =$4.75/s0Ibs. Time 20%prot.= 5.75/50Ibs. Corn: $5.9s/100Ibs. Barley: $3.35/80Ibs. Feed Pablum with water 4 x per day. 1 egg lx/day Approx. $50.00- Feed once $IOO/bird per yr. per day. Costs established from Hayden Grain Co., 198 E. Lincoln, Hayden Colo. Cost of Schmitt diet from Ed Schmitt, Denver Zoo. �65 Table 6. Year Food for Captive Cranes. Units Item Total Cost 1 5 Purina Game Bird Flight Conditioner, 50 lb bag, @$6.80 $ 34.00 35 Purina Game Bird Maintenance Chow, 50 lb bag, @$5.95· $208.25 2 48 Purina Game Bird Maintenance Chow, 50 lb bag, @$5.95 $285.60 3-10 40 Purina Game Bird Maintenance Chow, 50 lb bag, @$5.95 $238.00 8 Purina Game Bird Breeder Layena, 50 lb bag, @$6.90 Annual total costs: 1 242.25 2 285.60 3-10 293.20 $2,873.45 $ 55.20 �66 Holding Pens for Captive Cranes The various techniques to be considered in the reintroducion strategy require facilities for maintaining captive Greater Sandhill Cranes. The captives should be fed and observed on a daily basis. Although visits to the Arapaho and Monte Vista NWR's and conversations with their managers established the potential feasibility for holding captive cranes on these refuges, it was clear that neither currently possessed adequate holding facilities. In addition, both NWR's experience harsh winter, "temperatures having dropped to -430C twice this past winter at the Arapaho NWR. The CDOW Fort Collins Research Station (FCRS) currently has 21 suitable holding pens available. Heated poultry bases will be needed to prevent icing of water for drinking, and 8 ft (2.4 m) by 4 ft (1.2 m) by 4 ft (1.2 m) high shelters will have to be constructed for use during the w~nter months. Canvass fabric maybe needed for visual isolation between pens (see Table 7). �Table 7. Materials for Holding Pens (FCRS). Supplier Units Item Total Cost Northern Colorado Feeders Supply, Inc. (359 Linden, Ft. Collins, CO). 20 matal feed pans, @$2.25 $ 45.00 10 poultry heater bases, @$17.50 $175.00 Jax Surplus (1200 N. College, Ft. Collins, CO). 40 54 in width canvass, 1 yd, @$4.00 $160.00 0"> -....I Sutherland Lumber (1901 E. Prospect, Ft. Collins, CO). 30 1 4 ft X 8 ft X ~ in plywood, @$8.39 $251.70 nails, 1 lb box, @$0.55 $ .55 '$632.25 �68 Breeding Pens Breeding pen design is adapted largely from "Standard Permanent Fence" of Gunnison Area Supervi.sor Jim Houston. The 84 ft (25.6 m) by 60 ft (18.3 m) pen standing 8 ft (2.4 m) above the ground surface will have braced wood post corners with steel posts to support fencing material between the corners. Three strands of barbed wire, projecting outward 1 ft (0.3 m) in the horizontal plane, 9 in (23 cm) below the top, and an apron projecting 30 in (76 cm) outward in the horizontal plane 4 in (10 cm) below the ground surface will provide additional protection against predators. For simplicity, there are no gates, ingress and egress provided by ladders. These pens will be maintained annually to provide for their 7-yr use. Their location, to include suitable nesting habitat, necessitates their construction in late summer when stream flow at the reintroduction sites should be at a low and before winter conditions interfere (see Table 8). Construction of Breeding Pens Layout: Mark out 84 ft by 60 ft rectangle. Locate post positions every 12 ft. Corner posts: "Cup" 12 of the sturdier wood posts; center cups at 9 in and 4 ft 9 ins. from narrow end of post. On 4 of these posts repeat "cupping turn from the first 2 cups. Connect the remaining 16 posts (braces) to theses 12 posts with ring shank nails to make 4 complete corners. With post hole digger, dig 3-ft holes for these corners. Set corners in place and pour concrete. lt ~ T-posts: Drive T-posts 1 ft 8 in into ground with T-post driver. Side fencing fabric: Dig earth 4 in deep for 30 in outward from pen perimeter. After concrete has set (may take 3 days), staple welded wire fabric to wood posts every 2 ft of post length and connect to T-posts with fence clips every 2 ft of post length. Connect the 2 widths of welded wire fabric to each other with hog rings every 9 in. Top predator proofing Cut 1 in by 2 in by 8 ft lumber into 6-16 in lengths. Nail these 16-in lengths (pointing out from pen) on wood posts at 1 ft 9 in from top, and attach with bailing wire to T-posts at 1 ft from top. Staple 3 strands of barbed wire at 4-in intervals to bottoms of the 16-in lengths. Bottom predator proofing: Cut 30-in apron from 60-in welded wire fabric. Connect with hog rings every 9 in to bottom of 60-in welded wire fabric sides and bury with earth removed earlier. �Table 8. Materials for Breeding Pens. Supplier Units Item Westridge Fence (2000 N. CO. RD. 23, Bellvue, CO) 28 12 Ft (6 in diam) treated wood posts, @$7.83 $ 219.24 Stockyard Lumber 6990 Hwy 85, Denver, CO) 12 10 ft. steel T-posts, @$4.79 $ Total Cost 57.48 Jax Surplus 5 1 in X 2 in welded wire fabric, 60 in width, 100 ft length, @$82.50 $ 412.50 Jax Surplus 6 1 in X 2 in welded wire fabric, 48 in width, 50 ft length. @$22.75 $ 136.50 Wheelers (1000 N. Hwy 287, Ft. Collins, CO) 8 hog rings, 100/box, @$0.85 $ Wheelers 1 staples, 10 Ib box, @$4.55 $ 4.55 Wheelers 1 fence clips, 50/pkg, @$0.49 $ .49 Westridge Fence 4 8-in ring shank nails, 1 Ib box, @$0.60 $ 2.40 Westridge Fence 12 Reddimix concrete, 2/3 cu ft, @$2.66 $ 31.92 6.80 Q\ \0 Wheelers 1 barbed wire, ~ mi reel, @$27.69 $ 27.69 Sutherland Lumber 4 1 in X 2 in X 8 ft lumber, @$0.35 $ 1.40 Wheelers 1 bailing wire, 6500 ft roll, @$29.50 $ 29.50 Jax Surplus 2 14-ft aluminum extension ladder, @$27.50 $ 55.00 Total for 1 pen $ 986.02 Total for 5 pens $ 4495.961) �70 Table 8 (cont.). Materials for Breeding Pens. Tools 2) 1 2 2 1 1 1 1 50 ft. tape measure shovels fencing pliars post-hole digger T-post driver hatchet hand saw Notes: 1) Materials needed for 5 pens are 5 times those needed for 1 pen, except for the following items whose total units for 5 pens are: 22 29 1 4 1 2) These tools should be available units 60 in width welded wire fabric units 48 in width welded wire fabric unit staples units barbed wire unit bailing wire from the cnow Gunnison Area Office. �71 Reintroduction Strategy The role of territorial behavior, especially in the male, has been noted for Greater Sandhill Cranes during breeding activities (Drewien 1973). Its development may be encouraged by restricting captive 2-year old males to a reintroduction site encompassing suitable nesting habitat during the entire nesting season. Since knowledge of migration routes to and from the nesting site is necessary for migratory behavior, it would be expedient to provide for at least one of the pair-bonded cranes, in this case the female since the male will be restricted to the habitat, to have this information. If the captive male is located under the migration path and could attract a wild female during her spring migration from the wintering area, between them, the pair would have both the needed sense of philopatry towards,and knowledge of migration routes to and from, the reintroduction site. With the strong drive in the female to return to the area of successful pairbond formation (and hopefully nesting), and the strong drive of the potential offspring to return to the area of their birth (its fulfillment made possible by incorporating information on the new migration route learned from the female during their first fall and spring migration), a natural sustained population increase at the reintroduction site is predicted. Longley (1970) reported one wing-clipped male crane attracting a mate in migration. Over 5 nesting seasons, this pair remained bonded and produced 5 young, the female leaving her mate each fall and returning the following spring. Five cranes, later observed flying over the pair, may have been their progeny returning to the area. Konrad (1975) reported 2 instances of captive Greater Sandhill Cranes forming pair bonds with migrants, at least one pair ralslng young. In Japan, Takahashi ~ Konrad 1975), captured 5 wild male Manchurian Cranes (G. japonensis), 3 of them successful in attracting and mating with wild females. Since Sandhill Cranes are believed to pair for life (Walkinshaw 1949), it would be expedient to use males with no prior pair-bonding. Also, as adult wild cranes may take many years to be suitable for this reintroduction project (see Captives), it appears most expedient to capture juveniles for use in this project. If obtained from the wild at over 4 weeks old, problems of aggression and feeding should be reduced (Kraft 1975). To restrict the males to the reintroduction site, and yet allow for their calling in migratory females, open-topped breeding pens must be used. Some method for restraining the males from flight, at least until a strong sense of philopatry has developed, must be used. Kraft (1975) suggests that one wing (i.e., the primaries) be clipped, causing unbalanced spiral flight. An alternate, and perhaps simpler, method would be to tie the wing (s) with elastoplast, a sticky plastic tape (pers comm., George Archibald). (The following strategy for reintroduction of Greater Sandhill Cranes in southwestern Colorado is adapted from the methodology of Konrad (1975). During the annual northerly migration (about 1 April) of wild Rocky Mountain Greater Sandhill Cranes from their traditional spring migration �72 stop at the Monte Vista NWR, 5 captive male Greater Sandhill Cranes in their second year will be individually restricted to open-topped, predatorproof breeding pens located under the migration path on sites which include suitable nesting habitat. If these sites are still totally snow covered, the introduction of these captives to the breeding pens may have to be delayed. Each of these 5 captive males will have one wing secured with elastoplast. Daily inspection of the breeding pens should be conducted to maintain their ability to restrict the captives and exclude predators, Purina Game Bird Maintenance Chow will be provided, ad libitum. During this first summer in the breeding pens, these young males will have an opportunity to develop philopatric ties to the site, and, although capable of forming pair bonds, will probably be too immature for actual nesting. During the period of fall migration (around 1 October), these 2-year old captives will have their wings freed and be returned to their winter holding facilities at the FCRS. During the following year's spring migration, these same nearly 3-year old captive males will one again have their wings secured with elastoplast and be returned to the same individual breeding pens that they occupied the previous year. Daily pen inspection, as performed the previous year, will again be maintained and Purina Game Bird Layena will now be provided, ad libitum. These birds should now have strong philopatric ties to their respective breeding pens and should be both physiologically and behaviorally ready for nesting activities. With the use of the unison call, " ••.an elaborate visual and vocal display," (Walkinshaw 1949), the captives should attract migrating, unpaired wild females, resulting in pair-bond formation and subsequent nesting activity. . As the fall migration reaches a peak (about 1 October), the wild females should lead their young in migration from the reintroduciton site, and, by example, the young should learn the route. After their departure, the males will have their wings freed and be returned to their winter holding facilities. During each of the following years, the same procedure will be followed, using the identical captives, if possible. Additions from the pool of captive cranes maintained at the FCRS may prove necessary should mortality in the 5 cranes used at the reintroduction sites occur. It is predicted that the same wild females will return and mate each year withfuese captives, and their young are also predicted to summer, and nest upon achieving maturity, in the vicinity of the reintroduction site. The breeding pens will be used during 8 nesting season/then dismantled and removed. At the onset of the ninth nesting season, the 5 cranes will be taken from the FCRS and placed at the reintroduction site, but with the absence of breeding pens, will no longer have their wings tied. They will be free to migrate with their mates and young. For disposal of remaining captives see "Captives." Table 9 illustrates the reintroduction schedule and Table 10 the projected costs. �Table 9. Year 1 Reintroduction Schedule. 1 July-30 September 1 October-31 December 1 January-31 March 1 April-30 June 1 July-14 July:prepare holding pens at FCRS 15 July-7 August: capture chicks 15 July-30 September: feed Flight Conditioner 15 July-30 September: daily intense observation and care of captives 1 October~2 hr care & observation/da of captives 1 October: inspect prospective reintroduction sites for worthiness 15 October~feed Maintenance Chow 1 January: order all materials for breeding pens 15 April: reinspect prospective sites for worthiness -...J 2 1 August-15 September: construct breeding pens 1 April: segregate captives at FCRS by sex )1 April, when breeding pens at least partially snow free, introduce captives to site, initiate daily inspection of breeding pens 1 October: place camper convenient to breeding pens 15 March: segregate captives at FCRS by sex 3-9 1 October: return captives from reintroduction site to FCRS; terminate isolation by sex 10 As in Year 3, then dis- 15 March: release mantle and remove breed- those captives not ing pens used at reintroduction site or donate As in Year 2, but initiate feeding of Breeding Layena at breeding pens As in Years 3-9, but wings will no longer be secured W �74 Table 10. Reintroduction Costs Year 1980-81 1980-81 1980-81 1980-81 1981-82 1981-82 1981-82 1981-82 1982-83 1982-83 1982-83 1982-83 1982-83 1983-84 1983-84 1983-84 1983-84 1985-on Item Cost Researcher A 4x4 Pickup Travel Operating Total: $19,200.00 7,000.00 500.00 2,000.00 $28,700.00 Total: $22,080.00 250.00 1,500.00 10,000.00 $33,830.00 Total: $22,800.00 7,000.00 2,500.00 10,000.00 2,000.00 $44,300.00 Total: $24,000.00 2,600.00 5,100.00 42000.00 $35,700.00 Researcher B Capitol Outlay Travel Operating Researcher B Camp Trailer Travel Operating Part-time help Researcher C Travel Operating Part-time help Similar to 1983-84 with inflation costs extra. �75 LITERATURE CITED Archibald, C.W. 1974. Methods for breeding and rearing cranes in captivity. Int. Zoo. Yearbook 14:148-156. Bailey, A.M. and R.J. Niedrach. 1965. Birds of Colorado. Den. Mus. Nat. His. Vol. I. 454 pp. Bieniasz, K.A. 1978. Biology of Greater Sandhill Cranes in Routt County, Colo. M.S. Thesis. Univ. of Northern Colo. Greeley, Colo. 71 pp. Blake, D.J. 1974. A preliminary study of the Greater Sandhill Crane in Colorado. Unpubl. report, Colo. Div. of Wildlife. 15 pp. __________ 1975. A study of Colorado Greater Sandhill Cranes on their dancing grounds and nesting areas. Unpubl. report, Colo. Div. of Wildlife. 19 pp. Carter. 1876. in Bailey, A.M. and R.J. Niedrach. 1965. Birds of Colorado. Den. Mus. Nat. His. Vol I. 454 pp. Drew. 1881. in Bailey, A.M. and R.J. Niedrach. 1965. Birds of Colorado. Den. Mus. Nat. His. Vol. I. 454 pp. Drewein, R.C. 1973. Ecology of Rocky Mountain Greater Sandhill Cranes. Diss. Univ. of Idaho. 82 pp. & 70 pp. Konrad, P.M. 1975. Potential for the reintroduction of cranes into areas of former habitation. Proc. Int. Crane Workshop. 1:317-325. Kraft. R.H. 1975. Conclusion of five-year study to reintroduce the Greater Sandhill Crane at Fish Springs National Wildlife Refuge, Dugway, Utah. Littlefield, C.D. and R.A. Ryder. 1968. Breeding biology of the Greater Sandhill Crane on the Malheur National Wildlife Refuge, Oregon. Trans. N. Amer. Wildlife Nat. Res. Conf. 33:444-454. Longley, W.H. 1970. Sandhill Cranes at the Carlos Avery Wildlife Area, The Loon. 4:124-128. Peters. 1925. in Brent, A.C. 1926. Life histories of North American Marsh Birds. U.S. Nat. Mus. Bull. 135. 490 pp. Szymczak, M. 1970. Canada Goose restoration along the foothills of Colorado. Tech. Publ. No. 31, Colo. Div. of Wildl. 30pp. Taylor, W.E. 1975. Sandhill Crane habitat management on the Hiawatha Nat'l Forest. Proc. Int. Crane Workshop. 1:44-50. Walkinshaw, L.H. 1949. The Sandhill Cranes. Cranbrook Inst. Sci. Bull. 29. Bloomfield Hills, Mich. 202pp. Warren. 1904. in Bailey, A.M_~ and R.J. Niedrach. 1965. Birds of Colorado. Den. Mus. Nat. His. Vol. 1. 454 pp. Wheeler, R.H. and J.C. Lewis. 1972. Trapping techniques for Sandhill Crane studies in the Platte River Valley. U.S. Fish & Wildl. Svc. Resour. Publ. 107, Wash. D.C. 19pp. Prepared by: Approved by: Howard Geduldig \-lildlifeTechnician 1-B aI~~#~ Walter D. Graul Research Leader �76 JOB PROGRESS State of Project REPORT COLORADO --~~~~~---------------No. Endangered SE-3-3 Work Plan No. II Endangered Job Title Development Prairie Period Covered: Personnel: Birds of a Preservation Job No. Program Wildlife Investigations 6 for Three Species of Grouse. 1 July 1978 - 30 June 1979. W. Busby, R. Calderon, J. Cancalosi, J. Garcia, W.D. Graul} R. Kahn, C. Loeffler, G.C. Miller, J. Torres, T. Washington, B.D. Will. ABSTRACT The status and key habitats of greater pra~r~e chickens (Tympanuchus cupido pinnatus), lesser prairie chickens (T. pallidicinctus), and sharp-tailed grouse (Pediocetes phasianellus jamesi) were investigated. All had undergone range reductions accompanied by changes in land use. There was no reason to believe that changes will not continue to encroach upon the 3 species' occupied ranges. A greater prairie chicken census using line transect sampling techniques showed promise but gave an estimate with a very large confidence interval (327 to 2993 prairie chickens, p < 0.05) • A greater prairie chicken habitat restoration plan prescribed interseeding, sand sage (Artemisia filifolia) control; and seeding of depression areas on approximately 1820 ha of the South Platte Management Area. Information still needed to preserve prairie grouse in Colorado :includes: (1)ways and means of providing prime habitat on public lands and (2) types of private land use practices compatible with prairie grouse welfare. �77 DEVELOPMENT OF A PRESERVATION PROGRAM FOR THREE SPECIES OF PRAIRIE GROUSE Gary C. Miller , P.N. OBJECTIVES This program is designed to identify management options to stabilize population levels of greater prairie chickens, lesser prairie chickens, and sharp-tailed grouse. It consists of grouse surveys to document present distribution and abundance of these species, vegetation surveys to describe existing and potential areas of grouse habitat, and contacts with landowners to ascertain possibilities for habitat acquisition and/or land-use modifications directed toward preservation of Colorado's prairie • grouse. SEGMENT OBJECTIVES 1. Determine the present population levels and distributions of all three species of prairie grouse in Colorado: Greater prairie chickens, lesser prairie chickens, and sharp-tailed grouse. 2. Identify key habitat use areas - booming grounds, winter concentrations, nesting areas for all three species of prairie grouse in Colorado. 3. Determine the potential for the restoration chicken population on the Colorado Division South Platte Management Area. of a greater prairie of Wildlife's (CDW) INTRODUCTION Three species of grouse (Order: Galliformes, Family: Tetraonidae) once ranged over most of Colorado's eastern plains. Each has undergone severe range reduction since the early to mid-1900's. In each case, this range reduction has accompanied changes in land use (Evans 1964, Rogers 1969,'Hoffman 1973). The Colorado Wildlife Connnission has classified greater prairie chickens and P. p. jamesi as endangered species, and lesser prairie chickens as threatened species in Colorado. CDW employees have periodically conducted lek surveys for all three species. This report presents preliminary results of work to update previous information. a STUDY AREAS Greater prairie chickens were studied primarily in Yuma and Phillips counties in northeastern Colorado. Parts of Kit Carson, Logan, Morgan, and Washington counties were visited occasionally. Intensive studies and �78 took place on 798 km2 (308 mi2) in the sandhills north of Wray, Yuma County. Greater prairie chicken habitat restoration plans were designed for the South Platte Management Area, Logan County. The sharp-tailed grouse study area consisted of 1460 km2 (560 mi2) in Douglas and Elbert counties. The area was characterized by dissected foothills and buttes in the western part, and rolling foothills and plains in the eastern portion. Lesser pra1r1e chickens were studied in Baca and Prowers counties in southeastern Colorado. Areas in Bent, Kiowa, and Otero counties were evaluated as potential habitat. METHODS AND MATERIALS Temporary CDW employees Cancalosi, Kahn, and Will chickens, sharp-tailed grouse, and lesser prairie Field work was performed from January until early and key habitat areas were delineated by personal with local residents. Key habitat characteristics line transect sampling methods. studied greater pra1r1e chickens, respectively. May. Distribution observations and interviews were ascertained by Several census methods were attempted and evaluated. Winter flock counts, roadside surveys (including rural mail carrier surveys), baiting station counts (Amman 1957), and lek counts (Schwarz '1944) were conducted. Cancalosi compared the methods of Hayne (1949) and Emlen (1971, 1978) for line transect census of greater prairie chickens. Temporary employee Busby developed the habitat restoration plan. RESULTS Greater Prairie Chicken The present occupied range of greater pra1r1e chickens is compared with their past distribution in Figure 1. Primary range comprised about 430 sections (111,370 ha) in the sandhills of Yuma and Washington counties. There were unconfirmed reports from Kit Carson and Phillips counties. Recollections of informants for the period 1969-1978 indicated periodic prairie chicken presence in Sedgwick and Logan counties. Virtually all prairie chickens seen loafing or roosting during the study were in moderately to lightly grazed pastures within 1.6 km of a crop field. One ranch north of Wray, believed to contain the greatest concentration of greater prairie chickens in Colorado, grazed calves between May and September 1978 at 8 acres per calf to maintain pasture and 10 acres per calf to improve pasture. Roughly one-half of the intensive study area was under a light to moderate grazing regime apparently suitable for greater prairie chicken loafing, roosting, and nesting (Table 1). Yuma County experienced an II-fold increase in irrigated acreage between 1959 and 1974. Conversion of prairie to irrigated crops has continued; about 100 well permits had been awarded in the prairie chicken range of Yuma County since 1977. �LEGEND -. .0 •••• . Q Presumed historic western and southern limits (Aldrich and Duvall 1955) COLORADO Distribution in 1962-1963 (Evans 1964). • to(;A •., &C"'IC;< ,.." Distribution in 1979. * Habitat restoration area. Ii i'l'AJ"''''''(}N 1"0 d,,A,,,c() . ·• •1•• ••· c••ell • A""lU) rU"'S~H "HeOtH ~IJA 'N! rt • ""(}t{T~C.U ~ l''Jl4u) S"''''_~''II. • ........___ U(1HLcr I'I?C":/{S Of "' [)O(O/(IS M()",rLZIJ..M.A lAC'" ~A'HV'I Fig. 1. '''' AS Past and present distribution of greater prairie chickens in Colorado, and habitat restoration area. �80 Table 1. Land use characteristics sharp-tailed Land Use of greater prairie chicken and prairie grouse occupied ranges in Colorado, Percentage Greater Prairie Chicken 1/ 1979. of Occupied Range Prairie Sharp-tailed Grouse 2/ Agricultural 25 ± 9 17 Pasture 74 ± 5 73 Other 1/ Results from line transect sampling ~/ Results from planimetry of USDA Soil Convervation Use Map, Douglas County, Colorado, 1973. 10 Service Land �81 Winter census provided little information about greater prairie chicken numbers. Rural mail carrier sightings averaged fewer than 1 per 12,950 km (5,000 mi) of driving. Lek surveys were hampered by inclement weather. Cancalosi drove a 22.4 km (14ffii) transect on 23· and 25 April, but was unable to distinguish possible repeat detection with confidence. The strip census estimates for greater prairie chickens and, for comparison purposes, black-tailed jackrabbits (Lepus americanus) using the Hayne (1949) and Emlen (1978) methods are summarized in Table 2. The point estimates from the two methods showed closer agreement for jackrabbits than for prairie chickens. The estimate for prairie chickens was hampered by sample size (23 birds in 16 observations). Applying the Hayne estimate to the area considered primary greater prairie chicken range gave an estimate of between 327 and 2993 prairie chickens(p< 0.05). Sharp-tailed Grouse Figure 2 compares the present occupied range of P.p. jamesi with their past distribution. In addition to about 14,570 ha (36,000 acres) in Douglas County, sharp-tailed grouse occupied a small area in Elbert County. A small population may have been in the foothills of Boulder, Gilpin, and/or Jefferson counties, but their occurrence was not verified. Land use in the Douglas County sharp-tailed grouse range is summarized in Table 1. Some areas may be threatened by encroachment of housing developments. In Elbert County, the grouse primarily used 3 49 ha (120 acre) ungrazed pastures. Vegetation sampling of areas used by the grouse was not conducted. Roadside counts and baiting station counts provided little indication of sharp-tailed grouse abundance. Ten active leks were located, 4 of which had not been previously identified. Peak numbers of grouse at leks ranged from 2 to 13. There appeared to be no reason to alter previous population estimates of 100-200 P.p. jamesi in Colorado. Lesser Prairie Chicken Figure 3 compares the present occupied range of lesser prairie chickens with their past distribution. Lesser prairie chickens occupied about 120 sections (31,080 ha) in Baca County and about 10 sections (2590 ha) in Prowers County. Most of the occupied range in Baca County was administered by the U.S. Department of Agriculture, Forest Service, Comanche National Grasslands. Occupied range in Prowers County was privately owned by several parties. In general, occupied range was characterized by sand sage grassland communities. No land use survey was conducted in these areas, but center pivot irrigated grain fields occurred in the Prowers County range. Vegetation characteristics of the occupied ranges and potential habitats are shown in Figure 4. There was little difference in the heights of bunchgrasses among all areas, but more variation in the height of shrubs. Shrub densities, however, varied little among areas, and mean percentages of bare ground varied about 7.5% between high and low values. (Fig. 5). �82 Table 2. prairie Comparison of Hayne (1949) and Emlen (1978) estimates of greater chicken and black-tailed jackrabbit densities, Yuma County, Colorado, 1979. Density Estimate (per mi2) Hayne method Emlen method Greater Prairie Chicken 3.86+3.10 All observations Possible Black-tailed resightings excluded 5.93 15.81±14.05 16.43 15.28±12.76 15.84 Jackrabbit All observations Possible 2.34+2.19 resightings excluded �-LEGEND - • ----., Presumed historic western and southern limit (Aldrich and Duvall 1955). COLORADO wno It Distribution in 1979. lQC"." J:&:''''''I( • 1' •. ' ~ ~ ,,"; g(t:AI'I I i'YA;"I":'/~/"I 'IotA'!'A ""0 ~'A""'O ""l~ coeca:« kif U.~S':N co W ® c«: " ,,",Ol'l''*.c.st .TIO ",,(4l" J""w'~ JAt<I _,,,II. I"'"'" OCJ,g~fS I"""" tAr -",,''''AS L / ;Olm, lJ(II, • Fig. 2. Past and present distribution of sharp-tailed grouse, P.p. jamesi in Colorado. • �._. LEGEND Presumed historic western and northern limits (Aldrich and Duvall 1955, Hoffman 1963). COL 0 R ADO W'LD Distribution in 1979. lOGA'" ".1 Areas evaluated as potential habitat. * " I'I'•• SIf/IIi"O'" tu,ooA ~"I.I.J) "HeoLH "'-lJA 00 ~ J""WCMI JAil ...",,11. I' IlC~ l fi.; QD(O MO"'TI tllMA ~",r ,vI'MAS Ail'",,, I ''''' Fig. 3. Past and present distribution of lesser prairie chickens in Colorado. �Fig. 4. Comparison of vegetation density and height, lesser prairie chicken study areas, 1979 Bunchgrass r==J Shrubs co V1 Baca Co. Prowers Co. OCCUPIED HABITATS Kiowa Co. B.ent Co. UNOCCUPIED HABITATS WIDTH OF COLUMNS INDICATES RELATIVE DENSITIES (AVERAGE) Otero Co. �86 Fig. 5. Comparison of percentage bare ground, Lesser prairie chicken study areas, 1979. 71 -,- 70 -- 69 -- 68 -f- ,-.. ~ t1l Q) ;:;:: 6'1 - '-" "0 § 0 .~ d 66 _'- Cll ~ cd j:Q .p s:: Cll 65 - - U ~ .Q) p... 64 - ~ 63 -- 62 - f- o Baca Prowers Co. Co. OCCUPIED HABITATS Kiowa Bent Otero Co. Co. Co. urr~CUPIED HABITATS �87 A total of 175 prairie chickens were counted on 19 leks during the study. This is 19 more birds, and 3 more leks, than counted the previous year. Table 3 summarizes lek census data since 1960. These data indicate that lesser prairie chickens have not decreased in abundance the past several years. GREATER PRAIRIE CHICKEN HABITAT RESTORATION PLAN About 1820 ha (4500 acres) of CDW's South Platte Management Area may be restored to support a greater prairie chicken population (Fig. 1). The tract, which once supported the species, apparently had been heavily grazed prior to CDW acquisition in the late 1940's and early 1950's. It was grazed at varying intensities until 1978, and no grazing took place in 1978-79. Instrumental in the plan's development were the late Stu Adams, William Busby, Richard Rhoades, Warren Snyder, Harvey Sprockley, and Clinton Wasser. The tract was divided into three units based on soils and vegetation (Fig. 6). Unit A contained Dailey and Valent soils, vegetated with prairie sandreed (Calamovilfa longifolia), needle and thread (Stipa comata), sand bluestem (Andropogon hallii), blue grama (Bouteloua gracilis), sand dropseed (Sporobolus cryptandrus), and sand sage. Unit B contained Dailey loamy sands on 3-9% slopes, somewhat susceptible to wind erosion. Needle and thread, blue grama, and annuals were dominant plants. Unit C soils were Valent loamy sand on 3-15% slopes, extremely susceptible to wind erosion. Sand sage dominated the area, with varying amounts of grass understory. Unit A No immediate treatment was prescribed for which should produce dense stands of tall Interseeding may be needed in small areas, seeded (Appendix I,ll). Sand sage should part of section 19 (Appendix III). the hills and side slopes, and mid-grasses with time. and depressions should be be controlled in the southern Unit B Interseeding-was prescribed for much of this unit. Test seeding plots should first be monitored to ascertain optimum mixtures. Trial seedings should include: prairie sandreed, little bluestem, sand lovegrass (Eragrostis trichodes), yellow sweet clover, varieties of wheatgrass (Agropyron spp.), and switchgrass (Panicurn virgatum),_and the mixture shown in Appendix II. Depressions in the unit shou ld be treated as shown in Appendix I. Unit C Sand sage control was prescribed for parts of sections 15, 20, 21, and 22 (Appendix III). Interseeding was recommended for the depressions in the unit. �8fl, Table 3. Results of lesser prairie Year chicken lek surveys in Colorado, Number of Grounds Total Birds (High Counts) 1960 6 39 1961 11 84 1962 11 116 1963 10 125 1964 No Data It\.vailable 1965 No Data ~vailable 1966 No Data ~vailable 1967 1 1968 No Data ~vailable 1969 No Data ~vailable 6 1970 .3 42 1971 3 37 1972 7 82 1973 7 65 1974 11 107 1975 13 151 1976 15 158 1977 17 178 1978 16 156 1979 19' 175 . , 1960-1979. �v~. ./ t ,., 9' .. " "" Fig. 6. Greater prairie chicken habitat Letters denote management units referred restoration tract, South Platte Management to in text.• Area, Logan County, Colorado. �90 Other Recommendations 1. Rye (Secale cereale), wheat (Triticum aestivum) or triticale may be planted in 30.5 m (100ft) wide strips at right angles to prevailing winds. Use mini-till methods, replanting each September with one-sweep tillage. 2. Monitor vegetation at potential lek sites, especially with regard to sand sage. 3. Ascertain feasibility of repairing windmills for trickle irrigation of food plots. 4. Test brood-rearing habitat plantings of alfalfa (Medicago sativa), clover (Trifolium spp.), croton (Croton texensis) , and lead plant (Amorpha sp.). 5, Test plant rejuvenation methods that m1n1m1ze soil erosion. Methods may include: rototilling narrow strips, breaking ground with interseeder, mowing, burning small patches, or grazing. 6. Test snowfencing to retain moisture and enhance tall grasses. 7. Seed blowouts according to Appendix IV. 8. Evaluate feasibility of erecting fencing to prevent low-flying birds from crossing Interstate Highway 76. Do not allow power lines to cross the tract. DISCUSSION There is no reason to believe land use changes will not continue to encroach upon occupied ranges of prairie grouse in Colorado. The situanion seems especially tenuous for greater prairie chickens and prairie sharptailed grouse, where most of their present range is in private ownership. There is an immediate need for two types of information: (1) ways and means of providing prime prairie grouse habitat on public lands and (2) types of private land use practices that are compatible with prairie grouse welfare, including range management and agricultural practices. In addition, further modification of the strip census technique must be made to provide population estimates with smaller confidence intervals. �91 LITERATURE CITED Aldrich, J.W., and A.J. Duvall. 1955. Distribution of American game birds. U. S. Fish and \.Jildl.Serv. Circ. 34. 23 pp , Amman, G.A. 1957. The pra1r1e Tech. Bull. 200 pp. grouse of Michigan. Emlen, J.T. 1971. Population densities counts. Auk 88: 323-341. 1978. Estimating breeding Auk 94: 455-468. Michigan gallinaceous Dept. Conserve of birds derived from transect season bird densities from transect counts. Evans, K.E. 1964. Habitat evaluation of the greater pra1r1e chicken in Colorado. M.S. Thesis, Colorado State Univ., Ft. Collins. 98 pp. Hayne, D.W. 1949. An examination of the strip census method for estimating animal populations. J. \.Jildl.Manage. 13: 145-157. Hoffman, D.M. 1963. The lesser prairie Manage. 27: 726-732. chicken in Colorado. J. Wildl. 1973. Changes in populations and habitats of lesser prairie chickens in Colorado, 1963 to 1973. Proc. of Prairie Grouse Tech. Council 10: 10-11. Rogers, G.E. 1969. The sharp-tailed grouse in Colorado. Fish, and Parks Div. Tech. Publ. 23. 94 pp. Schwarz, C.W. 1944. Comm. 179 pp. The prairie chicken Prepared by: in Missouri. Colorado Missouri Go.xy C_.r'1~ Gary C. killer Nongame Researcher Game, Conserve �92 APPENDIX I Seeding Reconunendations - Depression Are.as Seedbed Preparation - Early spring (March) - work the soil once or twice to ~ill existing vegetation. Seedbed may need to be packed to firm the seedbed. Seedbed should be prepared at angles perpendicular to the wind direction (from the northwest). Seedbed should also be prepared in strips no wider than 100 feet, leaving strips of vegetation in between those to be seeded. After seeded strips are established, the alternate strips can be prepared and seeded. Fertilizer - Fertilizer requirements can be determined by laboratory analysis or, in lieu of this, 40 pounds of nitrogen and 40 pounds of phosphorus per acre can be applied. Seeding - Areas should be seeded in late March or early April. Grass drills should be used for the seeding. Grass drills are available from the South Platte Soil Conservation Service in Fleming and the Morgan SCS in Fort Morgan. Species Amount of Pure Live Seed Per Acre to be Seeded Sand bluestem - Woodward SWitchgrass - Nebraska 28 Yellow Indian grass - Holt or Llano Yellow sweet clover Consider adding to mixture: Alfalfa - Rambler or Rhizoma Sainfoin Hairy vetch Burnet Twice the normal seeding rate was used since additional available and a dense stand of plants is preferable. 2.5 1.6 3.5 1.0 moisture will be Mulch - Areas will need to be mulched to protect the soil and encourage growth of seedlings. Mulch should be composed of prairie hay or cereal grain straw. 50% of the hay or straw should be 10 inches or more in length. The mulch should be applied after seeding and crimped into the soil with a crimper or a disc plow with its discs set straight. The rate of application should be 4,000 pounds per acre. �93 APPENDIX II Interseeding The area should be seeded in late March or early April. Amount of Pure Live Seed Per Acre to be Seeded Species .6 Little bluestem - Pastura sand bluestem - Woodward switchgrass - Nebraska 28 prairie sandreed 1.3 .7 .6 Two thirds of the normal seeding rate was used. This is standard procedure for interseeding, because you are seeding into an established stand of plants. No preparation is needed for interseeding on sparse density or weedy sandy sites. For sites with dense sand sage, elimination of some sage is necessary. This can be by a) spraying in June or b) stubble mulch disc plowing and planting sudangrass or sorghum in 12-18" rows. Planting would preferably be in June to reduce any seedset which might compete with the grass. This will produce a stubble into which the grass mixture can be drilled in late August with surplus soil moisture or usually with greater safety early the following April. APPENDIX III Sand Sage Control Control of sand sage is recommended by the Soil Conservation Service when it forms a canopy cover in excess of 15% as measured by transect. Spraying - The herbicide 2,4-D is recommended for sand sage control under our conditions. It does not completely control sage. The first application may only kill ~ to 2/3 of the plants. A second application the third year may be necessary. Total eradication of the sage would be undesirable as the sagebrush does serve to catch snow, thereby improving moisture conditions for other plants, and it provides shade and cover for prairie chickens. One and one half pounds acid equivalent of 2,4-D has been recommended. Dilution and wetting agent rates are variable but 1.5 pounds 2,4-D combined with one gallon diesel and brought up to five gallons with water per acre if applied by plane should work well. For ground spraying 25 gallons of water should insure adequate coverage. Timing is important. For effective control twig growth must be at a maximum. This usually occurs from May 15 to June 20, often June 1-10. Soil moisture should be high to insure continued growth. Mowing - Mow in two successive years. Time of year would be the same as above. Due to the rough terrain, mowing would be unsuitable in many areas. See pamphlets on sage control included with proposal. �94 APPENDIX Revegetation IV of Blowouts Seedbed Preparation - Existing vegetation should be killed. Any steep slopes, ledges or cutbanks should be smoothed and shaped so that no slope is greater than 3:1, preferably 6:1 or flatter. Fertilization - Use laboratory soil analysis or 40 pounds per acre nitrogen and 40 pounds per acre phosphorus. Apply prior to seeding. Seeding - Seed in late March or early April. Use grass drill. Amount Pure Live Seed Per Acre to be Seeded Species Sand bluestem - Woodward Little bluestem - Pastura Prairie sandreed Switchgrass - Nebraska 28 Yellow sweetclover Mulch - See description under Appendix 3.2 1. 75 1.6 .9 .7 I - Seeding. �95 JOB FINAL REPORT State of COLORADO --~~~~~------------------ Project No. -=S~E~-~3_-~2 Work Plan No. II Endangered Job Title Acquisition Sharp-tailed Period Covered: Personnel: _ Birds of Habitat Endangered Wildlife Investigation 8 for the Preservation of the Prairie Grouse in Colorado 1 July 1978 to 30 June 1979 John Torres, Walter Graul, Bernard Goetze, Donald Colorado Division of Wildlife Schoonover ABSTRACT An effort was made to acquire 1,000 acres of habitat in Douglas County for the preservation of the pralrle sharp-tailed grouse. The land was not acquired due to the position of the landowner involved. �96 ACQUISITION OF HABITAT OF THE PRAIRIE FOR THE PRESERVATION SHARP-TAILED GROUSE IN COLORADO WALTER D. GRAUL P.N. OBJECTIVES 1. To preserve and perpetuate habitat sufficient 300 prairie sharp-tailed grouse in Colorado. 2. To design and implement a managment plan to optimize grouse habitat on 1,000 acres of land. to maintain at least sharp-tailed SEGMENT OBJECTIVES 1. Acquire 1,000 acres of prairie County, Colorado. sharp-tailed grouse habitat in Douglas INTRODUCTION Historically, the prairie sharp-tailed grouse (Pedioecetes phasianellus jamesi) resided in Colorado from the foothills in Douglas County, northeastward to the Nebraska border. As the native pra1r1e community was subjected to intensive agriculture arid intensive grazing by livestock, the native grouse populations declined steadily. As a result of the preceding land practices, this subspecies in Colorado is now limited to about 250-350 birds in Douglas County. Unfortunately, the remaining Douglas County habitat is now being lost rapidly as the result of the construction of intensive sub-developments. If the prairie sharp-tailed grouse is to be preserved in Colorado, it will require the preservation of habitat. Since the existing habitat is located on private lands in Douglas County such a preservation effort involves either acquiring or leasing these lands. One acquisition possibility developed when a Douglas County landowner, interested in the preservation of the sharp-tail, expressed a sincere interest in selling 1,000 acres of his land to the Colorado Division of Wildlife. This project was designed to complete this purchase. The acquisition was especially appealing in that the landowner was only wanting $10. 00 per acre for the land; the rest of the value being offered as a donation. METHODS Negotiations were conducted the project period. AND MATERIALS with the landowner periodically throughout �97 RESULTS AND DISCUSSION Despite lengthly negotiations, an agreement between the landowner and the Division of Wildlife could not be reached regarding the sale of the 1,000 acres. As time progressed, the landowner developed more and more restrictions as to the future use and operation of the land by the Division. These restrictions were such that the land could not be managed fully for the benefit of the prairie sharp-tailed grouse. Since at the end of the project period prospects for the completion of the acquisition were not good, the decision was made not to pursue the aquisition in 1979-80. If the acquisition prospects do change, a new project will be developed. Prepared by Walter D. Graul Nongame Research Leader �98 JOB PROGRESS State of REPORT COLORADO --------~~~~-------------- Project No. Work Plan No. Job Title Job No. 1 Population Period Covered: Personnel: Nongame FW-22-R Investigations ~3~ Surveys of Selected Bird and Mammal _ Species in Colorado 1 July 1978 through 30 June 1979 W. Graul, S. Bissell, C. Loeffler, C. Chase III, M. Moulton, J. Freeman, C. Fox, S. Vallejos, C. Reck1ing - Colorado Division of Wildlife. Dr. Ronald A. Ryder - Colorado State University. ABSTRACT From 1 July 1978 through 30 June 1979 intensive field studies were conducted on the following species or species groups: White-faced Ibis, Great Blue Heron, Snowy Egret, Bats, Eastern Prairie Mammals. Data were also collected on the following additional species: Black-Crowned Night Heron, Double-crested Cormorant, White Pelican, Least Tern, Snowy Plover, Mountain Plover, California Gull, Greater Sandhill Crane, and Black-necked Stilt. Based on the preceding work, tentative distributional data were compiled for the mentioned species. For a few of the species more detailed data, such as productivity and habitat associations, were also collected. �99 POPULATION SURVEYS OF SELECTED BIRD AND MAMMAL SPECIES IN COLORADO Walter D. Graul P.N. OBJECTIVES 1. Determine statewide distributions, population levels, and population trends for select bird and mammal species - those with relatively restricted habitat requirements. 2. For the same species, determine species-habitat associations. SEGMENT OBJECTIVES 1. Determine statewide distributions for select bird and mammal species. INTRODUCTION Historically, This approach For instance, classified as wildlife management has been orientated toward single species. is not feasible for the broad spectrum of nongame species. there are 347 species of birds and 73 species of mammals nongame in Colorado. An alternative approach, termed a species-ecosystem approach, has been proposed for management of all nongame species in Colorado (Graul, Torres, Denney, 1976). In fact, this approach now constitutes the foundation of the Division of Wildlife's management program for those nongame species not classified as threatened or endangered (Hess, 1977). The method basically consists of determining: (1) what species in a community have the most restricted habitat requirements, and (2) what are the habitat requirements of these restricted species. The latter habitat requirements can then be used to develop management guidelines for broad habitat types that will inSure the preservation of a wide spectrum of species - not just a few. Based upon preliminary data collected by the Division's nongame program (Kingery and Graul, 1978; Bissell, 1978), it is possible to determine what bird and mammal species in Colorado should receive top priority in terms of applying the species-ecosystem approach. Namely, these are the species that in terms of distribution in Colorado are relatively restricted while not being peripheral. Furthermore, those species whose distributions are unknown need to be investigated immediately. The species addressed in this study were chosen according to the preceding criteria. METHODS AND MATERIALS All the intensive field studies were conducted by temporary procedures of each major segment will be described herein. employees. The �100 The eastern pra1r1e mammal survey work was conducted by Michael Moulton between July, 1978 and June, 1979. Small mammals were trapped in four lowland and six upland prairie sites - a total of 6,249 trap/nights was completed (one trap set for one night equals one trap/night), utilizing both snap traps and live traps. Specimens were prepared as standard study skins and skulls and deposited in the Division of Wildlife vertebrate collection. The White-faced Ibis (Plegadis chihi) study was conducted by Stan Vallejos from May-August, 1978, with Federal Aid covering the work during July and August. This work was restricted to the San Luis Valley, since this is the only area where the ibis nests regularly in the state. In conjunction with the ibis study, nesting data were also obtained on the Snowy Egret (Egretta thula) and the Black-crowned Night Heron (Nycticorax nycticorax), since these species nest in mixed colonies with the ibis. All nests of the preceding species were monitored regularly at Trites Lake to determine productivity. In January, 1979 a colonial waterbird mail survey was initiated by Caryn Fox to locate the major waterbird colonies in the state. The survey form was sent to all Colorado Division of Wildlife field personnel, u.S. Fish and Wildlife Service refuge personnel, u.S. Forest Service and u.S. Bureau of Land Management area office personnel. Additionally, the survey was sent to all state parks, all chapters of the Audubon Society in Colorado, and those ornithologists associated with colleges and universities in the state. Previous mail surveys conducted by Dr. Ronald A. Ryder in 1965 and 1973 were also used to obtain preliminary data on waterbird colonies in the state. Using the preceding survey data as a starting point, field work was conducted by Stan Vallejos and Candy Reckling during the summer of 1979. The emphasis was on visiting all Great Blue Heron (Ardea herodias) heronries identified in the previous surveys. Additionally, potential colony sites were visited that were not identified in the earlier surveys. In conjunction with the Great Blue Heron field work, data were collected on other waterbird colonies, but a special effort to locate the latter colonies was not made. A statewide field study of select bat species was initiated by Jerry Freeman in the summer of 1978 and continued through the summer of 1979. Emphasis was placed upon documenting the distribution of all species and trying to verify the occurrence of possible new species. In addition to the preceding intensive field surveys, limited data were collected on other select species. Charles Chase III documented nesting of Snowy Plovers (Charadrius alexandrinus) in 1978 and 1979, Least Terns (Sterna albifrons) in 1978, and Black-necked Stilts (Himantopus mexicanus) in 1979. In July, 1978, 100 juvenile White Pelicans (Pelecanus erythrorhynchos) were banded at Riverside Reservoir in Weld County. In conjunction with other work, two nesting colonies of the California Gull (Larus californicus) were monitored. Also, plans for monitoring Colorado's Greater Sandhill Crane (Grus canadensis tabida) nesting population were made in conjunction with Routt Forest personnel (U.S. Forest Service). Finally, new information on the breeding range of the Mountain Plover (Charadrius montanus) was obtained via short-term field work by Walter Graul and Steven Bissell. �101 RESULTS AND DISCUSSION Eastern Prairie Mammal Survey Segment Site Description All sites located at Bonny Reservoir, A. B. Lowland Yuma County, unless otherwise noted. Sites: 1. Panicu~Distichlis - this site is essentially a tall grass site. Panicum virgatun and Distichlis stricta are two typical species of perennial grasses in this community. 2. Wild Hay - this site is also essentially a tall grass community, except this site is annually mowed. Typical grasses in this site include: Andropogon gerardii, Sporabolus cryptandrus and Agropyron smithii. 3. Grazed Riparian Woodland. 4. Ungrazed Riparian Woodland. Upland Sites: 1. Short grass communi ties - these include Buchloe dactyloides, Bouteloua gracilis, and Aristida~. as typical species. Two of these sites had been subjected to grazing before trapping took place and two sites had been protected. Twe two grazed sites were located at Bonny Reservoir, Yuma County, and on Mesa de Maya in Las Animas County. The two ungrazed sites were located at Flagler Reservoir State Recreation Area in Kit Carson County and Bonny Reservoir. 2. Sandsage - typical species in this site were Artemisia curtipendula, and Yucca sp. 3. Little Bluestem - this was situated (Schizachyruim scoparium). in a protected filifolia, Bouteloua stand of Little Bluestem Results In a 6249 trap/night effort individuals of 3 of the 6 select species were captured. In the selective trapping for Scalopus aquaticus none were collected, but a live specimen was captured by hand. A total of 213 individuals of 10 species was captured during these trapping sessions. The results are listed in Tables 1 and 2 in terms of total captures and in terms of captures per 1000 T/N • • Discussion Peromyscus leucopus: This species occurs throughout the eastern United States, but its distribution becomes dendritic in the western portions of its range (Fleharty and Stadel 1968). By most accounts, this species shows a marked preference for wooded habitats, which, in the western segment of its range, occur in streamside communities (see Fleharty and Stadel 1968, Kaufman and Fleharty 1974, and Choate and Fleharty 1975 for examples). �102 Table 1. Small Mammal Trapping Results in Lowland Communities 1. Grazed Riparian Species Peromyscus manicu1atus Reithrodontomys megalotis Reithrodontomys montanus Perognathus f1avus 2. Ungrazed Riparian Species Peromyscus manicu1atus Reithrodontomys megalotis Microtus ochrogaster Cryptotis parva 3. Pan;cum-Distichlis 4. Wild Hay July 1978 No. Captured T/N=1800 7 ( 4) 26 (14) 8 ( 4) 1 (.56) 42 3 8 7 1 19 T/N=599 ( 5.00) (13.00) (12.00) ( 2.00) June 1979 No. Captured Peromyscus maniculatus Reithrodontomys mega10tis Microtus ochrogaster T/N=1800 16 ( 9) 25 (14) 1 (.56) 1 (.56) 3 December 1978 No. Captured Peromyscus manicu1atus Reithrodontomys megalotis Microtus ochrogaster Cryptotis parva *Numbers in parentheses July 1978 No. Captured T/N=800 2 15 ( 3.0) (19.0). 1 ( 1.0) 18 = number of captures per 1,000 trap nights. �103 Table 2. Small Mammal Trapping Results in Upland Communities Shortgrass (grazed) Bonny R~servoir Species No. Captured ~romyscus maniculatus 2 (20) Onychomys leucogaster 2 (20) Spermaphilus tridecemlineatus 1 (10) Di podomys ordi i 11 (11 0) May 1979 Shortgrass (ungrazed) Species Peromyscus maniculatus Reithrodontomys montanus Onychomys leucogaster May 1979 T/N=lOO 16---------------- Bonny Reservoir No. Captured -----------------4 1 (10) 1 2 (10) (20) T/N=lOO Shortgrass (grazed) Mesa de Maya (Las Animas County) Aug. 1978 Species No. Captured Peromyscus maniculatus 8 (80) 'j (10) Onychomys leucogaster Perognathus hispidus 1 (10) 10 Shor tgra ss (un gra zed) Fla 9 1er S. R .A. (K it--=-Ca:.:.;r-=s-=o.;.;.n_C::..:o:...:u:.:.;n..:.:tYL-)~ _ ___.:.T.:.../.;..;.N=_1:. Speci es No. Captured Peromyscus maniculatus 2 (20) 2 Sandsage Species Peromyscus manicu1atus Reithrodontomys mega10tis Onychomys 1eucogaster Perognathus hispidus Dipodomys ordii Microtus ochrogaster Little B1uestem Species Dipodomys ordii Peromyscus maniculatus Reithrodontomys megalotis Onychomys leucogaster "July 1978 T/N=250 No, Captured W 9 2 4 '1. 1 jO' (40) (36) ( 8) (16 ) (16) ( 4) March 1979 No. Captured 12 (20) u (18.3) ( 6.7) 4 2 29 ( 3.3) T/N=600 �104 In Colorado ~eromyscus leu copus is known from riparian and shrub communities in the southeastern corner of the state (Armstrong 1972). Its status and distribution in the northeastern quarter of the state is unknown although suitable habitat does not seem to be lacking (Armstrong 1972). Despite a rather exhaustive trapping program in riparian woodland communities in the South Republican River drainage, no~. leucopus was captured. It seems unlikely that this species does in fact occur in this drainage at this time. Reithrodontomys montanus: Individuals of this species were collected in both tile grazed riparian site and in the ungrazed shortgrass site. Habitat preferences of !. mont anus in southwestern Kansas were described by Hill and Hibbard (1943), and in southeastern Wyoming by Maxwell and Brown (1968). Occurrence of this species is to be expected throughout eastern Colorado in grassy habitats (Armstrong 1972), but its exact habitat requirements remain unknown. Perognathus hispidus: Individuals of this species were captured in the ungrazed sand-sage site at Bonny Reservoir and in the grazed short grass site on Mesa de Maya in Las Animas County. This species is apparently inactive during winter months (Choate and Fleharty 1975). Perognathus flavus: A single individual of this species was collected in the grazed riparian site. P. flavus shows a preference for short grass communities in southeastern Wyoming-(Maxwell and Brown 1968), but none was collected in shortgrass habitats during this particular study. Perognathus flavescens: The ecology of this species is virtually unknown throughout its range (Armstrong 1972). Maxwell and Brown (1968) described habitat preferences of this species in southeastern Wyoming. During the present study no individuals were captured although a variety of prairie habitat types were sampled. Scalopus aguaticus: An individual was captured at Bonny Reservoir and observed for 15 days in captivity. During this period it fed on the carcasses of Microtus ochrogaster, Reithrodontomys megalotis, Peromyscus maniculatus and Dipodomys ordii. It also ate earthworms and ground beef. Special Note A portion of the findings in this study was published as follows: Moulton, M. 1978. Small mammal associations in grazed versus ungrazed cottonwood riparian woodland in eastern Colorado. In Graul, W.D. and S.J. Bissell, tech. coord. Lowland river and stream habita~in Colorado:a symposium (Greeley, CO, Oct. 4-5, 1978). Colo. Chap. Wildl. Soc. and Colo. Audubon Council, 195p. This report segment prepared Michael P. Moulton by: �105 White-faced Ibis Segment In association with initial surveys of all potential ibis nesting areas in the San Luis Valley, in south-central Colorado, three areas were identified in late May and early June, 1978, as likely nesting areas: (1) Adams Lake - 5 miles south of Alamosa, (2) Head Lake - 6 miles east of Mosca, and (3) Trites Lake 11 miles south of Saguache. Adams Lake Although from 2-30 ibises were seen feeding at this location through June, no nests were found. The absence of nesting may have been due to a rapid lowering of the water level in late May, caused by irrigation practices. This drop in the water level apparently caused the abandonment of a Snowy Egret colony (20 birds) and a Black-crowned Night Heron colony (60 birds). A small group of ibises did nest at this location in 1976. Head Lake Although ibises did nest at this location in 1976, no nests were found in 1978. A severe drop in the water level is thought to have been the cause of the lack of ibis nesting in 1978. Ten Snowy Egret and 20 Black-crowned Night Heron nests were abandoned in June, as the water level dropped. Despite the lack of nesting, from 3-30 ibises were observed at this location throughout the summer. Trites Lake This lake, with a 2.5 mile perimeter, had an average water depth of 2.5 feet in early May and a 1 foot average depth in August. Although emergent vegetation at this location includes hardstem bullrush (Scirpus acutus) and broad-leaved cattail (Typha latifolia), all ibis nests were found in the hardstem bullrush. The bullrush covered about 30% of the lake's surface area. The ibis nests were located in a heronry (Black-crowned Night Herons and Snowy Egrets) which covered an area of about 300 yds x 200 yds. From nests each This nests late May through early June 9 ibis nests were located and monitored. These contained 23 eggs or young when found, but only 4 young were fledged (2 nests produced 2 young). This yields an average of 0.44 young fledged per nest. is extremely low when compared to the 1976 Trites Lake data. In 1976, 11 produced 17 young - 1.55 young fledged per nest. Although the exact reasons for the low ibis fledging success at Trites Lake in 1978 were not known, details of the fate of individual nests were recorded. Four nests, containing a total of 12 eggs, were deserted. One nest contained 3 eggs and they were all cracked, and another nest contained only 1 egg, which was also cracked. One nest containing 3 young, was either abandoned or the adults perished - the young died in the nest. The Black-crowned Night Herons and Snowy Egrets also experienced problems at this location in 1978. A total of 81 night heron and 82 egret nests was found. Since time did not allow intensive monitoring of all nests, 13 heron and 18 egret were selected for detailed study. �106 The 13 Black-crowned Night Heron nests contained a total of 43 eggs or young when found, but only 3 young fledged (l from one nest and 2 from another). This yields an average of 0.23 young fledged per nest. The 18 Snowy Egret nests contained 57 eggs or young when found, and a total of 14 young fledges - 0.77 young fledged per nest. An estimated 27% of the 82 egret nests and 30% of the 81 heron nests perished because they were left unattended. This does not include nests found with cracked eggs, since the latter may have been caused by predation. The cause of the high number of nests with eggs and young left unattended is unkown, but it may be related to a conflict at a nearby private fish hatchery - a favorite feeding area for the herons and egrets. Namely, the owner of the hatchery apparently shoots large numbers of these birds each year to protect his fishery resource. An attempt was made to band most young in the mixed colony. Snowy Egrets and 54 Black-crowned Night Herons was banded. Statewide Waterbird A total of 110 Survey Segment Great Blue Heron Approximately 200 mail survey forms were distributed in January, 1979 (an exact count was not kept) and 57 were returned. The returned survey forms accounted for 27 active Great Blue Heron hero~ries and 14 inactive ones. Later field work resulted in an additional 11 active and 7 inactive heronries being located. Thus, a total of 38 active Great Blue Heron nesting sites and 21 inactive ones were located (Figure 1). Details for all active and inactive colonies are presented in Appendice I and II. Although the Great Blue Heron heronries are widely distributed throughout the state, the greatest concentration of these heronries is in north-central Colorado. The active heronries ranged from 1-85 nests in size, but 61% contained from 1-15 nests. The known years of existence for the heronries ranged from 1-65 years with 52% only known to be in existence fur less than 10 years. The estimated size of the inactive heronries ranged from 1-40 nests with 83% of them containing 1-10 nests. The precise causes of abandonment for all the inactive sites are not known, but some documented causes include removal of nesting trees, development of reservoirs with associated human recreation activity, and highway construction. Additional work is needed to determine what the heronries will and will not tolerate. The preceding data can be used to draw some tentative conclusions. The smaller heronries seem to be most subject to abandonment, but overall there does seem to be a lot of movement, i.e. a substantial number of sites have been abandoned and a majority of existing sites apparently have been in existence for less than 10 .years. This overall lack of stability is not surprising in that these birds tend to colonize old cottonwoods in wet situations. Also, these water areas are subjected to overall high levels of human activity, which may displace the birds. Perhaps this explains why several of the older heronries are in north-central Colorado (this area has many private irrigation reservoirs that are not subjected to high levels of human activity). �FIGURE 1. GREAT BLUE HERON (197-9)· COLORADO MO"'Ar Ro//rr * * * • RIO """'Q ~FllLJ) ""'l JA •• ** *.- ** O:)UGlAS /1""lIlI'J *1f'(O(GA ..•. ~: rc I Til.". L.&::'AJlrlt\·v!vN I (lS(IU * AITU..'?SOH LIf{CO~N * ucu«, ...Ai..UO(iI"C'-; r * * ** * CA I~OC: I'I(LI) I •... 1.* o -.J PASO 'H!f!NAJ!_ --- MQI/TIICS£ I IKm~ I'U(S,O 1c~''''''L{Y I p~CWf~" Bfllr SA"_tO"/!. t'IlIlrSO.JU 00LOl/15 MO"T{ tUM' e"" ,e._. IICII VI.. I f;.. ~ Active Heronries Inactive Her6nri~s * �FIGURE 2. DOUBLE-CRESTED CORMORANT (1979) COLORADO M:;" ..<T «u» .,su.; .... ,c.{ !tJ';,t,v '··'a,",] •••. IlIO I't'A;If/ .••·cr.;;N • • st» ""Q '("'GAN !,,".A • CA f"'lUJ ElS(,,'" L''-';::'.:J:''N IfI""U"S~'" ""l S. •....• a 00 CH~" CN:r~/"AJ! ,MOli r eas« I(IOrl. !'lI.ao'/L(Y I ~ p~c·"'~I.! litNT ",J DOLOlI.fl I A4AAIOS· MO,,11IUMA • Active Colonies ~.s .•. ",I,·\AS CA" �109 Double-crested Cormorant Eight Great Blue Heron heronries also contained nesting populations of the Doublecrested Cormorant (Phalaerocorax auritus). Interestingly, most of the cormorants were restricted to the northeast part of the state (Figure 2). The only exception to this is the one cormorant nesting area not associated with the Great Blue Heron. Namely, two nests were found on an island in Antero Reservoir (Park County - South Park). These two nests were in a nesting colony of California Gulls. Other Avian Select Species Snowy Plover, Least Tern, Black-necked Stilt Charles Chase III surveyed the reservoirs associated with the Arkansas River drainage in southeastern Colorado during the nesting seasons of 1978 and 1979. In 1978 he found 37 Snowy Plover nests distributed as follows: 2l-Cheraw Reservoir, 3-Horse Creek (Blue) Reservoir, 7-Adobe Creek Reservoir, 6 Nee/Noshe Reservoir. In addition, he found 1 Least Tern on Horse Creek Reservoir and 1 on Adobe Creek Reservoir. All of the preceding nests were associated with expanses of alkali flats, which are characteristic of the Arkansas River drainage. Because of the alkali condition, this area is apparently extremely important for the preceding two species. The Snowy Plover does not nest in this concentration in any other part of the state; in fact, nesting has not been confirmed in other parts of Colorado, although it is suspected that they sometimes nest in the 'San Luis Valley. The preceding Least Tern nests are important in that there was previously only one nesting record for Colorado--in 1949. In 1979 the Snowy Plover nests were distributed as follows: 47-Cheraw Reservoir, l5-Horse Creek Reservoir, 6-Nee/Noshe Reservoir. Two Least Tern nests were found at Nee/Noshe and 1 at Horse Creek Reservoir. The preceding 3 nests were all flooded, and apparently all three pairs involved renested at Horse Creek Reservoir 3 late nests were found. A Black-necked Stilt nest with 6 eggs was found at Cheraw Reservoir in conjunction with the Snowy Plovers. California Gull Antero Reservoir (Park County). A substantial nesting colony (about 200 pair) has been at this location for several years. Both in 1978 and 1979, young were banded by Charles Loeffler and Dr. Ronald A. Ryder. Riverside Reservoir (Weld County). A nesting colony has existed at this location since 1962 and young are banded by Division of Wildlife personnel and Dr. Ryder annually. In 1979 the colony was estimated to consist of 30-50 breeding pair. Eleven-mile Reservoir (Park County). In 1978 a small colony (no exact count) nested for the first time at this location. In 1979 the nesting island was under water (the usual condition) and, consequently, no nesting occurred. White Pelican A nesting colony has existed on Riverside Reservoir since 1962. In recent years the colony has consisted of between 300-400 nesting pairs. In July, 1978, 100 juvenile pelicans were banded. Technical advise was also provided by Walter Graul �110 on two major pelican management projects. One involved a substantial habitat stabilization effort by the U.S. Bureau of Land Management on the Riverside Reservoir nesting island. The other involved preliminary planning for the construction of a new island in conjunction with a proposed new reservoir (Wildcat Reservoir to be built by Public Service Company of Colorado). Greater Sandhill Crane Efforts in the spring of 1979 consisted of meeting with U.S. Forest Service personnel in an effort to develop a cooperative management plan between the Forest Service and the Division of Wildlife for the Routt Forest nesting population. The cooperative effort, involving intensive field studies and inventories, is to be implemented in 1979-80. Mountain Plover Nesting in Lincoln County (5 miles south of Limon) was confirmed in early June, 1979. Additionally, suspected breeding birds were found on Mesa de Maya in July, 1978. These locations will be added to the known distribution in the final report. Bat Segment The 1979 field season began in May and extends through October. For this season, the 1979 results will be presented in the next annual progress report. The 1978 results will be presented herein. Between 1 May 1978 and 30 August 1978, approximately 50 nights of work at 28 locations were completed. Habitats of various types were sampled. About 300 bats were caught and examined. One hundred and thirty-two bats were prepared as voucher specimens. The most significant finding was locating a colony of 10,000 Mexican free-tail bats (Tadarida brasiliensis). Evidence of breeding was noted. Specimens from this colony have been sent to the Patuxent unit of the U.S. Fish and Wildlife Service for analysis of pesticide residue; pesticides are thought to be a major factor in the population decline of this species. The colony was located in an abandoned mine shaft on the east side of the San Luis Valley. �t 111 LITERATURE CITED Armstrong, D.M. 1972. Distribution of mammals in Colorado. Nat. Hist., Univ. Kans. No.3, 4l5pp. Monogr. Mus. Bissell, S.J. 1978. Colorado mammal distribution latilong study. Div. Wildl., Denver, CO., 20pp. Colo. Choate, J.R. and E.D. Fleharty. 1975. Synopsis of native, recent mammals of Ellis County, Kansas. Dcc. Pap. Mus. Texas Tech. 37:1-80. Fleharty, E.D. and D.L. Stadel. 1968. Distribution of Peromyscus leucopus (Woods Mouse) in western Kansas. Trans. Kans. Acad. Sci. 71:231-233. Graul, W.D., J. Torres and R. Denney. 1976. A species-ecosystem nongame programs. Wildl. Soc. Bull. 4:79-80. approach for Hess, D. 1977. Today's strategy •.•tomorrow's wildlife: a comprehensive management plan for Colorado's wildlife. Colo. Div. Wildl., Denver, CO. 96 pp. Hill, J.E. and C.W. Hibbard. 1943. Ecological differentiation between two harvest mice (Reithrodontomys) in western Kansas. J. Mamm. 24:22-25. Kaufman, D.W. and E.D. Fleharty. 1974. Habitat selection by nine species of rodents in north-central Kansas. Southwestern Nat., 18:443-452. Kingery, H.E. and W.D. Graul. 1978. Colorado bird distribution latilong study. Colo. Div. Wildl., Denver, CO. 58pp. Maxwell, M.H. and L.N. Brown. 1968. Ecological distribution of rodents on the high plains of eastern Wyoming. Southwestern Nat., 13:143-158. Prepared by: IIL~ CJ. 9k~<A~ Walter D. Graul Wildlife Research Leader �APPENDIX I --- ACTIVE GREAT BLUE HERON COLONIES IN COLORADO (1979) Location Years in Existence1 Tentative Status Information Source3 Number of Nests4 Decreasing* Carolyn Armstrong 1-5 Stable* Carolyn Armstrong 36-40 Other Nesting Species in Colony5 Adams County 1. Barr Lake (East) 15 mi. N.E Denver TIS, R66W, Sec. 28, SW~. 20-25 2. Barr Lake (West) 15 mi. N.E. Denver TIS, R66W, Sec. 27, SE~. 40 3. Horse Creek Reservoir 6 mi. S.E. Hudson, TIN, R64W, Sec. 31, SE~. Unknown Double-crested Cormorant Double-crested Cormorant, Black-crowned Night Heron Unknown D.O. 16-20 Double-crested Cormorant Black-crowned Night Heron, Great Egret Boulder County' 4. Boulder Creek Between Hwy. 287 and 95th St., TIN, R69W, Sec. 16, SE~. 30-35 Stable D.O. 76-80 5. Panama Lake 6 mi. S.W. Longmont, T2N, R69W, Sec. 35 NE~. 40 Stable D.O. 1-5 Double-crested Cormorant ,_. ,_. N �APPENDIX Location Years in Existence1 I CONTINUED Tentative Status Information Source3 Number of Nests4 Other Nesting Species in Colony5 Douglas County 6. Chatfield Reservoir 5 mi So. Littleton, T6S, R69W, Sec. 12. 5-10 Increasing* D.O. , Hugh Kingery 81-85 D.O. 1-5 Black-crowned Night Heron Eagle County 7. Gypsum T5S, R85W, Sec. 6, NE~. Unknown Unknown •..... •....• LV 9arfield County 8. Colorado River 8.1 mi W. Rifle, T6S, R94W, Sec. 29, SW~. Unknown Stable 9. Colorado River at Grand Valley T7S, R9G\\f, Sec. 13, SE~. Unknown Unknown 10. Colorado River at Grand Valley, T7S, R96W, Sec. 13, SE~. Unknown Increasing D.O. 11-15 11. Colorado River 0.5 mi S.W. Silt, T6S, R92W, Sec 10 Unknown Unknown D.O. 31-35 N~.J~1;. D.O. Howard Green 16-20 1-5 �APPENDIX Location I CONTINUED Years in Existence1 Tentative Status 15-20 Stable* Lloyd Palmer 11-15 10 Unknown .D.O. Kevin Cook 21-25 John Wagner 1-5 D.O. 1-5 Information Source3 Number of Nests4 Other Nesting Species in Colony5 Grand County 12. Colorado River, 6 mi. E. Kremmling, TIN, R80W, Sec. 18, NE~. Gunnison County 13. Gunnison River 5.5 mi. SW Gunnison, T49N, RIW, Sec. 8, SW~. Jackson County 14. Walden Reservoir, 3 mi. W. Walden T12N, R80W, Sec. 24, SE~. 6 Stable* Kit Carson County 15. So. Republican Drainage 3 mi. S.E. Flagler Larimer Black-crowned Night Heron 20-25 Stable County 16. Lone Tree Reservoir, 2 mi. NW Berthoud T4N, R69W, Sec. 9, NE~. 5 Decreasing D.O. Camille Cummings 41-1+5 Black-crowned Night Heron •.... •.... ~ �APPENDIX Location Years in Existence1 I CONTINUED Tentative Status Information Source3 Number of Nests4 17. Fossil Creek Res., So. Ft. Collins, T6N, R68W, Sec. 9, SW!z; 10 Decreasing D.O. 61-65 18. Horseshoe Lake 1 mi. N.E. Loveland T6N, R68W, Sec. 31, SW!z;. 5 Increasing D.O. 1-5 19. Timnath Reservoir 4 mi. N.E. Timnath, T7N, R68W, Sec. 24, SW!z;. 5 Increasing D.O. 1-5 20. Wellington Res. 113 1 mi. N.E. Waverly, T9N, R68W, Sec. 18, SE!z;. 5-10 Stable Lee Franz 16-20 50 Stable D.O. 26-30 25-30 Stable D.O. 6-10 Other Nesting Species in Colony5 Black-crowned Night Heron •.....• •.....• V1 Logan County 21. Jumbo Reservoir 8 mi. W. Sedgwick, TUN, R47W, Sec. 7, SE!t;. 22. Sterling Res. 14 mi. N. W. Sterling, T9N, R53W, Sec. I, NE!t;. Double-crested Cormorant �APPENDIX Location Years in Existence1 Tentative Status Unknown Decreasing* I CONTINUED -- Information Source3 Number of Nests4 Other Nesting Species in Colonys Mesa County 23. Colorado River 4 mi. So. DeBeque, T9S, R97W, Sec. 18 SE~. Moffat 11-15 County 24. 13 mi. N.E. Craig, T7N, R89W, Sect. 3, SE~. 25. Green River ~ mi. S.E. Brown's Park National Wildl. Refuge Prowers D.O. Chuck Woodward 1-5 Stable* D.O. 11-15 Unknown .• Decreasing D.O. 1-5 Unknown Unknown D.O. 1-5 15 . Rio Blanco County 27. White River 7 mi. E. Meeker, TIN, R93W, Sec. 33, NW~. ,_. ,_. (j\ County 26. Two Buttes Creek 30 mi. So. Lamar, \\lestend of State Mgmt. Area (Two Buttes). Increasing 2 �APPENDIX I CONTINUED Years in Existence1 Tentative Status 28. White River 4 mi. W. Meeker, TIN, R94W, Sec. 30, SE%. Unknown Unknown D.O. 1-5 29. White River 0.5 mi. E. Rio Blanco State Mgmt. Area, TIN, R96W, Sec. 5, SW~. Unknown Unknown D.O. 6-10 Location Information Source3 Number of Nests4 Routt County to-' to-' -...J 30. Yampa River 5 mi , W. Steamboat Springs, T6N, R85W, Sec. 9, NW~. 10 Unknown D.O. 21-25 31. Yampa River 5.25 mi. W. Steamboat Springs, T6N, R85W. Sec. 9, NW~. 10 Unknown . D.O. 1-5 50 Stable D.O. 11-15 Washington County 32. Prewitt Reservoir T5N, R4W, Sec. 14. N~. Other Nesting Species in Colony5 �APPENDIX I CONTINUED Location Years in Existence1 Tentative Status Information Source3 Number of Nests4 Ron Zacagnini 41-45 D.O. 1-5 Other Nesting Species in Colony5 Weld County 33. Empire Reservoir 3 mi. E. Masters, T3N, R61W, Sec. 1, NE%. 50 Stable 34. Franklin Lake 0.5 mi. S.E. Severance, T6N, R67W, Sec. 1, NW!t;. 5 Increasing 35. Johnstown 1.25 mi. N. Johnstown, T5N, R67W, Sec.33, SW!t;. .- 00 60-65 Increasing Brad Kirshner 46-50 36. Milton Res. 9 mi. W. Platteville T3N, R65W, Sec. 10, SE!t;. 5-10 Decreasing D.O. 46-50 37. Riverside Res. 2 mi. N. Masters, T4N, R62W, Sec. 12, NE!t;. 10-15 Unknown D.O. 1-5 Stable D.O. Yuma County 38. So. Republican River 1.1 mi. E. Hale, T5S, R43W, Sec. 12, SE!t;. Double-crested Cormorant, Blackcrowned Night Heron 5 - 1-5 Double-crested Cormorant, Blackcrowned Night Heron Double-crested Cormorant �APPENDIX 1. The "years in existence" is based on interviews 2. The asterisk indicates that the assigned are, therefore, tentative). 3. D.O. refers to direct field observation 4. This value was determined 5. Same as 4. I CONTINUED and/or questionnaires category is based on detailed by personnel by direct observation from people familar with the sites. data (those without the asterisk in this study. and/or interviews, and/or questionnaires. ,_. ,_. \0 �APPENDIX II INACTIVE GREAT BLUE HERON COLONIES IN COLORADO (1979) If Location Information Source of Nests Prior to Abandonment1 Probable Reason for Abandonment2 Boulder County 1. Wonderland Lake NW edge Boulder, TIN, R71W, Sec. 13. SE~. Inis Baker 1-5 Trees cut for housing development 2. East LyonsT3N, R70W, Sec. 20,N~. Inis Baker Unknown Commercial development ,_. N o Delta County 3. 4 mi. N. DeltaT15S, R96W, Sec. 17, SW~. Hal Burdick 6-10 Unknown 4. Fruit Grower's Res. 1 mi. So. Eckert, T14S, R94W, Sec. 18, SW~. Merle Hodges 6-10 Unknown Howard Green 1-5 Highway Construction Eagle County 5. 6 mi. W. Gypsum, T5S, R86W, Sec. 5, NE~. �APPENDIX II CONTINUED Location Information Source # of Nests Prior to Abandonment! Probable Reasons for Abandonment2 Garfield County 6. 2.5 mi. SE Carbondale T7S, R87W, Sec. 36, Ann Loughteridge 6-10 Corral construction Ann Abbott 1-5 Nests removed by people NW~. Jefferson County 7. Standley Lake 88th and 100th Ave. T2S, R69W, Sec. 20, •.... N •.... SWl.t;. Kit Carson Larry Strode 6-10 Reservoir drained 1966, human activity at heronry. 9. Barbour Ponds 7 mi. E. Longmont, T2N., R86W, Sec. 3, S ~. Eva ~adenacher 1..,5 Gravel pit and recreation area near the heronry. 10. Bodecker Res. 3.25 mi. W. Campion, T5N, R69W, Sec. 20 Judy Sisler 1-5 Housing development nearby. 8. SW,Reservoir, 2 mi. SW Flagler Larimer County m.J'~. �APPENDIX II CONTINUED # of Nests Prior Location Information Source to Abandonment1 Probable Reasons for Abandonment2 11. Boyd Lake 1 mi. NE Loveland, T6N, R68W, Sec. 32, N~. Judy Sisler Unknown Housing development adjacent to heronry. 12. Terry Lake (Island) N. edge Ft. Collins, T8N, R69W, Sec. 26, SE~. Ronald Ryder Unknown Cutting down of trees l~. Terry Lake (Shore) T8N, R69W, Sec. 26, S~. Rene Ellis 14. Lonetree Res. 3 roiW. Campion, T4N, R69W, Sec. 9, N~. Camille Cummings 75 15. Colorado River 3.2 mi. So. DeBeque T9S, R97W, Sec. 8, SE~. Joe Gumber 1-5 Unknown 16. Colorado River 4 mi. So. Debeque T9s,R47W, Sec. 18, S~. Joe Gurober 6-10 Highway Construction nearby. 1-5 Unknown •.... N N lllegal shooting) recreational activity adjacent to area. Mesa County' �APPENDIX II CONTINUED # of Nests Prior to Abandonment1 Probable Reasons for Abandonment2 Location Information Source 17. Colorado River 2.25 mi. W. Fruita, TIN, R3W, Sec. 14, NE~. John Gray 21-25 Highway constructed through he ronry , 18. Colorado River 1.5 mi. W. Fruita, TIN, R3W, Sec. 13. SE~. John Gray 11-15 Unknown •..... Moffat County 19. Yampa River 1.5 mi. SW Craig, T6N, R91W, Sec. 11, N~. N W Samuel Scanga 6-10 Unknown Dal Schaefer 6-10 Nesting Trees cut with reservoir development, intensive boating. Charles Reichertt 6-10 Unknown Pueblo County 20. Arkansas River 4 mi. W. Pueblo, Pueblo Reservoir area. Rio Blanco County 21. White River 4 mi. E. Meeker, TIN, R93W, Sec. 19, S~. 1. Size of heronriesprior to abandonment was determined by interviews and/or questionnaires. 2. Probable reason for abandonment as determined by site inspection, interviews, and/or questionnaires. � https://cpw.cvlcollections.org/files/original/9ee845c66b4c0846eb0dc2f651348614.pdf a7f11f74d2fe7e1d58321fe7a74eb758 PDF Text Text 124 JOB PROGRESS REPORT State of COLORADO Project No. W-41-R-30:_ Work Plan No. 3 --- Job Title: Contagious Bighorn Job No. Ecthyma Shee__pand Mountain Goat Iny_~t_i.g§ttions 2 (Sore Mouth, Orf) in the Ruminant Big Game of Colorado Period Covered: July 1. 1979 through June 30, 1980 Personnel: W. Lance, J. DeMartini, C. Hibler, and L. Pearson, State University; R. Schmidt and W. Adrian, Colorado Division life Colorado of Wild- ABSTRACT Two hundred and three bighorn sheep from eight major bighorn sheep herds within Colorado were examined physically and serologically for contagious ecthyma. Clinical disease was seen in one herd (Saguache) and strong serological evidence of the disease was found in three others (Chalk Cliffs, Cottonwood and Tarryall). Controlled studies in captive adult bighorn sheep indicate that bighorn sheep are highly susceptible to contagious ecthyma but mortality in adults is not expected in cases uncomplicated by other conditions. Bighorn lambs exposed to the bighorn strain of contagious ecthyma developed extensive severe lesions which could lead to mortality in a free-ranging animal. Mule deer, white-tailed deer, elk, and pronghorn antelope exposed to the bighorn strain of contagious ecthyma developed only mild transient oral lesions. �CONTAGIOUS ECTHYMA (SORE MOUTH, ORF) IN THE Ru}lINANT BIG GAME OF COLORADO William Lance P.N. OBJECTIVES The objective of this study is to document the distribution and prevalence of a contagious ecthyma-like agent in the bighorns of Colorado and to give management agencies concrete information as to the potential of this agent for producing mortality in free-ranging bighorn herds. It will determine whether or not this agent, and its potential mortality effect upon bighorn sheep needs to be taken into consideration in the formulation of future resource management policies in Colorado. SEGMENT OBJECTIVES 1. To determine the distribution and prevalence of contagious ecthyma (and/or Saguache Agent) in the bighorn sheep herds of Colorado. 2. To define the pathogenicity and mortality agent for bighorn sheep lambs. 3. To define the species susceptibility and pathogenesis of the Saguache agent for the native ruminant big game species of Colorado. MATERIALS Distribution and Prevalence potential of the Saguache AND METHODS Study Serum was collected from 203 individual sheep from eight bighorn sheep herds within Colorado in conjunction with the normal trapping and transplanting activities of the Colorado Division of Wildlife during the 1979-1980 season. Each sheep bled was physically examined for active clinical lesions of contagious ecthyma or the presence of white to gray scars on the lips and muzzle which result from recent recovery from the disease. The serum samples were tested for antibodies to contagious ecthyma by the complement fixation technique, a common and widely accepted procedure. Host Range Study Four mule deer fawns, three white-tailed deer fawns, two pronghorn antelope fawns and four elk calves, all from free-ranging stock, were successfully hand reared at the Wild Animal Disease Center Foothills Facility. At approximately 12 weeks of age three mule deer fawns, two white-tailed deer fawns and three elk calves were exposed to the ground suspension of lesion scab material from the original case of bighorn contagious ecthyma. These were exposed by a method identical to the one used for the adult bighorn sheep. One from each group served as a control. Both pronghorn fawns were exposed, with no control available. These animals were monitored for evidence of disease for 45 days. �126 Bighorn Sheep Pathogenicity Study Five of a group of nine adult bighorn sheep were exposed to a ground suspension of scab material from the original case of bighorn contagious ecthyma from Saguache (Sept. 1978). The oral mucosa, nasal mucosa, and axillary skin was lightly scarified and exposed to two drops of scab suspension. These animals were monitored over the next 120 days to observe: 1) development, duration and regression of clinical lesions, 2) spread of the disease between individuals, 3) characteristics of the lesions and resultant scars, and 4) ability of observers to detect affected sheep in a group under simulated field conditions. Two 12 week old bighorn lambs, head reared in isolation, were exposed to scab material in a similar manner to document the effect of the disease on young sheep. These were monitored closely for the next 60 days. RESULTS AND DISCUSSION Distribution and Prevalence The serological data is summarized in Tables 1-8. Sixteen of 27 bighorn sheep examined from the Trickle Mountain herd (Table 1) had active clinical lesions of contagious ecthyma (CE) and positive serological titers. Eight serologically positive animals from a group of forty from the Cottonwood herd (Table 2) indicate that the disease recently had been present in that area. Similar results for the Chalk cliffs herd (Table 3) indicate CE is present in the contagious bighorn sheep herds from Saguache to Buena Vista, Colorado. Results from the Sugarloaf portion of the Tarryall herd (rable 4) indicates the disease has been present, but the fact that no lambs were serologically positive suggests that CE was probably active prior to June 1979. The data from the Gordon ranch trapsite in the Tarryall range is also suggestive of the presence of CE in this herd. The Grant herd results (Table 6) also are suggestive, but not conclusive, that the disease is present in that area. No conclusions can be made from the single positive animal found in the Basalt herd (Table 7). The Poudre River herd (Table 8) is free of any positive reactors. Serum samples collected prior to March 1979 from the Poudre River, Pikes Peak, Waterton Canyon, Chalk Cliffs, Tarryall, and Trickle Mountain herds prior to March 1978 were all negative (Lance, unpublished data). By some unknown mechanism, contagious ecthyma has suddenly appeared in the bighorn sheep herds of southern and central Colorado. The herds of the Front Range and the northern portion of the state (Poudre River, Waterton Canyon, and Pikes Peak) are free of the disease at this time. At present, it is impossible to explain its rapid appearance in the Saguache, Collegiate Range and Tarryall areas since 1978. The status of the small remnant and newer transplant herds in the state is unknown. �Bighorn Sheep Pathogenicity Study Seven days following exposure to bighorn sheep contagious ecthyma scab material, all exposed animals developed oral and nasal mucocutaneous lesions that could be identified up to 60 meters distance by a trained observer aided by a spotting telescope. At 10 days post exposure (PI) all five exposed sheep could be identified as having lesions by untrained personnel aided by a spotting scope to a distance of 60 meters. Thirteen days PI all four unexposed control sheep had naturally transmitted disease which could be easily detected at 60 meters with a spotting scope. As the lesions progressed in size and new metastatic lesions developed in the oral cavities and over the nasal area, the sheep became increasingly restless, constantly rubbing the lesions against inanimate objects and nearby sheep in an effort to relieve the intense itching. In some individual sheep, the proliferative lesions became confluent over the entire oral and nasal mucocutaneous junctions and complicated with moderate to severe secondary bacterial infections. In no instances did the lesions appear to impair normal feeding activity in the adult sheep. The most severely affected sheep in a group could be identified with the unaided eye due to their COllstant rubbing of the lesions and licking of the nasal lesions. Individual animals euthanized at stages of the disease gave a more complete picture of the pathology produced. In most cases, lesions in adult sheep were completely healed in 28 to 35 days PI. Animals with severe lesions consistently had large depigmented white to gray areas in the mucocutaneous junction of the oral cavity and muzzle area. The irregular depigmented areas persisted for at least six months following the disease. These scars in recovered sheep may be used as a field mark to identify sheep that have had severe clinical CE and recovered. When two 12 week old hand-reared bighorn lambs were exposed to the bighorn CE virus, both lambs developed severe clinical cases of the disease. In a ewe lamb, weight loss was apparent as the clinical disease progressed presumably due to decreased feeding ability and the constant irritation from the itching and discomfort of the developing lesions. This ewe lamb constantly rubbed the oral, muzzle and, later, eye lesions against any available object. At 24 days PI, hard crusty proliferative lesions of CE spread to the eyelids and at one point produced corneal abrasions. As the eyelids crusted shut, the lamb became functionally blind due to these lesions. At this time local and systemic antibiotic therapy was started to prevent the loss of this lamb due to the secondary effects of CEo The second ram lamb survived the clinical disease without any sup'portive therapy. By 60 days PI he was completely free of active lesions although large white to gray scars were present on the oral and nasal areas underlying the old lesion sites. The severe clinical response of the two bighorn lambs supports the opinion expressed in the published literature that CE may be a primary or secondary mortality factor in young bighorn lambs (Samuel et al., 1975). The predisposing factors necessary for CE to be a mortality factor in lambs are unknown, but may include such factors as high lungworm burdens, intraspecific stress (crowding), and nutritional or climatic stress. �128 Host Range Studies Three each three month old, hand-reared, mule deer fawns, whitetailed deer fawns, elk calves, and two pronghorn antelope fawns were exposed to the bighorn strain of the CE virus. All exposed animals developed oral mucocutaneous lesions seven to ten days PI that never exceeded 2 ern in diameter. Secondary metastatic lesions that developed were minimal. All lesions began to regress at 14 to 16 days PI and were completely gone by day 21 PI. Although the bighorn CE virus did produce small localized lesions in the other wild ruminants under the conditions of this study, it is unlikely that this would occur under natural free-ranging conditions. The controlled studies demonstrated that this virus is of no concern to these species. Prepared by: W. R. LanceY.M. jJ.$/f!~ .. �129 ,,< Table 1. Serological Age Sex Ear Tag Lamb M Lamb results, Saguache herd, Trickle Mountain 1980. CE Titre Comments Red 1 1:10 Active lesions of contagious ecthyma over mouth and incisors M Red 2 Negative No active lesions or scars Lamb M Red 3 Negative No active lesions or scars Lamb F Red 4 No sample Small lesion left side at commissure Lamb M Red 5 1:80 Multiple active lesions Lamb M Red 6 No sample Multiple regressing Lamb M Red 7 1:20 Lesions around base of teeth and lips. Scars on nose. Lamb M Red 8 Negative Clean- no scars Lamb F Red 9 1:80 Severe multiple lesions, pigment loss on lips Lamb F Red 10 1:40 Scars on nostril Lamb F Red 11 Negative Small, minimal lesions. stages of disease. Lamb F Red 12 No sample Severe lesions on lips oral cavity clean Lamb F Red 13 Negative Small single lesion, upper right nostril. Lamb F Red 14 1:20 Severe metastatic lesions. Lesions on hard palate and around incisors. Lamb F Red 15 Negative No lesions Lamb F Red 17 1:10 Large, thick lesions, left side, right clean, teeth clean. Yearl F Red 16 1:10 No active lesions 2 yr M Red 18 1:10 Prominent scars on medial septum. Small circumscribed area, right nostril. No oral lesions. Adult F Red 80 1:20 Small white spots on nose. Old regressing lesions present. Adult F Red 81 1:20 Adult F Red 59 No tests No lesions, no scars Adult F White radio collar 1:10 No lesions * Neck Collar site, February Interpretation: suspicious of previous lesions Early Old scar area over nose Complement-fixing antibody titers of 1:5 or 1:10 are considered exposure. Complement-fixing antibody titers of 1:20 or greater �130 Table 1. (Continued) Age Sex Ear Tag Adult F Adult Neck Collar CE Titre Comments Blue 30 1:10 Minimal lesions, 0.5 cm in diameter F Blue 29 1:40 No active lesions or scars Adult F Blue 28 1:10 No active lesions or scars Adult F Blue 27 1:10 No active lesions or scars Blue 4 �131 Table 2. Serological * results, Neck Collar Cottonwood Creek herd, April 1980. Comments Age Sex Ear Tag CE Titre Lamb F G 26 Negative Lamb F G 28 Negative Lamb F G 27 1:5 Lamb M G 29 Negative Lamb M G 30 Negative Lamb F G 31 Negative Lamb M G 32 Negative Lamb M G 33 1:10 Lamb M G 34 1:10 Lamb F G 34 Unknown Lamb M G 35 Negative Lamb M G 37 Unknown Not bled Yearl F G 38 Unknown Not bled Year 1 F G 24 Yearl WRC8002700 1:20 F B 69 Negative 2 yr F B 62 Non-specific 2 yr M WRC9002704 Negative 2 yr F B 64 Negative 2 yr F B 65 Negative 2 yr F B 67 1:10 2 yr F B 68 Negative 2 yr F B 70 Negative 2 yr F B 41 Negative 2 yr F 2 1/2 F B72 Negative 3 yr F B 66 Negative 3 yr F B 78 Negative 3 yr F B 76 Negative 3 yr F B77 Negative 3 yr F B 52 Negative 3 1/2 F B 74 1:10 4 F B 61 Non-specific 4 F B71 Negative 4 F B 54 Non-specific G 25 Depigmented area on internal surface of left nostril Not bled reaction Depigmented nostril area on right 1:10 G 39 reaction reaction *Interpretation: Complement-fixing antibody titers of 1:5 or 1:10 are considered suspicious of previous exposure. Complement-fixing antibody titers of 1:20 or greater are indicative of previous infection. �132 Table 2. (Continued) Age Sex Ear Tag 4 1/2 F B 53 Negative 4 1/2 F B 73 1:10 5 F B 63 Negative 5 F B 75 Non-specific reaction 5+ F G 23 Neck Collar WRC7002697 CE Titre Negative Comments White scar on nose �133 Table Age Sex Ear Tag Lamb F Lamb 3. Serological Chalk Cliffs, February 1980 CE Titre Comment RT 20 Negative No active lesions or scars F RT 29 Not bled No active lesions or scars Lamb M RT 22 Negative No active lesions or scars Lamb F RT 23 1:20 No active lesions or scars Lamb M RT 24 Negative No active lesions or scars Lamb M RT 25 Negative No active lesions or scars Lamb F RT 26 Not bled No active lesions or scars Lamb F RT 28 Negative No active lesions or scars Lamb F RT 29 Not bled No active lesions or scars Lamb M RT 33 1:10 No active lesions or scars Lamb M RT 34 Negative No active lesions or scars 1 yr F RT 30 1:10 No active lesions or scars 1 yr M RT 35 Negative No active lesions or scars 1 yr F RT 36 Negative No active lesions or scars 1 yr F RT 37 37 Negative No active lesions or scars 1 yr M RT 38 No active lesions or scars 1 yr F RT 52 Yellow radio 1:5 148.525 34 1:5 No active lesions or scars 1 yr M Blue 6] Negative No active lesions or scars L yT ~ R'I 1 No active Lesions or scars 2 yr F RT 3L 1: LU No active lesions or scars 2 yr M l{T Non -s pe ctf r c No active l.esions or scars Negat No activ~ L~sions or scars r. io No active l.eaions or scars Non-specifi( No active lesions or scars 1:10 No active lesions or scars Negative No active lesions or scars No active lesions or scars No active lesions or scars 1:10 No active lesions or scars Red 85 Not bled Not examined Red 89 Not bled Not examined Red 56 Not bled Not examined 2 yr 33 27 4i I{T )4 2 yr F 2 yr M Blue 62 3 yr F RT 49 3 yr F RT 39 4 yr F RT 31 5 yr F RT 21 5 yr Neck Collar results~ 36 Lli 1H Yellow radio Non-specific 148.625 Not bled RT 70 Interpretation: Complement-fixing antibody titers of 1:5 or 1:10 are considered suspicious of previous exposure. Complement-fixing antibody titers of 1:20 or greater are indicative of previous infection. �134 Table 4. Serological Age Sex Ear Tag Lamb M Lamb * results, Neck Collar Terryall herd, Sugarloaf site, February 1980. CE Titre Comments RT 40 Negative No active lesions or scars F RT 41 Negative No active lesions or scars Lamb F RT 43 Negative No active lesions or scars Lamb M RT 47 Negative No active lesions or scars Lamb M RT 48 Negative No active lesions or scars Lamb M RT 50 Non-specific 1 yr. M RT 45 Negative No active lesions or scars 1 yr. M RT 49 Negative No active lesions or scars 1 yr. M RT 51 Negative No active lesions or scars 1 yr. M RT 52 Negative No active lesions or scars 1 yr. F 1:5 No active lesions or scars 2 yr. F RT 44 Negative No active lesions or scars 2 yr. M RT 46 1:10 No active lesions or scars 3 yr. F WT 58 Negative No active lesions or scars 3 yr-. M BT 9 1:10 No active lesions or scars 4 yr. F Negative No active lesions or scars 4 yr. M 1:20 Small scar on upper left nostril 6 yr. F 1:10 No active lesions or scars 7 yr. 44 40 43 61 reaction No active lesions or scars F Blue 45 White 26 41 No sample 8 yr. F Blue 42 1:10 No active lesions or scars 8 yr. F WR 148.750 1:5 No active lesions or scars 9 yr. F Blue 46 White 25 Nonspecific No active lesions or scars RT 42 *Interpretation: Complement-fixing antibody suspicious of previous exposure. Complement-fixing are indicative of previous infection. titers of 1:5 or 1:10 are considered antibody titers of 1:20 or greater �135 Table 5. Serological results from Terryall herd, Gordon Ranch site, March 1980 Age Sex Ear Tag Lamb M G 3 Lamb M G Lamb F Lamb Neck Collar CE Titre Comment Negative No active lesions or scars Non-specific reaction No active lesions or scars G 7 Negative No active lesions or scars M G 10 Non-specific reaction No active lesions or scars Lamb F G 12 Negative No active lesions or scars Lamb F R 95 Negative No active lesions or scars Lamb F R 96 Negative No active lesions or scars Lamb M R 90 Negative No active lesions or scars Yearl F G 5 Negative No active lesions or scars Yearl F G 6 Negative No active lesions or scars Yearl M G 9 Negative No active lesions or scars Yearl F R 91 Negative No active lesions or scars Yearl F R92 Negative No active lesions or scars Yearl F R 93 Negative No active lesions or scars Yearl M R 94 Negative No active lesions or scars Yearl F R 97 Negative No active lesions or scars Yearl F R 98 Negative No active lesions or scars Yearl F R 99 Negative No active lesions or scars 2 yr F G 1 1:5 Scar on nostril 2 yr F l4/G 2 Negative No active lesions or scars 2 yr F BET 8/G 2 Negative No active lesions or scars 5 yr F 1:5 No active lesions or scars 6 yr F Negative No active lesions or scars Adult F BET 2/ Red 84 RT 89 Orange 76 Not bled No active lesions or scars Adult F Orange 77 Not bled No active lesions or scars Adult F Orange 79 Not bled No active lesions or scars Adult F Orange 87 Not bled No active lesions or scars Adult F Orange 81 Not bled No active lesions or scars Adult F Orange 75 Not bled No active lesions or scars 4 Red 82 BET/ll * Interpretation: Complement-fixing antibody titers of 1:5 or 1:10 are considered suspicious of previous exposure. Complement-fixing antibody titers of 1:20 or greater are indicative of previous infection. �136 Table 6. Serological Age Sex Ear Tag Lamb M Lamb results from Grant Mountain Neck Collar herd, Geneva Creek site, April 1980 CE Titre Comment G 13 1:5 No scars or active lesions M G 18 Negative No scars or active lesions Lamb F G 20 Negative No scars or active lesions Yearl F G 15 (837) Negative No scars or active lesions Yearl M .G 16 Negative No scars or active lesions Yearl F G 19 Negative No scars or active lesions Yearl F G 21 Negative No scars or active lesions Adult F G 17 1:10 No scars or active lesions Adult F G 22 Negative No scars or active lesions Adult F B 43 Not tested No scars or active lesions Adult F B 45 R 26 1:5 No scars or active lesions Adult F B 51 R 17 1:10 No scars or active lesions Adult F B 58 R 22 Negative No scars or active lesions Adult F None R 44 Non-specific Adult F B 56 R 23 Not tested reaction No scars or active lesions No scars or active lesions *Interpretation: Complement-fixing antibody titers of 1:5 or 1~10 are considered suspicious of previous exposure. Complement-fixing antibody titers of 1:20 or greater are indicative of previous infection. �137 Table 7. Age Sex Ear Tag Adult F Adult Serological Basalt herd, March 1980. CE Titre Conunent B 8 Negative No active lesions F B 12 Negative No active lesions or scars Adult F B l3 Negative No active lesions Adult F B 14 Negative No active lesions or scars Adult F B 15 Negative No active lesions Adult F B 16 Negative No active lesions or scars Adult F B 18 Negative No active lesions or scars Adult F B 19 1:10 No active lesions or scars Adult F B 20 Negative No active lesions or scars Adult F B 21 Negative No active lesions or scars Adult F B 22 Negative No active lesions Adult F B 29 Negative No active lesions or scars Adult F R 81 Negative No active lesions or scars Adult F R 82 Negative No active lesions or scars Adult F R 83 Negativw No active lesions Adult F R 84 Negative No active lesions or scars Adult M R 85 Negative No active lesions or scars Adult F R 86 Negative No active lesions or scars Adult F R 87 Negative No active lesions or scars * Interpretation: Neck Collar results, or scars or scars or scars or scars or scars Complement-fixing antibody titers of 1:5 or 1:10 are considered suspicious of previous exposure. Complement-fixing antibody titers of 1:20 or greater are indicative of previous infection. �138 Table 8. Age Sex Ear Tag Lamb M R Lamb Serological Poudre River Herd, March 1980. CE Titre Comments 55 Negative No active lesions or scars M R 57 Negative No active lesions or scars Lamb M R 58 Non-specific No active lesions or scars Lamb M R 59 Negative No active lesions or scars Lamb F R 60 Negative No active lesions or scars Lamb M R 61 Negative No active lesions or scars Lamb F R 62 Negative No active lesions or scars Lamb M R 63 Negative No active lesions or scars Lamb M R 64 Negative No active lesions or scars Lamb M R 70 Non-specific No active lesions or scars Lamb M R 71 Non-specific No active lesions or scars 1 M R 69 Negative No active lesions or scars 1 M R 79 No sample No active lesions or scars 1 M R 74 Negative No active lesions or scars 1 F R 73 No sample No active lesions or scars 2 F Black 16 Negative No active lesions or scars 2 F R 80 Negative No active lesions or scars 3 F R 72 Negative No active lesions or scars 3 F R 77 Negative No active lesions or scars 3 F R 78 Non-specific No active lesions or scars 4 F R 76 No sample No active lesions or scars * Interpretation: Neck Collar Results, Complement-fixing antibody titers of 1:5 or 1:10 are considered suspicious of previous exposure. Complement-fixing antibody titers of 1:20 or greater are indicative of previous infection. �139 LITERATURE CITED Samuel, W.M., G.A. Chalmers, J.G. Stelfox, A. Loewen and J.J. Thomsen. 1975. Contagious'ecthyma in bighorn sheep and mountain goat in western Canada. J. Wildl. Dis. 11: 26-31. �140 JOB PROGRESS State of Colorado . -------------------------- Project No. W-41-R-30 ----------------------- Work Plan No. Job Title Period Covered: Personnel: 3 Trapping, REPORT Bighorn Sheep and Mountain Investigations 1 Job No. Translocating, and Inventory Goat of Bighorn Sheep July 1, 1979 through June 30, 1980. R. Schmidt, G. Jones, M. Conners, R. Dix, R. Edmiston, T. Ostertag, J. Rodgers, P. Yates, W. Rutherford, plus DWM's,. Area Supervisors, and Wildlife Technicians in specific localities. ABSTRACT Bighorn sheep trapping, treating, and translocation activities during the segment year resulted in 1,182 sheep receiving chemotherapeutic drug treatment for lungworm control. This figure includes both free-ranging sheep treated with drugs in apple pulp bait, and captured sheep treated orally. The total number of sheep trapped was 255, in 8 different catches. Of these, 104 were translocated to 5 different release sites, 12 were taken to CSU for disease study, 12 were transported to the State of Arizona, 12 were transported to the State of Nevada, and 115 were marked and released at trapsite. Updated estimates of statewide bighorn populations, by individual herd, are given. This information was collected directly in conjunction with field work, and by interview with District Wildlife Managers. Pellet group samples from 14 separate areas were collected, and analyzed by the DOW research laboratory in Fort Collins for lungworm larvae output. �141 TRAPPING, TRANSLOCATING, AND INVENTORY Roaert L. Schmidt and William OF BIGHORN SHEEP H. Rutherford P. N. OBJECTIVE The principal objective of this study will be to increase the total number of bighorn sheep within the State of Colorado by either introducing bighorn into areas which were historically bighorn sheep ranges and where no animals are in evidence today, or by adding additional animals to small remnant herds. By expanding bighorn sheep from crowded areas into other areas it is hoped that they will be better able to utilize the range and increase their numbers. Also the introduction of new animals, especially rams, into remnant populations should improve the reproductive capacity and livability of the lambs by improving the gene pool and lowering the percentage of inbreeding. SEGMENT OBJECTIVES 1. Trap and trans locate between 2. Treat all captured bighorn sheep with chemotherapeutic drugs for the control of Protostrongylus spp. lungworm parasitism and other parasites or diseases (800-900 individual animals). 3. Monitor selected herds and all translocated bighorn sheep for the degree of parasitism by collecting fecal pellet groups and checking them by standard laboratory procedures. METHODS 100 and 150 bighorn sheep. AND MATERIALS Baiting, trapping, drug treatment, and monitoring procedures are described in detail in the Job Progress Report for 1976-1977 (Schmidt 1978) and will not be repeated here. RESULTS All trapping, treating, and translocating activities during the segment year and the three previous years are summarized in Table 1. Following this, data on all sheep marked during the segment year (captured, marked, and released at trapsite, and captured, marked, and translocated) are tabulated in Tables 2 through 10. Updated estimates of sheep population statewide by individual herd, provided by District Wildlife Managers and project personnel, are presented in Tables 11 and 12. Bighorn sheep pellet groups were collected from 14 separate areas. These coitections, with one exception, were either from untreated herds or from herds that had not yet been treated this year. Pellet collections from post-treatment herds have not yet been analyzed. The one exception is from the Rampart Range herd, where both pre-treatment and post-treatment collections were made. Comparison of larvae output data from these collections showed a marked reduction in output following treatment. �Table 1. Treating, trapping, and translocating data from three previous seasons. Number treated, both freerangJIl_gand captured 1976- 1977- 1978- 19791977 1978 1979 1980 Area Basalt Chalk Cliff s Cottonwood Cr. Dillon Mesa Georgetown Grant Little Hills Mt. Evans Ouray Pikes Peak Poudre River Rampart Range RMNP Saguache Cr. Tarryall Cr. Taylor River Waterton Totals 23 o 70 82 94 107 o o o o 9JJ o o 6 23 37 o o o 31 11 211 216 38 174 204 31 o o o 112 l35 208 301 171 149 71 70 o o o 978 1534 771 1182 o llAll 20 marked and released 1/13 marked and released i/20 and translocated l/20 marked and released of treatment at trapsite at trapsite o 38 o o 51 30 36 33 o o o o o o o o o 16 o o o 51 63 49 41 o o 22 232 o 15 268 122 331_l (See Table 2). Canyon (See Table (See Table 4). questionable. (See Table 2). Number translocated1976- 1977- 1978- 19791977 1978 1979 1980 o o o o 1~1 o o o o o o o o o 3.s.!QI o o o o o o o o 19 25 12 o o o 20 o o o 2911/ 28 20 20 24..!i1 o 17 21 o o 112 90 III -- 20 marked See previous and translocated o 20!!_1 20E_! o o o 59..!i1 255 o o 12 o o 9 o o o 20 sheep given for current year only. (See Table 2). to Sawpit ~/3 marked and released at trapsite o (See Table 2). to Carrizo at trapsite ~/20 marked and translocated IIQuality o 74 42 82 54 l/Disposition of trapped and translocated for data from previous years. marked 40~/ o o 19791980 o o o o o of 1977-80, with comparative II o o o o o II Number trappecF 1976- 1977- 19781977 1978 1979 2021 121 10 winter 36 13 0 0 98 91 150 55 0 215 234 0 0 10ill sheep, Colorado, 26 20 125 188 0 0 20 53 138 6 76 16 106 bighorn o o 122i o o o o o o o 2011/ 16 o o o Progress .pN 24.!1/ 4412/ o o 37 ,_. o o o 140 Reports to Button Rocky (See Table 6) ..!lIs marked and released at trapsite (See Table 2). 131 3);-- 24 marked and translocated to Hot Creek (Table 7). ..!illS 151 marked and released at trapsite (See Table 2). -- 20 marked and translocated to Brown's Canyon (See Table 8). 12 marked and transported to State of Arizona (Table 9). 12 marked and transported to State of Nevada (See Table 10). 2/12 marked and taken to CSU for disease study (See Table 5) ...!il All 24 marked and released lQ/1S marked and released at trapsite (See Table 2). at trapsite (See Table 2). �Table 2. Bighorn sheep trapped, treated, marked, and released at trapsite, Colorado, winter of 1979-80. Trapping Area Water ton Waterton Waterton Water ton Water ton Waterton Waterton Waterton Waterton Water ton Waterton Water ton Waterton Waterton Waterton Waterton Waterton Waterton Waterton Waterton Waterton Date Canyon Canyon Canyon Canyon Canyon Canyon Canyon Canyon Canyon Canyon Canyon Canyon Canyon Canyon Canyon Canyon Canyon Canyon Canyon Canyon Canyon 12-14-79 12-14-79 12-14-79 12-14-79 12-14-79 12-14-79 12-14-79 12-14-79 12-14-79 12-14-79 12-14-79 12-14-79 12-14-79 12-14-79 12-14-79 12-14-79 12-14-79 12-14-79 12-14-79 12-14-79 12-14-79 Waterton Canyon Sex Age F F F F F F F F F lamb lamb lamb lamb lamb lamb lamb lamb lamb 3 2 4 3 4 2 3 3 1 M M 1 1 F 3 12-14-79 F 2 Waterton Canyon 12-14-79 M 2 Waterton Canyon 12-14-79 M 4 Saguache Creek Saguache Creek Saguache Creek Saguache Creek Saguache Creek Chalk Cliffs Chalk Cliffs Chalk Cliffs 2-07-80 2-07-80 2-07-80 2-07-80 2-07-80 2-12-80 2-12-80 2-12-80 F F F F 1 Unk Unk. Unk. 4 5 lamb lamb M M F F F M F M M M M M M Collar color and number Ear tag color and number None Blue - 17 None Blue - 18 Blue - 19 None None Blue - 21 Blue - 22 None None Blue - 23 None Blue - 24 None Blue - 90 Blue - 91 None None Red - 1 None Red - 2 Red - 3 None Red - 4 None None Red - 5 None Red - 6 None Red - 7 None Red - 8 Red - 9 None None Blue - 92 Blue - 93 None Green radio Freq. 148.900 None Yellow Radio None Freq.148.800 Blue radio Freq. 148.700 None Orange/Blue radio Freq. 148.950 None None Red - 16 None Red - 81 None Red - 59 None Red - 80 Blue - 85 None Red - 21 None Red - 22 None None Red - 25 Remarks Not Not Not Not Not Not Not Not Not Not Not Not Not Not Not Not Not Not Not Not treated treated treated treated treated treated treated treated treated treated treated treated treated treated treated treated treated treated treated treated Not treated Not treated Not treated Not treated ------------------------------------------------------------------------------------------------------------- •.....• .j::- w �Table 2. Bighorn sheep trapped, treated, marked, and released at trapsite, Colorado, winter of 1979-80. Trapping Area Chalk Cliffs Chalk Cliffs Chalk Cliffs Chalk Cliffs Chalk Cliffs Chalk Cliffs Chalk Cliffs Chalk Cliffs Chalk Cliffs Chalk Cliffs Tarryall Creek Tarryall Creek Tarryall Creek Tarryall Creek Tarryall Creek Tarryall Creek Tarryall Creek Tarryall Creek Tarryall Creek Tarryall Creek Tarryall Creek Tarryall Creek Tarryall Creek Tarryall Creek Tarryall Creek Poudre River Poudre River Poudre River Poudre River Poudre River Poudre River Date 2-12-80 2-12-80 2-12-80 2-12-80 2-12-80 2-12-80 2-12-80 2-12-80 2-12-80 2-12-80 2-19-80 2-19-80 2-19-80 2-19-80 2-19-80 2-19-80 2-19-80 2-19-80 2-19-80 2-19-80 2-19-80 2-19-80 2-19-80 2-19-80 2-19-80 3-07-80 3-07-80 3-07-80 3-07-80 3-07-80 3-07-80 Sex F M F F M M M F F F M M M M F F F F F F F F F F F F F M M M M Age lamb lamb I 3 Unk. 5 2 Unk. Unk. Unk. 4 Unk. 3 lamb Unk. Unk. Unk. Unk. Unk. Unk. Unk. Unk. Unk. Unk. Unk. Unk. Unk. lamb lamb lamb lamb Collar color and number None None None None None None None Red - 85 Red - 89 Red - 56 None None None None Orange - 75 Orange - 76 Orange - 77 Orange - 78 Orange - 79 Orange - 81 Orange - 83 Orange - Unk. Orange - Unk. Orange - 86 Orange - 87 Old yellow Old yellow None None None None Ear tag color and number Red - 28 Red - 33 Red - 36 Red - 39 Red - 51 Red - 70 Blue - 63 None None None White - 61 White - Unk. Black - 9 Unk. (escaped) Black - 11 None None None None None None None None None None Red - 54 Red - 56 Red - 63 Red - 64 Red - 70 Red - 71 (Cont'd). Remarks Found dead ,_. ~ .c- Recapture Recapture -~~------------------------------------------------------------------------------------------------------------- �Table 2. Bighorn sheep trapped, treated, marked, and released at trapsite, Colorado, winter of 1979-80. Trapping Area Poudre Poudre Poudre Poudre Poudre Poudre Poudre Poudre Poudre Basalt Basalt Basalt Basalt Basalt Basalt Basalt Basalt Basalt Basalt Basalt Basalt Basalt Basalt Basalt Basalt Basalt Basalt Basalt Basalt Grant Grant Grant River River River River River River River River River Date 3-07-80 3-07-80 3-07-80 3-07-80 3-07-80 3-07~80 3-07-80 3-07-80 3-07-80 3-18-80 3-18-80 3-18-80 3-18-80 3-18-80 3-18-80 3-18-80 3-18-80 ·3-18-80 3-18-80 3-18-80 3-18-80 3-18-80 3-18-80 3-18-80 3-18-80 3-18-80 3-18-80 3-18-80 3-18-80 3-28-80 3-28-80 3-28-80 Sex Age Collar color and number Ear tag color and number (Cont'd). Remarks Due to recorder error, data on the remaining 9 sheep are not available. Some were recaptures of previously marked sheep, and some were newly marked. Fifteen sheep were released at the trapsite, but only 6 were recorded. Four ear tags were used - Red #53, Red #66, Red #67, and Red #68. Of these, three were released at the trapsite, and one was included in the release at Button Rock. F F F F M F F F F F F F F F F F F F F F F M M Unk. Unk. Unk. Unk. lamb Unk. Unk. Unk. Unk. Unk. Unk. Unk. Unk. Unk. Unk. Unk. Unk. Unk. Unk. Unk. 2 lamb lamb None None None None None None None None None None None None None None None None None None None None None None None Red - 81 Red - 82 Red - 83 Red - 84 Red - 85 Red - 86 Red - 87 Blue - 8 Blue - 12 Blue - 13 Blue - 14 Blue - 15 Blue - 16 Blue - 18 Blue - 19 Blue - 20 Blue - 21 Blue - 22 Blue - 29 Blue - 46 Black - 43 Green - 13 Green - 18 -------------------------------------------------------------------------------------------------------------- I-' .po. lJ1 �Table 2. Bighorn sheep trapped, treated, marked and released at trapsite, Colorado, winter of 1979-80. Trapping Area Cottonwood Cottonwood Cottonwood Cottonwood Cottonwood Cottonwood Cottonwood Cottonwood Creek Creek Creek Creek Creek Creek Creek Creek Date 4-08-80 4-08-80 4-08-80 4-08-80 4-08-80 4-08-80 4-08-80 4-08-80 Sex F F M F F F F F Age 3 lamb lamb lamb lamb I 2 4 Collar color and number Blue - 78 None None None None None None None Ear tag color and number None Green Green Green Green Green Green Green - (Cont'd). Remarks 31 35 36 37 38 39 40 •..... l:(J'\ �Table 4. Bighorn sheep trapped at Cottonwood Creek, Chaffee County; released at Sawpit, San Miguel County, April 8, 1980. Sex Age F F F F F F F F F F F 4 2 5 2 2 3 2 2 1 1 6+ 2 lamb lamb lamb lamb lamb lamb lamb lamb M F F F fiI M M M M Collar color and number Blue Blue Blue Blue Blue Blue Blue Blue Blue Yellow Yellow Yellow None None None None None None None None 61 62 63 64 65 66 67 68 69 radio frequency 148.800 radio frequency 148.700 radio frequency 148.900 - .. Ear tag color and number None None None None None None None None None Green Green Green Green Green Green Green Green Green Green Green - 24 23 25 26 27 28 29 30 32 33 34 Remarks ,_. "'" '-l �Table 5. Bighorn sheep trapped at Grant, Park County; taken to Colorado State University for research on Johne's disease, March 28, 1980. Sex Age Collar color and number F F F F F F 7 5 7 3 3 1 1 Unk. Red Red Red Red Red None None None None None None None M F F F F F I lamb I Unk. 17 22 23 26 44 Remarks Ear tag color and number Black _.51 Black - 58 Black - 56 Black - 45 None Blue - 37, Green - 15 Green - 16 Green - 17 Green - 19 Green - 20 Green - 21 Green - 22 Table 6. Bighorn sheep trapped in Poudre Canyon, Larimer County; released at Button Rock, Boulder County, March 7, 1980. Sex Age M M M M lamb lamb lamb lamb lamb lamb lamb lamb lamb 3 4 2 3 3 1 2 1 F M F M M F F F F F F ]I M M M ? 1 1 ? Collar color and number None None None None None None None None None Blue - 47 Blue - 48 Blue - 49 Blue - 50 Blue - 51 Yellow radio frequency 148.750 Orange radio frequency 148.850 Orange/black radio freq. 148.950 None .None ? Ear tag color and number Red Red Red Red Red Red Red Red Red Red Red Black Red Red Red Red Red Red Red ? 55 57 58 59 60 61 62 65 75 72 76 - 16 77 78 73 80 69 79 74 Remarks Due to recorder error, four ear tags that were used were not recorded. These are Red - 53, Red - 66, Red - 67, and Red - 68. Of these four, one was included in this release at Button Rock; the other three were released at the trapsite in Poudre Canyon. See remarks above. I-' .po. '" �Table 8. Bighorn sheep trapped at Tarryall Creek, Park County; released at Brown's Canyon, Chaffee County, February 19, 1980. Sex Age M lamb lamb 8 lamb 2 F F F F M M M M M M M M F F F F F F F I 2 lamb lamb I lamb I I 3 7 8 4 I 6 9 Collar color and number None None White radio freq. 148.750 None White radio freq. 148.550 White radio freq. 148.650 None None None None None None None Blue - 40 Blue - 41 Blue - 42 Blue - 43 Blue - 44 Blue - 45 Blue - 46 Ear tag color and number Red Red Red Red Red Red Red Red Red Red Red Red Red White None None None None None None 40 41 42 43 44 45 46 47 48 49 50 51 52 - 58 Remarks ,_. LIl 0 �Table 7. Bighorn sheep trapped at Trickle Mountain, Saguache Creek, Saguache County, released at Hot Creek, Conejos County, February 7, 1980. Sex Age Collar color and number M M M lamb lamb lamb lamb None None None None None None None None None None None None None None None F M M M M F F F F F F F F M F F F F F F F I lamb lamb lamb lamb lamb lamb lamb lamb lamb lamb lamb 2 2 4 3 3 2 6 Unk. None None Blue - 27 Blue - 28 Blue - 29 Blue - 30 White radio, frequency 148.600 Red radio, frequency 148.500 Yellow radio frequency 148.950 Ear tag color and number Red Red Red Red Red Red Red Red Red Red Red Red Red Red Red Red Red None None None None None None None Rema1:'ks 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 17 18 >-' Vl ,_. Found dead �Table 9. Bighorn sheep trapped at Tarryall Creek, Park County; transported to State of Arizona, Feb. 19, 1980. Sex Age F F F M M M F F F M M F Collar color and number Ear tag color and number Remarks 5 None Green - 11 2 2 2 lamb lamb 1 1 lamb 1 lamb lamb None None None None None None None None None None None Old black - 8; New green - 8 Green - 1 Green - 2 Green - 3 Green - 4 Green - 5 Green - 6 Green - 7 Green - 9 Green - 10 Green - 12 Recapture - old orange collar #82. Collar removed. Recapture >-' VI N Table 10. Bighorn sheep trapped at Tarryall Creek, Park County; transported to State of Nevada, Feb. 19, 1980. Sex Age Collar color and number Ear tag color and number F F lamb 6 None None Red - 88 Old black - 21; New red - 89 M lamb 1 1 1 1 lamb lamb 1 1 1 None None None None None None None None None None Red Red Red Red Red Red Red Red Red Red F F F M F F F F M - 90 91 92 93 94 95 96 97 98 99 Remarks Recapture - old orange collar #84. Collar removed. �153 Table 11. Current (1980) population estimates of bighorn Colorado, compared with 1970 population estimates. Area 1970 estimate Arkansas River Beaver Creek Battlement Mesa Blanco Basin Brush Creek Buffalo Peaks Cimmaron Peak Clinetop Mesa Collegiate Range Dinosaur National Monument Georgetown-Empire Glenwood Canyon Gore Range Lake City Mount Evans Mount Silverheels Mount Zirkel Mesa Verde National Park Ouray-Cow Creek Poudre Canyon-Rawah Pikes Peak Pole Mountain Rampart Range Roan Creek Rocky Mountain National Park Redstone Rifle Hogback Saint Vrain Drainage, South End R.M.N.P. Sangre de Cristo Range San Luis Peak Sapinero Mesa South Platte River Canyon Sheep Mountain Snowmass Tarryall Taylor River Trickle Mountain Vallecito Creek West Elk Mountains Wilson Peak TOTALS 20 20 35 ? sheep herds, 1980 estimate 30 30 35 ? o o 50 40 15 100 139 75 12 30 70 160 15 110 75 15 200 130 40 o 30 90 75 300 14 20 3 30 90 250 15 o 30? 100 250 200 14? 60 o o 190 25 150 60 o o 6 70 150 6 18 40 25 100 40 175 12 45 6 100 200 10? 60 40 75 250 30 500 30 45? ? o 2,212 3,263 �154 Table 12. Population estimates (1980) of new bighorn established by translocations during past four years, Area Alamosa River Apishapa Canyon Basalt Brown's Canyon Button Rock Carrizo Canyon Conejos River Cow Creek, R.M.N.P. Cross Mountain Greenhorn Mountains Grand Junction (desert bighorns) Hot Greek Lone Pine Creek Monument Creek Sawpit Soap Creek Lit tle Hills TOTAL Population estimate sheep herds Colorado. Years elapsed since release 40 1 65 50 2 3 29 o 30 30 50 45 50 25 12 34 o o 95 35 2 1 26 o 20 2 3 12 648 1 2 2 3 1 o �Table 13. Lungworm larvae output as determined Collection Date Location Ouray pre-treat1/ by laboratory Analysis Date analysis of bighorn No. of Sample Groups o 1 sheep pellet groups. Frequency 1/ By Output Category- 234 2/ Average5 Cross Mtn.. hapa-4/ AP1S 5/29/79 12/01/79 8 3 5 6/20/79 12/10/79 17 o 17 o o . kl·e M tn. pre-treat- 3/ TrlC 1/29/80 4/10/80 43 1 30 8 2 2 Lone Pinef!-/ 5/22/79 12/10/79 9 6 3 o 7/10/79 12/10/79 14 4 10 o o o Ford Cree~ 3/ ? /80 7/24/80 20 16 2 2 o o o o o o o o o o o o Mueller Ranch~/ 2/20/80 7/24/80 21 3 9 3 1 1 4 2.00 Rampart pre-treat 1/03/80 7/09/80 20 1 9 6 1 1 2 1. 90 post-treat 2/16/80 7/25/80 17 10 7 o 0.41 1/29/80 7/08/80 22 2 9 11 o o o Chalk Cliffs pre-treat1/ o o o 1. 41 Black Canyon pre-treat1/ 3/03/80 5/19/30 20 1 2 7 3 3 4 2.85 Waterto~/ 4/15/80 5/09/80 7 1 4 2 o 5/09/80 25 6 17 2 o o o 2.14 4/02/80 2/ ? /80 5/06/80 19 o o o o 8 6 1 4 3.05 Tarryall pre-treat- 3/ 4/ '4/ Cottonwood Poudre Cr. pre-treat- pre-treat- 3/ 3/ 2/ ? /79 12/12/79 18 o 10 6 o 2 2/l3/80 4/25/80 76 14 26 29 5 2 o o o o 1. 67 1.41 0.62 1.00 1.40 0.33 0.71 0.30 1.84 1/ Larval output category values are assigned as follows: Larvae absent - 0; 1-50 larvae/gram of feces - 1; 51-250 larvae/gram - 2; 251-500 larvae/gram - 3; 501-1000 larvae/gram - 4; 1000+ larvae/gram - 5. 2/ This is a weighted value established by multiplying category values by frequencies, .dividing the summation by the total number of sample groups. 3/ Post-treatment 4/ - No treatments data not available at time of writing. were done on these herds. summing, and then ,_. V1 V1 �156 LITERATURE CITED Schmidt, R. L. 1978. Trapping and translocating bighorn sheep. Federal Aid P-R Progress Report, Project W-41-R-27, January: Prepared by Colorado 57-75. �157 JOB PROGRESS State of Project No. COLORADO I Job No. Bald and Golden Eagle Nesting Period Covered: Personnel: Raptor Investigations W-124-R --------------------- Work Plan No. Job Title REPORT Harch _ 1 Studies 15, 1978 through February 28, 1979 Gerald R. Craig, Thomas Bohanon, Aldin Forbes, Joe Frothingham, James McKinley, Wayne Russell and Gordon Saville, Colorado Division of ~.Jildlife ABSTRACT Four hundred and thirty golden eagle nests were checked, largely by aerial survey, for nesting activity and productivity from April 15 to May 20, 1978. Nests determined to be productive were rechecked from approximately June 1-20 to document the number of young birds which survived the nest period to fledge. Results indicate that while the number ~f active nests per total nests checked has remained stable since 1975, the number of young birds produced per active nest has risen 216% since 1974, and 37% since 1975. The number of eaglets fledged per successful nest remained steady compared to 1977 figures, and shows an increase of 36% when compared to 1975. These increases are presumed to correspond with increased prey availability, which was also at a low in 1974. Considerable effort was made in 1978 to consolidate and organize records on the golden eagle survey from 1973 to present. These efforts resulted in county and statewide tabulations of various parameters involved in the chronological reproductive patterns of the bird. �158 BALD AND GOLDEN EAGLE NESTING STUDIES Gerald R. Craig and Thomas C. Bohanon Although the golden eagle (Aquila chrysaetosJ population in Colorado appears to be increasing since its low point in 1974 (Crowley 1975), its predominance in areas of the state which are rapidly being developed for water and energy purposes merits a close monitoring of population dynamics and reproductive status. Furthermore, the maintenance of permanent nest locations and the ease of finding them, the fact that the bird is present in statistically significant numbers, and its presence as a large carnivore at the end of its food chain lend significance to not only the study of man's effect on the Golden Eagle in specific, but to the long term effects of man on the bird's expansive environment in general. Only two active bald eagle nests have been documented in Colorado although the state is generally considered to be peripheral as nesting habitat. Recent reclassification has designated bald eagles as endangered species in Colorado thus each site should rec8ive protection frommsturbance and habitat degradation. Prior to such protection, it will be necessary to delineate important hunting areas and buffer zones adjacent to the nest sites. P.N. OBJECTIVES The objectives of this study are: (1) to estimate the breeding numbers and obtain production data of bald and golden eagles nesting in Colorado; (2) to identify important nesting areas and associated hunting habitats of bald and golden eagles in Colorado; and (3) to compile data and submit reports to associated state personnel and federal agencies for use in delineating and protecting eagle nest sites. SEGMENT OBJECTIVES lao Continue to locate and map nesting sites of golden and throughout Colorado. Nest searches will be conducted wing aircraft and in some instances with helicopter. field work will be done from the ground by vehicles. of nest sites will be taken and pertinent information physical features of the habitat. bald eagles with fixedAdditional Photographs recorded about lb. Nesting areas will be stratified by such features as climate, elevation and habitat type. Sample areas then will be delineated which are representative of important nesting areas. The sample areas throughout the state will be flown with a fixed-wing aircraft in late April and early May to ascertain the number of active sites. The same sites again will be checked from the air in June to determine the nesting success and productivity of the sample sites. The percent of active sites, percent of successful pairs, and total production will be extrapolated for each region of the state. �159 2a. When nests are visited in la, details ,viII be recorded as to characteristics of the nesting habitat. Nesting eagles also will be observed from a distance to locate and map key hunting areas. 2b. Radio transmitters may be attached to several young bald eagles and a sample of young golden eagles to follow their movements after they fledge. Color markers mayor may not be used to mark the eagles as well. 2e. A sample of sites will be selected and visited to determine the prey species present at the nest sites. This information will be valuable in assessing the prey composition and availability to eagles. 3. Analyze all data obtained and prepare a report of the findings. METHODS AND MATERIALS The 1978 golden eagle Survey was conducted aerially, employing the use of a Cessna 185 in the Northeast, Northwest, and Southwest regions of the state, while the Southeast was surveyed by helicopter. Nests were located with the aid of descriptive materials, maps, and photographs as they were available. Nests were inspected at a distance of 50-150 feet. Approximately 40 nests were inspected from the ground, either due to their proximity to major roads or the inability to view them aerially because of inclement weather or scheduling problems. No specific searches were made to locate new nests, but observations were made both from the ground and in route between nests in the air which led to the discovery of many of the 22 new nests found in 1978. In addition, information from local Wildlife Conservation Officers, ranchers, other raptor researchers, and BLM biologists were instrumental in the location of new sites. Although considerable variability exists in hatching time. an effort was made to observe all nests on record by mid-May to determine the number of eggs or young produced before nest mortality began to take its toll. Active nests are defined as those observed to contain either an incubating adult, eggs, or young birds. Neither the presence of flying adults in the area of a barren nest, nor the presence of fresh nesting material on an unoccupied nest was considered evidence of activity, but did stimulate further search for an occupied alternate site nearby, and note was taken of these conditions. Observations included presence or absense of adults, condition of nest, age of the eaglets, and possible disturbance factors. New nests were plotted on 1:250,000 U.S.G.S. maps, described and photographed to aid in future location, and photographs and descriptive materials for old nests were added when lacking. Those nests which were determined to be active were rechecked in late May and the first 2~ weeks of June, depending on the age of the young birds, to determine the number of eaglets which would fledge. Young golden eagles fledge at 9-10 weeks of age, and birds over 7~-8 weeks were considered to have attained fledging age since mortality in the 8-10 week period is insignificant (Snow, 1973; Baglien, 1975). �160 Following field observations, 2 weeks were spent consolidating the voluminous amounts of data collected since 1973. A new, 10 year observation sheet was designed and implemented into the survey. Also, a county summary was created which in a minimal amount of space describes the chronological history of each nest and offers for yearly comparison, on a county and statewide basis, 16 statistical facets of the survey. RESULTS AND DISCUSSION ~olden Eagle Nesting Result~ The results of the survey are presented in Table 1 below. The 12 nests not checked were neglected due to either time considerations, their isolation, the difficulty reported by former surveyors in their location, or their consistant inactivity over several years due to obvious disturbance factors. Produced young include eggs as well as young birds. Fledged birds are considered to be any birds which survive to 7~-8 weeks of age since mortality in the time remaining before fledging is insignificant (Snow, 1973; Baglien, 1975). The number of young fledged per active nest (Row 11) is a conservative figure in that active nests which were not rechecked for fledging success are not removed from the denominator, making the figure smaller than is actually the case. Twenty-one young, or 8% of the total were unaccounted for in terms of fledging success. The figure is retained in the calculations because it includes those adults which made a nesting attempt but produced no young and thus gives a comparative indication of fertility from year to year. While an active nest is defined as one containing either an incubating adult, eggs, or young, a successful nest is defined as an active nest upon which a count of both produced young and fledging age young have been made. This figure includes nests which have suffered 100% mortality but does not include active nests with no productivity. Comparison of the number of young produced and fledged per successful nest (Rows 14 and 15) therefore gives an accurate indication of mortality from the egg-fledging time period. The acitve nests per nest checked figure (Row 12) compares the adult breeding population status from year to year. Finally, the Reproductive Index is designed to buffer the biases and variability involved in individual catagories, and lends itself well to long-term comparison. It is calculated by multiplying the number of young fledged per successful nest with the percent of total nests checked which were active. This figure therefore involves both the adult breeding population and nest mortality data in establishing statewide reproductive trends of the golden eagle. Results indicate that the Colorado golden eagle population is continuing its upward trend after a low in 1974. In that time period the number of active nests per nests checked has climbed 87%, the number of young produced per active nest has increased 216%, the number of young fledged per successful nest has increased 19%, and the Reproductive Index has risen 118%. Although only 123 nest were checked in 1974, Crowley (1975), using separate data, also documented it as the poorest year for golden eagle reproduction in Colorado since his study began in 1951. He also noted a degree of recovery from the slump in 1975. Comparisons with 1977 figures indicate a more modest increase in eagle reproductive success, exemplified by a 5.1% increase in the reproductive index and virtually identical figures �161 Table 1. Statewide summary of golden eagle nesting productivity in Colorado: 1978 activity and 481 1. Nests on record. 2. Nests not found. 3. Nests not checked. 4. Nests checked. 5. New nests found. 6. Nests inactive 7. Nests active 8. Young produced 9. Young fledged. 10. Young produced 11. Young fledged per active nest. 1.22 12. Active nests per nests checked . . 0.43 13. No. of known successful 14. Young produced 15. Young fledged per successful 16. Reproductive . 34 17 . 430 22 .. . . . 243 . 187 .... 254 228 . per active nest index 159 nests. per successful . . . . . 1.36 nests. nests . .... 1.47 1.43 .61.49 �162 when the number of young fledged per successful nest are compared. This increase is most likely due to the statewide increase in lagomorph numbers, which were at a low in their cycle in 1974 (Donoho, 1979). Donoho's evidence is based on hunter success trends, and he attributes two-thirds of the variation in hunter success to rabbit density and one-third to weather, ground cover, and other factors. Rabbit populations were high in the winter of 1977 and 1978 based on information from Donoho and from numerous ~]CO's in western Colorado. Lagomorphs generally, and specifically black and white-tailed jack rabbits are the major prey item of the Golden Eagle O.,Toodgerd, 1952; McGahan, 1967; Boeker; 1971; Mollhagen et. aI, 1972; Snow, 1973; Seibert et. al., 1976). Plans were begun in late 1978 to initiate a statewide rabbit census to obtain accurate comparisons of eagle and rabbit population fluctuation. Due to the ever-increasing numbers of nests to check, the use of fixed wing air craft in their inspection is of increasing economic value. Hickman (1972) calculated that taking time, salaries, and transportation costs into account, search for Golden Eagle nests from the air was 327% cheaper than corresponding search from the ground, based on linear mile of cliff searched. The completeness and accuracy of aerial counts enhances even more the value of fixed wing air craft in Golden Eagle nesting studies. Although the helicopter allowed a longer and more detailed inspection of nesting sites, the increased time required for travel between nests and the frequent necessity of refueling made it less satisfactory than the Cessna 185 aircraft used by the Division. Restructuring of nest observation forms and establishment of the statistical parameters given in Table 1 was necessitated by the annual geometric increase in the volume of material associated with the survey and the impossibility of obtaining an overview of population trends from year to year. The old observation form, which required one page per year for each nest, was redesigned to allow 10 years of nest observation to be entered on a single page. This new format will be implemented in 1979. Finally the activity, productivity, and fledging success of each nest, organized by county, is now available in a single note book. Each county form contains 8 years of data, and the statewide summary contains 10 years of data. This new format should greatly facilitate the recognition of trends, not only throughout the state, but in the patterns of reproduction exhibited by a single pair of birds over many years. Bald Eagle Nesting Results In 1978, the two bald eagle nests again produced two young each. sites were observed from a distance in order to avoid disturbance might cause site abandonment. The which �163 LITERATURE CITED Baglien, John W. 1975. Biology and habitat requirements of the nesting golden eagle in southwestern Montana. M.S. thesis. Montana State University, Boseman. 53p. Boeker, Erwin L., and Thomas D. Ray. 1971. Golden eagle population in the southwest. Condor 73(4): 463-467. studies Crowley, Lawrence D. 1975. Reproductive trends of golden eagles in Colorado and Wyoming. Unpublished research paper. 36p. Donoho, Harvey. 1979. Personal communication McGahan, J. 1967. Quantified estimates of predation population. J. Wildl. Manage. 31(3): 496-501. Mollhagen, Tony R, R.W. Wiley, and R.L. Packard. golden eagle nests: Texas and New Mexico. 784-792. by a golden eagle 1972. Prey remains in J. Wildl. Manage. 36(3): Seibert, Donald J., R.J. Oakleaf, J.M. Laughlin, and J.L. Page. 1976. Nesting ecology of golden eagles in Elko County, Nevada. Bureau of Land Management, Denver, Colorado. 17p. Snow, Carol. 1973. Golden Eagle (Aquila chryseatos). Habitat management series for unique or endangered species. Report no. 7. Bur. Land Manage., Denver, Colorado 52p. Woodgerd, W. 1952. Food habits of the golden eagle. J. Wildl. Manage. 16(4): 457-459. Prepared by Gerald R. Craig Sr. Wildlife Biologist �164 JOB PROGRESS State of REPORT COLORADO ------------------- Project No. W-124-R Work Plan No. I Job Title Job No. ------ Bald and Golden Eagle ~..j'inter Population Period Covered: Personnel: Raptor Investigations March 15, 1978 through February 2 Surveys 28, 1979 Erwin Boeker, U.S. Fish and Wildlife Service; Gerald Craig, Joe Frothingham, and Wayne Russell, Colorado Division of Wildlife; c. Eugene Knoder, National Audubon Society. ABSTRACT Midwinter bald and golden ea~le flight were conducted on census areas in northeastern and northwesrernColorado as well as the San Luis Valley. The number of wintering golden eagles remained nearly the same on all areas except the northwest which experienced a greater than twofold increase. Bald eagle numbers also increased in the northwest, probably as a result of greater numbers of jackrabbits. �165 BALD AND GOLDEN EAGLE WINTER POPULATION SURVEYS Gerald R. Craig Golden eagle winter population trend information is obtained by annually censusing smaple areas and recording the number of eagles observed per 100 square miles. The censuses yield area estimates which are of use in extrapolating probable eagle populations throughout the region. The aerial census flights are designed to be compatible with similar study areas in other western states. The U.S. Fish and Wildlife Service then compares data from states throughout the region to obtain population information for golden eagles throughout the western United States. Bald eagle population information is obtained by censusing concentration areas, primarily river courses throughout the state. As with golden eagle trend counts, aerial flights are made in midwinter when the population reaches its peak. Rather than an area estimate, all bald eagles sighted are recorded since the census tends to be linear in fashion. The censuses also yield valuable information about those areas which are preferred by wintering bald eagles. P.N. OBJECTIVES The objective of this study is to obtain winter population trend information for bald and golden eagles on selected wintering areas in Colorado. The information will be used to estimate wintering populations of bald and golden eagles throughout the state. SEGMENT OBJECTIVES 1. Aerial Counts of Wintering Golden Eagles: Once annually an aerial flight will be made on census areas in the San Luis Valley, northeastern and northwestern portions of the state. These census areas were established in 1972 and the procedures will be essentially the same. Random transects will be flown throughout each study area with a Cessna 185 aircraft and all eagles observed within ~ mile of each side of the transects will be counted and classified as adult or juvenile. From the transects, an area estimate will be obtained as to eagles per 100 square miles. Identical census. areas are also flown in Wyoming, Idaho, Montana, North Dakota, New Mexico and Nevada by the U.S. Fish and Wildlife Service and cooperating agencies. Information forthcoming from these states are then collected and analyzed by the Fish and Wildlife Service to obtain population estimates for the West. Age ratio information which is obtained provides an indication of the previous breeding season's reproduction. �166 2. Aerial Counts of Wintering Bald Eagles: Since bald eagles tend to congregate primarily along river courses and impoundments, aerial flights will be made along major river courses throughout the state and a direct count will be made of all bald eagles observed. The eagles will be classified as to age in order to obtain information about reproduction. The flights will be made in January when the highest concentration of eagles are usually present. Flights will be made along the South Platte River, Yampa River, White River and Rio Grande River. It is proposed to expand the censuses to the Colorado River and possibly the Gunnison, Dolores and San Juan Rivers. 3. Compile data and prepare annual progress and final reports and submit them to appropriate personnel and agencies. METHODS AND MATERIALS Midwinter golden eagle flights were conducted in the census areas in the San Luis Valley, northeastern. and northwestern Colorado according to procedures described in Segment Objective 1. The northeastern census area constitutes approximately 3,000 square miles in Weld and Logan Counties. North~south transects totaling approximately 600 miles were established randomly throughout the area. Observers count all eagles observed within ~ mile of either side of the aircraft, so this accounts for actual coverage of 300 square miles or 10% of the total census area. The transects are flown at speeds averaging 100-120 mph at altitudes of 100 to 300 feet, depending upon the topography. Census areas in the San Luis Valley and northwest portion of the state are flown in a similar manner. Transects account for a 10% sample of the San Luis Valley. The area encompasses 2,500 square miles which is the majority of the valley lands of the San Luis Valley. The northwestern census area includes Moffat and Rio Blanco Counties and. is the largest with 4,100 square miles. Due to extended flight time, the number of transects were reduced so that 7% of the area is covered. Flights were conducted on the following dates: northeastern Colorado, January 24, 1979; San Luis Valley, February 8, 1979; and northwestern Colorado, February 13, 1979. Winter bald eagle counts are conducted in the San Luis Valley in conjunction with golden eagle flights mentioned above. Since bald eagles are generally distributed ~hroughout the San Luis Valley, it is possible to obtain an effective area estimate of bald eagles during the same flight for golden eagles. Bald eagles are also censused in conjunction with golden eagle flights in northwestern Colorado. A direct count is made of the South Platte River between Fort Morgan and Greeley in Weld County upon completion of the golden eagle flight of northeastern Colorado. The course of the river is flown at altitudes of 100 to 200 feet and all eagles are counted along the river from Fort Morgan to Greeley. RESULTS AND DISCUSSION Results of the aerial flights for wintering golden eagles are summarized in Table 1 and the winter bald eagle flight information is shown in Table 2. �167 The total number of golden eagles wintering in northeastern Colorado dropped to the lowest ever recorded while the population in the northwest increased markedly. It appears that the lagomorph population is on the upswing in Moffat and Rio Blanco counties which attracted more eagles to the area. Golden eagles wintering in the San Luis Valley remained at similar population. In 1978, Dr. James Fitzgerald at the University of Northern Colorado initiated monthly flights in the winter months along the South Platte River between Greeley and Fort Morgan. In order avoid duplication, the Division's flights were discontinued and Dr. Fitzgerald assumed the responsibility of accomplishing the survey. The number of bald eagles frequenting the upland areas of northwest Colorado increased slightly, probably as a result of the jackrabbit population increase in the region. Because of weather and aircraft scheduling difficulties, the major drainages of the western portion of the state were not flown in 1979. �168 Table 1. Golden eagle aerial census of Colorado, 1972-1979 Northeastern Colorado (10% sample of 3,000 sq. mi.) Date Adults Juveniles Unknown 16 19 22 17 18 18 14 8 3 5 3 1 3 4 0 0 0 0 0 0 0 1/24/1973 1/16/1974 1/22/1975 2/19/1976 1/13/1977 1/04/1978 1/24/1979 - - - - - - - - - - - ------ Eagles per 100 sq. mi. - - - Est. of Total Eagles 8.0 7.3 9.0 6.7 6.3 7.0 6.0 240 220 270 200 190 210 180 ------ Northwestern Colorado (7% sample of 4,100 sq. mi) Date 1/25 & 2/26,1972 1/23/1973 1/22 & 1/29,1974 1/29/1975 1/26/1976 1/19/1977 1/19/1978 2/13/1979 - - - - - ------ Adults Juveniles Unknown 86 35 6 11 29 29 23 46 80 14 8 8 3 5 5 25 97 70 47 17 10 2 0 - - - - Eagles per 100 sq .mi. Est. of Total Eagles 91.6 41.5 21.2 12.5 14.6 12.5 9.8 25.8 3,757 1,700 871 514 600 514 400 1,057 3 - - - - - San Luis Valley, Colorado (10% sample of 2,500 sq. mi.) Date 1/29/1976 2/16/1977 1/14/1978 2/08/1979 Adults Juveniles Unknown Eagles per 100 sq .mi. 17 14 10 9 9 3 5 4 2 1 0 0 11.2 7.2 6.0 5.2 Est. of Total Eagles 280 180 150 130 �169 Table 2. Bald eagle aerial census of Colorado, 1972-1979. South Platte River, Colorado (exact count) Date Adults Juveniles Unknown Total % Juveniles 18 1/24/1973 13 31 0 42% 1/16/1974 16 15 31 48% 0 1/22/1975 28 14 0 42 33% 2/19/1976 10 19 0 29 35% ----------------The South Platte River was not flown this year--------------1/4/1978 8 17 68% 0 25 Northwestern Colorado (7% sample of 4,100 sq. mi.) Date Adults 1/25 & 2/26/72 1/23/1973 1/22 & 29/1974 1/29/1975 1/26/1976 1/19/1977 1/19/1978 2/13/1979 *No distinction was -------- - - Juveniles * * *1 * * *0 7 4 1 0 4 0 8 2 made between - - - Total Eagles per 100 sq. mi. Est. of Total Eagles 12.2 500 35 1.4 57 4 71 L7 5 1 14 0.4 11 3.8 157 14 0.4 0 1.4 57 4 143 3.5 10 adults and juveniles on these flights - .- -- ------- - - - .- - - - - - - - - San Luis Valley, Colorado. (10% sample of 2,500 sq. mi.) Date 1/29/1976 2/16/1977 1/14/1978 2/08/1979 Adults Juveniles 28 12 18 31 12 6 8 8 Total Prepared by _-=C"""--",--,->g._;. _ _,.,(?"",,-,,,L -...:=,"""1----Gerald R. ~ Sr. Wildlife Biologist 40 18 26 39 Eagles per 100 sq. mi. 16.0 7.2 10.4 15.6 Est. of Total Eagles 400 180 260 390 �170 JOB PROGRESS REPORT State of COLORADO --~~~~-------------- Project No. Raptor Investigations W-124-R Work Plan No. II Job No. 1 Job Title __~O~s~p_r~e~y~N_e~s~t~1~'n~g~S~tu~d~i~e~s Period Covered: Personnel: March 15, 1978 through February 28, 1979 Gerald Craig, Frank Fiala, Steve Porter, and John Wagner, Colorado Division of Wildlife ABSTRACT Nest occupancy by osprey in 1978 declined to the lowest ever observed. The four active sites were visited during incubation and all contained normal clutches of eggs. Extremely poor production of only 1 young continues to be the rule for the Colorado nesting population. _ �171 OSPREY NESTING INVESTIGATIONS Gerald R. Craig and Frank Fiala Although migrant osprey are regularly sighted in Colorado, only two unsatisfactory nesting records are reported in the literature. In the late 1960's R. A. Ryder reported a pair of osprey nesti~g on Grand Lake in Grand County and another pair were observed breeding successfully at Electra Lake in La Plata County. Since initiation of a breeding survey in 1972, the number of known osprey nests have expanded to 18, most of which are located in Grand,Jackson and Larimer Counties. The present investigation is designed to determine nest site requirements and monitor breeding success. P.N. OBJECTIVES The objectives of this study are: (1) to locate breeding sites and monitor productivity of nesting osprey, (2) to identify nesting habitat requirements and implement measures to encourage occupancy by additional pairs, (3) to compile data and submit reports to appropriate state and federal agencies to assist them in delineating and protecting key nest sites. SEGMENT OBJECTIVES 1a. Continue to locate and map nesting sites of osprey throughout Colorado. lb. All known nest sites will be visited annually to establish reproductive success. All unhatched eggs will be collected and analyzed to ascertain the cause of hatching failure. 2a. Habitat features such as key hunting areas, topography, vegetative type, climate, and geology will be recorded for each nest site in an attempt to establish habitat requirements. 2b. Artificial nests will be constructed in localities which offer potential for occupancy by osprey .. The sites will be monitored annually to determine the success of the endeavor. 3. Analyze all data obtained and prepare a report of the findings. METHODS AND MATERIALS Known nesting sites were observed when possible in late April or early May to determine osprey activity. Sites in which osprey were present were documented and observed for mating behavior. Such behavioral aspects included nest construction or nest rehabilitation, mating . displays and copulation. In late May and early June, all active sites �172 were observed for incubation behavior. During incubation, each active nest was entered to determine clutch size and to obtain measurements of each egg. Nest were again visited in early July to determine hatching success. When possible, addled eggs and egg-shell fragments were collected for thickness measurement and chemical analysis. Those nests in which young were present were banded with Fish and Wildlife Service bands at an appropriate age. RESULTS AND DISCUSSION Only 4 of 18 known osprey nests were active in 1977 and a single young was fledged (Table 1). This represents a significant decline in site occupancy from previous years despite intensified field observation (Table 2). The four active sites were visited to determine clutch size and obtain egg measurements which are presented in Table 3. All four active sites contained clutches of 3 eggs each but all except 1 egg failed to hatch. While nest failure can be blamed upon wind in one case, abandonment of eggs also occurred which may be caused by human harrassment. �173 Table 1. Osprey nesting Site Activity GRI success in Colorado, 1978 Comments Eggs Productivity Active 3 0 GR2 Active 3 1 GR3 Inactive GR4* Inactive GR5 Inactive GR6 Active 3 0 Eggs abandoned GR7* Active 3 0 Eggs missing GR8* Inactive GR9 Inactive GRIO* Inactive JA1 Inactive JA2 Inactive JA3 Inactive JA4 Inactive JA5 Inactive JA6 Inactive LA1 Inactive LA2 Inactive * Artificial nest structures Nest destroyed by wind �174 Table 2. Osprey reproduction in Colorado, 1973-1978 1973 1974 1975 1976 1977 1978 Nesting localities 6 6 12 12 17 18 Active nests 3 6 6 8 12 4 Eggs produced 2 7 1 6 1 12 Young hatched 2 7 0 3 1 1 Young fledged 2 7 0 3 1 1 Table 3. Egg measurements of Colorado osprey, 1978 Length (mm) Site Width (mm) GR-1a 65.2 52.0 GR-1b 64.0 51.8 GR-1c 67.5 49.8 GR-2a 69.0 51.0 GR-2b 70.3 50.8 GR-2c 65.0 51.0 GR-6a 64.0 51.0 GR-6b 62.5 51.5 GR-6c 64.0 56.8 GR-7a 67.0 50.5 GR-7b 66.5 50.2 GR-7c 67.7 47.2 Range 62.5-70.3 Prepared by: Gerald R. craig\ Sr. Wildlife Biologist Range 47.5-56.8 �175 JOB PROGRESS State of REPORT COLORADO Project No. W-124-R Raptor Investigations Work Plan No. III ~--------------------- Job Title ~--------------------------- P_r~a~i~r~i~e~F~a~l~c~o~n~N~e~s~t~in~g~S~t~u~d~i~e~s Period Covered: Personnel: Job No. 1 March 15, 1978 through February Gerald Craig, Brandon Grebence, Colorado Division of Wildlife. _ 28, 1979 James McKinley, and Kent Woodruff, ABSTRACT Prairie located falcon nesting activities throughout Colorado. for 1978 are presented on 6 study areas �176 PRAIRIE FALCON NESTING STUDIES Gerald R. Craig In Colorado, the pra1r1e falcon ranks high in popularity for falconry and demand will undoubtedly increase since this is the only large fa1cQn which can legally be removed from the wild. As harvest pressures increase, it wil1be necessary to monitor the populations to assure that harvest is not detrimental to the well being of the population. Preliminary investigations indicate that the number of occupied nests have remained fairly stable over the past decade. However, detailed investigations are required to establish distribution and productivity on a statewide basis. P. N. OBJECTIVE The objectives of this study are: 1. To document the breeding range and estimate the number of breeding pairs of prairie falcons in Colorado. 2. To obtain production data at selected nesting areas throughout the state and estimate total production of prairie falcons. 3. To delineate nesting habitat requirements of prairie falcons. 4. To document movements and mortality of prairie falcons throughout Colorado. SEGMENT OBJECTIVES 1. Locate and map all known nesting sites of prairie falcons throughout Colorado. From data obtained through approach 3, extrapolate the number of breeding pairs potentially occupying similar habitat types throughout the state. 2. Establish study areas in select habitats which represent important nesting areas. Study areas will be selected that represent shortgrass prairie, foothills and mountain nesting populations and productivity will be determined and'compared between areas. 3. Physical and biological parameters of each nest will be recorded on appropriate field forms and those features which favor occupancy by prairie information will be gathered in conjunction with 4. When nests are visited to determine productivity, the young will be banded with Fish and Wildlife Service lock-on bands and their movements will subsequently be traced through reports filed with the Office of Migratory Bird Management. site which is visited analyzed to establish falcons. The field approach 1. Should the occasion permit, transmitters will be placed on several breeding adults to determine extent of hunting �177 ranges. Radio transmitters will also be placed upon young at fledging and their movements and activities will be monitored. 5. Compile data and prepare annual and final reports. METHODS AND MATERIALS Known nest sites were visited during courtship and early incubation to determine the presence of breeding adults. Invariably all sites could not be visited early in the season and some only received visitation once after the young were produced. Of those sites which were checked, a sample of active nests was selected and an attempt was made to return to each of the sample sites and determine reproductive success. Young were banded with U.S. Fish and Wildlife Service bands when nests were visited to determine brood size, locate prey remains, and record other information about the nest. In addition, the following information was recorded on field forms: elevation, topography, geology of the nest cliff, major vegetative communities, potential hunting areas, distance and direction to disturbance factors, location of nest site, height from nest to top and bottom of the cliff, dimension of the nest ledge, and presence of other raptors in the vicinity. RESULTS AND DISCUSSION In 1978, the state was divided into 6 geographic areas and nesting success was recorded within each area. Area 1 represents the northwest portion of the state; Area 2 is the northeast; Area 3 and 4 account for the east-central portion; Area 5 is the southeast; and Area 6 represents southwestern Colorado. Table 1 summarizes prairie falcon reproduction within, each of the areas. �178 Table 1. Prairie falcon reproductive seccess in Colorado, 1978. Area 1 Area 2 Area 3 Area 4 Area 5 Area 6 Total No. Sites Checked 18 55 34 12 26 17 162 No. Occupied Sites 12 47 25 5 10 13 112 Percent of Sites Occupied 66.7 85.5 73.5 38.5 76.5 Total No. of Pairs 11 43 20 5 9 7 95 No. Successful Pairs 7 33 10 4 6 5 65 Total Young Produced 30 140 40 13 19 13 225 41.7 69.1 Young Produced per Successful Pair 4.29 4.24 4.0 3.25 3.17 2.6 3.92 Young Produced per Total Pair 2.73 3.26 2.0 2.6 2.11 1.86 2.68 Young Produced per Occupied Site 2.5 2.98 1.6 2.6 1.9 1.0 2.28 Total Young Fledged 26 125 29 10 16 11 217 Young Fledged per Successful Pair 3.17 3.79 2.9 2.5 2.67 2.2 3.34 Young Fledged per Total Pair 2.36 2.91 1.45 2.0 1.78 1.57 2.28 Young Fledged per Occupied Site 2.17 2.66 1.16 2.0 1.6 1.25 1.94 Prepared by C. . a.. ~ Gerald.R. Craig Sr. Wildlife Biologist �179 JOB State of Project Work Job PROGRESS Colorado --~~~~~------Raptor No • ..:.:W_--=1.=2...:..4_-R~ _ Plan Title Period· Personnel: REPORT No. III Job No. Investigations II __ --=p~r:..:a:..:i:..:r:...:i:...:e=_.:F:...:a=l=_c=_o:...:n=__=_B.::.r.=e.:::e.:::d.:::i.::n:.!Oig~P_=o~p_=u;.::l:.: ::.s.::.t=i=c=s-=S..=t:.:u;.::d:;:i:..:e:..:s:__ Covered: Gerald Steve March Craig, Platte, 15, 1978 through February 28, and James McKinley, Colorado Brigham Young University. 1979 Division of Wildlife; ABSTRACT Twenty-six marked a breeding pral.rl.e falcons were observed on the study area in 1978. Some marker deterioration was observed which would bias survival estimates. Tape recordings of 26 individuals were made and are to be subjected to sonographic analysis. �180 PRAIRIE FALCON BREEDING POPULATION CHARACTERISTICS STUDIES Gerald R. Craig and Steve Platte Before any management activities or protective measures can be implemented, it is necessary to understand population dynamics of the target species. Prairie falcon life tables have been developed from Fish and Wildlife Service band returns, but the calculated adult lTIortalityis generally believed to be unrealistic due to inconsistencies caused by illegal shooting. Also, a large number of falcons must be banded to yield sufficient returns to permit analysis. An alternative method is being investigated which involves the monitoring of a population of marked prairie falcons. A disjunct prairie falcon population in northeast Colorado will be trapped, banded and color marked at breeding sites, then will be monitored annually to determine the loss of breeding adults from the population. Annual adult mortality can then be determined by direct observation of the sample population. P. N. OBJECTIVE The objective of this study is to document nest site fidelity and replacement of members of breeding pairs of prairie falcons. SEGHENT OBJECTIVES 1. A study area in Northeastern Colorado will be delineated which encompasses approximately 20 pairs of breeding prairie falcons. The area is physically isolated from the nearest breeding pairs by 10 to 15 miles, thus it will be easier to monitor recruitment and replacement of adults within the population. 2. Attempts will be made to trap all adults in the study area, place Fish and Wildlife Service bands and colored leg streamers on them to identify individuals. If some falcons are already banded, their age and banding location will be obtained from banding records. 3. Vocalization of individuals will be recorded and subjected to spectrographic analysis with the intent of distinguishing adults without capturing and handling them. The technique will first be tested upon identified individuals to prove its effectiveness. 4. The study area will be revisited annually and loss of mates or shifts in breeding pairs will be determined by observing color marked individuals, recording and comparing vocalizations and trapping falcons and noting banding numbers. Any unmarked adults will be trapped, banded and color marked. 5. Analyze the data and prepare annual progress reports and a final report. �181 METHODS AND MATERIALS Prairie falcon nests were visited in May and June and attempts were made to capture the adults by using various techniques. Care was taken to avoid prolonged disturbance of the adults while they were incubating eggs. Those adults that were captured were color marked with short nylon streamers on either tarsus. Streamers were placed on alternate legs of pairs which occupied adjacent nests in order to distinguish them should a switch of mates occur. U.S. Fish and Wildlife Service bands were also placed on the other tarsus of captured falcons to provide certain identification. While the birds were in hand various measurements were taken in an effort to develop techniques to distinguish individual falcons. On subsequent visits to the nests, vocalization of each adult was recorded with a portable tape recorder and parabolic microphone. The tape recordings were later analyzed spectrographically in an attempt to determine the feasibility of identifying the vocalizations of individuals. On succeeding years, each site was revisited to determine replacement of adults and possible exchange of mates with other sites. The leg streamers permitted partial identification, but most of the adults had to be recaptured to check band numbers. Previously unmarked adults were marked with streamers and bands as described above. Additional tape recordings were taken on subsequent years to compare vocalizations of the same individuals which were recorded on previous years. Description of Study Area The study area includes the Chalk Bluffs and Pine Bluffs in Weld, Colorado. The habitat is typical short grass prairie interspersed with sandstone outcroppings. A relict population of limber pine is present in scattered locations on to Pine Bluffs and a few cottonwoods are present along intermittent stream courses. RESULTS AND DISCUSSION Breeding adult pra1r1e falcons were marked on either the left or right leg wiLh a black herculite jess secured with two aluminum pop-rivets. Some 26 markers were observed in 1978 (Table 1). Five marked females were retrapped and the deterioration of markers varied considerably. The markers were replaced on three out of 'the five individuals. Some falcons seem to pay no attention to the marker while others appear to continually wear down the soft h.erculite, presumably with pulling force exerted through the mandibles. This material should not be recommended as a leg marker in the future. Marker loss (probably from individuals marked for 2 years) may have biased this years survival estimate downward. It is believed that at least two birds had lost their markers. The 1977-1978 survival sample was divided into two subsamples (birds surviving either one or two years). Survival appears to be higher for birds that have already survived one year (Table 1), thus, survival may be higher with increased age. �182 Two adult falcons were trapped this year that were already banded. One was banded in an area of winter wheat eight miles south of the Chalk Bluffs by Al Harmata on 24 December 1974. He did not indicate the age class (juvenile or adult) of the bird when banded. The bird is at least four years old but will not be used in calculations of average age of breeders. Three more one year old females (two were trapped) were again observed as breeders. One laid four eggs, two pipped, and none hatched. When they are lumped with the seven other young females observed in the last two years (Platt 1977) then 90% of the attempts by juvenile females were successful. There is little doubt that age at first breeding is one year for the prairie falcon in Weld Co., Colorado Tape recordings of 26 individuals were made. Access to a Spectrograph belonging to Dr. D. E. Miller, Department of Zoology, Washington State University, was obtained. There appears to be marked differences in most individuals. The quality of the recording will be the prime limiting factor in analyzing the data. As with finger printing, a minimum number of points of similarity must be established for positive identification. A computer program will be developed to sort before final direct comparisons are made between vocalizations. In addition, the turnover of many taped individuals from year to year has severly limited the number of Control Group (banded, taped and retrapped) comparisons that can be made. Analysis of these data groups will begin in January, 1979. Falcons that are trapped as breeders and were banded as nestlings yield three types of information; fledging location or origin, distance to breeding location, and age. A total of 15 falcons were already banded when trapped. One was banded in winter and will not be considered here. Data for two males is unavailable. This leaves seven females and five males. All of these remaining falcons were fledged in \·]eldCo., Colorado. The closes t that a bird was breeding from the fledging point was 3.0 km and the farthest was 55.0 km. Individuals banded as nestlings and recovered as breeding adults also represent a sample of the age structure of the population (Brower and Zar 1977). The data from each year (subsample) are lumped to obtain an age pyramid (Figure 1). I am also assuming that the left vs. right marking system is valid for indicating identification of individuals. These individuals would have survived one year from the time when trapped and were not necessarily retrapped. One year was added to the last known age and this new age is taken as a data point in the second subsample also. If that bird is again suspected of survival into the third year, it is again taken as a data point in the third subsample. It is then possible, after lumping, for three different ages for the same bird to end up in the pyramid. The effect of following these individuals through time should be negligible if we assume equal mortality between the various age classes of breeder~ As noted in Table 1, this assumption may not be valid. The sample of two yp.ar old �Table 1. Survival of Marked Prairie Falcons During 1976-77 and 1977-78 -. Year Category J 1977-78 Marked in 1976 I 1976-77 1977-78 Marked in 1977 Number Harked 20 41 14 27 Number of Survivals 14 26 10 16 % Survival 70 63* 71** 59** ,_. 00 w * - Marker loss may have biased downward ** - Survival of known prior breeders (probably older) is slighly higher than that of new breeders (probably younger). �184 7 cf ~ (n-16) (n=13) 6 5 ~ >. '-' ...-l til :> '"' Q) ~ ~ 4 H 'Q) M <!! 3 2 1 2S 20 15 5 Percent Figure 1. () 5 10 15 21l 25 of Population Age pyramid of a breedinp. population east Colorado. 1976-78. of the Prairie Falcon in North- �185 females (Figure 1) does not contain this possible bias. It may, however, be a result of fluxuating intensity of banding activity between the years of 1974, 1975, and 1976. The numbers of birds banded in those years have not varied greatly. Thus, this high percentage of two years old female prairie falcons in Weld County, Colorado, may indicate that the breeding population is increasing. The percent of eyries occupied by pairs may have increased from 1976 to 1977 but has then stabilized (Table 2). The percent occupied by one or more falcons has essentially been stable for all three years. The number of successful pairs has increased slightly. Breeders, at a minimum, must produce at least one replacement in their life time. If a population is to maintain itself it must produce enough recruits (replacements) to offset all losses. In the case of the prairie falcon, all eyries are not successful. This means that if the population is to remain stable, the successful breeders must produce enough recruits to replace the mortality of unseccessful breeders. When a population has declined, the successful pairs must also produce replacements to supply unoccupied nesting territories. The historical·data supplied by past workers in the study area and the limited nest ledge availability enable the total number of available territories (49) to be fairly well understood. This allows us to make a valid comparison between the projected minimum necessary fledging success for stability (1.92 fledglings/available territory) and a calculated one for each year (Stable Population Index, Table 2). Thus, fledging success was in excess of the minimum in both 1977 and 1978. The status of the population as indicated by occupancy rates, by stable population index, and by comparison with historical data from 1960-1975 all indicate a stable population. Literature Cited Brower, J.E. and J.R. Zar, 1977. Field and laboratory methods for general ecology. WID. C. Brown Co. Publishers, Dubuque, Iowa, P. 194. Prepared by a. R. C1a;, Gerald R. Craig Sr. Wildlife Biologist �Table 2. Occupancy and Production of the Prairie Falcon In Northeast Colorado In 1976-1978 Category 576 Year 1977 1978 -- Total sites checked 35 45 49 Number of occupied by pairs 22 35 35 % occupied by pairs 63 78 74 Number occupied by single falcons 7 2 4 % occupied by one or more falcons 83 82 83 Number of successful pairs 17 25 26 Total young fledged 57** 91 103 Stable population index* 1.63 2.02 2.10 f-' *Using Enderson's (1969) mortality rates and Henney's (1970) formula (age at first breeding one year. Platt 1977) minimum fledging success should be at least 1.92 if the population is to remain stable. **Some eyries had fledged and total is probably lower than actual value. co a- �187 JOB PROGRESS REPORT State of COLORADO Project No. W-124-R IV Work Plan No. Job Title Job No. Ferruginous Period Covered: Personnel: Raptor March Hawk Nesting Investigations 1 Studies 15, 1978 through February 28, 1979 William Andersen, Otero Junior College; Division of Wildlife. Gerald Craig, Colorado ABSTRACT Ferruginous hawk productivity are compared between three study areas in Colorado. Nests in each of the areas have been enhanced to improve nesting success through a variety of techniques. Approximately 2.23 young were produced per nest attempt on all study areas. Productivity on the southern two areas was similar while the northern area experienced reduced success from disturbance. �188 FERRUGINOUS HAWK NESTING Gerald R. Craig and Hilliam STUDIES C. Andersen Ferruginous hawks are strongly associated with the short grass pralrle community of eastern Colorado, although thev do occur throughout the state in limited numbers. Their populations have undoubtedly declined within the past century as large portions of their habitat have been converted to agricultural lands. Presently, approximately 100 nests have been located, the majority of which are on or adjacent to the Pawneee and Comanche National Grasslands. Ferruginous hawks are extremely sensitive to human encroachment and habitat degradation and additional investigations are underway to establish population trends, tolerance limits and develop techniques to enhance reproduction. P.N. OBJECTIVE The objectives of this study are: 1. To document the breeding eastern Colorado. numbers and productivity of ferruginous 2. To delineate habitat parameters and disturbances ferruginous hawks in eastern Colorado. 3. To evaluate potential techniques to enhance nesting expansion of ferruginous hawk breeding populations. impacting hawks in breeding pairs or encourage SEGMENT OBJECTIVES 1a. Locate, map and photograph all known nest sites of ferruginous the eastern Colorado plains. lb. Known ence prior will 2. hawks on nest sites will be observed from a distance to establish the presof courting or incubating adults. Occupied sites will be revisited to fledging and the young will be counted, banded, and an attempt be made to identify all prey items present. Habitat features such as vegetative type, type of nesting structure, topography, soil type, climate, vicinity of human habitation, roads, and other disturbances will be recorded for those nests identified in 1a and lb. The physical and biological features will be evaluated to establish those parameters which favor successful nesting of ferruginous hawks. 3a. A sample of a predetermined number of nests will be stabilized with wire baskets in an attempt to enhance nesting success. Production of the manipulated nests later will be compared with unaltered sites to determine the effectiveness of the efforts. 3b. Between one and two dozen artificial nest structures will be placed in All breeding suitable habitats which are not occupied by breeding pairs. pairs adjacent to each treatment area will be located prior to placement �189 of the nest structures and will be monitored after placement of the structures to be certain that the structures actually encourage pioneering by new pairs and do not cause relocation of adjacent pairs. 4. Analyze the data and prepare annual progress. reports and a final report. METHODS AND MATERIALS Four areas of primary investigation were established on the eastern Areas 1 and 2 encompassed the southern and northern portions of the National Grasslands, respectively, in southeastern Colorado. Area the eastern and western portions of the Pawnee National Grasslands, tively, in northeastern Colorado. plains. Comanche 3 included respec- Ferruginous hawks have proven to be prone to abandonment if their nests are visited during the courtship and incubation periods, but they will tolerate visitation after the young have hatched. This behavior mandated that the study be carried out in three phases: Phase 1. Nests within all study areas were checked in April and May to determine the presence of courting or incubating adults. Observations were made from distances of at least one half mile (.83 km) from a vehicle and observers remained stationary for a duration of no more than ten seconds. Activity and nest locations were recorded after departure from the area. No other data was taken at the time. Phase 2. Nests were revisited between June 1 and June 25 when the young were old enough to band but not developed enough to attempt to fledge prematurely. At that time, young were banded with U.S. Fish and Wildlife Service lock-on type bands. Sizes 7B and 7D were placed upon the tarsus of each young. Data collected during this visit included: age and general health of young, color phases of young and adults, lining material in nests, and prey items present in and below nests. Pellets present in and below the nests were collected and catalogued for analysis. No habitat data were collected during this phase to reduce the length of stay at each nest to an absolute minimum. Phase 2 began in Area 1 and proceeded sequentially through Areas 2 and 3 since previous studies have shown that nesting activities in the north tend to lag behind southern areas by five to seven days. At the conclusion of Phase 2, all study areas were rechecked for previously unknown nests of renesting pairs. Phase 3. Final visits were made to the areas from early through mid-August after the young had fledged and were no longer in vicinity of the nests. At that time, all the remaining data'we~~collected such as nest site measurements, habitat information, disturbance factors, and maps were drawn. In addition, modifications were made at selected sites to improve their stability or make them more suitable for future occupancy. �190 Modification Designation or management techniques used on nests are as follows: Technique o. Nest was not altered in any way. 1. Top the nest. This required removal of several previous years nest material to reduce the bulk of the nest and thereby improve its stability in strong winds. 2. Remove restricting branches. Tree growth and/or annual addition of nesting material cause obstructions which make it difficult for adults to approach or depart the nest. The restricting branches may also reduce the surface area of some nests. 3. Shift nest center. As additional "pads" of nesting material are added annually, the nest may begin to lean in one direction until it eventually topples over. The nest is realigned over a vertical axis to improve support. 4. Widen nest. As nesting material accumulates, some nests become cone shaped and the nest platform is reduced. In other instances, the supporting tree limbs and trunk did not permit the hawks to construct a wide nest platform. 5. Reinforce base. Wire netting and other materials are used to "shore up" nest bases that have degenerated or lack adequate support for the weight or bulk of the nests. 6. Provide artificial base. In some cases it is best to remove the old nest entirely, construct an artificial base of welded wire and replace a portion of the nest on the new base. 7. Move the nest to new site. Where human disturbance repeatedly causes nesting failure, or the nest tree is an inadequate support structure, extreme action must be taken to move the entire nest to a superior location. 8. Provide predator protection. In extremely rare circumstances, mammalian predators (such as packrats) may endanger the eggs or young. Depending upon the type of predation, different solutions are used. A fairly effective measure is to nail tin sheathing around the tree truck to make it unclimbable. 9. Provide alternate nest. 'This approach is less extreme than 7 and consists of constructing an artificial nest in the vicinity of the existing nest. The adults can then select the superior site, or renest at the new site should their original nest be destroyed. This is generally a good insurance policy for each site where possible. �191 RESULTS AND DISCUSSION Ferruginous hawks frequently construct large, bulky nests in solitary deciduous trees on the pralrle. Generally, the trees are not occupying optimal growing conditions and are stunted. The combination of poor supporting structure and large surface area appear to make many nests susceptable to high winds. Previous investigations tended to bear out this hypothesis, so in 1976 those sites in Area 1 were modified using the above mentioned techniques where necessary to improve nest stability. Nests in Area 2 and 3 received similar treatments in 1977 and 1978. Tables 1, 2 and 3 summarize the nesting success within the 3 study areas and recommends the management activities still required. The young produced per successful nest did not fluctuate significantly between Areas 1 and 2, and the young produced per nest attempt was identical for the two areas. Nesting success was slightly less in Area 1 and young produced per nest attempt was poor due primarily to human disturbance by mineral exploration crews which were drilling test holes in the vicinity of nest sites during the breeding season. �192 Table 1. Site No Ferruginous hawk productivity and nest site management in Area I, 1978 Young Managment Produced Techniques Required FH-14 (CS) 1 0 FH-15 (CS) 2 0 FH-16 (CS) 4 0 FH-17 (CS) 3 0 FH-18 (CS) 1 0 FH-19 (CS) 4 0 FH-20 (CS) 4 0 FH-21 (CS) 4 0 FH-22 (CS) 4 0 FH-23 (CS) 0 9 FH-24 (CS) 4 0 FH-25 (CS) 4 0 FH-26 (CS) 4 0 FH-27 (CS) 0 2 FH-28 (CS) 3 0 FH-29 (CS) 2 9 FH-30 (CS) 3 7 FH-31 (CS) 2 0 Number of active nests: 13 Number of successful nests 11 Percent successful: 85% Young per successful nest: 3.2 Young p~r nest attempt: 2.7 �193 Table 2. Site No Ferruginous hawk productivity and nest site management in Area 2, 1978 Young Management Produced Techniques Required FH-l (eN) 4 6 FH-2 (eN) 0 7 FH-3 (eN) 4 0 FH-4 (eN) 3 0 FH-S (eN) 2 1 FH-6 (eN) 3 1 FH-7 (eN) 3 0 FH-8 (eN) 3 0 FH-9 (eN) 3 0 FH-I0 (eN) 3 0 FH-ll (eN) 0 7 FH-12 (eN) 3 0 FH-13 (eN) 4 0 Number of active nests: 18 Number of successful nests: Percent successful: 89% 16 Young per successful nest: 3.1 Young per nest attempt: 2.7 �194 Table 3. Site No Ferruginous hawk productivity and nest site management in Area 3, 1978 Management Young Produced Technigues Reguired FH-1 (p) FH-2 (p) FH-3 (P) FH-4 (P) FH-5 (P) FH-6 (P) FH-7 (P) FH....:8 (p) FH-9 (P) FH-10 (P) FH-ll (P) FH-12 (p) FH-13 (p) FH-14 (P) FH-15 (P) FH-16 (P) FH-17 (P) FH-18 (p) FH-19 (P) FH-20 (P) FH-21 (P) FH-22 (P) FH-23 (P) FH-24 (P) FH-25 (P) FH-26 (P) FH-27 (P) FH-28 (P) FH-29 (P) 3 0 0 3 2 0 3 0 0 3 0 2 0 0 3 2 3 2 4 0 2 2 0 3 4 0 4 3 2 Number of active nests: 29 Number of successful nests: 18 Percent successful: 62% Prepared by: Gerald R. Craig L Sr. Wildlife Biologist 0 8 9 9 7 0 0 0 7 0 5 8 8 2,4,5 0 0 9 0 8 7 9 0 7 0 0 0 1 0 0 Young per successful nest: 2.8 Young per nest attempt: 1.7 �195 JOB PROGRESS State of COLORADO Project No. W-124-R Work Plan No. Covered: Personnel: Raptor Investigations IV Job Title: ~ulation Periold REPORT March 2 Job No. Surveys of Small Owls 15, 1978 through February Burce Webb, University Division of Wildlife. of Colorado; 28, 1979 Gerald Craig, Colorado ABSTRACT Confirmed breeding status of Flammulated Owls and two breeding attempts by saw-whet owls were documented during the 1978 season in three Colorado latilongs. Spotted owls were not located in their historical localities, giving cause to suspect their year-round residency status in the state. Additional sightings during the 1978 season by the present study contributed to the Colorado Bird Distribution Latilong Study in eight latilong blocks. Breeding densities and reproductive output by the five species was not determinable in 1978 due in part to weather related nest tree destruction and nest abandonment. Snow and windy weather conditions unfavorable to calling or hearing owls hampered surveys, and limited effective time in the field to 65 days. Cooperator use of pre-recorded owl calls on cassette tapes in 1979 should produce further nesting information for these species. When breeding status becomes documented, various physical and biological habitat parameters can be identified, and management recommendations can flow. �196 POPULATION SURVEYS OF SMALL OWLS Gerald R. Craig and Bruce Webb Virtually nothing is known about the distribution and population status of the forest-dwelling owls in Colorado. The present study was undertaken to gather distribution, habitat and density information on five species of owl found in Colorado: flammulated (Otus flammeolus), pygmy (Glaucidium gnoma), spotted (Strix occidentalis), boreal (Aegolius funereus), and saw-whetowl (A. acadicus). In their "Red Book," the United States Fish and Wildlife Service lists the spotted owl as threatened in part of its range (Gould, 1974). Intermittent observations in Colorado of the other study species indicate that they are at least uncommon and may be quite rare. On the other hand, a sufficient number of breeding adults and young of all but the spotted owl have been reported, thereby eliminating the possibility that they are only rare stragglers to the state. Recent surveys to establish the distribution of spotted owls have been conducted in California (Gould 1974, 1977), Oregon (Forsman 1975), and Utah (Boner, pers. comm.). It is essential to gather distribution and density information about owls because the recent studies in California and Oregon have shown that the spotted owl may be suffering from habitat loss. The destruction of suitable habitat coupled with an almost total lack of ecological and population data gives cause for serious concern about the future of these species in Colorado. In addition, increasing use of public lands for outdoor recreation and land development introduce other tolerable stress factors into their environment. Proper planning and wise management of public lands can help to avert these problems. P.N. The objectives OBJECTIVE of this study are: 1. To determine statewide distributions saw-whet and spotted owls. 2. To delineate important breeding habitats pygmy, saw-whet and spotted owls. 3. Where applicable, to obtain production of flammulated, boreal, of flammulated, pygmy, boreal, data at selected nesting areas. SEGMENT OBJECTIVES 1. Initiate surveys within the study areas to determine presence and abundance of the four owl species. Previously it has been demonstrated that most owls will respond to recorded vocalization. Therefore, one technique will be to play recordings of all four species in suitable habitats to establish the presence of owls. Mist nets and bal-chatri traps may also be employed to locate owls if the recordings are not effective. �197 2. Distribute tape recordings of the four owl species vocalizations to appropriate field personnel with the Division of Wildlife, National Park Service, Bureau of Land Management, Forest Service, and various bird clubs. The participants will be encouraged to play the recordings in an effort to locate the owls. Reports from field personnel will be confirmed by the investigator. 3. When the presence of owls are documented, various physical and biological features will be recorded such as climate, elevation, topography, slope, aspect, vegetative type, and probable prey species. It is hoped that general habitat parameters can be developed for each species. 4. When owls are located through 1b and 1c, initiate surveys during the breeding season to discover their nests. When nests are located, they will be visited to determine productivity and pertinent habitat features will be recorded. 5. Analyze the data and prepare a final report including statewide distribution of the four owl species. METHODS maps delineating AND MATERIALS Field surveys to locate flammulated, northern pygmy, spotted, boreal and saw-whet owls were conducted at various wooded montane areas from 3 March through 12 August 1978. Using United States Geological Survey topographic maps, suitable habitat was selected primarily on the basis of proximity to heavily wooded areas with minimal human settlement. Isolated vehicle roadways through selected habitat were located on United States Forest Service maps. During the day, additional habitats were surveyed for their suitability. Additionally, historical localities of spotted owl (Table 3) were selected for intensive surveys work in an attempt ,to identify suitable habitat and establish breeding site tenacity. Many historical breeding localities for flammulated, boreal and saw-whet owls were also surveyed. Owls were located more frequently by vocal imitations of species specific courtship songs, a technique that has previously proven effective (pers. ob.). Pre-recorded cassette tapes also were played according to a standardized census technique (Appendix 1). In most cases, calling was done at 0.5 mi. intervals along selected roads through suitable habitable. When a second observer was present, continuous surveys were conducted by alternating between driving and walking by each observer. A request for sightings form was circulated in the statewide Field Ornithologist's Journal (Appendix 2). Colorado RESULTS AND DISCUSSION Considering the more apparent diurnal habits avifauna, the supposed rarity of many montane of most of the montane owls is at least partially �TABLE I Present Survey Records of Flammulated Owl in Colorado Date 2 Locality Number County Latilong Source June 28, 1978 (I) heard 10 mi. North of Hwy, 160 on Road 235 Archuleta 23 B. Webb July 9, 1978 (2) observed Carson Hole, Uncompahgre Plateau Mesa 15 B. Webb at nest TABLE 2 Present Survey Records of Pygmy Owl in Colorado Date April 9, 1978 Locality Number (1) heard Queen's Canyon f-' <o County EI Paso Latilong 19 Source B. Webb 00 �TABLE 3 Historical Records of the Spotted Owl in Colorado Date Number Locality County Latilong Source June or July, 1873 (I) observed in Deadman's Canyon EI Paso Block 19 Aiken & Warren 1914 June 1873 ( I) specimen CC 6074 not given El Paso prob. 19 Bailey & Nicdrach 1965 *3 1886 (2) observed In Boulder Valley (by Gale) Boulder Block 4 Henderson 1909 4 Nov·?·1903 ( 1) specimen not given specifically Pitkin Block 9/10 B & N 1965 5 1·1·1906 (2) observed in vicinity of Ft. Lewis La Plata Block 22/23 Gilman 1905 6 ?-?-1907 ( I ) observed Not given spcciflcally Costilla Block 25 Sclatcr 1912 7 May 24, 1919 ( I) specimen Collected in Queen's Canyon ncar Colorado Springs EI Paso DIock 19 Bailey S: Niedrach 1965 8 Sept. I, 1941 (1) young taken about 40 mi. from Hartsel; bird was seen in animal exhibit *9 Jan.21, 1956 (1) one seen Near Loveland Larimer Block 4 *10 Feb.6,1956 (1) one seen At Platteville Weld Block 5 same 11 Dec.31,1958 (1) specimen DMNH 34437 Collected in Lakewood (3035 Grove St.) Jefferson Block II same 12 July 22,1961 (1) one seen At Red Rocks Park near Morrison Jefferson Block II samc 13 Aug.17,1962 (1) one seen In Upper Buckhorn Canyon Larimer Block 4 same 14 May 18,1963 (1) one seen Fort Collins area on Spring Count Larimer Block 4 samc June 3-5,1975 (J) observed Rocky Mountain Arsenal Adams Block 12 DMNH-CFO filcs Sept. 5, 1975 (1) observed 1 mile west of Silverthorne Summit Block 10 DMNH-CFO files Oct. 24, 1977 (1) observed At Ridgeway Ouray Block 16 CFO Journ. 34:20 2 15 16 17 *Reported as a Barred Owl Gadd 1942 Bailey & Niedrach 1965 •....• \0 \0 �TABLE 4 Records of Boreal Owl in Colorado (Historical and Present Survey) Locality Dall' Number Oct. 14, 1896 ( 1) specimen Crested Butte Gunnison 2 November, 1903 (I) specimen Not given Pitkin 9 or 10 same 3 Nov. 11, 1929 (1) specimen Fraser Grand ]J same 4 Aug. 14, 1963 (1) juvenal 1 mi. south of Deadman Mt. Lookout Larimer 4 same 5 Apr 1, 1970 (1) specimen 3.5 mi. South of Estes Park Larimer 4 CFO 6 Aug. 31, 1971 (1) juvenal observed 1.5 mi. northeast Rocky Mountain Biological Laboratory Gunnison 17 CFO, No. 12, p. 14 W. Calder, pers.comm. 7 June IS, 1973 ( I) specimen West slope of Rabbit Ears Pass Routt 3 G. 8 July 13,1974 (1) observed 5.0 mi. north of Chambers Lake on Larimie Road Larimer 4 CFO Report 9 Dec. 2,1977 (1) observed 4.5 mi. South of Grand Lake Grand 4 Denver Museum of Natural History, CFO Records me. 10 Feb. 2, 1978 (1) specimen DMNH Evergreen Jefferson 11 DMNH-CFO Files II Feb. 6, 1978 ( I) specimen CSU Estes Park Larimer 4 DMNH-CFO Files 12 July 15,1978 (1) observed-adult 0.5 mi. northeast Deadman Lookout Larimer 4 B. Webb (present survey) County Latilong 17 Source Bailey & Niedrach 1965 No. 9, p. R. Craig, comm. 25 pers. N a a �TABLE 5 Present Survey Records of Saw-whet Owl in Colorado 1978 Season Number Locality County Latilong Source April 25 (2) captured banded Castlewood, 1 mi. west of Franktown Douglas 12 B. Webb 2 April 26 (2) heard 0.5 mi. west of Alamosa Ranger Station San Juan National Forest Conejos 24 Same 3 April 26 (2) heard 0.5 mi. west of Alamosa Ranger Station San Juan National Forest Conejos 24 Same 4 May 6 (2) observed (nest hole) 1 mile south of Red Feathers Lake Larimer 4 Same 5 June 9 (J) heard I mile south of Deadman's Lookout Larimer 4 Same 6 June 13 (I) heard Golden Gate Canyon State Park South Entrance Jefferson 11 Same 7 June 28 (1) heard 2 mi. north of Hwy, 160 on Rd. 630 Archuleta 23 Same 8 July 21 (1) observed Juv. 0.5 mi. South of summit Rabbit Ears Pass West Summit Routt 3 Same 9 July 22 (1) heard 3.0 mi. South of Hwy. 40 on Buffalo Pass Road, Routt National Forest Grand 3 Same N 0 •.... �202 behavioral. However, the development of techniques for finding certain species of owls has increased the number of records. The use of prerecorded tapes prepared in 1978 for cooperating state personnel will be of great potential in 1979. The following summarizes the results of the 1978 survey and literature search. Historical records of flammulated, pygmy and saw-whet are too numerous to include in tabular form. Flammulated Owl Flammulated owls were found on two surveys, and the other in block 15 (Table 1). one in latilong block 23 On 28 June, one flamrnulated owl was heard calling in an extensive aspen grove in the San Juan National Forest. In block 15, a nest was found on 9 July in a hole in an aspen tree on the Uncompahgre Plateau, Uncompahgre National Forest. Noises from young birds in the hole were heard and two adults were observed carrying food. In a one-hour period, the adults fed moths (length = 3-5 cm) to the young on 27 occasions. A colleague working on flammulated owls in the Gunnison floristic basin in 1978 located 15 birds. In Colorado, the flammulated owl has been found nesting in old woodpecker holes in aspens and coniferous forests (Bailey and Niedrach, 1965). Bailey and Niedrach summarized 17 records of flammulated owls prior to 1900, but only 7 between 1900 and 1965. They pointed out that this probably represents a decrease in activity of egg collectors rather than decreases in the population of owls. In California, Winter (1974) determined that the flammulated owl's statewide distribution corresponded to the distribution of yellow pine stands (Pinus ponderosa and~. jeffrey:). Further Colorado survey work in 1979 may show a flammulated owl distribution that corresponds with aspen stands, rather than coniferous forests. Northern Pygmy Owl Only one pygmy owl was found during the survey, on 9 April in latilong block 19, approximately 10 miles west of Colorado Springs in Queen's Canyon (Table 2). On 22 May 1977, one adult was observed during the day in an aspen hole on the Uncompahgre Plateau. In 1978, that hole was occupied by nesting flammulated owls. The pygmy owl generally is though of as breeding from lower montane through upper montane in Colorado, up to 12,000 feet in elevation (Bailey and Niedrach 1965). The numerous Colorado reports of pygmy owls involve sightings during late fall through winter,whenbirds seek lower elevations. Often they are found near backyard bird feeders, but also frequent wooded ravines with low bushy vegetation (pers. obs.). The coexistence of pygmy and flammulated owls in aspen-coniferous forests wuld not involve competition for food. The flammulated owl is insectivorous and nocturnal; whereas, the pygmy owl feeds on small birds and rodents, and has both diurnal and nocturnal feeding habits. �203 Also, on 15 August 1977, one pygmy owl was observed in Pinyon-Juniper Woodland habitat at Lee Gulch, southwest of Meeker, Colorado. PinyonJuniper habitat is not generally recognized as suitable nesting habitat for this species. The mid-August record in this vegetation type may have been due to post-breeding wandering downslope. Intensive bird survey work every week prior to the mid-August record did not reveal the presence of pygmy owls there. Spotted Owl Historically there are four Colorado reports of spotted owls during the May through August breeding season which had specific localities described (Table 3, Nos. I, 2, 7, 12-13). Each of these possible breeding localities was intensively surveyed on foot using the standard nighttime survey method. In addition, daytime searches were conducted to look for possible roost sites in locations with canyon walls. In each of these historical localities, no spotted owls were found. It is interesting to note that five Colorado reports of spotted owls (No.3, 9, 10, 15) involve owls on the plains, away from suitable nesting habitat. This type of "vagrancy" is usually associated with migratory birds. It is possible that spotted owls are not a permanent Colorado resident in the northern part of their range. It may move south to winter, and return north in spring to nest in favorable habitat. Strix occidentalis caurina generally is found assiciated with the presence of woodrats (Neotoma ~.). Pellet analysis in New Mexico, Arizona and Utah have shown this to be the primary prey species (Ligon, 1926; Marshall, 1942; Kertell, unpub. ms.). In Colorado, there are several woodrat species found in montane regions. During daylight surveys through Queen's Canyon and adjacent Williams Canyon, woodrat workings were not uncommon on the rock faces. Nearby in eastern Utah, at Canyonlands National Monument, the Spotted Owl is found as a breeding species in the moist, cool, canyon bottoms (pers, Comm. T.C. Wylie). In parts of Western Colorado, Colorado National Monument and Mesa Verde National Park in particular are structurally similar to Canyonlands topography. The many deep canyons at Colorado National March. No spotted owls were found. Monument were surveyed in Personnel at Mesa Verde National Park list the spotted owl without specific records, as a permanent resident of the Park's canyon country. Daytime and night surveys in late June and early July did not reveal any spotted owls. Boreal Owl One adult boreal owl was observed on 15 July 1978, 0.5 mi. northwest of Deadman's Lookout at 10,600 feet elevation in Larimer County (Table 4). The characteristic stand-type of this location was dense spruce-fir forest with extensive fallen timber in the understory. An extensive, unseccessful daylight two day search was made to locate its nest hole, roost sites or already-emerged juveniles. On 9 August, the Virginia �204 Mine site near Crested Butte was transversed on foot with Dr. Calder, who had observed a boreal owl there in 1971. No surveys in that or other sites in the vicinity of high mountain passes were successful. These included: Berthoud, Guanella, Hoosier, Loveland, Molas Divide, Monarch, Mt. Evans, Rabbit Ears, Red Hill and Wolf Creek. Baldwin and Koplin (1966) found recently-fledged Boreal Owls in the vicinity of Deadman's Lookout. Based on this and other distributional records, they proposed that the boreal owl exists as a Pleistocene relict in high mountain portions of the state. Due to late lying snow at the higher elevations, few observers reach suitable nesting habitat during the vocal courtship phase of the species nesting cycle. This probably also accounts for the few numbers of breeding records. Saw-whet Owl A literature search showed numerous records throughout (Bailey and Niedrach, 1965). As such only new records survey are provided in this report. the state from the present Eleven individuals were recorded at nine localities in 1978 (Table 5). At the Castlewood Canyon and Red Feathers Lakes localities, nest holes were located, and subsequent nest activities were monitored. In both instances, courtship activity by both adults around the nest tree (both Ponderosa Pines) was noted for two days. A subsequent check at Castlewood Canyon four weeks later disclosed that the dead nest tree had been felled under the weight of the heavy snowstorm of 4-5 May. Both adults, banded on 25 April, were not found there after in the area. Also, no eggs or young were found in the nest hole of the fallen tree. At Red Feathers Lakes, a subsequent check three weeks later revealed that the nest hole had been abandoned; no owls were seen in the area. At both nest sites, no regurgitated pellets could be found. The elevational range of these sightings go from the lowest extent of Ponderosa Pine at Castlewood Canyon near Franktown, Douglas County, to a lone owl in Juvenal Plummage observed at 9,426 feet in Spruce-Fir vegetation. This species was most frequently heard calling spontaneously at dusk. Of the five species investigated, this one responded most readily to the survey methods. LITERATURE CITED Aiken, C. E. And E. R. Warren. 1914. The birds of El Paso County, Colorado College Science Series. 12:455-603. Bailey, A.M., and R. J. Niedrach, 1965. Birds of Colorado, Denver Museum of Natural History, 454p. Vol, Colorado. 1, Denver, Baldwin, P.H., and J.R. Koplin, 1966. The Boreal Owl as a Pleistocene relict in Colorado. Condor 68: 299-300. Colorado Field Ornithologists, Denver. ORC Files, Denver Museum of Natural History, Colorado Field Ornithologists. 1978. Colorado bird distribution latilong study. H.E. Kingery and W.D. Graul, eds. Colorado Division of Wildlife, Denver. �205 Gadd, S.W. 1942. Spotted Owl nesting Gilman, M.F. 1907. 194-195. in Colorado. Some birds of southwest Condor 44: Colorado. 35. Condor 9: 152-158, Gould, G., 1974. The Status of the Spotted Owl in California. Unpubl. report, California Department of Fish and Game. 36+xxp. (multilith) Gould, G., 1977. Distribution Western Birds, 8:131-146. of the Spotted Owl in California. Henderson, J. 1909. An annotated list of the birds of Boulder County, Colorado. University of Colorado Studies 6:219-242. Forsman, E. 1975 A Preliminary investigation of the Spotted Owl in Oregon. Unpubl. Master's Thesis, Oregon State University, Corvallis. 127 p. Ligon, J.S., 1926. Marshall, 44: Habits of the Spotted Owl Auk 43: J.T., Jr., 1942. 66-67. Food and habitat Sclater, W.H. 1912. A history Co., London, 576 pp. by: of the Spooted Owl. Condor of the birds of Colorado. Winter, J., 1974. The distribution Western Birds 5: 25-44. Prepared 421-429. C R. ~ c Gerald R. Craig ( Sr. Wildlife Biologist of the Flammulated Witherby and Owl in California. �206 MONTANE OWL SURVEY Appendix 1. 1. We ask that you keep a written record of the date, location, time and results for every time you play the tape. A brief discription of the type of habitat covered in the survey is also important. For this project, we are interested in the number of negative responses as well as the number of times you here owls. 2. We recommend two standardized methods for censusing owls. Using any portable tape casette palyer, at nearly full volume, play the full series of calls for the four species. First, if surveying along a road, make survey stops everyone-half mile and play the working series of owl calls (Side 2 of tape). Second, if walking along trails or roads, play calls once about every 100 yards. This "continuous" survey provides better coverage than the "stop" method and is better suited for working an area only accessible by trail. The "stop" method provides extensive covereage in a shorter time period. 3. Spend a few minutes listening for spontaneously calling owls before and after playing the owl calls of each species. 4. The calls of the four species are located on side two of the tape. The order of occurrance is: Pygmy, Saw-whet, Spotted and Flammulated Owl. 5. When using the tape casette player, it works best to set the player down and walk some distance away to escape the distraction of the hiss and static. This also enables you to better hear any elicited response in the distance. One can often hear owls calling back to the tape from nearly a mile away under excellent conditions. Set the player at maximum volume for maximum range. 6. The tape can be shut off temporarily in mid-tape if you want to try and pinpoint the direction and distance of owls calling in the distance. If an owl responds to the tape you can alternately approach the bird and play the tape. This often brings them to within flashlight range for visual confirmation. 7. The four species of owls are found in coniferous forests. For Spotted Owls, select a relatively quite stopping point such as along a ridge top or in a box canyon. When in the vicinity of flowing streams, hike upslope to get away from the noise. Saw-whet, Pygmy and Flammulated Owls can be found in pine, fir and spruce forests. 8. For any calling night birds which you cannot identify from the reference series of calls provided on side one of the tape, make a written description of their voice on the survey sheet. 9. Notify us as soon as possible when you get a positive response by an owl. Likewise, when you find a nest of any montane species 'of owl, contact us by mail at the address below. Gerald R. Craig Raptor Management Specials it Colorado Division of Wildife 317 W. Prospect St. Ft. Collins, CO 80526 �207 NOTICES FLAMMULATED, PYGMY, SPOTTED, Appendix BOREAL AND SAW-WHET 2. Oh~S The distribution and status of certain of Colorado's montane owls are poorly understood. The University of Colorado in cooperation with the Nongame Program of the Colorado Division of Wildlife is initiating a study to determine the population status and habitat requirements of Flammulated, Northern Pygmy, Spotted, Boreal and Saw-whet owls in montane regions of Colorado. By soliciting cooperator sightings from C.F.O. members and other birders we can bring together scattered sightings to formulate a clearer picture of the status of these owls in the state. Individuals wishing to cooperate in this project are asked to complete the Cooperator Sightings Report Form inserted in this issue and forward it to me at the addresss below. Frequently owls are found killed on the road. Such specimens can be a particularly valuable source of owl diet information if cooperators choose to retrieve the specimen, wrap it, and freeze it for later pick-up and analysis. These specimens can later be examined for stomach contents, reproductive state, and finally can be deposited in museums as prepared study skins. An owl collected in this manner should include date and locality of discovery. Pick-up of the frozen specimen will be prompt once I'm notified. The success of this project will depend largely on the cooperation individuals throughout the state. of -Bruce E. WebbDept. of E.P.O. Biology University of Colorado Boulder, Colorado 80309 OWL INFORMATION COOPERATOR SIGHTINGS REQUEST REPORT FORM The University of Colorado in cooperation with the Non-game Program of the Colorado Division of Wildlife is initiating a study to determine, the population status and habitat requirements of Flammulated, Northern Pygmy, Spotted, Boreal and Saw-whet owls in montane regions of Colorado. If you have seen or heard any of these species has seen or heard of these species (including cooperation is greatly needed. Please answer and return this sheet with your name, address will contact you. Species observed: Dates(s): Locality (as specifically as possible): or know of anyone who road-killed owls), your the three questions below and telephone number and I Your name: Address: Telephone: � https://cpw.cvlcollections.org/files/original/45c8ee6d8203c72057ab381363feefd1.pdf daf884c264ebcf8cec632e4a39f2035c PDF Text Text April 1 JOB PROGRESS State of Project Work 1980 REPORT Colorado No. Game Bird W-37-R-33 Plan No. 1 Survey 22 Job No. Job Ti t Le __ E_v'-a;_l_;cu=-..:.:.a..:.t.....;i..:.o-'--n-'---'--o_f__:N_;_e_s-'-t.;.__i;_n .•..• g__:C_o--'v_e_;r;;...._;P;;_r;:;.._::_e.=f-=e-=r_;e...::nc;_;c;;_e __ in Relation Period Covered: Personnel: April to Wheat Farming 1, 1979 through H. Leroy Haeffner, Warren Snyder Kirk Methods March Snyder, 31, 1980 Jack Corey and ABSTRACT Near average precipitation amounts were received in 1979, and this moisture was well distributed and apparently well suited for sustained nesting efforts through the spring and summer. However, spring stubble plowing was delayed and late due to wet field conditions and green wheat growth was also late with rather sparse open stands and parts of many fields destroyed. Wheat harvest, likewise, was delayed due to late ripening of wheat and cool damp harvest weather. Pheasant hens trapped on and adjacent to the Sand Draw Property in early March, 1979, and equipped with solar-powered transmitters, sustained considerable predation beginning in midApril, primarily from great-horned owls. Hens tended to disperse away from the Sand Draw Property, possibly in part, due to this Monitoring of hen activities showed that p r'edat o r harrassment. nearly, if not all, first nesting attempts were in wheat stubble. Nest predators destroyed approximately half of the known nest sites whereas stubble tillage destroyed the remainder, so that no nests hatched in residual wheat stubble. Hens moved to green wheat to renest and most were successful on their first renest attempt there. Two hens were flushed by wheat harvest ,(combining) activities. One returned to terminate her nest successfully whereas the other abandoned and subsequently renested to successfully bring off a brood on her fourth known nesting attempt in late August. Two other hens were successful on their third nesting attempts. Information on nest placement, clutch size, recycle time, movements of hens after nest predation, and other variables were obtained. Hens with broods tended to remain in wheat until it was harvested. They then moved to roadsides and then to green row crops and weedy locations. �2 No known nesting efforts were located in chemical fallowed wheat stubble preventing direct evaluation of the impact of this farming approach on pheasant production. However, if all stubble had been chemically treated and initial tillage had been delayed until late June, all nests not predator destroyed would have hatched successfully. Question arises as to how many second nests would be placed in stubble at what date, and how many would be placed in green wheat when wheat stubble was still available. Thirty-four hens were trapped and equipped with solar-powered transmitters in mid-March, 1980. A late March snow storm, with snow depths accumulating to approximately 20 inches, apparently did not cause direct hen mortality, but did expose the hens to increased avian predation. The spring, 1980 breeding population was observably much higher than in 1979. �3 EVALUATION OF NESTING COVER PREFERENCES OF PHEASANTS IN RELATION TO ~!HEAT FARMING METHODS Warren D. Snyder P. N. OBJECTIVE 1. To document the relative importance of wheat stubble, green wheat, and other vegetative cover for (1) pheasant nest site selection, and (2) successful production of young in the wheatlands of northeast Colorado. Other variables pertinent to understanding the basic ecology of the nesting pheasant in the Tablelands include determination of primary limiting factors to reproduction and brood survival. 2. Upon documentation that pheasants use wheat stubble extensively for initial spring nesting and that adequate sample sizes can be obtained in the primary study area, a second primary objective will be to determine if mini-till summer fallow, using combinations of post harvest applied herbicides and sweep tillage, increases pheasant nesting success when compared to conventional summer fallow methods. The economic and practical aspects of mini-till farming will also be determined as they apply to the farmer and to Division of Wildlife properties. SEGMENT OBJECTIVES 1. Design plot layout and apply treatments. 2. Nonitoring, analysis and evaluation. METHODS AND ¥ATERIALS Reference is made to Snyder (1979) for a sunrnary of preliminary methods and materials. Modifications or supplements to this are presented in the following sections. 1. Plot Design and Application of Treatments The entire stubble acreage on the Sand Draw property was treated with a 2,4-D and atrazine herbicide mixture during the first week of August, 1979. This change from initial design was necessitated because the private land, designated for treatment by the lessee, was located 1 mi. north of the study area and was considered too distant from the study area. It vJaS also hailed to a degree on June 10, 1979. Appropriate controls will be selected proximal to the study area. �4 2. Analysis and Evaluation Environmental monitoring a. Green wheat height was measured at 10 day intervals starting on approximately Hay 1, and continuing to early June. Both average height and a H-D (height-density) measurement, using procedures previously described for sampling residual cover, were obtained for 50 samples from each selected field. b. Stubble plowing progress was surveyed at weekly intervals on an established route running from just west of Holyoke, north and east to the study area and then back toward Holyoke. Over 80 fields were included in the sample. Stubble plowing progress was also recorded for all stubble fields within the 9-section study area. The sampled route and area are illustrated in Fig. 1. c. Dates of various field activities, first tillage of the mini-tilled land, and wheat harvest progress were recorded. d. Precipitation was recorded at the east end of the study area through the growing season. Fall-winter data were obtained at a location 1~ mi. north and 1 mi. west of the Sand Draw Property and from Holyoke and Julesburg weather station records. e. Grasshopper densities in various cover types were obtained by obtaining ocular 1 sq. ft. samples at 1 pace intervals while walking. Nine samples were combined to project an index of density per sq. yd. and a 25 sq. yd. sample was obtained from each cover type. f. The relative occurrence of green vegetation in treated and untreated wheat stubble fields was measured in mid Hay, 1979. Samples were tallied by recording hits and misses of green vegetation on a one inch diameter hoop placed approximately one foot in front of the right foot. Samples were taken at one pace intervals while walking somewhat at randolllthrough the fields. One hundred samples we re collected per field. g. Wheat straw, old weeds and other organic litter was removed from square foot samples on'both conventionally summer fallowed and sweep+t Ll l.ed fields in the study area in November, 1979. A square foot hoop was thrown to obtain a random sample. A total of 35 samples, bagged, air dried, and weighed to the nearest 0.1 gram, was obtained from several conventionally farmed fields, and a sample of 23 was obtained from sweep tilled sites. �5 .. ":: "" =====~;:::.:.=== " . "-'<.':..' 'P- i!! rii ._ ~~.:._ I" ><-.~_~.:. I\"'~'f •....•••• ~~.. /I /I rl •• II _h,,--- II ".,'f" • II . . " . rll :: " " ...~~,-," ~r- ._~./-I...JI. _ .>..._ LEGEND ~ • • • •• Figure 1. Stubble Plowing Survey Route Sand Draw Nine Section Study Area The transect route used to measure progress of wheat stubble plowing and the nine-section Sand Draw Study area. ~•• �6 RESULTS AND DISCUSSION Environmental Measurements Precipitation Approximately 11.79 in. of precipitation was received from April 1 through August 31, 1979 on the Sand Draw. This moisture was well distributed as noted in Table 1, however, much of the summer rainfall came as light showers which were not highly effective in building subsoil moisture. Moisture received in 1979 was considerably above that obtained during the preceding year, which was one of the drier years on record. Vegetative Height and Density Height and height-density index (Kirsch 1978) were obtained on wheat stubble and grass in March and April, 1979, and on green wheat in May and early June. Averages are summarized and compared in Table 2. Results show the mixed grass cover on the property with an H-D index of 0.57 (measured in decimeters) was poor. Wheat stubble provided a much higher H-D index and both the H-D index and height-only measurements were slightly better on the Sand Draw fields than in adjacent stubble fields. These heights were above average when compared with stubble measurements obtained from 1963 through 1968 (Table 2). Green wheat made slow growth in 1979 and did not provide H-D indices and height-only measurements comparable to those for wheat stubble until mid to late May (Table 2 and Fig. 2). Figure 3 compares 1979 wheat growth on the property and in adjacent fields with growth plotted in the region during the 1960's. The 1979 growth lagged behind that for most other years. Wheat growth was severely stunted on the property by a dense competitive stand of annual brome (cheatgrass) which sprouted in late winter. This resulted in very poor wheat on the property compared to that on adjacent areas, and some bias resulted. It also resulted in almost no standing stubble remaining in late winter 1980. Cheatgrass occurrence on the Sand Draw fields occurred primarily because Division contracts in previous years required the lessee to not initiate plowing of wheat stubble fields until after June 20. This permitted cheatgrass to mature and establish an abundant seed source in the soil. Adjacent tree rows also contain a cheatgrass understory which spreads to a degree into the adjacent fields. �7 Table 1. Precipitation recorded on and near the Sand Draw study area in 1978 and 1979.11 Year Month 1978 1979 Long-term Mean January 0.45 0.64 0.31 February 0.43 0.09 0.29 March 0.07 2.14 0.86 April 0.74 1.45 1.53 May 2.78 2.44 3.29 June 1.37 3.81 3.83 July 1.63 2.32 2.82 August 1.25 1.77 1.88 September 0.11 0.65 1.64 October 0.50 1.00 1.13 November 0.43 1.60 0.46 December 0.75 10.51 in. T 17.91 in. 0.36 18.40 in. II - Data from 1978 through March, 1979 and long-term mean information was obtained from the U.S. Weather Service Records for Holyoke. �8 Table 2. Comparative height and H-D indices on the Sand Draw and adjacent areas for perennial grass, wheat stubble and green wheat in spring, 1979. 1/ H e1g . h t-1/ Location Cover type Date Sand Draw Mixed grass Mid-March 0.57 No Data Sand Draw Wheat stubble Mar.-April 1.43 14.28 Adj. fields Wheat stubble Mar.-April 1.30 11.69 Combined Wheat stubble Mar.-April 1.34 12.40 PhillipsSedgwick Co. Wheat stubble 1963-69 No data 10.53 Sand Draw Green wheat May 1 No data 5.23 May 11 0.05 4.79 May 21 0.34 10.52 June 1 2.04 16.74 May 1 Near 0 7.17 May 11 0.53 9.12 May 21 1.80 16.49 June 1 3.95 24.90 ave. Adj. fields Combined ave. Green wheat Green wheat H-D Index- Near 0 6.44 May 11 0.33 7.26 May 21 1.18 13.93 June 1 3.13 21.40 May 1 l/The H-D index is expressed measurements are presented in decimeters in inches. and height �9 25 / / 20 15 5~---~ o~ 5 - 1 ~ ~ ~ 5-11 5-21 6-1 DATE Fig. 2. Growth of green wheat on and adjacent Property during the spring of 1979. to the Sand Draw �10 ,./ 30 ,/ / / / 25 / ~ / / / 20 ,/ ,/ {Il B / ... . . / ... / /' 15 ~ , ,/ ~ / ;!. ,/ 10 / /; 5 / ,/ . " }9q1 / ,;.-:. .. .../..~. .. OJr3!· •• ",,-y / :. ". ••• . . .. / . . .. . .. '" ..••••• ... • ~..... .»: / .,_~ ...~ . .. ., ••••...... .... .. ... ... .. .. .. ...... ' / ./ -_ - / / --/ / / o 20 §Rn.. Fig. 3. 30 20 10 30 MAY Comparisons of wheat. growth among years in the study region with 1979 wheat growth on and adjacent to the Sand Draw Property. �11 30 • • 25 20 Ul I.sl ::r: U ¢:i 15 z H ~ ~ I.sl ::r: 10 • 5 't = .964 y = -1.98 + .239 (x) o 1 2 3 4 H•.D INDEX (DECIMETERS) Fig. 4. A correlation of 1979 measurements and the H-D index. of green wheat height 5 �- 100 80 Q t.:I s ...1 III 60 t.:I ~ I ~ / ..... A/ N ra.. 40 0 ~ t.:I ~ III 20 O~~~--~----~----~--~----~--~~--~----~-22 8 15 17 24 1 10 29 APRn.. Fig. 5. MAY Comparison of spring stubble plo'ving progress 9-section study area in 1979. 5 JUNE on the Transect route and the �100 s _/ .;\.9ffo /~Lh 60 I ~ ~ ~ ~'\.919 • I ~ tzl u )..'•• 0161 80 ~ ~ - I Cl ! r / / 17 APRn. 24 )...,.t.r >: I-' W 40 b:I C4 20 o 10 Fig. 6. Progression 1970's. 1 of spring stubble plowing 8 15 MAY 22 29 5 JUNE in the study region during the 1960's and �14 not begin until July 21-21. The Sand Draw wheat was harvested on July 26. Initiation of harvest was about one week behind normal for the area and the slow progress delayed termination so it was approximately two weeks behind normal. Vegetative Cover Types Spring, 1979.-- The approximate acreages and percentages of cover types present during spring, 1979 in the 9-section study area are illustrated in Table 3. Note that about 95 percent of the acreage was farmed and that around 90 percent of this was in wheat stubble, greenwheator fallowed set-aside. When shortgrass pastures and occupied farmyards were excluded, less than 1.7 percent of the land remained unfarmed and containing annual and perennial forbs and grasses or shrubs and trees. When section 19, which contained the Sand Draw Property, was excluded, the remaining 8 sections contained less than 1 percent unfarmed cover of value to pheasants. Summer, 1979.-- Table 4 summarizes cover types available to hens and broods in late July and August, 1979. Increased acreages of row crops and millets planted in spring, 1979 replaced fallow, mulched stubble, and corn stubble. About 9 percent of the total land area was center pivot irrigated containing corn, beans, and grain sorghum. Weed-grass acreage increased from spring due to breaking out shortgrass sod to establish weed growth and to seed grass and grass-legume mixtures on the Sand Draw property. A portion of section 30 was also reverted from cropland to mid-grass pastures for future grazing. Wheat stubble present after July harvest was rather short and weedy. The millet and bean fields upon harvest in late August and September contained no significant cover. Corn fields were also disked after harvest and provided little feeding cover. Edge Areas.-- Table 5 summarizes the approximate amount of edge cover per section within the 9-section study area. Note that section 19, containing the Sand Draw property, included about twice the amount of edge of other sections and that much of the better quality edge was located on the Sand Draw. Table 6 summarizes an effort to categorize edge types and rank them, and to show the percentage of each in the study area. Edge types,change rapidly with seasons and with progression of farming. Corn stalks are destroyed in early spring, reverted to fallow, and then back to corn or other row crops by mid summer. Likewise, wheat stubble is transformed to fallow, whereas green wheat has little cover value until it starts its primary height growth in May. With the exception of shortgrass pasture, most unfarmed permanent cover edges have greater value for pheasants, especially edges between permanent cover and cropland. Study area roadsides were confined to a narrow 2 to 8 ft. wide mowed shoulder of low value for cover. Wider roadsides usually were dominated by cheat grass so that little residual erect cover was present �15 Green wheat wa s measured by growth (height) in years previous to initiation of this study (Fig. 3). The H-D (height-density) index (Robel et al. 1970) as modifeid by Kirsch (1977) provided a measure of cover value for nesting pheasants among various cover types and was initiated in this study. Therefore, both height and heightdensity (H-D) measurements were obtained on green wheat in 1979. A close relationship between the two, indicated by a high coefficient (r = 0.964), is illustrated in Figure 4. The H-D index will be used as the dominant measurement of cover value for nesting pheasants among cover types in subsequent years of study. It is of much greater value than height data used alone. Farming Activities Progress of Spring Stubble Plowing.-- A comparison of initial spring stubble plowing progress in the 9-section study area and the transect route is illustrated in Figure 5. Note that progress in the study area lagged behind that on the transect route by a few days throughout the spring. Comparisons of 1979 plowing progress with that of previous years is illustrated in Figure 6. The illustration shows that 1979 plowing was delayed and one of the latest on record. Wet field conditions from May 8 to the 20th were partially responsible for the delay in both surveys. A couple fields in the study area were started, but not completed until mid-June due to farmers' commitments to planting nearby irrigated lands. The stubble fields within the Sand Draw property, designated for conventional farming, were not disked until late May, however, the private land control was worked in mid May. Hini-till Division and private tracts we r e initially worked July 1 and completed during the first week in July. Green Wheat Destruction and Harvest.-- Poor stands of green wheat were destroyed in April and May in numerous fields throughout the study region and most that remained were sparse, weedy and nearly all were sprayed with 2,4-D herbicide in spring. Portions of several fields within the 9-section study area were destroyed and subsequently planted to millet. Approximately half of the field immediately to the southeast of the Sand Draw property was swathed and baled on June 21 to comply with set-aside requirements. No other wheat field were observed destroyed to comply with set-aside but some fields were planted to millet and others lay idle because of acreage restrictions. Some nests were reported destroyed and some hens were potentially lost elsewhere in the region because of June destruction of wheat to comply with set aside requirements. Wheat cutting began on July 12 on a couple fields, but major efforts did not get underway until about the 15th of July. Showers and damp waa t he r slowed harvest. Cutting of several large fields did �Table 3. Acres per section of various cover types on the 9-section extensive study area in March, 1979. 13 18 17 24 Section 19 20 25 30 29 Total 160.6 142.3 560.0 232.7 285.9 266.0 347.1 298.5 54.2 2347.3 40.8 197.4 3.0 202.0 3.5 103.0 72.0 396.8 173.8 76.8 2.3 5.4 4.5 288.9 Cover Type Wheat stubble Mulched stubble Green wheat 428.2 Fallow 40.3 Corn stubble 1.6 119.4 Millet stubble 101.1 283.3 463.1 2098.1 36.5 9.6 112.1 578.2 10.1 179.4 3.1 40.9 0.7 115.1 60.0 14.9 Percent 26.0 •.....• 0\ Total cropland 644.0 -- - - - - - 562.1 - - - Mixed gr. pasture 62.9 Occupied farmyds. 4.0 Weed & grass area Roadsides - 0.3 0.8 Trees 5.5 637.3 - 0.5 635.8 633.3 117.5 17.9 3.3 2.8 0.9 1.6 1.2 1.8 25.0 1.0 2.0 10.0 640.8 636.2 644.4 623.3 - - - - - - - 0.7 1.3 645.1 464.2 4.3 Shrubs Total acreage 634.9 -- -- - - -- 638.8 - - 617.4 - - - 629.4 - - - 5445.9 94.7 - - - - - - 7.5 0.8 189.2 3.3 5.8 6.9 16.7 0.3 5.1 4.5 36.1 0.6 2.2 2.3 3.8 21.1 0.4 0.2 2.2 31.5 0.5 12.0 0.2 639.0 634.7 637.7 5752.5 �Table 4. Acres per section of cover types present on the 9-section study area in late July and August, 1979. 13 18 17 24 Section 19 20 25 30 29 Total Percent Summer fallow 270.3 198.9 565.6 238.0 291.5 268.8 362.3 312.3 166.3 2674.0 46.5 Wheat stubble 194.6 103.2 62.0 358.0 172.6 59.6 100.1 270.5 368.1 1688.7 29.4 Millet 179.4 10.0 38.5 174.9 37.3 95.0 535.1 9.3 139.9 2.4 254.6 4.4 130.0 2.3 Cover Type & stubble Sorghums 130.0 0.5 Corn (irrig.) 130.0 Beans (irrig.) Total cropland 644.3 - - 562.1 - - - Mixed gr. past. 62.9 Occupied farmyd. 4.0 & 637.5 - grass mix. Roadside 124.6 130.0 ---- - Weeds 9.4 0.8 Trees 5.5 / - - 634.5 - - - 464.6 - 0.5 - - - 633.3 - - -- 85.0 4.0 0.4 50.0 3.3 2.8 1.6 1.6 1.2 1.8 25.0 1.0 2.0 10.0 640.8 636.2 1.3 Shrubs 624.3 - - - 592.2 - - - 5422.3 629.4 - - - - - - - - 94.3 - - 7.0 26.8 182.2 3.2 5.3 6.9 16.2 0.3 4.3 4.5 66.5 1.1 2.2 2.3 3.8 21.8 0.4 0.2 2.2 31.5 0.5 12.0 0.2 .--- Total acreage 645.1 635.8 644.4 638.8 639.0 634.7 637.7 5752.5 - ----_--------~--- ,_. -....J �Table 5. Linear miles of various spring, 1979. Edge type Green wheat - wheat stubble Green wheat - roadside Wheat stubble - roadside Green wheat - fallow Wheat stubble - summer fallow Green wheat - weeds & grass Wheat stubble - weeds & grass Fallow - weeds & grass Fallow - roadside Fallow - shortgrass Wheat stubble - trees & shrubs Green wheat - trees & shrubs Green wheat - shortgrass Wheat stubble - shortgrass Wheat stubble - corn stalks Trees & shrubs --roadside Corn stalks - shortgrass Corn stalks - summer fallow Roadside - shortgrass Roadside - millet stubble Green wheat - millet stubble Wheat stubble - millet stubble Fallow - trees and shrubs Trees & shrubs - weeds & grass Shortgrass - weeds Section totals edge types per section within the 9-section 13 18 17 24 Section 19 1.75 1.35 0.50 0.60 1.50 0.50 1.50 1.70 1. 15 0.25 3.75 2.50 1. 50 0.50 2.50 0.70 0.65 0.25 0.10 0.25 1.00 0.10 0.30 20 25 30 1.00 0.25 1.55 0.35 2.35 1.35 0.55 1.20 0.10 2.15 2.95 0.70 1.70 0.30 0.45 0.15 0.65 0.20 0.30 0.75 0.50 0.70 0.60 1. 65 1.55 0.45 study area in 0.75 0.10 29 1.70 0.75 1.70 0.10 0.60 0.50 0.50 1.85 2.00 1.40 0.50 0.35 0.90 0.35 0.15 0.05 0.50 0.55 0.15 0.40 0.25 0.30 . 1.20 1.65 6.90 9.25 5.50 6.00 13.60 6.50 7.70 7.30 5.85 Total 13.70 8.50 12.30 2.75 6.30 1.90 1. 15 0.65 2.40 0.20 3.30 2.50 0.60 3.05 1.55 0.95 0.35 0.90 0.50 0.20 0.90 0.80 0.30 1.20 1.65 68.60 ,_. 00 �19 Table 6. Ranking Moderate Approximate values and proportions of edge types present on the 9-section Sand Draw study area in early spring, 1979. and type Linear miles to High Quality Shrubs/trees Shrubs/trees Shrubs/trees Wheat stubble Wheat stubble Edge - Wheat stubble - Roadside - Weeds & grass - \<Jeeds& grass - Corn stalks Fair to Moderate Quality Green Fallow Green Wheat 11.88 13.70 12.30 2.50 3.05 0.30 1. 90 49.20 8.50 6.30 0.65 2.40 0.60 0.35 0.90 0.50 0.20 1. 65 32.14 2.75 0.20 0.90 0.80 6.78 Edge Green wheat - Roadside Wheat stubble - Fallow Fallow - Weeds & grass Fallow - Roadside Green wheat - Shortgrass Corn stalks - Shortgrass Corn stalks - Fallow Roadside - Shortgrass Roadside - Millet stubble Weeds - Shortgrass None to Poor Quality 3.30 0.95 1.20 1. 15 1. 55 Edge Green wheat - Wheat stubble Wheat stubble - Roadsides Green wheat - Trees/shrubs Wheat stubble - Shortgrass Trees/shrubs - Fallow Green wheat - Weeds & grass Poor to Fair Quality Percent Edge wheat - Fallow - Shortgrass wheat - Millet stubble stubble - Millet stubble �20 Seven transmitters were recovered in late April. Predation, primarily by great-horned owls, was suspected as the cause of several of five predations, however, actual cause was not known in most instances. In addition, one hen flew into a fence and was killed, and one harness apparently became untied and dropped from the hen. These recovered transmitters along with one repaired transmitter were placed on 8 hens trapped during the last week in April in wheat stubble fields. Thus, a total of 37 hens were equipped with transmitters during the late winter and spring of 1979. Trapping efforts were much more difficult in late April due to a partial dispersal of hens away from the Sand Draw property. Eight more known hen mortalities occurred from May 1 through July 26. Contact with 6 other transmitters was lost, primarily in April and May. Predation is suspected on some of these, but equipment failure or movement of the hens out of the study area could also have occurred. A summary of hen fates and nesting is provided in Table 8. Predation and Mortality Great-horned owls, which nested near the Sand Draw Property and used it as their primary hunting area, were believed responsible for most of the transmittered hen losses from early April through early June. Coyotes and possibly red faxes were suspected in four predations, two of which occurred after nest completion in summer. Feral cats and skunks were common and badgers and weasels were also present. Prairie falcons were occasional spring visitors, and one pair of Swainson's hawks nested successfully on the Property. Several pair of crows and magpies were also residents. Hen number 202, suspected of laying or beginning incubation, was flushed under sweep tillage machinery in wheat stubble on May 3 near the Property boundary. She flew into an adjacent plum thicket and apparently never moved. The carcass was partially eaten when found, a few days later, however, possible injury on the machinery, when she flushed, was also suspected. Hen number 337 was killed by an avian predator after she had been incubating for approximately two weeks. Another hen was a suspected coyote predation soon after nest completion, howev er , her transmitter was lost and her fate uncertain. A third hen was killed by a predator (coyote suspected) after her brood was approximately 2 weeks old. Figure 7 illustrates that most of the spring hen predations occurred on or near the Sand Draw property. The impact of this on other hens in the harems and subsequent dispersal is suspected but not certain. Figure 8 shows the 1979 nest locations which were widely distributed on and off of the 9-section study area. �21 in early spring. Roadsides, in summary, were of very low value for spring nesting but increased in nesting and brood rearing value primarily from mid-June through July. Grasshopper Density Moderately high densities of grasshoppers were present in the locality during the summer of 1979. Samples conducted in July showed their relative abundance in different cover types (Table 7). Annual forbs contained high densities, especially in roadsides. Wheatfields contained varied low to moderate hopper densities and those with cheatgrass and weeds held more. Perennial grass and alfalfa cover contained moderate densities. Corn and millet fields contained few hoppers, especially away from field edges. Residual Straw on Summer Fallowed Fields Square foot samples of surface litter, primarily straw from the previous year, showed that sweep tilled fields retained significantly greater surface residue than conventionally disked or plowed summer fallow (t = 7.117 with 56 d.f.). Conversion from grams/sq. ft. to lbs./ac. for the respective samples would indicate that conventional su~ner fallow contained approximately 255 lbs./ac., whereas the sweep tilled method retained 1185 lbs./ac. over winter. This increased surface litter reduces the threat of soil erosion, but also potentially provides more favorable nest sites for hens during the subsequent late spring nesting season. Observations indicated hens selected for nest sites containing above average quantities of litter in 1979. Pheasant Hen Monitoring Hen and Transmitter Losses Solar powered transmitters were placed on 29 night-light captured hens in late February and early March, 1979. All but three were trapped in wheat stubble fields proximal to the Sand Draw property. One was obtained in weedy cover on the property and two were trapped during a trial effort in southwest Phil'lips County. Nearly all remained on or near the property using shrub and tree rows and wheat stubble for daytime cover and wheat stubble for feeding and night roosting. The two imported hens remained on or near the property. Monitoring efforts were handicapped by faulty equipment and frequent windy, cloudy, and rainy conditions in March and April. �22 Table 7. Grasshopper density indices obtained cover types in July, 1979. in several study area Location Quad.-Quart.-Section Mean density index per square yard Cover type Roadside - annual brome kochia and sunflowers 12 - S.E. 30 l3 - N.E. 18 39.28 Weed strip - sunflowers 14-16 N.W. 19 35.48 Alfalfa-cheatgrass 14-15 N.W. 19 14.40 Tree row with cheatgrass understory 12 - N.W. 19 5.24 Perennial 13-14 N.E. 19 mix mixed grass 20 - N.E. 8 25 - N.E. 9 Corn Millet Green 1 - S.E. 20 8 - N.E. 13 (ripened) wheat Green wheat Wheat & cheatgrass stubble (new cutting) 4-5 - N.W. 24 15 - N.W. 30 14.48 0 0.48 2.96 1-4 N.W. 19 16.00 12-13 S.W. 18 24.32 4 - N.E. 29 1 - N.W. 13 2.28 �23 Table 8. Known nesting efforts, nesting success and fate of transmittered hens during the spring and summer, 1979.l1 Transmitter Number Known Nesting Attempts Successful Nest 850 863 2 o Yes No 887 903 913 919 935 962 971 980 013 032 052 087 107 146 163 186 202 239 258 286 309 337 341 361 387 415 439 448 Fate of Hen Retrapped and released 1st & 2nd hens both apparently predator killed 2 Yes Retrapped and released Signal lost at start of suspected incubation fate unknown 2 Yes Retrapped and released 1 Yes Retrapped and released 3 Yes Retrapped and released ? Killed by avian predator in mid-May 4 Yes Retrapped and released Transmitter signal lost on April 20 - fate unknown 1 Yes Retrapped and released 2 Yes Retrapped and released 1 Yes Retrapped and released 1 Yes 1st hen killed; 2nd retrapped and releasedll 1 (more Yes Retrapped and released suspected) 1 (more Yes Retrapped and released suspected) 2 Yes Retrapped and released 2 No 1st hen predator killed, 2nd retrapped and released ? (l susp.) No Unknown mortality Transmitter signal lost on April 12 - fate unknown 1 Yes 1st hen killed; 2nd retrapped and released 1st hen slipped harness; signal lost on 2nd hen fate unknown 2 Yes 1st hen pre.; 2nd hen with brood predator killed 1 No Killed by avian predator Transmitter signal lost on April 14 - Assumed predator killed Signal lost soon after release - signal picked up on a live hen 1 year later ? (2 susp.) No Hen killed by avian predator 2 Yes 1st hen pred.; 2nd hen lost at nest termination 1 Yes Retrapped and released ? (1 susp.) No Hen predator killed approx. May 10 l/Only known nesting attempts are listed. Several others were suspected but not confirmed, and others were probably started and predator destroyed or abandoned before they could be suspected. �24 ·--13--- 18-- --- I ----17- I I I 202 (April) 962 May) 48 (May) 20 337 (June) .-~ I i •• I ., •• 25- I I I Fig. 7. 30 - I I I - -- - 29 I 309 (July) ~ mi. south Location of hen pheasant mortalities ( ) and month of within the 9-section Sand Draw study area in 1979. �25 ~07b 186b • 17 18 13 • 186a ~5bl~5C SAND DRAW 146a --- 24 & P~OPERT' b \ 309a 20 -----19-----..1 971b 850. 258a+ 971a 415a ~5Ob 415 013a 971d 032 13a {fY09b 032b ~19 29 1 mi e south Fig. 8. Location of known or st~gly and successful nests (~) study area in 1979. suspected nesting attempts on the 9-section Sand Draw (.), �26 Transmitter Impact and Monitoring Disturbance Increased predation or hen mortality due to presence of the transmitters could not be detected if it occurred. However, impact if any, was believed minor. Nearly all predator killed hens had been instrumented for several weeks or months before mortality occurred. One hen instrumented in late April, had apparently been laying for about 11 days before she was night-lighted. She returned about 1 mi. to her nest site upon release and continued laying to complete a clutch of 14 eggs, and began incubation 8 or 9 days later. Some human disturbance of hens was necessary, primarily in the early pre-nesting phase of monitoring to check for mortalities and to verify locations. Hens moved out well ahead on foot in the large stubble fields so that close harassment was not necessary and monitoring impact was not great. Use of triangulation using antennas mounted on vehicles and permanent towers reduced need for significant walking and subsequent disturbance. It was not always possible to distinguish whether a hen was initiating a nesting effort, or had become a mortality, when repetitiously located in one spot. Therefore, several days often lapsed before predatored carcasses and transmitters were found and recovered. Nesting Activities Cover type selection for nesting.-- All April nesting initiations and most early May efforts were located in wheat stubble except for one found in residual millet. Ten nests, located before they were destroyed, were initiated during the interval of April 16 through May 14. Several other initiations were suspected, but not found before they were destroyed or abandoned. Sweep tillage plowing, which retains a major portion of the stubble above ground (Fig. 9) was conducted on wheat stubble in Section 18, to the north of the Sand Draw Property, in fall, 1978. It was repeated there and on stubble immediately east of the Sand Draw in late April and early May, 1979. Two nests were found in this tilled stubble soon after the spring treatment and a third nest was not located before the hen was killed by an avian predator. One additional nest was believed destroyed ,by the sweep tillage treatment. All nests begun after the early May treatment would have terminated, if they had not been predator destroyed, before the second tillage was conducted in June. However, frequent rains delayed this second tillage, and most years, such nests would not have been able to terminate before the subsequent tillage operation on the stubble. �27 Fig. 9. Sweep tillage leaves a partially standing residual which will be utilized by nesting hens with probably nest destruction during a subsequent tillage. This treatment if delayed to mid-summer, retains more surface residue to protect soil and to provide a litter base in seeded wheat the following year. �28 One hen began nesting in green wheat around May 7 and a 2nd hen began on about the 12th. Figure 10 illustrates the projected dates of nest initiation in wheat stubble and green wheat and compares this with the H-D (height-density) index obtained on these cover types. As noted, most nests, initiated in green wheat began in mid to late May after the H-D index approached or surpassed the index for wheat stubble and after hens were forced out of wheat stubble by tillage. The latest initiation in wheat occurred on July 16. That hen was flushed from the nest on July 26 by a combine, which passed over her, but she returned to terminate her nest successfully (Table 9). One non-transmittered hen was accidentally flushed from a large clutch in late April in residual weeds along a road shoulder. Green weeds were present around a successful nest established in uncut field of set-aside wheat where standing residual wheat predominated. Green weeds dominated in a roadside adjacent to harvested wheat where one hen's fourth known nesting attempt terminated successfully on August 26. Specific Site Selection.-- Monitoring permitted visual inspections of many nests primarily during incubation and post-hatch periods. Observations and photos showed most nests in wheat stubble were located where a small pile of straw, deposited the previous year by a combine, provided the base for a nest. These small straw deposits were used primarily in bowl, formation, but in several instances, they provided a partial canopy as well. In general, overhead concealment of hens in wheat stubble was poor. Much less straw remained in green wheat fields, but visual evidence indicated some nest bowls contained considerable residual or new green cuttings gathered by the hens. The amount of residual litter present at nest sites should be compared if possible to average amounts to determine if hens are selected for sites which contain increased residual. Litter for a nest bowl appears to be an important factor in site selection, and lack of it in green wheat fields is one reason these have been considered poor to marginal nesting habitat. Figure 8 illustrates tha t most hens placed second or subsequent nests moderately close to the first nest site. This occurred even though a change in cover types was often'noted. Nest Fate Related to Cover Type.-- None of the nests initially established in stubble, or sweep-tilled stubble, were terminated successfully (Table 9). Six of these were destroyed by spring disking and four were destroyed by mammalian predators. One was predator destroyed the night before it would have been disked under. Human disturbance in locating nests ahead of disking caused two desertions which would soon have been destroyed anyway. Three of the hens were flushed by monitoring personnel, but three other hens flushed i~nediately ahead of the tillage machinery so that none were disked under. Farmers report that some hens are destroyed during spring stubble tillage operations. �100 X' NEST INITIATED IN STUBBLE • NEST INITIATED IN GREEN WHEAT 80 ~ Q ~ tl w 60 d] 0 ell ~ •....• ~ til Q I ~ tal i 40 :x: -- 2.0 ~ r... 0 \.~y I N \0 til :z ~ ~ tal I ~ a. fj 3.0 20 ~ 1.00 t!) ~ 0 5~ I ..... ~;-:~ . ~~R~~. ~~.~~.~ ~?'*.~RASS 0 o 15 20 25 APRIL Fig. 10. 30 5 10 15 MAY 20 25 30 5 10 15 JUNE Comparisons of stubble plowing progress, and H-D indices of green wheat, wheat stubble, and residual grass cover with time of nest establishment per cover type. �Table 9. Hen no. Suuuna ry data on known nesting attempts and fates by transmitter equipped hens in 1979. Nest no. 850 Incubation onset onset Wheat stubble 4-17 6-03 5-06 6-19 4-30 5-17 Green whe a t Green Cover type Green Green 919 1 2 3 971 013 wheat Residual millet 913 935 wheat I,heat stubble 887 11 Laying- association wheat Days incubated Clutch 5-26 7-12 18-20 full term 14 12 d Lsked hatched 9 5-15 6-02 5-16 6-25 full term 13 13 deserted hatched 10 5-06 5-23 5-23? 6-23 9 full term deserted hatched o 6-01 5-12 5-28 6-20 full term hatched 12 date size 12 Roadside weeds 1? Green 5-26 wheat wheat uheat Wheat stubble Green wheat 6-01 or 2 7-13 8-01 5-22 0 6-19 or 20 19 7-14 1 8-23 full term 6 unk. 9 8 Hheat stubble I 2 I? hatched 6-19 full term deserted hatched 5-20? 6-03? 6-26 full term unk. hatched 8 or more hatched 4 Prior unk. 9 disked under hatched o 17-18 15 16 disked under predator des. o 9 6-11 full t erm 9 hatched 5-31? 6-19 6·-01 7-12 o to I full tem 13 5-09 5-24 7-10 15 full term 14 5-01? 5-31 5-18 6-11 5-23 7-04 full term IJheat stubble Green whea t 5-06 6-21 5-27 7-03 6-14 7-18 Green 5-07 5-19 5-14 6-05 4-16 6-07 Green wheat wheat St;eeped stubble Green wheat Hheat stubble Green ·.Il Dates still bllt d~s~rted whf Le incubating Flushed by combine Finally successful 9 8-11 flushed nesting & abandoned efforts suspected Abandoned due to disturbance & pred. destroyed 5-23, disked 5-24 Nest suspected in residual set-aside (wheat & «eeds) but never confirmed - believe predator destroyed. Set aside 6-01 6-18 7-11 full term 13 hatched 13 1 2 415 combine 13 11 o not Hen returned after being flushed by full term Nest suspected but never confirmed before wheat stubble fiel.d was disked Gr'een wheat 5-27 6-01 6-23 full term 4 309 Hen 5-20 7-12 I? 258 7 Fl.ushed prior to disking of field 7-01 107 186 2 Hen flushed in front of traclor 6-07 ";reen wheat 2? disked under predator des. deserted hatched Comments 5-03 6-05 1? 163 o o o o Green Green I? o Hheat stubble 2 3 4 2? under 1/ eggs hatched o 052 146 Nest fate Sweeped stubble 5-06 5-22 6-09 18-19 11 predator des. Believed laying in wheat stubble when field was dIsked. Nest not found prior to tillage. Ripened wheat 7-16 7-23 8-15 full term 5 hatched 5-13 5-23 7-01 7-21 032 Termination wileat 6-17 a rv projected based o n 1.3 days per egg laid. 5 11 8 Hen flushed ahead of tractor 5 9 lien d i.d not attempt r enes t o predator des. hatched 11 preda tor des. ha tched 8 o Dares should be considered estimates rather than actual. Destroyed night before field was disked (brood & hen disappeared) w o �31 Predators destroyed two nests established in green wheat and one hen deserted her third nest at the time of harvest. Wheat harvest was delayed by weather permitting additional hens to terminate successfully. All 16 known location successful nests were in green or ripening whea t or were placed on residual wheat or weeds closely associated with wheat. One additional transmittered hen, number 087, was located with her brood in wheat after being lost for an extended period. Presumably she nested in green wheat but this could not be confirmed. Monitoring of individual hens provided strong evidence that several additional nesting attempts occurred in wheat stubble, but they were predator destroyed or disked under before incubation started and before they could be located. Extensive predation, primarily by skunks in roadsides, had previously been confirmed (Snyder 1974), but nesting efforts in adjoining large stubble fields were believed fairly secure from predation. However, monitoring showed considerable nest predation in stubble fields but less in green wheat fields. Stubble fields contained greater quantities of alternate food sources (small rodents, insects, etc.) than green wheat fields did which attracted skunks and badgers to the stubble fields so their chances of finding pheasant nests there were greater. Alternate food sources also increase in abundance with the progression of summer, thus, potentially gradually alleviating the stress on nests. Clutch Size.-- Clutches initiated prior to May 16 averaged 12.5 eggs per nest in a sample of 11 nests. Clutch size averaged 10.3 from mid-May through mid-June in a sample of 9 nests. It dropped to 7.7 eggs per nest in a small sample of 3 nests initiated after mid June. Other Nest Statistics.-- More than 266 eggs were laid in 28 nests for an average of 9.5 eggs per nest. More than half the nests (15) were successful, but less than half the total eggs (approximately 120) in these 15 nests hatched successfully (later, smaller clutches had a higher success rate). These data combined with information from other unobserved nests indicated approximately 142 chicks hatched from 17 nests for an average of 8.35 chicks per nest. Only one of the 18 known surviving transmitter equipped hens failed to nest successfully. Her second known nest was predator destroyed (Table 9) and she spent most of the re~aining summer in row crops. Recycle Time Between Nesting Attempts.-- Table lQ summarizes approximate intervals between termination of one nesting attempt before initiation of a subsequent nest. These data indicate hens that were still laying when their nests were destroyed, usually immediately established a subsequent nest. Recycle time was lengthened for hens that had progressed into incubation. Most hens whose nests were destroyed by predators or machinery immediately moved distances of 1 or 2 miles but then returned to renest in a few days within the general locality of their previous attempt. �32 Table 10. Bird Number The approximate time interval for hen pheasant renesting during the spring and summer of 1979. Days Incubated Time Interval to Renest 913 0 1 887 0-1 1 971 0 1 971 1 7 309 1-2 4 163 5 8 415 15 14 186 18 7 971 19 12 850 19 9 �33 Nesting in Relation to Weather.-- Moderate temperatures and frequent periods of showers and rain prevailed throughout most of the 1979 nesting season. Only one week of very hot dry weather occurred in early August. Crops, roadsides, and other annuals remained green into late summer, and wheat stubble contained considerable green weed and annual grass in late July through September. Most hens were able to bring off a successful nest by their second knovffinesting attempt. The latest nest initiation occurred on approximately 21 July and it terminated successfully. Mditional data for comparison will be forthcoming in subsequent years. Hen Use of Cover Types Hen monitoring and location of known nests indicated nearly all April hen activities and nest initiations, and most early May efforts occurred in wheat stubble. One was placed in residual millet and one non-transmittered hen's nest was found in residual weeds along a road shoulder. Table 11 summarizes monitored hen occurrence in cover types by hen activity period. This shows that small grain stubble, primarily wheat stubble, was used extensively in spring during prenesting laying, and incubation when nests were placed in stubble. Likewise. green wheat became the primary cover used as it increased in height in May and became attractive for nesting. Meanwhile wheat stubble was being plowed under, which forced use of green wheat as about the only cover available. Hens with broods gradually shifted from wheat to roadside weeds and then to green row crops such as corn, sorghums, and to millet. This movement was in part caused by harvest of wheat in July. Most spring and early summer monitoring of hen locations occurred after termination of morning feeding and before evening feeding activity began. Figure 11 and Table 11 illustrate the progression of major cover type used through the spring and summer or 1979. Cover Type Use in Relation to Availability.-- Nearly all wintering pheasants in the 9-section study area were concentrated on or proximal to the Sand Draw Property, which contained extensive areas of trees and shrubs. A corn stalk-shrub-wheat stubble association was available along the north side of the Property and contained many pheasants. However, trapping and monitoring in late February and March showed many pheasants used wheat stubble-woody cover-weed associations elsewhere on and proximal to the property. Heavy snow did not occur during the interval. Transmittered hens tended to disperse away from the property in mid-April as previously noted. Wheat stubble was used almost exclusively by these hens until green wheat afforded adequate protective cover in early to mid-May. Tables 3 and 4 show percentages of various cover types available on the 9-section area in spring and summer. Activity patterns show weeds, roadside (weeds and grass), shrubs and trees were used in much greater proportion than they were available in early spring. Use of these decreased during the nesting interval when most activities were concentrated in wheat stubble and gradually shifted to green wheat until after harvest. �Table 11. Primary cover type use by hen activity recorded occurrences). 1/ Hen activity Prelaying stubble status Prelaying wheat Laying Cover type Grass & grass/ Roadweeds Weeds side Shrubs trees & Corn Green crops Millet Sorghum 43 6 6 6 7 22 81.• 35 5 5 5 15 66 1 26 121 7 3 7 14 15 192 1 4 4 7 14'1:'/ 284 1 2 2 116 1 8 2 2 2 1 9 148 1 24 18 5 5 7 7 23 24 5 49 15 15 31 36 34 in wheat w ~ in green in green wheat Incubating wheat Green wheat the summer of 1979 (number of in wheat Laying in wheat stubble Incubating stubble Small gr.stubble period during 1 in green Hatched to 1 wk. old brood 1 to 3 wk. old brood 3 wk. and older brood l/Small grain stubble included wheat stubble, of millet and oats stubble. l/occurrence in new stubble following recently sweep-tilled harvested wheat wheat. stubble and small amounts �~~~~~... 1\ 90 ~-- - - - -"\ 80 ,"' / / ~ I r.~ / 70 \ (V/ (§/ \ / \ / 60 ~ ~ I 50 ~ _. \~ '..' . , ,, 20 , • I ' .. 10 ,,' 0'*' 0_ - 0.' ~o • "~ DSlDe•••••• - • - - • / .... .... . ,--- •••• g _• • • • •• .A.ow-;'--....,_~.a....~ "', / _ •• o 21 22':" 5 APRn.. Fig. 11. 6 -"19 20 .:.. 2 MAY Cover type location w 3 -16 17 .:.. 30 JUNE of monitored It • • , V1 , x,"\ ., ---' .d, ;y' • ••.•• 8 ,'",~ ",. ,~ . ,~ ~~' OA ~S- O'fJ A,*",' ,~ , .•••lIS. "''3 • ,,~" 'G RAss & R .q~ ••••• ,,~~ ~ <':f' ' oj) I • ' ,AA' ,~~ , ~ \~ I '. ' 30 .-;y¢ • I I r -. \~ , .. s't\}~~1..~• 40 •••••••••• \1i I I •• ~ ~ \ I • O~D • ~ \ I ~ ~ \ / 1 I ," • _o, ••••••••• 1 .:..14 ° • • • •• 15':" 28 ••• e •••• 29':" 11 JULY hens from April through August, " •••••• ~ " 12 - 25 AUGUST 1979. �36 Hens with broods moved to green row crops, predominately corn and sorghums, and to roadside weeds and millets after wheat harvest. Some hens moved to locations outside the 9-section study area so an exact use preference index could not be tallied. Figure 12 illustrates percentage occurrence in green wheat in relation to growth and in wheat stubble in relation to spring plowing of stubble. Hen-Brood Activities Use of cover types by hens with broods has been previously stated in the text and illustrated in Figure 11. Hens generally remained near the site of nest completion for a week or so, or until forced to move by wheat harvest. Roadsides were increasingly frequented and these often were used as travel lanes for movements to green row crop or to millet fields. Some hens with broods moved one or more miles to locations in row crops where they remained resident in August when the transmitters were recovered and monitoring was terminated. Water was noted proximal to all broods either in the form of standing water or from irrigation by center-pivot sprinklers. Brood Sizes and Mortality Hens with broods were seldom bothered by monitoring personnel and broods were seldom observed when young. Highly reliable counts of broods, when they were a month or so old, were seldom obtained. Therefore, no significant data on brood mortality with time is available. In several instances, little decrease in brood size was noted, but, in other instances, decreased numbers mayor may not have been due to a poor count or dispersed brood. Increased effort will be directed toward brood mortality in forthcoming years. Evaluation of the Impact of Mini-till Summer Fallowing Approximately 45 acres of wheat stubble in 3 fields on the Sand Draw and one adjoining privately owned field were treated with contact and pre-emergent herbicides by the lessee in late summer, 1978. This was done as part of an attempt to evaluate the impact of this new mini-till farming method on Fheasant nesting. Few weeds or annual grasses were present in these treated stubble fields in 1978 either before or after treatment because of extremely dry weather conditions (Table 1). Little significant weed growth occurred on untreated controls or other fields in the study area. �100 80 60 ~ 25 ~ W til f:> c.l :x: ~ 20 H ~ ~ ~ OCCURRENCE --_ -HEN - -- --.- -- .- 40 ~WHEKT-S~BLE -...J " ..•.•.. / 0. ~ 15 ~ c.l :x: 10 20 •.. ~ "- ~ z 5 !j ffi o "'- " <, a 15 20 25 APRIL, Fig. 12. 30 5 10 15 20 25 30 1 10 15 JUNE MAY Hen occurrence in green wheat and wheat stubble to cover availability. 5 in spring, 1979 in relation 20 24 �38 The pre-emergent herbicide (atrazine applied at 1 lb. active ingredient per acre) was moderately effective in suppressing weed and annual grass growth during the spring of 1979. One field, out of the four treated, showed considerable weed growth for unknown reasons. The others remained nearly or completely weed-free until July 1 when they were disked by the lessee. A May 15 sample of vegetative density illustrates the greatly reduced presence of green vegetation in the treated stubble when contrasted to the controls (Table 12). The wide dispersal of monitored hens for nesting (Fig. 8), which possibly, in part, was induced by avian predation on hens (Fig. 7), eliminated an adequate nesting sample on either the treated or control plots. One nest was initiated in the private land control and was subsequently disked under in May. This nest was only a few feet from the treated stubble, however, no difference in appearance be twe en the treated and untreated stubbles could be visually detected. If other nests were started on the Sand Draw fields, they were predator destroyed before they could be located. Some information on the potential impact of mini-till wheat farming methods on nesting pheasants can be derived even though the treated plots did not provide it. All stubble fields in the study area were weed free in late summer, 1978, because of dry weather and soil conditions (Table 1), and most contained little new green growth at the start of nesting in spring, 1979. Therefore, there was no significant difference in cover value for nesting between treated and untreated fields. Hens tended to select specific sites containing increased straw litter rather than sites with increased green weeds. Therefore, if these fields had all been treated with herbicide and subsequently not tilled until late June or early July, several first nest attempts and some renest efforts would probably have terminated successfully. Question arises, however, when predators destroy nests in stubble, as to whether most renests will be placed in stubble again, or in green wheat instead? It is probable that in 1979, most second nests initiated in early May would be again placed in stubble (if not disked) and most would probably have terminated before subsequent stubble tillage assuming all stubble fields had been chemical fallowed. Second nests initiated after mid-May might or might not have been placed in green wheat when an abundance of stubble was also present. If they were placed in stubble at a late May or'early June date, they would subsequently be destroyed by tillage. Nest destruction at that late date would undoubtedly significantly reduce nesting success. �39 Table 12. Comparison of hits and misses of green vegetation on herbicide treated (mini-till) wheat stubble and adjacent untreated stubble in mid-May, 1979. Location Hits Treated Misses Hits Untreated Hisses South field 5 95 52 48 Middle 7 93 84 16 North field 0 100 87 13 Schuler 0 100 40 60 12 388 263 137 Total field field �40 LITERATURE CITED Kirsch, L. 1977 (Rev.). Instructions for use of height density pole for measuring residual vegetation on grassland wildlife habitats. 2p. Mimeo Rept. Robel, R. J., J. N. Briggs, A. D. Dayton, and L. C. Hulbert. 1970. Relationships between visual obstruction measurements and weight of grassland vegetation. J. Range Manage. 23(4):295-297. Snyder, W. D. 1974. Pheasant use of roadsides for nesting in northColo. Div. of Wildl., Special Rept. No. 36. east Colorado. 24pp. 1979. Evaluation of nesting cover preferences of pheasants in relation to wheat farming methods. Colo. Div. of Wildlife. Game Research Rept. April:l-l0. Prepared by __ ~~~~~~·~rV~~~~.~~~.~.~~=· ~~ Warren Wildlife D. SrlYd7r~.li) Researcher C �April 41 JOB FINAL State Work No. W-37-R-33 Job Tit I e Job No. Survey 9a E_v_a_l_u-=---a_t_i_;.o_;_n_;__;_o:._:f:..._tc::..h:..._e_;:;E_;_f_;:f_:e_:c:._:t::_:s:... n~H=u.=.:n:..:t:..:i:.::n:.<g.,__ _ Regulations Covered: Personnel: Game Bird 3 Plan No. Period REPORT Colorado -------------------------------- of Project 1990 1 April on Sage Grouse 1975 through Populations 31 March 1980 J. Dingman and M. Monahan, Univ. of Denver, R. Ryder, Colo. State Univ., C. Donner and F. Giese, U.S. Fish and Wildl. Serv., L. Adams, H. F. Alexander, T. D. I. Beck, D. Benson, C. E. Braun, T. Campbell, A. Chappell, D. Covic, C. Crawford, D. Crawford, P. Curtis, S. R. Emmons, H. Funk, K. Giesen, D. Gore, W. P. Gorenzel, J. Hobbs, R. Hoffman, J. Jackson, J ..E. Kautz, R. Leasure, T. Lynch, F. Marcoux, S. McElderry, K. Miller, T. Olson, S. Palm, B. E. Petersen, S. Porter, B. Quinlan, S. Quinlan, B. Renfrow, W. Russell, T. Schoenberg, H. Spear, R. Stark, S. Steinert, J. Wagner, and R. Zaccagnini, Colorado Division of Wildlife. ABSTRACT The objectives of this study have been achieved. Reports covering the various objectives have been published, prepared and submitted and others are in progress. Those in progress will be published under Work Plan 22, Job 1 and are listed in the project documents for W-37-R-34. Manuscripts published, submitted or in progress are listed below. Beck, T. D. I., and C. E. Braun. Condor 80:241-243. 1978. Weights of Colorado sage grouse. Braun, C. E. 1977. Autumn structure of sage grouse populations, Park, Colorado. Am. Assoc. Advance. Sci. 143rd Annu. Meet. Publ. 77-2 :Pap. 119. North AAAS t 1978. Effects of changes in hunting regulations on sage grouse harvest. Trans. Annu. Meet. Central Mtn. and Plains Sect., Wildl. Soc. 23:Abstract. _________ , and T. D. I. Beck. 1977. Population North Park, Colorado. Trans o· Annu. Meet. Sect. Wildl. Soc. 22:14. _________ , T. Britt, of sage grouse dynamics of sage grouse, Central Mtns. and Plains and R. o. Wallestad. 1977. habitats. Wildl. Soc. Bull. Guidelines 5:99-106. for maintenance �42 Emmons, S. R., and B. E. Petersen. 1979. sage grouse in North Park, Colorado. Biotelem. 2:215-218. Telemetry investigations Proc. Int. Conf. Wildl. Stabler, R. M., C. E. Braun, and T. D. I. Beck. 1977. Hematozoa sage grouse from Colorado. J. Wildl. Dis. 13:414-417. of in Braun, C. E., M. F. Baker, R. L. Eng, J. S. Gashwiler, and M. H. Schroeder. 1976. Conservation Committee report on effects of alteration of sagebrush communities on the associated avifauna. Wilson Bull. 88: 165-171. 1976. Emmons, S. R. Colorado. North Park's sage grouse. Colo. Outdoors 25(5):26-28. 1980. Lek attendance of male sage grouse in North Park, M.S. Thesis. Colo. State Univ., Fort Collins. Petersen, B. E. 1980. Breeding and nesting ecology of female sage grouse in North Park, Colorado. M.S. Thesis, Colo. State Univ., Fort Collins. Braun, C. E. Population dynamics Colorado. J. Wildl. Manage. tions. of sage grouse in north central, In Prep. Harvest characteristics of sage grouse under varying Wildl. Soc. Bull. In Prep. Characteristics of sage grouse populations Div. Wildl. Spec. Rep. In Prep. Petersen, B. E., and C. E. Braun. female sage grouse. Condor. Emmons, S. R., and C. E. Braun. grouse. J •. Wildl. Manage. in Colorado. Breeding and nesting In Prep. Lek attendance In Prep. regula- patterns ecology of of male sage A Stabler, R. M., N. J. Kitzmiller, and C. E. Braun. redescription Eimeria centrocerci from sage grouse in Colorado. Trans. Am. Micros. Soc. Accepted. Beck, T. D. I., and C. E. Braun. 1980. Variation in counts of male sage grouse on leks. Proc. West. As'soc. Fish and Game Comms. , Accepted • . ' Prepared by: Clait E. Braun Wildlife Researcher Colo. of �April 43 1980 JOB FINAL REPORT State of Project Colorado No. Game Bird Survey W-37-R-33 3 Work Plan No. Job Title: Evaluation Job No. 9b of the Effects of Changes in Hunting Regulations on Sage Grouse Populations: Evaluation of Censuses of Males Period Covered: Personnel: 1 January 1978 through 30 September 1979. D. Hein, P. Lehner, R. Ryder, Colorado State University; Clait Braun, Jim Dingman, Steve Emmons, Howard Funk, Ken Giesen, Sue McElderry, Brett Petersen, Steve Porter, Tom Schoenberg, Colorado Division of Wildlife. ABSTRACT Daily lek attendance patterns, breeding season movements, and habitat selection of male sage grouse (Centrocercus urophasianus) were investigated in North Park, Colorado from late March through mid-June 1978-79. Thirtyseven males (20 adults and 17 juveniles) were equipped with radio transmitters and studied on 5 leks. Peak male attendance occurred 25-37 days after peak female attendance on the 3 largest leks investigated. Lek attendance of radio-marked juveniles increased to 91-95% in mid-}fuy and decreased thereafter. Adult male attendance increased to 98-100% in mid-May and then decreased. A non-Iek attending segment of the male population was not observed. Juvenile males visited 2-4 leks, remaining on each for an average of 4.3 days. One juvenile visited 2 leks (4.5 km apart) in 1 morning. Most adult males (72.7%) visited only 1 lek but'2 visited 1-2 additional leks for 1 day and 1 moved to an alternate lek after 15 May. Juveniles and adults moved at least 23.9 and 10.0 km, respectively, during the breeding season. Four juvenile males moved from the study area, including a move of over 25 km in 5 days. Three adults moved to leks 12-21 km from the study area. Off-Iek locations were within 0.5 and 1.0 km of a lek for 39.9 and 62.6% of 160 locations, respectively. Males typically dispersed over 1 km in nonrandom directions from leks to feeding and loafing sites. Subsequent mid-day moves over 150-300 m resulted from disturbance. Over 60% of the juveniles (60.4) and adults (69.8) returned to leks in the evening to roost. �44 Sagebrush (Artemisia spp.) canopy coverage and height at 160 feedingloafing sites averaged 28.1% and 43.5 cm, respectively. Approximately 90% (89.7) of the roosting locations occurred in sagebrush with a canopy coverage of less than 20% (avg. 8.7) and height less than 40 cm (avg. 18.7). Recommended procedures for count Lng leks are given. �45 INTRODUCTION Sage grouse inhabiting grasses are widely dis tributed rangelands dominated by sagebrush and forbs with interspersed dowse Sage grouse all seasons dependence in western and subdominant native and cultivated on sagebrush has been documented North America, for food and COver in by Patterson (1963), Klebenow (1969), Peterson hay mea- (1952), Dalke e t al. (1970), Eng and Schladwei.ler (1972), and Wallesta.d e t a1. (1975). In mos t states agencies routinely with sage grouse collect tunity and harvest. Sage grouse based on counts of males estimates August. population of nesting success males to a particular i. e. black grouse grouse management grounds) (Pedioecetes that less in breeding daily attendance patterns been in July and sage grouse on population size. of individual Recent studies (Lyrurus tetrix) (Robel 1969) and sharp-tailed phasianellus) Data presented of other oppor- lek. lekking grouse. (Rippin and Boag 1974), than 50% of the m<;.:.!e population given time. hunter in April and and brood size obtained trends wildlife has primarily Although peak counts of male and female is known concerning state data and regulate on leks (strutting leks have been used to estima.te little populations, in this report is present suggest on a lek at any are from March through �46 July 1978 and 1979, and include information breeding grouse season movements, in North Park, The primary concerning breeding season. males objective Samples Hypotheses by age (juvenile and adult) and tested were: (1) juvenile than 3 days/7-day capture; 1 year (2) adult (both go to leks during the display on display on only 1 lek. and habitat « (3) males (4) juvenile males than 1 lek; and (5) adult males season movements period; season; trapped off leks rarely season of initial tion, breeding knowledge of male sage grouse during the stratified attend leks less adults and juveniles) more of this study was to increase attend leks daily during the breeding breeding of male sage on and off leks) were used to document daily lek of males. of age) males selection Colorado. daily activity patterns location (trapped attendance and habitat on lek attendance, selection In addi- were examined. �47 DESCRIPTION OF THE STUDY AREA The investigation County, Colorado. North Park, was conducted The study area approximately was near was within 40° 30' and 41 and 106 30' west longitude. 0 tion of the study area's The area River, 2 219.5 km , The area although separated eastward from habitat River (Fig. 2). exact boundaries was between was relatively by d r a inag es , River Mountain. 2400 and flat with low undulating Major drainages in- from north to south into Lake John, the North Fork of the Nort!:_ Platte al.I of the Park Mountain, south by the North Fork to 2964 m on Independence cluded Lake Creek which drains then turning descrip- on bird movements. of the sagebrush and ridges 10' 0 a detailed and east by the North Platte boundary 2585 m (Beck 1975). benches and 106 were from 2400 m along the North Platte of most 1). location. depending the northeast Elevation Gill (1965) presented was approximately Elevations near 00' north latitude Ridge and Sheep Mountain, of the North Platte were flexible of Walden (Fig. was bounded on the north by Independence west by Boettcher Total area Jackson Lake John in northwest 16.0 km northwest The area 0 in North Park, River draining until--joining the North Platte south to north. Lakes from north to south which drains in the study area included �48 Fig. 1. Lake John study area, North Park, Colorado. �49 __ = _, --s>: . 'MATTENBERG -_- 2 L§t< ;" Fig. 2. Location of active leks in the Lake John area, r t ,__ North Park, N Colorado. �50 Lake John, Alkali Lake, Boettcher ephemeral unnamed lakes Lake and nurne rous smaller scattered of the study area was more rolling of the area has been described throughout. The northern than the southern half. half Geotogy by Beekly (1915), Finch (1957). and Hail (1965). The vegetation bunchgrass tridentata of the area was dominated with scattered vaseyana) comprised type in North Park. area were greasewood (Chrysothamnus (Salix spp.), vegetation perennial spp.), The climate measurements moisture rabbitbrush sarothrae), -willows Herbaceous perennial forbs and - of North Park are available, is cold and dry. prevailing in spring noonally normally in the study with few annual forbs (Beck 1975). particularly accounts increases Although no wind winds are from the south- £01" in the study a r ea-t U, S. Department Precipitation 1960, Smith 1966, tridentata). of low-growing west with Erequ errt high velocities, Precipitation (Beetle vermiculatis), (Purshia vis cidula) , (A. nova) occupied limited snakeweed (Gutierrezia primarily bunch grasses (A. ~_ Other Irnpo r tant shrubs (Sarcobatus and bitterbrush consisted silver were favorable and Smith 1978). (Artemisia 900/0 of the sagebrush approximately and black sagebrush where soil conditions Terwilliger Big sagebrush Alkali (A. longiloba), coaltown (A. argilosa) areas hay meadows. by sagebrush- throughout in winter and spring. 23.30/0 of the annual of Commerce 1973). the spring (Table 1). �Table 1. Spring precipitation and temperature, Walden, 1978 -79. Mar 2.01 2. 16 8.03 1. 57 13.77 -1. 3 3.1 506 11.9 1979 3.58 1. 24 3.46 3.15 11.42 ~3,8 1.6 6,3 11.2 30-year avg.c 1. 27 L 83 2.59 2,82 8.51 -4.6 L8 7. 1 11. 6 a 1975 b -- "u.s, b c 'f Mean daily temperature Precipita!ion Apr May Year (cm) Jun Colorado, (C) ---Totals Mar Apr May Jun ~-....... -~~ ., Department of Commerce (1978). U.S. Department of Cornm e r ce (1979). U.S. Department of.Cornm e r c e (1973). I..n •.....• �52 Most of the study area is usually following snowstorms temperature snow-free during March, by mid-March April and May. Mean annual is 2. 5C, varying during the study period from March to 11. 6C in June. The average is 46 days from mid-June to early August. Spring precipitation delaying access both years. Junction lek occurred frost-free Late snowstorms Permanent access -4.6 in season in Walden was 61. 8% above the 30-year 1978 and 34.20/0 above in 1979. average in occurred frequently to Boettcher Lake on 22 March 1978 and 15 April 1979, and to Alkali Lake lek on 30 March 1978 and 30 April 1979. peratures except were approx;imately 2. OC higher Spring tem- in 1978 than 1979. �53 METHODS AND MATERIALS Sage grouse along roads. at night while roosting on leks and Most b ia-d s were located using a spotlight with a back- pack or hand-held (Pyrah were captured power source and captured 1959. Braun and Beck 1976). (Lacher and Lacher and plastic. (1979). birds. A vehicle-mounted 1964) was used on 1 occasion. banded and released leg bands (size with long-handled were marked 16. National colored cannon net Sage grouse serially Band and Tag Co, , Newport. bandettes Weights and primary with aluminum coded to the individual molt were determined Age and sex classification of captured nets numbered Kentucky) (1978) or year for all captured birds followed Eng (1955). Selected transmitters Center) males were equipped with 14-15 g. 164 MHz radio (U. S. Fish and Wildlife Service. in 1978. Some transmitters to 14-15 g and 21-25 g transmitters (Champaign, Illinois), respectively. of the birds' attached Illinois), to the tail clip described were reused obtained m Transmitter (range -- rec~~ces packages 2440-3460 1979 in addition from AVM Company and Wildlife Mat e r i a.ls , Inc. body weights to the central Denver Research (Carbondale, represented g). 0.4-0.9% Transmitters using a bolt and clarrrp device by Bray and Corner (1972). were similar �54 Counts of male and female daily supplemented with periodic Wildlife personnel 1976). sage grouse on leks were made counts by Colorado following prescribed procedures Division of (Braun and Beck Counts were made from 23 March through 30 May 1978 and 26 March through 4 June 1979 between 0430 and 0730 MST. Radio -marked each morning Carbondale, males using a portable Illinois) males was attempted. 3-element Birds Center). as being off -Lek, leks were located completion located periodically ascertain at 1 -2 hour intervals daily activity coverage. sagebrush patterns. Slope, canopy coverage with Selected throughout aspect. and height, If visual location Radio-marked of counts. locations Fish of radio signals. than 0.1 km from of a lek were recorded after Their to be on a lek, located more Inc., yagi antenna (U.S. by triangulation were determined with lek counts (Wildlife Materials, Denver Research to a lek were classified radio-marked concurrently receiver and hand-held and Wildlife Service, respect were located the periphery males males not on were chosen days to total vegetative - and snow coverage .:»: were-recorded Locations colored analysis. on a standardized .:»: in 1979 were marked form at the time of location. with 1.2.m pka s tl c su r vevo r ' s tape to facilitate Locations U. S. Geological to the nearest surveyor's relocation were .:plotted on 7.5 minute Survey topographic O. 1 krn. (scale lath and for vegetation = 1:24,000) .- maps with movements-measured �55 Vegetative using Canfield's examined, located site. A steel between directions. to 4 transects 1-3 central were examined depending sagebrush. was recorded chi-square roosting nificance analysis locations. (1979) measuring in 1978 and 15.0 at each location 3-4 transects which were to the nearest analyzed recorded by compass In i.n 19790 except on leks. from All vegetation according as 1 group. to Vegeta- 0.5 ern for big and silver and rabbitbrush. with the Student's !_ test was used to test for differences and direction was P = O. 050 tape was originating 0.5 em along transects black greasewood, The data were into the ground at a randomly upon size of the lek. to the nearest except for grasses tive height using In were made At ea ch location rod and a 2nd rod located was 15.2 was analyzed points was measured species (1978) or plastic the central locations technique. rod was driven Each transect Lek vegetation for radio (1941) line intercept a 1. 5 m steel stretched Three cover measurements of movements. except that in lek attendance, The level of sig- �56 RESULTS Trapping Male sage grouse transmitters were captured from 25 February through 4 May 1979 (Tables were related Capture roads and Transmitter efforts through resulting Junction leks in 1978. (Bighorn lek), Access resulted Wattenberg Late capture along Jackson Bighorn capture leks resulted in trapping to intensively both years storms. County (J. C. ) and Boettcher Lake on the Bighorn Ranch efforts 2 lek and along J. C. 9A (Fig. and the decision dates from late winter difficulties in shifting accumulation Junction 1 May 1978 and 21 March were concentrated 7 and 7A and on Alkali Lake, include and equipped with radio 2 and 3). to poor access Life in April 2). monitor effo r t s being reduced 1978 to Heavy snow birds on fewer to Boettcher Lake lek and along J C. 7 in 1979. 0 "_ Thirty-seven male equipped with 35 radio class and capture sage grouse transmitters site stratification caught on leks and 6 adults and 3). season. Few males (16 in 1978, 19 in 1979). ~ere: and 4 juveniles were Le ca t ad roosting This resulted clas s in the sample. (17 in 1978, 20 in 1979) were in raw representation 14 adults Age and 13 juveniles caught off leks (Tables off leks during 2 the breeding of that capture site �Table 2.. Capture data and t r an s rrri tt e r life for male Band nurnb e r Capture date Juveniles ---7035 7036 7043 7044 7051 7101 .\ 7147' 7183 25 2.2 3 3 5 ,II '22 29 Feb 1\la r Apr Apr Apr Apr Apr Apr J,C, 7 J, c. 7 Alkali. Alkali Boettcher J. C. 9A Boettcher Alkali Adults 2543 6759 6907 6943 7037 7039 7040 70·12 7190 4 29 14 31 28 29 31 31 1 Apr Mar Apr Mar Iv1ar Mar Mar Mar May _ Wattenberg Boettcher Boettcher Alkali J. C. 7 Boettcher Alkali Bighorn Boettcher a II Transmitter Capture location _.__ lost Date last located sage Transmitter life (days) Mal" Apr Apr Apr May Apr Jul Jul 76 27 19 24 53 20 89 a 129 8 Apr 3 May 9 Jun 22.May 12 Jun 25 Apr 27 Apr 8 Jul 5 Aug 4 67 66 52 31 14 13 21 27 24 8 8 by 7040 and reused on 71-83. g r ou s e , North 81 33 a 99 96 Park, Colorado, 1978. R ern a rk s Experimental bird Moved from study al"ea Moved from study area Moved from study area Shot 9 Sep 1978 Shot 9 Sep 1978 Shot 9 Sep 1978 Vl -...J T'r an arni tt e r failure Intermittent transmitter Moved f r orn study Killed (coyote? ) area; killed by eagle �Table 3. Capture data and transmitter life for male . Band number Capture date Juveniles 7297 7303 7310 7325 7391 7402 7407 7408 7410 10 17 20 24 24 1 4 4 4 Apr Apr Apr Apr Apr May May May May J. C•. 7 Boettcher Boettcher Boettcher Boettcher Boettcher Boettcher Boettcher Boettcher 16 29 9 17 13 9 17 17 13 Apr May Aug Jun Jul Aug Jun Jun Jul Adults 6338 6929 7053 7294 7296 7298 7304 7305 7308 7409 7411 7 17 10 9 10 11 17 17 18 4 4 Apr Apr Apr Apr Apr Apr Apr Apr Apr May May . J. C. 7 Boettcher J. C. 7 J. C. 7 J. C. 7 J. C. 7 . Boettcher Bo ett che r j'. Bo e tt ch er Boettcher Boettcher 10 9 25 13 14 20 13 9 31 9 17 Apr Aug May Jul Apr Jun Jul Aug May Aug Jun Capture location Date last located sage grouse, North Park, Transmitter life (days) Colorado, 1979. Remarks ':' 'I', 1 aTransmitter used on 6338 and 7296, 8 48 Transmitter failure Int e rm.i tt ent tr an srnl tte r III 54 80 90 44 44 70 a 114 47 94 a Il 73 87 114 43 87 44 Shot 8 Sep 1979 Vl 00 Killed by eagle Moved from study area; Transmitter failure Moved from study area; shot 15 Sep 1979 shot 20 Jun 1979 �59 Transmitters functioned an average from 4 to over 129 days (Tables were operational of 64.3 days, 2 and 3). beyond termination Fourteen transmitters of the intensive periods (9 June 1978. 17 June 1979). located from mid-June These birds ranging research were periodically through August. Breeding Activities Lek Attendance Counts of male and female Lake John study area through were initiated 30 May 1978. on 4' active 7 April through 4 June. 7 April declined on 23 March rapidly Peak female thereafter attendance attendance -_ (Figs. a situation and continued were made occurred Female from 3 to attendance 3 and 4). previously Lower reported 1963), Eng (1963), and Gill' (196"'5). Late in 1979 was related Peak male attendance on the 3 largest in both years 4 to 5 weeks later, by Dalke et ale (1960, on leks in the 107 counts were made on 3 leks from 1978 and about 20 April 1979 (Table 4). peaks occurred female present During this interval204.counts In 1979, leks. sage grouse occurred 25 to 37 days after leks studied (Table 4). attendance observed both y~ars late spring situation descri.bed to extensive (Figs. snow COver. the female The rapid increase 3 and 4) was similar by Jenni and Hartzler (19T8). peak in male to the �Table 4. Peak counts of sage grouse on 4 leks, North Park, Colorado, 1978-79. Males , Lek No. 1978 Date( s) -- No. 1979 Date( s) -----1978 ------No. Date( Females--_ ..•. _-s) 10 May 72 27 May 20 3 Apr a Bighorn 44 29 Apr 31 8 May 19 4 Apr a Boettcher Lake '~unqtion 84 1,3 May 19 May 84 4: Apr 120 Wattenberg 2 21 26 May 7 7 Apr b aNo access prior to 30 April counts discontinued Lake 107 . 1979 ._---------No. Date( s) 56 Alkali __ 20 Apr ()\ blntensive b 1979. after 1978. 0 �61 MALES FEMALES RADIO- MARKED JUVENILE MALES RADIO- MARKED ADULT MALES _._ en UJ C) ~ a:: iLl > Cl ~ 0 I 10 !&""-~--, 100 en ~ .I.... \ iLl ..J , l 0 en LU _J c( ~ iLl IL. •••••• .. I . \ :.. .• of' i \ ~ I ':: "i: 0 ;Z q: . '.. it: : ..•/ -c.:« ~ o· I ' \ :\ fit •• :\ J ~ \ :.\ . ~IOO ~ . -. /. \ i 2: I~ .\ ~ a LIJ l! Il: :.\! q: :i ~ I 0 0 ~ a:: .. en LU ~ IL. ' a:: w CD :1 :i :::> l /. ~ 0 I- j:" <3!C U 0 ...J \ I _J <\ 9 "l " z 50(1) Z 0 I- , ,. , 0 0 . .. .. . .. . .•.... • r -•.....••• :.I 50 u, ~ IU \ _J '\t . I , \ Z 0 •... , Z IAI U \, at: c, / , O~I---~---~~r-~-----r----~--~~----r----~----r-~==~O / / .: 21 2S MARCH Fig, 3, UJ 6 Sage grouse 1978, II IS APRIL attendance 26 on 2 leks, 5 II 16 MAY North Park, 26 Colorado, �62 MALES --FEMALES •••••• RAOIO- MARKED JUVENILE MALES _._ RAOIO- MARKED ADULT MALES U) UJ (!) «« a: UJ > cl >- cl , C 10 r·~· · .: ; 100 U) ~ 1 .. UJ _j i.: Z 0 i : . tn _. UJ 1 oct .a.... ., r>; ..•......•..•....-,(./ .~. .. ". I ". .: '.: ••• ~ LIJ u, , '''''. . o • <: • • e. o .. • 0 o 2 ct U) UJ ..J ct :E '. OJ . 50 ~ ~ 0 a:: \ \.. \~ lLJ CD :::£ ;:) z ... \ . _J ct I- 0 I- 6 Fig. 4. II 16 APRIL Sage grouse 1979. 21 26 6 II 16 21 .. .. 26 MAY attendance on 2 leks. North Park. Colorado, �63 Data from 33 birds (16juveniles evaluate daily attendance nated from analysis (7039). premature patterns. because failure ment from the study area after late observer arrival leks were monitored locations not determined because on the lel< prior for analysis dates both years. were possibly of access known to visit a 1ek prior for were attributed increased 11-20 April 1978 (Fig. observations 5). on leks; diffi.cul.tl.es , that numbers exact locations Neither from 41.7 to 83.3%. (Patterson attending bird was a lek supports other leks increase 1952:153-154, Thirty-seven were from 1-10 to 1963, Eng 1963, Gill 1965, Jenni and Ha r tz l er 1978). was 81.00/0 for 21-30 Aprill978. off Lek attendance (P < 0.05) of juvenile males following peak female attendance captured Two adults visited This increase in March Only 2 of 16 (12.5%) 4 locations from 26 to 31 March 1978 (Fig. 5). of 6 juvenile males to my arrivaL Two juveniles to 30 March. to of lek attendance from 22-31 March 1978. early morning depredation after 4 days (2543) s : and move- i1 off-Iek locations or disturbance of late capture were elimi- 1 to 4 days (7037. 7043) (Table 2). , Few data were available because Data from 4 birds of injury and subsequent transmitter Data were also eliIninated and 17 adults) were used to Dalke et al. Lek attendance of 50 (74.0%) loca- tions were on leks during 29 days of observations. ~Attendance of juveni~_s increased from 1-15 May and decreased (P > o. OS) to 95.10/0 (P < 0.05) to 53.60/0 in late May 1978 �( ) CJ m (1)100 'It ~ NUMBER OF LOCATIONS JUVENILES ADULTS NO LOCATIONS w _J z o 80 (/) z o t- c« o 60 s o- .p.. ~) , _J <{ •... 40 o ~ LA- O z 20 1- w U Q: W Q.. o · I{I'( "9 ~ ";""-1 21 26 L l! I.LA.L.:_~_~~.A--i..L...a...L-..f.L.-AA--~ (I 6 II ,16 1 21 26 APRIL' Peak of Femoles Fig. 5. Lek attendance '---f-U!6 (' 6 II t.: 6 ~ . VA t' flt..L l1n_ __ 21 26 MAY 1 JUNE Peak of Males of r a di o crn a r k e d male sage grouse, North Park, Colorado, 1978. 6 �65 (Fig. 5). However, during 16-31 May I was absent area for 4 days and a late snowstorm 2 additional magnitude days (Fig. 3). prohibited These factors of the decrease. Average for 3 radio-marked attendance (P decreased (50.0%) locations was Lek attendance locations in April decreased juveniles. observation. adults of 6 radio-marked increased Attendance lek Only 1 of 2 adults was 71. 9% of 96 per 5-day interval (!:_> ranged 0.05) (Fig. 5). (P < 0.05) to 100.0% in 1-15 May, and of 5 birds Lek attendance in early June. Juvenile on a lek. 1978. 87 (93.1%) locations the for May 1978 was > 0.05) during 1-10 June. < 0.05) to 80.6% thereafter (P to 1 lek for may have influenced from 62.5 (6-10 April) to 80.00/0 (11-15 April) Adult attendance access attendance 78.3% of 69 locations from the study (Fig. 5). Eighty-one of were on leks during 27 days of decreased rapidly None (0 of 2) was present (P < 0.05). for on a lek during 1-10 June. During the 5 -day period in 1978 (1 -5 April), adult males, 50.0 and 70.6% of radio -marked respectively, and 94.2% of the juveniles on leks during (26 April-lO which included peak ferfiale counts were on ~eks (Figs. and adults, juvenile 3 and 5). respectively, 4J. Ninety-two were present the 15 -day Eeriod which included peak male May) (Table and counts �( ) NUMBER OF LOCATIONS D JUVENILES (Z1) ADULTS NO LOCATIONS * 100 en ~ W ...J Z o en 80 z o ~ 60 o- o s '" _J ~ 40 o t- IL o z 20 t- w o ~ W 0.. 1 o * 'j i 1~~t:~l!)J!il1 [ILll I.} 6 III 16'. APRIL (paak Fig. 6, Lek attendance VJ 21 A....Jp J!. 26 ¥-11). III II ~L!~a 6 II of Femcles of radio -rn a r k e d rn a l o sage MAY ( ,L 16 21 41 fllih, ,l\it!ji 26 Peak of Moles g r ou s e , No r th Pad" Colorado, 1979. �67 Juvenile radio transmitters tively. (P and adult male sage grouse and monitored Lek attendance were not equipped with until 10 and 7 April of 5 radio-marked juveniles 1979, respec- increased < 0.05) from 33.3 (11-20 April) to 100.0% (21-30 April) following the peak of female averaged attendance in 1979 (Figs. 87.90/0 of 33 locations Attendance of 8 juveniles May and increased Lek attendance (P during decreased (P 4 and 6). 16 days of observations. < 0.05) to 73. 1% from 1-10 < 0.05) to 91.1% during 11-20 May (Fig. < 0.05) for 21-31 May. was 78.8% (P dance for May 1979 was 79.3% of 169 locations Nine adult males Lek attendance (16-30 April) throughout (P> April. 6). 6). Attendance Lek attendance obtained (P attendance of 7 radio -marked increased adults (P > 0.05) lor 153 locations (P < 0.05) for both in early Two of 1.0 (20.0%) locations 1-10 Juvenile June (Fig. and adult males 100.0% of the locations, rapidly 5% during age classes on leks during to 98.0% s r, was decreased were on leks. decreased < 0.05) to 82.7% from 21-31 May Lek attendance for 7 juveniles 1979. 77.4% of 62 locations May 1979. June. atten- on 28 days. on 17 days in April averaged 1-'10 May, and decreased Average Average 6). < 0.05) from 0.0 (1-15 April) to 100.0% (P 0.05) to 94.0% from in 11-20 May, (Fig. were monitored increased (Fig. Lek attendance None of 8 locations recorded of 4 adults was 6). were present respectively, during on leks for €b. 7 and the peak of female �( ) NUUBER OF LOCATIONS [=:J 1978 fZ2'J 1979 en 100 II) un (~l 16. 1;1 * NO LOCATIOtJS ~ w __J z o 80en II z o ~ 60 o o .J (2) _J 12. <t •... 40 o lLL. o ;- z 20 w U 0:: W n, (5) "~ -10 'fIll ~O -5 In! It4! 5 10 Peck of Females 1(4_' ,,15 [Ell 20 V(I. ..L-fLL' 25~30 Peak of Males DAYS FROM PEAK Fig. 7. Lek attendance 1978-79. rOt fn-'-I/4! 35f-... 40 of radio -rri a r k e d juvenile FEMALE F.cal t'..,., 1&1"" I Second Peak of Females ATTENDANCE rn a Io sage grouse, No r th Park, Colorado, f~ 65 o- co �4) 18) 10) NUMBER OF ( )LOCATIONS 01978 [ZZl1979 en ~ w _J '" z NO LOCATIONS o 15 (/) z o tio g ..J g 0\ '-" ~ u, o ~ ~ w u a:: w a. o 'f: I r-f... 1 -10 ~u_p.! -5rO ~Peak o~ 1-1(0 5 j: fLAI 10 15 DAYS 8. Lek attendance tLLI 20 I -'I F (41 f..LA 25t::'t> 30 35 40 \Pea~ of \Second l Q, ~-~---- 1 Malas Females 1/ Fig. fUll 50 45 55 60 65 Peak of Females F~OM PEAK FEMALE ATTENDANCE of radio-marked adult male sage grouse, North Park, Colorado, 1978 -79. �70 attendance (16-20 April which included juveniles peak male and adults, The 2 years which include April after respectively, attended the peak of female 7 and 8). 89. 7 and 93.70/0 of the leks (Figs. by superimposing attendance (1-5 April During the period were on leks. (93.9%) attended the female 10 May 1978, atten- (90.4%) and (25 -40 days counts (26 April- 11-25 May 1979). Sage grouse males captured while roosting attended leks during captured off leks and radio-equipped. the breeding prematurely prior moved from the study area and 2 birds locations. Six adult males (6338) was killed season. to the beginning thereafter, originally peak male 16 -20 and adult males, the 15 -day period peak) which included 1978, of peak female Over 90% of the juveniles leks during 4 and 6). the 5 -day periods 55.6 and 79.2% of the r adto vmaz-ked juvenile respectively, adults During the 15-day period counts (11 -25 May), were compared 1979) (Figs. dance, 1979) (Table 4). after (7036, by a raptor (7037, locations. Three 7053, Four juveniles One transmitter of lek attendance, 7101) visited were captured of these males were (7497) failed 1 bird (7035) a lek 2 leks £or~9 of 20 (45.0%) along roads. while moving 7294, regularly 2 days and may have visited banded in 1975, 1 transmitter and 4 birds off leks towards One bird the lek where (7296) failed prematurely, 7298) visited leks for 19 of 30 (63.3%) moved to leks not in th'e=-study area. �71 Movements Movements males between leks within the study area were common (3.9 each) but were infrequent each). No differences juveniles had up to 13 movements than 6 in 1979 (avg. outside > 0.05) occurred (P by juvenile for adults between years although (avg , 4.4) in 1978 and no more 3.6) •. This does not include moves the study area (0.6 and possible moves to leks on 5 days in 1978 when data were not collected. Interchange between leks by juveniles as the breeding season more of number the result 11 in May) than large leks visited visited (P (Fig. of birds changes increased Each juvenile progressed (P < 0.05) increased 9). This may have been equipped with radios in movement patterns. (8 in April, Number of < 0.05) from April (1-2) to May (2 -4). from 2 to 4 leks (avg , 2.8) during th~ breeding season. Distances and averaged between 5.0 krn leks in the study area (Table in t er Lek move was 4.0 krri, Alkali Lake and Boettcher movements), 5). The average Moves commonly ranged from 2.4-8.1 di s tan ce per juvenile occurred between Lake J'uncti on leks (49.0% of all interlek Alkali Lake and Wattenberg 2 (17.6%), and Boettcher -_ Lake Junction ments and Bighorn__(15. 7%). The number between Alkali La ke.-a.nd Boettcher pected because shorter distances of interlek Lake Junction and topography move- Was unex- should have increased �12 ( .) ( ~IO ) z CJ :E rJII] W W NUMBER OF RADIO - MARKED JUVENIL.ES 1978 1979 > ::E o 8 :.::: W _J 0::: .,_w6 -...J z N u, o 3 0:::4 w (D ::E :J Z 2 .0 1I I"" 26 1'1 MARCH Fig. 9. \""1''''' I ( (OVID) 6 (2',(0) /.\ I '. (2)~.4) II ·16 APRIL 21 ·F.1 26 fLO! I Time distribution of int e r l ek rno vern e nt s by juvenile Colorado, 1978 -79. 1££11 6 male IliA! 1'/41 II 16 MAY sage grouse, fUAL 21 North t UA 26 Park, �" Table 5. lnterlek I " distances (krn) Alkali Lake No. of rn o v ern cn t s Juv. Ad. Distance between and sage grouse rno vern e nt s , Lake Bighorn Distance between No. of rnov ern ent s J'uv, Ad. John area, No r th Park, Boettcher Lake Junction No. of Distance rn o vern e nt s J'uv , between Ad. Bighorn 5. 1 2. 0 Boettcher La.ke Junction 4.5 25 0 2.4- 8 1 Riley 2.7 4 2. 6.9 0 0 5.5 1 0 Wattenberg 2 3.2 9 2 8. 1 1 0 7.6 1 2 17 I; Colorado, 1978-79. R il c y Distance between 4.2 No. 0[-rri o v cin on t s Juv.~d:- 0 0 --..J w �74 movements between Alkali Lake-Wattenberg and Boettcher Lake Junction-Bighorn. change is unknown but may be related averaged at least The r ea s on for this interto lek size (Table 4). 15.6 krn (range 2.4-58.7) during the breeding (range 5.1-60.8) (Table 6). (P 2, Alkali Lake-Riley, season. Juvenile throughout Mean distance in interlek males the breeding Juveniles movements moved at least 23.9 km season for both years per move was 1. 9 krn , No difference > 0.05) existed between years. Juveniles attended 1 -36 non-consecutive with off-lek days. visit was 2.8, from The average with a maximum of capture throughout a 2nd lek in late May. remained to that lek were intermixed number of consecutive of 15 days. of individual between 2 leks (Boettcher Because days per of my absence a lek for over 20 sedentary most juveniles and remained of the breeding season, Three the lek £i~ally moving between to 4 leks and Other individuals these ext r ern e s including Lake Junction varied. on or near One bird moved 13 times on each lek for 3 days or les.s. intermediate ranging from 22 -37). movements were extremely visits of 4. 3 days, 3 birds may have visited days (range Interlek birds days where the study area, consecutive a lek an average were 1 bird which visited and Alkali Lake) in 1 morning 4.5 km between 0510 and 0610 MDT on 22 May 1979. moving �75 Table 6. Distances (krn) traveled by male sage grouse during breeding season, North Park, Colorado, 1978 -79. No. a moves No. birds the Mean distance per bird Mean distance per move 187.8 26.8 2.2 21. 7 b 23.9 1.8 Total distance Juveniles 87 1978 7 1979 9 III 195.0 16 198 382.8 Subtotals 1.9 Adults / 1978 0 84 99.7 16.6 1.2 1979 10 78 60.4 6.0 0.8 Subtotals 16 162 160.1 10.Oc 32 360 542.9 17.0 Totals alncludes interlek bNo difference movements. between years (P (P > 0.05). < 0.05). cDi£ference between years dDi££erence between age classes (P < 0.05). d 1.Oc 1.5 d �76 Only 3 of 11 (27.30/0) adult males the breeding visited more than 1 lek during season. Two adults (6943, leks for only 1 day. One of these visits attendance One other adult male (6929) displayed to a lek. lek where marked (Boettcher and then attended Btgbo rnLek, Adult males between season 1978, 0.8 k:m in 1979). This was attributed Twenty-two distance movements percent 1. 5 km (Table 7). 11. 1 and 8.8% of all o££-lek locations respectively. Over 50% (55.3) and 1. 00 krn from a lek. 3) < 0.05) existed Differences (P < 0.05) per move (1. 2 krn in than that of juveniles. (58 of 198) of relatively by juveniles. of all off-lek locations Off-lek distances (P 0.6-31. per adult movement (P < 0.05) to the large mnnber long (> 1.5 km) interlek 0.26 Differences for mean distance less through 15 May 1979 of 10.0 km (range (Table 6). Average (1. 0 km) was considerably on the r 1978 (16.6 krn] and 1979 (6.0 km) , also existed between years 1-2 additional followed 12 days of non- Lake Junction) moved an average during the breeding 7190) visited were beyond beyond 2.0 km comprised for juveniles and-adults, of 313 off-lek locations were between Only 7. 31!/0 of the locations were within 0.25 km of a lek. Long distance for 4 juvenile a minimum move:rpents from the study area and 3 adult rna.l e sage grouse. were recorded Juvenile 7031; of 25. 7 km in 5 days to the Spring Creek area moved southeast �77 Distances from leks to feeding-loafing sites grouse, North Park, Colorado, March-May, Table 7. Distance (km) Juveniles No. a 0/0 Adults No. of male sage 1978-79. b Totals 0/0 No. % 0.01-0.25 9 5.9 14 8.8 23 7.3 0.26-0.50 44 28.7 58 36.1 102 32.6 0.51-0.75 9 5.9 18 11.2 27 8.6 21 13.7 23 14.4 44 14.1 1.01-1.25 14 9.2 11 6.9 25 8.0 1.26-1.50 13 8.5 11 6.9 24 7.7 1.51-1.75 9 5.9 6 3.8 15 4.8 1.76-2.00 17 11. 1 5 3.1 22 7.0 2.0+ 17 11.1 14 8.8 31 9.9 153 100.0 160 100.0 313 100.0 0.76-1. 00 Totals aDistances measured from nearest lek. bDistances measured from normal lek attended. �78 of Walden. tion. Two juveniles Juvenile left the study area 5 days after 7043 moved 2.5 km north of Walden Reservoir 6.0 krn east of the study area, 6 km south of Wattenberg Juvenile about while 7044 moved approximately 2 lek. This bird then moved north to Alkali Lake lek and then to near Bighorn lek before lost. instrmnenta- 1st signals were 7035 moved a rnirrirnum of 80 k:m. between 25 February and 31 March 1978. Three of 6 adults the study area. captured off leks moved to leks outside Bird 7037 was captured J. C. 7 (Table 2). From after 12 k:m. southwest Bird 7294 moved to Cheyenne lek, 15 April, 6 May. 1978 along 31 March to 9 May this bird was near Aspen lek west of Delaney Butte, area. on 28 March of the study 13 km south-southwest, while 7298 moved to Migan lek, 21 km south, Both birds of were captured by along J. C. 7 in early Apr i.I 1979 (Table 3). Daily Activity Patterns Selected intervals ..: transmittered throughout during the breeding birds were monitored at 2 -3 hour chosen days to document daily activity season. Three to 7 locations patterns were ascertained for 2-4 rn.af e s between 0500 and 2120 MDT on 21 and 24 May 1978, and on 13, 21, 25, 27 and 30 May 1979. Activity patterns were also .::=- available for juvenile and adult males when 2 -3 daily locations obtained from 23 March to 9 June 1978 and 8 to 21 May 1979. were �79 Juveniles later present on a lek in early morning in the day on 13 occasions. from 0.4 to 3.2 km (avg. Distances 1. 2). were relocated from the lek ranged Midday moves Over 0.5 km were uncommon as only 6 of 22 (27.3%) were beyond 0.5 km. move during midday was 0.3 km (range < 0.1-1.4). average Juveniles Undisturbed moves travelled birds as far as 3.2 km when leaving subsequently over 1.0 km usually may be more mobile This was observed (P The moved only 150-300 rn , resulted from disturbance. during the day early with 1 juvenile Juveniles or intermixing remained averaged occurred in a particular Midday Juveniles in the breeding season. 2.1 krn (range < 0.1) between morning and afternoon locations March. a lek. 0.8-3.7) from 23 to 31 flock until disturbed at a lek with subsequent dispersal changing flock composition. Adult males Dispersal irom difference (P Subsequent adults. were relocated leaving a lek on 18 occasions. 1. 1 krn (range < 0.1-2.9). the lek averaged No > 0.05) existed between adult and juvenile dispersal. midday moves Adult male that of juveniles, with subsequent over 1.0 km. after averaged sage grouse 1. e., had a ct ivi ty patterns relatively short moves. 0.2 km (range < 0.1-1.0) long distance Disturbance similar dispersal usually for to from a lek resulted in moves �80 Daily dispersal (P < 0.05). males Forty-one dispersed Junction from a lek was non-random percent southwest percent to south-southeast 23. 1% west to southwest. in the same direction between Reasons daily. for the directions Non-random adult males after of dispersal of sprayed dispersal available dispersed for dispersal and to disperse km south of the 2) for 4 days o. 1-0.5 km east 0900 MDT (8-17 May 1979). are unknown but may be related (Fig. 2), association with other areas. was also observed lek as 85.7 and 72.7% of the radio-marked respectively. appeared 10). dispersed 23.1% east to southeast, Adult 7409 was located other leks in the area Lake (Fig. juveniles km west of J. C. 7 (Fig. on 6 of 7 days located and location to southeast Bird 6907 moved 0.4-0.5 14 and 21 May 1978. to topography, males, Individual to moving 0.1-0.6 to northeast northeast Lake Junction, adult from Boettcher (8 of 26) of the radio-marked south from Boettcher Lek prior (24 of 58) of the radio-marked lek while 39.7% dispersed Thirty-one in direction to the south (Fig. from Alkali adult and juvenile 11). Lake males, Few data were from other leks in the study area. Roosting Roosting locations roosting (after birds data were obtained 1700 MST). because out the field season. ..-'- from capture Capture trapping sites sites and evening were biased was most successful towards lek- on leks through- �81 o JUVE N ILES ADULTS s Fig. 10. -Di sp e r s a l of male sage grouse from Boettcher Lake Junction lek, North Park, Colorado, 1978 -79. Each concentric circle r~presents 0.5 km. Number of locations at each site is irid icat ed (no number indicates oney 1 iocation). Locati.ons (N 12) beyond L 5 km are not shown. = �82 N o JUVENILES • ADULTS s Fig. 11. __Dispersal of rn a l e sage grouse from Alkali Lake lek, North Park, Colorado, 1978 -79. Each concentric circle represents 0.5 km. Number of locations at each site is indicated (no number indicates only 1 location). Locations (N Z) .~ beyond L. 5 km arc not s iio wn, = �83 Twenty-nine males were on leks (Table 8). in April leks. (84.6%), Highest locations on-lek with 60.7% of the roost Adult males obtained May, of 48 (60.4%) roosting roosted 2 age classes of the roost sites occurred in May being on sites on leks in April (P > O. 05) existed No difference between and the (P < 0.05) when 16.7 and 87.5% except during March sites roosting on leks 69.8% of the total locations with 55.0 and 78.1% of the roost respectively. for juvenile of juveni.les and adults, respectively, were on leks. Table 8. Month Roosting sites of radio-marked Park, Colorado, 1978-79. On-lek Juveniles Off-lek male Totals sage grouse, On-lek Adults Off-lek North Totals Feb 0 1 1 Mar 1 5 6 7 1 8 Apr 11 2 13 12 10 22 May 17 11 28 25 ..•.7 32 Totals 29 19 48 44 19 63 Vegetation Sagebrush obtained Analysis height and canopy coverage at 160 daytime feeding and loafing rneasuremenJ.s si.tes, were and 88 roosting �84 sites. Measurements for all locations, of forb and grass including Approximately occurred were obtained the 5 leks. 56% (56. 3) of all feeding and loafing sites in sagebrush Sagebrush coverage with a canopy coverage canopy coverage averaged 28.1% of 20-50% (range (Table 9). 2.9 -68.8) for 160 sites. Table 9. Frequency distribution of male sage grouse feeding-loafing and roosting sites by sagebrush canopy coverage class, North Park, Colorado, 1978-79. Feeding-loafing Number of locations Canopy coverage cla s s" Roosting Nu:rnber of locations sites % sites b. % 0.1-10.0 25. 15.6 75 85.2 10.1-20.0 30 18.7 4 4.5 20.1-30.0 31 19.4 1 1.2 30.1-40.0 42 26.3 6 6.8 40.1-50.0 17 10.6 2 2.3 50.1+ 15 9.4 0 0.0 Totals 160 100.0 a b Expres sed as percent Includes 71 on-lek roosting Eighty percent were in sagebrush average of fully closed 8S-- 100.0 canopy. sites. of the daytime feeding and loafing locations with a height of 20-70 ern (Table rriax irnurn height for the 160 locations 10).·~he was 43.5 cm (range �85 9 -88). This may reflect the area instead actual of selection height distribution by the birds of sagebrush (Wallestad in and Schladweiler 1974). Table 10. Frequency distribution of male and roosting sites by sagebrush Colorado, 1978 -79. Sagebrush height a class (em) Feeding-loafing Number of locations sage grouse feeding-loafing height class, North Park, sites Roosting Number of locations % b sites 0/0 0.1-10.0 1 0.6 8 10.1-20.0 19 11.9 68 77.3 20.1-30.0 22 13.8 2 2.3 30.1-40.0 30 18.8 1 1.1 40.1-50.0 19 11.9 0 0.0 50.1-60.0 32 20.0 7 7.9 60.1-70.0 25 15.6 2 2.3 70.1-80.0 9 5.6 0 0.0 80.1-90.0 3 L8 0 0.0 160 100.0 88 100.0 Totals aMeasurement blncludes of height of tallest 71 on-1ek roosting Approximately living plant. sites. 90% (89.7) of the roosting locations 71 on leks occurred in sagebrush with a canopy coverage than 20% (Table The av'erage canopy coverage 9). tions was 8.7 and ranged 9. 1 from 0.4 (Alkali including of less for the 88 loca- Lake lek, Table 11) to �Table Plant 11. Vegetative species COver (%) of 5 leks, Alkali , Lake North Bighorn Park, Colorado, 1978-79. Lek Boettcher a Lake Junction . Riley Wattenberg 2 Averages Big sagebrush (Artemisia tridentata) 0.37 7.31 5.45 0.96 16.62 6.14 Fringed sagebrush (A. frigida) 5.28 1. 43 1. 82 2.73 4.82 3.22 Snakcweed (Guti~\'reiia 0.00 2.72 0.56 5.64 2.36 2.25 Winterfat (Eurotia sarothrae) 00 lanata) 12.32 0.10 0.66 0.36 0.59 2.69 Moss phlox (Phlox bryoides) 0.50 2.58 1. 22 0.31 3.84 1. 69 Bluebell (Mertensia bakeri) 0.00 0.29 0.02 0.00 0.07 0.08 Low Daisy (Erigeron pumilus) 0.00 0.11 0.09 0.00 0.25 0.09 5.57 6.01 6.28 11. 98 7.90 7.55 24.08 20.57 18.00 36.14 24.40 Grasses Totals b (/ aTwo separate blncludes areas additional of the lek were species cornb in ed to form not found on all leks. composite 23.23 values. '" �87 49.30/0. Eight of 17 (47.10/0) off-lek roosting with < 200/0canopy coverage. The average off-lek The large sites locations resulted and roosting was 22.40/0. in differences canopy coverage nurnb e r of on-lek for the roosting < 0.05) between feeding-loafing (P (86. 40/0)roosting than 20 ern in height (Table was 18.7 ern (range locations 10). 9-68) compared occurred Average in sagebrush sagebrush height with 38.7 em for the 17 off-lek (P < 0.05). roosts Sagebrush canopy coverage on leks. Grasses gracilis, Calamagrostis lettermani), lanata), (Agropyron fringed averaged montanensis, sage (Artemisia of 1. 00/0or more 24.40/0 (range height averaged averaged 6.10/0 (range srrri th ii, Koeleria phlox (Phlox bryoides), an average brush were in sagebrush sites. Seventy-six less sites and snakeweed coverage. 18.0-36.1) cristata, Poa secunda, irigida), Bouteloua and Stipa winter£at (Eurotia each contributed Total vegetative for the 5 leks (Table 15.3 em (range 0.4-16.6) cover 11). Sage- 9-27). Band Recoveries Nine radio-marked 25 April male 1978 and 15 Septew-ber 3) were depredated chrysaetos), sage grouse 1979. were recovered Three adults du r in gis pr mg , 2 by golden eagles and 1 possibly by a coyote (Canis (Tables (Aquila Ia t r an s}. between 2 and �88 Three transmittered in the Lake John area hunting season. In 1979, 1 juvenile on 8 September the study area wa s collected .' (Table 2) were harvested 1978 during the sage grouse and 1 adult were harvested One juvenile was shot in the Lake John while the adult was harvested on 15 September for parasite of the study area males on 9 September during the hunting season. area juvenile (Table analysis on 6 June 1979 • 3). 20 km south of One additional near Migan lek, adult 21 krn south �89 DISCUSSION Transmitter Transmitter days. life averaged 64.3 Life days, Five of 35 (14.30/0) transmitters problems due to premature temperature-induced Three additional Tail clip-mounted to eliminate restricted movement 1979) and excessive transmitters and Hight, or result (3) and cold slipped off the tail when preened. (Bray and Corner associated with back-pack of displaying mortality did not appear failure operational (2). transmitters transmitters problems experienced transmitter intermittency ranging from 4-129 males in high mortality harnesses (Emmons (Rothenmaier to adversely 1972) were used 1979). such as and Petersen Tail-mounted affect male strutting (only 3 of 37 males behavior were killed by predators). Breeding Seasonal lek attendance previously_reported patterns (Patterson Eng 1963, Gill 1965). techniques based on the patrern all males of sage grouse 1952:153-154, Patt-erson in a population Activities (1952:93-94) he observed regularly visit leks. have been Dalke et aI, 1963, recommended census and the assumption Jenni. and Hartzler that �90 (1978) confirmed testing the validity 'of the Patterson the a s surnpcion that all males et ale (1963) reported that relative males varied and all females Daily counts indicated higher regularly trends banded males leks for 1-3 days at a time. in their lek visitation, whereas. nurribe r s of male influencing the precision counts during stable of the high variation that not all males similar males attend leks. absent from were irregular were present 1963:820). in the on leks and noted problems of counts. when attendance changes because annual fluctuations These investigators of using lek counts to indicate because were ordinarily ••• " (Dalke et al. and accuracy Variation- in male was considered questioned in sage grouse in daily attendance Rothenmaier's the validity populations and the possibility (1979) findings to those of Braun and Beck (1976) as he observed between actual population census and_Lincoln-Peterson in no r thea s te rnWyom ing , been documented 190/0 Although no consistent sage grouse the 9 -day period was 24.30/0. technique "dominant Braun and Beck (1976) reported season. was actually Less dominant males almos t daily under all conditions maximum the breeding that the male population were observed, Dalke of adult and juvenile daily throughout did not attend leks daily. without attend leks. proportions than counts obtained by the Patterson some males most technique estimates Non-displaying for other lekking grouse were a disparity of the male mai~ have with only 500/0 of the males �91 present on leks when peak male Rippin and Boag 1974). obvious that the basic population According assumption attendance and 4) were similar Beck (1976), However, peak, recommended (P < 0.05) existed of the male in 1978-79 (Figs. reported by Gill (1965), (1978), and Rothenmaier attendance (Patterson and (1979). the female 1952:92-93). in lek attendance Braun 3 25-37 days after not during the 3 weeks following peak A difference of radio-marked adult males 1978 (74.6%) and 1979 (96.70/0) during the 3-week period following the peak of female 15 May 1979) (Fig. lek attendance 8). during attendance No difference this period (11-31 April (P 1978, 26 April- > 0.05) existed for juvenile in 1978 (82.1%) and 1979 (82.3%) 7). No difference for either female (P > 0.05) in attendance age clas s during attendance included the period (21 April-10 the male peaks (Figs. iavg. 84.8), (Fig. 7), and 89.4 and 93 •. 5.-0/0 (avg. 8). between 2 -5 weeks after 3 and 4). was 88.0 and 83.3% (Fig. existed years the peak of May 1978, 6-25 May 1979) which years males percent observed the peak of male for census "it is quite of peak lek counts is critical." patterns Jenni and Hartzler I observed the female (Fig. concerning to patterns (Robel1969, to Braun (1979:20), pres ent during periods Seasonallek between counts were obtained During this period, Lek attendance respectively, 92.0), respectively, for the 2 for juvenile for adult only I of 11 (9.1%) juvenile males �92 males was present was present hypothesis 1 (juveniles attend leks less for 5.6-6.2 the 2nd hypothesis days/7-day (adults attended locations for 2 juveniles, captured while roosting captured leks. than 3 days/7 -day period) visited leks daily while This indicates that possibly indicating breeding birds. off leks. Lek attendance Therefore. captured was 9 of 20 (45.0%) for 4 adults the 3rd hypothesis go to leks during season. the breeding season which did not attend Leks One adult male provided the temporary existence of groups Bird 6943 (Table 2) was absent 1978 while displaying Alkali Lake lek the remainder This period high (avg. of non- from a~lek between of the .period from 1 April to 2.2 May followed peak female when average 4.4 females/day). ."--_ accompanied data on Riley lek on 18 April and on of non-attendance (3 April) and was a period undetermined) sage grouse is rejected. the breeding 6-17 April male and 19 of 30 (63.3%) locations off leks rarely capture) throughout relatively Therefore. period. samples. I was unable to identify males 1.978. it and may be true. oi£ leks regularly of initial period; attend leks daily) cannot be rejected Although based on small (males days). Four of 10 (40.0%) adult males visited completely than 5 days;7-day on 8 of 17 days (3.3 days/7 is rejected. 6 adults on leks for less female attendance attendance remained Only 1 other maI~(age 6943 for 1 day during this 12-day period �93 m contrast to the flocks of non-breeding Robel (1969). adults) Other radio-marked did not attend leks for periods a temporary non-breeding Interlek in various reported ments portions more years. juveniles (juveniles In Idaho, than adults movements }?ymales Dalke et a1. (1960) were involved in interlek of males movements of North Park, ments ..of juvenile and Schladweiler Colorado, male 1974). during (1967) and May (1970) also noted interlek within and between years. were summarized Interlek interlek movements movements more moveCarr 0-'1 males of males in North by Braun and Beck (1976) who found that = respectively, movements. to be more visiting the banding year. (N were invo Ived in interlek movements Interlek Within the Lake John 51.9 and 14.6% of the banded juveni.les Interlek to in central birds Gill (1965) reported sage grouse in- and between 1 year. to be uncommon with only 1 of 13 (7.7%) radio-marked rrio ve distances on leks appeared in at least were reported than 1 lek (Wallestad and have been reported within the banding year in numbers with interlek by of 1-7 days and may represent Dalke et a10 (1963) reported correspond Park sage grouse sage grouse range. movements Fluctuations Montana, by male of their between years. reported male population. movements volved in interlek area male black grouse b.y juvenile common than reported 27) and adults and adult males (N = 48), wei=e found by Braun and Beck (1976). All « �94 13 juveniles remaining during the breeding bird. Therefore, in the study area season Three than 1 1ek during 0.6 (range 0-4) moves were unusual. Interlek size the importance day. Decreased (juveniles the breeding display season movements movements documented on more displayed (adults of display by adults in this study empha- of counting all leks in a complex attendance per for an average The 5th hypothesis although interlek leks 1 -13) moves of 11 (27.30/0) adult males per bird. on only 1 lek) is rejected 2 or more 3.9 (range the. 4th hypothesis than 1 lek) is accepted. on more averaging visited on 1 lek may be a result on the same of movements to other leks. Patterson of male (1952:92-93) sage grouse counts performed immediately between recommended in spring by taking the maximum on 3 different days during following the female i hour before (1978) presented and i hour from i accuracy, after period Rothenmaier to 2 hours. 3-week p e r i od following thereby sunrise. all birds until had departed Jenni and Hartzler Montana, lek Ii hours counts after sunrise ext end in g the dai.ly count (1979) observed the peak of mating, on 4 of 9 days had departed of 3 lek Counts are to be obtained hour before without sacrificing 1 occasion peak. populations the 3-week period data showing that in central could be performed males censusing that during at least i hours by 1 by that time. after the 50.0% of the sunrise and on This was attributed �95 to raptor predation the Patterson and Rotheronaier (1979) (1952:92 -9 3) method in areas reco:rru:n.ended using where raptors are c ornrrio n. Regressions against Park sunrise, of daily lek departure and data (Figs. when less hours after and Ii hours 12 and 13). sunrise sunrise. when less and 16 of 44 (36.4%) than 50% of the males of high raptor be obtained t within populations hour after Braun and. Beck (1976) during a 30 -day period during 21-30 April, Hartzler (1978) provided more within i occurred 8 Ma v, hour conclusions counts Ii that should sunrise. recorrrmend.ed counting leks 4 times 11-20 April, - 1-10 May. that increased 1 count Jenni and number of counts estimates. counts would have been obtained during 1978 but The count for the -I: Leksriri ve s t iga t ad in 1978 was 182 males Assuming Ii sunrise. not in 1979 using the t echn i qu e of Braun and Beck (1976). highest i within hour after (i. e. North Park) and 1 count during reliable within as all of the 18 occasions s (1979: 51-54) with 2 counts during also reported Acceptable within occasions occurred occurred were present support Rotheronaier' for the North 42 of 44 (95.5%) The 1979 data were similar results in areas sunrise were present and 7 of 18 (38.90/0) occurred. These after In 1978, than 50% of the males after hours, i times were calculated a count above 150 males would be acceptable on as �08001.5 HR AFTER SUNRISE w = - 0700- APPROXIMATELY 50% DEPARTED:: 0 1\ Y = 629.13-2.62X r -0.12 OVER 75% DEPARTED 0 1\ Y = 651.59-2.24X r -0.54 .- = = :E .._ o Il: 0600- c:{ o ~ ~ .aOO~ U> . --- . -- Z . -• 0500 ~ ~ o o Z ;:) o -. !"ooo , 0\ ~~ -- :E 0.5 HR AFTER SUNRISE 0400 '-0 "'_.'l.. ~r::__1liI.... ~ o \ •• o· tr(!j --._ 0300 { II 31 10 MARCH Fig. 12. Regression of sunrise North Park, Colorado, 20 APRIL and daily 1978. 30 departure times T T T 10 20 MAY 30 of rn a l e sage grouse f r orn 4 leks, �0800 w 0700 I _. - - 1\ 1.5 HR AFTER SUNRISE . I-=-:------___ Y r - . - Y •... a: = 598.54 = -0.67 DEPARTED - 2.04X - 0 0600 0 ~. = ftII = -1.00 -r DEPARTED = 698.79-3.90X OVER 75% 1\ ::!: <{ APPROXIMATELY 50% =0 _ ---- _ .... CJ) z I.D -...J o 0500 ~ z o :::> 0 ::!: 0.5 HR AFTER SUNf~ISE 0400 0300 { I( 31 MARCH Fig. 13. 10 20 APRIL 30 Regression of sunrise and dally departure 3 leks, North Pad" Colorado, 1979. 10 .-------- 20 MAY t irn c s of rn a l e sage grouse T 30 I r orn �98 the high count, during 20.0, 11-20 April, 30.0 and 70.0% of the daily counts obtained 21-30 April and 1-10 May 1978, respectively, were in the acceptable acceptable range (Table 12). counts would have increased 16-25 April, when 20.0, twice during 26 April-5 The probability by counting once during May, and once during 70.0 and 70.0% of the counts, 150 males. Higher counts increased 20 days of the count periods respectively, in occurrence were above during the last May) period. Nwnber of counts with acceptable numbers of male sage grouse using 2 techniques in North Park, Colorado, 1978. References Braun and Beck (1976) Dates 151-160 No. of males 161-170 171-180 181-190 11-20 Apr 1 1 0 0 21-30 Apr 1 1 1 0 May 4 1 1 1 16-25 Apr 1 1 0 0 26 Apr5 May 4 1 2 0 6-16 May 1 1 4 1 1-10 This study 6-15 May from 3 (15.00/0 during 21 April-lO to 7 (35.0% during 26 April-151v1ay) days/20-day Table 12. of obtaining '..;,...-. .~ �99 In 1979, the highest The acceptable range count for 3 leks was 175 males was established 10.0 and 30.00/0 of the counts during 1-10 May, improved respectively, to 20.0, 26 April-5 female May, peak). (11 April-l0 the latter Table 13. were above 140 males 21-30 April (Table 13). 6-15 May and 16-25 May (6-35 days after counts (161-175 males) May) to 8 (Z6 April-Z5 and This increased May) days/ZO-day the from period 2 in of the period. Number of counts with acceptable numbers of male sage grouse using Z techniques in North Park, Colorado, 1979. References Braun and Beck (1976) This study 11-20 April, and 0.0, 30.0 and 70.00/0 if counts were made during Higher part at 141-175 males on 16 May. Dates 141-150 No. of males 151-160 161-170 171-180 11-20 Apr 0 0 0 0 21-30 Apr 0 0 1 0 1-10 May Z 0 1 0 26 Apr5 May 1 0 1 0 6 -15 May 1 0 2 0 16-25 May 1 1 2 3 �100 The 1978 -79 data indicate be conducted mended later in the breeding (Patterson Hartzler 19S2:92 -93, 1978, Autenreith and 1 count during when females be conducted 16-25 April, 6-lS May. because early and females from spring peak from 16 April-lS May 2 counts during 26 April-S During a late spring 5 -3S days following 2 counts during of the 2 years the peak of male following the female the female May (i. e. 1979) lek counts should peak with 1 count during the 2nd 10 days and 1 count during investigated attendance peak. the optimum Radio -marked schedule can be considered occurred Therefore, may be too late during normal 11-20 April, more than 3 weeks my recommended years. normal count During an early ._ to count leks should be 1 count during 2 counts during 21-30 April and 1 count during juvenile 7S.0-100.0/S-days) tively, recom- 10 days. Neither spring occurs During an early do not peak until IS April or later, the l s t 10 days, periods 1979). lek counts should be conducted with 1 count during the last Season than previously should Braun and Beck 1976, Jenni and et al. (i. e. 1978) when snow melt 1-10 April, that lek counts in North Park and adult males and 84.2% (range of the locations obta..ined during visited leks on 86.8 (range 70.0-l00.0/S-days), this 30-day period respecin 1978. During a late spring,"""4 lek counts should be conducted the 30 days following the female 1-10 May. peak with 2 counts during during the 2nd �101 10 days and 1 count each during the Ls t and last 10 days. attendance was 85.9 (range of radio -marked 71.0-100.0/5 tively, May 1979. 85% of the juvenile data indicate were present Lek attendance These distances of 313 off-lek Gill (1965) reported from 0.4-1.6 locations exceeded maximum 1967, Wallestad may exceed krn during throughout the breeding the breeding distance radii cruising in the same area. of 1. 8 km (Wallestad (1979) reported within 1.0 krn in northeast of the off -lek locations during reported from leks increased radii of 1. 4-1. 8 krn Daytime rno ve > Montana with and Schladweiler off-lek distances 1974). to 2.5 krri with 64% Wyoming .. This was similar to the 62.6% within 1. 0 km of a lek in North Park, 1978-79 (Table 7). 1979). Season in North Park, (1967) found maximum season previously to 1. 3 km from a lek were common in central Colorado and adults, 1974, Rothenmaier that the daily cruising Carr Rothenmaier on Iek s ' were beyond 2.5 krn , distances and Schladweiler Colorado. a maximum that were present. Six percent ments respec- this time as 90.4 and 93.9% of the juveniles respectively, (Carr Telemetry and adult males during the peak of male attendance. 900/0during and adults days) and 97.2% (range 93.1-100.0/5-days), during 21 April-20 at least juveniles Lek �102 Vegetation Over 550/0 (56.3) of 101 feeding and loafing sagebrush trasts Analysis with a canopy coverage and Schladweiler in the 0.1-20.00/0 class Montana (Wallestad (1974). was 2.5 times and Schladweiler 1974). types. Sagebrush canopy coverage to the 28 and 32% for winter brush 0.4-49. canopy coverage (71 of 88) of on-lek (P < 0.05) between wi th similar reported Reasons 28.10/0 in for this difin vegetation (range cover, 2.9-68.8), respectively, 1972, Wallestad vegetative normal Average 18.7 cm (range of less and to roost sites in sageThe large ..;_- cover in heavier was in differences sites. lek position height for roosting 9-68) compared occurred resulted of winter Apparently, selection sagebrush sites and roosting parameters locations than 200/0(Table 9). roosting feeding-loafing by Beck (1975). to overcome for 88 roosting 3) and 89.70/0 of the locations with a canopy coverage number by 1974). The average 8. 7% (range reported than reported and spring Montana (Eng and Schladweiler Schladweiler This con- to differences averaged in use in North Park greater are unknown but may be related in central classes Sagebrush ference similar occurred of 20-500/0 (Table 9). with the 800/0use in these coverage Wallestad sites This contrasts in North Park is stimulus cover. on lek~was. with 38.7 cm for the 17 off-1ek enough �103 roosts (Table 10). This difference desire to roost on leks. (P < 0.05) reflected the apparent �104 RECOMMENDATIONS 1. Systematic annually searches throughout in Colorado. searches 2. North Park Aerial in the breeding Season (20 March-20 tion to the regular 4 counts of males These April) nour a ft er s un r i. s e , harassment are not considered can be extended to All active on leks. t last hour before where raptor a problem, It hours after predation and and the daily count sunrise (Jenni and 1978). leks should be counted 4 times 30-day period. days, In areas to identify counts would be in addi- All active leks should be counted between Hartzler and ground should be counted 1-2 times attendance. period ranges should be conducted in April and early May. 1 , S. sage grouse from helicopter) the peak of female z 4. leks should be made and other (preferably Leks with 40 or more males early 3. for new or relocated each spting One count should be made during 2 counts during during a. the Ls t 10 the 2nd 10 days and 1 count during the 10 days. During years when the female and 10 April, the 30-day count period and continue through 10 May. peak occurs between 20 March should begin Q~ll At lower elevations April than North �105 Park period (below 1500 m) it may be necessary 5 to 10 days earlier because to begin the count of earlier peak attendance of females. 6. During years (peak after immediately 7. with delayed 11 April), snow melt and late female the 30 -day count period area which could involve interlek which are closely movements Alkali, and Lake Junction, 2 and Riley should be considered Over 85% of the males should be present attendance. Wattenberg associated should be counted as a complex on the same day (i. e. Boettcher a complex). should begin following the peak of females. All leks in a particular Bighorn, arrival within the area on the leks during the period of a complex of peak male �106 SUMMARY Investigations from late March conducted through mid-June Objectives breeding Thirty-seven transmitters operational and studied on 5 leks. 64.3). Jackson were County, male lek attendance and habitat selection. were equipped with radio Transmitters functioned expe r ienc ed Five of 35 (14.3%) transmitters due to premature transmitter intermittency (2). failure Three for (3) and transmitters off the tail when preened. Peak female attendance 1978 and about 20 April the female of juvenile respectively. after. in Colorado Investigations of North Park, sage grouse cold temperature-induced after 1978-79. season movements, problems were conducted were to determine male 4-129 days (avg. slipped sage grouse in the Lake John area Colorado. patterns, of male on leks occurred 1979. peak on the 3 largest and adult males during increased counts (1-5 April), leks. (P peaked 25-37 days In 1978, lek attendance (P < 0.05) there- which included most high female 50.0 and 70.6% of the radio -marked respectively, 3-7 April < 0.05) to 95.1 and 100.0%, 1-15 May, and decreased During the 5-day period and adult males, Male attendance between were on leks. juvenile Over 90% ~the �107 juveniles (92.1) and adults of peak male attendance Lek attendance late April 1979. on 1-10 May, decreased adults (P (P for juveniles attendance the male (90.4) male visited (26 April-lO of males Juvenile (P < 0.05) increased attendance and adults, (93.9) attended counts for the 2 years Alkali and respectively, during Over 90% of the juveniles leks during the period of high caught while roosting a segment off Lek s visited of the locations of the male obtained. sage grouse which di.d not attend leks. All radio -marked 2 -4) during averaged of combined. and adult males identify Attendance the peak of female and 89.7 and 93.7%, peak (11-25 May) in 1979. and was 66.7 and 100.0% during leks for 45.0 and 63. 3%, respectively, (range to 73.10/0 through mid-May Attendance respectively, (16-20 April) I was not able to to 100.0% in (P < 0.05) decreased (P > 0.05) high (94.0-98.0%) < 0.05) thereafter. Juvenile the period May). < 0.05) to 78.80/0 from 21-31 May. and adults population leks during (P < 0.05) to 91. 1% on 11-20 May, increased remained decreased (94.2) the breeding 3.9 (range Lake-Boettcher and Boettcher from Boettcher juvenile males s e a s on , 1-13) per juvenile Lals~ Junction, Lake Junction-Bighorn Lake Junction visited more Interlek than 1 lek movements and were common between Alkali Lake-Wattenberg leks. 2, One j uv eni.Iefrnov ed to Alkali Lake lek (4.5 km distance) �109 Approximately m sagebrush coverage with a canopy coverage respectively. loafing sites less were in sagebrush b. sagebrush pared Average locations Sagebrush (including were 8.7% and 18.7 cm, Nine of 37 (24. 3%) radio -marked Three 1978. Three season and 3 males adult males juveniles Recommended male (2 adults and height for all respectively, are: counts should be conducted after sunrise to minimize of raptor or immediately after in spring spring lek counts before and harassment; i hour 4 counts (twice during 2nd 10 days) beginning at lower 1500 m) during a year when peak female 10 April, were were shot in 1979. i hour between should be made in a 30-day period (P < 0.05). sage grouse for conducting the effects on 11 April (or 5-10 days earlier roosts com- during the 1978 hunting and 1 juvenile) procedures than 20% and were killed by predators were harvested Approxi- 71 on leks) analyzed of less canopy coverage canopy feeding and with a height of 20 -70 cm. sites occurred 28.1% and of the daytime with 22.4% and 38.7 em for the 17 off-lek recovered. sites averaged with a canopy coverage than 40 em high. 88 roosting sites Eighty percent 90% (89.7) of the roosting occurred of 20 -50%. and height at feeding-loafing 43.5 cm, mately 56% (56.3) of 160 feeding-loafing elevations attendance the female such as 1200occurs before peak in a late year; all leks in a c Io s e Iy a s s oc i a.teel complex should be counted on the same c �110 day to minimize effects of interlek movements; and searches should be continued for new leks in the count area. �III LITERATURE CITED Autenrieth, R., W. Molini, and C. E. Braun. 1979. Recommended sage grouse management practices. 2nd Draft. We s tern States Sage Grouse Committee, Twin Falls, Idaho. 4lpp. Beck, T. D. 1. 1975. Attributes of a wintering population of sage grouse, North Park, Colorado. M. S. Thesis. Colorado State Univ., Fort Collins. 49pp. Beekly, A. L. Colorado. 1915. U.S. Geology and coal resources of North Park, Geol. Surv. Bull. 596. 121pp. Beetle, A. A. 1960. A study of sagebrush, the section Tridentatae of Artemisia. BulL Univ, Wyoming Exp. Sta , 368. 83pp. Braun, C. E. 1979. Evaluation of the effects of changes in hunting regulations on sage grouse populations. Colorado Div, Wildl. Job Progress Rep., Fed. Aid Proj. W-37-R-32, Work Plan 3, Job 9a. pp. 11~35. , and 1:, D. L Beck. 1976. Effects of sagebrush control distribution and abundance of sage grouse. Colorado Div. Wildl. Final Rep., Fed. Aid Proj. W- 37 -R, Work Plan 3, Job8a. pp.21-84. ----on Bray, O. E., and G. We Corner. 1972. A tail clip for attaching transmitters to birds. J. Wild!. Manage. 36:640 -642. Canfield, R. H. 1941. Application in sampling range vegetation, of the line i.nterception J. For. 39: 388 - 394. method on abundance, M. S. Thesis. Carr, H. D. 1967. Effects of sagebrush spraying distribution, and movements of sage grouse. 106pp. Colorado State Un i v,., Fort Collins. Dalke, P. D., D. B. Pyrah, D. C. Stanton, J. E. Crawford, and E. Schlatterer. 1960. Seasonal movements and breeding . behavi.or of sage gro~se i.n Idaho. Trans. North AJ:B. Wildl. and Nat. Res. Coni. 25:396-407. �112 Dalke. P. D., D. B. Pyrah, D. C. Stanton, J. E. Crawford, and E. Schlatterer. 1963. Ecology, productivity. and management of sage grouse in Idaho. J. Wildl. Manage. 27:810 -841. Emmons, S. R., and B. E. Petersen. 1979. Telemetry tions of sage grouse in North Park, Colorado. Int. Biotelem. 2 :215 -218. Eng, R. L. 1955. A method sex ratios from wings. investigaConf. Wildl. for obtaining sage grouse age and J. Wildl. Manage. 19:267-272. 1963. Observations on the breeding biology of male sage grouse. J. Wildl. Manage. 27:841-846. , and P. Schladweiler. 1972. Sage grouse winter moveand habitat use in central Montana. J. Wild!. Manage. 36:141-146. ----ments Finch, W. C•• ed , 1957. Middle Parks Basin, Denver. 157pp. Guidebook to the geology of North and Colorado. Rocky Mtn. Assoc. Geol., Gill, R. B. 1965. Distribution and abundance of a population of sage grouse in North Park, Colorado. M. S. Thesis. Colorado State Urii,v•• Fort Collins. 185pp. Hail. W. J. 1965. Geology of northwest North Park, U. S. Geol. Surv. Bull. 1188. 133pp. Colo.rado. .Lerini, D. A•• and J. E. Hartzler. 1978. Attendance at a sage grous e lek: implications for spring censuses. J. Wild1. l~anage. 42:46-52. Klebenow, D. A. 1969. Sage grouse nesting Idaho. J. Wildl. Manage. 33:649 -661. and brood habitat Lacher, J. R., and D. D. Lacher. 1964" A mobile trap. J. Wild1. Manage. 28:595 -597. May. in cannon net T. 4';:~· 1970. Effects of sagebrush control on distribution and abundance of sage grouse. Colorado Game. Fish and Parks Div , , Job Compl. Rep •• Fed. Aid Proj. W-37 -R-2 3, -~ Work Plan 3, Job Ba , 23pp. Patterson, R. L. 1952. The sage grouse Inc .• Denver. 341pp. in Wyoming. Sage Books, �113 Peterson, J. G. 1970. The food habits and summer distribution of juvenile sage grouse in central Montana. J. Wildl. Manage. 34:147-155. Pyrah, D. B. 1959. Sage grouse population trend and trapping study. Wyoming Game and Fish Cornrn; , Job Compl. Rep., Fed. Aid Proj. W-50-R-8. 27pp. Rippin, A. B., and D. A. Boag. 1974. Recruitment to populations of male sharp-tailed grouse. J. Wildl. Manage. 38:616 -621. Robel, R. J. 1969. Movements and flock stratification within a population of blackcocks in Scotland. J. Anim. Ecol. 38:755 -763. Rothemnaier, D. 1979. Sage grouse reproductive ecology: breeding season movements, strutting ground attendance and site characteristics, and nesting. M. S. Thesis. Univ. of Wyoming, Laramie. 97pp. Smith, E. L. 1966. Soil vegetation relationships of some Artemisia types in North Park, Colorado. Ph. D. Diss. Colorado State Univ., Fort Collins. 203pp. Terwilliger, C., and E. L. Smith. 1978. Range resource types in North Park, Colorado. Colorado State Univ, , Range Sci. Sere 32. 48pp. U. S. Department of Commerce. 1973. Monthly normals of temperature, precipitation, and heating and cooling degree days, 1941-70. Climatography of the U.S. 81(5). 12pp. 1978. Climatological data: annual summary, National Oceanic and Atmospheric Admin. 83( 13). Colorado. 14pp. 1979. Climatological data: annual surrrmary, Colorado. National Oceanic and Atmospheric Admin. 84( 13). 14pp. Wallestad, R. 0., and P. Schladwei1er. 1974. Breeding season movements and habitat selection of male sage grouse. J. Wildl. Mana ge , 38:634-637. , J. G. Peterson,"'-·and R. L. Eng. ----s-a-ge grouse in central Montana. 1975. Food~ of adult J. Wild!. Manage. 39:628-630. �114 Prepared t. ; -;): by __ ~,~~·(i~v~~~e~·_'~-?~' __ ~~~:Lt~.,~.~{,~·~~'~"7~~~----- Steven R. Emmons (//2) Graduate Research Assistant Approved by ---~~~~=..!."'-=~::....:'~~fLU-~=----- Clait E. Br~ Wildlife Researcher �us April 1980 JOB FINAL REPORT Sta te ·0 Project f _:C:.:o:..:l=..:o:.:r:..:a=-d=-o=-~- Job No. 3 Work Plan No. Job Title Game Bird Survey W-37-R-33 No. Evaluation of the Effects on Sage Grouse Populations: of Changes 9c in Hunting Evaluation Regulations of Censuses of Females Period Covered: Personnel: 1 January 1978 through 31 March 1980 D. Hein, P. Lehner and R. A. Ryder, Colorado State University; Jean Bourassa and Larry Kolz, U.S. Fish and Wildlife Service; Clait Braun, Steven Emmons, Howard Funk, Kenneth Giesen, Sue McElderry, Brett Petersen, Steve Porter, Tom Schoenberg, John Wagner, Colorado Division of Wildlife. ABSTRACT The breeding and nesting ecology of female sage grouse urophasianus) was investigated in the Spring Creek area Colorado from March through August, 1978-79. Forty-two and 23 yearlings) trapped on or near 2 leks were fitted radio transmitters and located daily from April through (Centrocercus of North Park, hens (19 adults with 14-15 g early June. Approximately 90% (89.9) of the adult females attended 1 lek with 10.1% attending 2 leks. Adults attended leks from 1 (44.4%) to 3 days (33.3%). Approximately 82% (81.8) of the yearling hens attended 1 lek while 18.2% vis1ted 2 leks. Yearlings visited leks from 1 (45.5%) to 5 days (9.1%). Prior to renesting adult females attended the original lek vi~ited for 1 day if the nest was lost during incubation. Lek revisitation did not occur when nests were lost during laying. Yearling females that renested attended 1 (33.3%) or 2 days (66.7%) on a lek that was visited prior to their 1st nesting attempt. Distances moved to nest sites from known leks visited were 2.3 arid 5.4 km for yearling and adult hens, respectively. Movements to feeding and loafing sites from nests by adults averaged 1.5 km during egg laying, and decreased to 0.1 km during incubation. Yearling hens moved an average of 0.7 km from their nests to feeding and loafing sites. These distances decreased to 0.1 km during incubation. Both age classes left their nests twice each to feed during incubation. Movements after completion of nesting, either successfully or after loss, were related to timing of vegetation desiccation. �116 Thirty-five nests were located of which 7 were renests. Completed 1st clutches of adults averaged 7.9 eggs (range 6-9), while yearling hens averaged 6.7 eggs (range 5-9). Renests of adults averaged 6.8 and yearlings averaged 7.0 eggs. Number of days between date of last lek attendance and onset of egg l~ying was 7.6 and 9.8 for adults and yearlings, respectively. Timing of egg deposition was 2 eggs 30 hours apart followed by a 1 day interval when no egg was laid. Initiation of incubation by both age classes occurred an average of 1.1 days after completion of the clutch. Length of incubation was 25-26 days with hens and broods leaving the nest the morning following hatching. Nesting success was 53.8% for adults and 41.2% for yearlings. Hatching success of eggs was 85.4% and fertility was 91.7%. Eggs from adult hens averaged 56.3 x 38.9 mm (length x width), while eggs of yearlings were 55.8 x 38.5 mm. Egg weights varied from 40-55 g upon deposition and decreased 11.4-11.7% in weight prior to hatching. Eggs from adult hens averaged 47.0 g, while those from yearlings averaged 45.2 ~. Chick sage grouse from adult hens averaged 31.3 g when they left the nest, while chicks of yearling females averaged 30.0 g. Chicks lost an average of 0.6 g/day until the 3rd day after hatching and then gained weight at approximately 1.7 g/day until 1 week of age. Weight gain for chicks increased to 6.1 g/day at 3 weeks of age. Replacement of juvenile primaries occurred at 23-, 31-, 38-, and 48-days of age for primaries I, II, III, and IV, respectively. Mating areas on leks had an average sagebrush cover (Artemisia spp.) of 7.3% with a height of 5.3 cm. Roosting sites had average sagebrush height of 7.6 cm, while loafing and feeding areas were mainly (55.7%) in areas with average height of 15.1-30.0 cm. Nests were located under sagebrush or complexes of sagebrush and rabbitbrush (Chrysothamnus spp.). Average canopy height was 52.3 cm (range 27-76 cm), while height of the surrounding sagebrush was 32.3 cm (range 11.1-59.8 cm). Slope at nest sites varied from 0~36% with 85.8% of all nests on slopes of less than 12%. Feeding-sites were located in areas < 15 cm in height. �117 INTRODUCTION Sage grouse historically sagebrush-dominated distribution Canada, rangelands west to eastern appears Washington and south into eastern Populations Washington). to have resulted overgrazing were extirpated 1974). The decrease With increased the status developed areas Patterson grouse, California in some in other localities in overall distribution alteration, especially of the sagebrush (Griner canopy (Patterson 1939, Patterson 1970, Eng and Schladwei1er rangelands, Their 1952). of sage grouse has been shown to be intiInately cia ted with sagebrush Peterson declined from vegetation and removal The distribution North Arne r i.ca, New Mexico into southern (New Mexico) and have markedly (California, throughout much of the of western extended from northern (Aldrich and Duvalll955). areas occurred development asso- 1952, Klebenow 1969, 1972, Walles tad and Pyrah of energy reserves and distribution of sage grouse under western i~ these will be altered. (1952) described while Scott (1942), -_ tensively studied displays of female sage grouse the -general life history of the sage Lumsden (1968), and Wiley (1973) in2.f males on leks. has .hinde r ed study; of their biology are not known. The secretive consequently, Many studies behavior 'i'ome aspects have investigated nesting �118 and brood requirements areas utilized Patterson and physiographic during these periods aspects (Rasmussen of habitats and Griner and 1938, 1952. Klebenow 1969, May and Poley 1969, Poley 1969, May 1970, Peterson investigations 1970, Wallestad dealt primarily movements, with vegetation, but not with behavioral has examined daily and seasonal grouse on leks. However, and Pyrah activities attendance 1974). These habitat types, of females. patterns and No study of female sage fidelity of sage grouse hens to leks was mentioned by Scott (1942), Dalke et al. (1960, 1963), Gill (1965), and Wallestad females and Schladweiler was also mentioned this thesis were collected and include information phenology, Park, (1974). Daily lek attendance by Lumsden (1968). Data presented from March through August, on breeding and habitat selection by season activities, in 1978-79, nesting by female sage grouse in North Colorado. The objective was to supplement of daily activity patterns Hypotheses of females which were tested were: during the breeding season, existing biological from early spring to summer. (1) adult females visit 1 lek (2) adult females for more than 1 day during the breeding unsuccessful visited, in their 1st n es ttng attempt (4) adult females during their renest period, attend the same lek season, (3) adult females return to the lek initially that have lost nests breeding knowledge attend the lek for 1 day (5) yearling females visit more �119 than 1 lek during the breeding each lek visited more yearling females season, (6) yearling females than 1 day during the breeding that have lost nests do not attempt attend season, to renest. and (7) �120 DESCRIPTION OF STUDY AREA The investigation Colorado (Fig. was conducted with the center 1). Boundaries of the study area of the area 14 km southeast 0 (Fig. 2). of Walden and Owl Ridge on the south, and Colorado Highway 125 on the west (40 32'-42' 0 County, Colorado Highway 14 and County (J. C.) Road 27 on the east, 106 04' -18' west longitude) Jackson were the Michigan River Johnny Moore Mountain on the north, Jackson in North Park, Total area north latitude and was approximately ? 222.2 km ~ and encompassed all known movements of instrumented birds. Elevation its confluence Topography varied from. 2463 m along the Michigan River with the Illinois was low-rolling and drainages major streams. patterns The majority which flowed northwest the Michigan and Illinois in the low alluvial edges of the area. to 2684 rn on top of Owl Ridge. to flat benches leading north. by Spring Creek, River near with most of the area to the Illinois rivers, small valleys was drained River. Two formed meandering flood plains .along the north and west Owl Creek, in the middle of the area, emptied into the Michigan River. Three county roads dl.vide the area. 14 and went south across Owl Ridge; J. C. 21 started at Colorado J. C. 25 began at Colorado 14 and �121 s• _\ . '.._za \ j--~----~~~ I.i __•. _ or. ,.' J ,'!...... ~ I I . /\'" I ! 8160 -. j"'°L"r ._, "...• \. ~.t\ •.•. (".. '. ;. "'•.J. •• \ 1.'_· f , _ (.- \ •• ... : '_'--"~ I ~--~J . • I • I , C •••.••• 1--. 6"'" Fig. 1. Female 1978-79. sage grouse study area, North Park, Colorado, �N r PEREGRINE lEK . .) ~ ~ •.... N N SCALE Fig. 2. - KILOMETERS Location of active leks in the Spring Creek area, North Park, Colorado, �123 and ended at J. C. 27; and J. C. 32 started went south-southeast 125 and went east ending at J. C. 25. Colorado throughout the area Many vehicle trails aided accessibility. Six leks were known to occur in the area. (Spring Creek at 1 and 2) were 2.4 k:m apart The 2 largest along J. C. 21. Creek 4 was west of J. C. 21, 2.4 k:m from Spring Creek lek was 1.9 km north-northeast leks Spring 1. Eagle of Spring Creek 4; Peregrine lek was located 0.9 k:m east of Colorado Eagle lek. Owl Creek lek was east of J. C. 25, 7. 3 km southeast Spring Creek I (Fig. in the area tridentata 900/0of the sagebrush (A. ~ (Beetle areas herbaceou~ and bitterbrush forbs and bunch grasses vegetation of North Park winter dominated (A. argilosa) silver occurred were favorable were rabbitbrush, (Purshia snakeweed ver~icu1atus), tridentata). comprised Perennial the majority (Terwilliger is cold, of the and Smith 1978). dry and windy .. No wind but prevailing and sprmg. longiloba), (Sarcobatus with few annuals are available, southwes t during of approximately and conditions Other shrubs s a r oth r a e }, greasewood The climate measurements Alkali (Artemisia where soil moisture willow (Salix spp.), low-growing which comprised and coaltown sagebrush 1960, Smith 1966). (Gutierrezia was sagebrush-bunchgrass vaseyana type. viscidula), in limited of 2). The vegetation by Artemisia 125 and 6.0 km northwest winds are fr~ Spring precipitation the consists �124 mainly of snow and accounts Mean monthly precipitation (Table 1). for 23.3% of the annual pr-ec ipi.tat ion, increases The mean temperature from March through July during the investigation varying from -3.8 C in March to 14.8 C in July (Table 1). was 64 C; �Table 1. Spring precipitation and temperature, Year Mar Precipitation Apr May a 1978 2 ..'o 1 2.16 8.03 1979 3.58 1. 24 30 -y r , avg.e 1. 27 1. 83 b (em) Jun Walden, Colorado, 1978-79. Mean daily Apr temperature (C) May Jun ·-flil Jul Mar 1. 57 0.96 -1. 3 3. 1 5.6 11. 9 14.8 3.45 3. 15 0.94 -3.8 1.6 6.3 11.2 14.8 2.59 2.82 3. 15 -4.6 1.8 7.1 11. 6 14.8 a b c U. S. Deparhnent of Commerce (1978). U.S. Department of Commerce (1979). U.S. Department of Comrneree (1973). ,_. N VI �126 METHODS AND MATERIALS Roosting were located on or near leks and along roads using vehicle-mounted or hand-held spotlights. were captured net (Pyrah 1976). female sage grouse with a long-handled Some hens were captured nets in April bands (size or year color-coded bandettes in 1979. molt recorded prior Females range, weighing 14-15 g. Transmitters Captured bandettes birds in 1978 were classi- 1975), and weight with wing near Transmitter U. S. Fish and Wildlife Ser~ice 1978 transmitters to attachment of 164 MHz of radios cloth to reduce were rnounted on the central end of the rachis 1972). Prior with a dark-colored The tail clip consisted (Bray and Corner Rebuilt plastic to release. were covered on the proximal the follicle. All hens were banded were equipped with VHF radio transmitters, in 1979, birds struggling. cannon nu.m.bered a.Iurn.inurn leg color-coded fied as to age (Eng 1955, Beck e t al. frequency 1950). Division of Wildlife serially 14) and individually 1959, Braun and Beck on 2 leks with 3-projectile 1978 (Dill and Thornsberry with Colorado Birds 2 rectrices the point of emergence q. from clamp and bolt arrangement units were supplied (Denver Research Center) by the in 1978. c;pnstituted 70% of the units utilized in �127 1979. Additional transmitters Company (Champaign, were obtained from AVM Instrument Illinois). Daily counts of female largest and male leks in the study area upon accessibility) attending the 2 were made from late March (depending until 31 May following techniques Braun and Beck (1976). 2 located sage grouse by on 4 additional leks including during 1978 were counted as time permitted. Supplemental counts of birds Birds present prescribed on these leks were made by Colorado Division of Wildlife personnel. Radio -equipped females of birds on leks each morning 3-element range. Locations in respect were determined Transmittered approached This procedure as determined cover, hens Hens were by receiver meter signal was continued until the 8th day after to' have been on a lek. were taken at 1- -to 2 -hour intervals percent by was attempted. or flushed until a nest was located. identify daily activity aspect, If radio-equipped visual location or until each hen was last determined locations tuned to the 164 MHz not on leks were located next. to within 50 m, with counts to being on a lek were determined to be on a lek, were then observed receiver of the radio signals. birds concurrently during April and May using a hand-held yagi antenna and a portable USe of triangulation strength. were located patteTns for different percent sagebrush capture Hens Additional during the day to radio-tagged and average hens. Slope, height were �128 recorded on standardized years. A written forms des cription form to aid in relocating with surveyors to the neares the area for each nest were: condition of nest date each additional date located, (construction was initiated, (±_ and width were measured g) on the day found. and at frequent was foraging Chicks nests, Nests were intervals measured in 1979 or when large summer loss and located or failure weighed during (:!:_ 2.0 (+ 1. 0 mrn), 1978. of eggs at clutch and date hen either or abandoned. examined were left the Egg length and carpal daily until incuba- captured while the hen the day they left on a spring scale color-coded expandable plastic enough to retain colored numbered Chicks were on brood hens. and primary (+ 2. 0 g). Each followed throughout at 1 - to 2 -week intervals of transmitters g), if during incubation females chick was banded with individually plas tic ba~.?ettes and egg covering. away. of radio-marked were of O. I rom) and each egg was weighed some distance and weights bandettes measured number egg was found, number date incubation tion started on 7.5 -minute and movements nest with a brood or the nest was destroyed (+ 2.0 being marked were plotted maps on the same t O. 1 km, eggs present, completion, Locations in both was placed with each location Survey topographic Data recorded any), were located of each location tape in 1979. U. S. Geological when birds coil the until brood dispersal Chicks lengths were or re-=Captured were measured �129 Measurements of vegetation ti on of the Canfield examined, (1941) line intercept a central in length was driven measuring point was randomly into the ground. tape was used between by compass directions. on each transect. supporting the 2nd rod. 15.0 m in 1979. A minimum At each location chosen and a steel additional 2 transects were used. and mating for big sagebrush, (1979) rod and a 2nd rod located to the highest was 15.2 m during centers to species, at all to the nearest an cm was black greasewood, to the nearest except by 1978 and on leks where sagebrush, was measured Only sagebrush was examined Vegetation silver Vegetation according rod 1.5 m (1978) or plastic the central of 3 transects sites pooled in 1978. technique. A steel Each transect except nest along transects using a modifica- Most sag of the line was eliIninated locations, and rabbitbrush. were made Height of the tape was adjusted vegetation recorded cover grasses was measured 0.5 ern whi ch-we r e along transects during 1979. The data were chi-square nest P = P = analysis success, o. 05 O. 10. analyzed with the Student's !_ test was used to test for differences and hatching ex c ep t for distance success. The level except that in clutch size, of significance was moved from leks to nest site which was �130 RESULTS AND DISCUSSION Trapping Forty captured sage grouse tail-mounted captured nets with 2 additional with cannon nets. radio transmitters on the 42 females; were females (1 adult All were equipped with between 26 March and 10 May except 1 hen that was equipped on 17 June. utilized Life hens (18 adults and 22 yearlings) using long-handled and 1 yearling) and Transmitter 3 radios Thirty-eight were placed transmitters were on 7 different birds. Three hens during, 1978 were equipped with the same transmitter due to its slipping transmitter off the 1st hen and death of the 2nd bird; remained Two other radios to movements on the 3rd female until transmitter were both placed on 2 different from the study area. moval of the transmitters Capture efforts hens were 1. captured Cannon-nets the apparently manner. allowed these radios were and along J. C. roads Subsequent concentrated 21, 32, and 25. near in 1979 due recapture and re- Spring Creek Forty-one percent 1 and 2 leks, (40.5) of the on the 2 leks with 12 (28.6%) on Spring Creek were utilized greater supplied failure. to be reused. s tress once on each lek during placed on the 2 birds use of this technique was discontinued. along roads birds the 1978. captured Capture 57. 1% (24) of all radio-equipped Due to in. this efforts females with �131 33.30/0 (14) captured along J.C. 21 and 14.3% (6) along J.C. Four hens (9.5%) were captured the study area "'Ia.S the I s t birds moving elsewhere while incubating in North Park. resulted to March, Early spring precipitation to on her nest and difficulty attachment. temperatures storms snow accumulating which delayed melting. and permanent ing Spring _~reek 1 in both years. days) for all radios used. the road. in In 1979, the lek times due did not occur until 20 April. a minimum of 5 days after Due to late access, the peak of hen attendance operated Mean on 26 March but the road was closed-3 access l). in March and April 1 lek was 1st reached to Spring Creek 2 occurred Transmitters 2 C colder did not close on 31 March, after into (March and April) was 34. 5% above the Spring Creek hens were captured in capAccess in 1978 and 55.5% above in 1979 (Table 1978 and subsequent Access One hen was captured in late transmitter were approximately to snowstorms were were staging prior to the study area and colder 30 -y ea r average was l s t reached 3 females in 1979 than in 1978 due to more prior 1). since no signal on 17 June 1979. the area was later 1979 (Table These in 1979 and apparently Lack of accessibility turing females to have left, the day following capture. captured temperatures along J. C. 25, but 2 moved from and 1 other was believed heard after 32. an average reach- 78.6% of the (Table 2). --= of 52.2 days (range 0 -151 Maximum radio life was determined from �132 Table 2. Transmitter attaclunent by ti:me period for female grouse, North Park, Colorado, 1978-79. Number of females 1978 Time period 26-31 Mar sage radio-marked 1979 o 1 o 1-5 Apr 6-10 Apr 5 11-15 Apr 3 16-20 Apr o 21-25 Apr 2 5 26-30 Apr o 9 1-5 May 1 3 6-10 May 1 1 Totals 20 apeak attendance on 5 April. of hens occurred on 4 April; attendance of hens occurred on 20 April. bpeak 2 cOne additional 2 radios radio was placed on an incubating were attached hen on 17 June. �133 date activated transmitters until the date signals were last received. Fourteen were still working beyond 15 June 1978 and 24 June 1979 when daily locations were terminated. After this time, tions were made at 1 - to 2-week intervals. Exact dates that some transmitters last functioned were unknown, and transmitter was possibly somewhat longer than calculated. radios The average Nine transmitters mittent signals, tures. Two radios known to operate probably because 1978 after loss by their hens. intermittance transistor was attributed than the battery of these 9 radios average life of (21.4%) had inter- of shut off due to low tempera-' at intervals These transmitters (14.3%) were used in 1978; in addition, in 1979. 63.9 to 54.4 days. testing. These radios in and They reduced average Transmitter was the most important problem transmitters life from since all data Intermittent experienced. in conjunction with late attachment collection more in 1978 than in 1979. reduced transmitter life was not critical Six 1 .trans- Only 3 intermittent on nesting ecology were collected by early July. radios were tested could supply at low temperatures. life from 95. 2 to 58.2 days. transmission were retrieved to a need for rno r e .voltage by the did not work after original (7.1%) occurred mittent life in 1978 was 58.2 days (range 0-151 days) and 54.4 days (range 1-122 days) in 1979. mitter loca- signal Inter- affected data �134 Five additional transmitters data for short periods. prior to transmitter Three failure; radio-equipped analysis after nest loss. on 17 June. This problem occurred in 1979 by filling the grooves Early attadunent. rubber because until no hole re- holding the transmitters was elimi- the hens with a dark cloth during radio Three hens in 1978 lost retrices May apparently c.larnprng of in 1978 and was eliminated with silicon loss of retrices nated in 1979 by covering used One r ad io slipped from the tail because in the tail clip were too large for adequate the retrices. mained. for parasite placed on the nesting hen had been previously and was attached the grooves hens were killed 2 were killed by unknown predators, while the 3rd hen was collected The transmitter (2 in 1978 and 3 in 1979) provided of struggling early in April and during radio attachInent. Lek Attendance Counts of female Spring Creek area 1 hour after and male sage grouse were made from 1 hour before sunrise. females -_ newly located only for nesting in 1979. in 1978; 1 was. visited other was found while locating initiated on 26 March to no-later than One lek (Owl Creek) not counted by myself was in an area not used by radio-marked marked on 5 of 6 leks in the hens in 1978 and used by Two small leks were by 1 radio -marked another marked 1978 when approximately female. hen and th~ CZlunts were 350/0 of the area was �135 snow free. The 1st count in 1979 was on 31 March when the area was 50/0snow free and 30 cm of snow was present sibility to the leks was unaffected = daily counts (N during (N of birds by snow storms during 199) were made until 1 June (Table 1979 blocked roads = 96) on leks. present 3 times 3). Acces1978 and Snow storms after plowing and daily counts on leks were made from 23 April until 1 June (Table 3). Peak attendance of hens on leks occurred 1978 and 20 April 1979 (Fig. initially Counts in 1979 began with no birds appeared were required Spring Creek in 1979 (Fig. years. on leks when 3). occurred 5 days after a major which began_ the afternoon constant spring conditions number of hens on that occurred the peaks between the 2 peak was observed in 1978 being more days in timing of peak attendance Sixteen days separated smaller on the lek; Thirteen to peak number Difference was due to the later present on 8 April. appearance 1 during 1979. A later occurring on 7 April and females from initial between years fairly Hens were present counted in 1978, and it was unknown when hens started attending. males 3). the weeks of 4 April pronounced both years (Fig. snow storm 3). ~ith the 1 This peak (20 ern total depth), of 3 May and continued until 6 May. of hens attending possibly due to poor weather period. A 3rd smaller conditions peak occurred The leks in May 1979 was throughout the n es't irig in 1978. 5 days after a storm �Table 3•. Peak counts of sage grouse on 5 leks, North Park, Colorado, 1978-79. -. Lek ,I Number of counts 1978 1979 Females High count Dates 1978 1978 1979 1979 Males High count 1978 1979 Dates 1978 1979 Spring Creek 1 62 44 55 65 4 Apr 20 Apr 76 56 8 May 16 May Spring Creek 2 58 36 29 52 4 Apr 11 Apr 27 Apr 63 70 16 May 9 May 4 34 7 0 0 -- -- 3 0 8 Apr Eagle (found 7 Apr 1978) 25 5 8 0 -- 11 2 9 Apr 15 Apr 30 Apr Peregrine (found 20 Apr 1978) 20 2,6, 8 Apr 13 May Spring Creek I 4 3 1 8 Apr 20 Apr 13 May 17 20 ,_. w. 0\ �137 70 • • SPRING SPRING .--~ SPRING /:r---A SPRING 0----0 60 CREEK I CREEK I CREEK 2 CREEK 2 1978 1979 1978 1979 50 (f) iJ.J _J ~ ::: !.LJ u, 40 l ? u, a a: l1J CD I I I 30 ,f. :E I \ => Z I \ I 'zt I 20 \ \ I I I I I 10 /i - 0 I <0 N If) 10 10 I , I'NI <-MARCH 0 N. I I <0 <0 It) N APRIL N I 0 If) I <0 N 10 It) 0 N I <0 10 N I <0 (\J If) I <0 N MAY .::== Fig. 3. Female sage grouse Colorado, 1978 -79. 5 -da y inte rval. attendance on 2 leks, North Park, Data represent the high counts for each �138 that occurred on 17 and 18 May. peaks was apparently nests to predation on leks were more The occurrence due to an increased or abandonment. pronounced Data on lek attendance This lack of data was not indicative radios. of late transmitter Average yearlings 1. 8 days (Table 4). Table 4. Lek attendance by female Colorado, 1978-79. a Total number of days on lek( s) 1 Totals Averages a Females bIncludes c Includes (days) attended 1.9 by females, for hens that visited 2 hens that attended but of leks was of 1. 7 days and sage grouse, North Park, of females Renesting Adults S Sc - attempts Yearlings 0 1 2 0 0 0 11 2 3 1.0 1.7 1.8 only 1 le.k except where noted. 1 hen that attended birds. and early failure Number First nesting attempts Adults Yearlings 9 of hens hens were available went to leks an average 4b 2 3 0 2 3 5 in numbers of attendance length of attendance adult females of hens losing of the 42 radio-equipped attachment 1. 8 days; smaller snowstorms. of radio-marked for 23 (11 adults and 14 yearlings) was because number Increases after of later 2 different 2 different leks. leks. 1 2 0 0 �139 Number (Table 4). of days each hen visited Prior to 1st nesting attempts, leks an average of 1.9 days (range 1. 8 days (range 1-5). days. period. 1-3), Attendance One y ea r l.ing hen visited Hens attempting revisiting a lek. renests variable Adult females incubation yearlings nest, revisited in 1979 attended but no renest time for renesting Number to number dominant to or during require closer without 1 lek for days. Three loss of their There probably 1st was not enough in 1979. appeared With large may be unable to service hens to return. should have required attended 1 lek for hens losing nests a lek for 1 day after all hens. in 1979. were leks for only 1 day, due of hens present, This would radio-marked Therefore, mo re, th~n 1 day for breeding. 2. 3 days on Iek a-dur-in g 1979. to be partly numbers Adult· females to the peak of hen attendance averaged and attended attended a lek for 2 consecutive of hens on the leks. unmated an 11 day incubation. that renested of days leks were attended males attended during laying renested females was located. attended attended 1 day (33.3%) or 2 days (66.7%) (X = 1.7 days); during 1 to 5 while yearlings were less that los t nests Yearling adult females a lek on 5 days during 1 day if the nest was lost just prior 3 adults from was usually not on consecutive a lek only 1 or 2 days (Table 4). Apparently a lek varied these hens Adult females Adults marked i;-1978 sugges ting that late radio attachment �140 after the peak of hen attendance Renesting adults required 1st nest probably sive behavior Since yearlings of proper males 1 (88.00/0 of radio-marked at 2 leks occurred visited 2.4 km apart. Another yearling 1 to Peregrine lek. sites Two of 3 yearling to both nesting prior visited were less females attempts. to the I s t nesting breeding attended a lek 2 3 times by other hens) or 2 (12.00/0) to 1st nesting the 2 largest leks, attempts. a distance of moved 6.3 krn from Spring Creek The 2 adult females apparently capture Aggres- attendance. One adult and 1 yearling prior hen that renested because mating was interrupted Attendance original attend leks for the 1st time, on leks may also deter One yearling on than 1 day of other hens present during her initial to renesting. more cues may be necessary. Hens visited leks. required behavioral by subdominants. days probably after loss of their of the small nurrib er of hens present Yearling hens possibly due to inexperience. some learning in the lower average. 1 day of lek attendance because leks at that tirn e, resulted that attended the originallek a lek prior attended. as the than 300 m from the i"ek revisited. that renested visited the same lek prior It was unknown which lek was visited attempt by 1 yearling female that had a brood patch at the tirn e of captur-e, More yearling leks. Yearlings females (14.30/0) are inexperienced than adults (9.10/0) vrnited 2 at mating and nesting and may �141 investigate an area prior the possibility to site attachment. of going to more than 1 lek. 1978, that was equipped with transmitters This may increase One yearling both years, hen in attended 2 leks in 1978 and only 1 of the same leks in 1979. Three yearling displaying males females away from known leks. quent1y examined to learn none was observed. leks, were observed It is possible next closest dates with These sites were subse- if they were regularly but these 3 hens were later the lek closest on different used by males, that mating occurred located on leks. to their off lek observation. away from All 3 hens attended One hen also visited varied from 1 to 3. The nest of 1 adult was destroyed and 3 mornings visited a lek either were lost. the lek. Number of days between loss of nest and attendance afternoon but later she revisited the next morning Three yearlings a lek. or 2 mornings in 1979 which lost their on a lek during the Yearling after females their nests 1st nests re- ..•. visited leks within 2 or 3 days with 2 revisiting during the evening display period. Of the hypotheses dance it w~s necessary tested during the investigation to reject 1) adult females on lek atten- only visit 1 lek, since 10.10/0visited 2 leksl.._and 2) adults visit a lek for more 1 day, as 45.5% visited fo:v_only 1 day. reject 3) yearling hens visited more It was also necessary than to than 2 leks since 81.8% of all �142 yearlings (45.50/0) only attended instead to renest were: 1 lek. of more Yearling than 1 day. when time allowed. 1) adult females hens also attended Yearling for 1 day hens were also found The 2 hypotheses that were accepted attend the same lek when r ene atirig and 2) they attend for only 1 day. Lek attendance Dalke et .a.l, (1960, has been mentioned 1963) being the 1st to mention being common for hens. that hens were observed attended on more time, Morse than 1 lek, in a row. hens observed 4 leks in 1 year. observed Wallestad interlek the next closest Number of days on leks was 1st mentioned (1948) who suggested Aggressive also reported by Lumsden mate (1968) examined hens each being mated the lek again. 1 marked (1974) as 1 hen 1975) to 42.0% of the 1ek. that females Lumsden reported data in Wallestad than 1 lek from 26.0 with movements and Pyrah movements, usually 7 marked interlek but did observe Eng (unpublished that hens used more attend a lek daily. studies Gill (1965) working in Colorado hen on the same lek 2 years using radio-marked in previous by Batterson and only once and do not only 1 lek and observed only once with none being seen on behavior between hens and males (1968) as causing interruption was of mating. �143 Movements Twenty-three they attended female from Leks to Nesting sage grouse leks until nests were monitored were located. Distance obtained for 11 adult hens which laid 15 clutches in 1978; 8 hens and 11 nests Sites traveled was (3 hens and 4 nests in 1979) and 12 yearling each year) which laid 13 clutches from dates (1 hen renested hens (6 in in 1978). The average dis tance from leks to nest sites for all hens was 4.0 krn, Average distance between nests while for 17 nests (P and leks attended in 1979 the average in 1978 was 2.2 krn, was 4.9 km, The difference < 0.05) between years was possibly due to timing of transmitter attachment, attendance since more hens were captured of females during of hens radio-marked close to leks"prior radio-marked The average 1978 than in 1979. in 1978 had already to capture. birds Because were possibly distance years was probably nest sites fbr adults Yearling (Fig~ 4). No difference nesting sites of earlier capture in 1979, randomly selected. to nest sites averaged 0.4-5.8), (P > (P < 0.10) respecbetween Dis tance moved to 5.4 km. distance respectively o. 05) froIll.;.leks by adult 0.9-11.0), to timing oj. capture. hens moved an average a majority selected The difference in both years 4.3) and 2.7 krn (range Perhaps and 6.2 krn (range (Fig. A). related more travelled hens was 3.2 (range 0.8-4.9) tively for the 2 years 1 week af't e r peak of 1. 6 (r~~e O~-6- in 1978 and 1979 between years was found, but �144 a ADULT 1978 o ADULT 1979 ••.YEARLING 1978 A YEARLING 1979 N s Fig. 4. Distance (krn) from leks to nest sites, North Park, Co lo r ado , 1978-79. The center represents the lek and ~ch c-oncentric circle equals 1 krn. Two nests of an adult located 10.8 and 11. 0 krn ENE of the lek are not plotted. Data are for females with known lek attendance. �145 yearling hens moved less radio-marked in 1978. yearlings The average in 1978 than in 1979. were selecting distance This suggests nesting sites for all yearlings that when captured in both years was 2.1 km , Distances (P between lek attended < 0.05) between age classes year, but only different between age classes hens have had greater areas adequate (P and nest sites for the combined years < 0.10) for the 1st year. could possibly experience Differences in nesting and more since adult knowledge of for nest sites. adult and 1 yearling the adult nesting visited other yearling equal distance nested each attended the 2 Ia r'ge rLeks from both. between the 2 leks but closer three nests and the 2nd be due to experience Three hens (1 adult and 2 yearlings) nested were different 2 leks. in the study area The yearling to the originallek to the lek attended. with female visi-ted. 0.6 km from the 2nd lek attended. were closest The. The Twenty- Seven other nests were ..;;. closest to a lek that was not visited. the nearest lek and 2. 3 k:m.from the lek actually of hens that attended Spring Creek 1. 0) of the_:_OwlCreek lek. adult female, actually The closest visited. was unknown. Lek attendance visited. Four nests 1 w~re within 1. 0 km (range The remaining were 1.2 k:m.closer was 0.3 k:m.from 2 nests, to the nearest of the remaining 0.5- both by the same lek than the 1 ---= 12 hens that nested �146 Renests Distances of 4 adult and 1. yearling between 1st nests and averaged 0.8.lcrn. females and renests ranged from 0.2 -1. 9 k:m. The 1st nest attempt of the yearling 1. 6 km from the lek visited while her renest same lek revisited placed their avg. 0.7 krn). as a yearling visited. female hen attended (1964) speculated years 1 of the original 2 between the was 2.8 km, Colorado. an average since 1 he~ marked to incubation. distance He also each year to nest. were 24.1-32.2 Rogers k:m.from Foley (1969) using radio-marked of 2.4 k:m (range 0.1-7.6) for could be biased by time of radio-attachment, while roosting with 6 eggs 1 day later. (1952), who reported a lek 3.2 k:m from her nest site. that some nesting areas This average in North Park located both years; The distance to the same area leks in Moffat County, prior she again attended in Wyoming by Patterson felt that hens returned 4 birds.· k:m., between lek attended and nest sites for sage grouse was 1st observed hens reported hen had 1st nests 3.0 km from that lek. in the 2 successive 1 marked All 4 adults she located her nest 0.6 km from 1 of 2 Lek.s In 1979 as an adult, Distance nest. to the lek attended (range 0.1-1.9 One additional leks and nested nests closer hen was was 1. 8 k:m.from the and 0.6 km from the original renests were located. on a lek was located This female was probably on a nest captured Distance moved by known-age birds'~as just reported in 1969 and in Montana during 1969 through 1972. (1970) found that 4 adult hens moved an average of 4.4 km (range May �147 2.5 -7. 2) from leks on which they were captured, averaged 8.2 km (range reported 2.2-13.7). Wallestad that 13 adults and 9 yearlings captured; a range by age class but the minimum previously due to either utilized mating distances location of nesting habitat during the investigation. centers or on vehicles may have altered possibly increased Wallestad and Pyrah leks after capture, nesting they did not present The differences in relation (Lacher either and Lacher and Pyrah for some period (1974) reported Morning locations in May's after capture and of leks attended. May to (1970) docufrorp. lek of of radio-marked hens were study and these hens could possibly have a tt.ende d and mated on other leks. leks could account for the -difference habitat on This type of 3 other leks prior within 1. 7 km of the lek where captured. not always identified located that some hens visited .additional with I female attending to nest sites. could be 1964), were used (1974). moved and number between to leks or methods men ted that hens bypass ed other leks while travelling capture of 2.5 was 0.8 km and 2 Cannon nets, behavior distances (1974) and those in my investigation by both May (1970) and Wallestad capture distance more than 4. 8 km from the 1ek. reported and Pyrah moved an average and 2.7 km from leks where originally hens nested while 4 yearlings Nesting habitat in distance. in Montana may hae e been closer in relation Suitable nesting to leks than in Colorado. to �148 Daily Activity Patterns Egg Laying Activity for distance movements There patterns for females moved/day and distance were monitored were no differences travelled during a day. (P> indicated locations. low light intensity morning location with an apparent Distances to an area probably was be- were made during the morning the hen was 0.8 km north of her of other hens followed after fresh area was traversed in the early morning in the day. and the point of roosting wandering in distances being within 0.3 km of each other. was made. than later were restricted distance Three locations Tracks that more Daily One adult moved 1.1 km during the This greater on this adult with all locations previous sites. 0.05) between age classes Daily movements 1 day that it was followed. When the next location moved from nest for 9 laying hens (5 adults and 4 yearlings). of O. i km for 88.90/0 of the hens. cause of being flushed. during egg laying were examined Distances or landing wes up to 0.4 krn pattern. 32 clutches. the greatest between the nest and a known use area. by adults (N = 12) averaged Movements for 28 hens which produced ments 1979. 'No difference were examined 1.5 km (range O.S-S.~g adult hens av er ag ed 1. 1 km in 1978. during during between original moved around the nest site were recorded distance snow and 9 adults averaged (P > 0.05) between years using Move- Three 1. 6 km was found. �149 The greater distance moved in 1979 was due to a single adult hen that moved 5.5 krn on 3 different days during 2 nesting These long movements to be for feeding and loafing, 2 occurred appeared attempts. since on days when the hen was known not to lay an egg. sagebrush area approximately vegetation encompassing 1 ha in size, for normal The the nest on Johnny Moore Mountain was and probably maintenance did not include adequate activities. Eight of 12 adults (66.70/0) restricted the area used around nests laying. Distances moved from the nest during laying decreased adults that renested. This decrease to::. 1. 0 km during averaged 0.4 km (range for 4 hens which were followed during both nesting 1 adult did not decrease Movements distance by yearling 0.5 -1. 3), and were identical moved> area 0.0 -1. 0) attempts. Only moved during laying. hens (N = 16) averaged in both years. 1.0 km from her nest. for 0.7 krn (range Only 1 yearling Most yearling (6.2%) hens restricted used during laying to within 0.5 km of their nests. the Movements . ~.:. from nests decreased An additional remained yearling from 0.9 to 0.5 km for 1 yearling hen (Page within a 0.5 km area 46 [2]), with only the renest area adults for both age classes with a radius located, of her nest. Obs e r va.tion s during laying also indicated areas that renested. of hens. an affinity One yearling of O. 1 krn, 6 of 8 days. female The areas each for 5 long feeding and loafing movements to specific used an us~ by 2 were within a �150 radius of 0.2 km, 4 to 7. Movements centered Nurnb e r of days between movements uniform, from during laying were not random and usually around 2 or 3 specific from nest sites varied varied areas per nest. which suggests and that hens selected Directions that the vegetation areas that met their moved was not requirements during laying. Hens required area aiter attending by a yearling only 1 day. to breeding. areas The maximum only 2 days. a suitable Three visiting yearlings ne at.ing amount of time was 4 days required from the lek to a nesting area This small amount of variability be due to most hens having .chosen a location captured along roads a lek to locate an area only The greatest attending close to a lek. prior required for nesting. site to a nest after Two of these hens nested a lek on non-consecutive to leks. time to locate Most hens (6 adults and 8 yearlings) moved from capture 0.4 km, little The longest movement could possibly 1 day after a lek. female. (11.0 km) required distance relatively a lek was Hens that visited - days moved back and forth from nest area Hens that took 2 or 3 days to move to a nesting between the lek and nest and did not return site used to those areas again. Incubation With the onset of incubation, were limited to feeding excursions. activity patterns Feeding sites of females were a maximum �151 of 0.3 krn. from nests, occurred period with most usually twice a day, prior was longer and 1 yearling) excursions. to sunrise and farther within 0.1 krrr, and after from the nest. than 15 minutes), nests One feeding Two females (1 adult being shorter twice in 1 day. (22.20/0) were observed All hens in 1979 were away from their which fed only once a day on 1 occasion each. of 2 yearling to the nest, feeding periods Because from the nest and close proximity it was difficult of greatest to observe duration hens Length of time away was as long as 30 and short as 15 minutes. the short absences (less fed once a day 2 times during most days with the exception from nests their longer in duration I felt that most hens apparently during the 1978 season as only 2 of 9 females off their nests sunset. moved 0.3 krn to feeding sites during Due to 1 of the periods Feeding of of feeding areas exact length of absences. The did not change from rnorrring to evening and stayed the same for each h en , Nelson (1955) collected precise data on timing of feeding for -...;_ incubating hens by using a recording that hens left their nests as 35 minutes. Absences twice daily for as little from nests 0430-0630..and 1800-1900 hours. were predictable females collected thermometer. usually He reported as 10 and as long occurred He also observed that some hens at leaving nests during both periods, were predictable between while other during only 1 period or not at all. in my study support Nelson's (1955) findings. Data �152 Post Nesting Data on activity successfully adults or after patterns nest loss, and 9 yearlings). from early locations nests after were available Locations to late summer completion of nesting, either for 17 females (8 of these hens were obtained at 1- to 2 -week intervals. Number for each hen varied from 1 to 8.. Distances and from last locations were grouped according of moved from to the amount of tizn a since nest departure. Successful stayed after females (6 adults and 4 yearlings) on or moved toward leaving nests. averaged ridge tops and benches Distances traveled 0.8 krn with no differences Length of time hens with broods hatching date in relation average spring quently movements conditions on ridges remained up to 9 weeks, eventually krn and depended upon desiccation Broods hatching prior while those hatching During this tUne movements· Movements varied on ridges during both years. 0.2 krn in 4 weeks to 1. 0 km in 2 days. broods from 0.3-2.3 0.05) between age classes. (~> occurred within 1- to 2-days in 1978 and 1979 may be attributed that prevailed. for 3 weeks. varied to onset of vegetation precipitation with broods •. Above Conse- to the moist to 1 J~ne remained after 18 June remained along ridges All successful varied from hens with moved to-rneadows. by unsuccessful females (2 adults and 5'yearlings) as did amount of time spent in an area. Initial movements �153 by these hens averaged yearling 1. 5 k:m (range made initial movements remained females in these areas distances One unsuccessful One adult and 1 away from their nest sites until transmitter moved the greatest to an area. O; 5-2. 7). failure. Four yearling and showed the least yearling. which abandoned on 22 May moved 2.6 k:m at 2 - to 3-day intervals of < 0.5 k:m in the interval. dows, All unsuccessful movements than 2 to approximately appeared to vegetation to be more related 5 weeks. (1969) reported her nest prior for differential Length of to timing of nest lost than females of their nests Griner of radio-marked were concentrated observed hatching. nests hen moved May (1970) working that 2 birds remained in the for short per iods until radio for approximately Wallestac;l..(1971) reported broods Poley within 0.3 k:m of while another (1939) and Gill (1965)' reported ne a r their ing to meadows. after with broods between years. hen remained to moving to a meadow, with radio -marked and its variability movements that 1 radio-marked toward a meadow immediately remained in meadows desiccation. in Idaho accounted failure. its nest hens moved into mea- Klebenow (1969) noted that precipitation vicinity fidelity with short moves with amount of time between nest loss and arrival varying from less and then that hens with broods 2. weeks prior average to migrat- daily movements of."(}.4-0. 8 krn and that late s urnrn.er activities around areas of succulent vegetation. �154 Nesting Phenology Thirty-five including nests nests 14 in 1978 and 21 in 1979 19 found while hens were laying (Tables were located an unmarked marked were located, hen was accidentally female. days prior during incubation, flushed while locating of the hen and transmitter were located a.ttachrrrerit, At radio at ta chrn.ent by finding transmitters. found when a radio was observed of the nest tail clip. Large numbe r s of body feathers the site indicating with the central possible predation 2 rectrices of the hen. entangled This nest was abandoned after the antenna became resulting in loss of the central signals, date of abandonment accidentally nests The other nest in the sagebrush rectrices. around overstory. wedged in sageBecause of inter- was unknown. were abandoned (1 in 1978 and 2 in 1979) when I flushed were s till laying, still held by the were scattered was found with the transmitter Three One was lying on top of the sagebrush overstory mittent a radio- on the 24th or 25th day of incubation. Two nests brush Fourteen checked for 7 consecutive this time ,4 of 7 eggs were pipping indicating occurred 6). 13 on the 1st day and 1 when This nest was visually to capture 5 and the hens. Two adult females while 1 yearling that abandoned' hen had just initiated incubation. �l,bll' .~. :,c·::.llng I· v I·IH~ ==.,_===== n.«. Ir ,I, ~. . :1:-:.11.1 .1l1-1I1.,·tI 01 r..JdIU-Jlt,ll· U ~.Jgl· [\'IIi.,Il! g r cu -- s c, North tit' ~,tt{'nd,·d 1,1:;1 1,:k r 1""":)1 .tt ,:nd,·d NV ;-';Il Park, CuluJ'o.Idv, . NL1111Uer uf 0.11,' .! AIJr Adull •..l _._._- _ .. Jut:..J I cd c'c~S:i wh c-n l o c a ted 14 ]\\~y -I D.lle ne s t - I'nti. _ ... ====;-':;_~':';-'~:'; .~~:; ... _,,-- .~=.;:.=;::",;=== n~lt(· r n cub a t rou iJq';'lll CLillh :-;i/..· of ncs t l',lt" NO rc n b Spring C'r-eck 2 t.. Ap r 2 7 Apr 9 n Apr 9 Ad"lt 10 "1'1' Spring Creek 1 b )I,!ay 15 May 5 19 lI.!ay t Chicks left nc fit on 15 Jun .\J,lIt !O Apr NO 30 Apr 7 30 Apr 7 Destroyed 5 MOlY on AcLilt 1U Apr B Ma y 22 May 2 30 May " Destroyed 12 Juri on NO 3 May 2 12 May 8 Chicks Idt nest on 8 Jun , Ad~lt ND Sp r ir.g Crc(.'~ 1 ND 24 Apr nest ou 2 3 ~Iay Y e;,s r Irn g 7 Apr Spring 2 Ib Apr 5 May b 5 May b Destroyed 13 May on Y,'arhnt l~ Apr Spring Creek I and Peregrine 20 Apr 7 1\·lay 8 7 May 8 Destroyed 23 May on Yearling 8 Ap r Spring Creek 1 20 Apr 10 May 6 10 May (> Abandoned 14 May on Yearling B Ap r Spring Creek I 15 May 26 May 4 29 May 7 Yec r l ing II Spring Creek 2 17 Apr 2 May 4 9 May 7 ~~ Apr Creek and Spring Creek I Yearling 10 May 22 May b Z2 May L 2 Spring I 12 May 28 May 5 30 May 7 (, May J II May l> AI-.r Creek Spring " Yt:,.trJing I 0 ~.Ia)' .! t,t.I)' Creck ~'-lD 0 I sl 8 l -I 2nd 0 IsI 0 2nu 6 1s t 0 lsi st ,_. 0 151 -Hen k il l e d 0 1s t Chicks left ncst on 25 Jut! 7 2nd Chicks Ic It n e st on 5 Jun 3 IsI 0 Ls t .j !nd 0 I c Chicks ( Ye.lrJ.I!'lg a(ll'I':pl left ;. .-1J.1C f Nl'6t ----------_._- .._-_._---------------_--_. Chi..:l;,,;; :\chai Numbe r o r cgg:j h at ch ed ~J IJ "[';0 = no data. b ' Abe nd onrne nt c a r s r-d by t r a n srnit t o r e nt a n g Lern r-n t in s a g eb ru s h, c Ab a ndcuru e nt CdUS('U by i nvos t i g c t o r . I~ft on Zb Jun 11('51 Ab audc nc d on 19 ~I'.)· -------- sI VI VI �156 T~hlr i,. ="r,c.in~ ~vcnts of radio-ft'tarkr.d female I>."ltr tr:lnSmltt~r .ltt.l.ch~d Age !.S Apr Adu.ll North Park. .~ge grouse. D~te Lckf oS) attended Spring (;reek 1 .ttP.nded last lck Z7 Apr Oate ne.t located & II~y Colorado. Number of ell_ when located 1979. D.a.tC': incu.bation began 1& May :"un\h.·r Clutch stze 9 Urstruyc·d !5 Apr Spring Creek 1 I May 15 May 7 15 May hatc.:hrd on :":,'st .ltt,,".;,1 1,., t Jun re Chick~ Adult fIr I'~~S Fate· ul n~st 1••. n(!st on I sf II Jun A<lull II Apr Spring Creek 1 Z7 Apr IZ May Adult ZS Apr Spri.ng Creek 1 l7 Apr 13 Ma.y & NO NO lien kiU~d on II ~i.l.y ND NO Drstroyrd May Chicks .!S Adult t\or S!)rin~ C eeek 1 Z7 Apr 21 May 3 1'> Apr Sr~ring Creek l b May !4 May l.o Apr Spri.ng Creek 1 I May ZO May 1st .!:"I Jun U<"struy("d Zol May 8 " Irlt j .!O Ma.y un Jun Chicks Adu.lt I'll n('slon Jun l'l Adull on to u 0 I ~t X I ~I 0 1:-.1 •• .!I:d 0 1:-.1 1(,,, nrst on 1(, Jun NO !.7 Apr .'duJt Adult .!1 Apr Adult .!7 Apr NO SprinR Creek Creek l Spring Creek aM Creek l. Yr,J,J"ii.n~ .!") Apr Spring Creek II May NO b b Z-I May 10~y II May NO NO 8 b Cb ick s 1("(t n("st on lCJ Jun 10 May b Jun bJun NO 1 NO l') Apr II May 1 ~nd Spring YearlinK ND Spring .!1 Apr Adult ND NO 1 30 Apr 13M.y b .!O May 18 May b Dc-atroyed .!4 Jun 9 Dcstroy("d 30 May on Io)n .!Iul 0 Chic:ks Irit n("Ii' on !O May lSI I.' t c Jun '!c.1.rhn't NO } May NO ZI May ~ Chicks 1('(f nrst on Zl May Is. tt!. Jun Y("..•. rhnf.; y,.,J,rhnt.: ? May Spri.n)rt Creek ., ,May 1 'J May NO NO lZ: May Abandoned .!8 Mav Ll May on \0 May 0 Chicks 1(·(t oe er on LJ M.y Is t I•• ~ Jun May Yr.a.rhnlo: -; May Sprinjl Crr.ck Z ~J Yrolrh"~ ~S Apr Spring Cre-ek l. 1 May Yr,J.riLnt: .!b Apr Spr,nR Creek Z .!7 Apr Y,' ••.rtlna: ..;.7Apr Yrarhnl( i 7 1'.1n Spring Creoek 1 50 Apr 10 May O('stroyrd o .Iu n 17 May H Deet r cved on 'i Jun I~ May 10 May 14May to May 6 NO May 1~ May Destroyed ':':1 :-10 NO 9 Jun NO O,.:ttroycd .!O May nrllt III ')liD:: no d.l' ••• b:\b.&ndo.tln,.nr c •.u.e<i by lnvt".tts:o&tor. -."'- On 0 I.' I •• 0 lsi May Chicks :-lU on on t st 1,"1t On Jun 1 ~t �157 These hens apparently did not return to their nests Sixteen other hens flushed from their nests Egg deposition were incomplete was determined when found. days approximately apart by 1 day when no egg was laid. repeated. morning for 19 hens whose clutches This cycle, did not vary. where 2 days elapsed (twice by 1 adult) and 3 yearling the cycle laid by hens that were disturbed The interval tion was identified (68.7%) initiated Two yearlings for 16 hens (6 adults a nest. prior again. These a renest eggs were site. and initiation and 10 yearlings). on riests after of incubaEleven completion between laying and start The average of incubation egg. of their of incubation. 2 days after length of time between for both years Two to laying or 'by hens and 1 adult-(.18. 7%) began incubation and the initiation by 2 adults the day following laying of the last (12.5%) remained clutch completion. in the Six instances commenced between clutch completion 2nd clutch _~ith no interval Two yearlings cycles 1st nest and had not selected incubation either was then One hen laid 1 egg then skipped eggs were found lying in the open, not near that had lost their days, 3 days later. between hens. 1 day when no egg was laid before possibly 2 eggs/3 followed A hen that laid late in the morn- ing would still lay an egg in late morning were observed on 2 consecutive or with a 6 hour lag time, The hour that an egg was laid by a hen, or afternoon, being flushed. did not desert. Eggs were deposited 30 hours after end of laying was 1. 1 days. �158 The onset of egg laying for nests was calculated by back-dating 3 days for each 2 eggs. nests located and individuals (P> day was added for the 13 on the 1st day of incubation were calculated for clutches from the day the nest was located using One additional in the onset of incubation. capture found with incomplete Averages individuals to account for the delay for adult and yearling with known dates of lek attendance. without known dates of lek attendance date as the last possible females breeding date. 0.05) were found between the 2 methods but using No differences and data for both groups were pooled. Dates were known when 25 of 35 hens with nests a lek. Nurnb er of days between lek attendance laying varied from 3-13 (Table 7). no difference (.!: > shortest period observed have lost her original still apparently revisit occurred nest prior physically leks for breeding ready and onset of egg Adults averaged 0.05) between 1st and renest 7.6 days with attempts. The when an adult hen believed to incubation to lay. when their nests Three revisited One hen initiated to the lek while other adults did not were lost during ing; 2 took 6 days between nest loss and 'deposition in the 2nd nest. last visited egg lay- of the 1st egg a 2nd clutch only 2 days after abandoning her 1st cl utchv-sYearling visitation hens a ve r ag'ed 9.5 days (range 4-13) from last lek to laying. Approximately 10 days (9.8) were required �159 Table 7. Number days Number of days between last lek attended and onset of egg laying by female sage grouse, North Park, Colorado, 1978-79. Number First nesting attempt Adults Yearlings of of nests Second nesting attempt Adults Yearlings 3 0 0 1 0 4 1 1 0 0 6 2 2 0 1 7 3 0 0 0 8 3 0 0 0 9 0 0 0 1 10 1 2 0 0 11 0 2 0 0 12 0 2 1 0 13 1 2 0 0 7.6 9.8 7.7 7.7 Averages (days) .L �160 for 1st nesting (Table 7). attempts, and only 7.7 days prior The length of time from mating yearling 4690 from 1st to renesting initiated 12 days after started 6 days after of her original female clutch. egg laying (Table Thirteen required Her 1st nest was while the renest was This was 8 days after percent loss (61.5) of all yearling 10-13 days from apparent were followed from initiation Length of incubation Four nests and broods a lek, for mating to 7). nests until hatching. Sixty-two to laying decreased attempt. lek revisitation. sage grouse (11 nests). last visiting to renesting leaving was 25 (2 nests) were observed Only 1 instance possibly left the nest after most hens and broods or 26 days during hatching the nest the morning hatched. of incubation was observed after with hens the 1st chick had when the hen and brood dark on the day the chicks hatched. left nests the day after hatching, Thus 26-27 days from s tart of incubation. Aspects of nesting in the literature. breeding The time interval The shortest while the longest estimate was 7-12--days (Girard 7 -14 days after mating or for combined was 4 days (Patterson 1952) 1935) and 2 weeks (Scott May and Poley (196'9) using radio-equipped hens required have been reported between last lek visitation and onset of egg laying has been estimated age classes. 1942). phenology of sage grouse hens estimated to begin nesting. Rothenmaier �161 (1979) speculated that 1 radio -marked after last being in the vicinity are within my observed Deposition by Girard Reported 20 (Girard 1948, Patterson greater the nest too few times Other investigators (1939) and Keller the following morning. protection has varied to determine (Batterson from predation This varia- date and Morse obse~vation that incubation et al. from of the nest at time of 1963) using frequent and_hens did not leave nests or possibly with the same in my investigation. due to the unknown status Griner nests 3 eggs that 1 egg 1939. Nelson 1955). reported (1948) but observed length for sage grouse 1952, Pyrah eggs were laid (1952) calculated He also observed and eggs of known status 25-27 days. broods Patterson in Wyoming and Morse every other day, 1935) to 29 days (Griner was initiated. estimates day and an interrup- Batterson days that was observed or checking incubation nests days. incubation tion was probably They believed delay each succeeding 1. 3 days. cycle of 2 eggs/3 location (1939). eggs were deposited was laid every eggs was examined laying resumed. laid on consecu.tive All published 4 days data. (1935) and Griner tion of 1 day before believed of a lek. of sage grouse daily with 1 to 3 hours adult hen required and length was (1941) reported that until dusk of the hatching Both investigators was provided day felt that with lessi"ight. �162 Clutch Size Twenty-nine = 16). 5-9) (Fig. 5). 1979 (N (range 6-9), The most completed Average nests number were found in 1978 (~= of eggs per clutch was 7.0 (range Twelve adult hens averaged while yearlings averaged common clutch sizes 41. 40/0of all nests Renesting Females nest, 5). of hens was documented. a renest 1st nests only. eggs per renest. and (3) comparison of birds the initial known to renest Adult females Yearlings, renested however, or nestihg 2 hens (1 ~dult and 1 yearling) which renested located. The adult hen decreased while the yearling increased other adult hens that renested and averaged 6.8 to renest No difference attempts re- to be attempting were documented 7.0 eggs (Table 8). at the and the other both years was found between age classes clutches captured All other hens were considered only in 1978 and averaged . Three (1) the nest after losing or abandoning tinl.e (1 of which was later turned to a lek). eggs, if: (2) the hen had a brood patch when .captured and was later found laying a clutch, same (Fig. (76.40/0). Seven egg clutches to have attempted hen was found with another 5-9) (Table 8). for adult hens were 7 and 8 eggs of both age classes were considered 7.4 eggs per clutch 6.8 eggs (range (66.6%) and 6 and 7 eggs for yearlings comprised 13) and (P > 0.05) in clutch size. Only had both complete clutch size from 7 to 6 .- its clutch size from 6to without revisiting 7 eggs • leks after �163 YEARLINGS ( N=17) 30-; I (J) l- 20 - 10 I I (/) W I f 5.9% Z LL o 11.80/0 ! o i i 5.910/0 ! I I ~4°l a.. 30 1[j DULTS (N==i2) I 20 10 o~--------~~~._~~~~~~~~~~~ 5 6 7 8 _ -9 ,----NUMBER OF EGGS Fig. 5. Percent of sage grouse North Park, Colorado, nests by age class 1978-79. and clutch size, �164 Table 8. Mean clutch size of adult and yearling sage grouse by nesting attempt, North Park, Colorado, 1978-1979. Nesting attempt by age class 1978 N Years N 1979 1978-79 N Adults 1st 8.0 3 7.8 4 7.9 7 2nd 6.0 2 7.3 3 6.8 5 All 7.2 5 7.6 7 7.4 12 1st 6.5 6 6.9 9 6.7 15 2nd 7.0 2 7.0 2 All 6.6 8 6.9 9 6.8 17 1st 7.0 9 7.2 13 7.1 22 2nd 6.5 4 7.3 3 6.9 7 All 6.8 13 7.2 16 7.0 29 Yearlings All Females �165 loss of their different 1st nests nests. cumulatively All 2nd clutches laid 10, 12, and 14 eggs in 2 (range 6 -8 eggs) were larger than the smalles t clutch found. Clutch size has been reported to 8.2 (Wallestad Pyrah and Pyrah 1974) (Table 9). (1974) using radio-equipped separately for both age classes. (range 7-11), Clutch sizes (Table 9). Patterson to vary from 6.8 (Griner and 9 yearlings averaged Renesting renesting to all previous investigations. that lost their and sizes 9.0 eggs 4-10). as being most documented, common although of small from my study was similar However, Batterson 1st nes t prior averaged might occur because Clutch size determined fied by clutch size alone. clutch 6.9 eggs (range has not been previously late clutches. nests. adults of 7 or 8 eggs have been reported (1952) believed in another Only Wallestad hens documented Thirteen 1939) renests and Morse cannot be identi(1948) believed to clutch completion would start hens laying nest and would lay a total of 12 to 14 eggs in the combined This hypothesis was supported by data from 3 adults that renes ted in my study. Length and Width. clutches - - Length and width of 217 eggs from (16 adult and 17 yearling hens) were measured a~ 33 averaged ,~- 56. 1 x 38.7 rnm with ranges tively. Greater variation of 51. 0 -62.0 and 36.0 -40. 9 rnrn, was noted in lengths respec- of eggs than in their �Table 9. Sage grouse clutch size in western North Arn.er ica , Authority Griner Location (1939) Utah Clutch size Mode Range 6.8 .7 1-9 Avg. - Number nests 147 Keller et al , (1941) Colorado (1940) 7.5 7 -8 6-10 34 Keller et a l , (1941). Colorado (1941) 7.6 8 5-9 35 ~ 1 of Patterson (1952) Wyoming (1949) 7.3 - 5-9 80 Patterson (1952) Wyoming (1950) 7.5 - 5 -13 74 Nelson (1955) Oregon 7.1 7 6-9 23 Walles tad and Pyrah (1974) Montana 8.2 - 4-11 22 This study Colorado 7.0 7 5 -9 29 ..'"'" �167 widths. A difference lengths (P < 0.05) was found between years for average but not for widths. Eggs from adults mean lengths 38.8 nun, (N = 105) averaged 56.3 x 38.9 mm with and widths in 1978 and 1979 of 55.8 x 39.2 and 56.6 x respectively (N = 112) averaged (Figs. 6 and 7). Eggs from yearling 55.8 x 38.5 mm with mean lengths for 1978 and 1979 of 55.0 x 38.1 and 56.4 x 38.8 mm, hens and widths respectively (Fig s. 6 and 7). Differences classes (P < 0.05) were found between years for age in both length and width, only found for the 1s t year differences adults and combined could be related and yearlings, on the hens during colder and wetter 1979. Therefore, but differences years to physiological The winters than normal for length. readines egg size could be related the winter. (P < 0.05) were With both to the stres s placed of 1978-79 were both with 1978 being less eggs laid during s, These severe than 1979 should have been smaller than those laid in 1978. However. creased. in size could be due to the increased This reversal of time which allowed yearling tract. Wit?- a longer following year, period yearling even with a more Severe egg size of yearling hens to develop to laying the hens would be able to lay larger in- length the reproductive of time from hatching winter. females eggs, �168 j 1978 ~ I S.D. ""0 -::JI o MEAN 2 S.E. ~ 55 T -=1 - ~ ( N=47) (N=33) ~ 50 _' ------------~------ ., J 01 I 6 ~ :l ~ 55 ....J '- 1979 i ~ ----liIii l<N=72l (N=65) ~ 50-~._---------------------------=-------1978 and 1979 T -- 55 (N= 105) (N=1I2) 50~-----..------..-------------------------------ADULTS ~Fig. 6. YEARLiNGS Length of eggs from adult and yearling North Park, Colorado, 1978 -79. sage grouse hens, �169 1978 1S.D. MEAN .. 38 1 2 S.E. ~: l (N=331 _L l 35~----------------------------------(N= 47) :~~- ~ E E 39 -:I: 38 5 37 ?: 36 1979 t (N = 72) ( N = 65) 35~-----------------------------------1978 and 1979 41 40 39 38 37 36 (N= 105) (N= 112) 35 ~------------------------------------ADULTS Fig. 7. YEARLINGS Width of eggs from adult and yearling ..--North Park, Colorado, 1978-79. sage gr.@se hens. �170 Measurements of sage grouse Wyoming and the United States et al , (1918) examined an average Museum (Table eggs in the U.S. National Museum 10). and reported Nelson (1955) working in Oregon reported Eggs from Wyoming measured All length and width means than those in my investigation. due to subspecies iN eight. variation egg by Patterson previously These reported differences were could be or to sampling. - - Two hundred t ion, and an additional average (1979) were 55.0 x 38.0 and 56.3 x 37.8 nun, (1952) and Rothenmaier respectively. Grinnell of 54.9 x 38.1 nun for length and width, size of 58.2 x 37.6 nun. smaller from Oregon, National measurement respectively. eggs were available eggs were weighed prior 16 were weighed near to incuba- the end of incubation. The preincubation mean weight of all eggs was 46.1 g (range with no difference (P 40-55) > 0.05) between years for the 71 eggs in 1978 and 129 eggs in 1979. Eggs from adult hens averaged with those (24) in 1978 being heavier 47.0 g for the combined (49.0 g. range 44-55) those (71) in 1979 (46.3 g, range 41-52) (Fig. (P < 0.05) between years since weight is related from I st and renest 3 renests Eggs from the renest than 8). The difference was due to the difference in egg width, more to width than length (Hoyt 1979). clu tch.ee of 3 adults were lighter 2 years were weighed. tha:r;r::--eggs in 1s t clutches of 1 additional from Eggs Eggs in the the same hen. adult hen were weighed; the �10. Sa g e grouse Libl" Authority egg length, Location Grinnell Avg. width and weight Length(mm) Max. Min. rn ca su r ern en t s I r crn western Avg. Width(mm) Max. Min. North Arn e r ic a , Preincubation 109 54.9 37.6 Wyoming 55.0 38.0 Oregon 58.2 37.6 Wyoming 56.3 60.3 53.6 37.8 40.6 35.5 Colorado 56. 1 62.0 51. 0 38.7 40.9 36.0 Patterson (1952 ) (1955) a 40.5 b 36.') lH 30 Ro th c nrn a ie r (1979 ) This study aWeights b Weights f r orn 18 eggs. fdim 40 eggs. CWeights from 200 eggs. II. ,_. ," _. et a l , (19(8) Nelson Nurnb e r of eggs Weight(g) Development Early Late 41.0 46.1 c 34 217 �172 1978 55-, j -t ::]§ (N=24) 40--1. ~~ ~(N~=_4~.7~) _ 1979 55-, 3 _ 50-=i ~45j - ~ ~ 3 4°1~ !!'• . i.' j_ (N__=7_'_) ~(N~=_5~8~) _ 1978 and 1979 55 50 45 40~~_~ (~N_=_95_) ~(N_=_IO_5_) ADULTS YEARLINGS . Fig. S.' Egg weights North Park, _ ..-'- from adult and yearling Colorado, 1978 -79. sage grouse hens, �173 average weight of these eggs (53.8 g) was the heaviest examined. Decrease in egg weights for renests since laying hens use large amounts for all clutches would be expected of minerals and energy for the 1st clutch. The mean weight of all eggs from yearlings 47 eggs in 1978 averaged 44.6 g (range weighed in 1979 which averaged difference Eggs of 1 yearling egg weight increased This increase clutches should require A pos sible reason of yearling in renesting required to initially experience. The the maximum (P the 1st to the renest and minerals produce as an adult. tracts eggs that are as large With maturation, reproductive larger eggs Since the 1st clutch may not have amount of minerals eggs in renests and energy due to smaller cou ld be slightly < 0.05) between age classes This difference laying 2 was that reproductive would be able to produce than in 1st clutches. combined years. difference. 8). were weighed, since yearlings as much energy females eggs and clutch size, difference was unexpected may be unable attempts 0.1 g from for this observation as a hen with previous tracts than the 58 45.8 g (range 42 -52) (Fig. from both 1st and renest clutch. of yearlings lighter The < 0.05) between years was due to width difference.' (P and average 40-50), was 45.2 g. larger. A was found for 1978 and the corresponds to the effect of width �174 Eggs of an adult hen exaxnined on the 19th day of incubation averaged 38.4 g (range unknown. eggs of s irn.i.Iar size from other clutches estiInate a probable 38-39). Since preincubation preincubation weights were were used to weight of 43.5 g. This suggests a loss oi 5.1 g of weight occurred during the 1st 19 days of incuba- tion. hen not exaxnined until the 24th Eggs in a nest of a yearling or 25th day of incubation preincubation average averaged being estimated a 10S S of 5.2 g (11. 40/0)occurred Water loss accounts 40.3 g (range 40-41), at 45.5 g. with the This suggests during the 25 days of incubation. for the majority of this weight loss (Rahn and Ar 1974). Little water loss apparently incubation with an average occurs in the late stages of of 11-120/0weight loss of eggs occurring during incubation. Sage grouse gations. egg weights have been reported Patterson (1952) and Rothenmaier (1979) working in Wyoming weighed eggs during early development of 40.5 and 41. 0 g (Table 10). in late development These authors and reported did not report time of weighing. Patterson in 2 other investi- and found averages (1952) also weighed eggs a mean of 36.9 g (Table 10). how far i.ncuba t ion had progressed Consequently, comparisons at are not feasible. Success ,~- Nesting of total nests success located was estimated for nurnbe r of nests and number of radio -tagged female hatching sage grouse �175 of all radio -marked hens that incubated nest was considered Fourteen hatching marked successful of 35 nests if at least of females lost their The difference 1st nest renested. success At least were eliminated, hens renested. since success Renesting success and success 33.3% of all hens that likely to renest as a rn in irnurn of 55.5 and 16.7% renested, killed by predators the success of radio- in 1978 (50.00/0) than Adults were more If hens that lost transmitters and yearling more Success between nest was due to rene sting. than yearlings adults with slightly in 1978 (42.9%) than in 1979 (38.1%). in 1979 (44.4%). 1st nests in an increase respec- and females 71. 4 and 20.0% of the adult was most important from 30.8 to 63.6%. rate of all yearlings resulted after Renesting increased A 1 egg hatched. (40.00/0) hatched hens was 46.7% with higher tively. a clutch to cornpl.ati on, improved from 29.4 in chick production for With renesting, to 46.'1%. of 28% (23 birds). Nest losses (N = 7). were caused by predation Destroyed of predation nests were examined following descriptions Ground squj r r el s (Spermophilus portant nest predators and coyotes respectively (N = 14) or abandonm ent for evidence in Einarsen richardsonii) (20.-0% of all nests). and method (1956) and Gill (1965). were the most Badgers im- (Taxidea taxus) (Canis Iat ran sr=caus ed 11.4 and 2.9% of the nest lost, (Table 11). Loss of hens to unknown predators accounted �Table 11. Sage grouse nest loss by source and year, North Park, Colorado, Source Predation of hen Ground squirrel Badger Number of nes ts 1 2 0 1 Ufo of all nests 7.1 14.3 0 7. 1 2 1 1 9.5 4.8 4.8 7 4 1 2 20.0 11.4 2.9 5.7 Year , Coyote 1978-79. Abandoned Totals 1978 1979 \ , Nurnb e r of nests 6 a 4 28.6 a 3 8 57. 1 13 •.....• Ufo of all nes ts 28.6 14.3 61. 9 1978-79 Number of n e s ts % of all nests -- alnc1udes 2 caused by the investigator. 7 20.0 21 60.0 -....J '" �177 for 5.70/0 of the nest los s. incubation was destroyed, The nest of 1 of 2 females apparently by a coyote; other hen lost during laying was not destroyed. successful nests disappeared eggs in 1 nest were believed was flushed by cattle after initiation in 2 unsuccessful nests disappeared (20.0% nest loss (Table 11). of incubation. Two The last since 2 additional to destruction eggs by predators. was an important interference) were abandoned. abandomnent snow (4.8%) were observed, eggs to be displaced eggs from cause of 1£hurnan-a ttr ibuted nest loss (transmitter (3 of 31) of all nests tracks prior of all nests) and investigator were: Three feeding in the vicinity of her nest. taken by a ground squirrel, tanglement the nest of the to have been kicked out when the hen egg was possibly Abandonment killed during was eliminated, Other factors from the nest. only 9.70/0 causing nest as the nest was covered and flushing by a predator en- (4.8%) and no cau-sing The cause of abandonment of 1 nest was unknown. Reported from 60.3 Morse 30.4 nest success in Utah (Griner 1948). 1939) to 23.7% Six other investigations «nu rsss), and 35.:3 (Keller and Pyrah based on nest searches 32.2 et al. (Dargan 1940), - has varied in Oregon (Batterson reported 34.0 nest success (Patterson 1952), 1941-), and 39.2% (Nelson 1955). (1974) working with radio -marked female and of 35.0 Wallestad .= sage grouse �178 (N = 14) in Montana reported hen success Causes John area et al. reported of North Park, of all nests Colorado that badgers However, taking single eggs from source of nest destruction only slightly less Wyoming reported tridecemlineatus) while badgers reported destroyed predator accounted magpies ravens being responsible badgers destroyed Abandonment (30.4%). (Phasianus Patterson Batterson ravens colchicus) were a major with badgers being (1952) working (including 42.60/0 of all nests including cause ground squirrels in Spermophi1us lost to predators and Morse (Corvus (Pica pica) and crows (Corvus 54.2% of all nests (1955) reported pheasant for 36.2%. of 41. 7% egg shell than known (i956) observed that ground squirrels destroyed Keller were not an important (34.80/0 of all nests) important in the Lake for destruction that ground squirrels that avian predators black-billed examined had 1 less ring-necked Gill (1965) reported and 44.4% for yearlings. accounted Einarsen nests. They also in 1940-41 and 1964. many nests to have been in the nest. of 63.6%. were extensively and that ground squirrels of nest loss. success of 76.9% for adults of nest losses (1941) stated nesting (1948) corax), brachyrhynchos), lost in Oregon with no single mammalian for more accounted than 7.6% of all nests. for 33.3% of all nests Nelson lost. while 7.8% of the nests. was an .-rrripo r tant factor in Wyoming as 20.0% of all nests affecting were deserted nesting (Patterson su c c e s s 1952). �179 In Montana, Wallestad tion wi th 2 (9. 10/0)caused by entanglement Griner (1939) found Batterson by Wallestad These rates as to completion. of abandonm.ent, nests nests of inaccuracy Patterson (1952) observed were collected in classifying squirrels be credited With secondary within 1 week. Another possible is secondary success. cause destruction. were examined after nest completion. destroyed scavenged by badgers predation, with nest destruction tion of nesting Destruction in my study since eggs in nest predation (42.9%) were later were classified 31. 00/0of the deserted that had hatched (14.30/0)and 3 nests nesting year. of the year, within 1 month. was not documented Nest sites in my investigation Two nests except that reported located during searches nests found in 1 year were destroyed deserted only 6.00/0and Nelson (1955) and Pyrah (1974) which used only nests could be low since all nests of deserted of the radio harnes s. 14.30/0 abandomnent of all nests in Utah while and Morse (1948) reported 9.80/0 in Oregon. 22.70/0 nest deser- and Pyrah (1974) reported Success incorrect resulting rates by ground during that species in incorrect may determina- based on nest search data may be low. Hatching S~ccess The percent of eggs hatching in 14 successful 96 eggs was used to calculate success hatching su cce s s , was 85.40/0for the entire investigation, nests .~ £_ontaining Average hatching with success in �180 1979 (50 of 54 eggs, 76.2%). Hatching was greater 92.6%) surpassing success fertility > 0.05) for yearling to be related fertile to snow storms developrn.ent. (73.7 -96.2%) and that occurred during laying, growth prior Hens with clutches of eggs that showed any embryonic when snow storms The remaining to death (8.3%). 8 eggs had some Thus egg fertility (P > 0.05) between adult and yearling completed hatching success occurred prior to snow storms (42.9 -75.0%). More snow storms variation in temperature also occurred Greater in 1978 which could have success and fertility were examined by Griner that 94.8% of all eggs were fertile (1952) found 96.9"% egg fertility and hatching Nelson (1955) stated that hatching success (1939) and hatched. success . 94.5%. daily death. in Utah who reported Patterson hens. (80.0 -100.0%) than hens still laying during the peak of laying in 1978 than 1979. embryonic was 91. 7% in both years occurred Hatching Egg All eggs that did not hatch (N = 14) were examined; with no difference caused in 1978 eggs did not hatch in 1978 than in 1979. only 6.3% (6 eggs) were infertile. had higher (86.7%) between The lower rate of hatching was based on the number embryo females than eggs laid by adults (84.30/0). Variation adult hens (78.3-89.3%). since more in 1978 (32 of 42, of eggs laid by yearling 1978 and 1979 was slight (P appeared success - of. was 97.60/0 in �181 Oregon. Data from my study indicate similar fertility, but lower hatching success. Nest Construction Width. depth and materials were classified (nearest for 31 nests. us ed in construction The depression and lining was measured twice cm) for width at 90° angles and for depth to the nearest 0.5 ern; Nests were round to slightly oval in shape with none being out-of-round by more than 1 ern, ranged from 13 to 22 em; Mean depth of the depression and ranged from 4. 0 to 7.5 ern, age classes No difference (P was 5.5 > 0.05) between was found in size of nests. Material used in construction included dried grass, The hollow appeared a central Average width was 16.8 and leaves, of nests was entirely and stems and small twigs of sagebrush. to be formed by moving litter point outwards litter and humus from by either use of the feet or body since the sides of the nest were slightly higher (1-2 cm) than the_surrounding -_;- exterior litter. by the thickness made. The depth of the depression of the litter layer and time the measurements Shallow nests were common in areas when nest; were measured number of body feathers (> 40). was poss ibly controlled Feathers of low sagebrush, while the hen was still laying. lining nests varied from few « added to nests were possibly loss during formation of the brood patch. were and The 1_Q) to-rrrany .::::- done passively due to Support for this hypothesis �182 was obtained by comparing First nests invariably hens were actively some 2nd nests material 1st and 2nd nests had many more adding feathers of the same hen. feathers than 2nd nests. into the nest depression, should have had as many feathers was noted as being brought If then as the 1st. No into the nest from outside the area. Nest construction reported Griner nests was 1st examined by Girard as being oval in shape measuring (1939:19) also reported similar slightly manner 7.6 cm. sagebrush mentioned approximately twigs and moss feathers. during egg-laying supports with use" and believed They also reported 30 to 40 breast II or any evidence of nests and Morse in a (1948:12) 20.3 cm wide with a depth of were "lined with dry grass, if available, Griner studies, 20.3 x that hens built nests Batterson nests and incubation most previous other material. He found that "capacity to dust bathing. found round nests 22.9 x 20.3 em. oval shaped nests measuring 17.8 ern with a depth of 5.1 ern; increased (1935), who and after (1939) believed incubation started, feathers were lost to form a soft lining. My data but I found no nest as large that indicated as a:ctive lining of nests with �183 Chick Growth Chicks from 12 broods (6 adult and 6 yearling captured the day they left the nest. hatching (N = 65) was 30.7 g. Average weight of chicks In 1978, 22 day-old 31. 0 g, and in 1979, 43 day-old hens) were chicks averaged chicks at averaged 30.5 g. Thirty- five chicks from adult hens had a mean weight of 31.3 g (range 28-36), 32.5 g in 1978 and 30.8 g in 1979 (Fig. (P < 0.05) between years chicks from yearling was probably hens averaged in 1978 and 30.1 g in 1979 (Fig. between years difference between have been due to more to yearlings. would occur hypothesis an automatic was experienced Less if hens remained was obtained incubator 9.9% greater 29.8 > 0.05) hens. egg water on nests corresponds behavi.orjof loss during longer. to may adults incubation Support for this when 11 eggs from adult hens hatched produced chicks of chicks from adult hens may enhance averaging hens. their A but not in The weight difference incubating g of chicks from pooled data, than those hatched in the wild. chicks from yearling (P of yearling weights of chicks and widths. Thirty 24-33); No difference in 1978 and the 2 -year that found in egg weights compared 30.0 g (range (P < 0.05) was found in mean weights The difference The difference due to egg size. was found in weights of chicks adult and yearlings 1979. 9). 9). 34.2 g. in This Heavie:t:_wetghts survival .~ over that of �184 1978 I S.D. - ( N=IO) ---..i.._ (N=12) - .... 5~ .::) 1979 f J ; j ~ 30j ~ 25~~: (_N_=_25_) (_N_=_18_) 1978 and 1979 35 - 30 I~ j 25~ ,._~__ (_N_=3_5_) ADULTS-Fig. _ 9. --(N=30) _ YEARLINGS Weights of 1 -day old juvenile Colorado. 1978-79. sage grouse, North Park, �185 Weight gain was estimated recapture wi th known weights weight of chicks leaving recaptured 46 times they left their nest. of day -old chicks the nest in 1978. of o. 6 tained prior females. chicks were Three o. 0-1. Weights then increased than 22 days. Weight gain/day differences was Sex could not be ascer- to 48 days of age and the data reflects Sex related of approximately 6. 1 g/ day (range 0.8 -2. 3) at 6 or 7 days of age to at ages greater day-old 0) from the date at a rate 12). at time of in 1979 and mean Thirty-six g/day (range 5.9 -6.9) at 21 to 22 days of age (Table not estimated weights from 3 -48 days post-hatching. chicks lost an average I. 7 g/ day (range by comparing in growth rates both males may occur and and could bias the data. Wing measurements were recorded to 48 days of age. Emergence II days, but more commonly emerged at 22 to 26 day s, of juvenile Replacement 23 days, at approximately Emergence about 38 days, while juvenile Juvenile of juvenile while juvenile P X P I with adult P It-was replaced of adult P III occurred P IV was 'replaced of adult pr~_maries was approximately from 6 P IX began as e a r.Iy as at 13 to 16 days. P I began at approximately 31 days. for juveniles 6 mm/day. at 45 days. These at Growth dates were for pooled s exes and may be bias ed, Pyrah rates (1963) working with captive of weight gain and primary birds replacement in Idaho r ecor-ded pattern by sex class. �T~bll' 12. Juvenile sage grouse At;" IV "ight ( da ys ] (g) _._-----(> tJ (, 3') 39 b 40 6 40 44 c 7 7 8 9 9 10 LO 10 II 11 11 13 13 13 13 13 13 14 14 15 18 18 18 21 Z2 22 Z2. 22 31 31 31 48 aE = gain (g/Jay) 3') 33 (, I. s ·16 44 49 52 53 57 66 57 75 76 31 83 84 79 84 80 77 86 88 In 149 L66 159 102 156 184 166 300+ 300t 300t 300t = quill. A Co lo rad o, 197!l-79. P'r im a r y 1l'llgth (lIUll)d. -X---IX----';i rr: --vrr ---Y·I---y----i'\;-----·iiC----·-il----·-i.. _------.--_. __ ._--_._._-----_._._---_._------------.. _._,-----_ .._.- -- 70 74 n 80 84 88 94 97 98 97 100 101 102 94 103 97 132 130 132 -- L41 143 151 142 177 181 181 232 = Park, Carpel l e ng t h (ruru] n ----- North data, 5') 51 -l9 60 58 6L 6·1 65 63 0.8 1.8 1.3 1.7 1.7 2.3 1.9 2.1 1.5 2.0 2.6 2.1 2.5 3.8 2.7 3.9 4.0 3.9 4.1 4.2 3.8 4.2 3.8 3.1 3.9 3.7 7.8 6.5 7.4 6. I 5.9 5.6 6.9 6. I H ernpt y, 0 wl~lght AV(!I'..AgC growth adult Icathor , L E _:tl E E E E I:! L (l( .l) E E E Ol~) (Wi) (~I 3) E (l( 5) E E E E E E E E 00) E E E E E E E E E E 30 25 24 11 LO 13 9 17 53 54 S4 I!O 0(1) 8 14 10 13 10 12 18 19 20 31 32 28 32 31 28 25 25 20 50 49 50 68 E E E E E E E E E E E E E E E E E E E E E E E E E E E E E E E E 0(2) 22 2-1 Z4 82 n 78 70 75 107 100 IOu IlI2 .!H l3 1 I) 31 30 33 33 32 31 -10 40 36 43 38 45 51 53 54 58 62 59 62 64 67 58 62 59 82 85 85 94 97 99 98 98 129 121 I! I Iv'! is :lH 3') :I') 2') 25 37 33 38 38 H 35 48 45 44 49 46 52 59 59 62 67 69 66 69 71 70 66 69 68 9Z 96 95 99 103 104 107 104 I 33 Il<1 ll~ Il'l 32 27 40 ,11 43 43 46 40 52 50 46 52 50 56 61 65 67 71 73 33 30 ·11 40 33 n 73 7S H 70 75 ·12 46 4(, 42 51 51 50 54 52 58 62 67 69 71 73 73 73 75 7S 70 76 n n 95 97 97 104 105 lOb 109 106 I' - I 1-,-I 1'- I 151 96 95 96 102 103 103 110 103 1!4 IlK lUI E 39 -H HI 33 31 ·10 38 ·10 H H )0 ~I 46 ,15 44 H 4') 51 50 53 52 57 60 65 68 69 71 70 70 n H 68 75 71 90 90 n 95 94 93 105 93 10<1 102 10! 5tiA 4tl 50 49 53 51 57 57 63 67 66 66 66 66 67 71 66 71 69 85 85 86 83 83 81 95 79 E 100 100 107A leI 32 32 40 37 39 42 -B 43 45 49 47 51 50 53 5S 61 64 58 59 61 59 61 66 59 67 64 78 78 79 76 75 -, ,~ 71l 71 53.-\ 5".-\ ~uA 141A •.... CO ()\ �187 He found no difference His records between sexes up to 21 show similar to 22 days of age. data for emergence, but slightly faster growth through 48 days of age. Behaviors Chick survival was not examined due to human interference. broods of Brood Hens during the investigation Since radio-marked hens and their were located at 1- to 2 -week intervals, brood scattering and increased were able to collect flushing the hen, chick distress mortality and measuring call was used to attract hens would fly toward the distress hens would rarely Many adults females fly towards after flushing concealed truder Adult hens than yearlings. all chicks, a tape-recorded the hen to the chicks. call and cackle whereas gave a broken wing display. displays Adult females brood and used a flutter adult hens still had most had chicks remaining remained yearling Yearling in the vicinity flight display to distract than of the the in- By late chicks while few yearling females with-them. Two hens (1 adult antf I yearling) left their nests Adult and most flew more when a chick ran from 1 hiding place to another. summer After the call and many would not cackle. did not give distraction 1 krn upon flushing. of chicks. their broods much easier capturing this may have caused the rrio r m ng after with newly hatched a storm deposited chicks 4 cm of snow. �188 The adult hen was found broodi.ng 40 m from the nest with all chicks that hatched. 10m The yearling was not brooding from the nest and apparently days later any chicks had lost her entire the adult still had at least 7 of 8 chicks. support the hypothesis through the 1st summ.e r than those of yearlings. to learned survival behavior that chicks of adults and experience. may be learned with broods during by unsuccessful f ied by short sites 13). (N sagebrush Eighty percent = rate This may be due important to brood hens associating 40) used by female were in sagebrush including ment of radios, pre-breedip.g. these The other with hens than 20 -crn, of 10 em or less and nesting hens attended were typi- in height 3.6-16.3) for periods. Since leks after attach- of 21 of th eLocat ions could be considered 19 sites to nesting. groups. height of less areas pre-breeding a minimum and prior sage grouse height was 7.6 em (range 54.8% of the transmitter-equipped between at a higher Analysis with average The average all locations, breeding examples Sites Roosting (Table Six the summer. Vegetation Roosting brood. These survive Behavior when found therefore No difference were areas (P used after > 0.05) was found �189 Table 13. Frequency Class (em) of sagebrush height classes used by female sage grouse for roos ring. feeding, and loafing and feeding sites, Colorado, North Park, 1978-79. Ro os ting sites N 0/0 Loafing and feeding sites N 0/0 0.0- 5.0 12 30.0 5.1-10.0 20 50.0 3 10.1-15.0 7 1705 15.1-20.0 1 2.5 Feeding N sites 0/0 5 10.4 2.5 11 22.9 19 15.5 16 33.3 29 23.8 6 12.5 20.1-25.0 19 15.5 7 14.7 25.1-30.0 20 16.4 3 6.2 30.1-35.0 8 6.6 35.1-40.0 10 8.2 > 40.1 14 11.5 122 100.0 48 100.0 Totals 40 100.0 �190 Leks Transmittered the study area. marked hens attended One of the 2 largest in 1978. attended Examination by radio-marked of mating areas last week of May 1978. Thus, the end of the breeding averaged (range 3. 0-10. 3) of the mating greater than 20 cm in height. small 5.3 cm (range islands of taller area sagebrush area some protection from avian predators the major on all leks. source Only snakeweed bushes and black sage- No sagebrush seemed were present This taller than 26% (range 21.1-2.5.8) was done in height and covered (Table 14). The taller during the description Big, alkali, of each breeding had less hens. the vegetative 3.2 -7.2) by 1 yearling was examined was accomplished season. brush, Peregrine and it was only visited areas near sagebrush for hens. phlox (Phlox byroides) was t}:lecenter may supply All mating ground cover. 7.3% important areas Gr"asses were of ground cover on 2 of the 3 leks (Table and moss in by all radio- the 2 years. Cons equentl y veg eta ti.on of mating on only the 3 leks visited since leks was visited hens known to attend leks during was the only smalliek near 3 of 6 leks known to occur 14). were found on all 3 leks. Loafing and Feedin~ Sites .~Localions identi.fied as loafing and feeding areas used by hens during egg laying. were those Data from the 122 sites examined �Table 14. V g ti species c et a v Percent vegetative cover on sage grouse mating e Spring Creek 1 sites, North Park, Colorado, 1978-79. Leks --:::---:=---:'~--------::::Spring Creek 2 Peregrine 1 Sagebrush (Artemisia 'I spp. ) 7.6 (5.5) a Fringed sagebrush (A: frigida) Snakeweed (Gutierrezia 3.4 s a r o th r a e ) Stonecrop (Sedum 10.3(7.2) 0.9 1.2 1.8 1.7 ,_. ,_. 3.0 pumilus) s tenopetalum) Grasses 4. 1 1.2 O. 1 0.3 O. 1 O. 1 11.9 10.2 8.8 25.8 21.1 23.4 {. Totals a b Average blncludes '// ' height species a \0 Moss phlox (Phlox byroides) Low daisy (Erigeron a 3.9 (3.2) (cm). not listed that occurred on only 1 Lek, �192 were variable. No areas than 5 ern and only 2.50/0 heights of less 10 cm or less. cm class (Table 13). During periods had average in areas of sagebrush These areas when fresh of hens were followed to identify areas approximately occurred use (55.70/0) The greatest in the 15.1-30.0 for feeding. were used where sagebrush with areas were preferred snow was present. used. was Sagebrush tracks bushes 30 em in height were conunonly walked around as the birds apparently The most conunonly used class 20.0 cm (23.80/0) fed on sagebrush (Table 13). leaves and old flower heads. of sagebrush Areas of taller height was 15.1sagebrush than 40.0 em) were conunonly used during inclement (greater weather. Nest Sites Nest sites were described slope and aspect. Average was 32.3 cm (range nests the nests sagebrush 11.1-59.8). were in sagebrush sagebrush as to sagebrush less height at nests height surrounding Approximately was 52.3 cm (range 27-76), of 35 nests greater were under sagebrush -found under big sagebrush Sagebrush cover at nests was not uniform. averaged However, and 600/0 of than 50 crn , bushes; except 1 found under silver One nest was under a corrrp l ex of big sagebrush 35 nests 520/0 (51.4) of all than 30 cm tall (Table 15). were under bushes with heights Thirty-four cover and height. all were sagebrush. and rabbitbrush. 23.6% (range 9.4-42.6), but Many nests had 1 or 2 sides open, possibly for �193 Table Class 15. (em) Average height (em) of sagebrush surrounding Colorado, nests, North Park, 1978-79. Canopy cover Number of nests 0/0 sage grouse Sagebrush Number of nests height % 10.0-14.9 0 0 2 5.7 15.0-19.9 0 0 3 8.6 20.0-24.9 0 0 6 17.1 25.0-29.9 1 2.9 7 19.9 30.0-34.9 1 2.9 3 8.6 35.0-39.9 4 11. 4 3 8.6 40.0-44.9 4 11. 4 5 14.3 45.0-49.9 4 11. 4 3 8.6 50.0-54.9 4 11. 4 1 2.9 55.0-59.9 6 17.2 2 5.7 60.0-64.9 4 11. 4 0 0 65.0-69.9 5 14.3 0 0 70.0-74.9 0 0 0 0 75.0-79.9 2 5.7 0 0 35 100.0 35 100.0 Totals �194 escape sites r ou.tes , No differences chosen by adults with more and taller Slope at nest on slopes of less This selection of the area > 0.05) were found between nest and yearlings. sagebrush sites Adults normally varied was apparent were on north facing slopes sagebrush on north more sagebrush of 6% or less was an indication nests was taller of the topography No selection occurred and more (1974) reported ern, because dense, therefore that average at nests 1969) to 50.8 ern (Patterson has varied 1952). from o"ther investiga- of 35.0 (May 1970), 41. 4 (Poley 1969), 42.9 (Gill 1965), and 48.3 that nests height of sagebrush than that of the surrounding height of sagebrush heights height of Data from my study.supports c ov e r was taller tors found intermediate also reported 10). 57.10/0 of all nests This apparently was only 23.4 Average 20.3 (Klebenow However, was 40.4 cm and average the Ii.ndi.ng that nesting sagebrush. 10). (Fig. sites. and Pyrah at nests surrounding slopes nest Wallestad (Fig. slopes. females. from 0-360/0 with 85.80/0 of all nests than 120/0and 60.00/0 on slopes for gentle used areas than those used by yearling as much of the land was gently rolling. for aspect providing (P cm (Nelson were under plants 1955). taller Klebenow (1969) than the surrounding vegetation. Comparison Wallestad of percent and Pyrah (1974), sagebrush cover could not b~ made with since they counted the number of sagebrush �195 • ADULTS .6 YEARLINGS Fig. 10. Aspect and slope at nest sites of adult and yea?1ing sage grouse, North Park, Colorado, 1978 -79. The center equa.ls 0% and each concentric circle 3% slope. Two nests of adult hens on 35 and 360/0 slopes are not plotted. �196 plants within 61 cm and 9.3 m nests were typically found no nests brush 2 around the nest. in open areas in dense or on edges of open areas sagebrush were used sparingly May (1970) stated areas. Dense stands and of sage- in my study if an opening was adjacent to the nest. Aspect conducted and slope of nest in North Park, May (1970) all reported found 91. were examined Colorado. Gill (1965), no preference oi less than 10%. of nest sites were on that the average slope Areas classified as feeding areas bating hens whi.Le away from nests. by following 14 different hens from (P > 0.05) between areas used, brush in these areas less Poley (1969) 85.7% of the nests Gill (1965) stated and was 2.0%. Locations ference in 3 studies Poley (1969), for aspect •. 70/0 and May (1970) reported slopes Feeding sites These 48 sites their nests. adult and yearling and the data were pooled. 13). females was no dif-in feeding height of sage- with 66.6% being in areas The average =: 3.8-27.9) were located There The average was 30 cm or less than 15 cm in height (Table 14.2 cm (range were those used by incu- for all locations. height was �197 RECOMMENDATIONS 1. Counts of numbers of hens on leks do not provide an indication of the number of females in an area. They are useful only for indicating the peak of breeding and when hatching will occur. 2. Nesting areas related should be identified, to leks. These areas since they are not closely are related to vegetation height and COver and not to dis tance from a lek. 3. 1£ estimates of production of juveniles is desired, be conducted after vegetation desiccation counts should and near meadows. �198 SUMMARY Intensive female investigations sage grouse were conducted Data were collected marked April of breeding early Adult females nest, Colorado. and 23 yearlings) and located 1 lek 89.90/0 attended 1 to 5 with adults yearlings ecology of radio- each morning from June 1978 -79. vis ited 1 lek 81.80/0 of the time. from in North Park, on 42 hens (19 adults with 14-15 g transmitters through and nesting visiting Number while yearlings of days on leks varied 1 (44.4%) to 3 days (33.3%) and 1 (45.4%) to 5 days (9.1%). the same lek was visited of the time, When hens attempted to re- for 1 day by adults .and 1 or 2 days by yearlings. Distance moved to nest sites from leks visited for adults and 2.3 km for yearlings. to feeding and loafing areas were within 1.5 km of nest decreased to 0.1 km during incubation. average of 0.7 krn from nests laying and 0.1 krn during of nesting parison were related to timing Movements Yearlings was 5.4 krn during egg laying sites moved an to feed,ing and loafing areas incubation. Movements to !J_ming of vegetation of hatchi!)g or nes t loss. and after during completion desiccation in com- �199 LITERATURE CITED Aldrich, J. W., and A. J. Duvall. 1955. Distribution of American gallinaceous game birds. U. S. Fish and Wildl. Servo Circ. 34. 23pp. Batterson, W. 1\11.., and W. B. Morse. 1948. Oregon sage grouse. Oregon Fauna Sera 1. Oregon Game Cornrn , , Portland. 29pp. Beck, T. D. 1., R. B. Gill, and C. E. Braun. 1975. Sex and age determination of sage grouse from wing characteristics. Game Inf. LeaH. 49 (revised). Colorado Div. Wildl. 4pp. Beetle, A. A, 1960. A study of sagebrush. The section of Artemisia. Wyoming Agr. Expo Sta , Bull. 368. Tridentatae 83pp. Braun, C. E., and T. D. 1. Beck. 1976. Effects of sagebrush control on distribution and abundance of sage grouse. Colorado Div. Wildl. Final Rep., Fed. Aid Proj. W-37 -R, Work Plan 3, Job 8a. pp. 21-84. Bray, O. E., and G. W. Corner. 1972. A tail clip for attaching transmitters to birds. J Wi Idl , Manage. 36:640 -642 .• 0 Canfield, R, H, 1941. Application in sampling range vegetation. Dalke, of the line interception J. For. 39:388-394. method P. D., D. B. Pyrah, Do C. Stanton, JoE. Crawford, and E. Schlatterer. 1960. Seasonal movements and "breeding behavior of sage grouse in Idaho. Trans. North Am. Wildl. and Nat. Res. Conf. 25:396-407. _- 1963. ,---.....•..•. , and Ecology, productivity, and management of sage grouse in Idaho. J. wuui. Manage. 27:810-841. Dargan, L. M. 1940. Cons. Comments Wings over North Park, 3(41:1-4. "'- Dill, H. H., and W. H. Thornsberry. net trap for capturing waterfowl. 14: 132 -137. Colorado. Colorado 1950. A cannon-projected J. Wild!. Manage. �200 Einarsen. A. S. field signs. Eng, 1956. Determination of some predator Oregon State Monog. Stud. 2001. 10. R. L. 1955. A method s ex ratios f r orn wings. 1963. grouse. J. by for obtaining sage grouse age and J. Wildl. Manage. 19:267-272. Observations on the breeding WildL Manage. 27:841 -846. , and P. Schladweiler. ----ments and habitat use in species 34pp. biology of male sage 1972. Sage grouse winter movecentral Montana. J. Wildl. Manage. 36:141-146. Gill, R. B. 1965. Distribution and abundance of a population of sage grouse in North Park, Colorado. M. S. Thesis. Colorado State Univ., Fort Collins. 185pp. Girard. Griner, G• 1935. Life history, habits, grouse, Centrocercus urophasianus. Wyoming, Laramie. 153pp. .!....I. and food of the sage M. S. Thesis. Univ. of L. A. 1939. A study of the sage grouse (Centrocercus ur opha s ia.nus ) with special reference to life his tory, habitat requirements, and numbers and distribution. M. S. Thesis. Utah State Agr. Co Il , , Logan. Illpp. Grinnell, J., H. C. Bryant, and T. 1. Storer. 1918. The game birds of California. Un iv, of Cali£. Pres s , Berkeley. 642 pp. Hoyt, D. F. 1979. Practical methods of estimating fresh weight of birds eggs. Auk 96:73-77. Keller, R. J., H. R. Shepherd, and R. N. Randall. 1941. Survey of 1941: North Park, Jackson County, Moffat County, including comparative data of previous seasons. Colorado Game and Fish Comm., Sage Grouse Surv. 3. 31pp. Klebenow, D. A. 1969. Sage grouse nesting Idaho. J. Wildl. Manage. 33:649-662. volume and and brood habitat Lacher, J. R., and D. D. ...Lacher. 1964. A mobile trap. J. Wildl. Manage. 28:595 -597. in cannon net Lumsden, H. G. 1968. The display of the sage grouse. Dep. Land and Forests Res. Rep. 83. 94pp. Ontario �201 May, T. A. 1970. Effects of sagebrush control on distribution and abundance of sage grouse. Colorado Di v, Wildl., Job Compl. Rep., Fed. Aid Proj. W-37-R-23, Work Plan 3, Job 8a. pp.115-138. , and B. ----female sage States Poley. 1969. Spring and sununer grouse in North Park, Colorado. Sage Grouse Workshop. 6:173-178. movements of Proc. West. Nelson, O. C. 1955. A field study of the sage grouse in southeastern Oregon with special reference to reproduction and survival. M, So Thesis. Oregon State Coll., Corvallis. 113pp. Patterson, Inc. R. L, 1952. The sage grouse Denver. 341pp. of Wyoming. Sage Books, Peterson, J. G. 1970. The food habits and summer distribution of juvenile sage grouse in central Montana. J. Wildl. Manage. 34:147-155. Poley, B. 1969. Effects of sagebrush control on distribution and abundance of sage grouse. Colorado Di v, Wildl., Job Compl. Rep., Fed. Aid Proj. W-37 -R-22, Work Plan 3, Job 8a. pp. 61-86. Pyrah, D. B. 1959. Sage grouse population -t r end and trapping study. Wyoming Game and Fish Comm., Job Compl. Rep., Fed. Aid Proj. W-50 -R -8. pp. 38 -64. Dep., Proj. Rahn, 1963. Sage grouse Wild!. Restoration W-125-R-2. investigations. Idaho Fish and Game Div., Job Compl. Repo~ Fed. Aid H., and Ao A'r, 1974. The avian egg: water loss. Condor 76: 147 -152. incubation time and Rasmussen, D. 1., and L. A. Griner. 1938. Life history and mana_gement studies of the sage grouse in Utah, with special reference to nesting and feeding habits. Trans. North Am. Wild!. Coni. 3:852 -&64. Rogers, G. E. 1964. Colorado Game, 132pp. Sag-e grouse investigations in Colorado. Fish arid Parks Depo, Tech. Publ. 16. �202 Rothenmaier, D. 1979. Sage grouse reproductive ecology: breeding season movements, strutting ground attendance and site characteristics, and nesting. M. S. Thesis. Uni v, of Wyoming, Laramie. 97pp. Scott, J. W. 1942. 472-498. Mating behavior of the sage grouse. Smith, E. L. 1966. Soil vegetation relationships of some Artemisia types in North Park, Colorado. Ph. D. Di s s , , Colorado State Urri.v,. , Fort Collins. 203pp. Te rwi Il igez-, Co, Jr., and Eo L, Smith. 1978. types in North Park, Colorado. Colorado Sci. Dep , Sci. Sere 32. 48pp. Auk 59: Range resource State Un iv, Range U. S. Department of Commerce. 1973. Monthly normals of temperature, precipitation, and heating and cooling degree days. 1941-70. Climatography of the United States 81 (Colorado). 12pp. 1978. Climatological data: National Oceanic and Atmospheric annual summary, Admin. 83(13). Colorado. 14pp. 1979. Climatological data: National Oceanic and Atmospheric annual summary, Admin. 84(14). Colorado. 14pp. Walles tad, R. O. 1971. sage grouse broods 35:129 -136. Summer movements in central Montana. and habitat use by J. Wildl. Manage. 1975. Life history and habitat requirements of sage grouse in central Montana. Montana Dep, Fish-and Game, Helena. 65pp. , and D. ----grouse hens Pyrah. 1974. Movement and nesting of sage in central Montana. J. Wild!. Manage. 38 :630 -633. , and P. Schladweiler. 1974. Breeding and 11abitat selection of male sage grouse. 38:634-637. ---_-' Wiley, season movements J. Wildl. Manage. R. H. 1973. Territorialityand non-random rnatfng in sage grouse, Centrocercus urophasianus. Ani m, Behav, Monog. 6:85-169. �203 Prepared I) \"") .." by __ ~$.~~~~~_t(~ __~~-~=:v~·_c~~_,~,v __ ~.~+- _ Brett Petersen (j'-/L) Graduate Research Assistant Approved by _=--,a::::::--'-"=~=L-::' ::-=L~. -L~....J.L.::==':_ Clait E. Braun Wildlife Researcher _ ��205 JOB PROGRESS State of April 1980 REPORT Colorado ------~~~~~----------------- Project No. W-37-R-33 Work Plan No. Job Title Game Bird Survey 3 Vulnerability Job No. and Population Characteristics 11 of Sage Grouse in Moffat County Period Covered: Personnel: 1 April 1979 through 31 March 1980 Kevin Berner, Clait Braun, Jack Corey, John Ellis, Howard Funk, Don Hoffman, Richard Hoffman, Laura Spess-Jackson, Brent Renfrow, Dave Roche, Ed Rodriguez, John Testa, and Duane Wagner, Colorado Division of Wildlife. ABSTRACT Population characteristics, harvest statistics and vulnerability to hunting were investigated for sage grouse (Centrocercus urophasianus) in Moffat and western Routt counties, Colorado. Counts of males on 69 leks averaged 53.4/ lek, substantially higher than the 37.2 males/lek counted in 1978. Twelve hundred and fifty sage grouse were banded in 1979 in 5 zones within Moffat County. This sample was comprised of 903 males (139 young of the year, 311 yearlings, 453 adults) and 347 females (177 young of the year, 93 yearlings, 77 adults). Trapping success was not indicative of actual sex and age ratios in either the spring or late summer population. Free permits were required of all sage grouse hunters in Moffat County in 1979 and 2,673 were issued. An estimated 2,041 permittees hunted and harvested 7,840 sage grouse, including crippling loss. Hunter success was good (81.0%) and each successful hunter bagged an average of 4.4 grouse. Approximately 56% (56.5) of the total harvest occurred during the initial weekend of the 16 (9 days in limited areas) day season. Check stations and volunteer wing collection barrels were used to obtain harvest statistics and grouse wings. A total of 826 hunters was checked at 3 check stations. These hunters harvested 2.3 grouse each; Hunter success'was best at Cold Spring Mountain and similar in the remainder of the area. Analysis of 3,424 wings from sage grouse bagged in Moffat and western Routt counties indicated that the harvest was comprised of 55% young of the year, 27.2% yearlings and 17.8% adults. There were 1.9 chicks per female in the harvest sample and overall production and nesting success was less than in 1978. The sex ratio at hatching approximated 1:1 with differential survival favoring females in older age classes. There were 2 females to each male in the adult and yearling segment of the fall population. Estimated turnover of adult males was 57.0%, while for adult females it was 44.5%. Chicks were slightly more vulnerable to hunting than other age classes but the differences were not significant (R> 0.05). The overall harvest rate in 1979 was 7.0%, somewhat less than the 9.5% documented in 1978. �206 RECONMENDATIONS 1. Counts of males on selected leks or complexes continued as a management function. of leks should be 2. Trapping and banding should be continued in 1980 with a goal of at least 100 males banded in each of 5 zones in Moffat County. 3. The free permit requirement for all sage grouse hunters County should be continued in 1980. 4. Check stations should be operated at 3 locations on the initial 2 weekends of the sage grouse season in Moffat County in 1980 (Cedar Mountain, Maybell, Dinosaur). 5. Collections of sage grouse wings from Moffat County after 1980 should be continued as a management function. 6. Field work should be terminated following data collection through the 1980 hunting season. Data analysis will take at least 1 additional year. in Moffat �207 VULNERABILITY AND POPULATION CHARACTERISTICS OF SAGE GROUSE IN MOFFAT COUNTY Clait E. Braun and Donald M. Hoffman Sage grouse are hunted annually in Colorado, with season lengths typically of 3 days and bag possession limits of 2 and 4 until recent years. Intensive bandings of adult and yearling sage grouse along with experimental hunting seasons in North Park, Jackson County, Colorado have indicated that only about 10% of the fall population of these segments is annually harvested. Data from this area indicate that yearling males are most vulnerable, with adult males and hens of both age classes having lower rates of exploitation. Knowledge concerning population characteristics and harvest rates are important if maximum allowable recreational opportunity through hunting is to be achieved. If turnover rates are moderately high (40-60%) and recovery rates (and by relationship, vulnerability rates) are low, conservative seasons and bag limits have little merit. This report covers the 2nd year of a 3 year study to investigate population characteristics and vulnerability to hunting of sage grouse in Moffat County and adjacent areas in western Routt County, Colorado. The period covered was from mid-March 1979 through mid-March 1980. P. N. OBJECTIVES 1. Estimate young of the year vulnerability adults and yearlings of each sex. 2. Provide reliable estimates 3. Provide reliable data on characteristics 4. Estimate turnover of harvest and survival to hunting rates in relationship (recovery to rates). of the harvest. rates. SEGMENT OBJECTIVES 1. Counts of males and females will be conducted on a mlnlmum of 28 leks within Small Game Management Units 16 and 18 during April and May. A minimum of 4 early morning counts will be made per lek. Additional new leks will be located through ground searches. 2. Adults and yearlings (both males and females) will be captured and banded Coloron or near strutting grounds (leks) during March, April, and May. A coded leg bands will be used to identify birds banded by location. sample of 300 males and 300 females is desired .. 3a. Chicks will be captured and banded within brood rearing areas during July, August and early September through use of cannon nets, drive traps and/or night lighting. A minimum of 300 (150 male and 150 female) chicks i~ desired. Aluminum and year color-coded leg bands will be used to identify birds by location. �208 3b. Adult and yearling sage grouse will be banded in conjunction with the banding of chicks whenever possible. Aluminum and year color coded leg bands will be used to identify birds and location. 4a. Sage grouse hunting will be by free permits, unlimited in number, in Moffat County. All hunters will be mailed a questionnaire within I week of the end of the hunting season. One follow-up letter will be sent 3 weeks later. 4b. Check stations will be operated at least during both days of the opening weekend near Cedar Mountain, Dinosaur and Cold Springs Mountain to collect harvest data, wings, and compliance with the permit requirement. 4c. Wing barrel collection stations will be located at 16 sites to sample the harvest within Moffat and western Routt counties. A sample of 600 wings is desired. 4d. Numbers and locations of banded birds shot will be recorded from field hunter checks, check stations and hunter questionnaires. 4e. Hunter success will be determined check stations. from hunter questionnaires and 4f. Age and sex composition of the harvest sample of wings will be determined in Fort Collins following the open season. 4g. Total harvest will be calculated 5. Vulnerability estimates 6. Data will be compiled, from the hunter questionnaire survey. will be made based upon band recoveries. results analyzed, METHODS and a progress report prepared. AND MATERIALS Counts of males and females on leks were made in early morning from late March to late May following procedures described by Rogers (1964) and Braun and Beck (1976). Intensive efforts to locate new leks were not made in 1979. Sage grouse were trapped in July and August at night where they roosted along trails and throughout the early merning and evening where they concentrated in meadows. Methods used involved spotlighting and capture with long-handled nets, wire drift fences with, a cannon net at the end of the fence, bumper mounted cannon nets, and modified lily pad traps (Braun and Beck 1976, Lacher and Lacher 1964, Guillion 1965). Birds captured were weighed, measured, classified as to age and sex (Eng 1955, Beck et al. 1975) and banded with serially numbered aluminum leg bands (size 16 for males, 14 for females) and plastic bandettes color-coded for year and location (5 different areas in Moffat County). �209 Sage grouse hunters in Moffat County in 1979 were required to have in their possession while hunting a free numbered permit. Permits, unlimited in number, were available from Division of Wildlife offices in Craig, Denver, Fort Collins, Colorado Springs, Grand Junction, Meeker, and Montrose, all license agents in the Moffat County area, and all project and management personnel working in the area. Questionnaires were sent to all permittees immediately following the end of the sage grouse hunting season in Moffat County. One follmv-up letter and questionnaire were sent in mid-October to all non-respondents to the initial mailing. Check stations were operated at 3 locations (Cedar Mountain, Maybell and Dinosaur) on both weekends (except Maybell wh Lch was not open during the 2nd Saturday) of the hunting season. Each station was operated from about 0800 to 1900 MDT, depending upon traffic load. Data obtained per party were: county of origin, number of hunters, hours hunted (total of all hunters in each party), birds observed, birds bagged, birds lost, number of banded birds and location where each was harvested, areas hunted, and information on previous sage grouse hunting experience. One wing was obtained for 1,692 of 1,898 birds checked (89.1%) with ovaries being taken or examined from a sample of yearling and adult hens. Efforts were made to ascertain sex by gonadal inspection for a sample of birds. These data were recorded on tags with wings being individually marked with corresponding information concerning actual sex of that bird. Wings were frozen and stored for later analysis. In addition to the 3 check stations, field checks of hunters were made on the opening weekend and 2nd Saturday at Cold Spring Hountain and wing barrels and signs (Hoffman and Braun 1975) were placed at 16 locations in Moffat and western Routt counties, including the 3 check station sites. Volunteer wing collection stations were in operation the entire season at all sites. Wings were collected following each weekend and midweek and frozen for later analysis. Collected wings were thawed and classified to age (chicks, yearlings, adults) and sex following procedures outlined by Eng (1955) and Beck et al. (1975). Hatching dates were calculated for chicks of the year using data from Pyrah (1963). Ovaries collected were stored in AFA (alcohol, formalin and acetic acid) and later examined for presence of ovulated follicles as described by Kabat et al. (1948). DESCRIPTION OF AREA Moffat County is located in extreme northwestern Colorado, bordered by Wyoming on the north, Utah on the west, Rio Blanco County on the south and Routt County on the east. The major drainages include the Green River which flows north to south through western Moffat County, the Yampa River which flows east to west in the southern part of the county to its intersection with the Green River, approximately 3 miles (4.8 km) from the Utah border, and the Little Snake River which flows northeast to southwest through the middle of the county where it joins the Yampa River in Lily Park. �210 The climate in Moffat County is semi-arid with 8 to 20 inches (20.3-50.8 cm) of precipitation annually. Precipitation occurs mainly in late summer to early winter. Mean annual precipitation for 6 weather stations was 12.99 inches (33.0 cm). The annual mean temperature for 5 stations was 43.3 F (6.3 C) during years of record. Elevation in Moffat County varies from approximately 4,600 to 11,045 feet (1,402-3,367 m). The county seat, Craig, located in eastern Moffat County, is 6,240 feet (1,902 m) above sea level. Sagebrush ranges comprise approximately 60% (2,841 mi2) of the total land area of Moffat County. Non-sagebrush ranges consisting of pinon-juniper, rough and rocky mountain shrub~ aspen, or spruce-fir types comprise most of the remaining 40% (1,902 mi ) of the total land area. RESULTS AND DISCUSSION Counts of Sage Grouse on Display Areas From late March to late May 1979, 205 counts were made of 69 active sage grouse leks in Moffat County and western Routt County (Table 1). The average number of males/lek was 53.4, substantially higher than the 37.2 males/lek recorded for 54 leks in 1978 (Table 2). The 53.4 males/lek was the highest number of cocks recorded in Moffat County since 1970 (Table 3). Numbers of males on leks has increased (I < 0.05) since 1977 when adequate numbers of leks were annually counted. Prior to 1977, numbers of leks counted varied from 17 (1975) to 29 (1969). Unless leks were randomly counted from 1969 through 1976, data on average numbers of males/lek have little value for documentation of long term trends. This is because it is not known whether only larger or accessible leks were counted. Comparable data (Table 2) for all leks and years for which count information is available indicate the lack of continuity for many leks even during the period of relatively intensive study. Data for all 4 years are only available for 20 of approximately 70 leks. Unfortunately, even these 20 leks were not counted systematically following recommended procedures (Braun and Beck 1976). Although intensive efforts to locate new leks were not made in 1979, 7 new leks were located. It is hypothesized that the number of known leks could be at least doubled and possibly tripled if intensive ground and aerial lek searches were conducted over a 2-year period. Due to changes in personnel, the original data forms for 1979 could not be inspected to ascertain peak of female attendance of leks. However, field observations indicated peak female lek attendance was the 1st week of April in northcentral Moffat County and about 1 week later in southern Moffat County. Peak hen lek attendance was probably during the 2nd or more likely the 3rd week of April at Blue Mountain and Cold Spring Mountain. Brood Counts Actual counts of sage grouse broods along defined routes were not conducted in 1979. However, all distinct broods seen in July and August were recorded. �211 Table 1. Peak lek counts of sage grouse, Moffat Lek Blue Mountain Bear Creek Escalante Haslim Cow Camp Karren Ranch Sixteen Road (new) State Line Cold Spring Beaver Basin Gee Flats Goodman Draw Summit Spring East of Baggs Highway Cowboy Reservoir Eighty Road Elk Mountain (new) Elkhead Creek Fan Rock Fly Creek Four Mile Creek 1 Four Mile Creek 2 Four Mile Creek 3 Sage Creek (new) Slater Park Twenty-nine Road 1 Twenty-nine Road 2 Northcentral-East Cottonwood Gulch Mud Spring Draw Pole Gulch Timberlake 1 Timberlake 2 Timberlake 3 Timberlake 4 (new) West Timberlake 2 Northcentral-North Big Hole Butte Big Hole Gulch Cox Ranch Dressler Gulch 1 Dressler Gulch 2 (new) Scandinavian Gulch Seven Mile Reservoir Thornburg Gulch Twenty-one Road (new) West Timberlake 1 County, No. of counts Males 3 3 64 112 1979. Date(s) 14 17 14 14 92 May May May May 21 May 22 May 1 6 6 April 4 18 2 2 1 37 14 7 April 27 April 23 May 24 Hay 23 April 16 May 9 April 6 April 21 April 1 May 2 97 3 3 3 80 77 o o o 3 1 5 5 12 152 41 4 46 85 128 2 41 o 4 4 o 153 1 17 3 4 93 99 55 5 4 4 4 104 5 111 2 61 42 23 33 18 68 4 1 2 1 3 108 43 2 5 4 40 2 1 26 1 All dates 19 May 21 May 10 Hay 1 May 16 April 19 April 6 May 10 May 26 April 20 April 11 May 20 April 25 April 25 April 17 April 20 April 1 May 30 March 20 April �212 Table 1. Peak 1ek counts of sage grouse, Moffat County, 1979. Lek Northcentral-South Big Gulch 3 Bord Gulch Grassie Reservoir Greasewood Gulch Lay Creek North Fork Big Gulch 2 Sand Creek (new) Spring Creek 1 Spring Creek 2 Spring Creek 3 Upper Nineteen Road South of U.S. 40 Axial Basin Boxelder Gulch 1 Boxe1der Gulch 2 Deception Creek Dry Lake 2 Duffy Mountain Horse Gulch Juniper 1 Juniper 2 Morgan Gulch 1 Morgan Gulch 2 Morgan Gulch 3 Round Bottom Temple Gulch Yellow Jacket Sunbeam West-North Coffee Pot Spring Cross Mtn. 1 Cross Mtn. 2 Cross Mtn. 3 East Sand Wash Lone Tree Powder Wash Hill Snake River West Thornburg Well No. of counts Males (Continued) Date(s) 1 11 4 50 25 18 18 1 2 11 24 24 1 18 19 4 3 4 4 5 72 71 62 43 117 19 23 20 2 2 April April April May May 5 1 4 45 2 43 78 43 57 38 28 23 19 23 1 24 27 7 April April April April May April April May 24 27 17 30 29 68 76 23 35 26 9 27 24 17 17 26 17 9 April April March March April April April April April 1 4 4 1 1 3 1 4 2 8 42 34 27 125 76 45 46 20 April April April May May May April April May April April o o 4 4 1 2 2 I 5 6 2 4 4 3 4 4 8 --------------------------------------------------------------------------- �213 Table 1. Peak lek counts of sage grouse, Moffat County, 1979. SUMMARY No. of active leks counted Avg. number of males Blue Mountain 6 87.0 Cold Spring 1 6.0 11 66.1 Northcentral-East 8 78.8 Northcentral-North 10 31.7 Northcentral-South 11 44.0 South of U.S. 40 13 52.2 9 36.6 69 53.4 Area East of Baggs Highway Sunbeam-West-North All areas (Continued) �214 Table 2. Peak lek counts of sage grouse, Moffat County, 1976-79. Lek Blue Mountain Bear Creek Escalante Haslim Cow Camp Karren Ranch Sixteen Road State Line 1976 Northcentral-East Cottonwood Gulch Mud Spring Draw Pole Gulch Timberlake 1 Timberlake 2 Timberlake 3 Timberlake 4 West Timberlake 2 Northcentral-North Big Hole Butte Big Hole Gulch Conway Spring Cox Ranch Dressler Gulch 1 Dressler Gulch 2 Scandinavian Gulch Seven Mile Reservoir Thornburg Gulch Twenty-one Road West Timberlake 1 1978 1979 64 62 26 l12 97 40 65 80 77 92 Cold Spring Beaver Basin Gee Flats Goodman Draw Summit Spring East of Baggs Highway Cowboy Reservoir Eighty Road Elk Mountain Elk Head Creek Fan Rock Fly Creek Four Mile Creek 1 Four Mile Creek 2 Four Mile Creek 3 Sage Creek Slater Park Twenty-nine Road 1 Twenty-nine Road 2 1977 101 64 13 19 92 153 55 6 42 18 11 20 18 37 14 113 12 116 12 8 34 36 27 40 18 136 152 41 46 85 128 41 36 4 o 62 153 8 38 17 37 76 87 29 55 60 43 53 66 21 20 93 99 55 104 108 5 47 111 17 19 ·34 34 37 37 42 20 36 3 16 27 20 23 25 33 18 45 42 68 14 7 5 17 40 43 61 26 3 1 �215 Table 2. Peak lek counts Lek Northcentral-South Big Gulch 1 Big Gulch 3 Bord Gulch Grassie Reservoir Greasewood Gulch Lay Creek North Fork Big Gulch 2 Sand Creek Spring Creek 1 Spring Creek 2 Spring Creek 3 Upper Nineteen Road South of U.S. 40 Axial Basin Boxelder Gulch 1 Boxelder Gulch 2 Deception Creek Dry Lake 2 Duffy Mountain Horse Gulch Juniper 1 Juniper 2 Mor gan Gulc h 1 Morgan Gulch 2 Morgan Gulch 3 Round Bottom Temple Gulch Yellow Jacket Sunbeam \.Jest-North Coffee Pot Spring Cross Mtn. 1 Cross Mtn. 2 Cross Mtn. 3 East Sand Wash Lone Tree Powder Wash Hill Snake River West Thornburg Well of sage grouse, 1976 Moffat County, 1976-79. (Cont'd.) 1977 1978 1979 23 6 52 37 42 34 20 27 34 22 57 37 125 24 42 46 32 24 31 16 33 12 30 11 50 28 58 34 44 72 3 8 32 21 30 32 31 25 60 22 44 14 57 76 45 20 71 62 43 117 22 35 17 27 4 14 31 45 45 8 8 15 4 39 43 24 55 36 50 32 43 78 43 57 18 54 24 18 26 56 35 2 38 24 26 29 25 31 27 13 5 17 16 28 30 29 22 2/+ 55 ·12 69 68 76 8 32 23 35 36 �216 Table 2. Peak lek counts of sage grouse, Moffat County, 1976-79. (Cont'd.) SUMMARya Area 1976 1977 26.0(1) 64.8(4) --(0) 87.0(6) 49.3(4) 67.0(4) 6.0(1) 113.0(1) 35.9(10) 50.3(4) 66.1(11) Northcentral-East 44.5(2) 28.2(6) 55.3(7) 78.8(8) Northcentral-North 23.0(4) 24.7(4) 23.5(8) 31.7(10) Northcentral-South 25.8(8) 26.8(9) 31.2(9) 44.0(11) South of U.S. 40 32.7(6) 34.0(11) 32.6(14) 52.2(13) Sunbeam-West-North 22.0(2) 25.6(8) 28.3(8) 36.6(9) All areas 31.9(24) 33.5(59) 37.2(54) 53.4(69) Blue Mountain Cold Spring East of Baggs Highway 1978 a Numbers in parentheses are number of active leks counted. 1979 �217 Table 3. Trends in peak lek counts of male sage grouse, Moffat County, Colorado, 1969-79. N Average number per lek 1969 29 74.6 1970 28 54.4 1971 25 52.0 1972 27 38.4 1973 22 45.8 1974 22 41.4 1975 17 31.7 1976 27 31.9 1977 59 33.5 1978 54 37.2 1979 69 53.4 a Counts by research personnel started in 1976 and continued through 1979. �218 During July, 195 hens were seen with 932 chicks for an averaged brood size of 4.8. Thirty-six distinct broods with 243 chicks (avg. = 6.8) were seen in August. The increase in average number of chicks per brood in August was a function of brood shuffling and formation of gang broods. This left some successful hens without any chicks and other hens with more chicks than the average clutch size. During August, 9 distinct broods were seen containing from 9 to 16 chicks. The average clutch size for sage grouse is 7-8 eggs. Capture and Banding Intensive efforts to capture and band sage grouse in 1979 were conducted from 27 March through 30 May and from 18 July through 30 August. In all, 1250 sage grouse were newly banded of which 823 were in spring and 427 were in summer (Table 4). Five zones were initially established in Moffat County for data anlaysis. The Sunbeam-Cold Spring Mountain zone was later subdivided to more fully clarify some differences. While banding goals were similar in each zone, banding effort and results were not uniform, especially in summer. This was because few concentrations of broods could be located outside of Cold Spring Mountain and Blue Mountain. Trapping of males in spring was fairly uniform as at least 100 males were banded in each of the initial 5 zones. However, within each zone, trapping was not uniform as success was much greater on some leks than on others (Table 5). Only I bird banded prior to 1979 was recaptured. Bird 4899 was originally banded I August 1978 as a chick female near Mud Spring Draw. It was recaptured on 30 April 1979 as a subsdult female between Fourmile Creek I and 2 leks, northeast of the original banding site. During summer, 6 additional birds banded in 1978 were recaptured (3 banded as chick females, 2 as chick males, I as a yearling male). All were correctly classified to sex when initially banded. Also, all 6 recaptures were on Cold Spring Mountain and all had been banded there in 1978. Hunting Season Data Collection The sage grouse season in Moffat County (Units 14, 16, 18, 20, 26) in 1979 opened one-half hour before sunrise on 8 September and closed at sunset on 16 September (Units 14, 26) or 23 September (Units 16, 18, 20). Thus, season length in most of Moffat County was 16 days. Bag and possession limits were 3 and 6, in the aggregate with'sharp-tailed grouse. The only change in hunting regulations in this area from 1978 to 1979 was the addition of 7 days in Units 16, 18, and 20. All sage grouse hunters in Moffat County were again required to obtain and have in possession a free permit. Purpose of the permit was to obtain names and addresses to which questionnaires could be sent. Questionnair~s were used to derive total hunter numbers, total harvest, increase band reporting rates and collection of other hunting statistics. Three check stations were operated in 1979 at the same locations and during the same time periods as in 1978. These stations were at Cedar Mountain (at the junction of M.C. roads 3 and 77), Dinosaur (junction of Harpers Ferry road and U.S. Highway 40) and Maybell (junction of Colorado Highway 318 �219 Table 4. Moffat County sage grouse bandings, 1979. Males 1- Females If- 2+ 12 31 1- If- 2+ 4 1 48 Totals Cold SEring Mountain Spring Sunnner 123 16 13 153 27 38 370 Subtotals 123 28 44 153 31 39 418 34 72 14 8 128 34 72 14 8 128 53 75 18 15 161 53 75 18 15 161 114 118 19 5 256 South of U.S. 40 Spring Sunnner Subtotals East of Baggs Highway Spring Sunnner Subtotals North Central Spring Sunnner 7 0 2 6 0 0 15 Subtotals 7 114 120 6 19 5 271 42 101 0 3 146 Blue Mountain Spring Sunnner 9 5 2 18 3 5 42 Subtotals 9 47 103 18 3 8 188 35 39 0 8 2 84 35 39 0 8 2 84 311 453 177 93 77 1,250 Sunbeam West and North Spring Sunnner Subtot.als Totals 139 �220 Table 5. Sage grouse banded by area and lek, Moffat County, Spring 1979. Males Area and lek Females 1- 2+ 1- 2+ Fan Rock 8 34 1 3 46 Four Mile Creek 1 9 13 3 I 26 Four Mile Creek 2 27 6 5 5 43 Four Mile Creek 3 9 22 9 6 46 53 75 18 15 161 Axial Basin 9 8 5 1 23 Boxelder Gulch 1 0 1 0 0 1 Boxelder Gulch 2 3 11 3 3 20 Dry Lake 2 9 25 0 2 36 Juniper 1 1 14 1 2 18 Morgan Gulch 2 2 5 0 0 7 Morgan Gulch 3 7 8 5 0 20 Round Bottom 3 0 0 0 3 34 72 14 8 128 3 14 0 1 18 Haslim Cow Camp 17 35 0 1 53 Karren Ranch 17 26 0 0 43 M.C. 16, Marker 25.2 4 7 0 0 11 State Line 1 19 0 1 21 42 101 0 3 146 Gee Flats 8 19 3 1 31 Goodman Draw 4 12 1 0 17 12 31 4 1 48 Totals % of Totals East of Baggs Highwa:l Subtotals 19.6 South of U.S. 40 Subtotals 15.6 Blue Mountain Escalante Subtotals 17.7 Cold SEring Mountain Subtotals 5.8 �221 Table 5. Sage grouse banded by area and lek, Moffat County, Spring, 1979. (Continued). Males 2+ 1- Females 12+ Cross Mtn. 1 2 5 0 0 7 Cross Mtn. 2 0 1 0 0 1 East Sand Wash 4 10 3 1 18 20 12 4 0 36 Snake River West 0 3 0 1 4 Thornburg Well 9 8 1 0 18 35 39 8 2 84 2 8 0 0 10 27 24 9 1 61 Timberlake 1 0 2 0 0 2 Timberlake 2 43 25 6 3 77 Timberlake 3 9 10 2 0 21 Timberlake 4 4 1 0 0 5 Big Hole Butte 2 4 0 0 6 Cox Ranch 4 0 0 0 4 Dressler Gulch 0 6 0 0 6 Scandinavian Gulch 3 6 1 0 10 Thornburgh Gulch 7 4 1 1 13 Bord Gulch 0 10 0 0 10 Grassie Reservoir 0 4 0 4 Lay Creek 9 5 0 0 '0 14 Spring Creek 3 3 3 0 0 6 Upper 19 Road 1 6 0 0 7 114 118 19 5 256 31.1 290 436 63 34 823 100.0 Totals % of Totals Sunbeam West and North Lone Tree Subtotals 10.2 North Central Mud Spring Draw Pole Gulch Subtotals Totals �222 and U.S. Highway 40). Check stations were operared from about 0800 to 1900 hours on 8, 9, 15 and 16 September at Cedar Mountain and Dinosaur and about the same hours on 8, 9 and 16 September at Maybell. Each station was staffed with 2 research personnel. Field checks by research personnel were conducted on Cold Spring Mountain on 9 and 15 September. Data obtained per party were: county of origin, number of hunters, hours hunted (total of all hunters in each party), birds observed, birds bagged, birds lost, number of banded birds and location where each was harvested, and area hunted within Moffat County. Most importantly, 1 wing was obtained from 1,692 of 1,898 birds checked (89.1%). Ovaries were collected from 71 adult and yearling females for laboratory analysis. Ovaries from an additional 56 adult and 106 yearling females were classified at Cedar Mountain as to ovulation history. Whole body weights were obtained from 247 uneviscerated sage grouse. Volunteer wing collection barrels and signs were placed at Axial, Deception Creek, Juniper Springs, M.C. Road 2, M.C. Road 101, California Park Road, Elkhead Road, Dinosaur (when the creek station was closed), Wolf Creek Road, Maybell (when the check station was closed), Lay Creek, Maybell City Park, M. C. Road 3, M. C. Road 4, Big Gulch and Cedar Mountain (when the check station was closed). Check Stations During the 4 days of check station operation (3 days at Maybell), 826 hunters with 1,898 sage grouse (2.3 birds per hunter) were checked. These hunters reported observing 14,639 sage grouse. Hunter efficiency was low, 13.0% (1,898 birds harvested + 14,639 birds observed) and 145 birds (7.1% of those retrieved + those reported lost) were reported crippled and lost (Table 6). Data in Table 6 suggest that hunter success and total harvest were higher in 1979 than in 1978. Table 6. Sage grouse harvest statistics, Moffat County, Colorado, 1978-79a. Check station Number hunters checked Number birds observed Hunter efficiency % Crippling loss % Birds per hunter Cedar Mountain 534 7,175 1,099 15.3 6.6 2.1 Dinosaur 162 3,237 3S7 11.0 9.6 2.2 Maybell 130 4,227 442 10.5 6.2 3.4 All Stations 826 14,639 1,898 13.0 7.1 2.3 1978 770 15,295 1,490 9.7 7.6 1.9 1979 826 14,639 1,898 13.0 7. 1 2.3 Number birds harvested a Only those statistics collected at check stations. �223 It is readily apparent that differences occurred in hunter pressure, birds seen, hunter efficiency and birds per hunter in the 3 areas sampled by check stations. Hunter efficiency was lowest and birds per hunter was highest in the Cold Spring Mountain area (Maybell Check Station). It is also apparent that where hunter pressure was highest, birds per hunter was lowest, suggesting that sage grouse in these areas were more difficult to approach and harvest. Small game management units in Moffat County were subdivided into harvest zones to examine hunter pressure and harvest by area (Fig. 1). Data available from check stations, while biased because of their location, illustrate that hunter pressure and harvest were not uniformly distributed in Moffat County in 1979 (Table 7). These data indicate that hunter pressure was low in zones 14A, 16C, 20B, and 26A. This is probably true for zones 16C, 20B and 26A but not for 14A. Harvest generally paralleled hunter pressure in all zones but 18A (Cold Spring Mountain). Birds per hunter was 4.3 in this zone suggesting that hunters had ample harvest opportunity. These data are similar to those from 1978 (Table 8). Table 7. Hunter pressure, harvest and hunter success by harvest zone, Moffat County, 1979a• Statistic 14A 16A 16B Harvest zones 18B 16C 18A Total hunters (%) 0.6 19.4 46.6 2.3 9.1 2.1 19.8 0.1 0.0 Total harvest (%) 0.4 21.1 38.2 1.8 17.0 2.5 18.9 0.1 0.0 Birds hunter 1.4 2.5 1.9 1.8 4.3 2.8 2.2 2.0 0.0 20A 20B 26A aCheck station data only. Table 8. Hunter pressure, harvest and hunter success by harvest zone, Moffat County, 1978-79a• Harvest zone Total hunters (%) 1978 1979 Total harvest (%) 1978 1979 Birds per hunter 1979 1978 14A 0.0 0.6 0.0 0.4 0.0 1.4 16A 22.7 19.4 16.4 21.1 1.4 2.5 16B 37.5 46.6 28.5 38.2 1.5 1.9 16C 2.5 2.3 3.0 1.8 2.3 1.8 18A 9.0 9.1 22.0 17.0 4.8 4.3 18B 2.3 2.1 0.9 2.5 0.8 2.8 20A 26.0 19.8 29.2 18.9 2.2 2.2 20B 0.0 0.1 0.0 0.1 0.0 2.0 26A 0.0 0.0 0.0 0.0 0.0 0.0 aCheck station data only. �:. T . . . . . . . . . .!: .;••~5·"""""p'~!'!!.",;1-"-·._ TI li~~::Ii" ~. ~~~M j 'n ~, "" .~ " r "I •• .•.. I, ~ ".,' 7 'hJ; <to.> ••--;, ,_ •• 'l i,;,~';t,!, «, , _ \ M" •.•. 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I , .'~k.i. ~ :~Iiii • •.•• : l :~;" : U ""m )t"; :' " '... 7J:!..~ :!!'r •• " ~,\, I;" ~I" i h1 .\ .•-r:» ~ "~i" ~ "I" .",L::___.f.'~\~r,.1'1"'JI", i'" •• " , " o 0 . •••• 1 Fig. 1. Harvest zones, Moffat County, Colorado. • , • ~~,'" '+:" • • • .: i J:, ~L'.'.LC:.:. {t.i '2UU.~;!O:;;_~':J.. ~h, ~. 'L~ ,~,.1'.' ,,, • '1' . , . .... :l ;; ZONES �225 Sage Grouse Hunting Experience Most hunters contacted at check stations were asked whether or not they normally hunted sage grouse in Moffat County. Of the 760 hunters responding, 538 (70.8%) reported they normally hunted sage grouse in Moffat County, 52 (6.8%) normally hunted sage grouse elsewhere while 170 (22.4%) were 1st time sage grouse hunters (Table 9). These data are similar to those collected in 1978 in Moffat County and in 1974-79 in North Park. Table 9. Previous sage grouse hunting experience Moffat County, Colorado 1978-79. of sage grouse hunters, Year Normally hunt in Moffat County No. % Normally hunt elsewhere No. % 1st time sage grouse hunter No. % 1978 502 69.1 81 11.1 144 19.8 1979 538 70.8 52 6.8 170 22.4 Hunter Origin Origin of hunters contacted at check stations was ascertained from vehicle license plates. The origin of hunters was similar to that documented in 1978 (Table 10). Most originated from the Western Slope with Moffat (31.0%), Mesa (12.6%), Rio Blanco (9.2%) and Routt (4.8%) counties being most important. About one-third of the hunters originated from the Eastern Slope with the Denver Metro Area being most important (21.2%). These data and those presented in Table 9 suggest that despite higher fuel prices, sage grouse hunters who normally hunt in Moffat County did not markedly change where they hunted in 1979. Table 10. Hunter origin, Moffat County sage grouse hunting parties, 1979 1978 County No. Moffat Mesa Rio Blanco Denver Metro Area Jefferson Adams Arapahoe Denver Douglas Boulder Garfield Routt All Others Totals 129 46 41 (74) 32 18 II 11 2 12 11 6 28 347 1978-79. % 37.2 l3.7 1l.8 (21.3) 9.2 5.2 3.2 3.2 0.6 3.4 3.2 1.7 8.1 100.0 No. III 45 33 (76) 32 14 11 19 0 13 8 17 55 358 % 31.0 12.6 9.2 (21.2) 8.9 3.9 3.1 5.3 0.0 3.6 2.2 4.8 15.4 100.0 �226 Analysis of Wings Origin--A total of 3,424 sage grouse wings was received (Table 11). These data indicate that few hunters (2.9% of all wings received) hunted sage grouse south of U.S. 40 in Moffat County with most hunters hunting sage grouse in the northcentral area of the county (53.8% of all wings received). With few hunters apparently hunting south of U.S. 40 (parts of Units 16, 20, and 26) and east of Colorado 13 (part of Unit 14), it does not appear necessary to have differing seasons in different small game management units in Moffat County. It is exceedingly doubtful that sage grouse east of Colorado 13 or south of U.S. 40 comprise different subpopulations. Consequently, the sage grouse resource in Moffat County should be managed as 1 population. Table 11. Origin of sage grouse wings, Moffat County, Colorado, 1979. South of U.S. 40 No. Dinosaur (Blue Mountain) No. Axial Deception Creek Juniper Springs Total % of Total = 2.9 a East of Colorado 13 25 47 27 99 Check Station Dinosaur Wing Barrel Wolf Creek Road Total % of Total = 13.2 323 115 15 453 No. Maybell (Cold Spring Mountain) No. Check Station Maybell Wing Barrell Cold Spring Field Checks Total % of Total = 19.2 378 168 112 658 M.C. Road 2 58 M.C. Road 101 198 California Park Road 29 Elkhead Road 1 Slater Grouse Camps 67 Middle Park Check Station 8 Other (Misc.) 13 Total 374 % of Total = 10.9 Northcentral Location Lay Creek Maybell City Park M.C. Road 3 M.C. Road 4 Big Gulch (North of U.S. 40 and West of Colorado 13) No. Location Cedar Mountain Check Station Cedar Mountain Wing Barrel Wing Envelopes Middle Park Check Station North Park Check Station Field Checks 57 130 143 117 11 Total 1,840 % of Total 53.8 GRAND TOTAL = 3,424 aIncludes extreme western Routt County. No. 989 297 4 47 28 17 �227 Origin of wings in 1978 and 1979 was similar (Table 12) with only the substantial decrease from Dinosaur (Blue Mountain) being of significance. These and check station data strongly indicate that harvest of sage grouse is minimal south of U.S. Highway 40. Table 12. Origin of sage grouse wings, Moffat County, 1978 Area % No. 266 11.0 374 10.9 Northcentral 1,169 48.3 1,840 53.8 Cold Spring 395 16.3 658 19.2 Dinosaur 531 22.0 453 13.2 57 2.4 99 2.9 100.0 3,424 100.0 Moffat Moffat County East County South 2,4.18 TOTALS a Includes extreme western 1978-79. 1979 % No. a Colorado, Routt County. Time of Harvest--All wings received were identified as to harvest period (Table 13). These data, when compared to those from 1978, indicate that the longer season in 1979 was accepted and used by sage grouse hunters. Quite obviously, either more hunters were afield or hunter success was better in 1979 than in 1978 as more wings were received during each time period that the season was open in both years. The data also suggest that the longer season in 1979 may have been responsible for the increased total harvest over that expected in a 9 day season by about 10-11%. Table 13. Time distribution Colorado, 1978-79. of sage grouse wings received, 1978 Time Period No. 1979 % No. % 1,745 75.0 2,291 66.9 First week 224 9.6 313 9.1 Second weekend 357 15.4 462 l3.5 First weekend Second week Season closed 75 2.2 Third weekend Season closed 283 8.3 3,424 100.0 Totals 2,326 100.0 Moffat County, �228 Age and Sex Composition--All wings received were useable for separation of age classes. The data are presented by area within Moffat County (Table 14). Percent young of the year in the harvest was similar in all areas except Blue Mountain (Dinosaur) in 1979. This suggests that nesting success and brood size were lower in the Blue Mountain area than in other areas but were similar in all other areas. All areas had good numbers of yearlings in the harvest reflecting good nesting success and production in 1978 and excellent survival overwinter. Table 14. Age composition of the sage grouse harvest, Moffat County, Colorado, 1979.a Young No. % Yearling No. % Moffat County Easta 215 57.5 104 27.8 55 14.7 374 10.9 1,051 57.2 516 28.0 273 14.8 1,840 53.8 Cold Spring 356 54.1 143 21.7 159 24.2 658 19.2 Dinosaur 204 45.0 143 31.6 106 23.4 453 13.2 56 56.5 25 25.3 18 18.2 99 2.9 Northcentral Moffat County South Totals Percent 1,882 611 931 55.0 Adults No. % Percent of total sample Area 27.2 Totals 3,424 17.8 100.0 alncluding western Routt County (104 wings) Age and sex composition data of the sage grouse harvest in Moffat County for 1976-79 are presented in Tables 15 and 16. These data suggest: 1. The sex ratio at hatching approximates 1:1 with some differential survival favoring females occurring before chicks are 3-4 months of age. 2. Differential survival favoring females occurs in the older age classes and is most pronounced in adults. 3. There are 2 hens to every male in the spring breeding population (4-year average = 65.8% females: 34.2% males of all adults and yearlings). 4. Production of young was excellent in 1976 and 1978 and good in 1979 and probably 1975 (based on percent yearlings in the 1976 harvest). 5. Recruitment of yearlings was poor in 1978 following the average production experienced in 1977. 6. Recruitment (overwinter survival) has been more than sufficient to maintain population stability. Substantial increases in average number of males on leks should have occurred in 1977 and 1979 with some increase in 1976. No increase would have been expected in 1978. �Table 15. Age and sex composition of the sage grouse harvest, Moffat and Western Routt Counties, Colorado, Irnrnatures Females Males Totals No.--- Yearlings Females Males % ~---%- 1976-79. Adults Females Males Totals Totals x Total no. of wings Year No. 1976 203 42.6 273 57.4 476 63.6 41 35.7 74 64.3 115 15.4 25 15.9 132 84.1 157 21.0 748 1977 121 39.3 187 60.7 308 43.0 77 45.8 91 54.2 168 23.4 89 36.9 152 63.1 241 33.fi 717 1978 825 49.6 837 50.4 1,662 68.7 77 32.1 163 67.9 240 9.9 141 27.3 375 72.7 516 1979 871 46.3 1,011 53.7 1,882 55.0 386 41.5 545 58.5 931 27.2 183 30.0 428 70.0 611 4-yr. avg. % 46.7 No. % 53.3 59.2 No. % 40.0 No. 60.0 % 19.9 No. % 28.7 No. % 71.3 No. I 21.4 2,418 17.8 3,424 20.8 ===========================================================================================================~ ~ �Table 16. Age composition (%) of the sage grouse harvest, Moffat Countya, Colorado, 1976-79. 1976 Young 1977 1978 1979 1976 Yearlings 1977 1978 1979 1976 Adults 1977 1978 1979 Moffat County Easta 46.3 39.6 77 .8 57.5 33.3 19.8 8.7 27.8 20.4 40.7 13.5 14.7 Northcentral 64.1 34.0 71.8 57.2 14.5 29.0 5.0 28.0 21.4 37.0 23.2 14.8 Cold Spring 73.1 59.8 62.3 54.1 7.7 17.6 16.7 21.7 19.2 22.5 21.0 24.2 Dinosaur 61.6 60.4 62.1 45,0 15.2 16.0 17.0 31.6 23.2 23.6 20.9 23.4 Moffat County South 71.6 25.8 70.1 56.5 13.4 29.0 5.3 25.3 14.9 45.2 24.5 18.2 Area N w 0 Four-year averages 63.8 alncluding western Routt County. 42.6 68.7 55.0 15.4 23.9 9.9 27.2 20.8 33.5 21.4 17.8 �231 Turnover--Estimated turnover (from wing analysis) of adult male and female sage grouse in Moffat County was 67.8 and 56.0%, respectively, in 1979 (Tables 17 and 18). These estimates are higher than all previous years and are undoubtedly higher than actual. This is the result of the excellent production in 1978 and survival of those young to 1979. It is probable that turnover of adult males and females in 1979 approximated or was lower than the 4-year averages of 57.0 and 44.5% respectively. The estimated annual survival rate (4-year averages) of 43.0 and 55.5% respectively for adult males and females are realistic (Tables 17 and 18). It is of interest that yearling males outnumbered adult males only in 1976 and 1979 following the good (calculated) and excellent production years of 1975 and 1978, respectively. Yearling females outnumbered adult females only in 1979. These data are indicative of a markedly increasing population in 1979. Nesting Success and Production---Molt of adult and yearling sage grouse was examined to estimate nesting success (Table 19). Differences in estimated nesting success closely paralleled differences in percentages of young in the harvest (Table 7). Considering all areas within Moffat County, less than one-half (46.1%) of the yearlings and slightly more than one-half (58.4%) of the adult females were successful nesters in 1979. About one-half (51.5%) of all hens were successful. These data are lower than the estimated overall nesting success of 60.8% in 1978 but are higher than the 30% nesting success estimated in 1977. Estimated nesting success was similar in 1976 and 1979. With the exception of 1977, estimated nesting success of adult hens has exceeded 50/~ each year. In contrast, estimated nesting success of yearlings has varied markedly, being consistent only in 1978 and 1979 (Table 20). Estimated nesting success was similar in the Northcentral Cold Spring and Moffat County South areas in 1979 and was lowest on Blue Mountain (Dinosaur) and east of the Baggs Highway (Table 19). Chicks per hen was lowest at Blue Mountain and highest in the Northcentral area. There was about I chick less per hen in 1979 than in 1978. These data suggest that there will be fewer yearlings in the 1980 spring population than in 1979. Hatching Dates--Ages and hatching dates were calculated for 1,882 chick sage grouse harvested in Moffat County in 1979 (Table 21). Hatching started the week of 11-17 May and continued until the week of 20'-26 July ,\lithan obvious peak between 8 and 28 June. Thus, the hatching peak in 1979 was about 10-14 days later than in 1978. Hatching was earliest in the east and south and latest at Blue Mountain and Cold Spring Mountain. These findings are similar to those from 1978. Those chicks hatching after 5 July were undoubtedly the progeny of hens'that had lost their initial clutch and had renested. However, the data indicate that renesting in 1979 contributed little to overall production as only 6.5% of the chicks was certainly the progeny of renesting hens. These data and those from previous years indicate that nesting is delayed in late springs (delayed snow melt or frequent storms in late March and early April). When nesting is delayed, renesting is relatively unimportant to overall production. Hunter Questionnair~ A total of 2,673 permits was issued for hunting sage grouse in Moffat in 1979. up from 2,006 issued in 1978. Based on 826 hunters contacted County �232 Table 17, Estimated turnover of adult male sage grouse, Hoffat County, Colorado, 1976-79. Year Percent in harvest Yearlings Adults N 1976 37.9 62.1 66 1977 53.6 46.4 166 1978 64.7 35.3 218 1979 32.2 67.8 569 4-Year Average 43.0 57.0 Estir:latedannual turnover for adult males in a stable population 57.0%. Table 18. Estimated turnover of adult female sage grouse, Moffat County, Colorado, 1976-79. Percent in harvest Yearlings Year Adults 1976 64.1 35.9 206 1977 62.6 37.4 243 1978 69.7 30.3 538 1979 44.0 56.0 973 4-Year Aver3ge 55.5 44.5 Estir::atedannual turnover for 3dult females in a stable population N 44.5% �Table 19. Estimated sage grouse nesting success and production, Moffat County, Colorado 1979. Area Moffat County East Yearlings No. successful % Adults No. successful % 4.1 250/ 54.7 2.3 4.3 60.6 116/209 55.5 1.7 3.1 38/71 53.5 65/144 45.1 1.4 3.1 10/15 66.7 18/34 52.9 1.6 3.1 51.5 1.9 3.8 22/48 45.8 Northcentral 136/272 50.0 114/185 61.6 Cold Spring 50/100 50.0 66/109 Dinosaur 27/ 37.0 42.1 Totals Averages 8/19 251/ 250/428 545 46.1 Chicks/ successful hen 1.7 37.0 Moffat County South % Chicks/hen 40.3 30/81 73 All hens No. successful 52/129 457 501/973 58.4 N w W �Table 20. Estimated Area sage grouse nesting success and production, 1976 Moffat County, Colorado, 1976-79. Yearlings 1977 1978 1979 1976 Adults 1977 1978 1979 1976 A1l hens 1977 1978 1979 1976 Chicks/hen 1978 1977 1979 ----Moffat County East 28.6 28.6 76.5 37.0 50.0 58.3 70.6 45.8 40.0 47.4 73.2 40.3 1.7 0.9 5.0 1.7 Northcentral 36.2 6.4 61.0 50.0 51.9 25.6 74.0 61. 6 46.1 18.4 71.8 54.7 2.3 0.9 3.4 2.3 Cold Spring 0.0 25.0 45.0 50.0 80.0 80.0 54.2 60.6 66.7 59.4 50.5 55.5 3.2 1.9 2.5 1.7 Dinosaur 20.0 28.6 39.7 37.0 50.0 35.7 48.1 53.5 40.6 33.3 44.4 45.1 2.2 3.0 2.3 1.4 Moffat County South 62.5 14.3 50.0 42.1 80.0 0.0 77.8 66.7 72.2 10.0 72.7 52.9 2.7 0.8 3.6 1.6 Four-year averages 35.1 14.9 50.3 46.1 55.7 39.6 65.3 58.4 48.3 30.0 60.S 51.5 2.3 1.2 3.1 1.9 N VJ -'=' �Table 2l. Estimated sage grouse hatching dates, Hoffat County, Colorado 1979.a Hales Cold Blue Spring Mtn. .M.C. South 11-17 May 0.9 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 18-24 May 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.3 1.6 0.0 0.0 0.5 25-31 May 2.7 1.7 3.0 0.0 4.4 2.0 3.9 3.1 2.6 0.0 6.1 2.9 1-7 June 15.2 12.2 6.7 4.1 21.7 10.9 11.8 6.6 7.3 0.0 9.1 6.6 8-14 June 27.8 24.1 12.7 16.5 26.0 21.6 39.2 36.0 17.3 12.1 33.3 30.1 28.5 24.2 18.6 43.5 26.4 23.5 24.Lf 25.7 32.7 30.3 25.6 15-21 June Totals M.C. East Females Blue Cold Spring Htn. N.C. Dates N.C. South -- H.C. East N.C. Totals N 22-28 June w 16.1 23.8 33.3 38.1 0.0 25.6 9.8 19.4 23.0 32.7 12.1 20.3 29 JuneS July 8.8 7.6 14.0 18.6 4.4 10.1 11.8 8.0 15.7 17.8 6.1 10.8 6-12 July 4.4 2.1 6.1 4.1 0.0 3.3 0.0 1.7 6.3 3.7 3.0 2.7 13-19 July 0.0 0.0 0.0 0.0 0,0 0.0 0.0 0.3 0.5 0.9 0.0 0.4 20-26 July 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.1 Sample sizes 112 474 165 97 23 871 102 578 191 107 33 1,011 a In percent. VI �236 at check stations, only 59 (7.1%) of those hunting did not have the required permit. This contrasts with 8.4% of 770 hunters contacted in 1978. If this percentage reflected the hunter universe in 1979, then it is possible that there were 2,863 sage grouse hunters (2,673 +190) in Moffat County in 1979. However, when it is realized that of the 2,673 permittees, only 2,041 (Table 22) hunted, then it is reasonable to conclude that there were no more than 2,200 sage grouse hunters in Moffat County in . 1979 (2,041 + 145). Origin of permittees was primarily from Moffat (34.4%), Rio Blanco (8.2%) and Mesa counties (8.1%). Counties in the Denver Metropolitan area (Adams, Arapahoe, Denver, Douglas, Jefferson) collectively comprised 24.0% of the total, slightly higher than the 19.7% these counties contributed in 1978. Other counties providing 1.0% or more of the'total hunters were Routt (5.0%), Boulder (4.2%) Garfield (2.9%), Larimer (2.5%), EI Paso (2.0%) and Montrose (1.0%). All other counties and states each contributed less than 1.0% of the'total permittees. These data are similar to those collected in 1978 and are not markedly different from those collected at check stations. Questionnaires were sent to all permittees immediately following the sage grouse season in Moffat County (28 September). Responses were received from 1,573 permittees. On 22 October, follow-up letters and questionnaires were sent to all non-respondents and 455 additional responses were received. In all, 2,028 permittees (75.9%) responded (Table 22). Sixty-seven questionnaires (2.5%) were undeliverable. Mean values calculated for permittees responding to the follow-up letter were used to project for the 645 (24.1%) non-respondents. This undoubtedly somewhat inflated total estimates as the available evidence indicates that non-respondents are even less successful than respondents to the follow-up letter. Data presented indicated that about 70% of the permittees hunted and harvested (including crippling loss) an estimated 7,840 sage grouse in Moffat ·County. About 78% of all hunters were successful and harvested 4.4 grouse each. Fewer permittees hunted in 1979 than in 1978 (70 vs 75%) but those that did hunt were slightly more successful (78 vs 75%, 4.4 vs 4.2 grouse per successful hunter). Time period of hunting was receIved for 1,691 hunts which resulted in 4,875 grouse being bagged. Based on this sample, 47.3% of the hunts were on the ~st weekend and resulted in 56.5% of the total harvest. The l~t 5 week days of the season attracted 15.4% of the hunts which resulted in 13.0% of the total harvest. During the 2nd weekend, 15.8% of the hunts resulted i~ 14.0% of the harvest. The 2nd week of the season attracted 6.5% of the hunts and resulted in 5.2% of the harvest. Surprisingly, 15.0% of the hunts occurred during the 3rd weekend but only 11.2% of the total harvest was bagged. Six hundred and ten of the 1,589 hunters responding (38.4%) reported achieving the bag limit at least 1 day during the season. Of this sample, 229 hunters achieved the bag limit on I day, 331 did on 2 days, 27 did on 3 days, 16 did on 4 days, 2 did on 5 days, 3 did on 6 days and I each achieved the bag limit on 7 and 9 days. These data are remarkably similar to those reported in 1978 when 39.0% of the hunters achieved the bag limit on .at least 1 day. �237 Table 22. Moffat County sage grouse hunter questionnaire data, 1979. Projected for 1,573 No. in sample Percent of total permittees Percent of total hunters of non-hunters No. of successful hunters Percent successful hunters Percent success of permittees No. of grouse bagged Grouse/hunter Grouse/successful hunter Grouse/permittee No. birds lost Birds lost/hunter Total harvest Percent 1,054 cr Lpp Lfng loss 248 2,028 75.9 1,589 78.4 439 21.6 1,302 645 24.1 452 70.1 193 29.9 351 2,673 100.0 2,041 76.4 632 23.6 1,653 83.0 77.7 81.9 77.7 81.0 67.0 54.5 64.2 54.5 61.8 2.2 Days/hunter 136 19.3 29.9 2,838 No. of hunter days 319 80.7 70.1 303 No. of non-hunters Percent 58.8 17.0 1,270 No. of hunters 455 Projected for 4,875 661 2.1 984 3,499 2.2 5,859 949 2.1 1,401 4,448 2.2 7,260 3.8 3.1 3.7 3.1 3.6 4.6 4.0 4.5 4.0 4.4 3.1 2.2 2.9 2.2 2.7 441 58 0.35 0.18 5,316 1,042 8.3 5.6' 499 0.31 6,358 7.8 81 0.18 1,482 5.6 580 0.28 .7,840 7.4 �238 The questionnaire was designed to examine harvest and hunter activity by Small Game Units and harvest zones (Fig. 1). Data presented (Table 23) LndLcat.e that 54.3% of the hunters were in Unit 16. These hunters harvested ·52.,1%of the total birds taken in Moffat County. The next most important ha.ryest units were 18~ 20 and 14. Few hunters hunted in and harvested extremely few birds in Unit 26 in Moffat County. These data are almost identical to those obtained in 1978. Ta.ble 23. Hunter activity and harvest by Small Game Management Units and harvest zones~ Moffat County~ Colorado 1979. Unit (Harvest Zone) No. of hunters Percent of total No. of birds harvested Percent of total 14(A) 184 10.9 590 10.1 16(A) 314 18.7 1,011 17.3 (B) 445 26.5 1,628 27.8 (C) 147 8.7 389 6.6 7 0.4 23 0.4 198 11.8 838 14.3 (B) 140 8.3 378 6.5 (C) 23 1.4 71 1.2 20 (A) 123 7.3 618 10.5 (B) 10 0.6 31 0.5 26 (A) 32 1.9 64 1.1 Unknown 58 3.5 218 3.7 100.0 5,859 100.0 (?) 18(A) Totals aTota1 does not equal 1,589 as some hunters hunted in more than 1 zone. �239 Vulnerability Bands from 115 sage grouse banded in Moffat County were reported in the 1979 recovery year. Of this sample, 109 were shot recoveries, 2 were road kills and 4 were found dead. These of those found dead were during the 1979 hunting season. Twenty-four of the recoveries were of birds banded in 1978 (11 males, 13 females) while 91 (63 males, 28 females) were banded in 1979 (Table 24). Table 24. Year Sage grouse banding and recovery data, Moffat County, 1978-79. Number recovered 1978 1979 No. banded Males-chicks 1978 217 1979 139 20 8 17 ".. Yearlings 1978 25 1979 313 4 1 18 -Adults 1978 24 1979 451 5 1 28 Females-Chicks 1978 204 1979 178 16 8 17 -Yearlings 1978 13 1979 93 1 3 6 -Adults 1978 23 1979 76 a 2 1 5 Two unknown age sage grouse (1 male and 1 female) banded in 1978 and recovered in 1979 are not included. �240 Data presented (Table 25) suggest that chick males and females are more vulnerable to hunting than other age and sex classes. Statistical analyses of the data indicate that the differences are not real (f > 0.05) and that those apparent may be due to chance or sampling error. Overall hunting mortality (direct recovery rate = harvest rate) of sage grouse in 1979 was less than 10% and was less than the 9.5% recorded in 1978. Table 25. Sage grouse banding and recovery data, 1979 bandings, Moffat County, Colorado, 1979. Age and sex class Number banded Number recovered Harvest rate (%) Females Chicks 178 Yearlings 93 17 6c Adults 76 5 347 28c 7.8a Chicks 139 Yearlings 313 17 lac 12.2 5.4a Adults 451 2Sd 5.Sb Subtotals 903 63 6.6a,b 1,250 91 7.0a,b Subtotals 9.6 6.6 5.4a Males Totals aExcluding I road kill. bExcluding 2 found dead near power line. cIncludes I road kill. d Includes 2 found dead. Direct recovery rates were compared by area, sex and age (Table 26). Recovery rates were low south of U.S. 40 (2.3%),east of the Baggs Highway (3.1%) and west and north of Sunbeam (4.8%). They were highest at Cold Spring Mountain (10.0%) and Blue Mountain (8.5%) with the important harvest area in northcentral Moffat County being intermediate (7.7%). Recovery rates were not high for any age or sex class or area. The rates of 16.7 and 14.3% for chick females and males, respectively, in the northcentral zone were based on few �241 bandings (6 females, 7 males) (Table 4). These data indicate that sage grouse harvest is not excessive or even near the potential in any zone in Moffat County for any age or sex class. Table 26. Direct recovery rates of banded sage grouse, Moffat County, 1979. Males Females 1- 2+ 7.1 13.6 South of U.S. 40 2.9 East of Baggs Highway Cold Spring Mountain 12.2 1- 8.4 2+ All birds 9.7 7.7 10.0 1.4 7.1 0.0 2.3 5.7 1.3 5.6 0.0 3.1 Northcentral 14.3 7.0 8.3 16.7 10.5 0.0 7.7 Blue Mountain 11.1 6.4 7.8 11.1 0.0 25.0 8.5 2.9 7.7 0.0 0.0 4.8 5.8 6.2 6.5 6.5 7.3 Sunbeam West & North All Areas 12.2 9.6 LITERATURE CITED Beck, T. D. I., R. B. Gill, and C. E. Braun. 1975. Sex and age determination of sage grouse from wing characteristics. Game Inf. Leaflet No. 49 (Revised). Colorado Div. Wildl. 4pp. Braun, C. E., and T. D. I. Beck. 1976. Effects of sagebrush control on distribution and abundance of sage grouse. Colorado Div. Wildlife, Final Rep., Fed. Aid Proj. W-37-R, Work Plan 3, Job 8a. pp. 21-84. Eng, R. L. 1955. A method for obtaining sage grouse age and sex ratios from wings. J. Wildl. Manage. 19:267-272. Gullion, G. W. 1965. Improvements in methods for trapping and marking ruffed grouse. J. Wildl. Manage. 29:109-116. Hoffman, R. W., and C. E. Braun. 1975. A volunteer wing collection station. Game Inf. Leaflet No. 101. Colo. Div. Wildl. 3pp. Kabat, C., I. O. Buss, and R. K. Meyer. 1948. The use of ovulated follicles in determining eggs laid by the ring-necked pheasant. J. Wildl. Manage. 12:399-416. Lacher, J. R., and D. D. Lacher. Wildl. Manage. 28:595-597. 1964. A mobile cannon net trap. J. �242 Pyrah, D. G. 1963. Sage grouse investigations. Idaho Fish and Game Dep., HildI. Rest. Div., Job CompL, Rep., Fed. Aid Proj. W-12S-R. 71pp. Rogers, G. E. 1964. Sage grouse investigations in Colorado. Dep. Game, Fish and Parks, Tech. Publ. 16. 132pp. Prepared by _~'-~.~({.=:'~:::::.!...-=c:::::..=:c::""?=-._-~~~-==-~~ Clait E. Braun Wildlife Researcher C Colo. �April 243 JOB PROGRESS No. Job Title 3 Potential Job No. Covered: 12 Impacts of Strip Mining on Sage Grouse Movements and Habitat Personnel: Game Bird Survey W-37-R Work Plan No. Period REPORT Colorado State of Project 1980 Use 1 January - 31 December, 1979 R. A. Ryder, Colorado State University; Clait Braun, Steve Emmons, Howard Funk, Sue McElderry, Brett Petersen, Tom Schoenberg, John Wagner, Colorado Division of Wildlife. ABSTRACT Investigations concerning sage grouse (Centrocercus urophasianus) movements and habitat use were initiated in North Park during 1979. Three male and 4 female sage grouse were followed using radio telemetry from 18 April to 1 July. Daily movements from the lek by males averaged 0.8 km (range 0.1-2.4). Ninety-six percent of all movements were within 2.0 km of the lek. The mean distance from the lek of 3 nest sites was3.0 km (range 0.2-6.4). Daily movements from the nest site of 4 hens during the pre-incubation period ranged from 0.3-1.7 km. Five feeding sites used during incubation ranged from 60-300 m and averaged 153 m from the nest. Daily movements of 2 hens with broods during June averaged 117 m and ranged from 100-1100 m. Mean sagebrush canopy coverage was greatest at male feeding-loafing sites (32.5%) followed by female pre-incubation feeding-loafing sites (29.2%) and nest sites (27.2%). Average sagebrush canopy coverage was greater at randomly located sites (26.3%) than either brood-rearing sites (23.1%) or feeding sites of hens during incubation (19.8%). The greatest average height of sagebrush was at nest sites (30.0 cm) followed by male feeding-loafing sites (28.1 cm), female pre-incubation feeding-loafing sites (22.6 cm) and brood-rearing sites '(19.1 cm). Only feeding sites of incubating hens had lower average sagebrush height (16.0 cm) than randomly located sites (18.9 cm). Pellet transect counts revealed a 74% decline in sage grouse use of the study area between winter and summer, 1979. Harvest data collected at check stations and wing collection barrels showed an increase of 0.5 birds/hunter from 1978 and an increase of 0.9 birds/hunter over the previous 5 year average. �244 RECOMMENDATIONS 1. Continue lek censuses within the study and throughout North Park. 2. Continue banding sample of 50 males in each quadrat of North Park. 3. Maintain transmitters on 5 males and 5 females from January through August to provide seasonal movement and habitat selection data. 4. Continue counting and clearing sage grouse pellet transects each spring and fall. 5. Operate check stations and wing collection barrels during the sage grouse hunting season. �245 POTENTIAL IMPACTS OF STRIP MINING ON SAGE GROUSE MOVEMENTS AND HABITAT USE Thomas J. Schoenberg Studies of sage grouse movements and habitat use have been conducted in Montana (Wallestad 1971, Eng and Schladweiler 1972, Wallestad and Pyrah 1974, Wallestad and Schladweiler 1974), Idaho (Klebenow 1969) and Colorado (Beck 1977). While these studies provide useful data concerning the sage grouse-sagebrush (Artemisia tridentata) relationship, more detailed studies are required in areas likely to be disturbed by strip mining of coal and other energy developments. It is especially important to better understand the sage grousesagebrush relationship in North Park since currently proposed coal mines will jeopardize historic breeding, nesting, brood-rearing and wintering habitats. The sage grouse population is locally migratory with wintering concentrations located in the northeast quadrat (Beck 1975) in areas proposed for mining. Habitat disturbance in the northeast quarter of North Park will impact the entire population. P. N. OBJECTIVES The objectives of this study during the predevelopment period are to: 1. Ascertain seasonal movement patterns and habitat preferences of male and female sage grouse within the area to be impacted. 2. Continue population monitoring on known sage grouse leks within the study area. Data from monitoring population trends in adjacent areas and throughout North Park continued by research and management personnel will be used for comparative purposes. 3. Prepare management plan for predicting, impacts on the sage grouse resource. 4. Based upon findings, prepare plan for rehabilitation of developed areas that would be attractive to and effective for maintaining and enhancing sage grouse populatio~s. Hypotheses reducing, and mitigating being tested are: a. Movements of sage grouse within sex class. b. Habitat class. selection (preference the study area differ for each and need) differs for each sex �246 c. Movement patterns for each sex class vary during critical periods of the annual cycle (prebreeding, breeding, nesting and brood rearing). d. Habitat use varies and brood rearing) seasonably (prebreeding, for each sex class. breeding, nesting, SEGMENT OBJECTIVES 1. Review available literature concerning niche description and measurement, habitat use by sage grouse, sage grouse movements, use of radio telemetry to identify habitat selection by birds, and western mineland reclamation. 2. Select and grid the study area identifying and winter use sites. 3. Randomly locate 20 sage grouse pellet transects within the study area. Transects 0.5 km long and 2.5 m wide will be searched in late Mayor early June and late September or early October depending upon snow cover. Droppings found will be classified as to color, consistency, number within 25 cm and rated as to size. All droppings found will be cleared from the transect. 4. Continue intensive counts (2-3 per week) of all sage grouse present on each known lek (Prague, Denmark, Pronghorn, Raven and Roth) within the study area each year. Emphasis will be placed on leks likely to be directly impacted (Denmark and Pronghorn). Counts will be in the 0430-0730 interval from about 20 March to 31 May. Birds present will be classified as males and females. Counts of sage grouse present on other leks (4/season of leks with more than 10 males) in North Park will be continued. 5. Continue banding samples of male and female sage grouse within the study area. Grouse will be located where they roost by spot lighting and will be trapped with long handled nets. Some trapping of hens with cannon nets on leks may be necessary. Birds trapped will be banded with serially numbered aluminum bands and color coded (location) bandettes. A sample of 20% of the high spring count on each lek is desired each year. Samples of 50 cocks banded in each of the other 3'quadrats within North Park are desired for comparative purposes. 6. Equip 10 male and 10 female sage grouse with radios to ascertain movement patterns and habitat use preferences within the study area. While year around documentation of habitat use is desirable, emphasis will be placed on spring (March-June) and late winter (January-February). Movements will be mapped and vegetation composition, height and canopy coverage will be ascertained for adequate samples (present sample size unknown) of habitats utilized by season for necessary functions of sage grouse. known leks, brood �247 7. Population size will be estimated utilizing counts of birds present on leks, established patterns of lek attendance and known sex ratios, and ratios of banded to unbanded birds in harvest samples. 8. Harvest data will be obtained from check stations and volunteer wing collection barrels. Check stations will be operated at Gould, State Line and Willow Creek Pass during at least the opening weekend of the hunting season. A sample of 500 wings is desired. 9. Number and location of marked birds harvested through field checks of hunters and voluntary will be obtained mail reporting. 10. Age composition of the harvest will be determined through examination of wings collected during field checks and through use of wing collection barrels. 11. Compile data, analyze results, and prepare progress and/or completion reports. Suitable findings will be submitted to appropriate technical journals for publication consideration. Management recommendations will be included in the final report. METHODS AND MATERIALS Seven leks in the study area were censused on 94 occasions between 22 March and 1 June. Counts were made between 0430 and 0730 MST (Braun and Beck 1976). Perdiz, a previously unknown lek, was discovered on 23 April south of the Wyoming Fuels coal mine while locating a radio-marked hen. Sage grouse were captured while roosting along roads and on leks throughout North Park. Trapping was accomplished using a hand-held spotlight and long-handled net (pyrah 1959). Sage grouse were marked with serially numbered aluminum leg bands and color coded plastic bandettes. Three males and 4 females trapped on or near Raven lek were fitted with 164 MHz transmitters mounted on tail clips (Bray and Conner 1972) attached to the 2 centralrectrices. Data on sex and age (Eng 1955), weight and primary molt were collected for each bird. Radio-marked sage grouse were located daily between 0500-2100 using portable receivers with hand-held 3-element yagi antennas. Feedingloafing sites were triangulated with prominent geographic features and plotted on a map of the study area. Climatic and physiographic features, and flock size and composition were recorded at each site. All sites were flagged for later vegetation description and measurements. �248 Vegetation measurements were made along 20 m of line transect using Canfield's (1941) line-intercept technique. Two 10 m lines were oriented along N-S and E-W axes with the bird location as the center of the plot. Measurements included intercept distance (nearest em) for foliated and unfoliated big sagebrush, forbs, grasses, litter and bare ground. Additional measurements of big sagebrush included dimensions (LxWxH) of the largest individual plant in each clump intersected. Soil samples were collected at each site. Twenty sage grouse pellet transects were randomly chosen from 200 possible points on a grid map of the study area. Transects 500 m long and 2.5 m wide were placed along a N-S axis and marked with steel fence posts at either end. Two red-painted and flagged re-bar pegs were placed along each t ransec t as well as flagging tape at 25 m intervals to aid in accurate alignment prior to counting and clearing. All sage grouse pellets encountered were classified to 1 of 4 age categories (1 = 1 year+, 2 = winter, 3 = spring, 4 = fresh) based on color and consistency. Three types of dropping groups were classified based on the number of pellets within a 25 cm diameter (1-5, 6-14, 15+). Sixty random vegetation analysis sites were chosen from 4 random points along each of 15 pellet transects. Wing collections were made during the sage grouse hunting season (8-23 September) at 4 volunteer wing collection barrels and 3 check stations. The wing collection barrels were maintained throughout the entire 16 day season at Gould, State Line, Willow Creek Pass and Muddy Pass. Check stations were operated at Gould and Willow Creek Pass on 8, 9 and 16 September, and at State Line on 8 and 9 September. Data collected from each party of hunters at check stations were given by Braun (1979) and included county of origin; number of hunters; hours hunted (total of all hunters); number of grouse observed, bagged, and lost; number of banded birds bagged and location where shot; and the area hunted. In addition, individual hunters were asked whether or not they normally hunted sage grouse in North Park or if they were 1st time sage grouse hunters. DESCRIPTION OF AREA The study area is in the northeast quadr-at of North Park east of Colorado Highway 125 and north of Colorado Highway 14. It is bounded on the north and east by the Canadian River and on the south and west by the Michigan River. The vegetation is characterized by the sagebrush-grassland type with native and irrigated meadows along major stream courses. Detailed geographic, geologic, vegetational and climatic features of the area have been described by Gill (1965), Carr (1967), Beck (1975), Braun and Beck (1976) and will be treated in the final report. There are 5 leks in the primary study area and 2 others north of the Canadian River. Areas of wintering sage grouse concentrations have been identified by Beck (1975). Movements and habitat use studies were concentrated on sage grouse from Raven lek since it will be the 1st to be impacted by coal mining. �249 RESULTS AND DISCUSSION Lek Census Seven leks in the study area were censused. Raven and Denmark leks were censused most intensively (2-3 times/week). Pronghorn and Perdiz leks were censused 1-2 times/week while Canuck, Prague and Roth leks were censused once every 2 weeks. Dates of peak lek attendance for males occurred between 17 April and 26 May (Table 1) with the peak for all leks occurring during the week of 13-19 May (Fig. 1). Peak hen attendance occurred between 19 and 25 April with the peak during the week of 15-21 April, 4 weeks prior to peak male attendance. No 2nd lower peak for females occurred as has been reported previously (Dalke et al. 1963, Eng 1963, Gill 1965). The late arrival on leks and rapid increase in lek attendance by both males and females during the 1st 2 weeks of April and the lack of a lower peak for females was attributed to heavy overwinter snow accumulation accompanied by a late spring thaw. Approximately 90% of the study area was covered with an average of 60 cm of snow at the end of March. By mid-April only about 25% of the study area was snow-covered. Table 1. Lek Denmark Raven Prague Canuck Perdiz Pronghorn Roth aTwo cocks Peak counts of sage grouse on leks, northeast Colorado, 1979. Males 136 63 43 21 16 10 Oa (1 strutting) Dates 15 May 17 April 23 May 21 April 26 May 2 May All dates North Park, Females 66 26 5 7 4 4 0 Date 22 April 22 April 28 April 23 April 24, 25 April 19 April All dates 0.3 km N of the lek on 25 April. High spring lek counts for 1978 and 1979 differed (Table 2). The apparent 20% increase in number of cocks in 1979 is due in part to the discovery of Perdiz. Since the study area was censused more intensively in 1979, the apparent increase may be an artifact of sampling rather than an actual population increase. �250 60 • / ._ I -: ./ -. -, I I / -, I/ ' "" -, / '. Males -, . " • I I I I I A_ I I J 10 / / I I I / I -6 / I ' '\ I \~ " Females <, '6--6 --0_ to N I N N APRIL Fig. 1. MAY JUNE Sage grouse lek attendance on 7 leks, northeast North Park, Colorado, 1979. �251 Table 2. Peak sage grouse lek counts, northeast 1978-1979. Lek Cocks 1979 1978 Denmark Raven Prague Canuck 80 94 35 21 10 1 136 63 43 21 16 10 0 241 289 Per d Lz+ Pronghorn Roth Totals Avg. a North Park, Colorado, 1978 Hens 1979 40 52 3 10 66 26 5 7 + 70 - 53 + 23 % b. + 65 - 50 + 67 - 30 L~ 6 0 4 0 III 112 -100 - 33 + 20 + 1 Found in 1979. The dramatic increase in 1979 in number of cocks on Denmark lek and decrease on Raven lek was explained from examination of the age ratio of birds captured on each lek (Table 3). Of 22 cocks captured on Raven lek, all were adults. The age ratio of birds trapped on Denmark lek was approximately 1:1 (13 juveniles, 15 adults). There was apparently little or no recruitment of yearlings to the Raven lek with excellent recruitment to Denmark lek. Reasons for the disparity are unknown. Based on 1979 census data, total numbers of birds on each lek increased at the same rate. Also, initial observations of strutting males on each lek occurred at approximately the same time. Table 3. Lek Sage grouse trapping success on 4 leks, northeast Park, Colorado 1979. Juvenile Cocksa Adult Totals Denmark Raven Perdiz Pronghorn 13 0 1 1 15 22 3 1 28 22 4 2 Totals 15 41 56 aNot including b%HSP %HSp15 recaptures. = % of the high spring count. 21% 36% 25% 20% Juvenile North Hens Total Adult 3 1 0 0 4 3 0 0 7 4 0 0 4 7 11 %HSP 11% 15% 0% 0% �252 Banding Sample Trapping was conducted throughout North Park on 46 nights from 21 March to 22 May. During this period, 257 cocks (113 juveniles, 144 adults) and 100 hens (48 juveniles, 52 adults) were banded. Thirtysix cocks and 7 hens were recaptured. A hermaphrodite, previously unreported in the literature, was trapped and collected in the southeast quadrat of North Park on 6 April. Four leks within the study area were trapped on 24 occasions between 16 April and 22 May (Table 3). Fifty-six cocks (15 juveniles, 41 adults) and 11 hens -(4 juveniles, 7 adults) were banded. Recaptures of birds on leks included 9 adult cocks. The desired goal of banding 20% of the high spring count of cocks on each lek was achieved with values ranging from 20% on Pronghorn to 36% on Raven. Including roadside captures, 79 cocks (23 juveniles, 56 adults) and 38 hens (17 juveniles, 21 adults) or 33% of the total banded sample for North Park was captured within the study area. Radiotelemetry Radio-marked Investigations Birds Three adult cocks and 3 hens (2 adults, 1 juvenile) from Raven lek, and 1 juvenile hen (3415) captured 2.9 kffisouth of Raven, were fitted with transmitters between 18 April and 9 May (Table 4). All birds were located daily between 0500 and 2100 until 11 June. Thereafter, radio locations were made only on weekends. There were 45 on-Iek locations and 99 feeding-loafing site locations during this period for the 3 radio-marked cocks and 146 locations for the 4 hens. There were 69 pre-incubation feeding-loafing site locations, 5 feeding locations from 3 hens during incubation, and 14 locations of 2 hens with broods. The remaining 59 locations were from 3 hens on nests during incubation. Excluding hen 3411, the average transmitter life was 57 days. Hen 3411 lost her transmitter between 18 and 20 May, after only 10-12 days of operation. The bent antenna and badly frayed rectrices indicated she had probably pulled her rectrices out to remove the transmitter package. Hen 4796 was depredated by a raptor between 18 and 20 June, after 61-63 days of transmitter operation. She had been rearing a brood for 14-16 days at that time. Movements of Males Daily movements from the lek by males averaged 0.8 km (range 0.1-2.4). Ninety-six percent of all movements were within 2.0 km of the lek. Movements from the lek were primarily northwest (39%) and southeast (33%) followed by southwest (19%) and northeast (9%). Adult male 7327 disappeared from the Raven lek area on 27 May and was not located until 10 June. He was found loafing with 25 other males approximately 1 km west of Denmark lek, a distance of 7 km from Raven. It is not known whether he returned to the Raven lek �Table 4. Sage grouse telemetry data, northeast North Park, Colorado, 1979. Number Sex Age Capture site Capture date Date of last location Total a locations 7181 M Ad. Raven lek 18 April 10 June 53 53 7327 M Ad. Raven lek 18 April 10 June 44 53 7367 M Ad. Raven lek 25 April 24 June 47 60 4796 F Ad. Raven lek 18 April 20 June 49 63 3401 F Juv. Raven lek 25 April 24 June 45 60 Transmitter b life (days) Remarks Raptor kill N VI 3411 F Ad. 3415 F Juv. "2.9 k.m So. of Raven Raven lek 8 May 9 May 20 May 1 July 14 12 38 53 alncludes both on and off lek locations but not capture site. bR epresents mlnlmum "l'f 1 e. Lost transmitter before incubation w �254 area since his transmitter had quit by the next location attempt on 17 June. Both other radio-marked males remained in the Raven lek area until their radios shut off. Movements of Females All 3 hens captured on Raven lek moved south to select nest sites. Hen 3401 nested 0.2 km from the lek alongside the William's Draw stream. Hen 4796 nested 6.4 km from the lek, 0.7 km southeast of Perdiz lek. Hen 3411 lost her transmitter before her nest was located but had confined her movements in an area 2.4 km from Raven lek. Hen 3415 nested 3 km WSW of Raven lek. It is not known where she was bred since she was trapped off lek after breeding. During the pre-incubation period daily movements from the nest site were within a 0.3 km radius for·hen 4796, a 1.2 km radius for hen 3401; a 1.5 km radius for hen 3411 and a 1.7 km radius for hen 3415. Five feeding sites during incubation ranged from 60 - 300 m from the nest site. Average distance from the nests to feeding sites was 153 m. Daily movements of 2 hens with broods ranged from 100 - 1100 m for 9 locations. During the 1st 6 days after hatching, hen 4796 had moved 0.7 km north of her nest averaging 117 m between successive daily locations. Sixteen days after hatching, her kill site was found 1 km east of her nest. Successive daily locations for hen 3415 with her brood were made on 2 occasions. Between 23 and 24 June she moved 1100 m with her 3-5 day old brood. Between 30 June and 1 July she moved only 90 m. Nest Success Nests of 5 hens (3 radio-marked, 2 unmarked) were located in the study area. One other radio-marked adult hen. (3411) lost her transmitter 10-12 days after attending Raven lek. While her movements became localized just prior to loss of the radio package suggesting she was laying somewhere within a restricted area, subsequent searches to locate her nest failed. Of·the 5 nests located, 3 hatched (60.0%) and 2 were depredated and/or abandoned. Adult hen 4796 initiated incubation of ~ eggs on 9 May, 20 days after attending Raven lek. On 4 June, the 26th day of incubation, 7·of 8 eggs hatched; the 8th egg had failed to develop. Juvenile hen 3401 initiated incubation of 6 eggs on 16 May, 20 days after attending Raven lek. She had not hatched her clutch by 11 June, the 26th day of incubation. Between 11 and 16 June, 3 of the 6 eggs were depredated by an unknown predator. The hen abandoned the remaining 3 eggs, 2 of which had just begun to pip. The 6th egg was infertile. Juvenile hen 3415 initiated incubation of 6 eggs on 24 May. A 7th egg was laid on 25 May. Since she was not trapped on a lek it was not possible to determine how many days after lek attendance she initiated incubation. All 7 eggs hatched between 17 and 23 June, 25 to 29 days after initiation of incubation. �255 Nests of 2 unmarked hens were located within the study area. One nest with 8 eggs, located on 9 May, was depredated by a badger or coyote on 22 May. Another nest containing 8 eggs was located on 11 June. All 8 eggs had hatched by 16 June. Habitat Selection Vegetation measurements were made at 60 random sites, 59 sites for males and 57 sites for females including 35 pre-incubation feedingloafing sites, 5 nests, 5 feeding sites during incubation and 12 brood-rearing sites. Data on species composition (canopy coverage) and height of sagebrush we re tabulated for all sites (Table 5). Average sagebrush height for random sites was 18.9 cm. Except for feeding sites of incubating hens, all other sage grouse use sites exceeded this height, The greatest average height of sagebrush was at nest sites (30.0 cm) followed by male feeding-loafing sites (28.1 cm) and female pre-incubation feeding-loafing sites (22.6 cm). Beck (1977) found that feeding-loafing sites for females during winter had greater average height of sagebrush than did sites used by males. The opposite occurred during the breeding season. Table 5. Species composition and mean sagebrush height at sage grouse use and random sites, northeast quadrat, North Park, Colorado, 1979. Sitea classification 1. 2. 3. 4. 5. Random Male FL Female PIFL Female N Female IF 6. Female BR "z Species composition N Sagebrush height (em) 60 59 35 5 5 12 18.9 28.1 22.6 30.0 16.0 19.1 Sagebrush 26.3 32.5 29.2 27.2 19.8 23.1 Forbs 9.~. 7.0 9.6 6.6 8.4 8.7 (%) Grass Litter Bare ground 1.1 3.0 1.4 1.4 1.8 1.8 2.8 7.4 4.6 12.2 7.4 7.8 60.4 50.1 55.2 52.6 62.6 58.6 male feeding-loafing, 3 = female pr e'-dncubat f.on feeding-loafing, 4 = nest site, 5 = feeding site during incubation, 6 = brood-rearing site. == �256 Mean sagebrush canopy coverage was greatest at male feeding-loafing sites (32.5%) followed by female pre-incubation feeding-loafing sites (29.2%). This was opposite of what Beck (1977) described for wintering sage grouse. Wallestad and Schladweiler (1974) found an average sagebrush canopy-coverage of 32% for males during the breeding season in Montana while Emmons (1979) found this value to be 29.3% for males during the breeding season in Colorado. Two of the 5 sage grouse use types had average sagebrush canopy-coverage below the 26.3% mean of the random sites. These were brood-rearing (23.1%) and feeding sites of incubating hens (19.8%). These sample sizes are probably inadequate for valid comparisons. Eighty-one percent of all locations for male grouse were in canopy coverage classes greater than 20% (Table 6). This compares with 82% reported by Eng -and Schladweiler (1972) in Montana. Similarly, 80% of the female pre-incubation feeding-loafing sites were in sagebrush canopy coverage greater than 20%. These values contrast markedly with the random sites where only 67% had a sagebrush canopy coverage greater than 20%. Female sage grouse appear to select for more open areas of sagebrush for feeding during incubation and brood-rearing than they do for preincubation feeding-loafing. Eighty percent of the feeding sites of incubating hens and 75% of the brood-rearing sites had sagebrush canopy coverage less than 30%, while only 63% of the pre-incubation feedingloafing sites had sagebrush canopy coverage less than 30%. Pellet Transects Pellet transects were counted and cleared between 25 May and 7 June and again on 27 and 28 October. Total numbers of droppings counted on all transects were compared for the winter-spring and summer-fall periods (Table 7). Using these totals as a measure of intensity of use indicated there was an overall 74% decline in use between winter and summer, 1979. This would indicate that sage grouse winter use of the study area was approximately 4 times greater (6886/1792 = 3.8) than summer use. This winter to summer decrease reflects a change in distribution of sage grouse throughout North Park. The population is locally migratory with winter movements toward the north where snowfall is lower and more sagebrush is available (Beck 1977). Of 20 transects counted, numbers of droppings decreased on 13 and increased on 4, while 3 transects had no droppings on either count. The average decrease was 82% and varied from 45 to 99%. The average increase was 62% and ranged from 12 to 97%. Of particular interest was transect 179 where the increase in total droppings was 97%. It lies immediately south of the Canadian River and borders native and irrigated meadows. Brood use of moist lowland areas with succulent vegetation is well documented (Patterson 1952, Petersen 1970, Wallestad 1971). This explains the dramatic increase in number of droppings on this transect. �Table 6. b Sagebrush canopy coveragea distribution for random and sage grouse use sites , northeast quadrat, North Park, Colorado, 1979. Canopy coverage class(%) Random (N = 60) Male FL (N = 59) Female PIFL (N = 35) Nest (N = 5) Female IF (N = 5) Female BR (N = 12) 0.1 - 10.0 13 0 3 40 20 25 10.1 - 20.0 20 19 17 40 40 17 20.1 - 30.0 27 31 43 0 20 33 30.1 - 40.0 22 20 17 0 20 8 40.1 - 50.0 18 20 14 0 0 8 50.1 - 60.0 0 7 3 0 0 0 60.1 - 70.0 0 3 3 20 0 8 a In percent. bMale FL = feeding-loafing, Female PIFL = pre-incubation feeding-loafing, Female IF site of incubating hen, Female BR = brood rearing. feeding N Vl -....J �258 Table 7. Numbers of sage grouse pellets encountered on 20 transects, northeast quadrat, North Park, Colorado. Winter and summer, 1979. Transect 3 4 48 57 60 74 78 81 83 88 99 115 129 136 166 174 179 194 196 197 Totals Winter accumulation Summer accumulation % change 0 0 232 7 328 159 1087 23 52 455 917 19 1179 1319 51 0 20 334 342 362 0 0 2 3 182 51 72 114 121 14 81 7 242 106 57 0 661 4 58 17 0 0 -99 -57 -45 -68 -93 +80 +57 -97 -91 -63 -79 -92 +12 0 +97 -99 -83 -95 6886 1792 -74 Droppings were not encountered on 3 transects during either count. This was understandable on transect 174 which lies north of the Canadian River traversing the East Sand Hills. Dominant vegetation on the transect is rabbitbrush (Chrysothamnus spp.) and bitterbrush (Purshia tridentata). Harvest Data Check Stations Sage grouse harvest data from Gould, State Line, and Willow Creek Pass check stations were tabulated (Table 8); During 8 check-station days, 521 hunters bagged 982 sage grouse or 1.9 birds/hunter. This represents an increase of 0.5 birds/hunter from 1978 and an increase of 0.9 birds/hunter over the previous 5 year average (1974-1978) as reported by Braun (1979). There were 57 banded birds reported at check stations representing 5.8% of the total bag. The total number of sage grouse observed (6,910) was higher than has ever been reported (6~062 in 1974) during the previous 5 year period (Braun 1979) although the total number of hunters checked (521) was less than the high (738) in 1975. �Table 8. Check station sage grouse harvest data, North Park, Colorado, 1979. Check Station State Line No. of hunters Hours hunted Birds seen Birds bagged Birds Lost Birds banded Birds/ hunter 42 292 485 85 2 5 2.0 Willow Creek 206 1824 2601 430 25 23 2.1 Gould 273 2002 3824 467 21 29 1.7 Totals 521 4118 6910 982 48 57 Avg. N VI \D 1.9 �260 Hunter and Harvest Distribution Sage grouse hunter pressure and harvest as determined from check station data have been consistently low in the northeast quadrat of North Park (Table 9). From 1974 to 1979, an average of only 10% of all hunters surveyed hunted in the northeast quadrat and bagged an average of 13% of the harvest. The northwest quadrat, especially the Lake John area, has traditionally had high hunting pressure as has the southwest quadrat. With the opening of the Arapahoe National Wildlife Refuge to sage grouse hunting in 1978 and 1979, hunting pressure and harvest have increased substantially in the southwest quadrat. Table 9. Sage grouse hunter and harvest distribution, Colorado, 1974-1979.a,b NE % NW % North Park, SE % SW % Year Hunt. Harv. Hunt. Harv. Hunt. Harv. Hunt. Harv. 1974 1975 1976 1977 1978 1979 12 13 14 14 11 9 8 9 15 15 9 38 45 36 37 28 30 18 14 11 35 42 44 46 49 35 8 6 13 13 6 6 8 3 9 35 30 34 36 38 44 35 32 47 40 53 51 10 13 42 36 12 8 36 43 6-year avg. aUnpublished 11 data (C. E. Braun 1979). bData from check stations only. Band Recoveries Recoveries of banded sage grouse during the hunting season reflect relatively light hunting pressure in the northeast quadrat since 1973 (Table 10). During the 7 year period, only 27 of 393 (7%) band recoveries have been in the northeast quadrat. During the 1979 hunting season, 11 of 69 (16%) band recoveries were from the northeast quadrat. This increase was the result of increased numbers of banded birds in the northeast quadrat during 1979 rather than increased hunting pressure. Thirty-three percent of the 1979 banding sample was in the northeast quadrat. Check station hunter survey data (Table 9) actually show a decline in percent of the total harvest in the northeast quadrat in 1979. �261 Table 10. Sage grouse band recoveries, Park, Colorado, 1973-1979. northeast Year recovered 1974 1975 quadrat, in northeast 1976 1977 North quadrat 1978 Year banded Total banded in North Park 1973 1974 1975 1976 1977 1978 1979 288 191 421 379 540 258 357 2 0 0 0 0 3 0 0 0 0 0 0 1 0 3 0 0 0 0 3 4 0 0 0 0 3 2 6 Totals 2434 2 0 3 0 4 7 11 1973 1979 Wing Analysis During the 1979 hunting season, 1081 sage grouse wings were collected in North Park. Examination of age structure (Table 11) shows the highest percentage of immatures ever recorded in the harvest indicating excellent production and survival in 19,79. The percent of yearlings in the harvest was also the highest ever recorded indicating excellent recruitment of 1978 production into the 1979 population. Both of these statistics indicate a healthy, increasing population of sage grouse in North Park. Table 11. Age structure of the sage grouse harvest, Colorado, 1974-1979.a Year Immatures No. 1974 1975 1976 1977 1978 1979 350 212 210 290 385 624 6-year avg. aUnpublished Adults Yearlings % 50.1 42.0 42.3 45.9 53.2 57.7 No. % No. 138 111 117 123 143 256 19.8 22.0 23.5 19.5 19.8 23.7 210 182 170 219 196 201 50.0 data North Park, (C. E. Braun 1979). 21.5 % 30.1 36.0 34.2 34.6 27.1 18.6 28.5 �262 Wings from 283 hens (129 adults, 154 yearlings) harvested in 1979 were classified as to primary molt to estimate nesting success (Table 12). Estimated nesting success was 55.8% for yearlings, 65.1% for adults, and 60.1% overall. This is the highest ever recorded and was reflected by equally high values of percent young in the harvest (57.7%), young/hen (2.2:1), young/successful hen (3.7:1), and the average number of birds/hunter in the harvest (1.9). Table 12. Sage grouse nesting success and production rates, and a hunter success, North Park, Colorado, 1974-1979. Estimated nesting success Adults Yearlings Total Year 1974 1975 1976 1977 1978 1979 64.5 53.2 52.9 59.3 59.7 65.1 aUnpublished 46.1 39.0 26.8 32.7 38.3 55.8 data 58.7 48.6 43.2 50.3 51.4 60.1 % young in harvest 50.1 42.0 42.3 45.9 53.2 57.7 .Young per hen Young per successful hen Birds per hunter 1.4: 1 1.1: 1 1.1: 1 1.2: 1 1.8: 1 2.2:1 2.3:1 2.3:1 2.5:1 2.0:1 3.6:1 3.7:1 1.1 0.7 0.8 1.1 1.4 1.9 (C. E. Braun 1979). LITERATURE CITED Beck, T. D. I. 1975. Attributes of a wintering population of sage grouse, North Park, Colorado. M.S. Thesis. Colo. State Univ. Fort Collins. 49pp. 1977. Sage grouse flock characteristics and habitat selection in winter. J. Wildl. Manage. 41:18-26. Braun, C. E. 1979. Evaluation of the effects of changes in hunting regulations on sage grouse populations. Colorado Div. Wildl. Prog. Rep., Fed. Aid Proj. W-37-R-32, Work Plan 3, Job 9a. pp . 11-35. , and T. D. I. Beck. 1976. Effects of sagebrush control on distribution and abundance of sage grouse. Colorado Div. Wildl. Final Rep., Fed. Aid Proj. W-37-R, Work Plan 3, Job 8a. pp. 21-84. ---- Bray, O. E., and G. W. Conner. 1972. A tail clip for attaching transmitters to birds. J. Wildl. Manage. 36:640-642. �263 Canfield, R. H. 1941. Application of the line interception sampling range vegetation. J. For. 39:388-394. method in Carr, H. D. 1967. Effects of sagebrush spraying on abundance, distribution and movements of sage grouse. M.S. Thesis. Colo. State Univ., Fort Collins. 106pp. Dalke, P. D., D. B. Pyrah, D. C. Stanton, J. E. Crawford, and E. Schlatterer. 1963. Ecology, productivity, and management of sage grouse in Idaho. J. Wildl. Manage. 27:810-841. Emmons, S. R. 1979. Evaluation of the effects of changes in hunting regulations on sage grouse populations: evaluation of censuses of males. Colorado Div. Wildl. Prog. Rep., Fed. Aid Proj. W-37-R-32, Work Plan 3, Job 9b. pp. 37-65. Eng, R. L. ratios 1955. A method for obtaining sage grouse age and sex from wings. J. Wildl. Manage. 19:267-272. 1963. Observations on the breeding grouse. J. Wildl. Manage. 27:841-846. ________ , and P. Schladweiler. and habitat use in central biology of male sage 1972. Sage grouse winter movements Montana. J. Wildl. Manage. 36:141-146. Gill, R. B. 1965. Distribution and abundance of a population of sage grouse in North Park, Colorado. M.S. Thesis. Colo. State Univ., Fort Collins. 185pp. Klebenow, D. A. 1969. Sage grouse nesting Idaho. J. Wildl. Manage. 33:649-662. Patterson, R. L. 1952. Denver. 341pp. The sage grouse and brood in Wyoming. habitat in Sage Books, Inc. Petersen, J. G. 1970. The food habits and summer distribution of juvenile sage grouse in central Montana. J. Wildl. Manage. 34: 147-155. Pyrah, D. B. 1959. Sage grouse population trend and trapping study. Wyoming Game and Fish Comm., Job Compl. Rep., Fed. Aid Proj. W-50-R-8. 27pp. Wallestad, R. o. 1971. Summer movements and habitat use by sage grouse broods in central Montana. J. Wildl. Manage. 35:129-136. ________ , and D. Pyrah. 1974. Movement hens in central Montana. J. Wildl. and nesting of sage grouse Manage. 38:630-633. �264 Wallestad, R. 0., and P. Schladweiler. 1974. Breeding season movements and habitat selection of male sage grouse. J. Wildl. Manage. 30:634-637. Prepared ,---r.~....J. by --- ~ Thomas J. Schoenberg Graduate Research Assistant Approved by ~d",,--,-=-~-,,-·_~_._~~--=CIa it E. Braun Wildlife Researcher C __ 7 �April 265 JOB PROGRESS State of Colorado Project No. W-37-R-33 Job Title: Population REPORT Game Bird Survey 9 Work Plan No. 1980 Job No. 5 Dynamics and Habitat Relationships of Blue Grouse Period Covered: Personnel: April 1, 1979 to March 31, 1980 D. Benson, C. Braun, L. Carpenter, J. Claassen, R. Clippinger, J. Corey, K. Duncan, R. Estes, H. Funk, J. Gerrans, W. Heicher, D. Hoart, D. Hoffman, R. Hoffman, B. Sigler, M. Smith, L. Spess, S. Steinert, L. Strong, and W. Woodward. ABSTRACT Investigations concerning the effects of hunting on blue grouse (Dendragapus obscurus) populations, stability of breeding population levels, and habitat relationships of blue grouse were initiated in 1975 and continued in 1979 on two areas in northwestern Colorado. Data obtained from breeding and production surveys over the past 5 years indicate that the populations under investigation were relatively stable and reproductively healthy. The ecological density of territorial males at Green Mountain and Eiby Creek has ranged from 7.7 to 9.2 ha/male and 9.1 to 10.1 ha/male, respectively. Sex ratio in the breeding population approximated 1:1. Evaluation of 8 nests located since 1975 indicated an average clutch of 6.0 eggs. Egg loss reduced the hatch by 33.3 percent (66.7% hatching success). Overall, 50 percent of the nests hatched successfully, 30 percent were partially successful, 10% were destroyed by predators, and 10% were deserted. Peak of hatch varied by as much as 3 weeks over the 5 years of study, but most chicks hatched within a 25-day period from 18 June to 12 July. Estimated nesting success based on wing analyses was 59.8 percent for Middle Park and 66.4 percent for the Eagle area in 1979; the 5 year average success rate was 68.4 and 68.9 percent, respectively. Adult hens exhibited less variation and an overall higher success than yearlings. Fifty-nine broods were counted in 1979 with a composite average of 4.5 (Green Mountain) and 5.5 chicks per brood (Eiby Creek). Production and mortality estimates suggest there are more chicks produced than necessary to replace natural losses in the breeding population; consequently, there are always surplus birds available in the fall population and in most years this surplus exceeds 40 percent. �266 Only a small portion (4%) of the fall population was removed by hunting compared to what could be safely harvested (23%) without adversely affecting the subsequent spring population. Justification and recommendations for the implementation of a statewide wing collection program as a means of monitoring blue grouse populations are discussed. Examination of 137 crops collected during check station operations resulted in the identification of 45 genera of plants and 5 orders of insects represented in the fall diet of blue grouse. Vegetation maps were prepared for both study areas and characteristics of breeding, nesting and brood areas were described. �267 POPULATION DYNAMICS AND HABITAT RELATIONSHIPS OF BLUE GROUSE Richard W. Hoffman P. N. OBJECTIVE Major objectives of this study are to (1) increase the harvest of blue grouse in Colorado (double present harvest estimates without harm to breeding populations in subsequent years, (2) to identify differences in breeding densities due to differing habitats, and (3) to document the stability of breeding densities over time. SEGMENT OBJECTIVES 1. Estimate breeding densities on the study areas through use of acoustical census and systematic search. 2. Estimate 3. Compile data, analyze results, prepare progress nesting success and production on the study areas. report. METHODS AND MATERIALS Reference is made to Hoffman (1976) for a detailed discussion of methods and materials used in this study. Any deviations or additions to these are discussed in the Results and Discussion section. DESCRIPTION OF STUDY AREAS Research is presently being conducted on t~o selected study areas of differing habitat types in northwestern Colorado. These areas are described in detail in previous reports (Hoffman 1976 and 1977) and are depicted in Figures 1 and 2. Vegetative characteristics are presented in a later section of this report entitled "Habitat Investigations". �268 GREEN MOUNTAIN AREA COLORADO I ( · · l '; \, - STUDY AREA BOUNDARY · .. . ~ "'. .""-. _,' .1· ' . ., " ' . .•... "., \. , ;.\ . "".' '. . ._ "', -", . -. ':--:- •.. ,. ' ,! \ .. . \ . " \ " ... '.- 1. __"._'\,.., : . SCALE IN KILOMETERS o I r---------------~--------------~ f~ Green Mountain " study area . �OJ' N 90~2 (j\ 1'" EBY CREEK AREA COLORADO N '''1'''' ~ - 73 Fig. 2. Eiby Creek study area. STUDY AREA BOUNDARY SCALE IN KILOMETERS I 0 I �270 RESULTS AND DISCUSSION Breeding Timing, Territoriality Survey and Density In all years, grouse were observed on the breeding range at both study areas in early April. First arrivals were mostly males. Few females were present during the first 2-3 weeks of April; thus, display was much curtailed, limited to the predawn hours, and associated more with territorial establishment than sexual responsiveness. The frequency and intensity of displays increased about the third week of April which coincided with an increase in the number of hens observed on the study areas. Activities associated with breeding and density of breeding grouse peak in early May. Timing of breeding events was similar in all years except 1977 when activities were advanced about one week. Display by males was concentrated in the early morning period between 0430 and 0600 MST. Occasional display occurred after dusk, but such display was sporadic and involved fewer birds. Males were effectively finished displaying by late June-early July, although flutter flights were heard into early August. Displays associated with breeding have been described in detail in a previous report (Hoffman 1979). Blue grouse males on the study areas were territorial and returned to the same territory year after year, though the boundary of the defended area changed occasionally. New territories were located in some years, but for the most part, the same territories were occupied each year. However, not all known territory locations were occupied by a male every year. A few yearling males were successful in establishing territories, but most of the yearling males captured and marked during the breeding season were classified as non-territorial birds. Only one adult male captured on Green Mountain was not territorial. All other adult males captured on the study areas were territorial. Territory size was determined for 9 banded males seen at least 5 times each on Green Mountain. Territory size ranged from 1.2 to 1.9 hectares, with a mean of 1.6 hectares. Harju (1974) reported a mean size of 0.7 hectares (range 0.4-1.1 ha) per territory in Wyoming. Bendell and Elliott (1967) found that sooty grQuse occupied territories of 0.4 to 0.8 hectares in a dense population and 1.6 to 2.8 hectares in a sparse population. Martinka (1972) reported a mean territory size of 0.8 hectares (range = 0.4-1.5 ha) for blue grouse in Montana, while Boag (1966) found that blue grouse males in Alberta occupied territories of 0.22 to 0.92 hectares. �271 Total number of territorial males recorded during the peak of breeding activities was used as an index to population size. Hens could not be accurately censused during the breeding season because of (1) greater daily movements, (2) more secretive habits, and (3) lower response rate to recorded calls. Since the study areas under investigation differed in terms of habitat type, ecological density rather than total males counted was used for comparison of densities between study areas. In the estimation of ecological density, any area on the breeding range where grouse were observed was considered habitable. /For Eiby Creek and Green Mountain, the estimates were 291.6 hectares (60.5% of total area) and 146.7 hecatres (80.9% of total area), respectively. Table 1 presents the change in number of territorial males and ecological density of breeding males over a 5-year period at Green Mountain and 4-year period at Eiby Creek. Breeding surveys were not conducted at Eiby Creek until 1976. Both populations have been relatively stable with gradual increases. A portion of the increase can be attributed to improved observer efficiency. Ecological density of males appeared to be slightly higher on Green Mountain than at Eiby Creek, but the difference was non-significant (P > .05). Table 1. Study Numbers and ecological density of territorial male blue grouse at Green Mountain and Eiby Creek, 1975-79. Size No. Territorial Males (ha) 1975 1976 1977 1978 1979 Ecological Density (ha/bird) 1975 1976 1977 1978 1979 Green 181.3 Mtn. 16 16 17 19 19 9.2 Eiby 482.0 Creek ND 29 29 30 32 ND 9.2 8.6 7.7 7.7 10.1 10.1 9.7 9.1 Number of hens on the breeding range could not be accurately ascertained from the breeding survey due to factors previously described. It was possible to estimate the number of hens on the study areas just after most of the broods had hatched. At this time, hens were very responsive to a tape recorded chick distress call and were easily located. These data are presented in Table 2 along with the total population estimate. The data are given from 1976 to 1979 for Green Mountain only. Due to the smaller size and isolated nature of the Green Mountain from surrounding breeding populations, the area was easier to search and the hens observed were probably resident birds. Data from 1975 are excluded because less time was spent searching for broods than in other years. �272 Population 1976-1979. Table 2. estimate of blue grouse on Green Mountain, Males Nonterr Total Year Succ Females Unsucc 1976 13 4 17 16 3 19 1.1: 1 36 1977 17 4 21 17 3 20 1.1: 1 41 1978 15 5 20 19 4 23 1.2: 1 43 1979 9 7 16 19 4 23 1.4: 1 39 Total Terr Sex Ratio Males:Female Total Population Some hens and non-territorial males undoubtedly escaped detection in all years, therefore, the total population estimate was minimal. The sex ratio indicated by these estimates did not deviate significantly from 1:1 (P > .05). Data collected in other studies similarly suggest a 1:1 sex ratio in the spring population (Zwickel 1972, Bendell et al. 1972) . Nesting Parameters, Hatching Dates and Production Natality Few nests were located during the course of study due to the difficulty in finding nesting hens and because no concerted effort was made to conduct nest searches on either study area. All nests were found while performing other duties. A total of 12 nests were located from 1975 to 1979. Site and floristic characteristics of all nests sites were recorded and are discussed under "Habitat Investigations". Clutch size and hatching success was determined for 10 nests (Table 3). The other 2 nests were found about 5-6 weeks after the peak hatching period and there were few remaining egg shells to determine the clutch size or fate of these nests. Sample size was too small for comparisons between years or age classes. Table 3. No. Nests 10 II Natality rate among blue grouse based on 10 nests examined from 1975 to 1979. Clutch size11 Mean Range 6.0 4-8 Total eggs laid Unhatched Depredated Deserted 6 10 2 54 - Based on 8 nests for which Hatching success (%) Hatched 36 66.7 total clutch size was ascertained. �273 Egg loss, including one nest that was deserted before laying was completed, reduced the hatch by 33.3 percent. Boag (1966) reported an approximate 18 percent egg loss of Alberta blue grouse, whereas Zwickel (1975), reported what he considered a conservative estimate of 54 percent hatching success or a maximum estimate of 46 percent egg loss. Overall, 50 percent (5) of the nests examined during this study hatched successfully, 30 percent (3) were partially successful, 10 percent (1) were destroyed by predators and 10 percent (1) were deserted. Hatching Dates Figure 3 presents the hatching curves for blue grouse Middle Park areas. Time of hatch was similar between data were grouped. All chicks were aged according to lined by Zwickel and Lance (1966). The estimated age for bias as described by Redfield and Zwickel (1976). in the Eagle and areas, so the procedures outwas then adjusted Peak of hatch varied by as much as 3 weeks over the 5 years of study, but most chicks hatched within a 25 day period from 18 June to 12 July. Examination of the hatching curves show that no second peak, indicative of renesting, was evident in any year. This does not imply renesting did not occur, but the contribution of renesting to production of young was insignificant. Nesting Success Since hens with broods are easier to locate than those without, this factor tends to produce an inflated estimate of nesting success when calculated from field observations. Consequently, analyses of the wing molt pattern of hunter harvested birds was used as the best estimate of nesting success (Braun 1971). The technique provides a minimum estimate as some successful hens will have completed their molt by mid- to late September and cannot be distinguished from unsuccessful hens. This problem is compounded during years when hens initiate nesting earlier such as in 1977. Few yearlings were identifiable in 1977 and many adult hens were in the advance stages of molt. Therefore, nesting success was probably better in 1977 than what the data indicate. Overall, 59.8 percent of the adult and yearling female wings collected in Middle Park in 1979 were classified as from successful nesters. This represents the lowest estimate obtained from wing samples since 1975 (Table 4) and was attributed to (1) a late nesting season, and (2) adverse weather conditions during the nesting period. However, even under these less than optimal conditions, nesting success was still good. Based on field observations of successful (9) and unsuccessful (7) hens in 1979, estimated nesting success on Green Mountain was 56.2 percent. �274 70 1975 1976 1977 1978 1979 60 (!) • • .-----. 0 0 0-----0 K ~ 50 ~ ::r: u ~ ::x: 40 • t--:- 1\ z w u / / a:: W c, 30 / I I I / 20 / / ,• / "<, / / 10 / "" " •I .,, , / I 1-7 8-14 15-21 22-28 ,,'e__ 29-5 0 . ......•. 6-12 ......•. .......;: -- 13-19 20-26 JULY JUNE WEEK OF HATCHING Fig. 3. Blue grouse hatching curves, Middle Park and Eagle, 1975-79. �275 Table 4. Year Estimated nesting success, Middle Park and Eagle, Middle Park Estimated Nesting Success Adults Year(N)l/ Total lings2/ Estimated Adults 1975-1979. Eagle Nest inf!:_ Success Y~ar- 2/ Total Lf.ng s - 1975 80.3 (71) 33.3 (21) 69.6 (92) ND~_/ ND ND 1976 68.3 (60) 82.1 (28) 72.7 (88) ND ND ND 1977 68.5(143) 68.7 (16) 68.6(159) 57.1 (63) IS!!__!(4) 57.1 (63) 1978 80.9(115) 65.2 (46) 76.4(161) 81.0 (58) 80.8 (26) 80.9 1979 66.7(150) 42.4 (59) 59.8(209) 67.2 (64) 65.1 (43) 66.4(107) 56.5(170) 68.4(709) 68.1(185) 71.0 (69) 68.9(254) Total 72.2(539) l/Total sample of wings examined. . . some years, estimated -2/Due to sma 11 samp 1e Slzes 1n not be representative of this age class. nesting (84) success may 3/ - No data. 4/ - Inadequate sample. With the exception of 1979, combined nesting success of adult and yearling hens was relatively constant from year to year (Table 4). Nesting success varied more within and between age classes than noted for combined success of both age classes. Adult hens exhibited less variation (66.7-80.3%) and an overall higher success (X=72.2%) than yearlings (X=56.5%, range = 33.3-82.1%). Reasons for this difference are uncertain, but may be an artifact of sample size. Zwickel (1975) detected no significant differen~e in nesting success of yearling and adult hens. Few wings were collected from the Eagle area in 1975 and 1976. In 1977, only 4 yearlings were identifiable from the wing sample so nesting success was based solely on the sample of adult hens. Wings from 107 females (adults=64; yearlings=43) were classified as to primary molt in 1979. Examination of the wing molt pattern revealed 43 adult hens (67.2%) and 28 yearling hens (65.1%) were probably successful nesters in 1979. Combined success for both age classes was estimated at 66.4 percent. Estimated nesting success on the Eiby Creek study area as ascertained from field observations in 1979 was 70.6 percent. Adult Turnover Estimated turnover (from wing analysis) of adult male and female blue grouse in Middle Park was 38.5 and 28.1 percent, respectively in 1979. The estimates are based on the assumption that in a stable population, percent yearlings must equal annual loss of adults. While these data suggest differential survival favoring adult females examination of the 4 year average annual turnover rate (adult males=28.3%; adult females=28.5%) indicates similar survival of males and females. �276 Yearly estimates varied more for males than females because of the compounded problem of separating yearling and adult males. Most likely adult males have a constant annual turnover rate as shown by females in Table 5. Overall the estimated annual turnover rates (4-year averages) are probably lower than the actual turnover rates, again, due to the fact that some yearlings were undoubtedly assigned to the adult age class. Sample sizes were too small for similar calculations using the data collected from the Eagle area. Estimated turnover of adult male and female blue grouse, Middle Park. Table 5. 1/ Year- Adults No. % Males Yearlings No. % Total Adults No. % Females Yearlings No. % Total 1975 30 85.7 5 14.3 35 41 73.2 15 26.8 56 1976 68 81.9 15 18.1 83 64 69.6 28 30.4 92 1978 121 77 .1 36 22.9 157 116 71.6 46 28.4 162 1979 131 61.5 82 38.5 213 151 71.9 59 28.1 210 AVERAGES 71.7 28.3 28.5 71.5 1/ - Data from 1977 excluded because of the unrepresentative sample of yearlings. Production and Juvenile Mortality Brood survey, conducted from mid-June to mid-August, resulted in complete counts of 231 broods (Green Mountain = 115; Eiby Creek = 116) from 1975 to 1979 (Table 6). Broods were counted more than once, but not within the same month. Yearly average brood sizes recorded were as follows: Green Mountain Eiby Creek 1975 1976 1977 1978 1979 1975 1976 1977 1978 1979 3.9 4.4 3.8 4.9 4.5 No data 3.4 3.9 5.1 5.5 �277 Table 6. Average brood size, range, and number of broods observed monthly intervals, 1975 to 1979.l/ Month 1975 Green Mountain 1976 1977 1978 1979 1976 Eiby Creek 1977 1978 by 1979 June Mean Nrl:-/ 5.3 5.1 6.2 7.2 5.0 4.6 6.5 6.0 Range ND 3-7 4-7 5-9 6-9 4-6 3-6 6-7 3-9 Sample Size ND 3 11 9 5 2 7 2 3 July Mean 5.2 4.1 4.0 5.4 4.8 3.7 4.1 4.9 6.5 Range 2-7 2-6 1-7 3-7 1-10 1-7 2-9 3-8 4-10 4 7 10 9 12 9 10 13 15 Mean 3.0 3.0 2.5 3.7 3.0 2.2 3.6 5.0 4.3 Range 1-5 1-4 2-6 1-6 1-4 1-10 3-8 1-7 13 14 11 5 20 17 13 Sample Size August Sample Size l/only 2/ distinct 6 1 broods with full counts are included. - No data, late hatching year, no full counts obtained. Average brood size was similar between areas in 1977 and 1978. The apparently smaller brood size at Eiby Creek in 1976 was attributed to a larger sample of broods counted in late July - early August, whereas primarily early to mid-July counts were made on Green Mountain. All evidence points to better production and survival of young at Eiby Creek in 1979 than occurred at Green Mountain. Rogers (1968) found average brood sizes of 4.1, 2.6 and 3.9 in 3 years of study in western Colorado. Since some doubt exists as to whether blue grouse broods can be accurately counted until after they are 4 weeks old (Boag 1966), late June and early to mid-July counts may be underestimated. Brood counts in August may also be misleading due to shuffling of chicks between broods, formation of gang broods, and brood breakup and dispersal. Therefore, obser'led differences in brood size from June to August may not accurately reflect loss of chicks during this period (Table 7). The gradual attrition in size of broods over the summer months is the least reliable method of evaluating chick mortality (Zwickel and Bendell 1967), but was the only data available for determining losses in this study. Average annual summer mortality of chicks was 47.4 percent for Green Mountain and only 21.2 percent for Eiby Creek. �278 Table 7. Mortality of juvenile blue grouse from late June to mid-August. Mean Brood Size June August Area Year Green Mountain 1975 5.2]) 3.0 42.3 1976 5.3 3.0 43.4 1977 5.1 2.5 51.0 1978 6.2 3.7 40.3 1979 7.2 3.0 58.3 1975-79 5.9 3.1 47.4 1976 5.0 2.2 56.0 1977 4.6 3.6 21.7 1978 6.5 5.0 23.1 1979 6.0 4.3 28.3 1976-79 5.2 4.1 21.2 Eiby Creek Mortality (%) l/Mean brood size for July as no full counts were obtained in June due to late hatch. Production and Breeding Populations Estimated total production in relation to breeding population levels is presented in Table 8 for both study areas. Calculations are based on the following assumptions which are supported by data collected in this and other studies: (1) A 1:1 sex ratio exists in the breeding population (2) Estimated percent nesting success is correct (3) Mean annual brood size and monthly estimated brood sizes represent the production of chicks on the study areas (4) Immigration equals emigration �Table 8. Area Estimation Year of the annual production Total..!_! Nesting Success Breeding Population (%) of young on Green Mountain No. Hens With Broods Average Annual Brood Size Total Production Mortality JuneAuzu s t and Eiby Creek. Total Production by midAugust Total Population by midAugust Percent Gain Green Mountain 1975 32 69.6 11 3.9 43 42.3 25 57 44 1976 32 72.7 12 4.4 53 43.4 30 62 48 1977 34 68.6 12 3.8 46 51.0 22 56 39 N -...J \0 1978 38 76.4 14 4.9 69 40.3 41 79 52 1979 38 59.8 11 4.5 49 58.3 21 59 36 1977 58 57.1 17 3.9 66 56.0 29 87 33 1978 60 80.9 24 5.1 122 23.1 94 154 61 1979 64 66.4 21 5.5 ll5 28.3 83 147 56 Eiby Creek l/Excludes non-territorial males. �280 Zwickel (1965) and Bendell and Elliott (1967) reported a mean annual death rate of approximately 30 percent for breeding populations of blue grouse on Vancouver Island, British Columbia. A similar estimate of 39 percent was obtained from reobservation of marked birds at Green Mountain and Eiby Creek. Assuming this estimate is constant from year to year, one can determine from the estimate of total production the minimum replacement requirements necessary to maintain a stable breeding population (Table 9). Production by mid-August rather than total production was used in the computations because mid-August figures more closely approximates production and survival of juveniles until fall. The replacement requirement then becomes an estimate of the fall to spring mortality of juveniles that should occur in a stable population. Both populations under investigation were essentially stable. It is apparent from the data presented in Table 9 that production was more than adequate to replace natural losses in the breeding population. For Green Mountain, the average annual fall to spring loss of juveniles required to maintain the population from 1975 to 1979 was 52 percent (65% for Eiby Creek from 1977 to 1979). These data suggest there are always surplus birds in the fall population and in most years this surplus exceeds 40 percent. Table 9. Area Estimates of the fall to spring loss of juveniles to maintain a stable breeding population. Year Total Breeding Population Annual Mortality Breeding 1/ Population- Total Production by mid-August necessary Fall to Spring Mortality of Juveni1es~/ (%) Green Mountain 1975 32 12 25 52 1976 32 12 30 60 1977 34 13 22 41 1978 38 15 41 63 1979 038 15 21 29 1977 58 23 29 21 1978 60 23 94 75 1979 64 25 83 70 Eiby Creek l/Number of birds expected to be lost from the population constant annual mortality rate of 39 percent. -2/ Morta 1·lty necessary . . to malntaln vooou La tI thee popu atl0n. assuming a �281 Harvest Hunting Season The 1979 blue grouse season in Middle Park opened on 8 September and closed on 7 October. Blue grouse hunting was also permitted between 13 October and 13 November in conjunction with the deer, elk, and combined deer-elk seasons. Season length was 57 days with daily bag and possession limits of 3 and 6 birds, respectively. Season structure, bag limits and length from 1975 to 1979 are presented in Table 10. Table 10. Blue grouse hunting seasons, Colorado, 1975-1979. Year Hunting season dates Season length (days) Bag Limit Possession limit 1975 9/13-10/5 23 3 6 Unit 80 and all units west of interstate 25 except portions of Unit 52 1976 9/11-10/10 30 3 6 Same as 1975 9/11-10/10 40 3 6 Unit 28 (Middle Park) only 9/10-10/09 30 3 6 West of interstate 25 and Unit 80 10/15-11/15 27 3 6 ~est of interstate 25 and Unit 80 when open to deer or elk hunting 9/09-10/08 30 3 6 Same of 1977 10/14-11/14 27 3 6 Same as 1977 9/08-10/07 30 3 6 Same as 1977 10/l3-11/l3 27 3 6 Same as 1977 and Open areas 10/16-10/26 1977 1978 1979 Middle Park For the third consecutive year a check station was operated by regional and research personnel at the Prairie Point Campground along Highway 9 at the south end of Green Mountain Reservoir. The station was operated opening weeken.j from about 1000 to 1800 MST, depending upon traffic load. All vehicles were stopped on the highway, but only hunters were directed to pull off at the check station. Data obtained per party �282 included: county of or1g1n, number of hunters, hours hunted, birds observed, birds bagged per hunter, area hunted, birds shot but not retrieved, and location where each bird was harvested. One wing was removed from each bird provided no wings were deposited in barrels. Whenever a bird was missing one or both wings, the hunter was questioned as to what he did with the wings. Thus, the status of all wings was recorded as follows: (1) hunter disposed of wings, (2) hunter deposited wing in barrel, and (3) wing collected at check station. Sex by gonadal inspection was ascertained for immatures whenever possible. Ovaries were collected from adult and yearling hens when present. Whole body weights were obtained for all grouse that were not eviserated. In addition to were available tions in Middle in the field. the check to hunters Park. A Some wings station, 14 volunteer wing collection stations throughout the entire season at various locafew hunters were contacted opportunistically were also obtained from the mail survey. A total of 946 wings were collected from Middle Park in 1979. Wing barrels accounted for 88.7 percent (839 wings) of all wings collected (Table 11). The remaining wings were collected at check station (84 = 8.9%) and from the mail wing survey (23 = 2.4%). Comparative data pertaining to the number and source of blue grouse wings collected in Middle Park from 1975 to 1979 is given in Tables 11 and 12. Table 11. Number and source of blue grouse wings collected Middle Park from 1975 to 1979. Year Wing barrels No. % 1975 120 (10)11 65.6 26 14.2 0 1976 292(13) 84.9 49 14.2 3 .9 1977 501 (15) 82.7 87 14.4 18 2.9 1978 875(16) 87.0 105 10.4 0 1979 839(14) 88.7 84 8.9 23 85.2 351 11.4 44 2627 Check stations No. % . parentheses -1/N um b er 1n represents Mail surve% No. 0 in Miscellaneous No. % Total 20.2 183 0 0 344 0 0 606 37 26 2.6 1,006 2.4 0 0.0 946 1.4 63 2.0 3,085 0 number of wing barrels used. �283 Table 12. Number of wings collected from wing barrels, Middle Park, 1975-1979. No. of Wings Collected Station Location 1975 1976 1977 City Reservoir Beaver Creek Williams Fork Corral Creek Troublesome Pinto Creek Lawson Ridge Spring Creek Blue Ridge Chimney Rock Gore Pass Trough Road Kremmling Rock Creek Willow Creek Cottonwood Pass Ute Pass 8 1 0 21 13 9 0 30 38 3 19 1 43 2 0 3 55 23 95 32 17 8 0 53 18 5 2 62 63 85 69 74 14 5 NB NB NB NB NB NB NB NB NB.!/ 1979 Total NB NB NB NB 26 57 16 19 62 8 20 28 60 24 257 66 37 11 477 237 513 412 233 47 5 119 24 77 501 875 839 2627 NB 4 120 TOTAL 1978 NB NB NB NB NB 12 292 28 10 80 16 23 3 169 50 136 154 85 29 4 13 60 17 0 3 161 63 197 157 74 1/ - No barrel at this location. During the 2 days of check station operations, 255 hunters with 342 blue grouse (1.3 birds per hunter) were contacted. Total birds reported observed by these hunters was 830. Some duplications are undoubtedly present in the observations. Of the 255 hunters checked, 53.3 percent were successful. Only 19 birds were reported wounded and not retrieved for an estimated crippling loss of 5.6 percent. Hunter efficiency (birds bagged .;birds observed x 100) was calculated at 41.2 percent. Comparative harvest statistics for all years is presented in Table 13. Table 13. Blue grouse harvest 1975-1979 ..!/ Year No. hunters checked No. birds observed No. birds bagged 1975 1976 1977 1978 1979 112 138 228 222 255 158 141 480 758 830 45 61 226 294 342 1/ statistics, Hunter efficiency % 28.4 43.2 47.1 38.8 41.2 Middle Park, Colorado, % successful hunters 30.4 33.4 39.5 54.6 53.3 Crippling loss % Birds per hunter 4.4 8.2 4.0 3.1 5.6 0.4 0.4 1.0 1.3 1.3 - Check stations operated on the Williams Peak road in 1975 and 1976 and at the Prairie Point campground from 1977 to 1979. �284 In order to evaluate the effectiveness of wing barrels for sampling, Middle Park hunters were questioned as to what they did with the wings from the birds they harvested. This information is summarized below with similar data from 1977 and 1978: ......... deposited .................... disposed ..................... of unknown status ............ % wings deposited % wings collected been % wings % wings in barrels that should have 1977 1978 1979 60.7 50.6 52.6 18.9 17.4 3.0 20.3 28.3 0.8 22.8 16.4 8.2 As in other years, distribution of harvest and hunting pressure in 1979 was not uniform within Unit 28 (Table 14). Chimney Rock was the leading harvest area in 1979 accounting for 23.5 percent of the harvest, but only 12.9 percent of the hunters. Spring Creek ranked second in terms of harvest with 19.2 percent of the kill and first in terms of pressure with 24.9 percent of the hunters. Gore Pass was next with 18.7 percent of the harvest and 23.6 percent of the hunters. Blue Ridge and Piney attracted a lot of hunters (15.3% and 11.6%, respectively), yet only accounted for 7.5 and 8.8 percent of the harvest. Corral Creek and Willow Creek had 7.1 and 7.4 percent of the harvest. All other areas in Middle Park each had less than 3 percent of the harvest and hunter pressure. Table 14. Blue grouse harvest Park, 1977-1979.11 and hunting 1977 pressure within Middle 1978 1979 % % % % % % hunters harvest hunters harvest hunters harvest 33.0 12.2 30.7 18.8 24.9 19.2 8.1 16.0 6.4 14.0 12.9 23.5 Gore Pass 14.8 14.1 19.3 17.2 23.6 18.7 Blue Ridge 23.4 12.7 16.5 7.4 15.3 7.5 Piney 11.5 15.5 15.2 9.7 11.6 8.8 9.1 4.1 8.0 2.1 7.1 Area Spring Creek Chimney Rock Corral Creek Willow Creek Ute Pass 5.7 1.9 7.4 6.9 1lTotals will not add up to 100 percent as only major hunting and harvest areas are included. All other areas in Middle Park each had less than 3 percent of the hunters and harvest. �285 Most hunters contacted in 1979 originated from the Denver metropolitan area. The counties of Jefferson (71), Denver (44), Adams (42), and Arapahoe (37) accounted for 194 of 252 (77.0%) hunters for which origin was ascertained. Local hunters from Grand and Summit counties made up 5.6 percent of the hunters contacted. Undoubtedly this figure is underestimated as many local residents are not contacted at the check station. All other counties each comprised less than 5 percent of the hunter contacts. Similar data have been collected in previous years. Distribution of wing collections according to time period is given in Table 15. Liberalization of the season in 1977 and 1978 tended to spread the harvest and hunting pressure over a longer period of time. This effect was not expressed by the data collected in 1979. Check station figures for opening weekend suggest that more blue grouse hunters were attracted to Middle Park in 1979. Consequently, more birds were shot opening weekend compared to previous years. Perhaps Middle Park has gained popularity in terms of blue grouse hunting due to additional publicity received as the result of intense research efforts in this area. However, no data are available to support this hypothesis. Table 15. Time distribution of blue grouse wings collected, Colorado, 1976-1979.l/ 1975 1976 Middle Park, 1979 1978 1977 No. % No. % No. 188 20 63 28 44 7 18 11 12 37.5 4.0 12.6 5.5 8.8 1.4 3.6 2.2 2.4 288 99 109 13 55 13 23 10 119 32.9 11.3 12.5 1.5 6.3 1.5 2.6 1.1 13.6 375 30 65 81 48 48 18 35 39 44.7 3.6 7.7 9.7 5.7 5.7 2.2 4.2 4.6 Deer season 37 7.4 67 7.7 54 6.4 Elk season 63 12.6 44 5.0 25 3.0 Combined season -- 10 2.0 35 4.0 21 2.5 96.2 391 78.0 729 83.3 739 88.1 3.8 110 16.7 100 11. 9 100.0 501 100.0 839 100.0 Time Period No. weekend week weekend week weekend week weekend week 53 9 20 0 8 11 19 1st 1st 2nd 2nd 3rd 3rd 4th 4th 5th % 44.2 7.4 16.7 0.0 6.7 9.2 15.8 weakend Experimental season (1976) 10/16 - 10/26 Total-regular season Total-Big season Total-Both seasons 120 100.0 game No. % 121 36 36 11 23 10 21 8 15 41.4 12.3 12.3 3.8 7.9 3.4 7.2 2.7 5.2 11 3.8 281 nY 120 100.0 292 l/Wing collections from barrels l/Experimental season only. only. 22.0 1Lf6 100.0 875 % �286 Of 946 wings collected from Middle Park in 1979, 920 were classified to age and sex (Table 16). Immatures (54.0%) and yearlings (15.3%) were well represented in the sample indicating that both production of young in 1979 and survival of young from 1978 to 1979 were good. Of importance is the fact that neither production nor survival was impaired by the severe winter of 1978-79. There were proportionally more yearlings in 1979 than recorded in previous years. Yearlings were probably under-represented in the 1975 through 1978 samples, not because of poor production and subsequent low recruitment, but because fewer yearlings could be separated from the adult age class. The problem is associated with the onset of primary molt and whether the key feathers (primaries 9 or 10) for distinguishing age classes are still present on the wing at the time of collection. If these feathers have already molted, as was the case with many wings examined from 1975 to 1978, then yearlings cannot be separated from adults and are assigned to the adult age class. This problem is most pronounced in "early" years (see 1977 data -- Table 16), and moreso for males than females because males initiate their primary molt before females. Sex ratio of adults and immatures approximated 1:1 in all years except 1978 when there were significantly more immature males than females. Reasons for the deviation in sex ratio are uncertain, but may be related to a greater harvest of male chicks late in the 1978 season. Yearlings had a balanced sex ratio in only 2 (1978 and 1979) of 5 years of data collection. From 1975 to 1977, there were significantly fewer males than females comprising the yearling segment of the harvest. This exemplifies the fact that more yearling males than females were incorrectly assigned to the adult age class. The actual sex ratio of yearlings is probably 1:1. Eagle A total of 530 blue grouse wings was collected from the Eagle area during the 1979 season. Ten wing collection stations accounted for 97.9 percent (519 wings) of the wings obtained. Few wings were collected from other sources in 1979 (Middle Park check station = 6 wings; mail survey = 5 wings). Number of wings collected from each barrel over the past 3 years is shown in Table 17. Note that some barrels were eliminated and relocated to more productive sites. As a result, number of wings collected from wing barrels increased 42.7 percent from 1977 (297) to 1979 (519). �Table 16. Year Age and sex composition Males % No. Adults Females % No. of the blue grouse harvest, Total No. % Males % No. Yearlings Females No. % Middle Park, Colorado, Total % No. Hales % No. 1975-1979. Immatures Females % No. Total % No. 1975 30 17.7 41 24.1 71 41.8 5 2.9 15 8.8 20 11.7 36 21.2 43 25.3 79 46.5 1976 68 19.9 64 18.8 132 38.7 15 4.4 28 8.2 43 12.6 74 21.7 92 27.0 166 48.7 1977 l33 22.1 144 23.9 277 45.9 3 0.5 16 2.6 19 3.2 164 27.2 143 23.7 307 50.9 1978 121 12.5 116 11.9 237 24.4 36 3.7 46 4.7 82 8.4 364 37.4 290 29.8 654 67.2 1979 131 14.2 151 16.4 282 30.6 82 8.9 59 6.4 141 15.3 242 26.3 255 27.7 497 54.0 N CX) " �288 Table 17. Number of wings collected from wing barrels, Eagle area, 1977-1979. No. Wings Collected Station Location Squaw Creek Lake Creek2/ Muddy PassMilk Creek Coffee Pot Springs Red Sandstone Eiby Creek So. Eiby Creek No. Brush Creek Cabin Creek Gypsum Creek Wolcott TOTAL 1./ No 1977 1978 1979 Total NlJ/ NB NB NB NB 68 73 67 63 26 78 28 5 8 38 83 78 69 52 14 59 30 8 44 82 5 5 214 174 215 172 59 l37 85 13 71 120 297 454 519 1270 5 5 63 23 79 57 19 NB 27 NB 19 barrel at this location. l/same as Red and White Mountain Road. Data presented in Table 18 suggest that liberalization of the blue grouse season in 1977 reduced the "opening weekend syndrome" which generally accompanies any type of conservative season. Whereas hunting pressure probably remained the same, more hunters took advantage of the longer season by hunting later in the season. This effect was evident from 1977 through 1979. Prior to 1977, over 60 percent of all wings collected were obtained opening weekend. Further examination of Table 18 indicates that harvest of blue grouse during the big game seasons has remained relatively stable since 1977. Most grouse were shot during the deer season with a substantial decline in harvest during the elk and combined seasons. This reduction in kill can be attributed to a cha~ge in the birds habits. By late October early November most birds have departed frqm the lower, more open habitat types and moved to higher, more inaccessible coniferous forest types. At the same time; they revert from primarily ground dwelling habits to aboreal habit;s; consequently, they become harder to find. All but 10 of 530 blue grouse wings collected in 1979 were identified to sex and age. This information is presented in Table 19 along with comparative data from 1977 and 1978. These data indicate that both production of young in 1979 and survival of young from 1978 to 1979 was good. Again there was no evidence that production or survival was impaired by the severe winter of 1978-79. As in Middle Park, more yearlings were present .in the Eagle sample in 1979 than either 1978 or 1977. This was attributed to a later onset of primary molt in 1979; therefore,more year.li.ngs coul9 still be distinguished from adults during the fall hunting season. �289 Table 18. Time distribution of blue grouse wings collected, Colorado, 1977-1979. 1977 No. of % of Wings Total Time Period 1st weekend 1st week 2nd weekend 2nd week 3rd weekend 3rd week 4th weekend 4th week 5th weekend Deer Season Elk Season Combined Season Total-Regular Total-Big Total-Both Season Game Season Seasons 1978 No. of % of Total Wings Eagle area, 1979 No. of % of Total Wings 127 48 21 27 10 10 10 5 11 39 15 19 37.1 14.0 6.2 7.9 2.9 2.9 2.9 1.5 3.2 11.4 4.4 5.6 126 62 54 9 17 18 29 14 44 62 27 3 27.1 13.3 11. 6 1.9 3.7 3.9 6.2 3.0 9.5 13.3 5.8 .7 173 44 37 67 16 16 17 30 3 73 37 6 33.3 8.5 7.1 12.9 3.1 3.1 3.3 5.8 .6 14.1 7.1 1.1 269 78.6 373 80.2 403 77 .6 73 21.4 92 19.8 116 22.4 342 100.0 465 100.0 519 100.0 Sex ratio of adults and immatures approximated 1:1 in all years. Yearlings exhibited an unbalanced ratio in favor of females in 1978, but the ratio was not significantly different from 1:1 in 1977 and 1979. Most likely the distorted ratio in 1978 was an artifact of sample size. Utilization Direct evidence from banding data and indirect evidence based on the attributes of the populations under investigation indicate that present levels of harvest have no measurable impact on grouse populations. Available data suggest that even though there is a high and variable loss of grouse during their first year of life, production of young still greatly exceeds the number necessary to replace natural losses in the breeding population. These excess birds represent a harvestable resource that can be removed without adversely affecting the subsequent spring breeding population. Few birds were banded, but the percentage of banded birds shot (4.0%) indicate hunters removed a negligible portion of the fall population (Table 20). Hens (probably successful hens with chicks) and chicks sustained the bulk of the kill (4.2%) followed by males (3.3%). Bendell and Elliott (1967) reported approximately 5 percent of the hens and chicks banded on their study areas on Vancouver Island were shot each year, while very few (0.7%) adult males were taken. Mussehl (1960) found that hunters removed 7 (1957) and 12 (1958) percent of the grouse banded in the Bridger Mountains, Montana, most of which were juveniles. �Table 19. Year Age and sex composition of the blue grouse harvest, Eagle area, 1977 to 1979. Males No. % Adults Females No. % Total No. % Males No. % Yearlings Females No. % Total No. % Males No. % Innnatures Females Total No. % No. % Total Wings 1977 87 25.8 65 19.3 152 45.1 10 3.0 4 1.2 14 4.2 81 24.0 90 26.7 171 50.7 337 1978 62 13.4 59 12.8 121 26.2 9 2.0 26 5.6 35 7.6 153 33.1 153 33.1 306 66.2 462 1979 75 14.4 64 12.3 139 26.7 38 7.3 43 8.3 81 15.6 153 29.4 147 28.3 300 57.7 520 N \0 o �291 Number of banded grouse available Middle Park and Eagle, 1976-79. Table 20. Year Males No. % Shot Avail. Females No. % Shot Avail. to and shot by hunters, Juveniles No. % Avail. Shot Total No. % Shot Avail. 1976 6 0.0 13 0.0 5 0.0 24 0.0 1977 22 0.0 17 0.0 14 20.8 53 5.7 1978 15 6.7 33 3.0 18 5.5 66 4.5 1979 17 5.9 32 9.4 58 0.0 107 3.7 Totals 60 3.3 95 Lf.2 95 4.2 250 4.0 Hickey (1955) states that gallinaceous birds can safely withstand a hunting kill equivalent to about one-half their annual mortality rate. Based on a 5-year average for Green Mountain (Table 8), production contributes to about a 45 percent increase in the population. In a stable population there must be an annual loss from fall to fall of this amount. Therefore, according to Hickey (1955), the population can absorb a harvest of 22.5 percent. This represents a minimum estimate because the annual mortality of adults is not taken into account. Yet, the calculated yield (22.5%) still greatly surpasses the estimated harvest rate (4.0%) and further substantiates the conclusion that hunting has no detrimental effects on blue grouse populations. These data provided managers with the justification to liberalize the blue grouse season statewide in 1977 (see Table 10). Season length was increased from 30 to 57 days by allowing blue grouse hunting in conjunction with the deer, elk and combined deer-elk seasons. A survey was conducted in 1977 and 1978 to determine hunter participation and response to the longer grouse season. The results of the surveys were presented in a previous report (Hoffman 1979). Application of Wing Collection Program One of the expected results of this study was to develop a means of systematically and routinely conducting breeding and production surveys in order to monitor population trends. While the means of conducting the surveys have been developed, their practicality and usefulness as a management tool are questionable. To obtain quality data, considerable time, manpower, expense, and training of personnel would be necessary and the surveys would not provide important data on age and sex structure of the population. Furthermore, there appears to be no direct relationship between breeding and production counts and harvest success because of other factors involved in blue grouse hunting such as accessibility, time of migration, food supplies, hunter pressure and interest, and extensive occupi.ed range of the species. �292 Therefore, survey. evaluation of the harvest would necessitate a separate Some of the deficiencies inherent in conducting breeding and production surveys can be partially overcome by collecting wings from hunter-harvested birds. From analysis of these wings, information can be acquired concerning sex and age composition of the harvest, nesting success of breeding females, hatching dates, and production and survival rates. An ongoing wing collection program can also be useful in monitoring population trends which is about the only value of breeding and production surveys. The primary drawback is that harvest data may not exactly reflect the characteristics and structure of the population from which they were collected; however, the data become more meaningful when used for comparisons among years, areas, and species. Traditional methods of collecting harvest data include manned check stations and mail wing surveys. For species such as the blue grouse, with low harvests, hunter densities, and interest, these methods have not been economically feasible for obtaining sufficient samples of wings to satisfy management requirements. Volunteer wing collection stations have been developed and successfully used in this study to inexpensively and efficiently increase samples of wings collected. From 1975 to 1979, 3,085 wings from blue grouse were collected in Middle Park, of which 2,627 (85.1%) were obtained from wing barrels. In 1979 alone, wing barrels accounted for ~,239 (87.5%) of 2,558 blue grouse wings collected primarily in northwestern Colorado. As a result of this success, it is believed that a statewide wing survey using volunteer wing collection stations is the most feasible and useful method of obtaining data on blue grouse (and possibly other species) for management purposes. Until 1979, the development, testing, and application of this technique has been primarily a research function. Before the program can be turned over to management, the research findings must be integrated into the system to insure that the program is properly designed and implemented. The first step would be to incorporate the information already available into one or several publications which would include (1) results of wing surveys conducted by research personnel, (2) procedures for conducting the survey, (3) techniques for classifying wings to age and sex, (4) methods of analysis and interpretation of the data, and (5) design of a statewide s~rvey to insure consistency in data collection procedures. The actual collection of data will be management's responsibility with research providing the organization and technical assistance necessary to make the program operational. �293 FOOD HABITS STUDIES Early fall food habits of blue grouse were studied by examining the contents of 137 crops collected during check station operations in northwestern Colorado from 1975 to 1979. The contents of each crop was removed and representative samples of every food item was placed in a small coin envelope. The samples were subsequently dried and stored for later identification. Food samples were identified with the aid of a disecting microscope, reference collections, plant and insect keys, and field comparisons. Since only representative food items from each crop were preserved, results could only be reported in percent frequency of occurrence. Gilfillan and Bezdek (1944) believed that "frequency of occurrence is an important index of the leading foods and affords a more accurate picture of food preferences than volumetric data." However, frequency data do not lend themselves to statistical analysis even though they are a good index to preferred foods. Frequency of occurrence for each food item eaten was calculated by summing the occurrence of each food item and dividing that sum by the number of food samples (i.e. number of crops examined). Samples were divided into 4 categories for analysis: (1) adult males, (2) adult females, (3) juvenile males, and (4) juvenile females. Table 21 presents a complete list of plants, animals and other items recorded in the crop analyses. Plant parts eaten and percent frequency of occurrence for each age and sex class are listed. Represented in this table are 45 genera of plants and 5 orders of insects. Of the 45 genera) 21 had a frequency of occurrence greater than 5 percent. Fifteen crops were empty and consequently excluded from the data given in Table 21. Vaccinium sppo leaves were the most common food item. being represented in 41 percent of the samples. The fruit and stems of Vaccinium spp. were also heavily used. Adult grouse utilize this plant 2-3 times more than juveniles. Vaccinium spp. has been found in ~he diet of blue grouse in other studies. but not at such high freq~encies. This plant was found in only 3 and 9 percent of the crops examined by Boag (1963) in \~ashington and Knapp (1962) in north-central Colorado, respectivelyo Members of the family Formlcidae (ants) appeared in 32 percent of the samples Adults and juveniles ate ants about equally as often. but females ate these insects more often than males. Ants were common in the diets of grouse in studies done by Knapp (1962) and Boag (1963), occurring in 5 and 20 percent of the crops examined, respectively. 0 Juvenile grouse tended to eat more Insecta, Mollusca and centipedes This is what one would expect due to their (Chilopoda) than adults. higher energy and protein requirements and coincides with what other researchers have found. �294 Table 21. Fall contents of 122 crops from blue grouse collected in northwestern Colorado from 1975 to 1979. % Frequenc~ occurrence Adult Juvenile Male Female Male Female Food item Plant part Vaccinium spp. leaves fruit stems 19 6 6 10 4 3 leaves flowers 4 6 " leaves seeds Lath~rus Spa Lupinus spp. Total 6 6 3 1 2 1 41 15 12 1 4 1 10 2 23 4 3 1 8 5 6 1 22 4 leaves 3 5 4 8 20 leaves seeds 9 4 1 1 1 15 1 Amelanchier spp. fruit 3 4 2 4 14 Senec io spp. leaves flowers 3 1 3 1 1 2 1 4 fruit leaves 5 2 1 1 3 Trifolium spp. leaves 2 1 4 Antennaria rosea leaves seeds 8 1 Picea engelmannii leaves 6 3 1 1 10 Poaceae leaves 2 1 2 4 10 Juniperus connnunis fruit 4 3 3 9 Erigeron spp. leaves seeds 2 1 1 1 1 1 1 5 4 Achillea lanulosa leaves 4 3 1 1 8 Fragaria spp. leaves fruit 2 1 1 1 1 1 5 2 fruit leaves 2 1 1 1 2 6 fruit leaves 1 2 1 1 fruit leaves 1 1 1 1 2 5 1 leaves 1 3 1 1 6 seeds leaves 1 1 1 1 1 4 1 1 1 1 2 4 1 " " " " Taraxacum spp. " " Eriogonum spp. " II " " " Ribes spp. II " " " II II II " Symphoricarpos sp • II " Tragopogon dubius II " Mentzelia spp. II " Oxyria digyna Arctostaphylos uva-ursi II Rosa spp. --,,- " -11- --,,- fruit leaves 2 9 9 3 4 11 9 1 1 1 1 6 1 �295 Table 21. (cont'd ,) % Frequency occurrence Adult Male Female Juvenile Male Female Food item Plant part Pinus contorta leaves 1 1 1 4 Poa spp. leaves 1 1 1 4 Prunus virginiana fruit Rubus spp. fruit Shepherdia canadensis Carex spp. " " Draba sp. " II Total 1 1 2 4 1 1 1 1 4 leaves 2 1 4 seeds leaves 1 1 1 2 fruit leaves 1 1 1 1 1 2 1 Campanula sp. " flowers leaves 1 1 1 " 1 1 porteri leaves 1 1 2 Ligusticum Pseudostuga menziesii " " Acer spp. Agoseris sp. leaves cones 1 seeds 1 leaves 1 Alnus tenuifolia catkins Artemisia leaves tridentata 1 1 1 1 1 1 1 1 1 1 1 1 1 Mahonia repens seeds Penstemon leaves 1 1 Populus angustifolia leaves 1 1 Populus leaves spp. tremuloides 1 1 1 1 Potent ilIa sp. leaves 1 Pterospora fruit 1 seeds 1 1 1 leaves 1 1 1 andromedea Rumex sp. Thermopsis divaricarpa Vicia spp. leaves Unknown plant parts If 1 " Insects Formicidae Orthoptera Hymenoptera Coleoptera Hemiptera. Lepidoptera Unidentified " leaves seeds 1 1 36 13 1 12 3 6 3 4 6 9 6 3 32 1 10 3 3 2 2 4 2 1 3 2 4 1 6 1 1 1 6 1 1 7 6 �296 Table 21. Food item Grit Feathers (cont'd. ) Plant part % Freguency occurrence Juvenile Adult Male Female Male Female 11 12 8 12 43 3 1 1 1 6 1 3 4 Mollusca Arachnida Chilopoda Total 1 1 1 1 1 �297 The leaves of sulphur f Lower (Eriogonum spp.) were represented in 22 percent of the crops. Knapp (1962) and Haggstrom (1966) did not report this food item in their results. Boag (1963) found Eriogonum sp. in 23 percent of his crop sample. Dandelion (Taraxacum spp.) leaves were utilized by 23 percent of the birds in this study. The leaves of this plant were used by 37 percent of the birds in Boag's (1963) study, but were not mentioned in Knapp's (1962) or Haggstrom's (1966) studies. Conifer needles made up a significant part of the diet of blue grouse in Washington, Oregon, California, Idaho and Montana (Beer 1943, Stewart 1944). Although 4 species of conifer needles were found in the crops of grouse in Colorado, the highest frequency of occurrence was 10% for Engelmann spruce (Picea engelmannii). Adult grouse preferred conifer needles more than juveniles. There were major differences in the food contents of blue grouse crops examined in this study compared to the results reported by Haggstrom (1966) for birds collected in alpine areas of north-central Colorado. WillovJ (Salix spp ,) was the principle food item of alpine blue grouse, occurring in 31 percent of the crops. Willow was not found in any crop from this study. Other plants found in Haggstrom's (1966) study, but not recorded in this study include: Lychnis spp., Oxypolis fendleri, Saxifraga spp., Ranunculus spp., Veronica spp., Arenaria spp., Stellari~ spp. and Cerastium spp. Many of these plants may have been utilized by grouse collected in this study, but were among the unidentified materials. However, there are enough differences to indicate variations in the food habits of blue grouse occupying different habitat types. When reviewing the literature it becomes apparent that there are considerable differences in the diet of blue grouse according to location. For example, some of the food items which were in the diet of grouse in Washington, but were not found in this study, include: Larix occidentalis, Sambucus melanocarpa, Hieracium spp., Stellaria spp., Arceuthobium campylopodum, Ceanothus velutinus, Epilobium paniculatum and Polygonum spp. Many of these plants do not occur in Colorado. Others occur in Colorado, but may be utilized during seasons other than fall. Several plants were utilized in Colorado, but not in Washington. They include: Senecio spp., Juniperus communis, Oxyria digyna, Mentzelia spp., Ligusticum porteri, Draba spp. These dietary differences show the variability in the diet of blue grouse. Blue grouse seem to be opportunistic feeders and are able to use whatever is available to them, depending on location and time of year. However, they seem to prefer succulent, herbaceous vegetation, seeds, and fruits during the fall period. �298 Habitat Vegetation Investigations Characteristics Green Mountain lies in the southwest corner of Middle Park, one of three major intermountain parks in Colorado. Unlike the other intermountain parks characterized by broad, rolling grass (South Park) or sagebrush (Artemisia spp.) (North Park) covered floors, Middle Park is mountainous and locally heavily forested; however, sagebrush types still predominate. The study area includes portions of both the sagebrush and Douglas-fir (Pseudotsuga menziesii) vegetation zones as described in Harrington (1964). Scattered to dense stands of Douglas-fir predominate above 2700 meters and extend downward to 2600 meters. Open areas below this elevation are vegetated primarily with big sagebrush (!. tridentata). Intermediate areas are best described as a mixed conifer-aspen (Populus tremuloides)-shrub association. This zonal pattern of vegetation is not distinct as extensions of various sagebrush types occur throughout the Douglas-fir zone creating a patchwork of vegetation types. Canopy coverage for the entire study area is about 40 percent. Seventy percent of the crown cover is Douglas-fir, 27 percent aspen, and 3 percent Rocky Mountain juniper (Juniperus virginiana) and mountain maple (Acer glabrum). Understory cover consists of a mosaic of shrub and grass-forb types. Snowberry (Symphoricarpos spp.), common juniper (I. communis), rose (Rosa acicularis), choke cherry (Prunus virginiana), and serviceberry (Amelanchier spp.) are the major understory shrubs; yarrow (Achillea lanulosa), pussy toes (Antennaria spp.), northern bedstraw (Galium boreale) and peavine (Lathyrus spp.) are the major forbs encountered in the understory, and sedge (Carex spp.) and bluegrass (Poa spp.) are the major grass and grasslike species. Unforested areas constitute about 60 percent of the study area of which in excess of 90 percent is dominated by sagebrush. Other shrubs, forbs, and grasses (or grasslike) commonly found in association with the sagebrush include: snowberry, rabbitbrush (Chrysothamnus spp.), serviceberry, rose, bitterbrush (Purshia tridentata), pussy toes, arrowleaf balsamroot (Balsamorhiza sagittata), paintbrush (Castelleja spp.) sulphur flower (Eriogonum umbellatum), common lupine (Lupinus argenteus), bedstraw, sedge, wheat grass (Agropyron spp.), Junegrass (Koleria cristata), needle-grass (Stipa spp.) and bluegrass. Plant nomenclature is according to Weber (1972) and Nelson (1969). Eiby Creek falls within a transition belt between the Pinyon-Juniper and Spruce-Fir zones and consists of a mosaic mixture of mountainshrub, aspen, sagebrush and riparian communities. Coniferous types are virtually absent within this belt, except in a few localized areas. Extreme variations in slope, aspect, soil type and moisture availability contribute to the complexity of vegetation on the study area. Open to dense stands of aspen are scattered throughout the area and occur in various growth forms from scrub thickets to over-mature, tall stands with a poorly developed shrub layer. The largest, continuous stand encompasses about 80 hectares. Most stands range in size from 2 to 10 hectares and occur in small patches that are separated by unforested areas alternately dominated by sagebrush, serviceberry, choke cherry, and snowberry. Major drainage is to the south via Eiby Creek �299 and its numerous tributaries. Thinleaf alder (Alnus tenuifolia), willow (Salix spp.), aspen, and red-osier dogwood (Cornus stolonifera) form fingers of dense thickets along these stream courses. Approximately 45 percent of the area was classified into forested types. Canopy coverage was estimated at 32 percent of which 91 percent is aspen, 7 percent alder, and 2 percent balsam poplar (Populus balsamifera) and Rocky Mountain juniper. The dominant understory shrubs are snowberry, currant (Ribes spp.), choke cherry, serviceberry, and rose. Conspicuous herbs are yarrow, sweet cicely (Osmorhiza spp.), meadow rue (Thalictrum spp.), dandelion (Taraxacum officinale), peavine, northern bedstraw, and chickweed (Stellaria spp.). Grasses most commonly encountered are bluegrass, brome (Bromus spp.), and wheat grass. The remainder of the study area (55%) is an assemblage of shrub types mainly dominated by sagebrush and secondarily by serviceberry and choke cherry. Rabbitbrush, snowberry and rose co~~only grow in association with these types. Many of the forbs and grasses previously mentioned as being abundant in the forested types also frequently occur in unforested areas. Other notable forbs and grasses (or grasslike) include: arrowleaf balsamroot, common lupine, fleabane (Erigeron spp.), larkspur (Delphinium spp.), sulphur flower, geranium (Geranium fremontii), mint (Ag~stache spp.), violet (Viola spp.), bluebells (Mertensia spp.), sedge and Junegrass. Several factors have markedly influenced the composition and structure of vegetation on both study areas. Between 1880 and 1890, the northern portion of the Green Mountain study area was set on fire by a local homesteader to create a source of firewood. Much of the area burned has reverted to sagebrush wh Lch charred logs, stumps, and patches of Douglas-fir dotting the landscape. Around 1930 the Douglas-fir was selectively logged as a local source of ties for constructing the railroad. Few trees were removed, but enough to enhance development of the understory. Both study areas were subjected to extensive disturbance in the early 1960's when sagebrush dominated areas were sprayed with 2,4D to enhance grass production for livestock. The degree of mortality in shrub species was not uniform throughout the sprayed areas, thus, vegetative composition and structure were altered moreso at some sites than others. Adequate time has since elapsed for the sagebrush to recover except in those areas where spraying was most e'ffective. Dead or partially damaged sagebrush and serviceberry plants were still an important component of ground cover in such areas. Perhaps the greatest impact on the vegetation at Eiby Creek and to a lesser extent on Green Mountain has been excessive livestock grazing. Eiby Creek is still heavily grazed while Green Mountain receives only light summer use by cattle mostly at the lower elevations near available water. Regeneration of aspen on the Eiby Creek study area has been seriously impeded or virtually eliminated and in some places the aspen stands are deteriorating and reverting to a brushland or grassland type. Mule deer use both areas from late spring until early winter and elk are commonly found at Eiby Creek in spring and early winter. Besides grazing, no other use is presently made of either area. �300 Vegetation Type Mapping Vegetative classification and description of the study areas was done in accordance with procedures outlined by Kuchler (1955). This method of vegetation mapping is essentially a derivation of Braun-Blanquet's (1951) system of floristic and physiognomic description of plant communities. Vegetation maps have been prepared elsewhere in Colorado using this method (Braun 1969, Medin 1962). Aerial photographs of the study areas were used as base maps. Before going into the field, the photos were carefully examined and every area of vegetation appearing dissimilar from adjacent areas was delineated by a line drawn directly on the photo. The minimum area of vegetation bounded by a line on the photo was approximately .5 hectares. No types were omitted within the study area boundary. Upon delineation of all areas apparent on the map, each area was then inspected in the field. The first task was to walk about the area being examined from end to end and across, observing the vegetation critically and field checking the boundaries shown on the photo. Salient features of the area were recorded on prepared data sheets. Consecutive numbers were assigned to each area inspected and were entered both on the data sheet and the photograph. Each area received its own number regardless of any similarities to previously inspected areas. To better facilitate preparation of the final map, a photograph was taken of each vegetative unit. Number of the individual exposure was recorded along with the corresponding number of the unit where the photograph was taken. The next step was to record the vegetative physiognomy and other physical features of the area including: slope gradient, exposure and character. The physiognomic classification (Table 22) reveals the appearance and structure of the plant community (vegetation unit), i.e., height and density of every item listed, and in addition, such special features as may be present. A detailed description of the classification scheme and its application is presented by Kuchler (1949). Slope gradient and exposure were measured with an Abney level and compass. After the physiognomic formula for an inspected area had been established, attention was focused on the floristic character of the vegetation. All species occurring on the ar~a were recorded by name and each species was assigned numerical values describing its coverage and sociability (Table 23)~ Coverage is understood to mean the percentage of ground that would be covered if the full spread of the species were projected vertically to the ground. Sociability refers to the distribution of a species within the area of vegetation under consideration. Undoubtedly, some species which only very few specimens were present in a given type escaped detection. This was also true for "early or late flowering" species that peaked prior to or after the period of field work, which was designed to coincide with the peak of plant development (mid-June to mid-July). �301 Ta bl e 22 . CAPITAL . c 1 aSSl.f'lcatlon Ph'YSlognomlc. 0 f vegetatlon.. 1/ LETTERS: Woody Vegetation: B: evergreen broadleaf D: deciduous broadleaf E: evergreen needleleaf N: deciduous needleleaf 0: without leaves Herbaceous Vegetation: G: graminoids H: forbs L: lichens and mosses SMALL LETTERS: Group I: Height: t: m: 1: Group tall; height height med ium tall; Height Height low; Haximum Maximum height height of trees: of herbaceous of trees: of herbaceous 25 m 2 m plants: 10-25 m Yz-2 plants: of trees: of herbaceous m 10 m plants: Yz m s: shrubs; Hinimum height: 1 m z: dwarf shrubs; Maximum height: 1 m II: Density: c: continuous i: interrupted; p: plants r: rare, yet conspicuous b: barren; Group III: Special 1/ Hinimum Hinimum growth plants scattered vegetation usually singly, do not touch or in groves largely or entirely absent Features: e: epiphytes u: palms j: lianas v: bamboos k: succulents w:' aquatic q: cushion y: - From Kuchler or patches plants (1955). tree ferns and tuft plants �302 Table 23. Floristic classification of vegetation.1/ Coverage: Sociability: + Very sparsely present; cover very small 1 Plentiful but less than 1/20 of the area 2 Covering 1/2 - 1/4 of the area 3 Covering 1/4 - 1/2 of the area 4 Covering 1/2 to 3/4 of the area 5 Covering greater Kuchler (1955). l/From 1 Growing singly 2 Grouped or tufted 3 In small patches 4 In extensive 5 In great crowds patches than 3/4 of the area Additional features such as the presence and amount of bare ground, stumps, logs and rocks were recorded for each vegetation unit. Comments on phygiography, soil, water condition, whether the area was logged or burned, and any other pertinent information were noted on the data form. The result of the laboratory and field activity was a set of lists and notes and a series of aerial photos on which the vegetation units were outlined. Each unit received its own number corresponding to the number in the lists where the particular type was analyzed. The next step was the preparation of the base map which shows the exact outline of each indi.vidual vegetation unit and its number. Preparation of the base map involved sorting through the data forms, notes and photographs and rearranging the vegetation units into combinations encompassing similar floristic and physiognomic classifications. Woody plants including trees and shrubs were given primary emphasis during the first phase in the sorting and combining process. Units within these major vegetation types were then manipulated to produce categories based on similar floristic and structural characteristics. After assigning each unit to a major ve~etation type and combining similar vegetation units, the final map was traced off the base map, lumping together all similar units which were adjacent to each other. All similar units were then reassigned the same number to simplify reading the map. Total area mapped as planimetered from U.s. Geological Survey topographic maps was 181.3 hectares for Green Mountain and 482.0 hectares for Eiby Creek. Area encompassed by each vegetation type and unit was calculated by the weight method (Welch 1948, p. 85). �303 Five major vegetation types and 26 vegetation units were recognized on the Green Mountain study area (Figs. 4 and 5) compared to 7 vegetation types and 27 units delineated at Eiby Creek (Figs. 7 and 8). Vegetative summary tables (Appendix) were compiled from the data sheets and are presented as supplementary information describing the physiognomy, floristics, and physiography of each vegetation unit. Examination of the vegetation maps might indicate that the vegetation types and units were clearly distinct. While the major types and some units were distinct, most units were not, with the majority of units gradually intergrading into adjoining units. In some cases the transition was so gradual as to warrant establishment of a separate unit. When the transition was more abrupt, the boundary line was placed midway through the area of overlap. Thus, admixtures of adjacent units were frequently present along their boundaries. "Islands" of dissimilar vegetation within a larger, individual unit were usually too small « .5 ha.) for recognition, and consequently, were ignored unless they were of known importance in terms of grouse utilization. Characteristics of Breeding Areas Blue grouse males on the study areas were territorial and defended chosen territories throughout the breeding season (mid-April to early July). All territories located (GM = 22, EC = 36) from 1975 to 1979 were marked on aerial photos. Territories were subsequently plotted on the vegetation maps to facilitate their description in relation to vegetative features of the area (Fig. 6 and 9). Delineation of territorial boundaries was limited to 9 territories on Green Mountain where the resident male was banded and observed 5 or more times within the same breeding season. Otherwise, only an approximate location of the territory was determined. No territory was located completely in anyone vegetation unit as portions of at least 2 and up to 4 units were encompassed within the boundaries of all territories on both study areas. Breeding territories were primarily located where Pseudotsuga (GM) or Populus units (EC) intergraded into typically more open areas such as the Artemisia units (GM and EC) and the Amelanchier, Symphoricarpos, and grass-forb units (EC). All territories at Eiby Creek were at least partially associated with the Populus units. Riparian units were not extensively utilized by territorial males except where the aspen extended away from the creek bottom and adjoined a more open unit. Douglas-fir units were the major vegetational components of 21 of 22 territories on Green Mountain, though aspen was frequently present but in small amounts. The only territory void of Douglasfir was associated with an aspen-sagebrush mixture, and this territor~ was occupied for just one breeding season. Territorial males avoided dense stands of Douglas-fir except along the edges adjacent to more open units. �304 Aspen also appeared to be an important part of breeding habitat on Green Mountain as it was much less abundant in the overall tree cover than at sites where territorial males were observed. This was attributed to the fact that (1) territorial males were most frequently found while performing their displays, and (2) aspens occurred along the edges of coniferous stands and openings which were the same areas preferred by males for displaying. Features found in cornmon among all territories examined included: (1) some form of tree cover, (2) shrub thickets, (3) edges, (4) open areas, and (5) some degree of openness in both the canopy and understory cover. The data do not imply that territorial males selected Douglas-fir and aspen over other species of trees for breeding habitat, but simply reflect the high incidence of Douglas-fir and aspen on the two study areas. What the data do suggest is that structural characteristics of the vegetation moreso than species composition are an important factor in breeding habitat selection. Within Colorado, and throughout its range, the blue grouse has been documented to breed in a variety of forest and mountain shrub types from the foothills to timberline with no apparent restrictions to any habitat type within this elevational range. Territorial males were seldom found more than 25 meters from an opening and avoided areas with dense canopy or understory cover except along the edges. During early to mid-April when most ground cover was beneath the snow, the birds on Green Mountain spent the majority of time in trees utilizing small, dense clumps of conifers in an otherwise open habitat. As phenological changes progressed and the birds reverted to ground dwelling habits, shrub thickets became important during resting and feeding activities while trees were primarily used as escape cover and for roosting. Conversely, the birds at Eiby Creek depended heavily upon shrub thickets for cover in early spring. The openness of the aspen stands afforded little cover from predators. Only in the early morning and late evening hours during feeding activities were the birds found in the aspen. unless disturbed while on the ground, then they would frequently fly into a nearby aspen. Some territorial males, both at Eiby Creek and Green Mountain, performed their displays considerable distances (750 m) from tree cover and therefore relied on nearby shrub thickets for cover. Areas characterized by an interrupted mixture of low growing vegetation, bare ground, stumps, logs and patches of tree and shrub cover were preferred over areas with continuous or homogeneous ground cover. �305 Characteristics of Nest Sites Twelve nests were found on the study areas (GM = 11, EC = 1) from 1976 to 1979. Each nest site was described in detail as to aspect, slope, altitude and vegetative cover. All nests had some type of cover immediately above and surrounding the nest. The cover was mainly in the form of shrub clumps, but two nests were partially protected by a log while another was adjacent to the trunk of a Douglas-fir. Three nests were found under sagebrush, three beneath saplings of Douglas-fir, one at the base of a larger Douglas-fir (approx. 6m high), two in the center of a sagebrush-snowberry clump, one in a sagebrush-rabbitbrush clump (EC nest), one under a log partially covered by snowberry, and one in a serviceberry-sagebrush clump. Maximum distance from the nest to the nearest tree was 42 meters. Excluding the nest under the large Douglas-fir, the height of vegetation immediately above the nest averaged 98 cm and ranged from 30 to 280 cm. General appearance of the habitat around the nest was open to semi-open. Of the 12 nests located, 6 were found in transition zones (edges) between the following units: (1) dense Pseudotsuga-Symphoricarposmixed shrub and Artemisia-Amelanchier-mixed shrub-scattered Pseudotsuga (2 nests), (2) semi-open Pseudotsuga-Populus-Symphoricarpos~mixed shrub and Artemisia-Symphoricarpos-mixed shrub-scattered Pseudotsuga (2 nests), (3) semi-open Pseudotsuga-Populus-Symphoricarpos mixed shrub and Artemisia-mixed shrub-scattered Pseudotsuga (1 nest), and (4) semi-open Pseudotsuga-Populus-Symphoricarpos-mixed shrub and dense Pseudotsuga-Symphoricarpos-mixed shrub (1 nest). Two of the other 6 nests were in the Artemisia-Amelanchier-mixed shrub-scattered Pseudotsuga vegetation unit and one nest was found in each of the following 3 units: (1) Artemisia-Symphoricarpos-mixed shrub-scattered Pseudotsuga, (2) scattered Pseudotsuga-Populus-Artemisia-mixed shrubrock, and (3) Artemisia-mixed shrub-scattered Pseudotsuga. The one nest location for Eiby Creek was situated in the Artemisia-Chrysothamnus-mixed shrub vegetation unit. Nest sites were in the same vegetation units selected for breeding, but only half the nests were located within the boundaries of a territory. Artemisia tridentata was associated with all nesting sites except one, but this may simply be related to the abundance of sagebrush on both study areas. However, hens apparently did select the more open units for nest sites and these same units were preferred brood use areas. All nests were between 2500 and 2740 meters elevation. Sample size was inadequate to evaluate the importance of slope and aspect, but neither feature was believed to be a significant factor in nest site selection. Exposure of nests included all major compass directions except south. The majority of nests faced in northerly and easterly directions which were the primary exposures for the Green Mountain study area where most of the nests were found. Slope measurements at the nest site ranged from 6 to 25 percent. �306 Characteristics of Brood Areas Two hundred and fifty-seven observations of blue grouse broods were made on Green Hountain (N = 121) and at Eiby Creek (N = 136) from 1975 to 1979. At one time or another, broods were observed within most all vegetation units shown in Figures 5 and 8. However, there were definite concentration areas. Nearly 90 percent of all brood sightings on Green Mountain were made in the open and semi-open habitats on the east face of the study area (Fig. 6). Broods tended to use more of the study area at Eiby Creek, possibly due to the greater availability of open areas and edges. Nevertheless, broods congregated in certain areas. These areas, as depicted in Figure 9, accounted for about 70 percent of all brood observations and together with the concentration sites on Green Mountain were the focal areas for evaluation of brood habitat. Broods used areas where vegetation had interspersions of plants of various life forms. Such areas were generally along the edges of brush or tree cover and provided a high degree of concealment. Forested areas with a dense understory or open areas with heavy herbaceous ground cover were avoided except along the edges. Almost invariably, any brood encountered in the field was originally observed on the ground. They seldom ventured more than 30 meters from brush or tree cover. Shrub thickets and edges of forested areas were used for resting and for escape when disturbed. Broods were frequently found along stream courses or near other wet sites at Eiby Creek providing the vegetation was not too den se . However, there was no evidence of the necessity for open water near brood cover. Only two sources of open water occur on Green Mountain, and no broods were found at either location. Evidently the broods obtained their water requirements from succulent foods and condensation of moisture on plants. I Summer brood range partially overlapped with breeding areas, but there tended to be a greater diversity of plant species, more ground cover (especially herbaceous), and a greater amount of open vegetation units in brood habitat than breeding habitat. Again, structural characteristics of the vegetation moreso than species composition appeared to be the critical factor in habitat selection by broods. Examination of Figure 6 indicates that a major portion of the brood range on Green Mountain was included wi.thin the semi-open PseudotsugaPopulus-Symphoricarpos-mixed shrub vegetation unit. Even in this predominantly Douglas-fir unit, broods were most often found near or in the many small clearings (Artemisia-Symphoricarpos-mixed shrubscattered Pseudotsuga) dispersed throughout this unit. Characteristically, these openings, as with other forest edges on the study area, were often bordered by aspen; thus, the distribution of broods closely approximated the distribution of aspen on the study area. At lower elevations broods were found in transition area~ between the coniferous and sagebrush types where aspen was commonly present. Vegetation units favored by broods in these areas included: scrub, low, and medium Populus-mixed shrub units, Artemisia-Chrysothamnus-mixed shrub, Artemisia-Amelanchier-mixed shrub-scattered Pseudotsuga, open Pseudotsuga-Populus-mixed shrub, and open Pseudotsuga-ArtemisiaSymphoricarpos-mixed shrub. �307 There was a great deal of variation in brood habitat at Eiby Creek. Broods were found in and along the edges of aspen groves, on open sagebrush hillsides, along alder-willow dominated stream courses, and in dense patches of choke cherry and serviceberry. Even so, there was a clear preference for edges where open vegetation units occurred adjacent to aspen stands or patches of tall shrubs, namely serviceberry and choke cherry. Aspen was the dominant tree in brood cover, but there was no apparent preference for certain aspen units as long as the stands were bordered by open areas. During daylight hours, broods were most frequently found foraging on open hillsides covered with shrubs, while at night they roosted in aspen stands or patches of serviceberry and/or choke cherry. Sites used for roosting also served as resting and loafing cover during the day; and when disturbed, the hen and chicks usually flew into the nearest aspen stand or shrub thicket for escape cover. The Artermisia-Symphoricarpos-Chrysothamnus-mixed shrub was the single most important vegetation unit in terms of brood use at Eiby Creek, especially where this unit bordered aspen. Other open units where broods were commonly located included: Artemisia-Chrysothamnus-Prunus-mixed shrub and Chrysothamnus-Symphoricarpos-Prunusscattered Artemisia (sprayed area). Several smaller, localized areas were identified as brood concentration sites. One such area that was heavily used by grouse throughout the spring, summer, and fall consisted of a mixture of the Amelanchier-Prunus-mixed shrub, medium Populus-Symphoricarpos-Prunus-Amelanchier-Ribes, and ArtemisiaCercocarpus-Amelanchier-mixed shrub vegetation units. Light to moderate grazing pressure had little impact on brood habitat at Eiby Creek. However, excessive grazing pressure virtually eliminated the herbaceous cover forcing the grouse to seek less disturbed sites. Such an influence was particularly noted in the forb-grass scattered shrub vegetation unit. Where herbaceous cover remained, broods were found in the same pastures with cattle, but usually on steep slopes or thickets less frequented by the cattle. �308 GREEN MOUNTAIN t I x •• )(~ "' " _'I. --_.j _ Pseudot suga-Populusmixed shrub (44.4) I I Populus - mixed shrub (13.2) ~ Carex-A gropyronsca t t er ed shrub (2.9) 1/ -c- Surfa ce area in hecta res PI N 4~ SCALE ~--------------~I--------------'J o Y2 .IK. ~~~gsgs~ I km ~ ~ ~ "'- .... Fig. 4. Major vegetation types, Green Mountain study area. �309 GM 1 Artemisia-mixed shrub-scattered Pseudotsuga GM 2 Artemisia-Symphoricarpos-mixed GM 3 Artemisia-Chrysothamnus-mixed GM 4 Artemisia-Amelanchier-mixed GM 5 Artemisia-mixed GM 6 Artemisia-Agropyron-Chrysothamnus GM 7 Dense Pseudotsuga-Symphoricarpos-mixed shrub GM 8 Dense Pseudotsuga-bareground-scattered shrub GM 9 Dense Pseudotsuga-Juniperus-mixed shrub-scattered shrub shrub-scattered shrub-rock-scattered Pseudotsuga Pseudotsuga shrub GM 10 Dense Pseudotsuga-Acer-mixed GM 11 Semi-open Pseudotsuga-Symphoricarpos-mixed GM 12 Semi-open Pseudotsuga-mixed GM 13 Open Pseudotsuga-Artemisia-Symphoricarpos-mixed GM 14 Open Pseudotsuga-Prunus-mixed GM 15 Open Pseudotsuga-Artemisia-mixed shrub GM 16 Open Pseudotsuga-Juniperus-mixed shrub-rock GM 17 Semi-open GM 18 Open Pseudotsuga-Populus-mixed GM 19 Dense Pseudotsuga-Populus-mixed GM 20 Scattered GM 21 Low Populus-mixed GM 22 Medium Populus-mixed GM 23 Scrub Populus-mixed GM 24 Medium Populus-Pseudotsuga-Prunus-mixed GM 25 Medium Populus-Carex-Lathyrus-mixed GM 26 Carex-Agropyron-mixed Fig. 5. Pseudotsuga shrub shrub-rock shrub outcrop shrub shrub Pseudotsuga-Populus-Symphoricarpos-mixed shrub shrub shrub-rock Pseudotsuga-Populus-Artemisia-mixed shrub-rock shrub shrub shrub grass-scattered shrub shrub shrub Vegetation units, Green Mountain study area (supplementary information presented in Appendix). �310 GREEN MO N SCALE o I km Fig. 5. Vegetation units, Green Mountain stu y information presented in Appendix). �311 GREEN MOUN Approximate of breeding 1][1 location territories Approximate boundary breeding territories r- '3 ,----. Brood concentration 1_ , d of areas N SCALE o Fig. 6. Major vegetation types and blue grouse use areas, Green Mountain. �312 EI Y CREEK shrub (184.5)l/ Artemisia-mixed Populus-mixed Amelanchier-mixed shrub (45.8) Symphoricarpos-mixed Prunus-mixed N shrub (179.6) + shrub (10.6) shrub Forb-Grass-meadow [ ~opulus-Alnus/Alnus-Salix riparian Fig. 7. types, Eiby Creek study area. Major vegetation (43.4) a SCALE I 1/2 ::JIkm �313 EC 1 Artemisia-Chrysothamnus-mixed EC 2 Artemisia-Chrysothamnus-Prunus-mixed EC 3 Artemisia-Cercocarpus-Amelanchier-mixed EC 4 Artemisia-Amelanchier-rock-mixed EC 5 Artemisia-Symphoricarpos-Chrysothamnus-mixed EC 6 Chrysothamnus-Symphoricarpos-Prunus-scattered EC 7 Artemisia-Amelanchier-mixed EC 8 Medium Populus-Symphoricarpos-Ribes-mixed EC 9 Medium Populus-Symphoricarpos-Prunus-Amelanchier-rock shrub shrub shrub shrub shrub Artemisia (sprayed area) shrub (sprayed area) shrub outcrop EC 10 Medium Populus-Symphoricarpos-Prunus-Amelanchier-Ribes EC 11 Medium Populus-Symphoricarpos-Prunus-Ribes EC 12 Medium Populus-mixed EC 13 Low Populus-Amelanchier-Prunus-Symphoricarpos EC 14 Scrub Populus-Symphoricarpos-Prunus-Rosa EC 15 Open Populus-mixed EC 16 Open Populus-Symphoricarpos-mixed EC 17 Amelanchier-Prunus-Symphoricarpos-mixed EC 18 Amelanchier-Prunus-mixed EC 19 Amelanchier-Prunus-Symphoricarpos-scattered EC 20 Symphoricarpos-Prunus-mixed EC 21 Symphoricarpos-Chrysothamnus-mixed EC 22 Prunus-Symphoricarpos-mixed EC 23 Forb-Grass-scattered EC 24 Populus-Ainus-mixed EC 25 Alnus-Populus-Salix-Ribes-Cornus-riparian EC 26 Alnus-Salix-Ribes-riparian EC 27 Salix-Alnus-Cornus-riparian Fig. 8. Vegetation units, Eiby Creek study area (supplementary information presented in Appendix). forb-grass shrub shrub shrub shrub Juniperus shrub shrub-scattered shrub-scattered Populus Populus shrub shrub riparian �314 EIBY CREEK N + SCALE I 0 Fig. 8. Vegetation units, Eiby Creek study area (supplementary presented in Appendix). I 1/2 information I Ikm �315 EIBY CREEK N ~ ~ c..:...::l i- Brood concentration .=-:-=1 Fig. 9. t Approximate location of breeding territories SCALE areas c_ _____,..I __ ~::J o Major vegetation 1/2 types and blue grouse use areas, Eiby Creek. I kro �316 LITERATURE Beer, J. R. 1943. 7(1):32-44. CITED Food habits of the blue grouse. J. Wildl. Manage. Bendell, J. F., and P. W. Elliott. 1967. Behavior and the regulation of numbers in blue grouse. Can. Wildl. Servo Rept. Ser. 4. 76pp. -------- , D. G. King, and D. H. Mossop. tion of blue grouse in a declining 36(4):1153-1165. 1972. Removal and repopulapopulation. J. Wildl. Manage. Boag, D. A. 1963. Significance of location, year, sex, and age to the autumn diet of blue grouse. J. Wildl. Manage. 27(4):555-562. 1966. Population attributes of blue grouse in southwestern Alberta. Can. J. Zool. 44:799-814. Braun, C. E. 1969. Population dynamics, habitat, and movements white-tailed ptarmigan in Colorado. Ph.D. Dissertation. Colorado State Univ. 189pp. of 1971. Determination of blue grouse sex and age from wing characteristics. Colorado Div. Game, Fish and Parks. Game Info. Leaflet. No. 86. 4pp. Braun-Blanquet, J. Wien, Germany. 1951. Pflanzensoziologie. pp. 58-66. Springer-Verlag, Gilfillan, M. C., and H. Bezdik. 1944. Winter foods of the ruffed grouse in Ohio. J. Wildl. Manage. 8(3):208-210. Haggstrom, D. A. 1966. Fall food habits of blue grouse in the northcentral Colorado alpine. Unpubl. rep., 9pp. Harju, H. J. 1974. An analysis of some aspects of the ecology of dusky grouse. Ph.D. Dissertation. Univ. Wyoming. 142pp. Harrington, H. D. 1964. Manual of the .plants of Colorado. Books, Denver. 666pp. Sage Hickey, J. J. 1955. Some American population research on gallinaceous birds. Pages 326-396 in A. Wolfson, ed. Recent studies in avian biology. Univ. of Illinois Press, Urbana. 479pp. Hoffman, R. W. 1976. Population dynamics and habitat relationships of blue grouse. Colorado Div. Wildl. Job Prog. Rept. Fed. Aid Proj. W-37-R. April 1976. p. 135-152. �317 1977. Population dynamics and habitat relationships of Colorado Div. Wildl. Job Prog. Rept. Fed. Aid blue grouse. April 1977. p. 83-104. Proj. W-37-R. 1979. Population dynamics and habitat relationships of Colorado Div. Wildl. Job Prog. Rept. Fed. Aid blue grouse. April 1979. p. 201-244. Proj. W-37-R. Knapp, D. B. 1962. September foods of blue grouse in north-central Colorado. Unpubl. rep., 6pp. Kuchler, A. W. 1949. A physiognomic classification Ann. Assoc. Amer. Geographers 39(3):201-210. 1955. A comprehensive method of mapping Assoc. Amer. Geographers 45(4):404-415. Martinka, R. R. territories 498-510. of vegetation. vegetation. Ann. 1972. Structural characteristics of blue grouse in southwestern Montana. J. Wildl. Manage. 36(2): Medin, D. E. 1962. An ecological investigation of the Cache la Poudre deer herd. Colorado Dept. Game and Fish. Job Completion Rept. Fed. Aid Proj. W-105-R. July 1962. p. 187-204. Mussehl, T. W. 1960. Blue grouse production, tions in the Bridger Mountains, Montana. 24(1):60-68. Nelson, R. A. 1969. Handbook King, Tucson. 331pp. movements, and populaJ. Wildl. Manage. of Rocky Mountain plants. Dale Stuart Redfield, J. A., and F. C. Zwickel. 1976. Determining the age of young blue grouse: a correction for bias. J. Wildl. Manage. 40(2):349-351. Rogers, G. E. 1968. Fish ann Parks. Stewart, R. E. 112-120. 1944. Weber, W. A. 1972. Press, Boulder. The blue grouse in Colorado. Tech. Publ. No. 21. 63pp. Colo. Div. Game, Food habits of the blue grouse. Rocky Mountain 437pp. Welch, D. S. 1948. Limnological Inc., New York. 381pp. flora. methods. Condor 46(3): Colorado Assoc. Univ. McGraw-Hill Book Co., �318 Zwickel, F. C. 1965. Early mortality and numbers of blue grouse. Ph.D. Dissertation. Univ. British Columbia, Vancouver. 153pp. 1972. Removal and repopulation of blue grouse in an increasing population. J. Wildl. Manage. 36(4):1141-1152. 1975. Nesting parameters of blue grouse and their relevance to populations. Condor 77(4):423-430. ______ ~' and J. F. Bendell. 1967. Early mortality and the regulation of numbers of blue grouse. Can. J. Zool. 45(5):817-851. and A. N. Lance. 1966. Determining grouse. J. Wildl. Manage. 30(4):712-717. Wildlife Researcher C the age of young blue �319 APPENDIX �320 DESCRIPTIVE SUMMARY OF VEGETATIONTYPE MAP, GREEN MOUNTAIN- Bl.Uf. GROUSE INVESTIGATIONS Legal Description: T2S, RBOW.Sections 2, 3, 10, 11 County: SUl:lClit Range: 2500-2865 m Surface Area: 181.) ha Aerial Photo No. U.S~CN 10-3-72 Elevational VEGETATIVE Classification Physlognomi and c GM 1 Fe a t ur e s !I BziDziElp GM-2 GM 3 bziDziElp GM-4 GM-5 BziOzpElp 350-90 160-180 180 25-27 10-12 3.8 2.2 3&-85 36-46 45-180 18-30 20-25 5-17 14.7 Ccv2/sncL' Plant Species List 3/ ~.Trees: Acer glabrum Juniperus virginiana Populus e reeu l of des Paeude t s uga eena ies t r Woody-Shrubs: xcc r glabrum ~lanchier alnifolia Arctostaphylos cvc-urs i ArteClisia cana Artemisia t rddent at.a Ccanot bus velutinus Cercocarpus eont anus Chrysothamnus spp. Holodiscus dumosus Juniperun communis Juniperus virginiana Mahonia repens Pechys t t ec myrsinites Populus t rceut o tdes Prunue virginiana Pscudot scgc menzies f i Purshia tridentata Rhus t r t Iobaca Rlbes spp . Rosa ac rcul ar t s Rubun de Hc Losus SnClbucus rncceosa Shcpherdla canadensis Sycrphoricarpos oreophi1us Tetradymia cane sccno Vaccinium app , Forbs: Achillea Ianujosc Anemone app • Antennaria app . Aquilegia caerulen Arabia spp. Arnica spp. Artemisia frigida ArteClisia ludovic1ano. Aster spp. Astragalus spp , Bnlsaoorhiza s ag Lt t a t a catccbor ccc gunnisonii Catlpanu1a rotundifolia Aasigncd according to Soc Cov 41.9 Soc Cov Soc BzpGUDzp FEATURES NUMBER GH-7 GM 8 £mcliDzp Eme )40-)5 14-28 EmiDlzp 125 310 28 10 10-20 17 .7 7.2 .7 18.3 FLORISTIC CHARACTERISTICS Soc Cov Soc Cov Soc Cov cov +-1 1-2 + 3 1-3 1-2 GM-IO GM-9 EmliOzp[zp Soc +-1 4 Cov Soc +-1 4 1-4 cov GM-I:! GM 13 . EmiDzp ElpBSi[)zi 20-32 315 20-90 18-25 25-35 17-22 4.0 2.9 2.2 Soc Cov Soc Cov Soc Cov +-1 +-1 4 3-4 3 +-1 1-2 3-4 3 +-1 1-4 2 1-2 3 1-3 t::1-11 EmHlzp 3 1-2 3 1 +-1 1-2 1 Soc 1 1-2 3 1-2 1-2 2-3 1-3 + 2-3 1-3 3-4 1-4 3-4 + 1-2 1-2 1-2 "__1 +-1 +-1 +-1 1-2 +-1 +-1 +-1 1-2 1 +-1 +-1 1 +-1 1-4 1-2 1 + +-"i 1 1 1 1-2 2 2-3 1-4 1-2 1-3 +-1 1 1-2 1 1 2-3 1 1 1-2 1-3 2-3 +-1 3 1 1-2 1-2 +-1 2-3 1 1-2 1-2 +-1 1 3 + + +-1 1 1-2 +-1 1-2 + 1-3 1 1-2 2-3 2 +-1 2 1-2 + 1-2 2-3 1-2 +-1 +-1 2-3 1 +-1 +-1 1-3 1 1-2 1-2 +-1 1-3 +-1 +-1 + +-1 1-2 1 1 +-1 +-1 +-1 + + 1-3 1-2 1 1 +-1 +-1 1-2 1-2 +-1 1 1-2 1 +-1 1-2 1 1-2 + +-1 1-2 +-1 +-1 1-3 1 1-3 1-2 1 "__1 1 +-1 1-2 1-2 1-2 1-2 1-2 2 1-2 +-1 1 + +-1 +-1 1 +-1 1-2 +-1 1-2 1 2-3 +-1 1-2 +-1 +-1 2-3 2 1-3 1-2 1 1 +-1 +-1 +-1 +-1 1-3 ] +-1 +-1 + +-1 2-3 1-3 1-2 1 +-1 +-1 +-1 3 +-1 +-1 +-1 1 +-1 1-2 1-2 +-1 +-1 1 +-1 +-1 +-1 1-2 +-1 +-1 1-2 1 +-1 1-2 +-1 1-2 +-1 +-1 +-1 +-1 1-2 +-1 +-1 1-2 "__1 +-1 +-1 +-1 1 1 +-1 +-1 +-1 1-2 +-1 +-1 1-2 +-1 1-2 +-1 2 1-3 1-3 1 1 1-3 1-2 + +-1 2-3 1-2 2 ccscc ttejc spp , Chcencc t Iu doug1asii Chenopodiuo npp , Cirsiuo upp , C1ematia coluobiana cfceae rs hirsutissica cceendre ucbe11ata Crepis spp. Cz-yp t ant.hu upp , Cynogoloaaum officina1e Delphinium. spp. . Descurainia epp , Dodecatheon pu1chelluo Ebilobiuc app , Erigeron app , Eriogonur:. spp , Erysimuc spp , Fugaria app , Frasera epee toea Callum bo renLe Gcraniuc. frccontii HupLopappua cpp , Hcucheca spp , Heleniuo hoopesii Hc1ianthe11a quinquenervis Ipomopsis agg regat;a LD.thyrus opp , Linue lewis1 i Litho9p~rmul:l spp • Lupinu9 arg(!;ntiuG xe r tensdc opp . Oxytropia spp. Penc t eecn upp , Phuce 1 La spp. Phlox opp. Potentilla spp , Pricu1a app. Psecdccveopcerce ecntcnuc Pulsatilla patens Saxifragll bronchialis Senecio app , Scilacina spp. Solidago opp , Taraxacul:l officina1e Tragopogon dubiuG Thalictruo GPp. Viola opp. Craoinoids: Agropyron app. arceue app. cares opp. Dcoch.o.cpsia caesp1tosa Fcatuca spp. Irio ciosouriensis Kalerio gracilis Oryzops1s hymenoides Ph1C!um pratense Poa spp. S1tanion longifoliu:: Stipa spp. 1/ Cov 5-25 10.0 4.5 UNIT BziDszpElp Exposure (negecce Azimuth) Area (Hectares) VEGETATION GM 6 BziDzi GrOldlent (Percent, CLASSIFICATION AND SUPPLEIiENTARY 1 3 1-2 1 +-1 +-1 1-2 +-1 1 +-1 ·2 +-1 +-1 +-1 +-1 + 1-2 3 1 1-3 3 1 +-1 1-3 1 1-3 1 +-1 1 1-2 3 +-1 1 1-2 1-3 + +-1 1 1-3 +-1 1-3 1 +-1 1-2 +-1 +-1 +-1 +-1 +-1 +-1 2 1-3 1 +-1 1 .••.1 1-2 "__1 +-1 1-2 +-1 3 +-1 + 1-2 +-1 1-3 +-1 2 +-1 +-1 2 +-1 +-1 .••.1 +-1 + + +-1 +-1 + 1-2 1 1 1-3 1-2 1 1 1 +-1 1-2 +-1 +-1 + + 1-3 2 1 1 +-1 +-1 1-2 1 1-2 2 1-2 +-1 +-1 2 + +-1 1-2 +-1 1 1-2 1 1-2 2 +-1 1-3 1-2 1 2 +-1 + +-1 +-1 +-1 +-1 1-2 1-2 +-1 1-2 +-1 +-1 +-1 +-1 + 1-2 1-2 +-1 1 1 1 1-2 , 1 +-1 1-2 +-1 1 +-1 +-1 .••.1 +-1 + +-1 1-3 1 1-3 +-1 +-1 2-3 1-2 1 1-3 +-1 2 1 1 1-3 1 1-3 +-1 1-2 + +-1 1-2 1 1-2 +-1 1-2 1-3 1 1-3 + 1-2 2-3 +-1 Kuchler +-1 +-1 +-1 +-1 1-2 +-1 1 1 1-2 +-1 +-1 1-2 +-1 2-3 1 +-1 +-1 1-2 1-2 1-2 (1955). 1-2 1 +-1 + 1-2 1-2 +-1 +-1 the + phy.iosnoaic 1-2 1-2 1-2 + +-1 foraulae 1-2 1 1-2 reads 1-2 1-2 2;..3 1-3 1-2 +-1 +-1 1-2 1-2 + 1-2 1-3 +-1 1-2 1-2 1-3 +-1 1 .••.1 +-1 1 +-1 +-1 +-1 2-3 1 +-1 1-3 2-3 1 +-1 1-2 +-1 1 +-1 1-2 1-2 +-1 +-1 + left 1-3 ri8ht: 1 +-1 1 + 1-3 1-2 +-1 1 1-2 to +-1 with the most + couspicuouo 1 +-1 type placed at +-1 +-1 +-1 the 1 +-1 beginning. 2 1 1-3 + 1 +-1 +-1 1-2 1 1 �321 DESCRIPTIVE SUMMARYOF VEGETATION TYPE MAP. GREEN MOUNTAIN- BLUE GROUSE INVESTIGATIONS Legal Description: T2S. R80W. Sections 2. 3, 10, 11 County: Summit Range: 2500-2865 III Surface Area: 181.3 ha Aerial Photo No. ~ F16CN 10-3-72 Elevational VEGETATIVE CLassification Phv s Lognomf c ]j Expos ur e (Degrees Gradif-nt (Percent) Area GM-14 GM-IS GM-16 EmpDszp ElpBzpDzp [mlpDszp 35 90-190 65-120 17 12-25 35 .9 5.3 1.5 and Features Azimuth) (Hcc t a rea) Plant Species List3! Woody - Trees: Acer glabrum Juniperus virginiana Populus t reeu l otdes Pseudotsuga eena tes t i Woody- Shrubs: Cov2!Soc2! Cov Soc COY CLASSIFICATION GM-IS GM-I7 EmizpDlzp 17-30 Soc AND SUPPLEMENTARY FEATURES EmiDlszpBzp EmlDlp DzpBzpEcpDlp 35-90 Soc 3-4 3 1-4 GM-21 GM-22 DlcszpHGIc GM-23 DmczpHGlc GM-24 CM-2S DlcziHGlc DmpEmpEIRpDszp GM-26 DmiGlpHlp ClpHlpOsp 2-20 305-10 20-45 25-70 20 30-)5 30-40 322-72 15-25 34 10-20 9-15 5-16 18 18-20 11-22 6-22 2.9 2.6 2.9 4.3 .4 2.5 1 2 9 23.0 Cov VEGETATION UNIT NUMBER GM-19 GH-10 Cov 15.9 FLORISTIC CHARACTERISTICS Cov Soc Cov Soc Soc COY Soc Cov Soc Cov Soc J COY Soc Cov 3-4 2-3 1-3 1 4-5 Soc Cov Soc 1-2 +-1 3 3 1-3 1-2 1 3 Acer glabrum 1-2 1-2 Amelanchier alnifolia Arctostaphylos cve-urs r Artemisia cana 1-2 1-3 Artemisia tridentata Ceanothus velutinus Cercocarpus montanus +-1 Chrysothamnus spp. Holodiscus dumosus Juniperus cceeunt s +-1 1 Juniperus virginiana +-1 1 Mahonia repena Pachystima myrsinites +-1 2-3 Populus t reee Iot.de s 2-3 Prunus virginiana Pseudotsuga menziesii Purshia tridentata Rhus trilobata Ribes spp. 2-3 +-1 Rosa acicular is Rubus deliciosus Sambucus racemosa Shepherdia canadensis Symphoricarpos oreophilus 1-2 1-2 Tet radymia canescens Vaccinium s pp , Forbs: +-1 Achillea 1anulosa Anemone spp. +-1 2-3 Antennaria spp. AQullegia caeru Len Arabis spp. Arnica spp. Artccisia frigida Artemisia ludoviciana Aster spp. +-1 Astragalus spp. +-1 8alsru:wrhiza sagittata +-1 Ca1ochortus gunnisonii Campanula rotundHolia Castelleja spp. +-1 Chaenactls douglasii Chenopodium spp. + Cirsium spp. +-1 1-2 Clematis columbiana Clematis hirsutissima Comandra umbel1ata Crepis spp. Cryptantha spp. Cynogo1ossum officimr1e Delphinium spp. Descurainia spp • Dodecatheon pu1chel1um Eb Hob tue spp. Erigeron spp. +-1 Eriogonum spp • Erysimum upp , Fragarla spp. Frasera speclosa + Gal1um borea1e +-1 Geranium fremontll +-1 Haplopappus spp . Heuchera spp. Helenium hoopesl1 Hel1anthe11a qumquene rvt s Lporeops Ls aggregata Lathyrus spp. 1-2 1-2 Llnum lewisii Lithospermum s pp , Lupinus argentlus Mertensla spp. +-1 Oxytropis app , .Penstemon spp. +-1 Phacel1a spp , Phlox app. Potentll1a app • Primula spp , Pseudocymopterua montanus Pulsatilla patens Saxifroga b roncb La Lf s Senecio spp. Smilacina app , Solidago spp. +-1 1-2 TaraxacuCl officlnnle +-1 1 Trngopogon dublua Thnl1ctrum spp. Viola app. Gramlnoids: 1-3 +-1 +-1 1-2 3 1-3 2-3 2-3 1-3 1-2 3-4 1-3 +-1 1-2 +-1 1-3 1-2 1-2 1-3 1-3 +-1 -+-1 +-1 +-1 1 +-1 +-1 +-1 1-2 1 +-1 +-1 +-1 +-1 2-3 +-1 +-1 1-2 + +-1 +:1 +-1 +-1 +-1 2-3 1-2 1-2 1-2 +-1 1-3 1 +-1 +-1 1-2 +-1 +-1 +-1 +-1 +-1 +-1 1 +-1 +-1 +-1 3 1-3 3 1-3 2 1-3 +-1 +-1 1-2 1-2 +-1 +-1 +-1 1-2 +-1 1 2 +-1 2-3 1 1-2 +-1 + 2-3 1 +-1 1-2 +-1 2 1 +-1 +-1 +-1 +-1 2-3 3 2-3 +-1 1 1 2-3 1 1-2 2 1-2 1 1-3 1 2 1-2 +-1 1 1-3 +-1 1-2 +-1 +-1 +-1 +-1 2-3 2-3 2-3 1-2 1-2 +-1 1 +-1 +-1 +-1 2 1-2 1 1 1 2-3 1-2 +-1 1-2 1-2 +-1 1-2 1-2 +-1 1-2 +-1 +-1 +-1 1-2 +-1 2-3 +-1 1-2 +-1 + +-1 +-1 2 1-2 1 +-1 +-1 +-1 1-2 1 1 +-1 1 + +-1 +-1 2 2 1-2 +-1 1-2 1 +-1 1 2 2-3 1-3 +-1 +-1 +-1 1-2 +-1 +-1 1-2 1 1 +-1 +-1 3 1-2 1-2 +-1 +-1 + +-1 + +-1 +-1 +-1 1-2 1 +-1 +-1 +-1 +-1 1 +-1 1 +-1 +-1 +-1 +-1 +-1 +-1 1-2 1-2 +-1 +-1 +-1 1-2 2 1-2 +-1 +-1 1-2 1-2 +-1 +-1 +-1 1-2 -+-1 1-2 +-1 +-1 +-1 +-1 +-1 +-1 +-1 +-1 2 +-1 +-1 +-1 + +-1 + +-1 +-1 1 1-2 +-1 1 1-2 3 1 +-1 +-1 2-3 +-1 +-1 +-1 1-2 3 1 +-1 +-1 +-1 + + 1-2 2-3 1 3 1-2 1-3 +-1 1-2 +-1 1-3 1 +-1 +-1 +-1 +-1 1-3 1 +-1 +-1 +-1 +-1 1-2 +-1 1-2 1 l-~ 1-2 1-3 +-1 1-2 1 + +-1 1 2 1-2 +-1 +-1 + +-1 1 +-1 +-1 1-2 1-2 1 1-2 1-2 2 1 +-1 +-1 1 +-1 +-1 +-1 1-2 1-2 1 1-2 +-1 + ,1 +-1 +-1 +-1 +-1 +-1 2-3 + + +-1 + +-1 +-1 1-2 1 1-2 1-2 1-2 1-2 1-2 �322 DESCRIPTIVE S1lHKo\R't LeS.l Oe.crlptlon: !levatloMI Raose: Q}' VEGETAnON TYPE HAP, EIlIY CnD:~!lLtrt GROUSE VECtTATIV! cu.5SIFICATIott ec-i and ' •• turea Cnldlenc ..." (P.rc~I'It.' 18.0 Arca (H.cUr •• ) Ih:1Dt1pz:1HC "S., 25-)0 ••.• riable 5-28 101.8 1-' "_I 2-) 'l/t.portatt.t 1-2 2·) 1-2 1-2 1 1-2 1 1 m 10-19 13.0 '.0 ec-s n.c.:bpHC 140-215 &-)0 '.2 OlAM.C'l'!!J.ISTICS Cov Soc Cov Soc: 2-' 2-, 1-3 1-3 see Cov 1-2 2 +-1 1-2 1 2-' 1-2 +-1 +-1 1-' -I +-I + +-1 2 2-) 1-2 2-) 1-2 2 1-) +-1 1-) 1 1 +-I +-I +-1 1-2 2-) 1-2 1-2 1-' 1 2-' 1-2 1-2 1-2 2-) 1-2 1-2 1-2 1-) 2 1-' 1-2 +-1 2-3 2-' 3 2-' 1 1-2 2 2 +-1 , -I 2-3 i-j 1-3 2 +-1 1 2 +-1 2-' +-1 2-) 1-2 1-) + + 1-2 2 1-) 1 2 1 2 -+-1 _I • 1-2 2 1-2 2-) + -I 1 +-I 1 H + 1 1 + 2 2-' 2-3 +-I 2 1-3 1-2 2 + 3 -I +-1 1-2 1 +-1 2 2 1-2 +-I + + +-1 1-) +-1 1-2 1-2 +-1 1 1-) .•.•• 1 • 2 1-2 2 1 + + + +-1 1-2 , , 2-) 2-' 2 1-2 2 1 2 2 1-2 2 2 1-) +-I +-1 1 +-I 1 2-) +-1 2-3 , 2-' 1-2 +-1 +-I I 1 2-) 1 1 +-1 2-) + +-1 1-2 +-I + +-I + +-1 1 2-' 2 1-3 1 + +-I +-I 1 +-I 1 +-1 +-I 2 +-I + 1 +-I 1 1 • 1-2 2-3 +-I 1 1 2 .....1 z 1 1-2 + 1-2 +-I 1 2-'2 +-1 +-1 +-I +-I +-I 1-2 1-2 3 1 1-) 1 + -I 1 1-3 1-2 1-2 +-1 1-2 1 1 1-2 2 -+-1 -I + .....1 1-2 2 2 1 1-2 1 1 2-) 1 1-2 -I 1 1 1 1-2 1 1-2 2 +-I .....1 +-1 +-I 1-2 +-1 2 1-2 I-) 1-2 1-3 1-3 +-I 1 +-1 -I 1-2 1 .••.. 1 2 I-j 1-) +-1 + 1 1-2 + +-1 +-I 1-2 .••..1 1-2 1-2 1-3 +-1 +-1 1-2 1-2 1-2 -I 1 2-) 1-) 1-2 2 1 1 1 1-3 1-2 1-3 3 2-) , +-1 1 1-3 2-3 1 1-2 + +-I 1-' 1-] 1 1-2 1-3 1-2 1 1 1-3 1-2 +-1 1 2-' +-1 +-1 + +-1 1-2 1-' accol'ding to luc:hlu .!/c:overase and aoc:l.bllity 1-2 I, Soc 8zpOzspHC 1 &1,.... •• 11Mat,nee! Cov Cov !C-8 o.c:zllbpllCll )5-190 1-17 68.2 £C-7 EC-6 Dz:1azplUcClc 210 6-25 Bz:pOdHC 140-180 IS-30 32.7 noRlSTIC Pbnt Speclcli L1l1tl' Woody-Treeo: AInulI tcnu1foU. Junlpenll; v1rglnlonll Populull balsa.Hera Populus tre.uloldcoo Woody-Shrub.: ker glabrUIII Alnua tenuitolia "-'elanc:hier aln1(o11a Arte-iat. tridentat. cerc:ocarpu.a_ntanua Chryaothaanua SlIP. Cornua atolonifere Juniperua co-nia Juniperua virginiana Lonicera involucrau Kahanie repenll • P.chyati_ .yreiniteB 'entaphyllo!dea tloribunda Popu.lulltreauloideG Pruaua vir,ini,1n. Purahi. trident.t. R.ibellIIpp. 11.0 •• acicularia ltubua delic:io.us Salix app. Saabucua raC:t!IIIOIIO SYllPhoricsrpoooreoph1lua Tetradymla eeeceeene POl:'b.: Achille. hnulo •• Acon1tUIIcoluabianum Act.ea I:'ubra A,zaetachc app. "'oalll1:'1aglauc. A1liUIIapp. Angelic. app. Antennaria .pp. Aquileaie caerul.a Ar8bia app. M'eual:'ia congeacli Aruic. app. Artaiaie dl:'acuncu!ue Artea1aia frigid. Artl:lllilli. !udoviciana Aatreg.lu. IIpp. aala..,rhiza .a,ituu Caloehortua gu.Dniaonl1 Capllella buraa-p.atoria Cardalline cord1foUe Caatilleja app. Chenopodiuaapp. tireiua app. el_Ua app. Collinai. p8l:'viflora Collomia linearia eo.andra I,IIIbellat. Crcpia IIpp. CynoglollDU1II oUicinaIe Delphiniua app. Deacurein1a Ipp. Dodecatheonpulchdlulll EpllobiU1llapp. EquiaetUIIapp. [l'iS81:'0napp. El'1080nulluabell.CUII Erya'.apet'l,~ FregariA ovalia Fr•• er. apecio •• Gallulll opp. GeuniUII fl'ellClntl1 Geu1II app. Hackella flol'ibunda Heleni•.••hoopeaU Deli.anthena quinquenet'Vis Hel'ac.l_ lanatu. Ileu.chel:'aapp. Ipomopia .sgl't!SaU Lappula rodowaldi Lathyrua app. L1guaticUIIapp. LinUII1ew1l11 Lithoapet'1lWlI. lI.ult1florua I..o.atiumapp. Lupinu. 1I.l'8enteue Mertansia app. l;eaophlla bn~vif1oro OaIiIOrbha.IIpp. O!eypol1ahftdlel'i Pedicul.ria IIpp. PenlltClDOn app. Phacelia IIpp. Phlox IIpp. Phyoat'l. vitulitera PlaslobothrYII acopulorUII PolygonumIIpp. Potcntl11a IIpp. Ranunc:uluaapp. RIIdbackialaclniat& SCl'ophularia 1anceolata .S_cioapp. Sldalcea c:.aDd.Ua s.1lacba app. SoI1d.aso.pp. Stell.d. app. Tat'axac:ua.0fUc:inale ns.J.ictru. app. ThLaapi app. .Tt'.S0pGl0D.lIIubi",. Tt'1foliua app. Urt:l.ca IIIlo1c. V.l'atn. tenuipat.lu. Vet'onica app. \'lcu1en ••.ltiflou Viola 'Pp. Z,.aad_. el."fI. Gt'..u.oU.: Aa,l'opyl"On app. "'roatta .pp. Bro.u•• pp. CaI'Q .pp. Dac:tylia .lo.ont. De.c:Jw.paia .aeaplto •• pp. Festuca .pp. Glyced .• app. Ho1'4eu. btachyantberu. Iria ai __ riefl.ta JGDC\a8 .pp. 1oel8l'la ll'.cllta Kalka bulbo•• Phl_ pl'.t __ e Po.a.pp. Stipa .pp. RUMIll:R !C-S adOd"" 25-40 20-)0 12.4 70-260 UtlIT rc-a £C-2 BdDzl.pHC 8z1DzIHC Expo.un (OeSree, "tiauth i ruruus AND SUP'LEMENTAJl.Y VP.CP!TATIOM ~~;:~!!!:~!!? INVESTlCATIOHS TJS, R841.1, S~C. 31 T4S, 1t84W,Sec. '.S,6 Col,lnty:!.aIle 2500-2896D Surfac. Area: 482 ha Aerial Photo Mo.: CS-Ul2-1J4 7-6~SI 1-2 +-I (1955), the ph,.aiognollic: fonub ruda left to ri.ht 1-2 1 + +-1 +-1 +-1 2-3 -I with the .,.t a.81,ned .ccol'dlns to Euc:tller (I9~5). plant apec.taa on the ba81. of covar." •• Plant _cl.t",l:'. followa Vebal' (1912). 2-3 2-' 3 , 1 2 2 1-3 1 -I 2-', 1 1-2 1 1-2 2-) 1-2 1-3 1 1 +-I +-1 + 2 1-2 !==onapic:uoua type placed .t , 1-2 1 tbe bell;inniD.J • 1 I-) �323 DESCRIPTIVE legal El eve r Icna l Sl'HIWtY OF VEGETATION TYP! Dellcdptlon: Ralnge: 1)S, 2500-2896 III U4W, Sec. Surface )1; MAP, erer T4S, Area: 482 cnu-elUE R8'W. ha GROUS! INWSTI'"\TIONS Sec. 4,5.6 Aerial VEGETATIVE CLASSIFICATION 'i,le County: Photo Mo.: GS-L.H2-IJ4 7-6-51 AMI) SUPPLEMENTARY FEAT1JR£S VEGETATION UNIT MtTHIIER Clil5Sif 10n icat and Phys1olncllllc.U Exposure (Degre",s Grad lent ~rea tC-IO Fe,,' "r"~ 24-40 ~,~,.~O ____ _._ EC-12 ~6!H~C DIDc'P)~~gliCl1 ':"~{:tIu!\ (Percent) (Hectares) EC-tI tC-1) o..i~~~cCIC 10-)0 ~12~.~7 tC-I' Dt~~!~~~G 5 ~2.~4 ~~~--~~-~'2'-" Alnus "~"'So"_, 1 __ .. __ ~~ Woody-Trees: Socl. tC-16 !C-P ?i~~~g lbi~ogHG )0 ~3~.7~ n.oRISTIC 'p'~'~~~'spe~ EC-U Dl~f&HG 10-)0 ~~ 5-10 ~'~.~I £C-18 Dnf~& .••• pIP DupHG ~9-220 5-)1 32.0 ~-30 '.4 5-11 ~7~.'~ _ .. CRARACTE.tIStICS __ ~Oo""'__5o""'"_' _ _'Oo""'___'5o""'__'Oo.,v'___'S"'' '_'__''Oo'' v~_'S' ' ' __''Oo""v'__'50,",,_-,,00",,_ _-_._._-----_ Soc Soc :enuifc113 j'Jn1perusvirJ,;1nianH Populus ba18alll1fera Populus 2-) t r e ••ulcides o.:~_dy-Shrubs: seer dabrufl 3-4 2-3 2-3 2-3 •. AInu" r,,"u1(0115 Alnelan(. •.•ter aln1follt 1-2 2-3 1-2 2-1 2-3 1 Artemls:a tridentata cerccce rpce fIOntanus Cornua '1tolon1fera Junlper ••se;olllllUnis luniperus , +-1 app , Chrysothalafl"s 1-2 2-) Z +-\ \-2 J-4 1-4 , \-1 \-3 1-2 ~I virglniana Lonicera involucrata Kahonh ~I revens P.,chyst1ma ••yrsinltes f Ior Ibunde Pentaphylloides Popukue 1-2 trt!lIIuloides Prunus 1-3 vlrginiana 1-2 2 2-'1- 2-3 2-) Purshlatrldentata lUbes R.oal!.aclcular1s Rubus deUcloaua Salix spp. Sambucus 1-2 2-3 1-2 app. 1-2 l 1-2 1-2 1 2 1-2 2 Tetrad,.la rorbs: 1-2 I 1-2 2-3 +-1 1-2 I-i racemosa SYlllpnoricarpos 1-2 Achillea 14nlllosa AconitulII toluablanulII 2-) 1-2 3 oreophilus e eeeeeeee 1-3 .. H Actaearubra ~I Agastachellpp. +-1 1-2 \-2 1 Agolleria slauca Alltu. app. Angelica 2 Ipp. Antennarla Arabb 2-3 epp , Aquiles1a eaerulea IIpp. Aren.ria congesta Arnica IiIpp. Arttllli.!a 4racuncululil Arttlllis1a frlalda Arte.1sia l ••dovlclana Mtragolull !IalaalDOrhh.a 1-2 1-3 bur.a-pa.tori. Card'lain. cordifolta Castilleja app. Chenopodiu •• epp . . 1 • H ••1II IIpp. Coll1naia . 3 1 H 1 2 1-2 H · • +-1 ~I parvUlora Collo.1.a lincaris Co_ndra UIIIbellata ~I 1 • +-1 2 1 • H 1 2 _I 1-2 H 2 Ipp. CynoglollotJa 1 1 officinale +-1 DelphtniulllsPP· oescurain!a app. Dodeeatheon plllcheU Eplloblua Eq••iset Erigeron EriogonulII 2 1 1-2 ••1II app. ••••bellatulll Fragada +-1 . ova1"ia Fr.aeraspecioso GaliUIII .pp. •••• rre.ontit · +-1 1 2-3 _1 1-3 +-1 +-1 floribunda _I +-1 2 Helenl"l11 hoopeaU HeIlsnthella q ••inq ••enel'Vis Heracleua I 1 H H GeUIII app. Hae;kel1a .+ +-1 app. P;ryaiflulllt!sperUfl Get.nl \-2 .pp. ••• 1-3 1 1-2 2 2 2 Cleflll.tisspp. Crepis +-1 aunnt.onU Capllello Cirsi 1-2 .pp. lasittata Calochortu. +-1 H '-1 H +-1 1-2 2-) 1 Ianatum Heuchera app. lpolllOpia I...app-ula · ssaregata redowsltil I...athytua opp. H 1-2 L18W1t1culll app. Lim_ lewla!.i Lithoaperal •••• ltif +-1 1-2 1 1 loru. lDmati\IQOpp. Lupinus +-1 argenteus ~I Nemophila brevlflora 0slllOrh1.l:a app. 2 2 1-2 hndleri 1 Hertenaia epp , Oxypolia Peclic:ularia app. Penste.on Phacella app. app. Phlox8PP· PhY83rl. vttulifera 1-2 . H +-1 1 H 3 +-1 1-3 2-3 1 . Plaglobothrya scopulorum PolygonUll opp. Potentilla Ranunculu. app. app. 1-2 3 1-3 • +-1 1 1-2 +-1 2-) I +-1 +-1 +-1 ~I +-1 _I +-1 +-1 H +-1 ·· 2 2-3 1 2-3 1-) 1 • 1 _1 2-3 _1 2 • 1 +-1 2-3 1 + H Rudbeckia laciniata Scroph ••larla lanceolata Senecio Sidalcea spp. candida S.ilacina 2-3 Ipp. Solidago • +-1 +-1 +-1 app. Stellaria app. Tlraxac ••• officinale Thalictnl •• Thlaapi H 1 +-1 pp. 1 1-2 1 +-1 1 2 2 1 +-1 1-2 1 _I 1 1 1-3 1 1-3 1-3 1 1 +-1 2-3 1-3 + 1 1 +-1 1-2 +-1 2 _1 H 1 1 2 app. Tta80poaon Trifol1US1 +-1 1-2 dubiu8 spp. Urt1c. 410ics VerstrUII terwipetslu. Veronica spp. Viguiera Viola ••• It.iflora 1-2 spp. Zygadanu. Cr_inoi4a: Agropyron spp. , 1-2 1-3 1-2 .pp. O&ct7li. 1-2 1-2 1-2 1 1-) +-1 1-2 1 2·3 1-2 1-2 Iris 1-2 1-3 Hellt:a 1-2 1-) 1-2 1-3 1 1 1-2 1-) 1-3 2-) 1-3 1-3 spp. spp. btachyanthatua 1-2 .i.aourian.is XDel.rU 1-2 cse.pitoaa !ly.us.pp. Juncua 3 1 +-1 ala.erat. Deacl\aaplilU 'eltuea Glye.ria Botd_ • +-1 1 app. Agroatia Bromuaspp. Carex 1-2 elegans · +-1 2-3 3 .pp. 1 +-1 gracilis bulbosa +-1 1-2 Phleu •• prat.enae Poaspp. 1 1-2 Sctpaspp. l' Assigned acco"rding 1.1Coveraae snd }/ lIIIportllnt to sociability plant. apeclc, Kuchltr (l95S). IIss1gne4 on the the accord basis ur phylliogn0lll1t tng to eeveeege K••cmer . Plant 2-] +-1 1-] 1 fOnNla resds +-1 +-1 left to right ••lth the (19S5). no.enc1atU"re folIo •••• Weber (1912). -ost conaplCJo ••" type +-1 H 2 2 1-) 1-2 1 I ---plated at the beglnntna. 1-3 1-2 2 1 2 1-3 1 �324 DESCRIPTIVE Legal Elcv.1tional SUMMARY OF VEGETATION DeDe~lp(lon: Range: T35, 2500-2896 In TYPE R84W, Sec. Surbec KA.P. 31; E18Y T45, A-rea: CREEK-BLUE R84W, 482 ha VECETATIVE Sec. GROUSE INVESTICAnOSS 4,5.6 Aerial CLASSIFICATION Cla8s1f1t:ation and PhY"iogno Exposure ••1cY (Oe"r"(,8 Cradient (Percent) lu:c;:I (Hec t areu Plant Species [C-19 Features D:r.ispHC 200-220 OszpElrHC Azimuth) 21. 20-)0 6.' >.1 2 Soc- 7-6-!ll F'EATUR£S NUKIIER £C-25 EC-16 Dlllin!HC ll7 Dl?S%.pHC 175-195 £C-24 rMlliepzUbpliGll 20-79 170-190 6-29 5-1) 6-15 s .• 11.7 25.1 FLORISTIC 2 Cov- L1s~ UNIT HIICliOzp Dsp;r.i-I.pHC 1)0-246 14. 6-20 Eaglt' CS-L~1-i}4 ec-aa £C-22 DzsPlaPHC 10-25 3.' j £C-21 No.: AND SUPPLEMENTARY VEGETATION [C-20 eo",nty: Photo 10-6 6-9 .__ '_.6 ~)~.• So, Cov 2-) tcnuHolia Juniperus btlla:ullifer~ Populus trellUlo1dea 2-) 1-2 virginiana Populus n s s-s 2.> _ CKARACTERISTlCS So, ~!!!!!: Alnus EC-Z7 DupRClc 1-2 2-3 !'!WO~:~~.:f;S~:~f'!:~~:!:'~",:C-----------------------------------------------------------I,----,------------...--Alnustenuifolia II.lnifoli~ Artemisia cercocarpcs 1 1-2 1-2 2-3 1-2 1-2 1-2 2-3 1-2 2-3 1 1-2 1 1-2 1-) montnnus 2-) spp • Olrysoth4l:lJlu!l Cornus 3 tridentr.ta 1-2 1-2 2-) 1-2 ) 1-2 atolonifera Juniperus co •••.• n1!l Juniperu!l virgini~ns Um1eern involucrllt~ +-1 Hahonia r-epene Pachyatilllll lIyrainiteB Pent~phyllo1deo Populus +-I floribundo traauloidea PrunU!l 2-) 1-2 2-3 Amebnch1er Pur!lhin 2-) 2-) virginbna 2-) 2-) tridentatn '-2 Ribea app. Roan ncicubr1s 1-2 1 2 2-3 .1-2 2-) 2 2-) 1-2 1-2 1-2 1-2 RubuG deUcioBuII Tetrad)'1:lia 2-3 +-1 1-2 Salix npp , Salllbueu9 raCelllOon Syophoricorpos oreophiluo 1-) 2-) 1-2 1-2 1-2 +-I ) 2 +-I +-I Forbo: Achillea +-I lonuloon AconitulII Actaea 1 1-2 eeneeeene 1-2 +-I +-I +-1 colulllbianum rubra Agsou.chc +-1 opp. Agoserio 1-2 1-2 +-1 glaucll. AlliUlli opp. 1-2 app- Angelica Anten!Ul:ria A.quUegio Arabio Arenaria +-I spp. c.erulea spp. congeota Arnicaspp. Artee.illia dracunculus A:rte.1aia A:rtea1aill frigid. ludoviciana J.str.ga4&a klsa.orhi:.s 1-2 app. ollgittata C.lochortua 1-2 1-) 1-2 '-2 1 gunniaonU Ca.psella 1-3 bursa-pa~torio cardaldne cordifolia +-I ium, app. +-I +-I 1 Chenopod 2 +-1 2 Ciraiua npp , +-1 1-2 +-1 1-2 CaotiUeja app. CleaatiB Collinaia app. parviflora CoIIOllio Comandta lineario uClbellata Crepis 1 +-I +-I . 1-2 1 · ·· +-I +-1 app. Cynoglosll •.•• officinole DclphiniuUl 1-2 app. Deocurainia 1-2 H opp. Dodecatheon Epl10bium puichellull !lpp. EquiaetuUl Erigeron +-1 spp. opp. Eriogonu. +-I +-I umbellstwn EryoitlWll aoperua Fragario ovalia 2-3 +-l 1-2 +-1 Hackelia floribunda Heleni ••••• hoopeoU +-I Helianthello +-I 1-2 . quinquenervia lanatUIII Heuchc.ra opp. lpomopio oggregata redoW'8k11 spp. LiguoticUf!l Linwa opp. lewia11 LithoapertlUlIl lIIultitlorUlil LolID.tiulII opp. Lupinua argenteuo Mertena1.a epp. NCIIOphilo brev1floro Oomorh1:.a Oxypolle +-I +-I + 1 +-1 1-) 1 1 ·· +-I 1 +-I +-I +-I +-1 Ranunculus 2-3 1-2 1-2 +-I 1 1 1 1-2 2 +-1 1 +-1 +-1 1-2 +-1 +-1 2 +-1 1-2 1-3 +-I Senecio · app. app. app. Stellar1a . +-I app. TaraxacUfll 1-) officinale ThallctrWl Thla.pi +-I Tragopogol:l 1 • +-1 +-I +-I 1 • 1 1 .. +-l +-I l-J 1-2 1-3 +-1 1 1-2 1-2 2 +-1 +-I 1 2-3 +-1 1 . 2 2-3 1-2 1 2 2-) +-1 1 +-1 1-3 +-1 1 +-I I .pp. 2-3 dio1c. +-1 +-1 1 2-3 2-) +-1 1-2 +-I +-1 • +-1 3 +-I 2-3 1 2"-3 2 tenuip.talUII Veronica .pp. Viguiera _Itinora Viola t-a , +-1 +-I dub1u. Trifoliwa VeratrUII 1-2 +-I 2 1 1-) ) 1 2 spp. app 1 2 2-3 lonceolata candida 1 1-2 2 +-I !Jlllilacina Solidago +-1 +-I +-1 +-1 Sid.lcu 1-2 1-2 bcinU-tIl Scrophulari4 1-2 1 1-2 • +-I opp. Rudbeckb +-1 1 1-2 +-I ecopulorum Poly&onum spp. Potent111aopp. Urtic. 1 +-I +-I v1tul1fera Plagiobothrys 1 +-I +-I · +-I +-I opp. Phyoari. 1 +-1 1 epp , Phlox +-1 +-1 +-I +-I +-l +-I +-I +-l app. Phacel1a · +-I 1-2 2-3 spp. Pen.tcoon 1-2 2 1-2 • +-1 1-2 app. fendleri Pediculor1a 2-3 1 1-2 Ccum opp. Lappula Lathyrus 1-2 1 +-1 1-2 +-1 frClll(lntii Heraclew. +-1 1-2 1-2 +-1 Frooera opecioso Cel1U111 spp. CcroniulII r-a opp. +-1 1-2 +-1 1-2 1-2 Zygsdenuaelegsnll Crollinoido: Agropyron app. Agroatia Dactylis 1 1-2 1-2 1 1-3 1 2-) glo_rata Dellcbaapaia !lyaua •• 1-2 1 2 2 1 1 2-3 2 1-) 1-3 +-1 1 2 caespito 1 1-2 1-3 •• 2-) 1-) +-1 2 1-2 2 3 2-3 2 pp. .pp. HordCUII brachyantherUII .ta.ourien.i. Juncua.pp. +-1 +-1 KDeleria gracilis Mmlica bulbo ••. +-I Phleua pratenao Poa .pp. SUpa 1-2 J 1-2 .pp. Feotuc Clyceria Iris 1-2 1 1-2 opp. BrolllU' opp. Carex .pp. 1 1-2 ) 2-3 1-2 1-3 1 +-1 2-3 3 +-1 1-2 +-1 !I A.aaigned according 'Yeovcr&g. aM to IOciabllity plant species ~chler (i955). •• aigned on the the phy.iogl'101llic according baoia of 1 cover to Kuchler •••• Plant fOrall. 1 +-1 2-3 3 2 2 2 1-2 app. lllllportaat 2-3 2-) 1-3 2 r-a 1-) 1 reada left to right vith the (l955). no_nclature folloV8 W.ber (1972). _It con.plDJou. type placed at the beginning. +-1 I +-I 2 1 1-2 2 l-J �April 325 JOB PROGRESS State of Project 1980 REPORT Colorado --------~~~~~----------No. Game Bird W-37-R-33 Work Plan No. 13 Job No. Survey 7 Job Title __-=D:.:i::.:s=-t=-r=-=i.::.b.::u-=t-=i=o-=n:__::a:::n::.:d=--S=t.::a-=t.::u-=s~o:_:f:..... Period Covered: Personnel: April 1, 1979 through March 31, 1980 Mike Bauman, Clait Braun, Van Graham, Jim Hicks, Donald Hoffman, Robert Mangus, and Charles Woodward, Colorado Division of Wildlife. ABSTRACT Numbers of first-hand sightings of mountain sharp-tailed grouse (Pedioecetes phasianellus columbianus) made by Division of Wildlife personnel and members of bird clubs in Colorado in recent years have been very limited. Only nine study skin specimens specifically labeled as mountain sharp-tailed grouse were found in museum collections in Colorado, these in the collection maintained at the Denver Museum of Natural History. Data on sharp-tailed grouse maintained in Colorado Division of Wildlife files gathered in recent years is largely limited to records of a few counts of dancing grounds made by interested District Wildlife Managers in Routt and Moffat Counties and wing survey information from miscellaneous wing barrel collections. A minimum of 123 mountain sharp-tailed grouse of both sexes were counted on 15 active dancing grounds in Routt and Moffat Counties during the Spring of 1979. Eighty-two males were counted on these 15 dancing grounds for an average of 5.47 cocks per ground. Of 32 dancing grounds located and mapped pr Lo r to 1977, only nine (28.1%) are known to still be active following the Spring of 1979 census. Nine wintering areas were identified and censused during the period October through December, 1979. A total of 312 mountain sharp-tailed grouse were counted within these nine wintering areas, averaging 34.67 birds per wintering area. Twenty-one individual wintering flocks within these nine wintering areas ranged in size from one to 101 birds (average 14.86 birds). �326 Reported sightings in recent years, numbers of sharp-tailed grouse wings collected annually in wing barrels during open seasons, and findings from this first year investigation all indicate populations of this game bird are low on a statewide basis with populations being scattered over extensive areas of seemingly suitable habitats. Best present day populations are to be found within Routt and Moffat Counties where extensive coal strip mining operations and homesite developments, mostly within mountain shrub plant communities, have resulted in a vast reduction in amounts of available range. It is expected that this trend will continue in future years because of the need for energy and the resulting human population boom. �327 DISTRIBUTION AND STATUS OF MOUNTAIN SHARP-TAILED GROUSE Donald M. Hoffman Mountain sharp-tailed grouse have been hunted annually in Colorado since 1953 with season dates coinciding with those of sage grouse (Centrocercus urophasianus) and bag and possession limits in the aggregate with those of sage grouse. Very little inventory work has been accomplished on mountain sharp-tailed grouse since the period 1962-1965. This report covers the initial year of a study designed to gather and update all available information in order to assess the current status of the population. P. N. OBJECTIVE To increase the knowledge of the distribution tailed grouse in Western Colorado. and status of sharp- SEGMENT OBJECTIVES 1. Review all literature on historical distribution (location) and status (number of known activity areas) of sharp-tailed grouse in Western Colorado (west of the main eastern mountain ranges in Colorado including Saguache, Lake, Chaffee and Park Counties east of the Continental Divide). 2. Circulate questionnaires to all Division of Wildlife field personnel working or recently working in the area described (#1). 3. Circulate questionnaires to members of the Colorado Field Ornithologists, Chapters of the National Audubon Society in Colorado, and other organized ornithological clubs within Colorado. 4. Interview individuals with firsthand experience or knowledge past and present distribution and/or status of sharp-tailed grouse in Western Colorado. 5. Visit museums in Colorado to examine location lection of sharptails from Colorado. 6. Obtain and summarize all data on distribution and status of sharptails in Western Colorado from the Denver and regional offices of the Division of Wildlife, District Wildlife Managers and research personnel. of and date of col- �328 7. Identify distribution of sharp-tailed grouse in Western on topographic county and state maps. 8. Identify known status of sharp-tailed grouse in Western Colorado for leks, brood areas and winter sites by location and date. 9. Locate and map new areas of sharp tail activity identified by contacts with knowledgeable individuals and/or found during field investigations. 10. Prepare maps, summarize data and prepare progress METHODS Colorado reports. AND MATERIALS Literature searches were accomplished in libraries of Colorado State University in Fort Collins and the Colorado Division of Wildlife Research Center in Fort Collins. A personal reference library maintained in Craig was also utilized for this purpose. Questionnaires designed to obtain population information, sighting locations and names of knowledgeable individuals in regard to sharptails were circulated. Completed questionnaires were summarized and some follow-up interviews were conducted with individuals with firsthand knowledge of past or present distribution and status of sharptailed grouse in Western Colorado. Visits were made to museums maintained by four-year colleges, universities and the Denver Museum of Natural History. All specimens labeled as mountain sharp-tailed grouse were examined as to location and date of collection and the name of the collector. Where personal contact could not be made with the museum curators, correspondence was initiated to obtain this information. Visits were made to offices of the Colorado Division of Wildlife in Denver, Fort Collins, Grand Junction and Montrose to obtain management and population information on sharp-tailed grouse in Western Colorado. A spring census of dancing grounds in Routt and Moffat Co~nty was accomplished in 1979 to count males an~ total birds on known dancing ground locations and locate new grounds whenever possible. Procedures used were similar to those described by Rogers (1969). Attempts were made to locate and count sharp-tailed grouse broods along trails and roads and in potential habitats during the summer of 1979. Winter period searches of potential habitats were accomplished during the period from mid-October, 1979 through mid-March, 1980 to locate and count sharp-tailed grouse on wintering ranges. �329 Locations of all known active dancing grounds, brood sightings, winter period sightings, and miscellaneous sightings were recorded on u.s. Geological Survey topographic maps for Routt and Moffat Counties and on other suitable maps for other counties of Western Colorado. Names of individuals securing the sighting or count, number of birds, and period of the year were recorded. RESULTS AND DISCUSSION Review of Literature Historical Distribution and Status Colorado Marsh (1931) in Bailey and Niedrach (1965) stated mountain sharptailed grouse were first noted in present-day Colorado at the mouth of the Blue River, Grand County, July 29, 1839, by Thomas Jefferson Farnum, leader of an emigrant party. Also, a man named Carter collected eight specimens in Summit and Grand Counties, including a female killed near State Hill, Summit County, and a chick taken July 2, 1877 on an island in the Colorado River, Grand County. Morrison (1888) reported mountain sharp-·tailed grouse were common on the mesas and among the scrub oak of La Plata County. Gilman (1907), whose base of operation was Fort Lewis in La Plata County, noted'a few scattered sharp-tailed grouse on the mesas at about 7,500 feet elevation. Sharp-tailed grouse were reportedly residents throughout the year, with tracks being seen frequently on top of the snow three or four feet deep, and evidence of roosting in the snow. The greatest number seen was a flock of 18, with the usual number being six to ten. He reported that a nest with 11 eggs was found by W. M. Peterson. Cary (1909) stated that mountain sharp-tailed grouse were tolerably common between Hahn's Peak and Slater, Routt County. In 1907, they were tolerably common on both the northern and southern slopes of the San Miguel Mountains and in the Lone Mesa region of Dolores Coun~y. He flushed one at 9,000 feet ~levation, three miles southeast of Lone Hesa and another eight miles south of Norwood in San Miguel County. Also in 1907, mountain sharp-tailed grouse were reported in the upper pine belt near Pagosa Springs and a very few from the scattered sage parks lying between McElmo Canyon in Montezuma County and the Abajo Mountains of Utah. Cooke (1909) stated that is was ascertained that mountain sharp-tailed grouse (P. p. columbianus) was the form that occurs in Colorado west of the main-mountain range. Felger, 1910, in Bailey and Niedrach (1965), stated mountain sharp-tailed grouse were noted along the White River basin of Rio Blanco County. Frank H. Mayer (Roth, 1963) stated that sharp-tailed grouse and sage grouse were abundant during early market hunting days on the mesas and in the valleys near his camp on the Grand River (now the Colorado River) just east of its junction with the Blue River. �330 Bailey and Niedrach (1965) stated their notes made numerous references to observations of mountain sharp-tailed grouse in Grand, Routt, and Mesa Counties. The open, scrub covered slopes of the Uncompahgre Plateau seemed to be especially favorite places for these grouse. A dancing ground was found on May 18, 1937, eight miles north of Steamboat Springs, Routt County, on an open knoll in sage, within one-half mile of a sage grouse strutting ground. Eighteen males and several females were counted on this sharp-tailed grouse dancing ground. New Mexico Ligon (1927) stated the range of the mountain sharp-tailed grouse in New Mexico was very limited, being confined to the high, grassy, broken-rimmed mesas lying east and northeast of Raton. It was not considered probable that these grouse ever occupied an extensive range in New Mexico, and expressed little hope for range expansion. Oldest settlers advised they were more numerous in earlier days. As an indication of a more extensive range in earlier times, Wetmore (1936) found the distal part of a tibiotarsus of a sharp-tailed grouse in a cave near Jemez Springs, New Mexico. This cave probably was used by Indians as a temporary camp about 1300 A.D. Utah The range of the sharp-tailed grouse in Utah apparently was never widespread but populations were considered abundant in localized areas (Hart et al. 1950). Population estimates made in 1948 (Hart et al. 1950) indicated 1,500 sharp-tailed grouse wer e present in Utah, with the greatest concentration found in southern Cache Valley, Cache County, and northern Ogden Valley, Weber County. Other populations were also found in parts of Cache, Weber, Box Elder, Morgan, and Rich Counties. Destruction of habitat by cultivation, improper grazing, and burning had been the principal factors causing the decrease in populations in Utah (Hart et al. 1950). Protection from hunting since 1925 had been the principal method followed to attempt to preserve the sharptailed grouse in Utah. Habitat According to Edminster (1954), the core of sharptail habitat is brushland, the one indispensable cover type for the species. Brushland habitat must be extensive enough and well enough distributed to give sharp-tailed grouse adequate living space. Even though the sharptailed grouse is eminently well fitted to cope with a rigorous climate, its behavior is influenced in many ways by weather conditions. It resorts to tree budding when snow covers its preferred foods. Rogers (1969) stated that during extremely severe winters in northwestern Colorado, sharptails sometimes move to towns and farmyards to feed. �331 Habitat of the mountain sharp-tailed grouse in Colorado is dominated by mid and tall grasses within areas with a wide variety of browse species (brushlands), including serviceberry, sagebrush, chokecherry, oakbrush, and snowberry (Rogers 1969). The mountain shrub plant community described by Harrington (1954) was determined during this investigation to provide the primary potential habitat needed by the mountain sharp-tailed grouse in Colorado. In northwestern Colorado, these brushlands, found at elevations varying from 6,000 to 8,000 feet, are dominant in the transitions between sagebrush-grassland vegetative types in their lower limits and aspens or conifer forests at their upper borders. Past Work in Colorado Earliest studies on the mountain sharp-tailed grouse in Colorado were conducted by R. N. Randall during the period September 25, 1941 through March 15, 1942 with the average size of flocks observed ranging from 2.2 birds in September to 15.4 in March (Dargan et al. 1942). Randall recorded fall and winter period sightings in four separate areas of Moffat and Routt Counties. Estimated populations for these four areas were: Dry Fork of Little Bear Creek Fourmile Creek Long Gulch Dunkley Area 15 10 65 20 birds birds birds birds The advanced Biology Class of Moffat County High School (Paul R. Dillon, instructor) located and mapped five active sharp-tailed grouse dancing grounds in Moffat County in 1958 and 1959 as a class project (Anon. 1959). Grounds located and mapped included sites on the Pelly, Taylor, Schneider, Winfrey, and Wiseman Ranches. In 1959, 12 counts were made on these five grounds and a maximum of 76 males and 76 total birds were counted. The average number of cocks as well as total birds per dancing ground counted was 15.20. Rogers (1969) reported findings of a survey-type investigation on sharp-tailed grouse conducted during the period 1959 through 1965 under Federal Aid Project W-37-R. During 1964 he reported 28 counts were made on seven active and six inactive mountain sharp-tailed grouse dancing grounds. A maximum of 32 cocks and 36 total birds were counted on the seven active grounds for an average of 4.57 cocks per ground and 5.14 total birds per ground. During 1965, he reported that 47 counts were made on 15 active and eight inactive mountain sharp-tailed grouse dancing grounds. A maximum of 87 cocks and 91 total birds were counted on the 15 active grounds for an average of 5.80 cocks per ground and 6.07 total birds per ground. Detailed maps of all known active and inactive mountain sharp-tailed grouse dancing grounds along with suggested procedures for securing counts were recorded by Rogers (1969). �332 Rogers (1969) reported mountain sharp-tailed grouse inhabited portions of nine counties including Dolores, Gunnison, Mesa, Moffat, Montezuma, Montrose, Park, Rio Blanco and Routt during the period 1959 through 1965. They may also have been present in six additional counties during the period. Routt County had the largest sharp-tailed grouse population in 1965 and Moffat County ranked second. Questionnaire Field Personnel of Division Surveys of Wildlife Questionnaires were mailed to 154 Division of Wildlife field personnel working or recently working in counties west of the main mountain ranges. Of these, 49 (31.8%) were returned and 32 contained only negative information. Even so, valuable information was secured from the 17 which contained positive information. A limited sampling of non-responding individuals determined that most were lacking in first-hand knowledge of the sub-species. Four individuals (Robert Mangus, Charles Woodward, William Roland, and Charles Hector) were interviewed to secure information similar to that secured through use of the questionnaires. Table 1 lists locations of sightings and population information for all areas except eastern Moffat and Routt Counties secured from questionnaires and interviews. The numerous sightings and population locations secured for eastern Moffat and Routt Counties are recorded on U.S. Geological Survey topographical maps. Colorado Field Ornithologists Camille Cummings, Secretary of the Colorado Field Ornithologists, suggested seventeen members as persons who might have first-hand knowledge of sharp-tailed grouse in Western Colorado. These persons were mailed questionnaires of which eight (47.1%) were returned and all eight contained only negative information. Eight additional names and addresses of individuals suggested to contact were also mailed questionnaires. Of these, negative responses were received from six, and two were not returned. Thus, nothing of value was secured from this survey other than th~t apparently no first-hand knowledge was available from this source. Western Colorado Audubon Society Ron Lambeth, President of the Audubon Society of Western Colorado, reported that none of the members of this organization had seen sharp-tailed grouse, but that a report of a sighting on the Naval Oil Shale Reserve northwest of Rifle was received by this group. The only names suggested to contact were Glenn E. Rogers, retired, Colorado Division of Wildlife, and two biologists of the Craig Office of the Bureau of Land Management. �Table 1. Locations Counties, of sharp-tailed grouse sightings and populations, other than Eastern Moffat and Routt a secured from questionnaires and interviews of field personnel of Division of Wildlife. County Location Dolores Salter Y Area of Glade Grand Muddy Creek Drainage, Gunnison Left Hand Needle Jackson Moffat (Western) Rio Blanco of Sighting Reported Population North of Kremmling Reported exists by Mike McLain sightings Gerald Claassen Richard Hoffman David Freddy Bruce McCloskey population exists Thomas Sherrill Treasure Creek, Aga te Creek Area, NE of Sargents Remnant population exists Thomas Sherrill Taylor Park Sighting Creek, SE of Doyleville Thomas Sherrill (Tom Lines) Borquist Ranch, 1 mile S of Lake John Sighting, early 1960's Perry Olson Cold Spring Mtn. , vicinity of Utah Line Sighting, one bird, 1972 Roger Lowry One-half mile above Sighting, one bird, 1977 Charles South Fork Campground Reichert Population exists Charles Reichert (Gus Amich,deceased) Nine Mile Ridge, NE of Meeker Population exists Charles Beaver Mesa, SE of Norwood Sightings-60's & early 70' s Robert Mangus Gurley Reservoir, Sightings-60's & early 70' s Robert Mangus S of Meeker S of Norwood Iron Springs Mesa, E of Norwood aAll sightings topographical Remarks Remnant L07 Ranch, San Miguel or Population and populations maps. in Eastern Moffat Sighting and Routt Counties - early are recorded 1970's Reichert Robert Mangus on U.S. Geol. Survey \..V \..V \..V �334 Museum Specimens of Mountain Sharp-tailed Grouse All universities and four-year colleges in Colorado and the Denver Museum of Natural History were contacted during 1979 to examine locations and dates of collections of study skins of mountain sharptailed grouse. Only nine specimens, specifically labeled as P. p. columbianus were found (Table 2), these in the collection mai~tained at the Denver Museum of Natural History. The few specimens to be found in scientific collections in Colorado probably reflect the relatively low densities of this sub-species, plus little effort specifically to collect specimens. Table 2. Specimens Colorado, of mountain 1979. sharp-tailed grouse in museums in Sex Age F Mature , Location of Specimen Denver Museum of Natural History Number of Specimen Date of Collection Location of Collection 18443 10/06/1887 Routt County Egeria Park 19591 4/17/1911 Routt County Deer Park 19592 4/17/1911 Routt County Deer Park 11125 2/02/1925 Moffat County F 11126 2/01/1925 Moffat County F 28597 9/10/1954 Park County E. Buffalo Peak F 28598 9/10/1954 Park County E. Buffalo Peak F 28599 9/10/1954 Park County E. Buffalo Peak M 28600 9/10/1954 Park County E. Buffalo Peak M Information Secured from Division of Wildlife Files Personnel in offices of the Division of Wildlife in Denver, Grand Junction, and Montrose were contacted to secure available information on the distribution and status of sharp-tailed grouse in Western Colorado. Table 3 lists the limited amount of information obtained. �Table 3. Summary of information Source Denver Information Office Northwest Regional Office - Grand Junction secured for Division of Wildlife Secured Offices, 1979. Remarks Summary of Sharp-tailed Grouse Wing Surveys (Reports prepared by Clait Braun and Richard Hoffman) Years of 1976 through 1979. Report - Sharp-tailed grouse in Moffat County (Prepared by Advanced Biology Class, Moffat County High School, Paul R. Dillon, Instructor) Year 1959. Copies - Sharp-tailed grouse dancing ground forms (Submitted by District Wildlife Managers) Year 1977. Two grounds in Routt County (Green Acres and Hinkle Ranch) counted (1 count each ground) by Jim Hicks. Year 1978. Two grounds in Moffat County (Morapos Gas Field and Pellys) counted (1 count each ground) by William Roland and Charles Woodward. Four grounds in Routt County (Calif. Park Rd. #2, Green Acres, Hinkle Ranch, and Yellow Jacket Rd. #2) counted (6 counts total) by Jim Hicks and Charles Hector. Year 1979. Three grounds in Routt County (Green Acres, Hinkle Ranch, and Yellow Jacket Rd. #2) counted (1 count each ground) by Jim Hicks. Southwest Regional Office - Montrose Report of a sighting made by John Ellenberger on Blacktail Mountain, south of Steamboat Springs, (Elk count flight - helicopter) Year 1977. (Spring). Record of sharp-tailed grouse wing picked-up in wing barrel on Uncompahgre Plateau Year 1977. 1 wing. (Fall) . 5 birds. Blue Grouse Season. w w V1 �336 Except for dancing ground counts obtained by interested District Wildlife Managers stationed in Routt and Moffat Counties during the period 1977 through 1979 and wing survey summaries prepared by Project W-37-R personnel from wing samples obtained in wing barrels in Moffat and Routt Counties during the period 1976 through 1979, very little work has been accomplished with mountain sharp-tailed grouse by Division of Wildlife personnel, in recent years. Wing Collections Mountain sharp-tailed grouse open seasons have been combined with those of sage grouse and bag and possession limits have been in the aggregate for many years. While emphasis on wing collections in Moffat and Western Routt counties has been placed toward collection of wings of sage grouse during the period 1976 through 1979, wings of mountain sharp-tailed grouse were also secured from hunters. Table 4 compares numbers of wings of sharp-tailed grouse and sage grouse collected in wing barrels and at grouse check stations during the period 1976 through 1979. Numbers of wings of sharp-tailed grouse collected from hunters varied from a low of 14 in 1976 to a high of 80 in 1979 and comprised an average of 2.50 percent of the combined sage and sharp-tailed grouse wing collections for the four-year period 1976 through 1979. Caution must, however, be exercised in attempting to compare populations of these two species of grouse from wing samples secured. Many sharptailed grouse areas in Routt County were not adequately sampled and sharp-tailed grouse are undoubtedly more difficult to harvest than sage grouse. The latter is due both to the natural wariness of sharptailed grouse and the brushy habitat which it occupies, affording excellent hiding and escape cover. Also many landowners prefer that flocks of sharp-tailed grouse on their property not be hunted. Census Dancing Ground Counts, 1979 A minimum of 123 mountain sharp-tailed grouse of both sexes were counted on 15 active dancing grounds in Routt and Moffat Counties during the spring of 1979 (Table 5) (Fig. 1). Eighty-two males were counted on these 15 dancing grounds for an average of 5.47 cocks per ground. Included in dancing grounds censused in 1979 were seven new grounds located by Federal Aid Project personnel or interested District Wildlife Managers in recent years. Of 32 dancing grounds mapped either by members of the Biology Class of Moffat County High School in 1959 (5 grounds) or Glenn E. Rogers during the period 1962-1965 (27 grounds), nine (28.1%) are knoW11 to still be active, 15 (46.9%) are probably abandoned, and eight (25.0%) are unknown as to status following this initial Spring census (Table 6). Maps of all new dancing grounds have been completed for future reference. �Table 4. Year Comparison of numbers of sharp-tailed and sage grouse wings collected Moffat and Routt Counties, 1976-1979.a Season Length (days) b Number Sharptail Wings Number Sage Grouse Wings Combined Totals Percent of Total (Sharptails) during open seasons, Source of Sharptail Wings 1976 3 14 748 762 1.84 1977 7 66 717 783 8.43 44 10 6 5 1 1978 9 27 2,418 2,445 1.10 23 wings from area north of Hayden 3 wings from Moffat Co. 1 wing from Routt Co. 1979 9 or c 16 Totals 80 .3,424 3,504 187 7,307 7,494 -a/ Source 0 f·wlng co 11· ectl0ns: c/ from area north of Hayden (Routt Co.) from unknown locations from Yampa Area (Routt Co.) from Moffat Co. from Oak Creek Area (Routt Co.) (Routt Co.) 67 wings from area north of Hayden (Routt Co.) 9 wings from Moffat Co. 4 wings from Yampa Area (Routt Co.) 2.50 Average ~/Sharp-tailed 2.28 wings wings wings wings wing wing barrels and grouse check station. and sage grouse seasons were in combination with bag and possession limits in the aggregate. - Season length was 9 days for areas east of Colo. Hwy. 13 and 23 days for areas west of Colo. Hwy. 13. w w -....J �Table 5. County Summary of highest cock and total bird counts on sharp-tailed grouse dancing grounds, Moffat and Routt Counties, Spring, 1979. Name of Ground Located By Number Counts Date Highest Count (Cocks) Time Number a Counter Cocks (am) Total Birds Moffat Bear Creek Fly Creek Noland Ranch D.Hoffman D.Hoffman D.Hoffman 1 1 1 5/10/79' 5/16/79 5/17/79 7:00 6:45 5:30 D.Hoffman D.Hoffman D.Hoffman 6 10 6 8 12 8 Routt Annan's 20 Mile #2 California Park Road #1 California Park Road #2 Cottonwood Creek Elk Mountain #2 Elkhead Road #3 Green Acres Hinkle Ranch McKinney Ranch Sage Creek Salt Creek #2 Yellow Jacket ti2 G.Rogers G.Rogers C.Hector G.Rogers G.Rogers G.Rogers J.Hicks J.Hicks G.Rogers G.Rogers V.Graham G.Rogers (Reloc. by J. Hicks) 1 1 1 1 2 2 2 3 1 1 1 2 5/14/79 4/27/79 4/27/79 5/15/79 5/11/79 4/27/79 4/18/79 4/29/79 5/15/79 4/30/79 5/23/79 5/18/79 6:50 5:20 5: 10 5:50 6:30 7:00 7:30 6:45 5:20 5:35 6:45 5:45 D.Hoffman D.Hoffman D.Hoffman D.Hoffman D.Hoffman D.Hoffman J.Hicks J.Hicks D.Hoffman D.Hoffman D.Hoffman D.Hoffman -- 2 2 14 2 8 6 7 2 8 10 17 16 3 2 14 82 123 -Totals SUMMARY: Number of active grounds counted Average number counts per ground Average number cocks per ground Average number total birds per ground ~/Times are MST to 4/29/79 and MDT after 4/29/79. 21 = 15 = 1.40 = 5.47 = 8.20 6 3 6 2 -- 10 15 -- - w w 00 �Table 6. Period of Location 1959-1965 Status of sharp-tailed grouse dancing grounds, Moffat and Routt Counties, 1979. Status Active Name of Ground 1. Annan's 20 Mile #2 2. California Park Road #1 3. Cottonwood Creek 4. Elk·Mountain 112 5. Elkhead Road 1f3 6. McKinney Ranch 7. Morapos Gas Field 8. Sage Creek 9. Yellow Jacket Road 1f2 (Ground moved - relocated at 2 sites) Abandoned 1. Annan's 20 Mile Ifl 2. Dry Gulch 3. Elk Mountain III (last active in 1977) 4. Elk Mountain 113 (last active in 1977) 5. Elk River Cemetary 6. Elkhead Road III 7. Elkhead Road In 8. Foidel Creek _ 9. Mud Springs 10. Rock Creek 11. Salt Creek III 12. Schneiders 13. Winfreys 14. Wisemans 15. Yellow Jacket Road 1f2 Unknown 1. Dry Elkhead Ridge 2. Gillilands 3. Hayden Divide 4. lIes Dome Oil Field 5. Pellys 6. Smiths 7. Taylors 8. Wingates Located and/or Mapped by G. Rogers G. Rogers G. Rogers G. Rogers G. Rogers G. Rogers G. Rogers G. Rogers G. Rogers J. Hicks G. Rogers G. Rogers G. Rogers G. Rogers G. Rogers G. Rogers G. Rogers G. Rogers G. Rogers G. Rogers G. Rogers Students, Students, Students, G. Rogers G. Rogers G. Rogers G. Rogers G. Rogers Students, G. Rogers Students, G. Rogers MCHS MCHS MCHS MCHS MCHS Year(s) 1962-1965 1962-1965 1962-1965 1962-1965 1962-1965 1962-1965 1962-1965 1962-1965 1962-1965 (1979) 1962-1965 1962-1965 1962-1965 1962-1965 1962-1965 1962-1965 1962-1965 1962-1965 1962-1965 1962-1965 1962-1965 1959 1959 1959 1962-1965 1962-1965 1962-1965 1962-1965 1962-1965 1959 1962-1965 1959 1962-1965 w w \C) �Table 6. Period of Location 1977-1979 (cont'd.) Status Active Name of Ground 1. 2. 3. 4. 5. 6. 7. Bear Creek California Park Road #2 Fly Creek Green Acres Hinkle Ranch Noland Ranch Salt Creek #2 Located and/or mapped by D. C. D. J. J. D. v. Hoffman Hector Hoffman Hicks Hicks Hoffman Graham Year(s) 1979 1978 1977 1977 1977 1979 1978 W .J::- o �341 Other possible dancing grounds, which were not counted in 1979, included a site one-half mile south of the Soash Ranch Headquarters, along RC Road 46 in Routt County, where seven cocks were observed in a wheat field in April 1979 by Van Graham; the Morapos Gas Field Ground in Moffat County mapped by Glenn E. Rogers during the period 1962-1965, which repoytedly was active both in 1978 according to William Roland, and in 1979 according to Larry Osborn; a ground located by Joe Gerrans, south of Round Mountain in Routt County north of Steamboat Springs, while stationed in Steamboat Springs; and a ground located by Charles Roberts south of Yampa in Routt County, while stationed in Yampa. While numbers of dancing grounds counted in 1979 (15) was the same as the number counted by Glenn E. Rogers during the last year of his study in 1965, and the average number of cocks per ground in 1979 (5.47) was only slightly less than in 1965 (5.80), the number of grounds listed as abandoned in 1979 (15) may indicate fewer sharptailed grouse were to be found in Moffat and Routt Counties in 1979 in comparison with 1965 (Tables 5 and 6). Additional census work is needed, however, to recheck the 15 dancing grounds listed as abandoned and to check the eight dancing grounds listed as unknown status in Table 6. Wintering Period Flock Counts Wintering period searches of potential habitats "Jere started in midOctober, 1979, however, the split and combined deer and elk rifle hunting seasons extending from mid-October, 1979 through mid-November, 1979 prevented intensive field surveys in many of the sharp-tailed grouse areas due to numbers of hunters in the field. Many potential wintering areas in Moffat and Routt Counties became inaccessible to vehicular travel past mid-November, 1979 due to heavy early snowfall and lack of back-country road maintenance. Table 7 summarizes flock counts made in Moffat and western Routt Counties during the fall period of 1979 on nine wintering areas identified and counted during the period October through December. A total of 312 sharp-tailed grouse were counted within these nine wintering areas (range 4 to 138 birds), averaging 34.67 birds per area. Twenty-one individual wintering flocks within these nine wintering areas ranged in size from one to 101 birds (average 14.86 birds). Most observations (12) were secured within mountain shrub vegetative types, followed by wheat fields (8) and big sagebrush with cottonwood trees (2). Wheat fields were, however, found in the vicinity of two-thirds of the sites where wintering flocks were observed. Active dancing grounds are known for only one-third of the nine wintering areas listed. Wintering areas with no known active dancing grounds should become prime areas for intensive efforts to locate new or relocated dancing grounds during the spring of 1980. �Table 7. Sharp-tailed grouse wintering period populations, Moffat and Western Routt Counties, Fall, 1979. County Wintering Area Moffat Fourmile Creek Great Divide Long Gulch McInturf Mesa 2 2 2 Number Birds Highest Count 11/16/79 am & 12/12/79 pm 11/27/79 8:45 am-2:15pm 12/l3/79 2:15 pm 10/-/79 (M.Bauman) Vegetative Type No Mtn. shrub Wheatfield Big sagebrush w/cottonwoods Yes Yes Yes 45 No 1 101 36 Mtn. shrub Wheatfield Wheatfield Yes Yes Yes 138 No 10 Mtn. shrub No 10 Yes 10 10 25 N Yes 4 Yes No No No No 12 No Big sagebrush w/cottonwoods & wheatfield Yes 8 No 3 2 Wheatfield Htn. shrub Yes Yes 5 Yes 42 16 17 312 14.86 Wheatfield Wheatfield Wheatfield Yes Yes Yes 75 312 34.67 No Mule Creek 3 12/7/79 1:30 pm 8 2 1 1 Mtn. Mtn. Mtn. Mtn. 8 2 4 24 12/14/79 9:00 am 12/6/79 11:00 am-12:10 pm w .I:'- Mtn. shrub Elkhead Creek Active Ground 15 4 11/20/79 & 12/12/79 1:00 pm 9:15 am Total Birds No No Yes 12/5/79 10:30 am 4 Grainfield in Vicinity Mtn. shrub Mtn. shrub Mtn. shrub 6 6 3 1 Mud Gulch Totals Average 4 Date(s) & Time(s) Highest Count Milk Creek Willow Creek Routt Number Searches shrub shrub shrub shrub �343 While wintering flocks of sharp-tailed grouse appeared ~ettled into the nine wintering areas (Table 7) during the early portion of the winter of 1979-80, deep hard-packed snows had covered the grainfields by mid-January, 1980. This resulted in the grainfields becoming far less attractive to the wintering flocks and most birds associated with the grainfields dispersed. Many of these birds were observed to move back into brushland habitats during January, 1980. Flocks were observed ranging up to a distance of four miles from four of the wintering sites which were associated with grainfields, following the dispersal in January, 1980. It is not known if these were the same birds that were earlier using the grainfields since none were marked or banded, but this may have been the case. Wintering flocks not associated with grainfields usually remained within the general area but ranged over wider areas of brushland habitat in smaller groups of birds. Some of these sites increased slightly in total numbers of birds, probably resulting from birds leaving the grainfields. Reports were also received of several small flocks of sharp-tailed grouse moving to cattle feedyards during January and February, 1980. Rogers (1969) stated this behavior was associated with extremely severe winters. Deep snows, however, appear to be the rule rather than the exception within sharp tail ranges in Routt and Moffat Counties, even in winters which are near normal in amounts of precipitation and snow depth. Deep snows during the mid-winter period of 1979-80 prevented vehicular travel into most of the known wintering areas and some of these sites could be checked only with extreme difficulty using foot or snowshoe travel. Snow depths during the late winter of 1979-80 and the lateness of the snowmelt during the spring of 1980 indicated weather conditions within the brushland areas of Routt and Moffat Counties were comparable to the severe winter of 1978-79. For this reason, vehicular access into most kno~~ mountain sharp-tailed grouse areas is anticipated to be delayed until early May, 1980, similar to conditions experienced during the previous spring period. Habitat and Populations within Routt, Moffat, and Rio Blanco Counties During the initial year of this study (1979-80), field work was largely concentrated within Routt and Moffat Counties because highest populations of mountain sharp-tailed grouse are found within these counties. Limited field work was also conducted in Rio Blanco County because potential mountain sharp-tailed grouse habitat extends into this county. �344 All mountain sharp-tailed grouse dancing ground and wintering flock sites within Routt and Moffat Counties observed during 1979-80 were within the mountain shrub plant community, described by Harrington (1954), and complementary types (wheatfields and edges of big sagebrush ranges, aspens, or forested types) within the elevational zone from 6,800 to 8,000 feet with only one exception. This exception was a wintering flock site where eight birds preferred to spend at least portions of the past two winters in an atypical range consisting of big sagebrush-grassland vegetative type with a scattering of cottonwood trees at approximately 6,500 feet elevation. While the elevational belt from 6,800 to 8,000 feet appears to be the primary sharp tail range within Routt and Moffat Counties, reports in recent years indicate they may range below 6,800 feet in winter periods and above 8,000 feet in summer periods. All sightings during 1979-80 were in locations east of Colo. Hwy. 13 with the exception of several sightings in the Divide Area northwest of Craig, where preferred habitat consists of the mountain shrub vegetative type and complementary range types above the 7,000 feet elevation contour. Summer period sightings during the period 1968-70 of mountain sharp-tailed grouse were also recorded by Federal Aid Project W-37-R personnel on Isles Mountain at elevations above 6,800 feet, while searching for Hungarian partridges. Within extensive areas of seemingly potential habitat, mountain sharptailed grouse appear to be found only in isolated locations. The reason why one site is inhabited by the species and another area of habitat which appears suitable is not has not been determined. Extensive coal strip mining activities in areas south of the Yampa River and south of Steamboat Springs, Milner, Hayden, and Craig has eliminated large areas of potential range, some of which formerly contained populations of the species. Extensive homesite development of brushland acreages within Routt and Moffat Counties has also eliminated many areas of potential range. Routt County This county contains the largest population of mountain sharp-tailed grouse of any county in Colorado. Populations are known to exist within Eigeria Creek, Elk River, Elkhead Creek, Fly Creek, Foidel Creek, Grassy Creek, Little Snake River, Long Draw, Mud Gulch, Oak Creek, Sage Creek, Salt Creek, Slater C~eek, Trout Creek, Williams Fork, Willow Creek, and Yampa River drainages. Moffat County This county contains the second largest population of mountain sharp-tailed grouse of any county in Colorado within its eastern mountain shrub areas. Populations are known to exist within Bear Creek, Big Gulch, Fly Creek, Fourmile Creek, Fortification Creek, Milk Creek, Morapos Creek, Mule Creek, Putt Creek, Slater Creek, Willow Creek, and Yampa River drainages. �345 Rio Blanco County This county may rank third in populations of mountain sharp-tailed grouse based upon amounts of potential habitat. Additional field surveys are, however, needed to determine population levels. Populations of mountain sharp-tailed grouse have been reported along South Fork of the White River, Milk Creek, and the Williams Fork drainages and in the Egry Mesa, Nine Mile Ridge, and L07 Ranch areas. LITERATURE Anon. CITED 1959. Sharp-tail grouse in Moffat County. Moffat County High School Advanced Biology Class, Paul R. Dillon, Instructor. Class Report. 16p. Bailey, Alfred M. and Robert J. Niedrach. 1965. Birds of Colorado. Denver Museum of Natural History, Denver, Colorado. 1:274-277. Cary, Merritt. 1909. New records and important of Colorado Birds. Auk 26:180. Cooke, Wells W. Auk 26:411. 1909. The birds of Colorado range extensions - Third Supplement. Dargan, L. M., R. J. Keller, H. R. Shepherd, and Sharp-tailed grouse studies, Craig Area, in Report. Federal Aid Project Colorado 4-R. Fish Department, Denver, Colorado 4 (May). R. N. Randall. 1942. Sage Grouse Survey Colorado Game and lOp. Edminster, Frank C. 1954. American game birds of field and forest. Charles Scribner's Sons, New York. pp. 135-167. Felger, Alva H. 1910. Colorado Notes. Auk 27:89. Gilman, M. F. 1907. Some birds of southwest 152-158; 194-195. Colorado. Condor 9: Hart, C. M., O. S. Lee, and J. B. Low. 1950. The sharp-tailed grouse in Utah, its life history, status, and management. State Department of Fish and Game: Pub. No.3. 79p. Harrington, H. D. 1954. Manual of the plants of Colorado. Books, Denver. 666p. Utah Sage Ligon, J. S. 1927. Wildlife of New Mexico, its conservation and management. New Mexico Department of Game and Fish, Sante Fe, New Mexico. pp. 125-127. �346 Morrison, Charles F. 1888. A list of some birds of La Plata County, Colorado, with annotations. Ornithologist and Oologist 13:139. Rogers, Glenn E. 1969. The sharp-tailed grouse in Colorado. Colorado Division of Game, Fish, and Parks, Denver, Colorado. Tech. Publ. No. 23. 94p. Roth, Charles. 1963. 12(6):30-39. Diary of a market hunter. Colorado Outdoors Wetmore, Alexander. 1936. The range of the sharp-tailed New Mexico. Condor 38:90. Prepar ed by _~4o.==-(f-;,:":,t,-,-a/."",-,d,,,"-_'()1~-,--_:Jvt.;__,;_'O--~·ItF1~J-· Donald M. Hoffman l~ Wildlife Researcher C ,-,/)_:'..=:a~r<_/ ( __ ,Ad! / ) grouse in �w ~ --..J . ,. Fig. 1. A male sharp-tailed grouse on transient dancing ground used because of lateness of snowmelt, Moffat County, May, 1979. (Photo by D. M. Hoffman.) ��349 April 1980 JOB FINAL REPORT State of Project Colorado --------~~~~~----------NO. Work Plan W-37-R-33 Game Bird Survey 21 ---------------------------- Job No. 1 Job Title __~M=i=n=i=m~u=m=_T~~=·I=I=a~g=e~T=e=c=h=n=i~q~u~e=s~f~o~r~E~s~t=a=b=I=~=·=sh~i=n~g~S=h=r _ in Clump Plantings Period Covered: Personnel: April Howard 1, 1979 through D. Funk and Warren March 31, 1980 D. Snyder ABSTRACT Writing of a final report for this study to be published as a Division Special Report was completed along with preliminary editing and revision. Final editing and publication will be completed under Project . W-37-R, Work Plan 22,Job 1. The approximate citation for this publica~ tion is as follows. Snyder, W. D. 1980. shrub thickets. Minimum tillage techniques for establishing Colo. Div. of Wildlife Special Report. Findings from this study also are being incorporated in a comprehensive habitat modifications manual currently being prepared by the Division.' Prepared by -'=~=-"'4•.•k<!.4~,<..::,""--,,Lf::"·_-lJ~· ':..L;~~' 1-'La:e-~'_;_/-,-----,-. Warren D. Sny{i;r (~ Wildlife Researcher C ) ��April 1980 351 JOB PROGRESS State of Project REPORT Colorado No. W-37-R-33 Work Plan No. Game Bird Survey 22 Job Title Job No. 1 Upland Game Publications Period Covered: Personnel: April 1, 1980 through March 31, 1980 C. E. Braun, K. M. Giesen, A. G. Haskins, O. W. Olsen, and R. M. Stabler. N. J. Kitzmiller, ABSTRACT Publications planned are as follows: for and accomplished under this job for Segment 33 Giesen, K. M., and C. E. Braun. 1979. Nesting behavior of female white-tailed ptarmigan in Colorado. Condor 81:215-217. ________ , and Colorado. ~-1979. Renesting Condor 81:217-218. of white-tailed ptarmigan in Stabler, R. M., A. G. Haskins, N. J. Kitzmiller, O. W. Olsen, and C. E. Braun. 1979. Two new species of coccidia, Eimeria leucuri and E. oreoecetes (Protozoa:Eimeriidae), in grouse from Colorado. J. Parasitol. 65:272-274. i\, 2><(,.'~..b.e\ c 7 Prepared by :,~ :{./t·.(]'--! Howard D. Funk Section Chief Small Game Research .. :.b � https://cpw.cvlcollections.org/files/original/1a520333be92947cf1d68f83a9e75cae.pdf 74f2629ff15fedb55673b60a80c97d6a PDF Text Text July, -1- 1980 . JOB FINAL REPORT State of Colorado Project No. W-125-R Deer-Vehicle Accident Investigations Work Plan No. 1 ----------------- Deer-Vehicle Accident Investigations Job No. 1, 2 , 3, 4, 5, and 6 and Methods and Devices Period Covered: Personnel: January Deer Vehicle Accidents Statewide to Reduce Them 15, 1968 - November 11, 1979 Dale F. Reed, Bruce C. Giunta, Jerome Claudia A. Doose, Charles L. Merrell, T. Myers, William B. Zimmerman, James McDonnell, Kenneth R. Kincaid, Thomas Thomas D. I. Beck. J. Cebula, Dennis L. Money, Kenneth C. Dillinger, Gary D. Fleming, Sharon L. M. Pojar, Thomas N. Woodard, ABSTRACT Methods, devices; or structures related to reducing the number of deer-vehicle accidents were evaluated or experimentally tested after obtaining preliminary data on study areas and methodology. These methods, devices, or structures were highway lighting, underpasses, overpasses, 2.44-m fences and one-way deer gates, and deer guards. During the highway lighting study, 84 deer-vehicle accidents occurred, 45 and 39 with lights off and on, respectively. Behavioral responses of deer to the Vail deer underpass did not change substantially over the 10 years (1970-1979) of study. During the overpass studies, including experimental tests of bridge width and an overhead netting, 570 crossings were examined by video replay. An average of 78.5 percent fewer accidents occurred adjacent to sd.x 2.44-m fences after installation of the fences. Use of one-way deer gates located in four of the six fences was variable. Five deer guard prototypes were evaluated. Highway lighting, as tested in this study, did not result in significantly fewer accidents. Observations indicated that strategically located underpasses and overpasses with acceptable dimensions and characteristics were effective in providing deer with relatively safe passage. Adequately constructed and regularly maintained 2.44-m fences and one-way gates were efficacious in modifying deer movements or keeping deer off highways. Of the deer guard prototypes tested in this study none were effective in precluding deer movements. ��-3- DEER VEHICLE ACCIDENTS STATEWIDE AND METHODS AND DEVICES TO REDUCE THEM Dale F. Reed P. N. OBJECTIVE Locate and examine potentially critical deer vehicle accident areas in Colorado and recommend methods or structures which may reduce deer vehicle accidents in these areas. Subsequently, measure the effects of methods recommended and investigate deer responses to various experimental structures. SEGMENT OBJECTIVES Complete project-end data analysis and prepare for publication and for the job final report. All other sections Prepared by of this report are covered C·· /.i AI iQ 4(/:<, J Dale F. Reed Wildlife Researcher C and submit manuscripts in the following report. �-4- TABLE OF CONTENTS INTRODUCTION . 5 STUDY AREAS 7 Underpasses and Overpassn • Deer Guards • Deer Fence Length • Highway Lighting Animated Deer Crossing Sign METHODS 7 8 8 9 9 9 Underpasses and Overpass. . Deer Guards • Deer Fence Length . Highway Lighting Animated Deer Crossing Sign • 9 10 10 11 12 12 DATA ANALYSIS Underpasses Overpass. Number of crossings Crossings: approach ratios Duration of hesitations and crossings Behavioral responses: crossing ratios Deer Guards . Deer Fence Length • Highway Lighting Animated Deer Crossing Sign • DISCUSSION AND RECOMMENDATIONS . Underpasses • Overpass. Deer Guards Deer Fence Length . Highway Lighting Animated Deer Crossing Sign . 12 13 13 14 14 15 16 16 17 18 18 18 18 19 19 19 20 20 CONCLUSIONS . 21 LITERATURE CITED • 22 TABLES 1-19 25 FIGURES 1-9 45 ACKNOWLEDGEMENTS �-5- REGIONAL FINAL REPORT DEER-VEHICLE ACCIDENT Dale F. Reed, Thomas N. Woodard, RESEARCHa and Thomas D. I. Beckb TNTRODUCTION The problem of deer (Odocoileus spp.)-vehicle accidents has been documented (Thompson 1967, Puglisi et al. 1974). Besides the loss of the biotic resource, considerable personal property damage is incurred (Woodard and Reed 1974). The problem arises when highways are constructed through habitats where deer are concentrated or when highways bisect deer migration routes. Successful attempts to mitigate this problem were not initiated until the 1960's (Pojar et al. 1972, Reed et al. 1975). Some techniques such as reflectors have not been successful (Gordon 1969). Other techniques have produced contradictory results. Mansfield and Miller (1975) concluded that 76- x 76-cm symbol-type warning signs reduced deer-vehicle accidents in 11 of 19 study areas in California. Pojar et al. (1975) found that lighted, animated deer crossing signs did not significantly reduce deer-vehicle accidenbs in Colorado. This study utilized an evaluation technique which eliminated biases due to annual variations whereas the California study did not. This may explain some of the differences in results. Muller (1967) found that 80 percent of vehicle accidents involving game in Switzerland occurred from sunset to sunrise. More than 92 percent of a sample of 1,441 deer killed on Colorado highways were killed from 1700 to 0900 MST (Myers 1970). It is plausible that such accidents could be reduced by use of highway lighting which would increase motorist response and decrease the number and severity of deer-vehicle accidents. Helms (1969) discussed factors which affect visual acuity. Brightness is equated with light reflecting from the object. Excessive brightness is detrimental to visual acuity. Contrasts in seeing surface detail and in outline delineation are important. Farber et al. (1971) further discussed the complexity of visibility on illuminated highways. Factors which interact to affect visibility, contrast of a target object with its surround, include amount and uniformity of fixed illumination, amount of vehicle lighting, target object's location in relation to fixed illumination sources and vehicle, pavement and object reflectance properties, and level of disability glare. Gallagher et al. (1972), after a review of aCovers a period from November 11, 1974 to November 11, 1979. The contents of this report reflect the views of the authors who are responsible for the facts and the accuracy of the data presented herein. The contents do not necessarily reflect the official views or policies of the Federal Highway Administration. bAll were wildlife researchers with the Colorado Division of Wildlife. �-6- pertinent literature, concluded that urban roadway lighting reduced occurrence of the more serious accidents, especially those involving pedestrians. Reductions from 30 to 80 percent were found. Rumar (1975) indicated many investigations demonstrated roughly 30 to 65 percent accident reduction under road lighting conditions. There has been no research to determine effect of highway illumination on occurrence of deer-vehicle accidents, but factors involved in pedestrian-vehicle accidents are similar. Installation of fixed illumination would enhance night visibility of the motorist and possibly reduce occurrence of deer-vehicle accidents. Fences that restrict deer from getting onto highways may be used when other methods are not effective. Thompson (1967) reported that the use of fencing for critical deer-vehicle accident areas increased in 1967. Various types of fencing have been used. Puglisi et al. (1974) reported on the relationship of deer-vehicle accidents to four types of fencing installed adjacent to Interstate 80 (1-80) in Pennsylvania. The fences were 1.2, 1.5, 1.7, and 2.3 m high. The greatest number of deer-vehicle accidents occurred adjacent to the 2.3-m fence, apparently because this type of fence was installed in areas where the most critical deer-vehicle accident areas occurred. Falk (1975, unpublished data, Pennsylvania State University, University Park, Pennsylvania) observed deer jumping through this fence between wire strands near the top. In another study fences 2.13 m in height along the New York Thruway reduced collisions with white-tailed deer by 44.3 to 83.9 percent in the fenced areas and 12.9 to 24.7 percent beyond the ends of the fences (Free and Severinghaus, undated). More recently, l.vardet al. (1979) reported on 12.6 km of 2.44-m fencing on both sides of 1-80 in Wyoming. While some fences prevent animals from going where not desired, a problem arises when it is necessary to permit vehicle access through the fences. When gates hinder vehicular traffic flow, structures such as modified cattle guards have been used and recommended. The physical requirements of guards to preclude deer crossings have not been established. Used primarily in conjunction with sufficiently high and adequately maintained fences, underpasses and overpasses may provide alternatives to deer from getting onto highways and from having their movements and migration disrupted. Child (1974) reported on the reaction of caribou (Rangifer tarandus) to simulated pipeline over-head structures. Mansfield and Miller (1975) reported light deer use through 18 highway metal pipe culverts and a 1.83- x 1.83- xIS. 24-m (height x width x length) concrete box underpass. 2.44-m fencing was not used in conjunction with these structures. A study of a highway underpass (Reed et al. 1975) indicated a reluctance of deer during use. It has been surmised that overpasses (no substantial overhead structures) would be used more readily by deer. Klein (1971) reported on bridges used by reindeer (Rangifer tarandus) in Norway. The width of the bridge was reported to be dependent upon the size of the herd to be moved (i.e. 10-15 m wide for herds numbering up to 2000), since reindeer tend to bunch up when crossing and may force each other over the edges if the bridges are too narrow. Bridge surfaces were covered with soil. Location was critical to the success of the bridges. Child (1974) reported on caribou response to two gravel ramps of 30.5-m and 22.9-m lengths and 2.4-m heights over a simulated pipeline. The dimensions and design of underpasses and overpasses that are most effective for wild ungulate use are largely unknown. �-7- The purpose of this study was to evaluate and test the effectiveness of methods, devices, or structures related to reducing the number of deervehicle accidents. Consistent with this purpose was the need to locate and examine potentially critical deer-vehicle accident areas and recommend methods or structures which could have reduced deer-vehicle accidents in these areas. Subsequently, measurement of the effects of methods recommended and investigation of deer responses to various experimental structures was to be conducted. Specifically, methods, devices, or structures outlined in the study proposalc were as follows: A. Underpasses and Overpasses B. Deer Guards C. Deer Fence Length D. Highway Lighting E. Animated Deer Crossing Sign STUDY AREAS Underpasses and Overpass The underpass and overpass studies were located in several areas in western Colorado. Reed et al. (1975) described the study area of the Vail deer underpass. Eleven other underpass structures were monitored for deer use during various periods of time. Their approximate locations are as follows: Avon 0.5 km east of Avon Interchange, Eagle East I 5.2 km east of Eagle Interchange, 1.... 70 Eagle East 2 Eagle West ld 3.7 km east of Eagle Interchange, 1-70 3.2 km west of Eagle Interchange, 1-70 Eagle West 2d 5.5 km west of Eagle Interchange, 1-70 Mamm Creek 6.4 km east of Rifle Interchange, 1-70 Dry Creek 3.2 km east of Rifle Interchange, 1-70 Arch Deer d 10.6 km west of Hesperus, U.S. 160 Salida I 6.0 km southeast of Howard, Colorado 291 Salida 2 6.6 km southeast of Howard, Colorado 291 Chaffee Gulch 13.7 km north of Ridgway, c Attachment to the Contract Agreement vehicle Accidents Project HPR-3(3). dConstructed 1-70 specifically for a Cooperative for deer use. U.S. 550 study of Deer- �-8- The overpass study was conducted 7.5 km west of Vail adjacent to 1-70. The structure spans Gore Creek and was used by vehicles before the completion of 1-70. It was located just north of the 1-70 right-of-way on the Dale Mikkelson property. Approximately 4.8 km of associated 2.44-m fencing generally parallels the highway in both directions east of the overpass. Deer apparently move around the west ends of this fencing and cross the bridge overpass rather than Gore Creek. Deer Guards Two deer guards were installed in 2.44-m fences, one in the fence adjacent to Interstate 1-70 near Avon, Colorado and the other in a Bureau of Land Management wildlife exclosure fence at Trail Gulch·between Dotsero and Burns, Colorado. Both were monitored for deer use under field conditions (Reed et al. 1974b). The ~uard at Trail Gulch was used for controlled tests (prototypes I-V). Deer Fence Length The 2.44-m fencing study areas were located in six areas between Vail and Aspen. They were the Vail, Avon, Edwards, Eagle, Diamond S, and Carbondale 2.44-m fences. The Vail 2.44-m fence was located between the west Vail Interchange and somewhat east of the Dowd Junction bridge. The 5.6 km section of interstate that includes the fences and associated deer underpass was accepted for completion by the Federal Highway Administration on August 26, 1970. The location utilizes well-established deer migration trails and a natural drainage referred to as Mud Springs Gulch. Approximately 4.8 km of fencing generally parallels the highway in both directions from the underpass. The fencing on both sides of the highway joins chain-link fences near the West Vail Interchange. One-way deer gates were strategically located in the fences (Reed et al. 1974a). The Avon study area consisted of a segment of the interstate from the Avon Interchange east 3.6 km to the Eagle River Bridge and was opened to traffic October 1, 1971. The 2.44-m fence was completed along the north highway right-of-way on October 5, 1972. Open sagebrush areas and various browse species are common on the north side of the highway. Alfalfa fields are prevalent south of the b Ighway,. Deer inhabit this area from August until snow depth precludes use, usually by late December. The Edwards study area consisted of a segment of the interstate from the Edwards interchange west 3.6 km and was opened to traffic October 1, 1971. The 2.44-m fence was completed along the north highway right-of-way in July, 1972. Deer utilize the area adjacent the highway to the north during the winter. Sagebrush flats and pinyon-juniper communities are common. The Eagle study area consisted of a segment of the interstate from. near the Eagle Interchange east 7.7 km. The interstate highway was opened to traffic October 5, 1972. The 2. 44-m fence was completed along the north side of the highway right-of-way October,S. 1973. Alfalfa fields are prevalent south of the highway. Pinyon-juniper and some big sagebrush occur on the north. �-9- The Diamond S study area consisted of a 4-lane 1.8 km long segment of Highway 82 adjacent to the Diamond S Ranch, approximately 1.6 km northwest of the junction with Highway 133. A field of crested wheatgrass (Agropyron desertorum) is located on the east side of the highway and is one of the earliest grasses to green up in the spring' (Reynolds and Springfield 1953). Sagebrush is abundant around the perimeter of the crested wheatgrass field and is replaced by pinyon-juniper type as elevation increases to the east. Alfalfa fields line the west side of the highway. Deer concentrate in the crested wheatgrass field and sagebrush east of the highway in late winter and early spring. The Carbondale study area consisted of a segment of Highway 82 from about 1.3 km southeast of the junction with Highway 133 to about 0.2 km up Crystal Springs Road. The 2.44-m fence was completed along the north side of the highway right-of-way October 17, 1974. Alfalfa fields are prevalent north of the highway with big sagebrush occurring on an abbreviated south facing slope behind the fence. Highway Lighting The highway lighting study area was located 4.8 km south of Glenwood Springs, Colorado on State Highway 82. The 1.2 km segment of highway had 4 lanes and a posted speed limit of 88.5 km per hour. The average daily traffic volume within 4.0 km of the study area was 5,706, 5,111, 6,221, and 6,483 during the 1974, 1975, 1978, and 1979 January-March periods, respectively. Deer generally winter in the vicinity of the study area from mid-January to late March. A more extensive description of the study area was provided by Reed et al. (1977) and Reed and Woodard (1981). Animated Deer Crossing 'Sign The animated deer crossing sign study area encompassed the same area described above for the highway lighting study. The 2.4 km segment of ~ighway had a posted speed limit of 96.5 km per hour. The average daily traffic volume within 4.0 km of the area was 4,283 and 4,836 during 1972 and 1973 January-March periods, respectively. A more extensive description of the study area was provided by Pojar et al. (1975). METHODS Underpasses and Overpass The methods used in studying the Vail deer underpass were described by Reed et al. (1975) and Reed (1981a). The eleven other underpasses were checked for deer tracks periodically throughout the year and weekly or more often during deer concentration periods. Trackbeds were maintained when possible by raking the soil at the entrances' of the structures. Only one overpass became available for study. On June 7, 1974 it was discovered that deer were using a sub-standard bridge, 3.2- x 4.9- x 13.4-m height (underbridge clearance) x width x length (direction of traffic), over Gore �-10- Creek. Approaches to the overpass were checked for deer tracks and a video time-lapse surveillance system (Reed et ale 1973) was used to record imagery of crossings and overt behavioral responses during spring-summer (June-July) and fall (October-November) migration periods. During 1976 the bridge was modified (deck removed and re-built) so the width of the structure could be varied. It was designed to have the width (control = 4.93-m, variable = 2.48-m) (Fig. 1) changed for alternate three day periods. During 1978 an overhead netting, supported by arched plastic tubing (Fig. 2), was designed to be assembled (variable) or disassembed (control) for alternate three day periods. The number of video-recorded deer approaches,entrances, exits, and behavioral responses including muzzle-to-ground (Reed et ale 1975:366). hesitation (cessation of forward movement for 1.0 second or more), and crossing mode (walk, trot, or bound) were tallied during video tape replay. Deer Guards The first guard design (prototype I) utilized 3.05- x 3.66-m guard sections constructed with flat mill steel 1.3- x 10.2-x 304.8-cm (width x height x length) rails. A 3.05- x 17.98-m (width x length) runway was constructed with 2.44-m fencing attached to one end of the guard at Trail Gulch. The test involved releasing deer in the runway and observing their response as they attempted to escape via their only exit across the guard (Reed et ale 1974b). Other guard design prototypes (II, III, IV, and V) were tested using the same location and methodology. Prototype II was an alternate black and white pattern with 30-cm long alternating black and white sections painted on each rail of the first 3.05- x 3.66-m section of the prototype I guard. Prototype III was rubber tubing (five large tire innertubes cut and sectioned longitudinally to form elongated rectangles when stretched) stretched across and 15 cm above the prototype II guard. Prototype IV was rubber straps (93 tenspeed bicycle tire tubes) stretched parallel and next to each other across and 15 cm above the prototype II guard. Prototype V was a black and white scintillation or "ray pattern" (Teuber 1974:104) originally developed by Mackay (1957) painted on a 3.05- x 3.66-m tarp and placed over the prototype I guard at the end of the runway. The principal behind the use of this design was the stimulation of the line-detecting mechanism of form perception which causes a scintillation effect in human vision (Teuber 1974:104). It was hypothesized this effect would be similarily detected by deer and averse reactions potentially elicited. Deer Fence Length Deer density adjacent to the 2.44-m fences except for the Vail fence was estimated using the method described by Reed (1969). Locations of vehicle-killed deer were documented in relation to quarter-mile markers or known structures for all the fences. Track counts were made in the median of Highway 82 adjacent to the Diamond S fence. Conditions were normally favorable for maintenance of soil in the median during March, April, and May when most of the deer activity occurred. Deer were marked with numbered neck bands or automatic tagging devices (Siglin 1966) at the Vail and Avon fences. The automatic tagging �-11- devices were often placed on one-way deer gates (Reed et al. 1974a) located in the 2.44-m fences. A drop-net was used in 1971 to capture deer when they frequented areas behind the 2.44-m fence that lead to the Vail deer underpass. Clover traps (Clover 1956) were used to trap deer on the winter range adjacent to the Edwards and Eagle fences. A Cap-Chur gun using succinylcholine chloride Pneu-Darts was used to capture selected animals. Radio-tracking transmitter collars (Model MK 3, Telonics, 1300 W. University, Mesa, AZ 85201), were placed on does trapped or captured behind the 1-70 Eagle fence. Attempts were made to locate these animals by establishing bearings from several prominent observation points. These bearings were plotted on copies of a U.S. Geological Survey quadrangle map (scale: 1:24,000). Telemetered animal locations were estimated by determining the area where two or more bearings converged. Locations or sightings of marked deer in the vicinity of the 2.44-m fence were used to estimate their movements in relation to the fence. One-way deer gates (Fig. 3) were located in the Vail, Avon, Edwards, and Eagle 2.44-m fences. These gates were checked periodically for passages and activity by checking tracks in the raked soil at the gate entrance and exit. The methods used in calculating benefit-costs methods was described by Reed et al. 1981. of fencing and other Highway Lighting Thirteen, 37,OOO-lumen, 700-watt, clear, mercury vapor luminaires (lamps) mounted on 3.05-m arms at the top of 12.2-m metal poles were used to light the highway. Nine lamps were spaced at approximately equal intervals (59.2 to 68.9 m) to illuminate about 0.50 km of highway (full lighting). At the ends of the full lighting were areas of transition lighting, created by additional lamps spaced approximately 119 m and 302 m from the last full lighting lamp (Fig. 4). The lights were alternately turned on and off for one week periods during January through March or into April of 1974 through 1979. Horizontal illumination levels were measured with a General Electric SL480A light meter. Luminance measurements were taken at sites where the accidents occurred. Luminance values (foot-lamberts [fL]) were recorded with a spotmeter (Spectra Model UBA) (Fig. 5), and were taken on a target at the kill site and on the background to approximate the view of an approaching motorist when the lights were on. The target was a simulation of a female mule deer (transverse section of full taxidermy mount). Target readings were taken from a height of 1.3 m and a distance of 15.0 m, resulting in a measurement area diameter of 26.0 cm midway between the shoulder and hip. Background readings were taken at the same height and a distance of 60.0 m, resulting in a measurement area diameter of 102.0 cm. The taxidermy mount was removed during this background measurement. Measurements were taken between 0300 and 0600 MST to minimize traffic interference. No measurements were taken when vehicle headlights or other spurious light sources were present. Background luminance (Lb) and target luminance (Lt) measurements were transformed into visibility indices (VI) by the following equation (Gallagher and Meguire 1974): �-12- VI where C C (RCSLb) (DGF) 5.74 Lt-Lb Lb Relative contrast sensitivity background luminance. and DGE Disability glare factor for the recorded 1.0 RCS values were obtained or extrapolated from published tables and Meguire 1974, Technical Committee of the CIE 1972). Additional methods used were described and Woodard (1981).· Animated Deer Crossing (Gallagher by Reed et al. (1977) and Reed Sign The lighted, animated deer crossing sign (Fig. 6) had a reflectorized yellow, diamond-shaped background (1.83 x 1.83 m) with four silhouettes of deer made of neon tubing lighted in sequence from right to left across the sign. Two signs were constructed so they could be easily swung away from approaching traffic. In the on position, the signs were locked into place facing traffic and the neon lights were activated. The signs were turned on and off for alternate weekly periods during January-March periods. Other methods were similar to those used during the highway lighting tests and are described further by Pojar et al. (1975). DATA ANALYSIS Underpasses Data analysis covering the Vail deer underpass was reported by Reed et al. ( ~7))a d Reed (1981a). Eleven other underpasses were checked for deer use during the period covered by this report. Numbers of deer passages recorded varied considerably (Table 1). Moderate numbers (30-90) of passages occurred per annum through the Avon underpass since deer use of the structure was discovered on October 25, 1975, except for 1979. During 1979 the construction of the Avon airport and related buildings may have precluded regular use of the structure. Observations of a neck-banded doe (No. 27) indicated that most passages occurred as a result of nightly use of the structure by a small sub-group of deer. Deer passages through a small structure such as this are probably limited to few deer and a low percent of the nearby population. Of those structures under 1-70, only three (Vail, Eagle West 1, and Eagle West 2) were constructed specifically for deer use (Table 1). The reluctance of deer at the Vail structure has been discussed (Reed et a1. 1975' and Reed 1981a). Such reluctance did not appear to be present at the Eagle West structures. Trails established by deer passage through the underpasses were distinct and there was no "milling about" detected at the entrances. �-13- The primary stimulus of a given underpass structure to approaching may be termed the "openness effect". Calculated as follows: height x width (or open-end length deer surface area) the openness effects of the Vail and Eagle West underpasses were 0.31, respectively (Table 1). There are several factors to consider before relating openness effect to deer behavioral response. Additionally, any reasonable attempt to relate openness effect to deer use must consider deer density and motivation at the structure. Statistical analysis of openness effect as related to deer passage success (i.e. regression analysis, etc.) would require more data points than obtained in this study. 4.57, and 5.57 (metric measurements), Overpass The video time-lapse surveillance system was operated at the overpass 1) during four seasons, fall 1974, spring-summer and fall 1975, and spring-summer 1976, to collect pre-experimental crossings and behavioral data, 2) during four seasons, fall 1976, spring-summer and fall 1977, and spring-summer 1978, to collect experimental width crossings and behavioral data, and 3) during two seasons, fall 1978 and spring-summer 1979, to collect experimental overhead netting crossings and behavioral data. A total of 570 crossings were examined during video replay (Tables 2-4). Number of Crossings There were 329 deer crossings over the bridge during the pre-experimental period (Table 2). For purposes of examining variability in the number of crossings during alternate three-day periods (control-variable scheme to be used in experimental tests), these data were categorized into alternate three-day periods and tested for independence. They were significantly different X2 = 19.6, df = 4, P < 0.005). During the experimental width tests (control width = 4.93 m, variable width = 2.48 m), more crossings occurred under control than variable width during each of the four seasons (Table 3). A test for independence shows that the difference between the number of control and variable crossings during the alternate three-day periods was not significant (X2 = 8.0, df = 4, P > 0.05). During the experimental overhead netting tests (control = netting disassembled, variable = netting assembled), more crossings occurred under control than under variable during each of the two seasons (Table 4). A test for independence shows that the difference between the number of control and variable crossings during the alternate three-day periods was not significant (X2 = 1.4, df = 3, P > 0.50). The difference between the number of control and variable crossings.in the pre-experimental seasons was sufficient to indicate that the variability �-14- was due to number of animals within periods rather than the structure. Basically, the number of animals within periods was independent (significantly different) before the experimental tests and dependent (not significantly different) during the experimental tests. It is uncertain whether this dependence can be related to reluctance of deer to use the narrower bridge or the bridge with overhead netting. Crossings:approach ratios During the experimental width tests, fewer approaches of both types (Table 3) occurred during control (15) than during variable (35). The control crossings per approach ratios were significantly (p < 0.025) greater than that for variable. Generally, this would be expected if there were a greater reluctance of deer to cross a narrower structure (i.e. more reluctant animals either approached and failed to cross, or made more than one approach before crossing). During the experimental overhead netting tests, more crossings and approaches (Table 4) occurred during control (43 and 24 respectively) than during variable (18 and 17 respectively). However, variable crossings per approach ratios were not significantly (P > 0.50) smaller than that for control. Generally, it would be expected that the variable ratio would be smaller if there were a greater reluctance of deer to cross under a net arch (i.e. more reluctant animals either approached and failed to cross, or made more than one approach before crossing). The importance of ratio differences, however, is diminished if the differences in the number of crossings were due to unexplained variability. Duration of hesitations and crossings During the experimental width tests, the control and the variable means of the combined seasons (Table 5) of both the duration of hesitations and the duration of crossings are not significantly different (P > 0.20 and P > 0.10, respectively). Generally, if the narrower width was expected to result in more reluctance to cross, then longer hesitations would be expected. The means, except for the spring of 1977, support this trend. Reasons for the exception in the spring of 1977 are unknown. In all of the seasons except one (fall of 1976) the variable had shorter mean crossing times. Animals being more reluctant in crossing the variable (narrower width) would be expected to do so more hurriedly (more and/Gr faster trotting and bounding). During the experimental overhead netting tests, the control and the variable means of the combined seasons (Table 6) of both the duration of hesitations and the duration of crossings were not significantly different (P > 0.20 and P > 0.40, respectively). Generally, if the net arch was expected to result in more reluctance to cross, then longer hesitations would be expected. The means for the duration of hesitations for the fall of 1978 were not significantly different (P > 0.20). �-15- However, the variable mean was significatnly (P < 0.05) less in the spring of 1979. Also animals being more reluctant in crossing under the netting (variable) would be expected to do so more hurriedly (more and/or faster trotting and bounding). The duration of crossings under the netting (variable) for the fall of 1978 were significantly (P < 0.05) greater than the control, whereas the duration of crossings for the spring of 1979 were not significantly (p > 0.50) different. Differences between the spring and fall migrations are perplexing. The variable mean of duration of hestiations and the variable mean of duration of crossings were both significantly (P < 0.05) greater in the fall of 1978 than in the spring of 1979, whereas the control mean of duration of hesitations and control mean of duration of crossings were not significantly (P> 0.05 and P> 0.50, respectively) different. Possibly the data of the two seasons should not be combined. The differences may be related to the observed and postulated dissimilarities between the seasons, some of which are as follows: Spring Fall First overhead net experience Potentially second overhead experience Good physical condition Poor physical High wariness (hunting related) Moderate Most mature at side Maternal-fawn females with fawns bond strong condition wariness Most mature females Maternal-yearling parturient bond weak Dawn activity No dawn activity Very high motivation to migrate (more direct movements and less time consumption likely, casually related to increasingly inclement weather) Low stream flow in Gore Creek Behavioral net .High motivation to migrate (more indirect movements and more time consumption likely, casually related to weather) High-turbulent stream flow in Gore Creek responses:crossing ratios During the experimental width tests, there were more animals exhibiting behavioral responses (BR) and more instances of behavioral responses per crossings (C) during variable than control for each of the responses studied (Table 7). Although the ratios for instances of behavioral response per crossing are larger for variable width in each case, they were not significantly larger (P > 0.20). Therefore, differences in these behavioral responses may be due to chance and not to the reluctance of animals in crossing a narrow bridge width. �-16- During the experimental overhead netting tests, there were more animals exhibiting behavioral responses (BR) and more instances of behavioral responses per crossing (C) during variable than control for most of the responses studied (Table 8). Although the ratios for instances of behavioral response per crossing were larger for the variable except for muzzle-to-ground, they were not significant (P > 0.50). Therefore, differences in these behavioral responses may be due to chance and not to the reluctance of animals in crossing under a net arch. Deer Guards The first deer guard (prototype I) was tested during 1972-1973. Sixteen of 18 deer released in the runway attached to the guard crossed voluntarily (Reed et ale lS74b). Four other prototypes were tested, prototype II and III in 1975, prototype IV in 1976, and prototype V in 1978. Five of seven deer released in the runway attached to prototype II guard crossed voluntarily. They elicited only four instances of investigative behavior in apparent response to the structure. Two walked, two trotted, and one bounded across the guard (Table 9). Nine of 14 deer tested with prototype III crossed voluntarily, exhibiting 29 "band-necklook" instances of investigative behavior and 17 "walk-on" responses. Six of the animals walked, one trotted, and two bounded across the guard (Table 10). Five of eight deer tested with prototype IV guard crossed voluntarily, exhibiting eight "bend-neck-Iook" instances of investigative behavior and three "walk-on" responses. One of the animals walked, two trotted, and two bounded across the guard (Table 11). All four deer tested with prototype V guard crossed voluntarily, exhibiting 27 "bendneck-look" instances "of investigative behavior and six "walk-on" responses. Three of the animals walked and one bounded across the guard (Table 12). Ratios, including "bend-neck-Iook" per crossing, "walk-on" per crossing, and crossings per involuntary or no crossing, for the deer guard prototype are presented in Table 13. Deer Fence Length Six 2.44-m fences were evaluated as to the reduction of deer-vehicle accidents after installation of the fences. The range of reduction was 67.8 to 86.5 percent with a cumulative average of 78.5 percent (Table 14). Since this evaluation is based upon year-to-year comparisonse the cummulative average has fluctuated annually throughout the evaluation. The winter of 1977 was exceptionally mild and probably resulted in fewer kills at some of the fences. Conversely, the winter of 1979 Was eAn acknowledged common fallacy in biological investigations is t~ compare groups separated by time. For evidence so obtained, cause-and-effect should be attributed cautiously. �-17- exceptionally severe and probably resulted in more kills, especially at the Eagle 2.44-m fence. Data from severe winters were not discarded from the sample, similarly, data from mild winters should not be discarded either. Weather conditions, fence length (Table 14), and number and behavior of deer associated with the fenced areas were probably the major factors resulting in the varying degrees of accident reduction. Adjacent to the Diamond S 2.44-m fence, deer normally concentrated in crested wheatgrass fields northeast of the highway in late winter and early spring. Mean number of deer crossings during March-May periods continued to increase (Table 15) in spite of the decrease in numbers of deer seen during spotlight counts. Deer were neck banded with numbered collars (Fig. 7), automatic tagging devices, and telemetry collars in the Vail, Avon, Edwards, and Eagle 2.44-m fence areas. Numerous sightings and locations of these banded animals and direct observations of unbanded animals resulted in data on the movement of deer lateral or parallel to the Vail, Avon, and Eagle fences (Table 16). Females and males moved lateral to the fences for mean distances of 0.578 and 0.709 km, respectively. Twenty-eight one-way deer gates were located in the Vail, Avon, Edwards, and Eagle fences (Table 17). Reed et al. (1974a) reported on deer use of one-way gates in the Vail 2.44-m fence during 1970-1972. Since then the use of these gates have diminished substantially as deer apparently have adpated to the fencing-underpass complex and the increasing recreational-residential development. Deer use of the one-way gates in the Avon 2.44-m fence diminished in 1979 possibly because of the development of the Avon airport and associated buildings and chain-link fence. Benefit-cost analysis of 2.44-m fencing and other methods was reported by Reed et al. (1981). Additionally, specifications and maintenance of deer (2.44-m) fences and associated structures were reported by Reed (1981b). Highway Lighting Deer crossings and accidents occurred in the study area in 1974, 1975, 1978, and 1979. The lights were evaluated for 56 weeks during these four years, 28 weeks with the lights off and 28'weeks with the lights on. The estimated deer crossings per accident with lights off was less than the ratio with the lights on, 55.1 and 66.9, respectively. However, the difference was not significant for anyone of the years (X2 = 0.252 - 1.133, P>0.25), or for the composite of the four years (X2 = 0.781, P > 0.25). Lb and Lt measurements were made at each of the 39 accident sites, 26 in transition lighting and 13 in full lighting (Tables 18 and 19, respectively). It was hypothesized that these measurements would result in visibility indices that were lower in transition lighting than in full lighting, and that the former might have had to be discarded from the sample (i.e. transition �-18- lighting would not provide sufficient illumination for an adequate test). However, the means of the visibility indices (using absolute values) were 1.845 (± SD 1.538) and 1.754 (± SD 0.849) for transition and full lighting, respectively. The difference was not significant (P > 0.50). Hence, the transition lighting data were retained in the evaluation. Additional description of data analysis was provided by Reed et al. (1977) and Reed and Woodard (1981). Animated Deer Crossing Sign Signs were evaluated for 15 weeks during 1972 and 1973; eight weeks with the signs off and seven weeks with the signs on. The deer crossings per accident (deer kill) ratio with signs off was nearly identical to the ratio with the signs on, 56.5:1 and 56.9:1, respectively. By chi-square analysis, there was no significant difference (P > 0.50) between the crossings per kill ratios during 1972, 1973, or for the composite of the two years. A more detailed description of data analysis was provided by Pojar et ale (1975). DISCUSSION AND RECOMMENDATIONS Underpasses Generally, it was found that mule deer continued to be reluctant in using relatively small underpasses. This reluctance ultimately worked against highway safety and the deer resource. Consequently, it is recommended that larger (openness effect> 0.6, metric measurements) underpasses be constructed where deer passage under the highway is needed. This recommendation is made for deer having high motivation to cross the highway alignment. It follows that deer having light to moderate motivation will require larger structures such as open-bridge underpasses (Fig. 8) where our data suggest little reluctance occurs. Additional discussions and recommendations regarding underpasses were reported by Reed et ale (1975), Reed et ale (1981), and Reed (1981a, 1981b). Overpass Based upon the overpass data analyses, it does not appear that an important level of reluctance of deer to bridge width was reached. Further experimentation on the parameter of width was not considered practical since physical constraints did not allow a narrower « 2.48 m) width to be readily constructed and subsequently changed to a "control". Theoretically, there would be a point at which an overpass would be so long and narrow as to preclude deer crossings. Similar to the openness effect of underpasses, the primary stimulus of a given overpass structure to approaching deer may be termed the "bridge-effect". Calculated as follows: width V height length �-19- the bridge effects of the control and variable were 0.65 and 0.34 (metric measurements), respectively. In addition, it appears that deer were not reluctant to pass under an overhead netting designed to simulate a pedestrian-type overpass structure and wire mesh to prevent deer from jumping off and falling onto the roadway. Generally, deer crossed the overpass with somewhat less reluctance than that exhibited during passages at the Vail underpass (L.e , the look-up behavior was essentially absent at the overpass, but common at the underpass [Reed et al. 1975]). Caution should be used when comparing these two structures since different dimensional parameters are involved. Deer Guards The first deer guard (prototype I) had limited effectiveness in preventing deer movements through openings in 2.44-m fences (Reed et al. 1974b). The other four prototypes were equally limited in preventing or discouraging deer movements through an opening in a 2.44-m fence under the conditions of these tests. Several other concepts for deer guard prototypes (e.g. Fig. 9) were not tested. However, based upon the responses of deer to various aspects of the five prototypes, it is estimated that they would not be effective. Where 2.44-m vehicle gates (manual, "push-type", or automatic) are not feasible, no effective guard can be recommended as a result of this study. Deer Fence Length Six 2.44-m fences having an average length of 3.5 km were effective in that fewer accidents occurred after installation of the fences. Although cause-and-effect should be attributed cautiously because of separation of groups by time, number of years (5-10) and different areas (6) studied tend to represent some of the variability that exist for application of this methodology. In order to maintain approximately 75 percent fewer accidents after installation, 2.44-m fences must be adequately resistant to deer passage. This can be accomplished by construction and maintenance (Reed 1981b) where adequate basal closure and permanency are proVided. Based upon the mean lateral movements of deer to 2.44-m fences, such fences should extend approximately 0.8 km beyond deer concentration areas, and pass structures (underpasses and overpasses) should be located at least every 1.6-km along the fence where deer passage or crossings are needed. One-way gates were effective in allowing deer to escape the highway rights-of-way when they were strategically located. When one-way gates are recommended for installation in 2.44-m fences, they should be located near drainages or vegetative cover. Recommendations for spacing of the gates and other details were covered by Reed et al. (1974a) and Reed (1981b). Highway Lighting Visibility index means calculated where 70 percent of the motorists in the lighting study are near the level can see a target at satisfactory separation �-20- • distance (Gallagher and Mequire 1974). To attain a level where 85-95 percent of the motorists can see a target at satisfactory separation distance, a visibility index value of 2.6-3.6 is required. Only 6 of the 39 indices were within or above this range (Table 18 and 19). Probably inherent in this problem was the drab pelage of deer which readily blended with the lighted highway surface when in certain locations. For example, accident 24 (Table 19) invQlved a target and background with very low contrast despite its occurrence under full lighting. At these locations, just beyond the lamps and when the background was relatively well lighted (~ 1.0 fc) it is estimated that an increase in horizontal illumination would not substantially increase the contrast or likelihood of motorist visual discrimination. Since highway lighting was not effective as tested under the conditions of this study, it is not recommended as a method to reduce deer-vehicle accidents. Additional discussion is provided by Reed and Woodard (1981). Animated Deer Crossing Sign The lighted, animated deer crossing sign was not effective in reducing deer-vehicle accidents as tested under conditions of the study. Additional research should be conducted before signs are recommended as methods to reduce deer-vehicle accidents. Although additional sign research was provided for in the study proposal (i.e. test sign with advisory speed reduction on "educational" sign below animation), no additional research was conducted bec~use the study area was occupied with the highway lighting portion of this study during each available season (1974-1979). Additional discussions and recommendations regarding the lighted, animated deer crossing sign were reported by Pojar et al. (1975). CONCLUSIONS Studies 6f the first three methods, devices, or structures (underpasses and overpasses, deer guards, and deer fence length) .were undertaken to determine the efficacy of modifying deer behavior, or more specifically, of keeping deer off highways. Both qualitative and quantitative observations indicated that strategically located underpasses and overpasses with acceptable dimensions and characteristics were effective in providing deer with relatively safe passage to needed resources. Of the deer guard prototypes tested in this study none were effective in precluding deer movements. If simple and economically feasible guards are important to highway safety programs, additional research along new conceptual lines may be necessary. Segments of highway having deer fences were shown to have fewer accidents after fence installation. Generally, these fences have been effective in modifying deer movements, especially when used in conjunction with underpasses. It is imperative that such fencing be adequately constructed and regularly maintained. Studies of the last two methods, and animated deer crossing sign) of modifying motorist behavior. hypothesized that, by lighting a devices, or structures (highway lighting were undertaken to determine the efficacy In the highway lighting study, it was deer crossing area, motorists' visibility �-21- would be sufficiently enhanced to allow timely target (deer) discrimination and consequent accident avoidance. Similarly, in the animated deer crossing sign study, it was hypothesized that animated warning signs would increase motorist awareness sufficiently to allow for accident avoidance. Apparently, any increased motorist awareness did not result in sufficient behavioral change to reduce accidents. Of the five methods, devices, or structures tested, deer fencing (2.44 m in height), used in conjunction with strategically located underpasses and one-way deer gates, was the most effective. ACKNOWLEDGMENTS Five states, Colorado, Idaho, Nevada, Oregon, and Wyoming, participated in the HPR-3(3) pooled fund study under auspices of the Federal Highway Administration. Their participation is hereby acknowledged and appreciated. We thank the Federal Highway Administration, Colorado Division of Highways, Colorado State Patrol, Public Service Company of Colorado, and land owner Leo F. Jammaron for their cooperation. Denis E. Donnelly of the Colorado Division of Highways measured and analyzed the horizontal illumination levels and provided administrative direction. Burrell B. Gerhardt, then. of the Colorado Division of Highways, provided direction, herewith acknowledged with appreciation. Jim Fleming, Kenneth R. Kincaid, and Claudia A. Doose assisted with field work. �-22- LITERATURE CITED Child, K. N. 1974. Reaction of caribou to various types of simulated pipelines at Prudhoe Bay, Alaska. Pages 305-812 in V. Geist and F. Walther, eds. The behaviour of ungulates and its relation to management. Int. Union Conserv. Nature and Nat. Resour., Morges, Switzerland. 940pp. Clover, M. R. 201. 1956. Single gate trap. Farber, E., V. Gallagher, and A. Cassel. and vehicular illumination systems: Highway Admin. Rep. 109pp. Calif. Fish and Game. 42:199- 1971. Interaction between fixed Phase I - interim report. Fed. Free, S. L., and C. W. Severinghaus. Undated. Report on the effectiveness of a "deer proof" fence on the New York State thruway. Unpubl. Rep., P-R Proj. W-89-R-3, New York Dept. Environ. Conserv., Albany. 22pp. Gallagher, V. P., M. S. Janoff, J. G. Blubaugh, and P. L. Vetere. 1972. Interaction between fixed and vehicular illumination systems: Interim report on phase II. Fed. Highway Admin. Rep. FffiJA-RD-72-23. 114pp. , and P. G. Meguire. 1974. Contrast requirements Fed. Highway Admin. Rep. FHWA-RD-74-76. 72pp. ---- Gordon, D. F. 1969. "Deer mirrors" -- a clearer picture. Game, Fish and Pa rks, Game InL Leafl. 77. 3pp. Helms, R. N. 1969. Light or death. Paper presented Univ. Colo. Highway Eng. Conf. Feb. 14pp. Klein, D. R. 1971. Reaction of reindeer Science 173(3995):393-398. MacKay, D. M. patterns. Colo. Div. at 42nd Annual to obstructions 1957. Moving visual images produced Nature 180(4591):849-850. of urban drivers. and disturbances. by regular stationary Mansfield, T. M., and B. D. Miller. 1975. Highway deer kill District 02 regional study. Caltrans Environmental Branch, Sacramento, Calif. 49pp. Muller, S. 1967. Road traffic and wildlife. 53 (3) :121-129. Strasse und Verkehr. Myers, G. T. 1970. An investigation of deer-auto accidents. Colo. Div. Game, Fish, and Parks. Game Res. Fed. Aid Proj. W-38-R-24. Game Res. Rep. July, Part 3:403-437. �-23- Pojar, T. M., D. F. Reed, and T. C. Reseigh. 1972. Highway construction -motorist and deer safety. Proc. Western Assoc. State Game and Fish Comm., July 16-19. 268-271. --- , R. A. Prosence, D. F. Reed, and T. N. Woodard. of a lighted, animated deer crossing sign. 87-91. 1975.. Effectiveness J. Wildl. Manage. 39(1): Puglisi, M. J., J. S. Liridzey,and E. D. Bellis. 1974. Factors associated with highway mortality of white-tailed deer. J. Wildl. Manage. 38(4):799-807. Reed, D. F. 1969. Techniques for determining potentially critical deer highway crossings. Colo. Div. Game, Fish and Parks. Game Inf. Leafl. 73. 3pp. 1981a. Mule deer behavior at a highway underpass exit. Wildl. Manage. (In process) ____ J. • 1981b. Highway deer fencing and associated structures - specifications, maintenance, and effectiveness. (In process) , and T. N. Woodard. 1981. Effectiveness of highway lighting in reducing deer-vehicle accidents. J. Wildl. Manage. (In process) ---- ----:- , T. M. Pojar, and T. N. Woodard. Wildlife Surveillance. 3pp. , and ---J. Wildl. deer. 1973. A Video Time Lapse System for Colo. Div. of Wildl., Game Inf. Leafl. 94. 1974a. Use of one-way.gates by mule Manage. 38(1):9-15. ____ , and 1974b. J. Range Manage. 27(2):111-113. Mule deer responses to deer guards. , T. N. Woodard, and T. M. Pojar. 1975. Behavioral response of mule deer to a highway underpass. J. Wild1- Manage. 39(2) :361-367. ---- ____ , and T. D. I. Beck. 1977. Highway lighting to prevent deer-auto accidents. Colo. Div. of Highways Rep. 77-5. 26pp. ____ , T. D. I. Beck, and T. N. Woodard. 1981. Methods of reducing deer-vehicle accidents: benefit-cost analysis. (In process) Reynolds, H. G., and H. W. Springfield. 1953. Reseeding southwestern range lands with crested wheatgrass. Farmers' Bulletin No. 2056. U. S. Dept. of Agriculture. U. S. Government Printing Office. 20pp. Rumar, K. 1975. Causes and prevention of night driving accidents. Man-Environment Systems 5(3):171-174. Siglin, R. J. 1966. Marking mule deer with an automatic tagging device. J. Wildl. Manage. 30(3):631-633. �-24- Technical Committee of the ClEo 1972. A unified framework of methods for evaluating visual performance aspects of lighting. Pub. ClE (Commission lnternationale de L'Eclairage) No. 19. Paris. 90pp. Teuber, M. L. 1974. Sources of ambiguity in the prints of Maurits C. Escher. Scientific American 231(1):90-104. Thompson, F. A. 1967. Deer on highways 1967 supplement. Game and Fish. 7pp. Typescript. New Mexico Dept. Ward, A. L., N. E. Fornwalt, S. E. Henry, and R. A. Hodorff. 1979. Effects of highway operation practices and facilities on elk, mule deer, and pronghorn antelope. Fed. Highway Admin. Rep. FHWA-RD-79-143. 48pp. Woodard, T. N., and D. F. Reed. of deer-vehicle accidents. Soc. Conf. 19:18 (Abstr.). 1974. Economic considerations in reduction Trans. Central Mtn. Plains Sect. Wildl. �-25- Table 1. Highway underpasses, height and width dimensions, number of deer passages, openness or tunnel effect, and deer activity adjacent to the underpasses. Only the first two structures were intended for regular deer use, Underpass Height X Width length 1/ (m) Vail deer 'l:_/ 2 Eagle West 2 Hanun Creek Dry Creek Arch deer Salida East 1 Salida East 2 Chaffee Gulch 1/ Openness of tunnel effect (m) Adjacent deer activity/2,44-m fencing adjacent structure 59.8 0.03 Concentrated, highly motivated migration/ both sides Moderate/one side 3.0 0.42 Moderate/one side 1.5 0.10 Moderate/one side 4.57 Moderate/both sides 5.57 Moderate/both sides 16.7 3.75 Light/both sides 21.0 6.61 Light/both sides 66.5 0.61 Light/both sides 345.1 3.05 X 3.05 30.48 'IT(1.06) 108.7 Eagle East 1 4.27 X 4.27 45.58 Eagle East 2 2.44 X 2.44 4/ 59.54 (3.75 X 25.24)0.67 ]j Eagle West 1 Avon Number deer passages ner year 32.0 i/ ]_/ 0.31 (4.57 ~3;:~24)~.67 ~/ 13.87 . 5/ (3.66 X 20.73)0.67 13.56 5/ (5.49 X 24.38)0.67 13.56 3.05 X 6.10 23.77 5.49 X 14.63 'i_/ 11.25 144.0 124.0 5.17 Moderate/none 7.32 X 14.63 19.83 5/ (3.66 X 9.14)0.61 13.56 35.0 5.39 Moderate/none 15.4 1. 51 Moderate/none 'i_/ !.!Width (or open-end surface area) and length are measured parallel and perpendicular to direction of traffic, respectively. Formula is used to calculate openness or tunnel effect. 2/ - All except the Arch, Salida, and Chaffee Gulch structures were adjacent to Interstate 70. 1/Seasona1 mean resulting from a 4-year study (Reed et a1. 1975). 4/ - Eagle West 1 and 2, Mamm Creek, and Dry Creek structures are twin bridges. 'i_/Adjustedfor irregular inside topography. 6/ - - Most passages occurred without completion of the 2.44-m fencing. l/Based upon use during one month. The 2.44-m fencing was completed about October 1, 1979. to the structure �Table 2. The number ~f deer crossings and approaches examined on video replay at the overpass and the crossings per approach (C:a) ratios for four mi~ration seasons (Fall 1974 - Spring 1976). YearlSeason CrossinRs Distant 1/ Approach 21 Approach C:a 1974 Fall 99 - 2 49.5:1 1975 Spring 93 - 19 4.9:1 I 1975 Fall 77 - 3 25.7:1 1976 Spring 60 - 7 8.6:1 329 - 31 10.6:1 Tota1s/Avg. 11 - Denotes a deer that entered the overpass area not encompassed by an entrance and exit trackbed and that was oriented toward the structure. This overpass area was not covered by the video surveillance system during this period. 2/ -:'Denotes a deer that entered either the entrance or exit trackbed area without crossing the overpass. N 0'1 I �Table 3. The number 'of deer crossings and approaches examined on video replay at the experimentalvariable-width overpass and the crossings per approach (C:a) ratios for four migration seasons (Fall 1976 - Spring 1978). Crossings ._-" Control Distant 1/ ApproacJ/ ApproaclF C:a Crossings Variable Distant Approach Approach C:a Total Crossings 1976 Fall 5 25 5:1 5 19 3.8:1 44 1977 I N "'-I' Spring 41 4 10.2:1 19 4 4.8:1 60 1977 Fall 31 2 o 15.5:1 13 4 7 1.2:1 44 26 o 4 6.5:1 14 7 8 0.9:1 40 123 2 13 8.2:1 65 11 24 1.9:1 188 1978 Spring Totals/Average 1/ - Denotes a deer that entered the overpass area not encompassed by an entrance and exit trackbed and that wa~ oriented toward the structure. The overpass area was that area covered by the video surveillance system during the fall of 1977 and the spring of 1978. !/ Denotes a deer that entered either the entrance or exit trackbed area without crossing the overpass. I �Table 4. The number of deer crossings <111d .ipp rouchc s examined on video replay at the experimentalnet-arch overpass and the crossings per app roo ch (C:a) r a t Los for two migration seasons (Fall 1978Springs 1979). CONTROL Crossings 1978 Fa11 25 VARIABLE Distant 1/ '2/ Approach- Appr6ach- C:a 2 2 6.2:1 Crossings Distant Approach Approach C:a TOTAL Crossings 4 5 6 0.4:1 29 I N 00 I 1979 Spring 18 8 12 0.9:1 14 4 2 2.3:1 32 Totals/Avg. 43 10 14 1. 8:1 18 9 8 1.1 :1 61 l'Denotes a deer that entered the overpass area not encompassed and that was oriented toward the structure. by an entrance and exit trackbed l'Denotes a deer that entered either the entrance or exit trackbed area without overpass. crossing the �Table 5. The mean duration (seconds) of hesitations and crossings during control and variable widths at the experimentai-variable-Width overpass. 17 1976F- 1977S 1977F 1978S TOTAL p2_! Hesitations Control 3/ 4.5±1.STn=10) 13. 2±14.2 (n=29) 8.3±7.7(n=34} 9.2±6.9(n=30) 9.6±9.8(n=103) p > 0.20 Variable 7.4±4.8(n=14) 4.9±3.5 (n=6) 7.4±4.9(n=25) 12.1±lS.7(n=41) 11.7±11.1(n=18) lS.7±14.7(n=26) 11.8±11.8(n=64) Crossings Control 8.4±5.3(n=27) 8.9±5.8(n=26) 9.6±10.3(n=119) p > Variable 8.3±3.8(n=19) 8.8±8.9(n=19) 1/ . - Fal! or spring (F ~r S) migration periods. 2/ - Independent t statistic. 3/ - Standard deviation. 4.8±4.9(n=11) 6.8±5.7(n=14) 7.5±6.3(n=63) 0.10 I N \0 I �Table 6. The mean duration at the ~xperimenta1-net-arch (seconds) of hesitations overpass. FALL 1978 and crossings SPRING 1979 during control and variable pll TOTAL Hesitations Control 10.2±12.4- 2/ (n=42) 16.3:1;19,4(n=49) 13.5±16.7(n=91) p 14.3±14.3(n=20) Variable 7.2±4.6(n=20) > 0.20 10. 2±10. 4 (n=4 7) I Crossings VJ 0 I 7.2±12.4(n=25) Control 8.7:1;10.2(n=18) 8.0 ±ll. 4 (n=43) P > 0.40 Variable 24.2±19.8(n=4) 11 - Independent ~/Standard t statistic. deviation. 6.4±8.2(n=14) 10 ,1\ ±13. 3 (n=18) �-31Table 7. The number of selected behavioral responses exhibited by deer crossing an experimental-variable-widt~ overpass during four ndgration seasons (fall 1976, spring and fall 1977, and spring 1978) and calculated behavioral response ratios. Behavioral Responses (BR) Muzzle-to-Ground No. of Animals No. of Exhibiting Instances BR (BRa) of BR (B~) No. of Crossings BR :C1/ a (C) B~:C BR.~ :BRa '2/ - Control 32 55 123 0.26:1 0.45:1 1.72:1 Variable 1/ 18 33 65 0.28:1 0.51:1 1.74:1 Control 20 24 123 0.16:1 0.20:1 1.20: 1 Variable 15 22 65 0.23:1 0.34:1 1.47:1 Control 60 103 123 0.49:1 0.84:1 1.72:1 Variable 40 95 65 0.62:1 1.46 :1 2.38:1 Control 62 103 123 0.50:1 0.84:1 1. 66: 1 Variable 42 66 65 0.65:1 1.02 :1 1.57 :1 Control 33 63 123 0.27:1 0.51:1 1.91:1 Variable 34 57 65 0.52:1 0.88:1 1. 68: 1 Muzzle-to-Structure ~/ Low-Head 'if Hesitation §_! Alert Stance l/c 1.1 denotes the number of crossings during the four.seasons. l'Muzzle-to-ground denotes a deer lowering its muzzle to the ground. 1lControl and variable widths were 4.93 m and 2.48 m, respectively. ~!Muzzle-to-structure denotes a deer lowering its muzzle to the bridge deck or raising its muzzle to the bridge railing. 1/Low-head denotes a lowering of the head where the axis of the neck declines (posterior to anterior) below the horizontal. §_ICessation of forward movement for 1.0 second or more. Z/Alert stance denotes a position where the head and neck are above horizontal and the ears are erect with the vertical axis of the ear either perpendicular to horizontal or inclined forward. �-32Table 8. The number of selected behavioral responses exhibited by deer approaching or crossing an experimental-net-arch overpass during two seasons (fall 1978 and spring 1979).and calculated behavioral response ratios. No. of No. of Animals No. of Exhibiting Instances Crossi ngs BR ·.cll (C) a ·of·BR (BR.) BR .(BRa) a Behavioral Responses (BR) Muzzle-to-Ground BR. :BR ~ a ~I 15 33 43 0.35:1 0~77:l 2.20:1 6 9 18 0.33:1 0.50:1 1.50:1 16 25 43 0.37:1 0.58:1 1.56:1 8 15 18 0.44:1 o . 83 :1 1.88 :1 Control 39 102 43 0.91:1 2.37:1 2.62:1 Variable 18 55 18 1.00: 1 3.06:1 3.06:1 Control 48 91 43 1.12: 1 2.12:11.90:1 Variable 22 47 18 1.22: 1 2.61:1 2.14:1 Control 7 9 43 0.16:1 0.21:1 1.29:1 Variable 2 7 18 0.11:1 0.39:1 3.50:1 Control Variable 11 Muzz1e-to-Structure Control Variable Low-Head 41 'il . . 61 Hes]_tat~on - Alert Stance !/c 2/ denotes the number of.cr~ssings during the two seasons. !/Muzzle...,.to-ground denotes a deer lowering its muzzle to the ground. llControl was a 2.48 m vide bridge and the variable was the same bridge with a net arch similar to those used on pedestrian overpasses. ~/Muzz1e-to-structure denotes a deer lowering its muzzle to the bridge deck or raising its muzzle to the bridge railing. ~/ Low-head derio t as :a Lowe rLn g of the head where the axis of the neck declines (posterior to anterior) below the horizontal. ~/Cessation of forward movement for.1.0 second or more. l/Alert stance denotes a position where the head and neck are above horizontal and the ears are erect ~ith the vertical axis of the ear either perpendicular to horizontal or inclined forward •. �T.abLe 9. Responses of seven mule deer to pro t otvne II deer guard, 3,r)5 m wide X 3.66 m long and constructed with flat mill steel rails painted in 30-cm long alternating black and white sections. Result Crossed Crossed No crossing Involuntary crossing Crossed Crossed Crossed Predominate mode of crossing 1/ Number of bend-neck-looks bound trot 0 ---- f) walk trot walk walk 5 1 1 2/ Number of approaches 3/ Sex/age - 1 3 1 ~/ 3 5 11/~ 2 1 0 1 F/ ~/F M/F F/ F/ Time from release to completed crossing (sec.) 67.2 354.1 1()O.6 107.0 30.4· 67.0 1.1 Bend-neck-look and.apparently 2/ denotes a visible sensory inspection exhibited by bending neck, moving ears forward, looking at guard. - Approach denotes instances when subject comes within 1 m of guard and goes onto or turns away from the structure. 1/ Male or female is indicated by an M or F before the slash (/). slash. All others were yearling or mature. ~awn is indicated bv an ~ after the I W W I �Table 10. Responses of 14 mule deer to prototype III deer guard (five truck tire tubes cut, sectioned longitudinally, and stretched across and 15 cm above prototype II). Result No crossing Crossed Crossed Crossed Crossed Involuntary crossing No crossing No crossing Crossed Crossed No crossing Crossed Crossed Crossed 11 - Predominate mode of crossing ---walk walk bound trot Number of bend-neck-looks Number of approaches I 0 9 0 5 3 o . 11 Number of walk-ons 2 11 3 1 1 () 1 Sex/age F/ MIT< FI F/ M/ ---- 10 ---- f) ---- 0 walk walk 12 2 ---- 0 o 0 F/F walk walk bound 3 3 3 4 2 3 0 1 1 MI M/ F/ 3 1 27 6 3 1 () F/F MI f) F/ 4 1 FIT< F/ Walk-on denotes instances when subject walked onto the stretched returning to solid ground. Time from release to completed crossing (sec.) 83.8 1,653.0 7 J) 41. o 790.() --- 380.0 476.6 835.0 50S.2 83.6 tubing and backed off or turned around I VJ .I:I �Table 11. Responses of eight mule deer to prototyne IV deer guard (93 ten-sneed bicycle and stretched across and 15 cm above prototvne II). Result Crossed Crossed No crossing Crossed No crossing Crossed Crossed No crossing Predominate mode of crossing trot trot ---bound Number of bend-neck-1ooks Number of approaches Number of walk-ons 2 3 1 I) 1 1 I) 0 I) 1 2 1 ---- 0 I) () walk bound 5 3 I) f) 1 0 ---- 7 9 I) Sex/age ~/ ""1/ ~/ ~/~ ~/~ ~/ ~/ M./~ tire tubes cut Time from release to comn1eted crossing (sec.) 9Qf).n 15.6 174.5 492.3 301).() -- I W V1 I �Table 12. Responses of four mule deer to prototype V deer guard (plack and white scintillation or rotary motion ~attern described by McKay [1957]). Time from release to completed crossing (sec.) Result Predominate mode of crossing Number of bend-neck-1ooks Number of approaches Crossed walk 7 2 1 F/Y 1,765.0 Crossed walk 3 2 1 M/Y 124.3 Crossed bound 0 1 0 F/M or Y 5.2 Crossed walk 17 6 4 F/N or Y 647.3 Number of walk-ons Sex/age I w 0- I �-37- Table 13. The prototype, number of deer tested, bend-neck-looks (investigative behavior) per crossing (xing), walk-on per xing, and xing per involuntary or no xing ratios for five deer guard designs. Prototype No. Ratios wa1k-on:xing xing: involuntary or no xing n bend-neck-100k:xing 18 3.1:1 8.0:1 7 0.8:1 2.5:1 III 14 3.2:1 1.9 :1 1.8:1 IV 8 1.6 :1 0.6:1 1.7:1 V 4 6.8:1 1.5 :1 };/ I II !/ As reported by Reed et al. (1974b). �-38- Table 14. The mean annual number or pre- and post-installation deer highway kills and percent reduction for six 2.44-m fences adjacent to Interstate 70 and Highway 82. Mean annual preinstallation mortality Mean annual postinstallation mortality Percent reduction mortality Fence (Hwy) Length of hwy fenced km (Miles) Vail (1-70) 2.4 (1.5) 36 (3)1../ Avon (1-70) 3.6 (2.3) 28 (1) 3.9 (7) 86.1 Edwards (1-70) 3.6 (2.3) 27 (1) 5.7 (7) 78.9 Eagle (1-70) 7.7 (4.8) 167 (1) 22.5 (6) 86.5 Diamond S (82:: 1.8 (1.1) 10 (3) 1.8 (8) 82.0 Carbondale (82) 1.8 (1.1) 14 (5) 4.2 (5) 70.0 Cumulative Avg. 11.6(10).Y 67.8 78.5 1/ - (n) denotes the number of years of pre- and post-installation data. �-39- Table 15. Mean number of deer observed on spotlight counts and mean numh er of deer crossings be~een quarter-mile section markers 25 to 30 on Highway 82 during March-}~y for 1968 through 1976 (n = number of counts or number of 24-hour periods). Mean Mar .-Nay Crossings Year March April Mean Total 1968 134.8 (0=4) 73.0 (n=4) 103.9 (n=8) 1969 151.2 (n=4) 34.0 (n=5) 86.1 (n=9) 1970 104.5 (n=4) 56.0 (n=5) 77 .6 (n=9) 1971 66.8 (n=4) 51.4 (n=5) 58.4 (n-9) 11.7 (n=32) 1972 102.2 (n=4) 4.5 (n=4) 53.4 (n=8) 2.1 1-/ (n=38) 1973 137.4 (n=5) 47.0 (n=4) 97.2 (n=9) 5.5 (n=34) 1974 143.5 (n=4) 52.3 (n=3) 104.4 (n=7) 10.3 (n=38) 1975 126.8 (n=4 ) 93.0 (n=4) 109.9 (n=8) 17.8 (n=73) 1976 78.0 (n=4) 61.8 (n=4) 69.9 (n=8) 21.0 (n=28) 1..1 1.77 km of 2.44-m fence constructed during summer of 1971. �-40- Table 16. Female and male deer mean lateral movements (km) aIong or adjacent to the field side or highway side of 2.44-m fences at Vail, Avon, and Eagle as deterininedunder'several conditions. Vail Side of Fence Female Eagle Avon .Female ,Male Female Male Field Side Apparent II 0.900(n=3)~/0.500(n=2) Observed 11 0.272(n=8) 0.578(n=2) 0.588(n=107) 0.762(n=34) 0~350(n=l) 0.280(n=1) Highway Side Apparent Observed 0.617(n=3) O.350(n=l) 0.500 (n=5)4JO.506(n=4) -- Observed and ~/ harassed 0.975(n=4) 0.480(n=4) 1.207 (n=2) O.646(n=5) !I Movements estimated by noting the locations of radio-collared or neckbanded deer at selected points along the fences at widely separated time periods. 2/ (n) denotes number of lateral distances. 1./ !!_I Movements observed during their duration. Included one distance that was the "net" lateral movement:'occurring during a period of 45 minutes '. Twenty-two changes' in direction or mni-lateral movements occurred during this "net" movement.' 'The mean mini-·lateral movement was 140.9±154.8(Sn) m. 51 Animals were usually chased with a vehicle at night until they either escaped from the fenced side of the highway or "broke" back against the vehicle. �-41- Table 17. The number of one-way deer gates and mean annual number of passages through one-way gates located in four Interstate 70 2.44-m fences. 2.44 Fencing Location Number of One-way Gates Mean Annual Oneway Gate Passage Vail - both sides 1/ 7 (7) 2/ 73.6 (9) Avon - one side 6 (9) 70.6 (7) Edwards - one side 5 (7) 6.4 (7) 10 (10) 13.5 (6) Eagle - one side 1/ - (N) denotes the number of gates originally installed in the 2.44-m fence. In this case, two ~ates were removed and two installed at new locations. In cases where one-way gates received little use ( < 3 passages per year) and/or considerable human interference, they were removed and the openings fenced. 2/ - (n) denotes number of years of post-installation One black bear" passage occurred during 1974. data. �-42- Table 18. Visibility index measurements from accident sites in transition lighting. No. (fL) (fL) Contrast Relative Contrast Sensitivity (Percent) 1 0.019 0.025 0.311 2.21 _1,_/ 0.120 2 0.010 0.115 10.058 1.37 _3_/ 2.395 3 0.066 0.006 - 0.916 5.52 - 0.882 5 0.103 0.011 - 0.898 7.30 - 1.142 6 1.480 0.008 - 0.995 27.70~_I - 4.800 7 0.124 0.012 - 0.899 8.23 - 1.289 9 0.043 0.032 - 0.244 4.10 - 0.174 10 0.226 0.003 - 0.987 11.80 - 2.029 11 0.330 0.003 - 0.990 14.40 - 2.484 12 0.061 0.009 - 0.846 5.25 - 0.774 13 0.235 0.026 - 0.891 12.05 - 1.870 14 0.060 0.005 - 0.920 5.20 15 0.285 0.004 - 0.986 13.32 - 2.289 16 0.080 0.016 - 0.794 6.20 - 0.858 18 0.086 0.130 0.512 6.50 19 0.180 0.011 - 0.939 10.40 - 1.701 21 0.180 0.026 - 0.856 10.40 - 1.551 0.284 - 0.600 20.31 - 2.134 Background luminance· 22 (0.690)1/ Target luminance Visibilityl/ Index - 0.833 0.580 23 0.690 0.012 - 0.983 20.31 - 3.477 26 0.253 0.247 - 0.024 12.53 - 0.052 ----------------------------------------------------------------------------- �-43- Table 18. Visibility index measurements from accident sites in transition lighting. No. (Continued). Background luminance (fL) Target luminance (fL) Contrast Relative Contrast Sensitivity (Percent) V·1S1LbLLLt; 1/ 1 1 yIndex 27 0.066 0.260 2.939 5.55 2.842 28 0.279 0.136 - 0.512 13.20 -1.179" 31 0.041 0.006 - 0.839 4.00 -0.585' 32 0.280 0.009 - 0.968 13.22 -2.228 33 0.400 0.020 - 0.951 15.85 -2.625 36 0.045 0.475 9.556 4.25 7.075 ]/ Positive and negative values indicate frontlighting and backlighting, respectively. 2/ -- Derived from data presented by Technical Committee Report of the CIE 1972. 3/ -- ' The background at this site involved a snow covered emergency lane and right-of-way. The snow was lost before measurement. A value for similar conditions (No. 23) was used. �-44- Table 19. Visibility index measurements from accident sites in full lighting. No. Background luminance (fL) Target luminance efL) Contrast Relative Contrast Sensitivity (Percent) Visibilityl' Index 4 0.630 0.390 - 0.381 19.62 -1.303 8 1.100 0.209 - 0.810 24.44 - 3.449 17 0.240 0.470 0.958 12.20 2.036 20 0.197 0.069 0.650 10.95 -1. 240 24 0.185 0.178 0.038 10.57 -0.070 25 0.237 0.455 0.920 12.10 1.939 29 0.075 0.120 0.600 5.95 0.622 30 0.500 0.110 - 0.780 17.67 -2.401 34 0.420 0.140 - 0.667 16.24 -1. 886 35 0.580 0.330 - 0.431 18.98 -1.425 37 0.620 0.270 - 0.564 19.51 -1.919 38 0.440 0.108 - 0.754 16.63 -2.186 39 0.225 0.480 1.133 11.77 2.323 - ~/Positive and negative values indicate front1ighting and backlighting, respectively. �I L1I <r I Fig. 1. The control (4.93 m) and variable (2.48 m) widths (left and right, respectively) of the deer overpass over Gore Creek. Photos by D. F. Reed. �I '" ...::t I -.. _4~ __~_.._._.... __~ ..:;_ --.,.'" - ... .• - ~ ~ - , ---:.~ -, t ~j ! Fig. 2. Reed. The overhead netting asse~bled (variable) on the deer overpass. Photo by D. F. �-47- Fi~. 3. A one-way deer gate showing a deer bounding through the structure. Photo by D. F. Reed. �I 00 ...-:t I Fig. 4. A panorama of the lighting (13 luminaires) located (left to right, top to bottom, northwest to southeast of the study area) on Highway 82. Photos by D. F. Reed. �I 0'1 ..;j" I Fig. 5. The Spectra Model UBA spotmeter sholYn on the right was usee to measure the target (deer simulation shown at center) and background (highivaysurface a~d dark background beyond the simulation) luminances. Photo by D. F. Reed. �I o lI"\ I .... ~~.. , .» ::_,_.;." _"." - _- ~.~--i':~\: - •.':: .., -. Fig. 6. The lighted, animated deer crossing sign is sho\~ turned toward traffic with the lighted silhouette sequence beginning. Photo courtesy of the Colorado Division of Highways. �-51- .. .. ~ '! I I i , '" .. I .•I I I I .• T! " I I , I I \ .. ':' I. \ .• .• \ .,. ~,.-..••. - .. ...--- _ .....,.......-... .• Fig. 7. Neck banded doe number 50 near the 2.44-m fence east of Eagle and adjacent to 1-70. Photo by L. L. Green. �I N lJ') I Fig. 8. The Eagle Wes~ 2 bridge underpass under the west bound lanes of 1-70. fencing adjoins the bridge abutments. Photo by D. F. Reed. The 2.44-m �-53- Fig. 9. Reportedly plastic strip curtains have kept animals inside game parks in West Germany's Harz Forest Region. Effectiveness was based on light reflection from the plastic strips. AP photo. ��-55- July, 1980 JOB PROGRESS REPORT State of Colorado Project No. W-126-R-3 Work Plan No. Big Game Investigations 1 --~---- Job No. 1 Multispecies Investigations Guidelines for Wildlife Habitat· Manipulation Period Covered: Personnel: July 1, 1979 through June 30, 1980 W. H. Rutherford, W. D. Snyder ABSTRACT All information obtained during previous segments from literature review and personal interviews has been compiled, organized, and written into a manuscript for a guideline manual. Critique and review, revision, and submission for editorial review have been done, and the draft is now in the hands of the Division's technical editor. Publication will follow completion of the editorial work. GUIDLINES FOR WILDLIFE HABITAT MANIPULATION William H. Rutherford and Warren D. Snyder P. N. OBJECTIVE To publish a comprehensive manual of directional habitat management guidelines for sustaining and enhancing wildlife populations for consumptive and non-consumptive uses on Division of Wildlife properties. SEGMENT OBJECTIVES 1. Complete the preparation of a draft version of a habitat manipulation manual. 2. Submit the manual to selected reviewers for criticisms, comments, and suggested improvements. 3. Revise and rewrite the critiqued draft version. 4. Publish the manual as a Colorado Division of Wildlife Special Report. �-56- METHODS AND MATERIALS Activity during this segment was limited to writing, submitting drafts for editorial review, and revising. Methods and materials are self-explanatory. RESULTS The draft version was completed and submitted for review to the Assistant Director for Operations and the Chief of Game Research. Suggested revisions were made, the draft was re-written, and is now in the hands of the Division's technical editor. The schedule for publication of the manual is contingent on completion of the editorial work. The complete Table of Contents for the manual is as follows: A TECHNIQUES MANUAL FOR MODIFICATION OF HABITAT TO BENEFIT WILDLIFE Chapter I. II. III. INTRODUCTION TECHNIQUES FOR USE IN HABITAT MODIFICATION PLANNING, DESIGN, AND LAYOUT GENERAL MODIFICATION TECHNIQUES A. Mechanical Treatments 1. Timber and brush control 2. Renovation of perennial herbaceous cover 3. Dredging and diking 4. Blasting . 5. Sunnner fallowing and pre-plant cultivation B. Seeding and Planting 1. Shrub and tree establishment 2. Perennial herbaceous plant establishment 3. Food and food-cover estab Lf.shment; c. Prescribed Burning Herbicide Treatments Soil Tests and Use of Fertilizers Grazing Provision of Supplemental Drinking Water Farming Methods Compatible with Wildlife Habitat Timber Stand Management Riparian Habitat Management D. E. F. G. H. I. J. IV. HABITAT MODIFICATION PROCEDURES FOR SPECIES OR GROUPS OF SPECIES A. B. C. D. Big Game Plains Upland Game Birds Mountain Upland Game Birds Small-Game Mammals �-57- E. F. G. H. V. VI. Waterfowl Furbearers Raptors Non-Game Species EVALUATING EFFECT OF TREATMENT PROPERTY ADMINISTRATION CONSIDERATIONS A. Fencing B. Haying C. Grain Crop Production D. Share-Crop vs. DOW Operation E. Grazing Leases F. Incidental Income-Producing Procedure G. Use of Water Rights H. Weed Control I. Management Next to Private Property J. Property Size K. Access Trade-Offs L. Mineral Rights M. Mitigations ./l.Prepared by: ,f // _,.z/ j\ -~~/. [.t.~l{tl.Cz'·' I'. .. /::.£('/.,.' :'Y(L'-{ William H. Rutherford ;' Wildlife Researcher C t) ��-59- July, 1980 JOB PROGRESS REPORT State of Colorado ------------------------ Project No. W-126-R-3 Big Game Investigations Work Plan No. 1 Multispecies Job No. 2 Deer and Elk Management Period Covered: Personnel: Investigations Study July 1, 1979 through June 30, 1980 Colorado Division of Wildlife: Regional Wildlife Biologist Staff members; John Ellenberger, John Gray, Jim Olterman, Marc Elkins, Jack Vayhinger, Gene Schoonveld, Andre Duvall; Area Supervisors and District Wildlife Managers; Bob Hernbrode, Thomas M. Pojar. ABSTRACT The main thrust during this segment was to assist the Regions in becoming independent in their modeling programs. All four regions have access to a remote computer terminal in their Regional Offices. The NW and SE Regional Biologist Staff have the expertise to operate the ONEPOP model through their remote terminals. Both of these regions conducted their own modeling workshops where the simulations were updated and any desired modifications made. The SW region just acquired their remote terminal during this segment and have yet to be trained in its use. Assistance was given to the NE region in updating all simulations for setting the current management program. Some changes were made in DAU boundaries. The responsibility for management was discussed by the Big Game Supervisor. The results of these the Minutes of Biologist Meeting - June 24-25, the Big Game Supervisor's office in Denver. DAU's and regional the Regional Biologists and decisions are contained in 1980 and is available from ��-61- DEER AND ELK MANAGEMENT STUDY Thomas M. Pojar P. N. OBJECTIVE Devise and test a statewide deer and elk management system. SEGMENT OBJECTIVES 1. Divide the state into a system of data analysis units (DAU's) that are deemed practical for gathering and analyzing population data. 2. Based on currently available information, estimate population parameters, by DAU, through the simulation modeling approach. 3. Conduct training workshops involving management personnel to familiarize them with the interpretation of the output from the model and to incorporate empirical data from fieldmen directly associated with the DAU into the simulation to produce a candidate management simulation. 4. Identify data that are most useful for improving population parameters. estimates of RESULTS The following manuscripts editorial review. were prepared and revised in accordance Pojar, T. M. A management perspective of population modeling. Fowler, C., and T. Smith (eds.). Dynamics in Large Mammal Populations. John Wiley and Sons, New York. (In press). with In: Salwasser, H., and T. M. Pojar. Simulation modeling of pronghorn populations. In: Yoakum, J. (Ed.) Pronghorn Management. Wildlife Management Institute (Second draft). Wildlife ar Researcher ��-63July, 1980 JOB PROGRESS REPORT State·of COLORADO Project No. W-126-R-3 Big Game Investigations Work Plan No. 1 --------~---------- Multispecies Job No. for Deer-Elk 4 Animal and Pen Support Facilities Research Period Covered: Personnel: Investigations July 1, 1979 through June 30, 1980 Paul H. Neil, Lynn Sexton, Barry VanSant, Janice Reeves, Steve Torbit, Vicki Jameson, Carol Smith, Richard Parachini, and R. Bruce Gill ABSTRACT All of the major construction projects of the Research Support Facility were completed during the segment and all pens are occupied and functioning. Minor modifications to some pens and digestion cages still remain. The Foothills Campus pen facility presently houses 5 adult elk, 31 adult mule deer, 18 deer fawns, 3 adult antelope, and 13 antelope fawns. The Wildlife Research Center·pens are occupied by 5 mule deer fawns, 6 Rocky Mountain bighorn sheep, and 6 Rocky Mountain goats. A study is being conducted at present regarding identification and treatment of colostrum deficient fawns. A total of 18 fawns are in the study. Eleven have been identified as passive failures (received no 'colostrum after birth), six as passive successes (received colostrum after birth) and one unclassified. Attempts have been made to boost antibody levels in the passive failures by the use of plasm~ transfer therapy. Our results are inconclusive, however, preliminary analysis indicates we are boosting gamma globulin concentrations by 0.5 g/IOO mI. Thus far plasma therapy 'does not appear to be a successful treatment for preventing septicemia in fawns. ��-65- ANIMAL AND PEN SUPPORT FACILITIES FOR DEER-ELK RESEARCH Paul H. Neil P. N. OBJECTIVES To provide facilities and maintain populations of captive big game animals to support big game research programs. and pen SEGMENT OBJECTIVES 1. To continue to expand the big game research 2. To coordinate rearing, training, maintenance, and research activities with captive wild and tame animals under one research support facility manager. 3. To integrate animal and physical plant support monetary requirements under a single budget. METHODS and animal holding facility facility. manpower and AND MATERIALS Materials and equipment were ordered and various aspects of construction were placed on bid in order to complete several portions of the expansion program. The Toledo digital read-out scale was recalibrated to kilograms and the weigh room itself completed. Lumber and other materials were ordered for the construction of a large storage shed for a tractor and related equipment and miscellaneous supplies. The Young Adult Conservation Corps of the Bureau of Reclamation was enlisted to assist in completion of a tree nursery, road repair, and construction of a cement pad under the new digestion cages. Eight mule deer were transported to the Fort Collins facility from the Middle Park facility for summer holding. An additional eight mule deer that were being used in the Piceance Basin grazing studies also were transported to the Fort Collins facility. Five wild does were captured and released into the research facility pens to obtain fawns for the fawn rearing study. Thirteen orphan mule deer fawns were obtained from Division of Wildlife field personnel. Eighteen fawns were born to captive does within the facility. All of the fawns are being hand reared for recruitment into ongoing research programs. �-66- RESULTS AND DISCUSSION The second set of metabolism cages was completed in the spring of 1980 and the only remaining work to be done is installation of heat coils under the collection pans and pouring a cement pad in front of the cages. Winter work on the mule deer body composition study also was completed during the spring of 1980. Results and progress of this study are described under W-126-R, Work Plan 2, Job 1. Pregnancy scans were conducted in the spring of 1980 on 3 mature mule deer does and 3 mature antelope does. Comparison of the ultra-sound equipment and x-ray techniques for determining pregnancy was conducted. Detailed information on the techniques and results of these tests are described under W-126-R, Work Plan 5, Job 1. Eighteen of the 31 fawns being raised at the facility were incorporated into a study regarding identification and treatment of colostrum deficient fawns. We are developing a gluteraldehyde coagulation test, zinc sulphate turbidity test and total protein/gamma globulin estimation technique for the identification of colostrum deficient fawns (passive failures). With these tests we have correctly identified 11 of the fawns as passive failures, 6 as passive successes, and one which we could not classify. The mortality rate in the passive failures is 100% at present, and only 16% in the passive successes. Preliminary necropsy reports indicate the fawns died of septicemia characterized by diarrhea. Through plasma therapy we have tried to boost the antibody levels of the passive failures. The results here are incomplete, however, initial analysis indicates we boosted some of the fawns gamma globulin concentrations by as much as 0.5 g/100 mI. We may not have boosted to levels required for survival because the animals injected with plasma did die. Plasma therapy as a treatment in fawns that already have diarrhea has not been successful in preventing septicemia and death. The remainder of the fawns are being hand-reared research projects. Prepared by: Q~H~Dc*"Q Paul H. Neil -~ Wildlife Technician III and trained for use in various �July, 1980 -67- JOB PROGRESS REPORT State of COLORADO ----~-------------------- Project No. Big Game Investigations Work Plan Multispecies W-126-R-3 ----------------------No. 1 --------------------- Job No. 6 Investigations Digestible Nutrient Content of Deer and Elk Winter Forage Plants Period Covered: Personnel: July 1, 1979 through June 30, 1980 Roland C. Kufeld and Marilyn Stevens P. N. OBJECTIVE To estimate average nutrient content and digestibility values ~d their degree of variation within selected range forage plants during winter. SEGMENT OBJECTIVE To determine the degree of variation in nutrient content and digestibility of selected range forage plants during winter. METHODS AND MATERIALS A manuscript entitled "Winter Variation. in Nutrient and Fiber Content and In-Vitro Digestibility of Gambel oak (Quercus gambellii) and Big Sagebrush (Artemisia tridentata) from Diversified Sites in Colorado" was prepared and has been accepted for publication in the Journal of Range Management. Chemical analysis of nutrient content and digestibility of serviceberry (Amelanchier alnifolia) and mountain mahogany (Cercocarpus montanus) collected during the previous segment (Kufeld 1979) is currently underway. LITERATURE CITED Kufeld, Roland C. 1979. Digestible nutrient content of deer and elk winter forage plants. Colo. Div. Wildl. Game Res. Rept. July (1):47-51. Prepared by ~,ud c" ~ Roland C. Kufeld Wildlife Researcher C ��-69- July, 1980 JOB PROGRESS REPORT State of Colorado Project No. W-126-R-3 Big Game Investigations Work Plan No. 1 --~~------------ Multispecies Investigations Job No. 7 Period Covered: July 1, 1979 through June 30, 1980 Personnel: Big Game Research Publications A. E. Anderson, D. L. Baker, Dr. L. H. Carpenter, D. J. Freddy, Dr. N. T. Hobbs, R. C. Kufeld, T. M. Pojar, and R. B. Gill ABSTRACT Twenty-one articles were published or accepted for publication in scientific journals or symposia proceedings during the segment. One paper was accepted for presentation at a professional society meeting. Three Division of Wildlife internal publications were published in Segment 3 of W-126-R. ��-71- BIG GAME RESEARCH PUBLICATIONS R. Bruce Gill P. N. OBJECTIVE· To publish the results of research conducted under the auspices of Federal Aid Project W-126-R in a variety of professional journals and other indexed publishing media to insure widespread dissemination and availability of this information to natural resource managers and ecological scientists. SEGMENT OBJECTIVES Following is a list of publication working titles that will be prepared for publication under this job in Segment 3. The publication outlet is tentative depending upon acceptance by the particular periodical. 1. Big Game Investigations Federal Aid Job Progress Reports. 2. Guidelines for Habitat Manipulation - CDOW Special Report. 3. Simulation Modeling of Big Game Populations - J. Wildl. Manage. 4. Piceance Basin Deer Population Dynamics - CDOW Special Report. 5. Seasonal Diets of Tame Deer in the Piceance Basin, Colorado - J. Wildl. Manage. 6~ Estimating Deer Population Size from Counts of Fecal Pellet Groups CDOW Special Report. 7. Literature Review of Mountain Lion Population Dynamics - CDOW Special Report. 8. Miscellaneous unanticipated publications. PUBLICATION PROGRESS Publications Printed or Accepted for Publication Baker, D. L., and N. T. Hobbs. 1979. Nutritional quality of elk summer diets in Rocky Mountain National Park. Amer. Inst. BioI. Sci. Conf. Scient. Res. Natl. Parks, Proc. 2: In Press. Baker, D. L., and N. T. Hobbs. Outdoors 27(5):14-19. 1979. Elk are what they eat. Colo. �-72- Bowden, D., and A. E. Anderson. Sampling plans for sex and age ratios of mule deer. J. Wildl. Manage. (Accepted pending revision). DeYoung, C., and H. L. Geduldig. Parturition behavior in a wild mule deer. J. Wildl. Manage. (Accepted). Ellis, J. E., D. M. Swift, N. T. Hobbs, R. G. Woodmansee, and D. L. Baker. 1980. Estimation of nitrogen transport by large animals in seasonal ecosystems. Ecol. Soc. Amer. Bull. (Abstract). In Press. Freddy, D. J. 1979. Measuring heart rates of mule deer using a repeater type telemetry system. pp. 144-155. In F. M. Long (ed.). Proceedings of the 2nd International conference on wildlife biotelemetry. Univ. Wyo., Laramie. 259p. Geduldig, H. L. (Accepted). Home range of mule deer fawns. J. Wildl. Manage. Hobbs, N. T., D. L. Baker, D. M. Swift, and J. E. Ellis. 1977. Composition and quality of elk diets during winter and'summer: a preliminary analysis. pp. 47-63. In M. S. Boyce, and L. D. Hayden-Wing (eds.). North American elk: ecology, ,behavior, and management. Univ. Wyo., Laramie. 294p. Hobbs, N. T., D. L. Baker, D. M. Swift, and J. E. Ellis. 1980. and nitrogen based estimates of ungulate carrying capacity. Amer. Bull. (Abstract) In Press. Energy Ecol. Soc. Hobbs, N. T., D. L. Baker, D. M. Swift, and J. E. Ellis. 1981. Composition and quality of elk diets in Colorado. J. Wildl. Manage. 45(1):In Press. Hobbs, N. T., D. L. Baker, J. E. Ellis, and D. M. Swift. 1979. Nutritional evaluation of elk winter habitat. Amer. Inst. Biol. Sci. Conf. Scient. Res. Nat!. Parks, Proc·. 2:In Press. Kautz, M. A., W. W. Mautz, and L. H. Carpenter. 1980. Heart rate as a predictor of energy of mule deer. J. Wildl. Manage. In Press. Kautz, M. A., G. M. VanDyne, L. H. Carpenter, and W. W. Maub.z. Energy costs for some activities of mule deer fawns. J. Wildl. Manage. (Accepted pending rev,ision). Kufeld, R., J. Olterman, and D. Bowden. Colo. Outdoors 27(6):12-14. 1979. A new way to count deer. Kufeld, R. C., J. H. Olterman, and D. C. Bowden. 1980. A helicopter quadrat census for mule deer on the Uncompaghre Plateau, Colorado. J. Wildl. Manage. 44(3):632-639. Kufeld, R. C., M. L. Stevens, and D. C. Bowden. 1981. Winter variation in nutrient and fiber content and in vitro digestibility of gambel oak (Quercus gambellii) and big sagebrush (Artemisia tridentata) from diversified sites in Colorado. J. Range Manage. In Press. �-73- Milchunas, D. G., and D. L. Baker. In vitro digestion with respect to wild ruminants and sources of within and between trial variability. J. Range Manage. (Accepted pending revision). Pojar, T. M. A management perspective of population modeling. pp. . In C. W. Fowler, and T. D. Smith (eds.). Dynamics in large mammal populations. John Wiley and Sons, Inc., New York, NY (Accepted). Salwasser, H., and T. M. Pojar. Simulation modeling of pronghorn populations. pp. In J. Yoakum (ed.). Pronghorn antelope management. Wildl~anag~ Inst. (Accepted). Swift, D. M., N. T. Hobbs, and J. E. Ellis. 1980. Energy and nitrogen metabolism of cervids in winter. A simulation study. Reindeer and Caribou symp., Proc. 2. In Press. Swift, D. M., N. T. Hobbs, D. L. Baker, and J. E. Ellis. 1980. Strategies of diet selection by a large herbivore. Ecol. Soc. Amer. Bull. (Abstract). In Press. Internal DOW Documents Gill, R. B. 1978. Big game research program 1978-1988. Colo. Div. Wildl. Denver, CO. 72p. (Reprinted). Gill, R. B. 1979. (Ed.). Game research report. Div. Wildl. Denver, CO. pp. 1-230. July, Part One. Colo. Gill, R. B. 1979. (Ed.). Game research report. Div. Wildl. Denver, CO. pp. 231-502. July, Part Two. Colo. Papers Presented At Society Meetings and/or Workshops Carpenter, L. H., and C. E. Braun. 1980. A new approach to the dilemma of federal cost-sharing and technical assistance programs that alter wildlife habitat. Centro Mtns. Plains Sect. TWS. Ann. Conf. Gretna, NB. (\ U)~.~"\\ , Prepared by ~_~~ __ ~~__ .~__~.~ -~~~~ ~ R. Bruce Gill Big Game Section Chief 1\ _ ��July, 1980 -75- JOB PROGRESS REPORT State of COLORADO Project No. W-126-R-3 Big Game Investigations Work Plan No. 1 Multispecies Investigations Job No. 8 Big Game Publication Editing and Library Services Period Covered: Personnel: July 1, 1979 through June 30, 1980 M. W. Herschopf, Dr. O. C. Cope, R. B. Gill, A. E. Anderson, D. L. Baker, R. M. Bartmann, G. D. Bear, T. D. I. Beck, Dr. L. H. Carpenter, D. J. Freddy, Dr. N. T. Hobbs, R. C. Kufeld, P. H. Neil, T. M. Pojar, D. F. Reed, W. H. Rutherford, T. N. Woodard, T. V. I. Dailey, K. A. Schneider, T. A. Grotzinger, S. H. A. M~ Crump. ABSTRACT During Segment 3 of W-126-R, 2 Division of Wildlife Special Reports, 1 Division of Wildlife Division Report, and I Division of Wildlife Game Information Leaflet were final edited and published. One Division of Wildlife Special Report was submitted for preliminary editing and one more was nearly ready for preliminary editing at the segment's completion. Twenty-six publications were purchased and placed on file at the CDOW Research Center Library. Approximately 45 reports and short publications were requested and received free of cost or at minimal charge from federal and state agencies and from other sources. Twenty-nine theses were ordered on Interlibrary loan upon request from big game researchers. Eight literature searches were complete for various big game researchers. The Research Center Library located approximately 35 references per big game research employee and duplicated and delivered them to the requesting individuals. ��-77- BIG GAME PUBLICATION EDITING AND LIBRARY SERVICES R. B. Gill P. N. OBJECTIVE To provide a centralized support program for Big Game Research technical editing and library services so that Big Game Research scientists can allocate additional time to the conduct of actual research. SEGMENT OBJECTIVE To provide coordinated, efficient, economic editing and library services to all Colorado Big Game Research programs (Federal Aid Project W-126-R). SUMMARY OF SERVICES Publications Submitted for Editing and DOW Publication 1. Bear, G. D. 1979~ Evaluation of bighorn transplants in two Colorado localities. Colo. Div. Wildl. Spec. Rep. No. 45. 12p. 2. Carpenter, L. H., R. B. Gill, D. J. Freddy, and L. E. Sanders. 1979. Distribution and movements of mule deer in Middle Park, Colorado. Colo. Div. Wildl. Spec. Rep. No. 46. 32p. 3. Kufeld, R. C. 1979. History and current status of the mule deer population on the east side of the Uncompahgre Plateau. Colo. Div. Wildl. Div. Rep. No. 11. 24p. 4. Rutherford, W. H., W. D. Snyder. 1980. Habitat modification techniques for the benefit of wildlife. Colo. Div. Wildl. Div. Rep. No. 12. In Press. Publications purchased with W-126-R funds and placed in Research Center Libarary American Society of Animal Science, Western Section. 1979. Proceedings of the Annual Meeting, V. 30. University of Arizona, Dept. of Animal Sciences, 317p. Boyce, M. S., and L. D. Hayden-Wing (eds.). 1979. North American elk: ecology, behavior and management. Univ. Wyoming, Laramie. 294p. French, N. R. ed. 1979.. Perspectives in grassland ecology. Results and applications of the U.S./IBP Grassland Biome Study. SpringerVerlag, New York. 204p. �-78- Howard, G. S., and M. J. Samuel. 1979. Atlas of epidermal plant species fragments ingested by grazing animals. U.S. Dept. of Agric. Tech. Bull. 1582. 143p. Innis, G. S., ed. 1978. New York. 298p. Grassland simulation model. Springer-Verlag, Krebs, J. R., ed. 1978. Behavioural ecology; an evolutionary approach, edited by J. R. Krebs ann N. B. Davies. Blackwell Scientific Publ. Oxford. 494p. Pichard, D., and G. Robert. 1977. Forage nutritive value. Continuous and batch in vitro rumen fermentations and nitrogen solubility. Ph.D. Dissertation, Cornell Univ., Ithaca. 305p. Pielou, E. C. 1977. Mathematical ecology. Revised ed. of "An introduction to mathematical ecology". Wiley-Interscience, New York. 385p. The Sagebrush Ecosystem: Logan. 251p. A symposium, April, 1978. Utah State Univ., Sullivan, P. T. 1976. Genetic studies of the blood groups of Rhesus monkeys, (~~caca mulatta). Ph.D. Dissertation, Univ. of Wisconsin, Hadison. l31p. (Microfilm) Craighead, F. C., Jr. 1979. San Francisco, 261p. Track of the grizzly. Fleck, L. 1979. Genesis and development of Chicago Press, Chicago. 203p. Sierra Club Books, of a scientific fact. Univ. Long, F. M., ed. 1979. Proceedings Second International Conference on Wildlife Biotelemetry, July 30, 31, Aug. 1, 1979, Univ. of Wyo. IntI. Conf. Wildl. Biotel., Laramie. 259p. Maynard, L. A. et al. 1979. Book Company, New York. Animal nutrition. 602p. 7th ed. McGraw-Hill Mehrer, C. F. 1975. Some aspects of reproduction in captive mountain Felis concolor, bobcats Lynx rufus and Lynx canadensis. Ph.D. Dissertation, Univ. of North Dakota, 'Grand Forks. 142p. lions Mitchell, G. J. 1965. Natality, mortality and related phenoma in two populations of pronghorn antelope in Alberta, Canada. Ph.D. Dissertation. Washington State Univ., Pullman. 205p. North American Moose Conference and Workshop. 1978. Proceedings North American Moose Conference and workshop, Number 14. Halifax, Nova Scotia. 269p. Pasture and range plants. 1963. Phillips Petroleum Oklahoma. 176p. (R. Desilet) Company, Bartlesville, Rosenthal, G. A., and D. H. Janzen, eds. 1979. Herbivores: Their interaction with secondary plant metabolites. Academic Press, New York. 718p. �-79- Stonehouse, B., ed. 1978. Animal marking; recognition marking of animals in research. (Proceedings of the R.S.P.C.A. Symposium, London) McMillan Press, Ltd., London. 257p. Tanner, J. T. Tennessee 1978. Guide to the study of animal populations. Press, Knoxville. 186p. Van den Bosch, R. 1978. The pesticide City, New York. 226p. conspiracy. Doubleday, Univ. Garden Varland, K. L., A. L. Lovaas, and R. B. Dahlgren. 1978. Herd organization and movements of elk in Wind Cave National Park, South Dakota. USDI National Park Services, Natural Resources Report No. 13. 28p. Westra, R. 1978. The effect of temperature and season on digestion and urea kinetics in growing wapiti. Ph.D. Thesis, Univ. of Alberta, Edmonton. 175p. Yoakum, J. D., et a1., compo 1979. American pronghorn antelope; articles published in The Journal of Wildlife Management 1937-1977. The Wildlife Society, Washington, D.C. 244p. Hinman, R. A. 1960. Antelope populations in southwestern Utah, with special reference to golden eagle predation. M.S. Thesis, Utah State Univ., Logan. 66p. Publications obtained free or at low cost In addition to the books purchased with W-126-R funds, about 45 reports and short publications from state or federal agencies, and from other sources, were located, ordered, and obtained for use by big game research personnel. Theses obtained on interlibrary loan for use by researchers Barrington, M ..R. 1978. Habitat on the southern high plains. Stillwater. 165p. factors related to pronghorn productivity M.S. Thesis, Oklahoma State Univ., Bell, B. C. 1974. A comparison of age determination techniques for white-tailed deer from differing Louisiana soils. M.S. Thesis, Louisiana State Univ., Baton Rouge. 157p. Berger, J. 1978. Social development and reproductive strategies in bighorn sheep. Ph.D. Dissertation, Univ. Colo., Boulder. 157p. Boyer, K. B. 1976. Food habits of black bears (Ursus americanus) in the Banning Canyon area of San Bernardino National Forest. M.S. Thesis, California State Polytechnic University, Pomona. 63p. �-80- Brown, D. 1973. Some phenolic constituents of Artemisia tridentata Vaseyana. Ph.D. Dissertation, Univ. of Wyoming, Laramie. ssp. Bryant, F. C. 1977. Botanical and nutritive content in diets of sheep, angora goats, spanish goats, and deer grazing a common pasture. Ph.D. Dissertation, Texas A & M Univ., College Station. 103p. Conolly, E. J. Jr. 1949. Food habits and life history of the mountain lion (Felis concolor hippolestes). M.S. Thesis, Univ. Utah, Salt Lake City. 180p. DeBock, E. A. 1970. The behavior of the mountain goat, (Oreamnos americanus) in Kootenary National Park. M.S. Thesis, Univ. of Alberta, Edmonton. 173p. Ellis, J. E. 1970. A computer analysis of fawn survival in the pronghorn antelope. Ph.D. Thesis, Univ. of California, Davis. 63p. Goldberg, P. S. 1977. The use of infra-red scanning systems for the census of big game animals. M.S. Thesis, Utah State Univ., Logan. 68p. Hackett, E. J. 1978. Vital characteristics of white-tailed deer from selected areas of Mississippi. M.S. Thesis, Mississippi State Univ. SSp. Heet, G. C. 1977. Habitat utilization of white-tailed deer in southeastern Ohio as determined by radiotelemetry. M.S. Thesis, Ohio State Univ., Columbus. 86p. Hibben, F. C. 1936. A preliminary study of the mountain lion, Felis oregonensis. M.S. Thesis, New Mexico Univ., Albuquerque. Hudkins, G. G. 1976. Experimental cervine Sarcocystis infections in mule deer. M.S. Thesis, Oregon State Univ., Corvallis. 67p. Kauffeld, J. D. 1977. Availability to coyote predation of domestic Reno. 1S6p. of natural prey and its relationship sheep. M.S. Thesis, Univ. of Nevada, Kellyhouse, D. C. 1975. Habitat utilization and ecology of black bear in northern California. Unpublished M.S. Thesis, Humboldt State Univ., Arcata. 60p. Lund, R. C. 1979. Suitability of the cementum annuli technique for determining the age of white-tailed deer in New Jersey. Thesis, Rutgers Univ. M.S. �-81- Lynch, G. W. 1977. Nutritive value of forage species in the Rio Grande plain of Texas for white-tailed deer (Odocoileus virginianus texanus) and domestic livestock. M.S. Thesis, Texas A & M Univ., College Station. McCauley, M. N. 1977. Current population and distribution status of the panther (Felis concolor), in Florida. M. A. Thesis, Univ. S. Florida, Tampa. 58p. McFetridge, R. J. 1977. Strategy of resource use by mountain Alberta. M.S. Thesis, Univ. Alberta, Edmonton. 148p. goats in Mosha, G. T. 1976. Estimates of mule deer densities and browse conditions of the Sacramento Mountains. M.S. Thesis, New Mexico State Univ., Las Cruces. 48p. (Courtesy - T. Pojar). Osburn, W. S. Jr. 1958. Ecology of winter snow-free areas of the alpine tundra of Niwot Ridge, Boulder County, Colorado. Ph.D. Dissertation, Univ. of Colorado, Boulder. 77p. Shackleton, D. M. Dissertation, 1973. Population quality and bighorn sheep. Univ. Calgary, Calgary, Alberta. 227p. Ph.D. Sommerville, R. J. 1965. An evaluation of the 1961-1963 Alaskan brown and grizzly bear management program. M.S. Thesis, Univ. of Montana, Missoula. 117p. Springer, J. T. 1977. Movement patterns of coyotes in southcentral Washington as determined by radio telemetry. Ph.D. Dissertation, Washington State Univ., Pullman. 109p. Springer, M. D. 1977. The influence of prescribed burning of nutrition in white-tailed deer on the coastal plain of Texas. Ph.D. Dissertation, Texas A & M Univ., College Station. Trainer, C. E. 1971. The relationship of physical condition and fertility of female Roosevelt elk (Cervus canadensis roosevelti) in Oregon. M.S. Thesis, Oregon State Univ., Corvallis. 93p. Urbston, D. F. 1976. Descriptive aspects of two fawn populations delineated by reproductive differences. Ph.D. Dissertation, Virginia Polytechnic Institute, Blacksburg. 103p. as Zwank, P. J. 1978. Reduced recruitment in Utah mule deer relative to winter condition. Ph.D. Dissertation, Utah State Univ., Logan. 83p. �-82- List of literature searches Research Center Library Methods to detect Immobilization performed early pregnancy for Big Game Researchers in domestic by the or wild ungulates of game animals Coccidiosis Scrapie Aspen in relation Cambendazole to wildlife and Fenbendazole Lungworms Chemical Reference methods to repel deer document location and delivery In addition to the above listed services, the Research Center Library located about 550 references on request for the Big Game Research Section during this segment; 54 of these were obtained through interlibrary loans. These approximately 35 articles per Big Game Research scientist and technician Prepared we~ d.~icated by ~ \ \< \).J. ~ '.J':~ R. Bruce Gill Big Game Section delivered .... :>:. _: Chief \_Q to them. �-83- JOB PROGRESS COLORA=D~O State of Project No. July, REPORT _ W-126-R-3 Big Game Investigations Work Plan 2 Deer Investigations Job No. 1 Nutritional Quantifying the Capacity Period Covered: Personnel: 1980 Basis for of Winter Ranges to Support Deer July 1, 1979 through June 30, 1980 Len H. Carpenter, Steve C. Torbit, David J. Freddy, Larry 1.. Strong, Martin C. Fowler, Whitcomb M. Bronaugh, Kevin L. Berner ABSTRACT Two differ~nt experiments on nutritional characteristics of mule deer are described. One experiment deals wi.th determination of maintenance energy requirements of adult mule deer and presents weight pe rf o rmance values over an 8-week winter period for 3 groups of deer as related to f(lod intake. In vivo digestibilities for 15 deer during the 1979 winter are presented. Values for urine protein and urine volume also are presented for each group of deer. A separate experiment is described that presents body composition estimates for 9 mule deer on 3 feed levels and relates these estimates to food intake. An abstract of a M.S. Thesis is also presented that describes an experiment conducted to measure herbage yields in 10 separate habitat types in the mule deer critical winter range. Comparisons of herbage yield determined by remote sensing techniques and ground measurements are presented. ��-85- NUTRITIONAL CAPACITY BASIS FOR QUANTIFYING OF WINTER RANGES TO SUPPORT DEER Len H. Carpenter P. N. OBJECTIVE Develop procedures to support deer. for quantifying the capacity of Middle Park winter ranges SEGMENT OBJECTIVES 1. Estimate energy costs associated with selected 2. Develop sampling methodology for estimating availability of deer forage in winter. 3. Test the efficacy of a computer capacity of winter ranges. 4. Estimate energy requirements 5. Determine body composition simulation of pregnant activities of mule deer. annual production and model for estimating carrying does. of mule deer on various nutritional levels. METHODS AND MATERIALS Segment objectives 2 and 5 are being investigated by graduate research assistants Larry L. Strong and Steve C. Torbit respectively. An abstract of Strong's M.S. Thesis is presented in Appendix A and the yearly progress of Torbit's work is included in the following discussion. Work on objectives 3 and 4 was minimal during this segment. The ruminant nutrition computer model was updated as data were obtained from various aspects of the research effort as well as from published literature. No progress was made on objective 4 due to the lack of female mule deer. Most of the effort of the principal investigator was expended on Objective 1. Description of this work follows. Estimating Energy Requirements of Adult Mule Deer in Winter Methodology of this experiment closely followed that of Baker et al. (1979) and as further described by Carpenter (1979). For a detailed description of methods please refer to these sources. Work during this segment was essentially a repeat of work done during the past segment, but certain changes �-86- were made to supplement the study was conducted Colorado. and improve upon the experiment. This portion of at the Junction Butte Research Center near Kremmling, Nine adult mule deer were divided into 3 groups of 3 deer each. Six of these deer were the same as used in 1979 experiments. The 3 additional deer were the same age and background as the other 6 but had not been used in any previous nutritional trials. As in 1979, deer were ranked from lightest to heaviest and randomly allotted into 3 successive trios. The 3 treatments were feed levels and were as follows: 1) Ad libitum (as determined from 11 days of pretrial intake for all 9 deer);- 2) 2/3 of ad libitum, and 3) 1/3 of ad libitum. The 2/3 and 1/3 offerings were used in placeof the 1/2 and 1/4 offerings in 1979 attempting to ameliorate the severe weight loss of deer on the restricted intakes. Deer were maintained on these feed levels for 8 weeks as compared to 10 weeks in 1979. Measurements were made from January 7 until March 2, 1980. Beginning on January 21, one deer from each treatment was randomly assigned for 10 days to a digestion cage for complete nitrogen and energy balance measurements. The 10-day digestion trials were repeated beginning February 4 and again on February 18 to complete measurements for each of the 3 deer in the 3 treatments. Three days were allowed for acclimatization to the digestion cages only and no data (other than intake) were recorded. During the next 7 days all feces and urine were collected daily. Seven day collections were made in 1980 attempting to reduce some of the individual variation in daily intake and urine and feces output experienced in 1979. Every Monday during the experiment all deer were weighed. Procedures for handling urine and feces samples were the same as those previously described (Carpenter 1979). From the II-day pre-trial it was determined that mean ad libitum intake for the 9 deer was 1228 g + 37 (SE) g. Consequently deer receiving 2/3 ad libitum (Treatment B) re~eived 819 g daily and deer receiving 1/3 ad libitum "(Treatment C) received 409 g each day. In 1979 mean intake was 1165 g and the 50 and 25 percent levels were 583 and 291 g respectively. As was done in 1979, deer in the ad libitum treatment were offered 2500 g daily during the actual experimental period. Digestible and metabolizable energy and protein and dry matter digestibility coefficients will be determined for each deer as described by Baker et al. (1979). Results of the two winter experiments will be compared and if similar will be combined into one data set describing maintenance energy requirements of adult mule deer. Due to work loads in the Division of Wildlife Research Lab many of the analyses for 1979 or 1980 have not been completed, therefore certain results are pending. Body Composition Measurements Three food levels were offered to 3 groups of 3 mule deer used in this experiment. Each treatment (food level) group had 1 male castrate and 2 "females. Females were randomly chosen for each treatment. �-87- Estimation of body composition required animals that tolerated prolonged periods of confinement in metabolic cages. Mule deer fawns which had been trained to accept handling, confinement and collection of body wastes were used. This experiment was conducted at the Division of Wildlife's Wild Animal Research Facility in Fort Collins. Levels of feeding were determined in the following manner; body composition was estimated for each of 9 mule deer, these estimates served as input data for a ruminant nutritional simulation model developed by D. M. Swift of the Natural Resources Ecology Laboratory. Model experiments were conducted to ascertain what feeding levels of the ration would lead to specific weight losses. The ration fed was concentrated alfalfa pellets (Table 1). These modeling experiments produced predicted weight losses of 9 percent for the high intake group, 22 percent for the medium intake group, and 28 percent for the low intake group (Table 2). Gross energy intake fed per animal per day varied from 2650-3400 Kcal during the course of the 20 week trail (Table 2). Table 1. Analysis Composition analysis (Dry Matter of pelleted ration. Percent Basis) Dry Matter 90.59 Crude Protein 19.34 Ether Extract 2.84 Ash 7.86 Cell Wall Constituents 52.82 Acid-detergent 13.58 fiber 3.58 Lignin Apparent in vivo dry matter digestibility 72.07 4.60 Gross Energy Kcal/g Apparent digestible energy Kcal/g Apparent metabolizable energy Kcal/g 3.45 3.05 �-88- Table 2. Intake rates, predicted weight losses and actual weight for 9 mule deer on 3 food levels for 20-week period. Treatment Intake Predicted Weight Loss losses Mean Weight Loss High 3400 Kcal/day 9% 15.7% Medium 2800 Kcal/day 22% 23.6% Low 2650 Kcal/day 28% 32.7% During the course of the experiment, individual animals were maintained in isolation pens where the only available feed was a controlled offering of the pelleted ration. At 30 day intervals, animals were placed in metabolic cages for estimation of body composition. The first 3 days of cage occupancy were considered a pretrial period, allowing animals to become adjusted to the. cage environment. Body composition was estimated by an intravenous injection of tritiated water (HTO) and complete urine collection following procedures outlined by .Knox et al. (1969). To make an accurate and complete injection, animals were tranquilized with a combination of two drugs--M-99 and Rompun. Upon injection of HTO, M-50-50, an antidote, was administered and animals recovered in approximately 90 minutes. Fifteen hours after injection of the tracer, the first urine sample from each animal was collected. This period was allowed for complete equilibration of the HTO within the animal and also permitted total recovery from any metabolic effects of the tranquilization. Urine was collected every 12 hours for the next 7 days. Urine volume was measured and an aliquot of approximately 40 ml was frozen for assay. After a trial was completed, urine was assayed for tritium activity. This was accomplished by decolorization of the urine in 4.0 grams of activated charcoal, 50 ml of the supernatant was then placed in 20 ml, of a diocane based counting fluid, duplicate samples were counted on a Mark II Nuclear Chicago Liquid Scintillation Counter. Total body water was estimated from the logarithmic decay of tritium activity in the urine. Estimates of total protein and-total fat were made by applying relationships developed by Robbins et al. 1974 between total body water and these components. In addition to HTO assay, a blank urine sample was collected from each animal before injection of the tracer. These blank samples will be analyzed for total urinary nitrogen and creatinine. This analysis should provide further insight into the nutritional state of the animal. Levels of these waste products are directly related to deg~ee of �-89- protein catabolism occurring in the animal. This information combined with estimates of total protein stores allows a more complete understanding of this catabolic process. During this segment, a proposal was submitted to the National Science Foundation for acquisition of a whole carcass grinder. This proposal was funded, a grinder purchased and the equipment is currently being installed on the CSU campus. Three animals, one from each treatment, randomlY chosen, were euthuanized at the end of the experiment and a fourth died during the last phases of the trial. These animals were frozen and will be ground. Chemical analysis will be performed to determine actual body composition. These results will be compar-ed with tritium estimates and used to validate the in vivo technique. Final estimates of body composition will be based on the HTO technique, but will be corrected as necessary using results of the grinder study. RESULTS AND DISCUSSION Energy Requirements of Adult Mule Deer A summary of apparent dry matter digestion coefficients for the 1979 experiment was completed (Table 3). There were no significant differences (P ~ .05) in digestibility between treatments. Highest average digestibility was in Treatment A and lowest in Treatment C with overall digestibility being 69.62% 1.51 (SE). This is not statistically different from an overall apparent dry matter digestibility of 69.28 (SE + 1.50) determined for 13 mule deer fawns fed .the same ration during the winter of 1975 (Baker et a1. 1979). ± Table 3. Apparent dry matter digestibility of feed during winter of 1979. (%) for 15 mule deer on 3 levels Treatment A B C Deer % DDM 105 78.68 103 69.47 107 71.72 III 74.70 115 72.80 109 58.72 121 74.62 120 69.50 119 68.67 151 72.57 123 72.93 155. 60.08 154 60.00 179 71.83 162 67.97 X 72.11 71.31 65.43 3.18 0.77 2~54 SE All Deer Deer % DDM Deer X SE % DDM 69.62 1.51 �-90- An analysis of urine protein in samples collected in 1979 revealed no significant (P ~ .05) differences between treatments. Urine protein averaged 6.04 + 1.04 (SE), 6.65 + 1.14 (SE), and 5.10 + 0.86 (SE) percent for Treatments-A, B, and C deer respectively. The increase in daily urine volume for deer in Treatment C observed during the 1979 winter (Carpenter 1979) was not apparent in 1980 measurements (Table 4). Samples where snow dillution was suspected were deleted from the summary, resulting in unequal sample sizes. Five and 6 samples were deleted in 1979 and 1980, respectively. In 1980 greatest daily urine volumes were measured in Treatment B deer. Urine volumes for Treatments A and C were less in 1980 than in 1979. Across both winters average daily urine output for all deer in all treatments was 897.7 ± 42.9 (SE) mI. (Table 4). Nutritional aspects of these volumes will be better understood when all protein and energy balance measurements are completed. It is suspected that the less severe nutritional stress imposed on Treatment C deer in 1980 as compared to 1979 could account for the differences in urine volume of deer in this treatment between the 2 years. Results of urinary creatinine measurements could elucidate these factors. As in 1979, there were no significant differences (P ~ .05) in weekly intakes among the 3 deer in Treatment A. Average daily intake for the 8 weeks was 1178.8 ± 40.2 (SE) g or approximately 50 grams per day less than ad libitum intake measured during the II-day pre-trial. There were no significant differences in intake among weeks for the ad libitum deer in 1980. This intake was approximately 250 g greater tha;-was measured for Treatment A deer during 1979. Average daily intake of Treatment B deer was 737.2 ± 16.2 (SE) g while average intake for Treatment C deer was 400.2 ± 3.1 (SE) g. Table 4. Daily volumes of urine (ml) from mule deer on 3 levels of food for 1979 and 1980 winters. Samples with possible snow dilution deleted. Urine Volumes (ml/day Treatment Group ± SE) N 1979 N A 23 840.9 + 82.0 19 625.4 + 38.1 B 25 725.3 + 56.5 19 1093.9 + 98.6 C 22 1360.1 +156.6 19 734.1 + 48.5 70 962.8 + 67.3 57 817.8 + 46.5 127 897.7 + 42.9 Totals Grand Total (Both Years) 1980 �-91- Daily intake of Treatment A and B deer in digestion cages was considerably decreased. Treatment A deer consumed 486.0 + 52.6 (SE) g and Treatment B deer ate an average of 579.5 + 67.5 (SE) g while deer in Treatment C ate 406.7 ± 0.7 (SE) g daily duri~g the 10-day trials. This same pattern of decreased intake was measured in 1979. Treatment A deer in 1979 ate 654.4 + 71.4 (SE) g, and Treatment Band C deer consumed 576.5 ± 2.2 (SE) g and 288.5 + 0.8 (SE) g respectively. It appears that less nutritionally stressed deer (Groups A and B) can afford to avoid food during their confinements in the cages. In 1979 deer in all treatments significant decreased body weight (Carpenter 1979). In 1980, group A deer lost only 2.0% of their body weight during the 8-week measurement period while group Band C deer lost 10.3 and 16.1% body weight respectively (Fig. 1). This contrasts to 1979 measurements where Treatment B deer lost 17.1% and Treatment C deer lost 28.1% (Carpenter 1979). In 1979 Treatment A deer lost 132.3 + 24.5 (SE) g of body weight per day while Treatment Band C deer lost 145.1 + 9.7 (SE) g, and 287.6 + 29.2 (SE) g daily. In 1980, Treatment A deer lost-only 25.0 + 4.1 (SE) g ~f body weight each day and Treatment Band C deer lost 127~4 + 64.7 (SE) g and 194.0 ± 17.0 (SE) g each day. The reduced weight loss in Treatments Band C deer in 1980 as compared to 1979 is attributed to the less severe food restrictions for each treatment as well as the decrease in trial length from· 10 weeks to 8 weeks. Group A deer consumed about 250 g of feed more each day during 1980 than during 1979. However on the basis of grams of food eaten each week in relation to body weight, deer in Treatment A were very similar each winter (Table 5). This was also true for deer in Treatments Band C. It is also suspected that differences in weight performance between the 2 winters resulted from temperature. In 1979, average daily minimum temperature was -19.7 ± 1.0 (SE) C while in 1980 it was -13.6 + 1.8 (SE) C. Average daily maximum temperatures were -5.4 + 0.9 (SE) C-and 1.8 ± 0.7 (SE) C in 1979 and 1980 respectively. In contrast to 1979, when 2 deer in the low intake treatment died and 1 was removed from the experiment and refed all Treatment C deer in 1980 survived the 8-weeks. However, 1 deer from this treatment did die during a wet snow storm in the week following the experiment. As observed in 1979, deer in 1980 exhibiting the greatest weight losses consumed large quantities of hair from their sides and flanks. This resulted in the animal being most vulnerable to wet cold weather. Deer in digestion cages each week lost weight more rapidly than during other weeks. This was probably attributable to a more severe thermal environment in the digestion cages compared to isolation pens. Body Composition Measurements All laboratory analyses are not complete at this time, but body composition estimates for the first three trials have been made. Ration and feeding �Table 5. Weekly summary of metabolic body weights and food intake for mule deer on 3 levels of feed for 1979 and 1980 winters. 1980 1979 Average Kg BW Treatment Trial Week .75 G intake Kg BW .75 Treatment Average Kg BW Treatment .75 G intake Kg BW .75 Treatment A B C A B C A B C A B C 1 21.67 21.40 20.75 44.69 26.45 13.93 24.06 23.84 23.97 54.14 33.45 15.50 2 21.54 21.16 20.34 42.54 26.95 14.43 24.04 23.58 23.75 36.52 34.75 17.19 3 21.52 21.06 19.94 41.54 27.76 14.87 23.99 23.52 23.35 46.91 34.52 17.42 4 21.34 20.67 19.37 48.10 26.38 15.19 24.10 23.49 23.01 65.54 34.85 17.76 5 21.14 20.32 18.83 58.20 28.34 15.77 24.04 23.22 22.68 44.02 26.23 18.03 6 20.62 20.09 18.25 31.90 28.95 16.39 24.03 22.90 22.34 62.12 25.88 18.47 7 19.97 19.84 17.57 30.23 29.58 16.59 23.93 22.42 21.95 40.96 29.53 18.78 8 19.84 19.59 18.63 44.81 29.93 15.85 23.65 22.09 21.39 41.53 36.58 19.37 9 19.46 19.42 18.63 47.21 30.22 16.01 10 19.30 19.16 18.00 55.10 30.60 16.66 x 20.63 20.27 19.10 44.83 28.52 15.45 23.98 23.13 22.81 48.97 31.97 17.81 SE 0.3 0.3 0.5 2.8 0.5 0.4 0.2 0.3 0.6 3.7 1.8 0.5 I \0 N I �-93- regime yielded predictable weight loss responses. The high intake group simulated a set of animals which would not be in severe negative energy balance and would survive winter with minimal loss of body weight. The medium intake group represented animals experiencing a more severe weight loss characteristic of significant negative energy balance. The low intake group characterized animals which are in a profound catabolic state. Animals experiencing this degree of negative energy balance would not be expected to survive long periods. A trend in depletion of energy reserves for the first 60 days of the experiment was noted (Table 6). For animals in negative energy balance, fat reserves are initially depleted in an attempt to meet this energy deficit. However, protein depletion is apparently occurring before fat reserves are completely exhausted. This trend has important implications for the survival of the animal since protein catabolism has more deleterious effects on the animal's health than simple use of fat. Disease resistance and reproductive success may be severely compromised by extensive protein catabolism. All animals lost weight during the experiment with animals in the low intake group experiencing dramatic weight loss (Fig. 2). Animals in the medium intake group paralleled the weight loss exhibited by the low group with an apparent gain in weight at the end of the trial. This group exhibited the most variable loss of any group and the 1 animal which died was from this group. This animal lost considerable weight and the apparent gain at the end of the trial is an artifact of the death of this animal. The last 2 means were calculated from only the 2 surviving animals. LITERATURE CITED Baker, D. L., D. E. Johnson, L. H. Carpenter, O. C. Wa1lmo, and R. B. Gill. 1979. Energy requirements of mule deer fawns in winter. J. Wildl. Manage. 43(1):112-119. Carpenter, L. H. 1979. Middle Park deer study - deer habitat evaluation. Colo. Div. Wildl. Game Res. Div., Fed. Aid Proj. W-126-R-2, July, Part 1. p. 67-73. Knox, K. L., J. G. Nagy, and R. D. Brown. 1969. deer. J. Wildl. Manage. 33(2):389-393. Robbins, C. T., A. N. Moen, and J. T. Reid. 1974. while-tailed deer. J. An. Sci. 38(4):871-876. Prepared by J.._~=-,-::-I/_. (6_6~r~<;._ Len H. Carpentaf Wildlife Researcher • _ C Water turnover in mule Body composition of �Table 6. Preliminary body composition estimates for 9 mule deer on 3 food levels. Weight of protein and fat components for each deer and percent fat and protein tissue loss at the end of Day 70 as compared to pre-trial. Trial 2 Trial 1 Initial Protein (Kg) Fat (Kg) (Day 1 - Day 30) Protein (Kg) Fat (Kg) (Day 40 - Day 70) Fat (Kg) Protein (Kg) Treatment Deer High 62 9.82 6.54 9.07 5.53 8.41 4.71 High 63 11.58 8.05 10.24 7.12 9.32 5.86 High 64 9.32 5.86 8.66 5.02 8.24 4.52 15.15% % Tissue Loss Compared to Initial Estimates + 3.29 26.02% + 2.25 Medium 58 11.15 8.56 9.32 5.86 8.90 5.32 Medium 61 8.41 4.71 7.49 3.69 5.82 2.16 Medium 78 11.32 8.85 10.17 7.04 10.08 6.91 20.64% % Tissue Loss Compared to Initial Estimates + 8.11 37.97% +13.15 Low 60 10.90 8.16 9~16 5.65 7.32 3.50 Low 65 7.98 4.22 6.90 3.10 6.24 2.51 Low 76 10.99 8.31 9.32 5.86 8.06 4.31 % Tissue Loss Compared to Initial Estimates 27.10% + 4.52 48.58% + 6.78 I \0 .j::'I �-95- , Treatment A " ,, , 0-. -- ----0- - - --u., ,Treattnent ..•..•. B , 'D-_ ---~, ,, ,, w .~ 10 '0--:---0 « J: U 15 Treatment C 1 2 6 7 T~IAL WEEK Figure 1. Mean weekly weight change (percent) compared to initial weight of adult mule deer on 3 levels of food intake for 1980 winter. 8 �-96- o '"' ~ ffi 0... ......., ~ t::J ::x >~ -20 C) CQ :z: t-t h5 :z: g _~I~--~--~~--~--~---4--~ o 20 40 60 80 100 120 DAYS Figure 2. Mean monthly weight change (percent) of 3 groups of mule deer in body composttion study as compared to initial weight. �-97- APPENDIX A �-98- ABSTRACT ESTIMATING HABITAT PHYTOMASS PRODUCTION TYPES ON SAGEBRUSH OF STEPPE Methodology was developed for estimating phytomass production of 10 previously described and defined habitat types on the mule deer critical winter range of Middle Park, Colorado. Estimates of total, shrub, grass, and forb phytomass production were obtained during July and August of 1978 and 1979 on 3, 0.1 ha primary sample units for each habitat type. Phytomass estimation techniques varied between habitat types with changes in structure of shrub phytomass and physical characteristics of habitat types. Methods included double sampling clipped observations in conjunction with an electronic capacitance meter, herbage weight unit technique and dimension analysis, and simple random sampling employing clipped observations only. Spectral reflectance variables derived from microdensitometry of large scale (1:1200) 70 mm color infrared photography and gray scale panels of known reflectance, were used to develop phytomass estimation models for 4 habitat types. The accuracy of the remote sensing techniques WaS evaluated using phytomass estimate and spectral reflectance variables for 3, 65 ha areas dominated by Wyoming big sagebrush. Highly significant differences in total, shrub, grass and forb phytomass production were found between habitat types. No year effect or habitat type X year interaction was detected for phytomass production of any life form. Variances for phytomass production were heterogenous between habitat types and within habitat types between years. Total phytomass production of all habitat types was estimated with greater precision than phytomass of any of the life forms individually. Double sampling techniques had low relative costs and correlations with clipped observations were sufficient to reduce the variance of the mean phytomass estimate. A high positive correlation between spectral reflectance variables appeared spurious. An inverse relationship was found between phytomass production and reflectance. Linear relationships of red reflectance accounted for 56 and 59 percent of the variation in total phytomass production for 2 habitat types. Data illustrates soil reflectance greatly influences measurements of scene reflectance. Range in reflectance for the 3, 65 ha areas exceeded range in reflectance for the regression models generated from the 0.1 ha primary sample units. Estimates of phytomass predicted from regression models for the 3 large areas averaged 23 percent lower than phytomass estimates made on the ground. In future experiments, areas selected for development of phytomass estimation models should be large enough to encompass the range in reflectance expected for rangeland sample units being inventoried. �-99- July, 1980 JOB PROGRESS REPORT State of Colorado Project No. Big Game Investigations W-126-R-3 Work Plan No. 2 ----~----------- J~ ~. Deer Investigations 2 Snowmobile Harassment of Deer On Cold Winter Ranges Period Covered: Personnel: July 1, 1979 through June 30, 1980 D. Freddy, W. Bronaugh, K. Berner, M. Fowler, L. Carpenter. ABSTRACT During winters 1979 and 1980, marked mule deer were harassed by 1 or 2 people walking and I or 2 snowmobiles during 67 controlled harassment bouts. Observed overt reactions of deer were classified into 1 of 5 response classes ranging in severity from no reaction, to strong, purposeful movement. Maximum behavioral responses of deer were more severe for people walking than for snowmobiles. Threshold distances at which overt responses were elicited were about 417 m for people walking and 670 m for snowmobiles. Increasing severity of response was associated with a declining distance between stimuli. and deer. Deer tended to respond to people for longer time intervals than to snowmobiles over all response classes. When moving away from stimuli, deer tended to move greater distances over longer time intervals and intercept more elevational change when responding to people as compared to snowmobiles. Considerable data analysis awaits completion. ��-101- SNOWMOBILE HARASSMENT OF MULE DEER ON COLD WINTER RANGES David J. Freddy P. N. OBJECTIVE Evaluate whether snowmobile activity on winter ranges inhabited by deer decreases the ability of deer to survive winter by modifying activities of deer so as to significantly increase energy expended. SEGMENT OBJECTIVES 1. Ascertain activities of deer responding to prescribed harassment. 2. Evaluate reactions of deer to harassment in terms of energy costs. 3. Evaluate energy cost of harassment to deer in relation to total estimated energy budget of deer (concurring study). Investigations into effects of harassment on wild mule deer was initiated in winter 1979. The approach of this work was to conduct controlled harassment bouts whereby stimuli, both people walking and snowmobiles, followed routes designed to intercept neck-banded deer located prior to initiating a bout. Reactions of individual deer were then visually monitored by secluded observers. Work was conducted at the Colorado Division of Wildlife's Junction Butte Research Center located 3 miles south of Kremmling, Colorado. Junction Butte is an isolated mountain dominated by sagebrush vegetation (Artemisia tridentata wyomingensis). This vegetation as opposed to deciduousconiferous forests where other harassment studies have been conducted (Dorrance et al. 1975; Richens and Laviane 1978; Eckstein et al. 1979) offered the opportunity to visually observe overt reactions of deer responding to stimuli. The study area of approximately 5 km2 consisted of steep southerly and southwest-facing slopes flanked by a moderately rolling sagebrush flat. Elevations ranged from 2257 m to 2672 m. METHODS AND MATERIALS Mule deer were trapped and marked on Junction Butte during December 1978. and January and February, 1980. In 1978, 4 mature female deer, yearling or older, were equipped with telemetry neck-collars and 4 mature female were also neck-banded for individual identification. During January and February, 1980 4 mature females·were equipped with telemetry neck-collars while 7 adult females, I yearling male, and I male fawn were neck-banded. �-102- Because deer have been trapped and marked for several successive years up to 17 marked deer were potentially available for monitoring during harassment bouts. Selected deer were systematically harassed during January, February, or early March by I or 2 persons walking and 1 or 2 snowmobiles moving on prescribed routes. Harassment bouts were conducted weekly and up to 4 bouts were conducted within I day. Individual deer were often involved within successive bouts. Marked deer were selected for a bout based on the opportunity to harass tha~ deer and monitor its reactions. Reactions of deer were visually in 2 observation huts placed at Locations of huts allowed nearly by either I or both observ~rs. during bouts. and telemetrically monitored by observers strategic locations on Junction Butte. continuous observation of selected deer Observers remained out-of-sight to deer Overt reactions of deer were assigned to I of 5 classes: OR) no overt reaction, IR) mild alert reaction while bedded and animal remained bedded, usually noted by the animal turning its head towards stimuli with ears upright, 2R) mild alert reaction while up and active whereby animal's activity could be briefly interrupted, such as when feeding with momentary interruptions, 3R) moderate reaction whereby a definite change in animal activity occurred, such as from bedded to standing or cessation of feeding, 4R) strong reaction, whereby a change in animal activity was followed by strong purposeful movement. A gridded photograph of the study area denoting geographic and cultural points was placed in each observation hut allowing observers to locate banded deer and sources of stimuli during bouts. Observers noted locations of stimuli and banded deer, reactions of deer, and clock-times of locations and reactions using 7 X 50 binoculars, 15-40X spotting scopes, and stopwatches. Observations were tape recorded during bouts and later transcribed. Distances between deer and stimuli and distances deer moved and elevational changes of moving deer were determined from transcribed observations coupled with gridded topographic maps. Maps were 1:24,000 scale with 40 feet contour intervals. All distances and elevations were converted to metric equivalents. Stimuli were systematically alternated between bouts in insure equal numbers of bouts for each stimulus. Stimuli followed routes likely to intercept deer. Routes varied with location of deer and terrain and therefore, how stimuli approached deer was highly variable. Simuli started at a location distant from deer and decreased the distance to the point where deer were approached directly. The most direct route to target deer was not necessarily followed although stimuli were usually visible to deer throughout a bout. Degree to which deer were pursued was dictated by constraints of terrain, snow, and movements of deer during bouts. Snowmobiles were usually restricted to established trails because of terrain and snow conditions. These trails transected many of the areas used by deer within the study area. On occasion when snow conditions permitted; �-103- snowmobiles diverged from established trails. Attempts to chase deer with machines were made on only 3 occassions primarily because uneven terrain restricted situations where snowmobiles could be operated. Operators of snowmobiles were instructed not to stop machines or get off machines during bouts. Speed of snowmobiles was approximately 10 mph for all bouts. During most bouts, deer were exposed to snowmobiles on 2 passes as routes were circuitous in layout. By necessity, deer were exposed to snowmobiles other than during harassment bouts in order to maintain trails and other supportive endeavors. Snowmobiles used in 1978-79 were a 1976 Ski-doo 340 TNT and a 1972 Scorpion 440 Stinger and in 1979-80 a 1979 Polaris Apollo 340 and the 1976 Ski-doo 340 TNT were used.l People afoot had fewer terrain-related travel restrictions than snowmobiles, and therefore they attempted to intercept deer more often than snowmobiles. People afoot could be placed in terrain conditions which totally excluded persons riding snowmobiles. But people afoot also walked established snowmobile trails to compare the 2 types of stimuli on the same routes and when possible with the same deer. Bouts generally were initiated at the building facility on the study area and terminated when stimuli returned to the facility. People harassing deer maintained communication with observers via 2-way radios. Snow depth was measured and snow hardness was classified within the study area at weekly intervals. There were 4 transects having a total of 50 permanent sampling points in 1980 while only 2 of these transects with 20 sampling points were established in 1979. RESULTS AND DISCUSSION Data analyses are incomplete because all statistical comparisons have not been made. Statistical comparisons discussed in text were based on t-tests at a probability level of P 2 0.05. In 1979 and 1980, 27 and 40 harassment bouts were conducted respectively (Table 1). Seventy-four marked deer were encountered in the 1979 harassment bouts, and 160 were encountered in 1980 (Table 1). Duration of bouts ranged from 15 to 64 minutes. Both winters were considered moderate to severe. In 1979, snow depths exceeded 30 cm from January 15 through March 15. In 1980, snow depth was 20 cm during January and exceeded JO cm during February and March. Maximum snow depths were 40 cm in 1979 and 55 cm in 1980. Behavioral responses of deer were more severe for people walking than for snowmobiles (Table 2). During both years, 80% of the maximum responses elicited by people walking were classed as 4R while only 24% of the maximum responses prompted by snowmobiles were classified 4R. Considering only the stimuli which completly altered the nature of deer activities, people afoot elicited 3R and 4R responses 95 percent of the time. People riding snowmobiles elicited 3R and 4R responses only 44 percent of the time. Deer ITrade-names are listed only for readers information and does not consitute endorsement by the Colorado Division of Wildlife. �-104- were either more sensitive to people afoot and/or people afoot could approach deer more closely than snowmobiles because terrain was less restrictive to people afoot. Table 1. Number of harassment bouts conducted for each type of stimuli, numbers of deer involved, average duration of bouts, and average ambient temperature, 1979-80. Stimuli 1979 Bouts Conducted Number Marked Deer Involved Duration of Bouts Ambient Temp. + °c Total Deer Involved (Minutes) X SE X + SE One person walking 6 19 173 62 + 9 - 1 + 2 Two persons walking 7 23 226 55 + 10 - 1 + 1 One Snowmobile 8 15 223 25 + 5 - 6 + 3 Two Snowmobiles 6 17 129 41 + 9 - 3 + 3 27 74 751 One person walking 10 53 192 64 + - 3 + 2 Two persons walking 11 42 229 46 + 5 One Snowmobile 10 39 112 17 + 3 - 6 + 2 Two Snowmobiles 9 26 101 15 + 2 - 4 + 2 40 160 634 Totals 1980 Totals 5 - 4 + 2 �-105- Table 2. Frequency of maximum individual behavioral responses of marked deer responding to 4 types of stimuli, 1979-1980. Maximum Behavioral Response Individual Deer During Bout Bouts Marked deer Involved One person walking 6 Two persons walking Stimuli 1979 1980 OR lR 2R 3R 4R 19 0 0% 0 0% 0 0% 1 5% 18 95% 7 23 0 0% 0 0% 0 0% 2 9% 21 91% One person walking 10 53 1 2% 5 9% 0 0% 1 19% 37 70% Two persons walking 11 42 1 2% 0 0% 0 0% 8 19% 33 79% 34 137 2 1.5% 5 3.6% 0 0% 21 109 15.3% 79.6% One snowmobile 8 15 Two snowmobiles 6 17 One snowmobile 10 Two snowmobiles Totals 1979 1980 Totals 1 7% 3 20% 5 33% 0 0% 6 40% 2 12% 3 18% 1 6% 4 23% 7 41% 39 6 15% 10 26% 3 8% 13 33% 7 18% 9 26 1 3% 17 65% 2 8% 3 12% 3 12% 33 97 10 10.3% 20 20.6% 23 23.7% 33 34.0% 11 11.4% Distances at which deer reacted .to stimuli varied with stimuli and behavior response class (Table 3; Fig. 1). The OR response class in Table 3 and Figure 1 represents the distance at which deer behavior changed from "no overt response" to 1 of the 4 other response types. This distance can be considered a threshold distance and was approximately 417 m for people walking and 670 m for snowmobiles. Snowmobiles tended to provoke milder responses by deer at greater distances than people during both years. However, in 1979 both snowmobiles and people prompted a 3R response, or a change in deer behavior, at a similar distance of approximately 320 m. In 1979, 4R responses tended to occur at greater distances for people walking than for snowmobiles. In 1980, distances at which 3R and 4R responses occurred appeared similar for people walking and snowmobiles (Fig. ~; Table 3). �-106- Comparing response distances within each response class, there were no significant differences between 1 and 2 people walking in 1979 or in 1980. A similar comparison for 1 and 2 snowmobiles also yielded no significant differences in 1979 or 1980 (Table 3). Table 3. Mean distances (m) at which behavioral responses of adult female deer were elicited by people walking and snowmobiles, 1979-1980. Stimuli OR lR ResEonse Class 2R 3R 4R 1979 One person walking SE n Two persons walking SE n One snowmobile Two snowmobiles X X X SE n X SE n 435 16 98 446 39 5 376 27 39 351 25 46 311 29 51 440 26 70 421 113 7 413 52 23 317 34 42 262 38 41 766 53 52 561 106 10 572 70 14 288 56 9 177 44 9 638 49 44 637 106 8 420 76 12 335 74 10 126 50 8 393 13 254 467 26 49 424 32 50 280 16 118 200 25 85 399 17 196 392 30 37 440 54 19 309 17 102 234 26 63 614 31 147 650 55 34 590 67 38 406 68 22 153 36 6 661 63 76 526 50 32 477 192 3 387 141 7 253 93 4 1980 One person walking Two persons walking One snowmobile Two snowmobiles X SE n X SE n X SE n X SE n �1979 0 1M ."Z eJ 6 0 A. 5 ." w_ ~ ·0 4 .•. 0 0- • " ~ ~3 I 1980 6 I 0 I • ~E *'''~ Z-2 c ~ -" ." 1 L I i i OR 1R 2R i I 3R 4R RESPONSE I I OR . i 1R I I 2R 3R 4R CLASS Figure 1. Average response distances of adult female mule deer to people walking and snowmobiles for 5 behavioral response classes, 1979-1980. Symbols represent: () 1 person t'lalking3 t:, 2 persons walking~ 0 1 snowmobilej f:t. 2 snowmobiles. See text for discussion of "ORr esponse" distance 0 I 0 "-J I �-108- Duration of behavioral responses to stimuli was defined bY' 2 methods: 1) Duration of a distinct response within a bout was the time from beginning to end of a response class. There were several distinct responses for each deer within each bout and multiple entries for the same response class could occur for each deer in each bout (Table 4), and 2) total time each deer spent in each response class throughout a bout. This value represented the sums of the distinct responses defined above (Table 5). Table 4. Average duration (seconds) of distinct behavioral responses of individual deer for each stimuli, 1979-1980. More than I distinct response and response type could occur per bout per deer. Values are for adult female deer. Duration (sees) of response time IR 2R 3R 4R Stimuli 1979 One person walking SE 143 44 n 5 Two persons walking X SE 174 60 n X One snowmobile 118 16 39 146 15 45 189 21 51 9 97 18 21 171 17 40 297 31 41 222 62 10 78 21 16 72 14 12 43 12 SE 238 63 8 141 23 10 116 28 n 62 11 12 X 184 28 48 80 10 51 149 16 123 100 12 87 n 146 36 37 55 14 21 150 14 102 87 11 60 65 26 10 X SE n Two snowmobiles 1980 X One person walking SE Two persons walking SE One snowmobile Two snowmobiles n X X 73 52 SE 9 8 n 35 38 68 11 22 X 86 19 53 SE 9 9 n 31 4 14 10 9 8 23 6 4 �-109- Table 5. Average total time (seconds) spent by individual deer in each behavioral responses category during an entire harassment bout, 1979-1980. .Values are for adult female deer. Duration (sees) of ResEonse Type 3R 4R lR 2R Stimuli 1979 One person walking SE n Two persons walking SE n One snowmobile Two snowmobiles 1980 One person walking Two persons walking One snowmobile Two snowmobiles X X X SE n X SE n X SE n X SE n X SE n X SE n 179 39 4 383 58 12 386 65 17 535 90 18 224 73 7 159 26 13 359 56 19 .580 91 21 443 274 5 124 41 10 172 50 5 65 17 6 475 158 4 106 37 7 157 24 9 132 34 7 353 74 25 205 32 20 479 63 37 302 56 29 339 81 16 82 21 14 462 61 33 204 48 25 196 33 13 131 29 15 93 10 16 78 23 5 167 24 16 25 13 3 76 26 35 10 7 3 There were no significant differences in duration of distinct responses within each response class between 1 and 2 people walking in 1979 or 1980. There were also no differences in duration of distinct lR and 2R responses for land 2 snowmobiles in 1979. However, 2 snowmobiles did cause significantly longer 3R and 4R responses than 1 snowmobile in 1979. In 1980, there were no differences in duration of distinct responses within each response class between land 2 snowmobiles. Data from 1979 suggests people walking cause longer 3R and 4R responses than snowmobiles. In 1980, people walking tended to cause longer behavioral responses than snowmobiles in all response classes (Table 4). �-110- There were no differences in total time spent by individual deer within 1R, 3R, and 4R responses between 1 and 2 persons walking in 1979 or 1980 (Table 5). However, in both years, more time was spent in a 2R response due to 1 person walking than to 2 persons walking (Table 5). No differences in time within each response class were detected for 1 and 2 snowmobiles in both years. Total time spent in 3R and 4R responses tended to be greater for people walking than for snowmobiles during both years. When exhibiting a 4R response, deer tended to move greater distances for longer time intervals and intercept more elevational change when responding to people as opposed to snowmobiles (Table 6). Velocities of deer moving in response to people ranged from 1.2-1.8 m/sec. Velocities of deer moving away from snowmobiles varied from 0.8-4.4 m/sec (Table 6). There were no differences in distances moved and time spent moving in a 4R response between 1 and 2 persons walking within 1979 or 1980. Similarly, there were no differences in distances moved or time spent moving in a 4R between land 2 snowmobiles within both years. In 1979, deer moved greater distances when responding to 1 person walking as compared to 1 snowmobile, and also for 2 persons walking as compared to 2 snowmobiles. However, these differences did not occur in 1980. Total time spent by deer in a 4R in 1979 was also greater for land 2 persons walking as compared to land 2 snowmobiles, respectively. These differences did not occur in 1980 (Table 6). LITERATURE CITED Dorrance, M. J., P. J. Savage, and D. E. Huff. 1975. Effect of snowmobiles on white-tailed deer. J. Wildl. Manage. 39(3):563-569. Eckstein, R. G., T. F. O'Brien, o. J. Rongstad, and J. G. Bollinger. 1979. Snowmobile effects on movements of white-tailed deer. A case study. Environ. Consv. 6(1):45-51. Richens, v. B~, and G. R. Lavigns. to snowmobiles and snowmobile Naturalist 92(4):334-344. 1978. Response trails in Maine. of white-tailed deer Canadian Field- �Table·6. Total distance moved, time to move, and elevational change intercepted during a harassment bout by individual deer exhibiting 4R behavioral responses, 1979-1980. Values are means + standard errors for adult female deer. Stimuli Year Number Deer Responses One person walking 1979 18 856 + 109 65 + 16 46 + 7 1980 29 476 + 82 60 + 12 27 + Two persons walking 1979 21 711 + 110 1980 25 One Snowmobile 1979 1980 6 5 Time to Move (sees) Velocity (m/sec) 110 + 16 541 + 90 1.6 6 88 + 14 258 + 48 1.8 14 + 4 70 + 15 84 + 15 578 + 90 1.2 460 + 90 56 + 10 21 + 9 77 + 14 200 + 43 2.3 257 + 120 148 + 14 20 + 9 2 + 2 18 + 18 7 + 3 38 + 21 9 + 2 60 + 18 81 + 24 4.3 1.8 Distance Moved (m) Elevation changes (m) Positive Negative Total ..-....I I Tw.o Snowmobile 1979 1980 7 3 103 + 20 155 + 28 12 + 5 24 + 14 2 + 2 0 14 + 4 24 + 14 132 + 34 35 + 10 0.8 4.4 ��July, 1980 -113- JOB PROGRESS REPORT Colorado State of Project No. W-126-R-3 Big Game Investigations Work Plan No. 2 ----~------------ Deer Investigations Job No. 3 Estimating Parameters of Piceance Basin Deer Population Dynamics Period Covered: Personnel: July 1, 1979 through June 30, 1980 R. M. Bartmann, and Dr. L. H. Carpenter ABSTRACT The deer banding data have been put on punch cards for computer summary. The 1980 winter deer population estimate of 15,428 + 3,458 (P < .10) is about ~ of the 1979 estimate due largely to a high winter loss~ The non-random and stratified random quadrat methods for estimating herd structure produced similar results for the second year. The buck:doe (7:100) and fawn:doe (50:100) ratios estimated from the quadrat method were both lower than last year's while precision of both estimates was slightly poorer. The estimated preseason production of 6,104 fawns is less than ~ the 1979 estimate. The hunter harvest estimate of 2,536 deer was down about 75% from 1978. ��-115- ESTIMATING PARAMETERS OF POPULATION DYNAMICS R. M. Bartmann P. N. OBJECTIVE To develop and test a method for estimating density of over-winter mule deer populations in the Piceance Basin. SEGMENT OBJECTIVES 1. To analyze and publish the results of Piceance Basin deer distribution studies. 2. To estimate the size of the wintering deer population in Game Management Unit 22. 3. To estimate the sex and age structure of the wintering deer population in Game Management Unit 22. 4. To estimate the annual productivity of the Piceance deer population. 5. To estimate the annual hunter harvest of the Piceance deer population. METHODS AND MATERIALS Deer Banding See Bartmann (1972). Deer Density See Bartmann (1974a). Population Structure See Bartmann (1979). Productivity See Bartmann (1975). Hunter Harvest See Bartmann (1974b). �-116- 2 Table l. Number of deer counted on 120 ~-mile2 (0.6 km ) quadrats on the Piceance winter range in Game Management Unit 22, January 12-16, 1980. Quadrat Deer Quadrat Deer Quadrat 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 22 5 0 6 27 6 0 44 1 0 13 10 25 4 0 6 0 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 9 6 13 5 0 7 11 0 0 0 0 0 0 8 0 0 0 0 3 0 13 0 0 0 2 4 0 0 0 0 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 '78 79 80 81 82 83 84 85 86 87 88 89 90 3 10 8 0 0 20 0 12 0 23 0 0 4 n = LX = X s = = 120 691 5.8 8.56 = c.v.= sx 0.78 148% Deer Quadrat 0 4 9 1 12 1 15 0 0 0 0 0 0 0 10 14 7 5 0 4 0 4 0 0 16 0 0 0 8 1 90% Confidence Interval 5.8 ± (1.658) (0.78) = 5.8 Total POEulation Estimate 15,428 ± 3,458 91 92 93 94 95 96 97 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 ± 1.3 Deer 12 3 0 3 19 2 7 0 0 1 0 0 0 6 0 0 17 16 20 0 4 37 23 19 0 8 29 24 0 0 �-117- RESULTS AND DISCUSSION Deer Banding Results of the deer banding work have been sent to Dr. Gary C. White of the Environmental Services Section, Los Alamos Scientific Laboratory, Los Alamos, New Mexico. Dr. White has put all the banding information on punch cards for computer summar Laat Lon , Several outputs have been generated to "debug" the program, but little else was accomplished towards data analysis during the year. Deer Density The 1980 winter deer census was conducted January 12-16. Counting conditions were generally poor as south exposures over much of the basin were snow-free. Observers were the same for the third consecutive year; me and Dr. L. H. Carpenter. It was necessary to use a Bell Jet Ranger helicopter in lieu of a Hiller 12EJ3 with Soloy turbine conversion. An advantage was less observer fatigue as the Jet Ranger provides more comfort. A disadvantage was that I observer had to sit in the rear seat. This necessitated a change in quadrat flight patterns which contributed to the increase in the mean time per quadrat from 5.8 minutes in 1979 to 7.4 minutes in 1980. 2 2 Only 691 deer were counted which converts to 23.2 deer/mile (9.0 km ) or 15,428 + 3,458 total deer (90% confidence level) on the Piceance winter range (Table 1). This estimate is about ~ that of last year due largely to a high winter mortality. In 1979, the census estimate fell short of the predicted population by 8,000 deer. While it was assumed the 1979 population should have increased over that of 1978 instead of declining, the estimate seems .reasonable in light of this year's data. The 1980 predicted population, based on the 1979 census estimate, is within 2,800 deer of the estimate and falls within the 90% confidence interval (Table 2). Population Structure Results of the comparison of the non-random and stratified random quadrat methods for estimating herd structure paralleled those of last year. Buck:doe and fawn:doe ratios were similar between methods (Table 3), although both were lower than those for 1979. About 2~ times more deer per hour were classified by the non-random method. If only flying time on quadrats is considered, however, this difference is greatly reduced. Precision of both the fawn:doe and buck:doe ratio estimates, + 12% and + 29% (P < .10), respectively, was slightly poorer this year. Sinc; deer distribution-seemed typical for that time of winter, reduced deer numbers basin-wide may have contributed most to the change. Productivity The post-season fawn:doe ratio of 50:100 was adjusted for the estimated. antlerless harvest to yield pre-season ratio of 49:100 and a pre-season �-118- Table 2. Calculations to predict the 1979-80 winter deer population in Game Management Unit 22. Est. 1978-79 winter pop. Bucks Does Fawns Total 2,204 16,961 9,829 28,994 571 4,459 11,302 16,332 1,633 12,502 0 14,135 0 0 1,633 12,502 Est. 1978-79 winter mortality 1979 summer population Fawns apportioned 50:50 Adjusted 1979 summer pop. 1979 fawn prod. est. 49 fawns: 100 does 14,135 6,104 1979 pre-season pop. 1,633 12,502 6,104 20,595 1979 Harvest estimate 2,149 356 31 2,536 0 12,146 6,073 18,219 820 11,605 5,794 18,219 Predicted 1979-80 winter pop. Predicted population corrected for 1979 post-season buck:doe: fawn ratio (7:100:50) Table 3. Comparison of non-random and stratified random quadrat methods for estimating the sex and age structure of the mule deer population in Game Management Unit 22, December 1979. Parameter Total flying time Non-Random 14.0 hours Method Stratified Random Quadrats 15.7 hours Total flying time on quadrats 9.2 hours Total deer classified 1,433 634 102 40 Deer classified per hour Mean time per quadrat 11.5 min. Total bucks 49 29 Total does 908 404 Total fawns 476 201 Buck:doe ratio 5: 100 Fawn: doe ratio 52:100 7 + 2:100, 50 ~ 6:10~(P ~ .10) �-119- production of 6,104 fawns. This is less than ~ of the estimated 1978 production due in part to fewer does in the population and to a lower fawn:doe ratio. Hunter Harvest The 1979 regular deer seasons were antlered-only with the separate season October 13-17 and the combined season November 3-13. One thousand either-sex permits were issued for Units 11 and 22 combined which is a reduction from the 5,500 allowed for Unit 22 alone last year. The archery season was either-sex from September 1-23 and the muzzle-loader season antlered-only from September 8-16. The total estimated harvest in Unit 22 was 2,536 deer which is about ~ of the 1978 harvest (Table 4). This was mainly due to reduced deer numbers and, to a lesser extent, fewer either-sex permits. Table 4. Summary of the 1978 deer harvest in Unit 22 for all seasons.ll Season Bucks Adult Females Archery 4 Muzzle-loader 9 Separate Antlered-only 671 Either-sex 46 Combined Antlered-only Either-sex All o Does Adult Females 19 o o Total 23 9 671 25 231 o 302 1,391 1,391 28 3 106 3 140 2,149 28 356 3 2,536 l/All figures include an arbitrary inflation of 25% to acknowledge wounding loss and illegal kill. Reduced deer availability is reflected in the 29% Success for antlered-only hunters which is down from the 58% estimated in 1978. It is also reflected in the percentage of yearlings field-aged at the Idaho Springs check station, 0% for females and 14% for males compared to 22% and 59%, respectively, in 1978 (Table 5). �-120- Table 5. Deer check station results for the 1979 separate and combined seasons in Game Management Unit 22. Sex Hale Female Mature Yearling Fawn 164 28 4 52 248 57 0 3 10 70 ~_----:=--.._Prep ared by _,_(---,-;-,':I_·'·-,--=-/_?/(_/:..:..~_.I_.'.:..;-_:r~"'=. t~._-~_';:;;'," R. M. Bartmann \Vildlife Researcher C Unknown Total �-121- July, 1980 JOB PROGRESS REPORT State of Colorado --~~~~~--------- Project No. W-126-R-3 Big Game Investigations Work Plan No. 2 ---.;::;._------ Deer Investigations Job No. 4 Forage Preferences of Piceance Basin Deer Period Covered: Personnel: July 1, 1979 through June 30, 1980 R. M. Bartmann, T. G. Baumann, P. H. Neil, W. Erickson, D. L. Baker, N. T. Hobbs, M. Stevens, J. Ritchie, S. C. Torbit, D. Butler, L. Bridges, C. Perron, T. Beavers, and D. Reichert. ABSTRACT Rough drafts of 2 publications of results of previous tame deer food habits technique development research are being revised. Nine tame mule deer were maintained in the Little Hills pastures until September, 1979 when I disappeared. Another disappeared after the March 1980 trials. One hundred ninety-four new plant species from the Piceance basin were added to the herbarium. Six sets of foraging trials were completed on 2 study areas from September, 1979 through April, 1980. Preliminary analysis of the 1978-79 forage trial data show browse as a major diet component early in the winter. It was slowly replaced by forbs as the winter progressed until April when grass became an important forage. The in vitro digestible dry matter content was estimated for forage samples from the 1978-79 winter trials. ��-123- FORAGE PREFERENCES OF MULE DEER IN THE PICEANCE BASIN R. M. Bartmann P. N. OBJECTIVE To identify major forage species in the winter-long diets of mule deer on the Piceance pinyon-juniper winter range and to estimate the nutritional characteristics of the diets. SEGMENT OBJECTIVES 1. Analyze data from previous food habits technique development work (Project W-38-R, Work Plan 16, Job 5 and W-126-R, Work Plan 2, Job 4) and publish pertinent results. 2. Maintain and train tame deer.' 3. Prepare an herbarium of plants from the Piceance Basin. 4. Conduct deer foraging trials. 5. Estimate forage quality. METHODS AND MATERIALS Publication of Previous Research Rough drafts of 2 publications from previous food habits technique development work were prepared and sent to co-authors for comment. drafts were returned and rewrites are in progress. Both Tame Deer Maintenance and Training Nine tame deer were kept in the Little Hills pastures on native forage at all times except during foraging trials and the deer hunting season. One deer disappeared and was assumed dead from unknown causes just prior to the September, 1979 trials. Another deer wandered away from the group as they were being moved back to the pasture after the March, 1980 trials and has not been seen since. During the summer, each deer was given about 1/3 kilogram of concentrate 3 times per week. After the trials started in September, about ~ kilogram of alfalfa hay was added to the concentrate ration. Free water was provided at all times except when snow was available. �-124- Plant Collections Plants were collected throughout the Piceance basin during summer, 1979. Tentative identifications were verified by Dr. Dieter H. Wilken, Curator, Colorado State University Herbarium. All specimens were mounted and labeled. Foraging Trials With 2 exceptions, methods for conduct of foraging trials were similar to those used last year (Bartmann 1979). As the deer have become older, they have become more independent and do not respond to handlers as well. This winter they could not always be led away from the temporary pens during grazing trials, so no effort was made to influence the starting direction of each trial as before. The loss of I deer in March necessitated a reduction in the number of trials from 8 to 6 per site in April. Forage Quality In vitro digestible dry matter was estimated for the 1978-79 forage samples during March, 1980 using procedures of Tilley and Terry (1963) as modified by Pearson (1970). Four deer were live-trapped at Little Hills and transported to Fort Collins as rumen fluid donors. Forage samples from the September and November, 1979 and January, 1980 trials have been dried and stored in glass jars. Samples from the last 3 sets of trials are still frozen in plastic bags. In vitro and protein determinations for all the 1979-80 winter samples are scheduled for November, 1980. RESULTS Plant Collections There were 194 new plant species added to the herbarium during 1979 (Table 1). Several others need to be collected at different development stages for positive identification. Foraging Trials Bite/weight conversions were applied to the 1978-79 winter deer forage selection data. Forage category composition based on bites and forage weight differed little except when pinyon pine (Pinus edulis) and Utah juniper (Juniperus osteosperma) were taken in more than token amounts. Bite weights of these 2 species were about 6 to 9 times greater than those for most other species. The greatest effect on diet composition, increase in occurred in February on the pinyon-juniper-sagebrush study area where bites of pinyon-and juniper increased in the browse forage category by 16%. �-125- Table 1. Plant species collected in the Piceance Basin during summer, Trees and Shrubs Acer glabrum Alnus tenuifolia Artemisia spinescens Chrysothamnus parryi Cornus stolonifera Grayia spinosa Holodiscus microphyllus Juniperus communis Populus fremontii Ribes lacustre Ribes leptanthum Ribes mogollonicum Salix monticola Salix rigida Sambucus racemosa Graminoids Agropyron caninum Agropyron dasystachyum Agrostic exarata Agrostis stolonifera Bromus erectus Carex aquatilis Carex atherodes Carex emoryi Carex foena Carex geophila Carex geyeri Carex vallicola Dactylis glomerata Distichlis stricta Festuca dasyclada Glyceria striata Hilaria jamesii Juncus balticus Juncus bufonius Juncus marginatus Phleum pratense Poa sandbergii Poa scabrella Stipa columbiana Stipa lettermanii Forbs and Lower Plants Aconitum columbianum Agastache urticifolia Allium cernuum Allium geyeri Allium textile Ambrosia acanthicarpa Angelica _ampla Angelica pinnata Aquilegia caerulea Aquilegia elegantula Aquilegia micrantha Arctium minus Asclepias asperula Asclepias cryptoceras Aster engelmannii Aster glaucodes Aster hesperius Astragalus bisulcatus Astragalus flavus Astragalus kentrophyta Atriplex heterosperma Atriplex hortensis Balsamorhiza hispidula Beckmannia syzigachne Bidens cernua Brickellia scabra Calylophus hartwegii Cardamine cordifolia Catabrosia aquatica Ceratocephalus testiculatus Ceratophyllum demersum Chenopodium dessicatum Cicuta douglasi~ Cirsium calcareum Cirsium scapanolepis Claytonia lanceolata Conium macula tum Crepis occidentalis Crepis runcinata Cryptantha flava 1979. �-126- Table 1. Plant species collected (continued) in the Piceance Forbs and Lower Plants Cryptantha stricta Cymopterus acaulis Cystopteris fragilis Delephinium ramo sum Dugaldia hoopesii Echinoceras triglochidiatus Eleocharis palustris Epilobium angustifolium Equisetum arvense Erigeron glabellus Erigeron subtrinervis Eriogonum hookeri Eriogonum intermontanum Gaillardia aristata Galium boreale Galium triflorum Gayophytum ramosissimum Geranium fremontii Geranium richardsonii Geum aleppiuum Geum macrophyllum Gilia pinnatifida Gilia sinuata Glossopetalon meionandra Glyceria striata Relianthus nuttallii Hippuris vulgaris Iva axillaris Lemna minor Lemna trisulca Lepidium campestre Lesquerella alpina Limosella aquatica Lithophragma glabrum Lithospermum incisum Lomatium dissectum Lupinus ammophilus Lupinus brevicaule Lygodesmia grandiflora Machaeranthera canescens Machaeranthera leucanthemifolia Machaeranthera venusta Madia glomerata Malcomia africana Medicago sativa Melilotus alba Mentha arvensis Mentzelia densa Mertensia arizonica Basin during summer, (continued) Mirabilis multiflora Monolepis nuttalliana Myosurus aristatus Myriophyllum spicatum Najas guadalupensis Nemophila breviflora Osmorhiza chilensis Osmorhiza depauperata Osmorhiza occidentalis Pedicularis grayi Phacelia demissa Phacelia neomexicana Phacelia sevicea Physaria floribunda Plantago major Plantago patagonica Polemonium foliosissimum Polygonum amp hi bum Polygonum persicaria Potamogeton berchtoldii Potamogeton pectinatus Potamogeton pusillus Potentilla gracilis Puccinellia airoides Ranunculus aquatilis Ranunculus cardiophyllus Ranunculus glaberrimus Ranunculus macounii Ranunculus scleratus Rorripa nasturtium-aquaticum Rudbeckia laciniata Rudbeckia occidentalis Rumex hymenosepalus Sagittaria cuneata Scrophularia lanceolata Scirpus americanus _~cirpu~ ?evadensis Scirpus vallidus Sedum lanceolatum Senecio mutabilis Senecio serra Sidalcea candida Sidalcea neomexicana Silene menziesii Sisyrinchium angustifolium Smilacina racemosa Solidago canadensis Stachys palustris Suaeda intermedia 1979. �-127- Table 1. Plant species collected (continued) in the Piceance Forbs and Lower Plants Sullivantia purpusii Thalictrum fendleri Thelypodium integrifolium Thermopsis rhombifolia Thlaspi arvense Townsendia incana Typha latifolia Triglochin maritimum Basin during summer, (continued) Valeriana edulis Valeriana occidentalis Verbena Bracteata Veronica americana Veronica anagallis-aquatica Viola canadensis Viola nuttallii Zannichellia palustris 1979. �-128- Browse was the major diet component on both study areas during early and mid-winter (Table 2). As the browse percentage declined through the winter, forbs showed a compensatory increase until April when green grass was consumed in large quantities on both areas. Table 2. Perc.ent composition of forage classes in winter diets selected by 8 tame mule deer on 2 study areas on the Piceance winter range, September-April, 1978-79. Total number of bites are in parenthesis. Study Area Forage Class September Diet Composition (% by weight) November February March (42,681) (17,104) (14,001) April (19,938) (44,021) Pinyon- Browse 94 96 82 59 41 Juniper- Forbs 6 4 15 36 18 Mixed Graminoids 3 5 41 < < 1 1 Browse (46,054) (15,842) (10,549) (20,904) (67,594) Pinyon- Browse 83 96 61 41 18 Juniper- Forbs 16 2 17 46 12 Sagebrush Graminoids 1 2 22 13 70 Six sets of foraging trials were completed during the 1979-80 winter on the 2 study areas. The data are on forms ready for key-punching and computer summary. Forage Quality In vitro digestible dry matter percentages for the 1978-79 forage samples are presented in Table 3. No effort has been made at this time to evaluate the results. LITERATURE CITED Bartmann, R. M. 1979. Deer investigations--food habits technique development and forage preferences of mule deer in the Piceance Basin. P. 91-98. In Game Research Report. Colo. Div. of Wildl., Denver 3(Part 1):1-230. (Proc.). �-129- Pearson, H. A. 1970. Digestibility trials: in vitro techniques. 85-92. In Range and wildlife habitat evaluation--a research symposium. USDA For. Servo Misc. Publ. No. 1147. 220p. Tilley, J. M. A., and R. A. Terry. 1963. A two stage technique for the in vitro digestion of forage crops. J. Br. Grassl. Soc. 18:104-111. /' Prepared by: ··7··· /11' /{ -J"'~// 'c: /, ~ . R. M. Bartmann Wildlife Researcher C ....••• "I' U •• -,. I ~ I I.'" _--- P. �Table 3. In vitro digestible dry matter content, of major forage species eaten by 8 tame mule deer on 2 study areas on pinyon-juniper winter range in the Piceance basin, September-April, 1978-79. Month and Year 9-78 Pinyon-JuniEer Mixed Browse lVDDM (%) % of Diet Taxa by Weight x s.e. Amelanchier utahensis Purshia tridentata CercocarEus montanus Quercus gambellii SymEhoricarEos oreoEhilus Eriogonum umbellatum 34 21 20 17 1 1 40 46 36 27 64 51 6.3 1.0 0.7 2.2Pj 2.1 1.0 Pinyon-JuniEer Sagebrush IVDDM (%) % of Diet Taxa by Weight x s.e. Purshia tridentata 68 Amelanchier utahensis 10 Eriogonum lonchoEhyllum 4 Eriogonum umbellatum 3 CercocarEus mont anus 2 SymEhoricaq~os 2 oreophilus Commandra timbellata 2 LUEinus caudatus 2 Penstemon caespitosus 2 38 55 18 30 44 39 3.1 7.1 2.9 2.9 2. 1 7.3 46 59 33 7.r#/ 1.2 1.9 Cercocarpus montanus Purshia tridentata Juniperus osteosEerma Amelanchier utahensis Artemisia tridentata 47 26 11 3 3 43 42 69 39 66 2.4 1.4 0.4 -~/ 2.9E_/ JuniEerus osteosEerma 18 Chrysothamnus nauseosus 15 Sarcobatus vermiculatus 11 Pinus edulis 8 8 StiEa comata 7 Oryzopsis hymenoides HymenaEaEEus filifolius 6 Artemisia tridentata 5 Eriogonum lonchoEhyllum 3 3 AgroEyron species 2 AgroEyron inerme 1 Purshia tridentata 1 LeEtodactylon Eungens 52 35 57 43 30 68 58 73 18 66 44 46 39 0.4 I 11-78 2-79 CercocarEus montanus 29 Amelanchier utahensis 28 Quercus gambellii 20 Purshia tridentata 6 Chrysothamnus viscidiflorus4 Artemisia tridentata 2 Symphoricarpos oreophilus 1 51 30 46 38 59 81 20 Pinus edulis 23 Chrysothamnus nauseosus 8 Eriogeonum lonchoEhyllum 7 Artemisia frigida 3 Artemisia tridentata 2 Artemisia ludoviciana 2 AtriElex confertifolia 2 SympharicarEos oreoEhilus 2 OryzoEsis hymenoides 1 41 56 12 30 79 33 55 36 54 2.3 6.4 2.9 2.3E_/ 0.5 O. zE_/ 7.6E_/ 2.4 2.7 1.2 10.9b/ 0.2E.../ 8.1E_/ - "a/ 0.82.5 2. zE_/ 3.0E_/ 0.9 12.0E_/ 2.6 _a/ o. ]E_/ - ~/ 1.0 4.7 _ a/ 1.1~/ I-' W o I �Table 3. In vitro digestible dry matter content, of major forage species eaten by 8 tame mule deer on 2 study areas on pinyon-juniper winter range in the Piceance basin, September-April, 1978-79. (Cont'd) Pinyon-Juni2er Mixed Browse IVDDM (%) % of Diet Taxa by Weight x s.e. Month and Year 3-79 4-79 Artemisia ludoviciana 16 Amelanchier utahensis 15 Pinus edulis 13 Eriogonum loncho2hyllum 8 6 Juni2erus osteos2erma Artemisia tridentata 5 Tetradymia canescens 4 Sym2horicar20s oreo2hilus 3 Chrysothamnus viscidiflorus3 Artemisia frigid~ 3 Purshia tridentata 2 Chrysothamnus nause.osus 2 2 HymenoEa22us filifolius 2 Poa species Haplopappus nuttallii 1 30 24 53 29 55 79 36 37 36 44 33 57 43 52 50 24 15 12 9 5 3 3 2 53 59 53 42 48 59 40 57 Poa species Cercocar2us montanus Amelanchier utahensis Carex species CrY2tantha sericea Artemisia tridentata Phlox hoodii Mertensia lanceolata a/ - Only 1 sample. ~/OnlY 2 samples. 5.6 2.8El 5.3 1.8 1.4 3.6 1.5£'/ 1.1 3.3 0.6£./ 1.0£./ - ~/ 3.0 5.0£'/ 0.8£./ 2.3 0.4 1.3 3.7 4.2 0.1 2.5 1.2 Pinyon-Juni2er-Sagebrush IVDDM (%) % of Diet Taxa by Weight s.e. x 2.6 0.7 Hymeno2appus filifolius 14 Purshia tridentata 12 Juniperus osteos2erma 9 Ha2lo2appus nuttallii 9 Chrysothamnus nauseosus 5 Eriogonum loncho2hyllum 5 4 Tetradymia canescens Gutierrezia sarothrae 4 Artemisia tridentata 4 Cercocarpus mont anus 4 Penstemon osterhoutii 4 Stipa comata 3 Carex species 3 2 Koeleria cristata 37 46 67 60 40 36 46 65 60 49 57 31 55 62 3.9£'/ 1.6 1.7£./ 2.9£'/ 1.3 2.0 0.9 3.4 5.5 1.0 0.3 Poa species Purshia tridentata Koeleria cristata Artemisia tridentata Zygadenus venosus Agropyron species Amelanchier utahensis 81 54 67 75 75 70 71 2.3 1.9 5.8 1.3 2.1 3.5 1.1 55 7 5 4 3 3 2 o. JE_/ I •..... LV •.... I ��July, -133- JOB PROGRESS State of: Colorado Project ~']-126-R-3 No.: Job No.: ~5~N=E~ REPORT -Big Game Investigations Deer Investigations 2 Work Plan: 1980 _ Experimental Deer Inventory NE Region Period Covered: Personnel: June 30, 1979 - July 1, 1980 Peggy Abbot, Allen Anderson, David Bowden, Debbie Dorsch, Randy Fischer, Howard Geduldig, Nike Goodson, Cathy Sanz, Cathy Simon, Roger Sleeper, Carol Smith. ABSTRACT Based on counts of pellet groups on 40, permanent, randomly located, 107.64 ft2 (10 m2) circular plots within 23 stratified proportionally allocated and randomly selected square miles (2.59 km2) from 309 mile2 (800 km2) of G.M.D. 20 winter range; mean densities during the 1979-80 "winter period" with 95 percent confidence limits were 32.75 (15.2650.24) deer and 5.45 (2.97-7.93) elk per mile2 (2.59 km2). These estimates are similar but more variable than in 1978-79. Buck:d6e ratios (± SE) and fawn:doe ratios (± SE) based on 1,054 mule deer classified on 20, all-day, walking routes were .2535 + .0381 and .5764 + .0442, respectively. Neither ratio differed significantly (P < O~10) from those obtained for the similar (November-December) sampling period of 1978. The 1977-79 trend to significantly (P < 0.05) smaller mean group sizes in classified mule deer is discussed in relation to the literature. ��-135- EXPERIMENTAL DEER INVENTORY Northeast Region Allen E. Anderson P. N. OBJECTIVE To design appropriate sampling and reliably estimate deer numbers and based on a preliminary sampling of of the four administrative regions analytical procedures necessary to buck:doe:fawn ratios on winter range a problem management unit within each of the state. SEGMENT OBJECTIVES 1. Clear and read pellet group count plots in Game Management Unit 20. 2. To sample buck:doe:fawn ratios on 20 randomly located, all-day walking routes on "winter range" of Game Management Unit 20 • .METHODS AND MATERIALS Counts of Pellet Groups 2 Counts of deer and elk pellet groups on 13,180 temporary, 10-m , circular plots during 1976 and 1977, analyses of resultant data, and some empirical decisions formed the bases for the sampling plan adopted for installation of permanent pellet group plots (Anderson 1977, Anderson and Bowden 1978). The sampling plan is stratified, random, multistage and has been described in detail by Anderson and Bowden (1979). Definitions and procedures used in counting pellet groups are in Appendix A. Deer and elk pellet groups were counted and removed from 1,040-1,200 permanent sample plots by two, two-person crews; September 11 to November 8, 1978, April 24 to May 30, 1979, October 5, 1979 to November 19, 1979, and May 16, 1980 to June 19, 1980 thus sampling pellet group deposition rates during the "winter" of 1978-79, the "sunnner" of 1979, and the "winter" of 1979-80. The chronology of the 1979-80 pellet group counts are detailed in Table 1. This report addresses the estimates of average deer and elk numbers on winter range calculated from these rates during the "sunnner" of 1979 and the "winter" of 1979-80 and compares these to a similar estimate for the "winter" of 1978-79. Field procedures have been reported (Anderson and Bowden 1979). Data were recorded on prepared forms (Appendix B). Mule Deer Herd Structure Field and analytical procedures used to estimate buck:doe and fawn:doe ratios on 20 all-day walking routes with 2 observers have been described (Anderson 1977, Anderson and Bowden 1978) as well as the procedures to randomly select the locale of each route (Anderson 1977:233). To emphasize one important procedure; two methods were used in deer classification and entered separately on prepared forms (Appendix C). Method 1 included both �-136- unclassified single deer and groups in which one or more individuals could not be classified as an antlered buck, doe, or a fawn of the current year and unspecified sex. Method 2 included only classified single deer and those groups in which all deer could be so classified. We used a modification (Appendix D) of the criteria proposed by Dasmann and Taber (1956) to help identify sex and age classes. Just prior to actual sampling, observers spent one day making and comparing simultaneous independent classifications of the same groups of mule deer. Approximately 30 groups of mule deer were cla.ssified in this training exercise. Unfortunately, it has not been possible to have the same observers each year. From 1976 to 1978 I participated in all routes with different observers during each of those years. During 1979, however, I was able to participate in only five routes. Table 1. Chronology of counts of deer and elk pellet groups on G.M.U. 20 winter range, April 24, 1978 to June 19, 1980. Time Interval 2/ Dates-- (Days) of Pellet Group Deposition Period ~/ Mean 4-24-79 to 11-19-79 "summer" 80 166.72 10-5-79 to 6-19-80 "winter" 92 215.27 1/ 2/ Number of days between plots each. counts of individual 1...7 Min Max 14.55 147 190 17.68 184 244 SD transects (N) of 10 sample The dates pellet groups were counted and removed on the first andlast transect. RESULTS Counts of Deer and Elk Pellet Groups The large variance associated with the mean numbers of deer calculated for G.M.U. 20 winter range during the winter of 1978-79 was due mainly to the very large variances associated with the counts of deer pellet groups within strata 5, 6, and 9 (Anderson and Bowden 1979:106). In an effort to reduce those variances, two additional sample units were randomly selected for stratum 5, one for 6, and one for 9. Plots were established on those sample units for strata 5 and 6 during the October 4 to November 19, 1979 counting period but because of unusually early snowfall only 13 of the 40 plots established on the stratum 9 sampling unit could be cleared of pellet groups. In addition, the property owner of land containing sample unit 4-26 ordered the removal of all plot stakes on June 2, 1980. That sample unit was replaced with the random selection of sample unit 4-22. Plots were �2 2 (800 km ) Table 2. Estimated densities of deer during the "winters" of 1979-80 and 1978-79 on 309 mile of G.M.U. 20 winter range based on counts of deer pellet groups on 40, permanent, randomly selected, 107.64 ft2 (10 m2) circular plots within each of 23 stratified, proportionally allocated and randomly selected square miles (2.59 km2). . 1979-80 2/ Strata ~/ n- 1 2 3 4 5 6 7 8 9 10 11 26 25 34 27 29 30 30 28 32 22 26 309 2 2 2 1 3 3 2 2 3 1 2 23 Deer Per Sguare Mile ~?59 Mean SE 95% Conf. Interval l/ Mean Pe LLe.t Groups Per 10 Plots Per Day .004445 .014190 .003285 .002290 .014767 .023337 .025755 .021960 .026093 .030390 .016010 1978-79 s200-5) .990215 7.663220 .244205 0 38.02190 13.862.10 81.56760 60.97030 7.18476 0 2.72322 2/ n- Mean Pellet Groups Per 10 Plots Per Day 2 2 3 2 2 3 3 2 3 2 2 26 s200-5) 6.090106 4.923084 3.317595 0.280987 73.528455 27.260672 15.511025 11.572833 26.812108 13.883112 0.447872 .0077305 .0119177 .0108380 .0029583 .0191740 .0228510 .0127344 .0226472 .0266496 .0195380 .0143766 2 km ) 32.75~_I 5.83 15.26-50.24 31.2&~/ 5.00 17.37-45.15 Reduced from 310 square miles in 1978-79 because the owner of one sample unit within to remove plot stakes on June 2, 1980 thus reducing the sampling universe by one. stratum 4 told us 2/ The number of sample units differ during 1978-79 and 1979-80 because early and continuing snow during Oct. and Nov. 1979 precluded counting pellet groups on 6 sample units and 3 new sample units were established 1/ during that same period. Based on a mean defecation Neff (1968). rate of 13.0 groups per deer per day as generalized from information in I •....• w ....• I �Table 3. Estimated densities of elk during the "winters" of 1978-80 and 1978-79 on 309 mile2 (800 km2) of G.M.U. 20 winter range based on counts of elk pellet groups on 40 permanent, randomly selected, 107.64 ft2 (10 m2) circular plots within 23 or 26 stratified, proportionally allocated and randomly selected square miles ,(2.59 km2) or sample units. ,_,_ 1978-79 1979-80 Strata 1 2 3 4 5 6 7 8 9 10 11 1/ N n- 26 25 34 27 29 30 30 28 32 22 26 309 2 2 2 1 3 3 2 2 3 1 2 23 Mean Pellet Groups Per 10 Plots Per Day s2 (10-5) .004445 .002830 .000550 0 .248645 .064980 .060500 0 005005 0 .002705 .005320 .001650 .009670 0 .892003 0 1.46341 3.51122 .81675 0 0 1/ n- 2 2 3 2 2 3 3 2 3 2 2 26 Mean Pellet Groups Per 10 Plots Per Day .0050955 .0020905 .0028236 .0027105 .0006155 .0013733 .0018583 .0055685 0 .0006250 0 s2 (10-5) 5.19282 0.15824 0.18597 0.00530 0.07576 0.56581 0.29329 4.05090 0 .07812 0 Elk Per Sguare Mile (2.59 km2) Mean SE 95% Conf. Interval 5.4~_/ 1.06 2.97-7.93 4.121:./ 1.20 0.30-7.94 1/ See Tables 2, 4 for explanation of the differing sample sizes. 2/ Based on a mean defecation rate of 12.0 groups per elk per day as generalized from information in 'Neff (1968). •...I w 00 I �2 2 Table 4. Estimated densities of deer and elk during the "summer" of 1979 on 309 mile (800 km ) of G.M.U. 20 winter ranges based on counts of deer and elk pellet groups on 40 permanent, randomly selected,107.64 ft2 (10 m2) circular plots within 20 stratified, proportionally allocated and randomly selected square miles (2.59 km2) or sample units. Elk Deer Strata 1 2 3 4 5 6 7 8 9 10 11 1/ N rr- 26 25 34 27 29 30 30 28 32 22 26 309 2 2 2 1 1 2 2 2 3 1 2 20 Mean Pellet Groups Per 10 Plots Per Day .001575 .005640 .004050 .005810 .018460 .017945 .017305 .025800 .014426 .005490 .004045 2 -5 s (10) o o .003125 .089780 . .089780 1/ n- 2 o 2 2 2 3 1 .001050 2 s2(10-5) .105125 .571220 .784080 .000725 .001690 .009840 2 2 1 1 8.307610 8.281850 83.558700 .263243 Mean Pellet Groups Per 10 Plots Per__~ o o o o o o o o o o o o o o o o 20 2 Animals Per Square Mile (2.59 km ) 2/ 22.4rMean 4.10 SE -2.69-47.56 95% Conf. Interval 1/ 2.6s11 .53 0.12-4.97 Pellet groups could not be counted on 6 sample units because of snow during October-November 1979. 2/ Based on a mean defecation rate of 13.0 groups per deer per day as generalized from information in Neff (1968). 3/ Based on a mean defecation rate of 12.0 groups per elk per day as generalized from information in Neff (1968). I I-' W \0 I �-140- 2 Table 5. Number of square mile (2.59 km ) sample units required to be within specified percentages of the true density with confidence (I-a) for deer and elk on G.M.U. 20 winter range, 1979-80. (l-a) Percent .95 .90 .80 5 10 15 20 25 14244 3561 1599 890 573 5343 1336 600 334 215 1859 465 209 116 75 5 10 15 20 25 3326 832 370 208 133 1879 470 209 117 75 936 234 104 58 37 5 10 15 20 25 42432 10608 4715 2652 1697 18797 4699 2089 1175 752 7576 1894 842 473 303 5 10 15 20 25 1161 290 129 73 46 638 159 71 40 26 310 77 34 19 12 Species and Extreme Dates of Pellet Group Deposition Deer - "summer" April 24 - Nov. 19, 1979 Deer - "winter" Oct. 5, 1979 - June 19, 1980 Elk - "summer" April 24 - Nov. 19, 1979 Elk - "winter" October 4, 1979 - June 19, 1980 �-141- established and cleared thereon during June 18 and 19, 1980. Early snowfall also prevented counts of pellet groups on 6 of 26 total sample units during the October 4 to November 19 counting period. Hence, estimates of deer and elk numbers were based on 20 sample units during the "summer" of 1979 and 23 sample units during the "winter" of 1979-80. Obviously, comparisons of these estimates with those derived in the past or future must be regarded with caution. The basic statistics and pellet group derived estimates of deer and elk densities are given for deer (Table 2) and elk (Table 3) during the "winter" of 1979-80 and 1978-79 and for both deer and elk during the "summer" of 1979 (Table 4). Estimates of sample size requirements based on these samples is given in Table 5. These data suggest similar densities of deer and elk during both winters but much greater variation during 1979-80. The smaller sample sizes and the record snow depths during the winter of 1979-80 were probably contributing factors to that variation. The relatively large variation associated with mean deer and elk densities during the "summer" period of 1979 was probably influenced by both sample size and relatively low densities of both species on winter range during the "summer" period. The magnitude of yearlong variation was such that none of the estimated sample sizes, except for elk during the "winter" were adequate to be within even 25 percent of the mean density at P < 0.20. The increase in number of sample units from 26 to 30 during 1979-80 (Fig. 1) may reduce this variation. Mule Deer Herd Structure Mule deer classified by two methods as antlered bucks, does, and fawns of each sex by 3 observers during 1979 are combined by stratum and route in Table 6. A total of 1.,054mule deer classified during 1979 by both methods compares favorably with previous sample sizes of 1,067 (1977) and 1,221 (1978). Buck:doe and fawn:doe ratios and their standard errors by two classification methods and 3 observers are given in Table 7. Employing the standard error as a percentage of its ratio as a measure of relative variation, the buck:doe ratio had greater variation in every category (15.0 to 41.0%) than did the fawn:doe ratio (7.7 to 21.1%). The effects of observers and methods of classification on these ratios were assessed by 18 comparisons (Table 8). Five comparisons detected significant (P < 0.10) differences between ratios: (1) Buck:doe ratio between methods 1 and 2 for observer 2 since that observer detected no bucks by method 2. (2) Fawn:doe ratios between methods 1 and 2 for observer 3 which may have been associated with the small number (5) of routes sampled by that observer. Fawn:doe ratio of observers 1 and 2 with observer 1 obtaining higher ratios for method 1 (3), method 2 (4), and methods 1 and 2 combined (5). Observer 1 had both a longer training period and greater experience than observer 2 and these factors may have made the difference. These comparisons differ from similar comparisons for 1976, 1977 and 1978 wherein observers and methods of classification had little or no significant (P < 0.10) effect on either ratio. �-142- N t---; I MILE TOOTH RESERVOIR i ! I, e ~ Figure 1. BERTHOUD Thirty proportionally allocated and randomly selected square mile (2.~9 km2) sample units among 11 strata sampling about 309 mile (800 km2) of G.M.U. 20 winter range whose upper limit is approximated by the 8,500 ft (2,591 m) contour line. During the spring and fall, deer and elk pellet groups are counted and removed from 40, 107.64 ft2 (10 m2), circular, permanent plots randomly located in 4, 10-plot segments within each sample unit. �-143- Table 6. Summary of mule deer recorded on 20 all-day walking routes by stratum and method of classification, Nov. 28 to Dec. 24, 1979 on G.M.U. 20 winter range. The classifications of 3 observers are combined. Method 12./ D F L Route Stratum 1 2 2 2 3 3 B ~I D B Method 2~_/ F Uncl. 0 o o o 0 o o o o o 4 2 2 8 1 4 o o o o 0 o 0 o o 0 o o o o o o o 7 29 9 1 3 4 3 11 3 5 2 7 2 o 0 o o o o 12 38 26 76 28 1 9 5 6 21 3 8 o o o 0 o 0 o o o o 10 8 8 17 14 39 19 o o 2 1 1 4 1 11 9 15 31 11 57 20 3 10 2 7 22 5 12 9 26 91 51 168 57 5 7 4 7 23 6 13 10 7 41 37 85 22 o 1 1 1 3 1 14 10 1 33 15 49 12 o 6 4 8 18 2 15 15 8 31 21 60 29 1 1 o 2 4 2 16 15 21 51 18 90 37 1 3 o 4 8 3 17 12 11 92 59 162 25 2 9 8 2 21 2 18 12 1 22 11 34 10 o 0 o o o o 19 13 5 25 15 45 13 4 12 4 12 32 7 20 14 3 11 7 21 5 0 o o o o 128 509 297 934 295 67 35 55 175 36 o o 2 2 1 o o 1 1 4 o o o 0 o 4 5 2 4 3 9 5 5 o o o 6 6 5 17 7 7 o 8 7 9 L o. 18 2:..1 Method 1 includes only those groups in which all deer were classified while Method 2 includes groups in which 1 or more deer were unclassified. bl Number of groups of one or more deer. �-144- Table 7. Sunnnary of mule deer buck:doe and fawn:doe ratios obtained 20 all-day walking routes on G.M.U. 20 winter range, November 28 to December 24, 1979. Observer(s) II Method- on Buck:Doe SE Fawn:Doe SE 1 1 .2805 .0568 .6199 .0578 1 2 .3056 .1253 .6667 .1108 1 1 and 2 .2840 .0588 .6265 .0546 2 1 .1654 .0589 .5940 .0496 2 2 0 0 .4444 .0939 2 1 and 2 .1549 .0549 .5845 .0513 3 1 .2839 .0463 .5226 .0634 3 2 .3043 .0640 .3043 .0816 3 1 and 2 .2865 .0445 .4944 .0635 I, 2 and 3 1 .2515 .0398 .5835 .0434 I, 2 and 3 2 .2687 .0638 .5224 .0922 I, 2 and 3 1 and 2 .2535 .0381 .5764 .0442 ~I See Table 6 for explanation. �-145- Table 8. Comparisons of mule deer buck:doe and fawn:doe ratios sampled within G.M.U. 20 winter range, November 28, 1979 to December 24, 1979. OBS. 1 2 3 Ratio 1 and 2 Method 2 Diff. 1 2 1 and 2 1/ df 90% c.r)_! .3056 -.0250 .112131 19 (-.2189,.1689) F:D .6199 .6667 -.0468 .115570 19 (-.2466,.1530) B:D .1654 0 .1654 .058900 14 (.0617,.2691) F:D .5940 .4444 .1495 .064752 14 (-.0205,.0660) B:D .2839 .3043 -.0205 .065974 4 (-.1611,.1201) .3043 .2182 .076678 4 .5226 ---- - ----- ------- OBS 1 vs. OBS. 2 (.0547,.3817) --- - - - - - B:D .2109 .1654 .0455 .049121 14 (-.0410,.1320) F:D .6939 .5940 .0999 .051794 14 (.0087 ,.1911) B:D .1"538 .0000 .1538 .042200 14 (.0795, .2281) F:D .8077 .4444 .3632 .124130 14 (.1446,.5818) B:D .2023 .1549 .0474 .046312 14 (-.0342,.1290) F:D .7110 .5845 ------------OBS 1 vs. OBS. 3 .1265 .047309 14 (.0432,.2098) ------- B:D .4189 .2839 .135 .081384 4 (-.0385,.3085) F:D .4730 .5226 -.0496 .097831 4 (-.2582,.1590) B:D .7000 .3043 .3957 .372346 4 (-.3981,1.1895) F:D .3000 .3043 -.0043 .096691 4 (-.2104, .2018) B:D .4524 .2865 .1659 .092216 4 (-.0307, .3625) F:D .4524 .4944 -.0420 .094034 4 (-.2425,.1585) ----Method SE .2805 Method 2 vs. B:D F:D 1 Method 1 ------- Ratios differ significantly (P = 0.10) if interval does not encompass zero. �-146- However, as mentioned above, -1979 was the first year in which I did not participate in all routes. None of these factors appeared to affect ratio variation since the 1979 fawn:doe ratio again approached 15 percent of the true ratio at the 95 percent confidence lpvel (Table 9). As in previous years, the buck:doe ratio remains an intractable sampling problem. The 5 significantly (P < 0.10) differing buck:doe and fawn:doe ratios of 1979 suggest caution in comparing these ratios with their counterparts of 1977 and 1978 (Table 10). Moreover, the significantly (P < 0.10) higher fawn: doe ratio of 1977 cannot be explained with available information. Table 9. Numbers of all-day walking routes to be within a specified percentage of the true ratio and confidence level as calculated from the Nov. 28 - Dec. 24, 1979 sample. Confidence Level (I-a) Percent .95 .90 .80 Buck:Doe 5 10 15 20 792 198 88 50 540 135 60 34 319 80 35 20 Fawn:Doe 5 10 15 20 206 52 23 13 141 35 16 9 83 21 9 5 Ratio Table 10. Annual comparisons of mule deer buck:doe and fawn:doe ratios sampled during the breeding season on G.M.U. 20 winter range with 20 all-day walking routes. Sample sizes with methods and observers combined totaled 1,067 deer in 1977, 1,221 in 1978, and 1,054 in 1979. 90% Confidence Ratio Buck:Doe Fawn: Doe 1/ Comparison d. f. IntervaL!.! Difference SE of Diff. 1977 vs. 1978 .1716 .2241 -.0525 .03250 19 -.1087, .0037 1977 vs. 1979 .1716 .2535 -.0818 .12233 19 -.2933, .1297 1978 vs. 1979 .2241 .2535 -.0294 .04123 19 -.1007, .0419 1977 vs. 1978 .6970 .4860 .2110 .05770 19 .1112, .3108 1977 vs. 1979 .6970 .5764 .1206 .05256 19 .0297. .2115 1978 vs. 1979 .4860 .5764 -.0904 .16590 19 -.3773, .1965 Ratios differ zero. significantly (P = 0.10) if interval does not encompass �-147- Another interesting facet but perhaps not as inexplicable as the annual difference in the fawn:doe ratio, is the general trend (1977-79) in significantly (P < 0.05) smaller mean group sizes of mule deer (Table 11). Moreover, the strata with largest decreases in mean group size are also those with the more open habitats and generally larger sample sizes. The record snowfall and subzero temperatures which characterize much of the 1979 sampling period may have had an effect but the specific mechanisms reducing mean group size remain speculative. DISCUSSION For reasons discussed in the text, the effects of increasing the number of sample units in those strata with the largest variance during 1978-79 could not be evaluated during 1979-80. However, the strata with the largest variances were only partially identical in 1978-79 and 1979-80. These "high variance shifts" among strata may be associated with both the relatively severe winter and the smaller sample size during 1979-80. If, however, such shifts are characteristic of annual deer use of G.M.U. 20 winter range; counts of pellet groups may not yield useful annual estimates of deer densities. Current time and lahor considerations preclude a larger number of sample units than the 30 presently installed. Additional sampling during 1980-81, 1981-82, and 1982-83 will be necessary to adequately test the pellet group count technique on G.M.U. 20 winter range. During the summer of 1978, I recommended that the NE Regional management staff incorporate the present herd structure sampling plan into their data gathering process. No specific action on their part has been proposed and I recommend that this portion of Work Plan 2, Job 5 be dropped as an act ive research project. Group sIae as a parameter of potential usefulness in studies of the population dynamics of mule deer has received scant attention. Rodgers (1977) reported on group size parameters in 5, large, gregarious African herbivores and presented voluminous data to support his thesis that group size fluctuated seasonally and was correlated with social and reproductive behavior. This author also found that in 2 of the species, group density (Y) and group size (X) were negatively and significantly (P < 0.10) correlated (r = - 0.845, - 0.872). However unlike this study, successive annual means did not vary significantly for any of the 5 species studied over a 4 year period. It would seem, therefore, that testing of discrete hypotheses relative to group size in mule deer is an appropriate reserach topic. �Table 11. Mean group sizes of mule deer by strata obtained on 20 all-day walking routes on G.M.U. 20 winter range during the late Nov.-Dec. breeding season 1977, 1978 and 1979.!/ 1977 Stratum N X 1978 95% C.1. Y Sig •. Level N X 95% 1979 c.r , S'19.2/ Level 6 7 N - X 95% C.l. 20 2.0 0.8 - 3.2 23 4.0 2.8 - 5.2 .05 42 2.6 1.5 - 3.8 8 21 3.0 1.5 - 4.5 N.S. 31 3.2 2.4 - 3.9 .001 42 1.1 0.6 - 1.5 9 43 5.4 4.1 - 6.6 N.S. 56 4.2 3.3 - 5.0 .001 154 1.8 1.5 - 2.2 10 29 6.1 4.8 - 7.4 .01 45 4·2 3.1 - 5.2 .001 66 2.4 1.7 - 3.1 ~ 00 I 15 41 4.0 3.2 - 4.8 .01 67 2.6 2.2 - 3.1 .001 132 1.3 1.0 - 1.6 12 69 4.3 3.5 - 5.1 N.S. 69 4.3 3.4 - 5.2 N.S. 70 3.4 2.3 - 4.5 13 23 3.9 3.0 - 4.8 N.S. 22 4.1 2.5 - 5.7 N. S. 26 3.0 1.8 - 4.2 1/ Only those stratum with 20 or more sightings of one or more deer are presented. l/ I •..... Mean group sizes within strata tested for significant (P < 0.05) differences between years; 1977 vs. 1978 and 1978 vs. 1979 with the t test for unequal variances (Li 1964:143). �-149- LITERATURE CITED Anderson, A. E. 1977. Experimental deer inventory, northeast region. Job Progr. Rep., Pp. 227-250 In Colo. Div. Wildl. Game Res. Rep., July Part 2. 135-305p. (Processed). Anderson, A. E'f and D. C. Bowden. 1978. Experimental deer inventorynortheast region. Job Progr. Rep ,, Pp , 225-244 In Colo. Div. Wildl., Game Res. Rep., July Part 2. 137-298p. (Processed). Anderson, A. E., and D. C. Bowden. 1979. Experimental deer inventory, northeast region. Job Progr. Rep., Pp. 101-192 In Colo. Div. Wildl., Game Res. Rep., July Part 2. 1-230p. (Processed). Dasmann, R. F., and R. D. Taber. 1956. Determining structure in Columbian black-tailed deer populations. J. Wildl. Manage. 29(1):78-80. Li, J. C. R. 1964. Statistical inference. Ann Arbor, Michigan. 658p. I. Edwards Brothers, Inc., Neff, D. J. 1968. The pellet group count technique for big game trend, census, and distribution: a review. J. Wildl. Manage. 32(3):597614. Rodgers, W. A. 1977. Seasonal change in group size amongst five wild herbivore species. E. Afr. Wildl. J. 15(3):175-190. Prepared by: at&Je~~ Allen E. Anderson Wildlife Researcher C �-150- APPENDIX COUNTING A DEER PELLET GROUPS Definitions and Procedures Deer Pellet Group - Five or more deer pellets of the same general size, shape, hardness and color judged to have been continuously voided at the place where observed. Pellets redistributed by water of other agents to within the plot are not recorded. Pellets strewn across the plot are counted as a group if about one-half of the total linear distance or its midpoint falls within the plot as measured with a steel pocket tape. Groups occurring on the plot periphery are counted if about one-half of their total area falls within the plot. "New" Deer Pellet Group - Pellets are typically deposited during the previous year. shiny, often soft and "Old" Deer Pellet Group - Pellets are typically not shiny, generally hard and sometimes wholly or partially concealed by litter. Search Procedures - The area of search is defined by the light metal chain, 1.784 meters in length revolving about the metal rod driven firmly into the ground. The plot size is 0.001 hectare or 10 square meters or 107.64 square feet. On steep slopes the chain is held horizontal. When necessary, the plot boundary is defined by dropping a pebble from the 1.784 meter mark on the chain. Each plot is searched twice (clockwise and counterclockwise) with the two observers changing positions with the change in search direction. �-151- APPENDIX B FORM FOR RECORDING PELLET GROUP COUNTS G.M.U. 20 PELLET GROUP COUNTS W-126-R WP2 - J5 STRATUM~ DATE _ SAMPLE UNIT~ TRAN. Plot '------ DE'ER New Old TIME FINISHED ELK New _ Old _ ---- TRAN. Plot 1 11 2 12 3 13 4 14 5 15 6 16 7 17 8 18 9 19 10 20 Total Total Comments, Weather, etc. TRANSECT(S) OBSERVERS --------------- TIME BEGAN _ DE R New Old MIN. HR TOTAL ELK New Old --------------------------- _ �-152- APPENDIX C FORM FOR RECORDING DEER HERD STRUCTURE Date: Time: Area: Legal Descr.: -------- to Observer --------------------- 1/4, S , T Strata B D F Unc. N, R W. Random No. I: TIME B D F I: Unc. I TIME �-153- APPENDIX D SOME CRITERIA FOR DISTINGUISHING ATTRIBUTE ANTLERED MALES SEX AND AGE OF MULE DEE~/ DOES BODY SIZE FAWNS SMALLER THAN MOST ADULTS ESPECIALLY IN LENGTH OF HEAD AND SIZE OF WHITE RUMP PATCH HEAD SHAPE DEEP MUZZLE OFTEN APPEARS VERY THICK LESS DEEP MUZZLE. DORSAL SURFACE OF MUZZLE MAY APPEAR CURVED WITH VISIBLE VEINS SHORT MUZZLED. HIGH CROWNED. LONG EARS IN PROPORTION TO HEAD SIZE NECK THICK OFTEN LONG, THIN RELATIVELY SHORT AND THICK PELAGE DARK FOREHEAD CONTRASTING WITH LIGHT MUZZLE OFTEN LESS COLOR CONTRAST THAN MALE ON HEADMUZZLE AREA EARS AND NECK MAY APPEAR "FUZZY" OR "WOOLY" BEHAVIOR OFTEN SOLITARY OR SOMEWHAT APART FROM GROUP USUALLY TAKES LEAD IN GROUP TRAVEL, SOMETIMES AGGRESSIVE TO FAWN USUALLY CLOSE TO DOE. EARS OFTEN CLOSE TOGETHER AND NECK ARCHED WHEN ALARMED AND RUNNING OR TROTTING. 1/ Modified from Dasmann and Taber (1956). ��July, -155- JOB PROGRESS REPORT State of Colorado Project No. W-126-R-3 Big Game Investigations Work Plan No. 2 -------- Deer Investigations Job No. 5 Period Covered: July 1, 1979 through June 30, 1980 Personnel: 1980 Experimental Deer Inventory - NW Region D. Freddy, K. Berner, D. Bowden, and T. Baumann ABSTRACT In pinyon-juniper woodland, temporary pellet transects estimated mule deer pellet group densities as effectively as permanent transects in 2 of 3 years. Significant differences in numbers of new pellet groups per square-mile based on comparing totals of temporary and permanent transects occurred in only· I year (P ~ 0.10). Permanent transects were more time consuming to complete than temporary transects. During pre-treatment measurements, permanent transects took 140.4 + 4.21 (SE) minutes to install and enumerate pellet groups whereas temporary transects averaged only 67.7 ± 1.82 (SE) minutes to complete. After initial establishment, permanent transects took 82.4 minutes and temporary transects 68.7 minutes to complete. On a yearly basis, less than 1 percent of the permanent plot stake markers were missing. ��-157- EXPERIMENTAL DEER INVENTORY-PICEANCE BASIN-NORTHWEST REGION David J. Freddy P. N. OBJECTIVE To design appropriate sampling and analytical procedures necessary to reliably estimate deer numbers and buck:doe:fawn ratios on winter range based on a preliminary sampling of a problem management unit within each of the four administrative regions of the state. SEGMENT OBJECTIVE Compare the efficiency of permanent vs. temporary plots to estimate pellet group densities on 10 mi2 of winter range in Game Management Unit 22. INTRODUCTION Annual estimates of deer numbers via deer pellet group densities depends upon accurate discriminations of old pellet groups from new groups and an accurate estimate of the number of days deer occupy the sample area. Permanently staked pellet plots are usually established and cleared of pellet groups at least once annually to aid in discerning new from old groups and delimiting the number of days over which groups were deposited. Mule deer wintering in the Piceance Basin in northwestern Colorado are migratory and essentially vacate their winter range during summer. Arrival and departure times of deer on the winter range could be empirically estimated by systematic aerial or ground searches for deer during fall and spring. This would eliminate one reason for establishing permanent pellet plots. Subjective criteria of surface sheen, color and texture have been used to differentiate new pellet groups, those believed to be I year old or less, from old pellet groups (Freddy 1977). However, the validity of these criteria has not been tested adequately. If such criteria were valid, then temporary pellet plots could be used instead of permanent plots. While designing a permanent sampling system for estimating pellet group densities in the Piceance Basin (Freddy 1978), a question arose concerning the necessity to establish permanent pellet plots and transects. A permanent plot test was designed to ascertain the quantitative advantages or disadvantages of permanent and temporary plots. METHODS AND MATERIALS Ten sample units each I square-mile (2.59 km2) in size and having ~ high deer pellet group density (Freddy 1977) were selected for the permanent �-158- plot test (Figure 1). Vegetation of these units was dominated by pinyonjuniper-mixed shrub communities. Primary shrub species were sagebrush (Artemisia tridentata), rabbitbrush (Chrysothamnus sp.), serviceberry (Amelanchier utahensis), and mountain mahogany (Cercocarpus montanus). Terrain varied from moderate rolling hills to steep-faced gullies and cliffs. A grid of 10 transects at 0.1 mile (0.16 km) intervals each with 10 plots at 0.1 mile (0.16 km) intervals was delineated on topographic maps for each sample unit (Freddy 1977). Intersections of grid lines denoted proper locations of pellet plots and were used as an aid in placing pellet plots in their correct locations. Five pairs of transects were denoted for each sample unit with one permanently marked transect and one temporary transect in each pair. Permanent transects contained 10 permanently staked pellet plots while temporary transects also had 10 plots but there were no markers to locate plots. Permanent and temporary transects alternated in location within the grid with the first transect in each sample unit randomly selected temporary transect. Transects were oriented on true east bearings and intercepted maximum topographic relief within each sample unit. With this sampling design, estimates of new pellet groups per sample unit based on temporary and permanent transects could be compared using a paired t-test (Sokal and Rohlf 1969). Permanent and temporary plots were located by pacing with the aid of a hand-held compass and topographic maps. Permanent plots were marked with a 2 ft. steel stake painted orange and yellow flagging was tied to trees or bushes to aid in relocating these plots. No markers were installed to relocate temporary transects or plots. Temporary transects and plots thus could vary slightly in location from year to year. During the pretreatment year 1977, numbers of new and old pellet groups were determined for both temporary and permanent plots. In subsequent years only numbers of new pellet groups were determined for each plot. Permanent plots were cleared of pellet groups each year as measurements were made. For the initial establishment of plots in 1977, new pellet groups were subjectively differentiated from old groups on both types of plots. Pellet groups with sheen on some surfaces and noticeable color (usually brown or black) were considered new, whereas groups without sheen and of faded coloration were judged to be old. A pellet group was arbitrarily defined as 5 or more pellets appearing similar in size, shape, texture, and color and in close proximity to one another. A pellet group was considered on the plot if the mid point of an imaginary line drawn through the long axis of the group was within the plot and/or one-half of the geographic area covered by the group was within the plot. Plots were circular and 107.6 ft2 (.001 ha) in size. Plots were examined twice by 2 persons, once in a clockwise and then in a counter-clockwise direction. Aging plots were established on 5 microsights representing different aspects and vegetation types within pinyon-juniper community to objectively differentiate new and old pellet groups found on temporary plots. New pellet on �GAME MANAGEMENT UNIT 22 -PICEANCE Scale o 12 34 5 miles Meeker I t-> .U1 1.0 I Figure 1. Locations of square-mile Piceance Basin, Colorado. sample units (hatched area) sampled for deer pellet group density, �-160- groups were collected from wild deer and tame deer on native forage after recent snowfalls from October into April each year. Groups were placed on each of 3 aging plots within each microsight. These groups were inspected for physical characteristics of sheen, color, and texture in April each year before permanent and temporary plots were examined. Groups remained on plots for 1 year to develop criteria for differentiating known new and old groups (Table 1). Table 1. Criteria pellet groups. used to differentiate Type of Group New Old new pellet groups from old Criteria 1. Color usually dark, intense, and uniform over entire pellet surface. In some instances color may be faded on portions of the pellet surface. 2. Sheen usually present on at least one-half of the pellet surface. 3. Cracking of pellet surface not uncommon, surface texture varied from smooth to rough. 4. Occasionally, fungus or mold visibly present on pellet surface. 1. Color usually faded and often "bleached" gray on at least one surface resulting in a two-toned color effect. 2. Sheen not usually present on pellet surfaces, but may occur on bottom or protected surface of a pellet. 3. Cracking of pellet surface common, texture usually rough. 4. Fungus or mold not present surface. 5. Pellets often embedded surface on pellet in duff or soil. In 1977, pretreatment data were collected from 6 July through 17 August. From 1978-80, measurements were made between 20 April and 31 May. The author was an observer each year while the second observer changed annually. From 4 to 6 transects were completed daily. Measurements were avoided when pellet groups were wet after recent rains or snows. �-161- Table 2. Sampled pellet group densities on temporary and permanent transects on 10, square-mile sample units, Piceance Basin, Colorado, 1977-1980. Squaremile Sample Unit 4-18 4-19 4-30 Transect~/ 1977 Pre-Treatment New Old Groups Groups 1978 New Pellet GrouEs 1979 1980 1978-80 1 3 5 7 .9 I: 64 40 24 42 34 204 5 15 8 5 7 40 18 22 24 18 24 106 15 16 9 9 12 61 11 14 14 13 11 63 230 2 4 6 8 10 I: 60 33 32 58 30 213 9 17 14 12 8 60 33 10 11 24 23 101 10 9 6 17 11 53 29 13 9 12 4 67 221 1 3 5 7 9 L 27 38 26 49 59 199 5 7 5 6 6 29 18 25 13 16 12 84 10 3 8 9 16 46 17 11 11 11 7 57 187 2 4 6 8 10 I: 41 43 27 52 38 201 9 14 5 2 8 38 17 18 18 23 18 94 17 9 14 10 12 62 10 17 13 14 24 78 234 1 3 5 7 9 31 16 32 36 20 135 3 4 4 18 1 30 17 13 24 20 14 88 17 12 26 15 9 79 10 9 14 9 14 56 223 39 24 45 31 31 170 11 4 10 16 14 12 11 63 11 17 16 17 19 80 12 9 23 17 .10 71 214 L 2 4 6 8 10 L 8 10 3 36 ------------------------~~----------------------------------------------------- �-162- Table 2. Sampled pellet group densities on temporary and permanent transects on 10, square-mile sample units, Piceance Basin, Colorado, 1977-1980. (Cont'd). Squaremile Sample Unit 4-31 1977 Pre-Treatment a/ Transect1 3 5 7 9 L: 2 4 6 8 10 L: 4-12 1 3 5 7 9 L: 2 4 6 8 10 L: 4-13 1 3 5 7 9 L: 2 4 6 8 10 L: Old Groups New Groups 1978 43 29 19 19 22 132 16 16 7 3 6 48 27 9 22 2 24 84 11 21 8 5 11 56 15 3 3 6 2 29 169 28 23 24 15 18 108 11 13 4 5 7 40 11 9 8 7 14 49 2 8 12 8 13 43 4 4 3 2 13 26 118 29 32 39 37 37 174 12 7 3 11 8 41 17 9 7 16 7 56 9 5 5 10 11 40 2 5 3 13 11 34 130 23 52 38 30 47 190 5 7 8 7 22 49 13 13 7 4 16 53 6 10 17 16 25 74 6 5 8 17 15 51 178 36 50 17 29 27 159 8 3 3 4 4 22 15 19 12 14 6 66 9 11 11 8 9 . 48 11 7 6 12 1 37 151 29 10 19 20 30 108 5 1 4 4 5 19 14 11 11 6 5 5 38 9 4 4 1 7 25 110 6 6 1 20 47 New Pellet GrouEs 1980 1979 1978-80 ------------------------------------------------------------------------------ �-163- Table 2. Sampled pellet group densities on temporary and permanent transects on 10, square-mile sample units, Piceance Basin, Colorado, 1977-1980. (Cont'd). Squaremile Sample Unit 4-24 T;ansect~/ 1 3 5 7 9 ~ 2 4 6 8 10 z 4-25 1 3 5 7 9 z 2 4 6 8 10 ~ 4-14 1 3 5 7 9 ~ 2 4 6 8 10 z 1977 Pre-Treatment Old New Groups Groups 1978 New Pellet GrouEs 1978-80 1979 1980 33 26 57 34 34 184 7 15 6 9 8 45 10 11 19 9 16 65 13 15 21 11 19 79 12 9 16 8 12 . 57 201 46 39 57 37 33 212 13 15 6 11 4 49 13 14 10 17 12 66 14 12 10 12 12 60 12 18 16 10 15 71 197 21 26 37 30 40 154 6 5 5 7 12 35 9 16 10 4 14 53 8 13 7 13 17 58 5 10 8 6 7 36 147 17 45 25 35 48 170 3 9 1 14 12 39 15 15 7 9 8 54 13 11 9 6 17 56 8 10 8 8 8 42 152 39 25 19 . 30 41 154 9 5 3 4 7 28 19 23 19 8 17 86 13 15 5 9 9 51 3 8 8 11 11 41 178 33 26 24 32 29 144 8 4 5 8 9 34 9 8 7 28 23 75 10 9 7 18 10 54 8 6 9 8 3 34 163 �-164- Table 2. Sampled pellet group densities on temporary and permanent transects on 10, square-mile sample units, Piceance Basin, Colorado, 1977-1980. (Cont'd). Squaremile Sample Unit 4-23 1977 Pre-Treatment a/ Transect1 3 5 7 9 E 2 4 6 8 10 E Old Groups New Groups 1978 New Pellet GrouEs 1979 1980 1978-80 8 17 12 52 10 12 7 5 13 47 5 8 7 4 15 39 13 5 12 8 4 42 128 31 32 27 30 38 158 9 3 6 14 6 38 6 8 7 8 13 42 13 8 8 12 14 55 3 7 11 5 16 42 139 9 34 28 34 47 152 6 9 Totals Transects 1-9 1,647 370 735 557 452 1,744 Transects 2-10 1,674 402 644 575 507 1,726 ~/Odd-numbered transects contained 10 temporary plots, even-numbered transects contained 10 permanently staked plots. �-165- RESULTS AND DISCUSSION Numbers of pellet groups summed over all three years (1978-80) were similar between permanent and temporary transects for all square-mile (2.59 km2) sample units (Table 2). Significant (P ~ 0.10) within year differences between permanent and temporary transects occurred only in 1978 when the mean difference between temporary and permanent plots was 9.1 pellet groups per square mile (2.59 km2) (Table 3). During 1979 and 1980, slightly more new groups were found on permanent transects, but differences were non-significant (P < 0.10) (Table 3). The difference in 1978 could be attributed to: 1) sampling bias due to plot type or location, 2) more pellet groups being deposited on temporary plots, 3) inadequate criteria for distinguishing new and old groups. Because new pellet groups placed on aging plots established during winter 1977-78 had not yet weathered for I complete year when examined in spring of 1978, definitive criteria for distinguishing new and old groups could not be developed. Whereas, in 1979 and 1980, old groups of known ages existed on aging plots allowing better distinguishing criteria to be developed. However, there was no difference (P ~ 0.10) in numbers of new pellet groups by plot type for pretreatment measurements in 1977 (Table 3) when aging plots did not exist and only subjective, but similar, criteria were used to discern ages of pellet groups. Thus, the difference in 1978 probably was due to more groups being deposited at the locations of temporary plots. Number of permanent plots which could not be relocated either because stakes .had been pulled or plots could not be found, was 4, 2, and 1 in 1978, 1979, and 1980, respectively. On a yearly basis, missing plots amounted to less than I percent of the number of permanent plots established. Missing plots were restaked near their original locale since flagging usually remained in the general locations of plots. Table 3. Mean differences and variances of numbers of new pellet groups on temporary and permanent pellet plots per square-mile sample unit 19771980, Piceance Basin, Colorado. A negative (-) indicates more groups on permanent plots. D 1977 1978 1979 1980 - 3.2 9.1 - 1.8 - 5.50 SD 9.5196 13.4578 16.1382 11.068 SD 2.8965 4.2557 5.1033 3.500 df 9.0 9.0 9.0 9.0 t s 1.104#1 2.1383-aI 0.3527E' 1.571~' a b Significant difference at P < 0.10 but not at P No significant difference at P < 0.10 < 0.05 �-166- Amount of time needed to complete temporary and permanent transects varied between years and transect type. In 1977, when plots were initially established, permanent transects took 140.4 + 4.21 (SE) minutes to complete as compared to 67.7 ± 1.82 (SE) minutes for temporary transects (Table 4). Permanent plots had to be staked, numbers of new and old groups determined, plots cleared of all pellet groups, and flagged for future relocation, whereas on temporary plots only numbers of new and old groups had to be determined. During ensuing years, significant differences in times per transect type continued (P < 0.01) (Table 4). In 1978 and 1979, more time was needed to complete permanent than temporary transects (Table 4) as again permanent plots had to be relocated, cleared of pellet groups, and reflagged. However, in 1980 less time was needed to complete permanent transects than temporary transects, reflecting that permanent plots were not cleared or reflagged because the study was terminated. Times to complete permanent transects were not different between 1978 and 1979, averaging 82.4 minutes, but significantly less time was needed in 1980, 64.2 ± 1.38 (SE) minutes (P ~ 0.01). Comparing these times, approximately 18 minutes were needed to reflag and clear 10 permanent plots comprising each permanent transect. There were no significant between year differences in time needed to complete temporary transects (P ~ 0.05; F = 1.0705) (Table 4). Overall average time to complete temporary transects was 68.4 minutes. It is interesting that in 1980 permanent transects took less time to complete than the overall average time for temporary transects. This probably reflects the time needed for decisions regarding minor adjustments in final locations of temporary plots whereas permanent plots only needed to be located. Table 4. Average time (minutes) to complete temporary and permanent pellet plot transects, 1977-1980, Piceance Basin, Colorado. Ten plots per transect. 1977 Temp. 67.7 X 1.82 SE 50 n SE % X 0.027 140.4 4.21 50 0.030 1980 1979 1978 Perm. Temp. Perm. Temp. Perm. Temp. Pe rm; 68.8 85.1 65.8 79.7 71.4 64.2 1.43 50 0.021 3.04 50 0.036 2.26 50 0.034 2.13 50 0.027 1.52 50 0.021 Efficiency in running permanent transects tended to improve from 1978 to 1980 as the standard error expressed as a percentage of the mean decreased each year (Table 4). This likely reflects increasing familiarity with plot locations and decreasing numbers of missing plots each year. Preliminary data analysis indicates temporary transects estimated pellet group densities as effectively as permanent transects in 2 of 3 years. Under the circumstances of a very mobile deer population which essentially vacates winter 1.38 50 0.022 �-167- ranges in summer, temporary plots should be considered a viable alternative to permanent plots. Temporary plots were less expensive when judged by time needed to initially establish permanent transects and complete permanent transects in subsequent years. Temporary plots also offer advantages of flexibility in sampling designs or strategies. Sampling designs could be altered without abandoning a "permanently" established system of plots and transects. Disadvantages of temporary plots include the necessity to establish objective criteria to distinguish new and old groups. In the author's opinion, this mandates establishing a system of aging plots. Also, the ability to discern new and old groups declines when groups become wet from rainstorms, thus retarding completion of temporary transects during inclement weather. Further analyses will be conducted to compare numeric qualities of temporary and permanent plots more completely. Frequencies of zero plots, constancies of plots and transects, frequency distributions of pellet groups per plot and transect, comparisons of variances of transect types, and covariance analysis of treatment and pretreatment data will be examined to better understand how different plot types estimate densities of pellet groups. LITERATURE CITED Freddy, D. J. 1977. Experimental deer inventory-Piceance Div. Wildl. Game Res. Rep. July, Part 2:251-273. Basin. Colo. Freddy~ D. J. 1978. Experimental deer inventory-Piceance Div. Wildl. Game Res. Rep. July, Part 2:245-263. Basin. Colo. Sokal, R. R., and F. J. Rohlf. San Francisco. 776pp. 1969. Biometry. W. H. Freeman and Co., ��July, 1980 -169- JOB FINAL REPORT State of Colorado ---------------------- Big Game Investigations Pro j ec t No. ..:.,:W_-.::1;::.2.;:,.6-_,;R:..;,-_,;3=_ Work Plan No. 2 Deer Investigations Job No. 5 Experimental Southeast Period - Region Covered: Personnel: Deer Inventory July 1, 1979 through Bill Kendall, Paul Morency, June 30, 1980 Thomas M. Pojar ABSTRACT Permanently marked pellet group plots that had been cleared the previous summer were revisited and all new pellet groups recorded. A total of 5,120 plots were counted, 160 plots each on 32 square miles of the 1,026 square-mile study area. The number of pellets in each group of less than 30 pellets were enumerated so the population estimate could be based on various criteria of what constitutes a group. The results of this experiment and a study of the number of pellets in "obviously new" groups are reported in the two manuscripts appended to this report (Appendix A). ��-171- EXPERIMENTAL DEER INVENTORY SOUTHEAST REGION Thomas M. Pojar P. N. OBJECTIVE Design appropriate sampling and analytical procedures necessary to reliably estimate deer numbers and buck:doe:fawn ratios of selected management units in the southeast region of the state. SEGMENT OBJECTIVE 1. Clear and read pellet group count plots in Game Management and 581. METHODS Units 58 AND MATERIALS The sampling design and methods have been described previously in Pojar (1977) and Pojar (1978). Additional information on the determination of the most time-efficient sampling scheme is presented in Pojar (1979). Previously a pellet group was arbitrarily defined as 5 or more pellets of similar size, shape, and color. The area around Canon City characteristically lacks vegetative ground cover and is subject to very heavy summer rainstorms. We theorize that these factors plus the fact that the plots are visited only once a year increases the probability that groups of 5 or more pellets could be washed onto the cleared plot. To get an indication of how serious a problem this might be in affecting the density estimate, we counted the number of pellets in each group that contained 30 or fewer pellets. This allowed an evaluation of the data based on different criteria for the definition of a pellet group. In conjunction with the pellet group search, a specimen of each plant encountered across the entire study area was collected and pressed. A small number of the plants could not be keyed out because of their phenology at the time of collection. The near complete list of plant species that occur on the study area is included in Appendix B. RESULTS AND DISCUSSION The definition of how many pellets constitute a group and the affect on the population size estimate are reported in the first manuscript in Appendix A. The second manuscript in Appendix A reports the results of estimating the number of pellets contained in obvious intact pellet groups. These manuscripts will be revised and submitted for consideration to the Journal of Wildlife Management. A third manuscript reporting on the effectiveness of the sampling methods explored will be prepared during the next segment. The working title is: �-172- Pojar, T. M., D. C. Bowden. Use of the pellet group census technique and sampling strategy on year-round mule deer range. This paper will also be submitted Prepared to the Journal bY~~~~~ __~~~ __ 17~~~~ M. Pojar Wildlife Researcher C _ of Wildlife Management. �-173- Appendix A �Kenneth W. Kendall 500 West Prospect 28-J Fort Collins, Colorado 80526 -174- EFFECT OF PELLET GROUP DEFINITIONS ON THE POPULATION ESTIMATE KENNETH W. KENDALL, Colorado Division of Wildlife, Fort Collins, Colorado 80526 THOMAS M. POJAR, Colorado Division of Wildlife, Fort Collins, Colorado 80526 Abstract: When the pellet group census technique is used to estimate numbers of mule deer (Odocoileus hemionus) the number of pellets that constitutes I pellet group must be defined. Pellet group counts for several criteria classes were collected from 5,120 circular sample plots to evaluate the effect various definitions can have on the subsequent population estimate. Defining a group as 5 or less pellets resulted in an estimate of nearly 50 percent more deer than an estimate based on a definition of 30 or more pellets per group. Number of pellets defined as a group can effect the population estimate and should be carefully considered before making assumptions about population density based on the pellet group census technique. Further research is needed to clearly define the number of pellets in a pellet group. Density of fecal pellet groups has been widely used to estimate numbers of mule deer (Odocoileus hemionus). Many useable definitions and search procedures have been reported (Bowden et al. 1969, Eberhardt and VanEtten 1956, Neff 1968, Robinette et al. 1958, Smith 1968) but the number of individual pellets defined as I group in each of these studies is quite arbitrary. Strong and Freddy (1979) estimated the mean number of pellets per group deposited by wild deer on natural range to be 132 + 12 (SE) for "new" groups and·84 ± 3 (SE) for "old" groups. Criteria used in previous pellet group studies range from defining a group as one isolated pellet (Bennett et al. 1940) to a definition of 30 or more pellets of similar size, shape and color (Neff 1968). Accurate definition of what constitutes a group becomes important when the objective is to estimate actual density of animals rather than trend. The purpose of this study was to estimate the effect of various criteria for defining a pellet group on the subsequent population estimate. Understanding these effects will help in developing a clear definition of a pellet group and hopefully increase the utility of the pellet group census technique. METHODS 2 Fecal pellet group counts were conducted on 2,657 km of mule deer range in south-central Colorado near Canon City (Fig. 1) from 21 11ay through August 1979. A total of 64 1.609 km linear transects and 5,120 .001-ha circular plots were randomly selected for sampling. Each transect contained �-175- 80 plots at 20.l-m intervals permanently marked by a 25-cm steel rebar stake in the center. Pellet group density was estimated on each plot following search procedures described by Neff (1968). Individual pellets were grouped according to size, shape, color and juxtaposition. Groups possessing more than 30 similar pellets were classified as "alpha" groups and arbitrarily assumed to be complete groups. Groups with 30 or fewer pellets were considered incomplete subgroups and placed into 6 "beta" classes (Table 1). Total number of deer on the study area was estimated procedures (Overton 1971). (1) Calculate t: number of deer-days 1 t (2) Calulate N: number N na 'f.y d.r. with the following use per hectare y n a d.r. number of pellet groups number of sample plots area of one sample plot assumed defecation rate of deer on the study area tA P A P total study area (ha) assumed study period (days) Data from each pellet group class was systematically substituted for the variable 'f.yin a summation process beginning with the alpha class and proceeding through all beta classes. RESULTS AND DISCUSSION Table 1 shows the number of pellet groups counted for each classification type. Table 2 shows the resulting population estimates. As the number of pellets defined as a group decreased the number of pellet groups counted and the estimated number of deer in the population increased (Fig. 2). Defining a pellet group as 5 or less pellet resulted in a population estimate that was nearly 50 percent higher than an estimate based on a definition of 30 or more pellets. A higher number of beta-6 groups was expected because the probability of 5 or fewer pellets being translocated into a cleared plot through physical and biological dispersion (Ferguson 1955, Wallmo et ale 1962) is greater than for all other classes. The same is true for pellets that previous observers miss during sampling and fail to remove when plots are cleared. The number of pellets defined as a group can have a significant effect on a population estimate based on the pellet group census technique. When the number of pellets constituting a group is reduced the possibility of bias due to dispersion or observer error increases. This information should be considered before making assumptions about population density based on the pellet group census technique. Further research is needed to verify the mean number of pellets deposited per group and to evaluate rates of pellet dispersal in relation to physical and biological factors. Understanding these factors could help to determine a clear definition of a pellet group and increase the utility and acceptance of the pellet group census technique. �-176- LITERATURE CITED BENNETT, L. J., P. E. ENGLISH, and R. McBAIN. 1940. A study of deer populations by use of pellet group counts. J. Wildl. Manage. 4(4): 398-403. BOWDEN, D. C., A. E. ANDERSON, and D. W. MEDIN. 1969. Frequency distribution of mule deer fecal group counts. J. Wildl. Manage. 33:895-905. EBERHARDT, L., and R. C. VanETTEN. 1956. group count as a deer census method. Evaluation of the pellet J. Wildl. Manage. 20(1):70-74. FERGUSON, R. B. 1955. The pellet-group count method of censusing mule deer in Utah. M.S. Thesis. Utah State Agricultural College, Logan. 94pp. NEFF, D. J. 1968. The pellet-group count technique for big game trend, census, and distribution: A review. J. Wildl. Manage. 32(3):597-624. OVERTON, W. S. 1971. Estimating the numbers of animals in wildlife populations. Pages 430-432 in R. H. Giles, ed. Wildlife management techniques. 3rd Ed. The Wildlife Society, Washington, D.C. ROBINETTE, W. L., R. B. FERGUSON, and J. S. GASHWILER. 1958. Some problems in the use of deer pellet group counts. Trans. 23rd N. Amer. Wildl. Conf. 411-425. SMITH, A. D. 1964. 28:435-444. Defecation rates of mule deer. J. Wildl. Manage. SMITH, R. H. 1968. A comparison of several sizes of circular plots for estimating deer pellet-group density. J. Wildl. Manage. 32(3): 585-592. STRONG, L. L., and D. J. FREDDY. 1979. Number of pellets per mule deer defecation. J. Wildl. Manage. 43(2):563-564. WALLMO, o. C., A. W. JACKSON, T. L. HAILEY, and R. L. CARLISLE. 1962. Influence of rain on the count of deer pellet groups. J. Wildl. Manage. 26:50-55. �-177- 'i'able1. Pellet group counts and classification scheme. Group Classification Class Boundaries No. of Groups Alpha 30 2,212 2,212 Accumulative Total Beta 1 26-29 2 2,214 2 21-25 12 2,226 3 16-20 36 2,262 4 11-15 78 2,340 5 6-10 144 2,484 6 1-5 691 3,175 Table 2. Cumulative effect of various criteria classes on the population estimat~/ . Class Alpha Beta Estimate Increase % 24,192 0 0.0 1 24,257 65 0.3 2 24,389 197 0.8 3 24,783 591 2.0 4 25,636 1,444 6.0 5 27,210 3,018 12.0 6 34,768 101576 44.0 a/Percent increases are in relation to arbitrarily defining a pellet group as more than 30 pellets. �-j'-' MOFF~r .=._._. i i MORGIIN i ~C- •• _ i RIO e,UlAIC.O i-r I !I Klr LIHc.olrJ C4R$O, i fflESA I "''-I'JISD -.IE-YEflnE. j , ,--- I KiOWA I PP.ol'leltS BE.I'IT onltD , I •.... ....• i 00 I r oneil tllS /IN/triM \ I r-'-'-'-'l'-'---'- i .-.-._.~.. SCIlOOI. SERIES O.,u,;M.p ,_ -:~;:~~~~!:~•..." •... COLORADO I....l:- • ••• ,." •. JOO' jO, Pig- 1.. -lit. The study area, located in HiGh Park on the Arkansas river druinace near C~non City, Colorado. �-179- -- . 1~..enClcL.-'- •....•.. ~~ hO '-" .8 r_' ~ ,-. ,.!.. h 8 ." ~< 30 ~ ;2.: 0 H 8 ~ ...... ~ 20 ::::t.. 0 ."..... ...".. ,.....; H 10 ~ U2 <: ;~ ,..._. 1-'-< 0 ,..".. F-i I H 6-10 1-5 11-15 :ZUHBER OF PELLETS Effects of ?ellet DEFnmD 21-25 increases I 26-30 AS A GROUP group definitions po pu'l atLon estimate •. Percentage to defining 16-20 on the are in relation a pellet group as more than 30 pellets. �-180Kenneth W. Kendall 500 West Prospect 28-J Fort Collins, Colorado 80526 MEAN NUMBER OF PELLETS PER MULE DEER PELLET GROUP KENNETH W. KENDALL, Department of Fishery and Wildlife Biology, Colorado State University, Fort Collins, Colorado 80523 THOMAS M. POJAR, Colorado Division of Wildlife, Fort Collins, Colorado 80526 Abstract: A clear definition of how many fecal pellets constitutes I group has not been established but must be considered when using the pellet group census technique of estimating numbers of mule deer (Odocoileus hemionus). The mean number of pellets deposited per group by mule deer on natural range was estimated as 157 ± 12 (SE). The minimum number of pellets counted in any group was 69. The data seem to indicate definitions of fewer than 30 pellets are unnecessary. Using definitions of 30"or more pellets can significantly reduce the complexity and expense of sampling. Wildlife managers have often estimated numbers of deer (Odocoileus sp.) using the pellet group census technique (Bennett et ale 1940, Harris 1959, Dzieciolowski 1976). Pellet group densities provide an index to population level that can be calibrated to estimate population number (Overton 1971). When estimating total number of deer is the objective, an accurate pellet group counting procedure becomes important. Several scientists have evaluated various aspects of the counting procedure :(Bowden et ale 1969, Eberhardt and VanEtten 1956, Neff 1968, Robinette et al. 1958, Smith 1968, VanEtten and Bennett 1965, Wallmo et ale 1962). However, no clear definitions of how many individual pellets constitutes 1 group has been determined. Anyone who considers using the pellet group census technique must ultimately confront the problem of defining a pellet group. Definitions used in previous studies'have been arbitrary and vary a great deal. Anderson (pers. comm.) uses 5 pellets of similar size, shape and color as the criteria for a group; Ryel (1972) used 10 pellets and Neff (1968) used 30 pellets. Figure 1 shows the effect various definitions can have on the population estimate (Kendall unpubl. data). Defining a pellet group as some specific number of pellets has been difficult because little is known about the mean number of pellets deposited per defecation or decreases in number of pellets over time due to physical and biological dispersion (Wallmo et al. 1962). Smith (1964) found the number of pellets deposited per group by penned mule deer (Odocoileus hemionus) varied dramatically. Strong and Freddy (1979) estimated mean number of pellets per group, for wild deer on the Piceance Basin winter range, as 132 + 12 (SE) but age of groups was subjectively determined and no attempts were made to minimize loss of pellets from dispersion. �-181- The purpose of per group, for wild when dispersion of establish the basis a clear definition this study was to estimate mean number of pellets deer on annual range, from groups of known age and pellets is minimized. Such information could help for further investigations and ultimately lead to of a pellet group. I gratefully acknowledge the Colorado Division of Wildlife for providing funding. Paul Morency for his assistance in data collection. I also wish to thank the many landowners who cooperated with this study. METHODS Pellet count data were collected in conjunction with a pellet group census conducted by the Colorado Division of Wildlife from late May until September 1979. The study area (Fig. 2) consisted of 32 randomly selected 2.59 km2 quadrats, representing 2,657 km2 of annual mule deer range, located in High Park near Canon City, Colorado. Two 1.609-km linear transects were randomly selected from each quadrat. Each transect contained 80 circular sample plots at 20.1-m intervals. The center of each .001 ha plot was permanently marked by a 25-cm rebar stake. All pellets were cleared from the sample plots once each year for 2 years before sampling. Therefore, all groups were assumed to be new and deposited not more than 1year before sampling. Loss of pellets from dispersion was minimized by purposely selecting groups for sampling from plots that occurred on relatively flat terrain with stable soil conditions or vegetative cover. Groups that were compact and appeared complete and undisturbed were selected more often. Individual pellets were identified by similarities in size, shape and color according to search procedures reported by Neff (1968). RESULTS AND DISCUSSION A total of 89 pellet groups were selected for sampling and the mean number of pellets per group was 157 + 12 (SE). Comparison with results reported by Strong and Freddy (1979)-shows a higher mean number of pellets per group for this study (Table 1). The difference in sample size may be ~espohsible but variations in the sampling procedures are more likely to have resulted in a higher mean for this study. A higher mean was expected because pellet groups were purposely selected on the basis of appearance and could have resulted in larger groups being selected more often. Conversely, sampling groups at random, as Strong and Freddy (1979) did, does not minimize loss of pellets from dispersion and could result in fewer pellets than w~re originally deposited. Another possible factor that must be considered is the difference between study areas. The High Park area is occupied by deer all year and their diets vary with seasonal changes in available browse. The �-182- Piceance area is primarily deer winter range and diets of deer there would be restricted to only those types of browse available during winter. Therefore, similarities in the diets of deer between studies could be greatly reduced. Significant differences in diet can affect the number of pellets deposited per defecation (Ferguson 1955, Smith 1964). CONCLUSIONS Table 1 suggests that defining a new pellet group as any number less than 30 individual pellets is unnecessary. Definitions of fewer than 30 pellets riot only effect the population estimate (Fig. 1), but also increase the difficulty and expense of sampling. Additional research to document rates of pellet dispersal, in relation to physical and biological factors, could help verify this conclusion and establish a clear definition of a pellet group. Clearly defining a pellet group as some number of pellets greater than 30 could reduce the complexity and expense of sampling and thereby increase the utility and acceptance of the pellet group census technique. LITERATURE CITED BENNETT, L. J., P. E. ENGLISH, and R. McBAIN. 1940. A study of deer populations by use of pellet group counts. J. Wildl. Manage. 4:398-403. BOWDEN, D. C., A. E. ANDERSON, and D. E. MEDIN. 1969. Frequency distribution of mule deer fecal group counts. J. Wildl. Manage. 33:895-905. DZIECIOLOWSKI, R. 1976. Roe deer census by pellet group counts. Theriol., 21(24-31):351-358: EBERHARDT, L., and R. C. VanETTEN. count as a deer census method. Acta 1956. Evaluation of the pellet group J. Wildl. Manage. 20:70-74. FERGUSON, R. B. 1955. The pellet group count method of censusing mule deer in Utah. M.S. Thesis, Utah State Agric. College, Logan. 94pp. HARRIS, J. T. 1959. Total mule deer population estimates from pellet counts. Proc. 39th Annual Conf. W. Assoc. State Game & Fish Comm. 237-247. NEFF, D. J. 1968. The pellet count technique for big game trend, census, and distribution: A review. J. Wildl. Manage. 32:597-624. OVERTON, W. S. 1971. Estimating the numbers of animals in wildlife populations, Pages 430-432 in R. H. Giles, ed. Wildlife management techniques. 3rd ed. The Wildlife Society, Washington, D.C. ROBINETTE, W. L., R. B. FERGUSON, and J. S. GASHWILER. 1958. Some problems in the use of deer pellet group counts. Trans. 23rd . N. Amer. Wildl. Conf. 411-425. �-183- RYEL, L. A. 1972. Evaluation of a pellet group survey in Michigan. Diss. Abstr. Int. B. Sci. Eng. 32:6763-6764. SMITH, A. D. 1964. 28:435-444. Defecation rates of mule deer. J. Wildl. Manage. SMITH, R. H. 1968. A comparison of several sizes of circular plots for estimating deer pellet group density. J. Wildl. Manage. 32:585-592. ·STRONG, L. L., and D. J. FREDDY. 1979. Number of pellets per mule deer defecation. J. Wildl. Manage. 43:563-564. VanETTEN, R. C., and C. L. BENNETT, JR. 1965. Some sources of error in using pellet group counts for censusing deer. J. Wildl. Manage. 29:723-729. WALLMO, O. C., A. W. JACKSON, T. L. HAILEY, and R. L. CARLISLE. 1962. Influence of rain on the count of deer pellet groups. J. Wildl. Manage. 26:50-55. �-184- Table 1. Comparison of pellet counts per group between High Par~/and Piceance Basi~/study areas. High Park Piceance Basin 89 34 Mean (X) 157 132 Standard Error (SE) +12 +12 69-303 42-320 Sample size (n)£/ Number of Pellets Range (Min.-Max.) ~/Study area located on the Arkansas River drainage near Canon City, .Colorado. £/Piceance Basin study area, results reported by Strong and Freddy (1979). £/Number of pellet groups sampled. �-185- •......• ~ ....._., 40 ,- p;:J E-t ~ H E-t CI) 30 ,- p;:J z 0 H E-t <t! H. 20 ~ p.. 0 Pi Z H p;:J 10 '- CI) <t! p;:J ~ 0 Z H v , 1-5 6-10 j 11-15 16-20 I ! 21-25 ! I 26-30 NUHBER OF PELLETS DEFINED AS A GROUP Fig. 1. Effects of pellet group definitions on the population estimate (Kendall, unpublished). Percentage increases are in relation to defining a pellet group as more than 30 pellets. �Kendall -186- RIO 1 ~ i bl.AN('o i--1' I i /l'IE.SA 1 .J I The study area, located in High Park on the Arkansas river drainage near Canon CitYJ Colorado. �-187- I Appendix B �-188I CANON CITY DEER INVENTORY VEGETATION LIST �-189- FAMILY Agavaceae (Agave) Alliaceae (Onion) Anacardiaceae Anacardiaceae Asclepiadaceae Asclepiadaceae Betulaceae Betulaceae Boraginaceae (2)Brassicaceae Cactaceae Cactaceae Campanulaceae Cappidaceae Caryophylaceae (5)Chenopodiaceae Chenopodiaceae Chenopodiaceae Chenopodiaceae Chenopodiaceae Compositae Compositae Compositae Compo sitae Compo sitae Compositae Compo sitae Compositae Compo sitae Compositae Compositae Compositae Compositae Compositae Convolvulaceae Cyperaceae Ericaceae Equisetaceae Fagaceae Grossulariaceae Iridaceae Juncaceae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Malvaceae GENUS Yucca Allium Rhus Toxicodendron Asclepias Asclepias Alnus Betula Lappula Erysimum Coryphantha Opuntia Campanula Polanisia Arenaria Atriplex Halogeton Kochia Salsola Sarcobatus Ach.illea Ambrosia Ambrosia Artemisia Chrysopsis Chrysothamnus Cirsium Erigeron Erigeron GrindelIa Helianthus Hymenoxis Solidago Tragopogan Convolvulus Carex Arctostaphylos Equisetum Quercus Grossularia Iris Juncus Astragalus Astragalus Lupinus Melolotus Robina Thermopsis Trifolium Spharalcea SPECIES glauca cernuum trilobata rydbergii englemanniana speciosa tenuifolia fontanalis sp. asperum vivipara compressa sp. dodecandra fendleri con-fertifolia glomeratus scoparia kali vermiculatus lanulosa psilostachya trifida sp. villosa nauseosus sp. (3) sp. marcanthus squarrosa annus richardsonii missouriensis sp. arvensis sp , (2) uva-ursi sp. gambelii sp. missouriensis sp , (2) crassicarpus sp. (2) sp. officianlis pseudoacacia divaricarpa sp. (2) coccinia COMMON NAME * soapweed nodding onion skunkbush poison ivy milkweed showy milkweed narrow leaf alder river birch stickweed w. wallflower ball cactus prickly pear harebell clammy weed sandwort 4-wing saltbush * * * * * fireweed russian thistle greasewood yarrow w. ragweed giant ragweed sage golden aster rubber rabbitbrush thistle daisy showy daisy curly gumweed common sunflower rubberweed goldenrod salsify small bindweed sedge bear-berry horsetail gamble oak gooseberry wild iris rush groundplum milkvetch milkvetch bluebonnet yellow sweetclover black locust golden banner clover scarlet globemallow * * �-190- FAMILY Onagraceae Onagraceae Onagraceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Polygonaceae Polygonaceae Polygonaceae Polygonaceae Polemoniaceae Polemoniaceae Polemoniaceae Pinaceae Pinaceae Pinaceae Pinaceae Pinaceae Pinaceae Pinaceae Pinaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Ranunculaceae GENUS Epilobium Epilobium Oenothera Agropyron Agorpyron Agrostis Blepheroneuron Bouteloua Bouteloua Bromus Bromus Bromus Buchloe Cenchrus Echinochloa Elymus Hordea Koeleria Muhlenbergia Panicum Phlem Poa Schizachrium Setaria Sitanion Sporobolus Stipa Rumex Rumex Eriogonum Eriogonum Gilia Ipomopsis Polemonium Juniperus Juniperus Pinus Pinus Pinus Pinus Pinus Pseudotsuga Cercocarpus Fragaria Geum Potentilla Prunus Rosa Rubus Anemone SPECIES angustifolium sp. (3) caespitosa sp , (3) cristatum sp. trichlepis gracilis curtipendula japonicus marginatus tectorum dactyloides sp. crusgalli sp. sp. cristata montana sp. pratense pratensis scoparium sp. hystrix airoides robusta sp. crispus alatum jamesii aggregata aggregata delicatum scopulorum communus aristata contorta edulis flexilis ponderosa menziesii mont anus americana triflorum fruiticosa americana woodsii strigosus sp. COMMON NAME f Lr eweed willow-herb gumbo lily wheatgrass crested wheatgrass bentgrass pine dropseed blue grama side-oats grama japanese brome mtn. brome cheatgrass buffalo grass sand burr barnyard grass rye grass foxtail junegrass mtn. muhly witchgrass common timothy kentucky bluegrass little bluestem bristle grass squirrel-tail alkali sacaton sleepy grass dock curly dock buckwheat buckwheat skyrocket gilia scarlet gilia jacobs ladder Rocky mountain juniper common juniper bristlecone pine lodgepole pine pinyon pine limber pine ponderosa pine douglas fir mtn. mahogany strawberry Prairiesmoke shrubby cinquefoil wild plum rose rasberry windf1_ower * * * * �-191- FAMILY Ranunculaceae Ranunculaceae Ranunculaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae Salicaceae· Salicaceae Salicaceae Saxifragaceae Selaginellaceae Tamaricaceae Umbelliferae Urticaceae Violaceae Viscaceae * GENUS SPECIES Clematis Pulsatilla Thalictrum Castilleja Pedicularis Penstemon Verbascum Veronica Populus Populus Salix Heuchera Selaginella Tamarix Heraculum Urtica Viola Arceuthobium Plants are known to occur within collected for practical reasons sp. patens sp. sp. parryi sp. (6) thapsus americana sargentii tremuloides sp. sp. weatherbiana gallica lana tum diolca nutallii sp. the study area but were not () Number of specimens in that particular could not be identified further family or genus that COMMON NAME virgins bower pasque flower meadow rule indian paintbrush lousewort beard tongue velvet mullein american brooklime Plains cottonwood quaking aspen willow alum root little clubmoss salt cedar cow parsnip stinging nettle nutall violet mistletoe * * * ��July, -193- 1980 JOB FINAL REPORT State of Project C=:_;O::;_;L=:_;O;;.;:RAD=:...:O:;...__ _ Big Game Investigations W-126-R-3 No. Work Plan No. 2 Deer Job No. 5 Experimental Southwest Period InvestiQg~a~t~i~o~n~s~ Deer Inventory _ - Region Covered: Personnel: July 1, 1979 through Roland C. Kufeld, June 30, 1980 Jim Olterman, David C. Bowden P. N. OBJECTIVE To design appropriate sampling and analytical procedures necessary to reliably estimate deer numbers and buck:doe: fawn ratios on winter range based on a preliminary sampling of a problem management unit within each of the four administrative regions of the state. CURRENT STATUS A manuscript entitled "A Helicopter Quadrat Census for Mule Deer on Uncompaghre Plateau, Colorado" has been accepted for publication in the Journal of Wildlife Management. Responsibility for continuation of the census as a management technique has been assumed by the Southwest Region. Maps and other materials pertaining to the census were turned over to them, and assistance was provided to the Southwest Region by the Research Section in flying part of the winter census during February, 1980. Prepared by 1[;._~~C.. 1{l-/d.cI Roland C. Kufeld Wildlife Researcher C ��-195- JOB PROGRESS State of REPORT COLORADO Project No. Big W-126-R-3 ----~--~~------------ Work Plan No. 2 ---------------------- Job No. Activity July, 1980 6 Patterns Game Investigations Deer Investigations Winter Habitat Selection of Mule Deer in Front Range Shrubland and and Forest Habitats Period Covered: Personnel: July 1, 1979 through June 30, 1980 Roland C. Kufeld P. N. OBJECTIVES 1. To test Telonics telemetry equipment to determine its accuracy at various distances in locating a transmitter on the Horsetooth Mountain Study area, and the ability of an observer using that equipment to correctly detect deer activity patterns by habitat type. 2. To determine habitat selection and activity patterns of mule deer winter habitat types on the Horsetooth Mountain area during winter. SEGMENT OBJECTIVE 1. Prepare a detailed study plan. 2. Prepare an environmental assessment. METHODS AND MATERIALS A Program Narrative was written for the study. A study area was selected on Horsetooth Mountain about 4 miles west of Fort Collins. Land ownership patterns on the study area were mapped, and the major landowners contacted for permission to conduct the study on their property. Habitat types are currently being classified and mapped .• Prepared -«. ~~=-....=fP.(d~:tY/~/_/ _ __ by __:.~ __ . --:-/6_.., __ Roland C. Kufeld Wildlife Researcher C �� https://cpw.cvlcollections.org/files/original/a43c5a38ef6990661ff06bd4597ea4fe.pdf ab758dd089cada9bf3980d52926684ba PDF Text Text -197- July, 1980 JOB FINAL REPORT State of Colorado --~~~~~----------- Project No. W-126-R-3 Big Game Investigations Work Plan No. 3 Elk Investigations Job No. 1 Simulations of the Carrying ----~~~---------- Capacity of the Rocky Mountain National Park Elk Winter Range Period Covered: Personnel: July 1, 1979 through June 30, 1980 D. L. Baker, Dr. N. T. Hobbs, D. M. Swift, J. E. Ellis, J. Ritchie, L. Stevens, -c. Finch ABSTRACT 1. Nitrogen and energy based carrying capacities were estimated for Rocky Mountain National Park elk winter ranges. Quantification of forage energy and nitrogen supply plus extant knowledge of elk energy and nitrogen requirements provided basic information for baseline carrying capacity estimates and allowed examination of model parameters most sensitive to change in animal requirements and range supply. . . While much less nitrogen was available to elk than energy, estimated elk nitrogen requirements were also much smaller; consequently during both years, energy and nitrogen based estimates of carrying capacity were similar. Large annual differences were observed. Baseline carrying capacity calculated from forage energy supply was 1481 elk during Year 1 and 991 (33% fewer) during Year 2. Carrying capacity based on nitrogen showed greater variation; during Year I the winter range could support an estimated 1674 elk at maintenance while the following year 41% fewer (994) could be carried. Results of the sensitivity analysis showed the range-supply animal-demand algorithm to be most sensitive to changes in metabolic fecal nitrogen excretion rates; a 25% increase resulted in 74-86% reductions in estimates of carrying capacity. 2. Botanical composition and nutritional quality of diets of tame elk were affected by advancing season and plant communities chosen for feeding on alpine-subalpine summer ranges. Graminoids dominated summer diets in all habitats sampled during both summers. At comparable sampling periods, graminoids contained higher cwe levels, higher proportions of hemicellulose, and cellulose and less lignin than shrubs or forbs. Digestion coefficients were highest for graminoids and forbs, while forbs contained the highest percentage of cell soluables, crude protein and lowest fraction of cell wall as indexed by acid-detergent fiber. Forage and diet quality were similar in all habitats sampled during both summers. Diet quality showed different patterns of change between years for diet crude protein and in vitro digestible dry matter (IVDDM). During Year 1, IVDDM in diets declined from a mean of 54% during July �-198- to 43% in September. The second year, diet IVDDM did not decline over the same time interval but remained relatively constant. Diet crude protein declined over summer during Year 1 from 17% to 10% and from 15% to 10% the following year. �-199- SYSTEMS MODELING BIG GAME POPULATIONS: SIMULATIONS OF THE CARRYING CAPACITY OF THE ROCKY MOUNTAIN NATIONAL PARK ELK WINTER RANGE D. L. Baker, and N. T. Hobbs P. N. OBJECTIVE To develop and test a model simulating carrying capacity of elk winter ranges using Rocky Mountain National Park elk as a study population. SEGMENT OBJECTIVES 1. Complete nutritive evaluation of elk forage samples from Rocky Mountain National Park. 2. Run model simulations to estimate elk carrying capacity of Rocky Mountain National Park winter range. 3. Examine the dietary relations of elk, bighorn sheep, and mule deer, and assess the effects of those relationships on the carrying capacity of each species in Rocky Mountain National Park. 4. Prepare final reports, publications, and plan the next phase of study. 1. Estimate Carrying Capacity of RMNP Elk Winter Ranges. METHODS AND MATERIALS Investigations were conducted on upper montane winter range on the east slope of the continental divide 8 km west of the town of Estes Park, Colorado. Total area of this winter range is about 4000 ha, 2000 ha of which provide substantial amounts of forage. Elk generally occupy the range from October through April. For detailed discussion of weather conditions, vegetation, and topography see Hobbs and Baker (1979). Carrying Capacity Model - Carrying capacity for the winter range was calculated according to a modified formula of Mautz (1978): m K = L i (Bi x Fi) (Rq • Days)-En where K m = b. F: Rl. = q en Days = number of elk which the range can support for the winter period. number of elk forages. biomass of forage species i. nutrient content of forage species i. individual elk requirements; energy or nitrogen requirements for daily maintenance. endogenous reserves of nutrient. number of days elk occupy the winter range. �-200- Thus, data required for this estimate included identification of elk forage species, estimates of their quantity and quality, and information on elk nutritional requirements and nutrient reserves. Forage Quantity and Quality - Forage species consistently included in elk diets were identified by observing diet choices of tame, trained elk grazing on the winter range during each month from November through March of 1976-77 and 1977-78 (Hobbs and Baker 1979). Principal foods were defined as species contributing 2% or more of observed diets during any month; in total these species accounted for 92% of overwinter diets. Biomass of principal forages was estimated at the end of the growing season during 1976 and 1977. Thirty-two 1 ha stands of vegetation, stratified by habitat type, were each sampled with 30 ~ m2 plots for herbs and fallen leaves, and 102m2 plots for current stem growth of shrubs. Herbacious material was clipped at ground level; shrub production was collected between ground level and 2.5 m high. All species were individually separated, dried at 100 C for 48 hour, and weighed to the nearest 0.1 g. Mean values for forage biomass within a vegetation type were mUltiplied by the area of that type and summed across types to estimate total winter range biomass. Habitat areas were estimated by planimeter measurement of vegetation maps prepared from aerial photos and ground surveys (D. Stevens, unpublished data). In vitro digestible dry matter (IVDDM) and crude protein content of individual forages were determined according to procedures described in Hobbs and Baker (1979). Bomb calorimetry was used to determine caloric values which were expressed as gross energy (GE). Metabolizable energy (ME) content of forage species was estimated by multiplying gross energy times IVDDM times 0.85. This calculation provides a reasonable estimate of ME since IVDDM and percent in vivo digestible energy can be assumed to be related approximately 1:1 (Mould, 1980, Milchunas et al. 1978, Moir 1961, Rittenhouse et al. 1971, Ruggerio and Whelan 1976), and }lli is consistently about 85% of digestible energy (Simpson 1976, Mautz et al. 1974, Smith 1971, Thompson et al. 1973). Truly digestible nitrogen was estimated as the product of nitrogen content and estimated true nitrogen digestibility, 0.90 (Robbins 1973:Table 26). Range supply of ME was calculated as the sum of products of forage biomass values times their ME content. Nitrogen supply was determined similarly, except that forages which contained less than 5.6 g N • kg-1 of dry matter (3.5% crude protein) were excluded from the summation. These forages were eliminated from consideration because metabolic fecal nitrogen production for elk is about 5.6 g N . kg-1 dry matter intake (Mould and Robbins 1980). Since the nitrogen cost of digesting these forages exceeds their nitrogen contribution, such forages could provide no net nitrogen to the animal. Endogenous Energy and Nitrogen Reserves - Based on studies of several North American and European cervids (McEwan 1975, Anderson et al. 1972, Robbins et al. 1974, Simpson 1976) we inferred that elk entering the winter have fat reserves of 15% of their body weight, and that 1 g of fat yields 6 kcal of ME (Mautz et al. 1976). Nitrogen reserves were much more difficult to estimate. However, it is known that when dietary protein is �-201- defjcient, lean body is readily catabolized to meet nitrogen requirements (reviewed by Swick and Benevega 1977). Although nitrogen reserves in wild ruminants are not well characterized, we assumed (for use in sensitivity analysis) that elk could mobilize 10% of lean body to meet their amino acid requirements. Elk Energy and Nitrogen Requirements - We assumed no costs for lactation or growth were incurred by elk during winter. Energy and nitrogen requirements for maintenance were based on a weighted mean elk body mass (200 kg) derived from data on the age and sex structure of the resident herd (D. Stevens, personal communication) and elk body weight data of Dean et ale (1976). Carrying capacity estimate are reported in units of 200 kg animals. Daily energy requirements for maintenance were calculated from energetics data of Gates and Hudson (1978) coupled with elk activity budgets. Activity patterns of elk (time spent feeding, resting and travelin~were determined by continuous monitoring of 2-5 elk fitted with Te10nics activity collars. Each elk was monitored for 3, 24 hour periods during each month from .November through April of 1979. Time spent in each activity was multiplied by the estimated energy cost of that activity, and the costs summed over 24 hours (Table 1) •. Simulation modeling studies (Swift et a1. 1980) indicated that during the 2 winters studied, thermoregulatory costs for elk were insignificant. Table 1. Calculations of daily energy requirements Time Spent in activity (hr Activity F ee dd~ng-b/ for 200 kg elk. a/ . [1) Energy Cost. (kcal, . kg-l • h-1) Daily Cost (kca1) 11.38 2.37 5394 Bedding 9.77 1.03 2013 T rave 1" arig+c/ 2.85 3.12 1778 9185 Total ~/Data from Gates and Hudson b/ - Assume c/ - Assume 1978 2 kph walk 4 kph walk Nitrogen requirements for maintenance were equal to the amount of nitrogen of metabolic origin in the urine and feces. Metabolic fecal nitrogen was approximated as 5.6 g • kg-l dry matter intake (Mould and Robbins 1981). We assumed that elk voluntary intake was limited by rumen fill on diets of �-202- the nutritional quality we observed (Hobbs and Baker 1979); intake levels were estimated at 5 kg • d-1 for the calculation of metabolic fecal nitrogen (Hobbs 1979:82, Geis 1950, Hungerford 1948). Endogenous urinary nitrogen was estimated as .16 g • d-1 wtkgO.75 (Mould and Robbins 1980). Thus, total maintenance nitrogen requirements for a 200 kg elk were approximated as 36 g N • d-1. Gestation requirements for energy and nitrogen were estimated according to formulations of Moen (1973:340, 353) assuming 180 days of gestation on the winter ranges and 12 kg fetus at term. Sensitivity Analysis - A baseline prediction of carrying capacity was calculated according to the formula described above. For this estimate, we used energy and nitrogen requirements for maintenance, and assumed no weight loss by wintering elk. The baseline estimate was then compared to recalculated carrying capacity based on adjusted model parameters to examine sensitivity of the model predictions to changes in animal requirements and range supply. These adjustments were as follows: 1. To examine the importance of endogenous reserves, the caloric value of 90% of elk fat reserves and the nitrogen content of 10% of. lean body was subtracted from overwinter requirements. Since elk almost certainly lose weight during winter this is our best estimate of supportable animal density. 2. Energy and nitrogen costs of gestation were added to maintenance requirements of the adult female portion of the herd. 3. Metabolic intervals possible. 4. To compare model sensitivity to changes in energy costs relative to nitrogen requirements, activity and bedded energy requirements were incremented by 25%. 5. To simulate effect of plant succession and overgrazing of willow and aspen (Olmstead 1979), energy and nitrogen content of these shrubs was subtracted from range supply. fecal nitrogen rates were increased by 25%. Confidence on estimates of these rates suggest an increase is quite RESULTS AND DISCUSSION Principal elk forage species included 9 grasses, 5 shrubs, and a single forb; these species contributed 2.45 x 109 cal of HE and 11. 0 x 103 kg nitrogen to range supply during 1976-77 and 1-;64 x 109 kcal HE and 6.6 x 103 kg nitrogen the following year (Table 2). Seventy-five percent of elk forage nitrogen and 86% of energy was contained in graminoids. During both years, energy and nitrogen based estimates of carrying capacity were strikingly similar. However, we observed large annual differences. Baseline carrying capacity calculated from forage energy supply was 1481 during Year I and 991 elk (33%) during Year 2. Carrying capacity based on nitrogen showed greater annual variation; during Year I the winter range could support an estimated 1674 elk at maintenance, while the following year 41% fewer, 994 animals could be carried. These estimates are plausible given existing population levels of 1500-1600 animals (Bear and Green 1980). �-203- Table 2. Contrl.butionof principal elk forages to winter range metabolizable energy (~~E)and nitrogen supply in Rocky Mountain National Park, Colorado, during 1976-1978. Forage~j GRAHINOIDS Bromus inermis b BiomassSl Year-' (kg x .03) Range Supply fl Forage Concentration "'1Fi!7 Nitrogeri_gl ME Nitrogen(kca.l x g-l) (g x kg-I) (kcal x 106) (kg) 6.0 16 60 1.4 1.6 6.6 6.2 70 64 330 248 320 210 1.5 1.5 6.8 5.7 480 315 2176 1197 1977 1978 500 400 1.6 1.5 6.2. 6.6 800 600 3100 2640 1977 1978 ~I ~I 30 1.4 4.4 42 %.J 1977 1978 10 1.6 ~I ~I Bouteloua gracilis 1977 1978 50 40 Calamagrostis canadensis 1977 1978 Carex spp. Juncus balticus Muhlenbergia montana 1977 1978 240 100 1.5 1.6 5.7 5.0 360 160 1368 500 Phleum pratense 1977 1978 80 10 1.8 1.8 6.0 5.6 144 18 il Poa pratensis 1977 1978 40 30 1.7 1.6 7.2 5.8 69 48 288 174 Stipa comata 1977 1978 60 60 1.7 1.7 7.1 4.9 102 102 426 294 6(}1/ 60 1.8 1.6 8.1 5.6 108 96 486 BROWSE Populus tremuloides 1977 (leaves) 1978 480 il Potentilla fruticosa (stems) 1977 1978 6 5 1.1 1.1 7.8 6.6 5 47 33 Purshia tridentata (stems) 1977 1978 80 40 1.5 1.2 12.0 9.8 120 48 960 392 Rosa woodsii ~tems) 1977 1978 10 10 1.5 1.4 7.1 6.6 15 14 71 66 Salix spp. (leaves) 1977 1978 2(j 20 1.3 1.1 8.6 8.9 26 22 172 178 Salix spp. (stems) 1977 1978 60 50 1.7 1.5 11. 7 8.6 102 75 702 430 FORBS Eriogonum umbellatum 1977 1978 3#' 1.0 1.0 11.2 9.1 30 30 336 273 2449 11002 1639 6536 Total Supply 1977 1978 hI 30 7 �-204- Table 2. (Continued) -a/S·peCles J 2% or more to e lk· wlnter ddlets (H0bb s et a 1 • 1980) • contrlLbut utlng £/1977 = Footnotes 0 Winter of 1976-77, £/Total biomass on winter and Baker (1979). 1978 = winter range for individual d/ - ME = gross energy x in vitro dry matter IVDMD data from Hobbs et al. (1980). ~Nitrogen of 1977-78. N content x .90. habitat digestibility data see Hobbs (IVDMD) x .85. N content data from Hobbs et al. (1980). i/Includes only those species which contain greater than 5.6 g N kg-1 N. Values not reported for forages containing less than this amount. _g_/Notin diet. h/Estimate based on 1977-78 data. However, there are sources of bias in these predictions. At voluntary -1 intake levels of 5 kg • d-1 elk diets would have to contain 1.8 kcal • g ME and 7.3 g • kg-1 nitrogen to allow maintenance based on our estimates of nutrient requirements. Although most shrubs contained adequate nitrogen, and many grasses, adequate energy, a substantial portion of forage supplies, provided concentrations of nutrients inadequate for maintenance on diets limited in quantity by rumen fill (Table 2). These findings support the hypothesis of White (1978) and concur with the findings of Wallmo et al. (1977) that the relative quality of foods may be as important as their absolute abundance in determining supportable animal density. Consequently, our baseline estimates of carrying capacity overestimate the number of animals which could be supported at energy and nitrogen equilibrium. Realized nutritional status will depend on how forages are mixed in the diets (Hobbs 1979) and the length of winter (Wallmo et al. 1977). An additional bias results from probable declines in forage biomass during the winter period due to trampling, shattering, and wind losses. Such decrements can be substantial (review by Pieper et al. 1974); since we do not account for these losses, our baseline estimates are inflated. Results of the sensitivity analysis (Table 3) showed the range-supply animal-demand algorithm to be very sensitive to changes in metabolic fecal nitrogen excretion rates; a 25% increase in this parameter resulted in 74-86% reduction in estimates of carrying capacity. In comparison, proportionally equal changes in energy costs of activity and resting had relatively small effects, reducing carrying capacity 10% and 15% respectively. The large influence of metabolic fecal nitrogen is due to its simultaneous effect on animal requirements and range supply. A 25% increase elevates this rate to 7 g N • kg-1 dry matter intake. If the animal excreted nitrogen at this rate the only forages which could contribute to nitrogen balance were browses and a very few �-205- grasses. This effect is exacerbated because total nitrogen requirements are elevated from 36 to 50 g N • d-1. Consequently, the animal needs more nitrogen and has less food which can provide it. Table 3. Elk carrying capacity estimtes for winter National Park, Colorado, during 1976-1978. Estimated Carrying CaEacityC! Energy Nitrogen bl YearBased Based Condition range in Rocky Mountain % Change From Baseline Energy Nitrogen Based Based B ase 1· al arie+ 1977 1978 1481 991 1674 994 Add gestation~!/ costs 1977 1978 1480 991 1656 1043 0 0 - 1 Lose 90% fat, 10% lean body 1977 1978 1657 1109 1900 1128 +12 +12 +11 +11 Increase metabolic fecal N by 25% 1977 1978 1478 989 388 141 0 0 -74 -86 Increase activity energy costs by 25% 1977 1978 1330 890 1674 994 -10 -10 0 0 Increase bedded energy costs by 25% 1977 1978 1404 939 1674 994 - 5 - 5 0 0 Eliminate willow shrubs and aspen 1977 1978 1338 874 1446 851 -10 -12 -12 -14 - 1 ~/Estimate based on measured energy and nitrogen supply and best aproximations by elk requirements, assuming no weight loss. All other conditions are specified relative to baseline parameters. ~/1977 = Winter of 1977-78, 1978 = winter of 1977-78 ~/Assuming 180 day occupancy hectare divide by 2000. of winter i/Assuming range. To convert to elk per 65% of herd is female. Catabolism of endogenous reserves increased carrying capacity by 12% based on energy and 11% based on nitrogen. Thus, these reserves had a relatively small impact on total supportable animal density; maximum standing crop of animals in poor condition could only be slightly larger than those at maintenance. This does not mean that these reserves are not important to individual animal survival; they doubtlessly are, but it is important that these large differences in animal condition have small effects on carrying capacity. Seral shrubs appeared to offer minor forage reserves to elk populations in Rocky Mountain National Park. Estimates of carrying capacity �-206- without energy and nitrogell contributed by aspen and willow were 10-14% less than when these forages were included in carrying capacity calculations. Thus while it is likely that foraging by elk exerts a substantial impact on the ecology of these species (Olmstead 1979) it appears that they play a minor role in the trophic ecology of elk. Weins (1977) pointed out that a potent force affecting animal populations is periodic deficits in resource supply "bottlenecks".· We demonstrate that such bottlenecks do occur for elk on winter range because predicted supportable animal density was highly variable between years. This variation has important implications for management and for implementation of the carrying capacity model. When resource supply fluctuates from year to year, managers should expect variation in animal numbers unless populations are maintained at densities well below carrying capacity. Thus, carrying capacity should be viewed as a markedly labile rather than static characteristic of the habitat. This variability must be remembered in designing future research for use in carrying capacity estimates. That is, in the face of large annual changes in resource supply it may not be necessary to measure other model inputs as precisely as is sometimes advocated. Thus, we disagree with Robbins et al. (1979: 452) that ~much remains to be accomplished before precise analysis and evaluations of range carrying capacity ••• can be accomplished from an energetics standpoint". In a serial calculation like carrying capacity, the outcome is only as precise as the least precise input; we contend annual variation in resource supply will usually limit the precision of long-term carrying capacity estimates. II •• This does not mean that accuracy of estimates of animal requirements and range supply should not be improved. But, some of these parameters, are far more significant to model prediction than are others as was demonstrated in the sensitivity analysis. For example, we recommend that studies of elk maintenance nitrogen requirements and energy nitrogen interactions precede further study of energetics alone. This follows from the logic of Mautz (1978:322-323) who contended we should study parameters about which we know least; but in addition, we hold that accuracy in some inputs is disproportionately more important to accuracy of carrying capacity predictions. We conclude that in its current state of development nutritionally based estimates of carrying capacity offer a valuable procedure for evaluation of habitat. That is, these approaches allow decisions on the relative quality of ungulate ranges based on measurable attributes of the habitat which are related in a demonstrable way to animal population density. However, we caution that annual variability in resource supply and possible inaccuracy in model input parameters may limit the predictive value of these procedures in the absence of long-term studies and improved input data. 2. Complete Nutritional Analysis of Summer Elk Forage In RMNP and summarize diet composition and quality data. METHODS Methods and materials employed have been previously described AND MATERIALS in summer elk food habits (Baker and Hobbs 1978). investigations �-207- RESULTS AND DISCUSSION Diet Botanical Composition During this study our experimental animals took a total of 110,061 bites of at least 83 species. Relatively few species were taken in significant amounts. Thirteen species accounted for 84% of total bites (Table 4). Average forage class composition of diets (calculated across animals and habitats) was similar between years (Fig. 1). Diet composition with respect to major forage categories was not significantly different (f < 0.05) among habitat types during either year. Graminoids dominated summer diets. Graminoids were the most frequently chosen food item in all habitats sampled during both summers (Fig. 1). Principal grass and grass ..•. like species eaten included sedges (Carex spp.), tufted hairgrass (Deschampsia caespitosa) and rushes (Juncus ba1ticus). Shrub species were most often encountered and eaten in willow park and krummho1z ecotone communities, while forbs were primarily selected on alpine tundra. Leaves stems and inflorescences of willow (Salix brachycarpa Salix planifolia) and blueberry (Vaccinium spp.) were the most commonly selected shrubs. White-marsh marigold (Caltha leptosepa1a) alpine avens (Geum rossii), and clover (Trifolium spp.) were of importance in alpine communi tie s. Forage Quality In general, forage classes showed distinctive nutritional characteristics (Table 5). At comparable sampling periods, grasses contained higher CWC levels, higher proportions of hemicellulose (CWC-ADF) and cellulose (ADFlignin) and less lignin than shrubs or forbs. Digestion coefficients were highest for grasses and forbs while forbs also contained the highest percentage of cell soluables (l-CWC) and crude protein and the lowest fraction of the cell wall as indexed by ADF. In vitro digestible dry matter coefficients of shrub and forb tissues were consistently lower than cell soluable percentages. All soluables of shrubs ranged from 35 to 60% lower than IVDDM values while forbs varied from 5 to 55% below their respective digestion coefficients. While the relatively high lignin content of shrub species has been shown to inhabit microbial degradation of CWC and digestion of cell soluables (Van Soest 1970, Robbins 1973, Blair et al. 1977:674) it is not understood why forbs with low lignin content also exhibited this characteristic. Estimates of forage quality were similar both years (f > 0.06) with the exception of graminoid crude protein and structural constituents of shrubs. Averaged across graminoids summer habitats contained 12% more crude protein during Year 2 than during the previous year (P = .005). Shrub cwe were 11% less; ADF 19% less and lignin 30% less (P < 0.03) the second summer. Forbs did not change in nutrient quality between years. Measurements of forage quality showed different patterns of change .between years. During Year 1, forage classes showed similar rapid declines in nutritive quality with advancing season. Crude protein content of graminoids, shrubs and forbs declined on the average of 41%, 40% and 58% from July to �-208- Table 4. Mean percentage contribution of major forage species to the total number of bites taken by tame elk on alpine-subalpine summer ranges in Colorado during July-September 1977 and 1978, summarized by habitat type~/. Willow Park HABITAT TYPE Krummholz-Ecotone Al:eine Tundra b/ Taxa-- c/ Year- X% SE X% SE X% GRAMINOIDS Calamagrostis canadensis 1977 1978 2 1 2 1 2 2 tr 0 tr Carex spp. 1977 1978 45 42 10 7 45 49 10 9 40 27 7 8 Descham:esia caes:eitosa 1977 1978 4 5 2 4 2 8 1 3 9 16 4 6 Juncus balticus 1977 1978 10 13 4 4 3 5 4 4 6 16 3 8 KoblS:esia myosuroides 1977 1978 tr 0 9 4 5 3 3 1 1 1 All Graminoids~/ 1977 1978 64 62 5 9 68 67 5 8 61 63 4 9 1977 1978 6 4 4 14 10 4 4 2 2 3 1 2 Salix :elanifolia 1977 1978 14 9 3 4 7 9 2 5 4 11 3 5 Vaccinium spp. 1977 1978 9 13 2 3 2 3 2 2 0 0 1977 1978 26 27 6 7 22 24 5 6 8 14 3 6 1977 1978 2 8 1 5 tr 3 3 5 8 3 4 1977 1978 tr tr 3 1 2 1 9 5 2 Polygonum bistortoides 1977 1978 tr tr tr tr 2 Trifolium nanum 1977 1978 0 0 tr 0 4. 2 1+ 1977 1978 tr tr tr tr 9 3 4 1 1977 1978 10 11 32 22 4 1977 1978 18,051 16,149 SHRUBS Salix brachycar:ea d/ All ShrubsFORBS Caltha le:etose:eala Geum rossii Trifolium :earryi d/ All ForbsTOTAL BITES~/ (See footnotes on next page) 3 5 10 9 18,740 23,288 1 3 3 21,055 17,493 SE 4 0.5 3 2- 7 �-209- Table 4. (Continued) Footnotes. a/ - Means are ealculated across animal's diet percentages were pooled for each animal across all months. £/Species include those which contributed any month. £/1977 Sample size d/ - Includes ~/Sum f/ = 4 elk. = 2% or more of total bites during 1978 Sample size = 3 elk. all species selected not just those in the table. of total bites across animals. - trace where percentages < 1%. �Table 5. Nutritional 1977 and 1978. composition of principal forage species consumed by elk on alpine-subalpine summer ranges in Colorado during July-September COnt2osition as a Percent of Oven-Dr~ Weight CWC£/ ADFS./ 1978 Lignin 1977 1978 Crude Protein 1977 1978 Ash IVDD!j~/ 1978 Taxa~/ 1977 1978 1977 GRAMINOIDS Calamagrostis canadensis July August September 61. 2 Y 32.2 Carex sp. July August September 58.9 59.3 62.2 64.4 60.6 56.0 30.4 31.1 28.1 27.4 24.6 25.1 3.8 4.4 4.2 3.2 2.5 1.5 17.2 14.5 10.7 14.9 15.3 9.8 7.0 5.8 6.3. 5.8 5.1 6.4 69.0 62.6 46.5 58.9 65.1 62.0 Descham2sia caes2itosa July August September 64.7 63.7 61.2 67.8 58.3 55.0 36.2 37.2 31.3 33.1 29.0 27.8 3.1 3.5 2.8 2.2 1.7 2.0 12.8 8.7 8.9 11.5 15.3 14.6 7.0 4.6 4,8 6.4 6.1 4.9 55.6 51.8 52.7 54.6 68.1 68.0 Juncus balticus July August September 62.8 64.6 65.0 61.1 61.3 56.3 32.1 34.4 32.1 23.2 27.5 24.2 5.0 6.1 3.3 2.0 2.3 1.9 16.1 11.8 7.0 14.6 15.2 11.3 5.0 4.9 4.3 5.1 5.0 5.0 60.6 56.5 45.0 67.6 66.9 66.6 Kobresia m~osuroides July August September 3.6 10.8 1977 1978 4.9 1977 . 57.4 •.... 0 I 59.0 54.4 y,/ 56.8 55.0 32.8 30.4 23.4 27.4 4.1 3.3 1.9 2.5 13.9 12.0 14.3 14.1 5.5 5.1 4.6 4.3 15.0±l.0 11.8±1. 2 9.4±0.9 13.8±0.4 15.0±0.3 11.9±1.4 6.1±0.5 5.1±0.3 5.1±0.4 5.5±0.8 5.l±0.4 5.4±0.5 52.0 54.0 61.1 60.5 f../ X Graminoids July August September I N 61.4±1.4 60.5±2.3 62.4±0.9 62.5±2.4 58.8±1.4 55.8±0.4 SHRUBS Salix brachycar2a July August September 35.2 34.5 43.9 30.6 33.8 35.2 27.2 27.8 34.6 20.5 23.7 26.1 13.5 14.2 21.0 9.3 12.4 13.4 16.4 11.7 8.6 17.9 13.4 8.9 4.3 4.5 4.0 4.4 3.9 3.4 42.0 39.1. 26.4 47.4 36.6 30.5 Salix 2lanifolia July August September 34.8 25.0 1.4.5 34.7 31. 6 26.7 26.9 17.9 35.4 27.5 21. 5 17.8 15.7 19.8 23.8 14.'0 12.6 13.0 16.8 13.2 8.8 14.1 14.1 10.7 3.6 6.5 8.6 3.3 3.1 2.7 32.4 28.6 24.7 35.0 33.6 31. 4 Vaccinium spp. July August September 58.3 61.7 65.0 51.6 59.1 52.8 48.7 49.7 50.6 40.5 41.5 38.0 26.5 27.6 28.7 18.7 21.7 18.9 10.4 9.4 7.9 9.5 10.3 7.9 2.0 3.4 2.5 2.9 3.3 2.5 19.7 16.5 17.3 22.2 15.9 19.5 32,9±1.2 33.3±1.6 30.5±1.2 26.8±1.2 27.1±0.6 25.7±1.3 4.0±0.4 4.3±0.6 3.5±0.3 2.3±0.2 2.3±0.2 1.8±0. 2 59.3±3.7 56.2±2.3 50.4±2.9 60.6±2.7 65.2±1. 7 64.3±1.8 --------------------------------------------------------------------------------------------------------------------------------------------------- �Table S. Nutritional composition 1977 and 1978. (Continued). of .principal forage species consumed by elk on alpine-subalpine ComEosition Taxa!!.! 1977 CWC~.! 1978 as a Percent of Oven-Dr! Weight ADFcl 1977 Lignin 1978 summer ranges in Colorado during July-September 1977 1978 Crude Protein 1977 1978 IVOOMi.l Ash 1977 1978 1977 1978 31.4±6.S 28.1±6.5 22.8±2.8 j4.9±7.3 28.7±6.S 27.1:':3.8 ._--- X Shrubs July August September 42.8± 7.8 40.4±11.0 51.1± 6.0 33.2:!:1.4 41. S±8. 8 38.2±7.7 20.3 25.5 30.8 29.3 29.7 25.5 17.3 16.7 al bistortoides 16.1 15.1 al FORBS Caltha leptosepala July August September Geum rossi! ~l-y-August September Pol!gonum July August September !/ Trifolium nanum -July August September 30.7 al al Trifolium Earryi July August September al 33.7 al X Forbs July August September ~/Species 22.8± 4.0 19.7± 7.6 30.8 contributing 34.3:1:7.2 31.8±9.4 40.2±5.2 29.5±5.9 28.9±6.3 27.3±5.9 18.6±7.0 20.5±3.9 24.5±2.3 14.0±2.7 15.6±3.1 15. I:!: 1.9 14.5±2.1 11.4±1.1 8.4±0.3 13.8±2.4 12.61:1.2 9.2±0.8 3.3±0.7 4.8±0.9 5.0±1. 8 15.4 17.5 21.5 14.8 12.4 12.7 6.0 6.4 6.3 5.7 2.7 1.9 19.9 12.7 8.3 17.6 14.7 9.1 10.6 8.1 11. 7 8.7 9.8 10.1 75.97 64.8 1,0.5 74.9 60.1 48.4 12.5 11.2 13.3 9.9 3.2 1.6 2.8 2.4 17.3 11.4 15.5 13.6 5.4 4.9 5.1 4.9 43.7 38.4 43.5 36.4 al 28.7 al Y 20.4 dl el - Kcal per gram. 16.2 64.3 5.5 •.... •.... 22.4 7.6 22.7 6.7 67.6 23.2 7.3 16.6 8.6 56.2 I y 22.7±5.4 24.5±4.7 25.5 I N 16.8±2.9 l7.3±3.5 21. 5 14. 1±0. 75 14.2±5.5 12.7 S. 6± 1.3 5.1±1.8 6.3 2% or more of total bites observed during any month. ~/cell wall constituents. c/ - Acid detergent fiber. - In vitro digestible 11.00 3.S±0.5 3.4±0.3 2.9±0.3 dry matter. I !/Species did not occur in the diet that year. alSpecies did not occur in the diet that month. 4.3±1.2 5.4±2.9 1.9 20.0±1.6 13.6±1.6 8.3 16.6±O.9 14.8±0.8 9.1 7.6± 1.6 7.2± 1.2 11. 7 3.0±2:<' 6.7±1.9 10.1 62.4±9.7 53.1:':7.9 40.5 60.9±9.2 53.6±8.7 48.4 �-212- September, respectively. In vitro digestible dry matter decreased 32% 24% and 47%. In contrast to Johnson et ale (1968) cellulose and other structural components (CWC, ADF, Lignin) were variable throughout summer and displayed no predictable relationship to declining digestibility. For Year 2 declines in quality were less rapid for graminoids and shrubs while forbs were similar to Year 1. Peak crude protein levels for most graminoids and shrub species occurred during August, while IVDDM was relatively constant throughout summer. This annual variation in forage quality may have been partially related to late spring snows during Year 2 which covered much of the summer range until mid-July. Diet Quality Nutritional quality of elk diets was estimated by integrating botanical composition and nutrient concentration data. While overall quality of summer diets were similar among habitat types and between years (Table 6) considerable temporal variation in diet quality among habitats within the summer season was observed. Diet quality like forage quality showed different patterns of change between years. While dietary structural components (CWC, ADF, ADL) remained relatively constant over both summer rates of change in dietary crude protein (Fig." 3) and IVDDM (Fig.::4) we'.redifferent. During Year 1, IVDDM in diets declined significantly (P = 0.001) from a mean (across animals and communities) of 54% during July to 43% in September. The second year diet IVDDM did not decline over the same time interval (year x month interaction, P = 0.001) but gradually increased. Diet crude protein showed considerably less variation between animals than did diet IVDDM. Crude protein in diets declined linearly (P = 0.001) over summer during Year 1 from 17% to 10% and from 15% to 10% the following year. Based on these data and on studies of nutritional requirements of wild and domestic ruminants, we infer that elk diets from summer range in Rocky Mountain National Park provide digestible energy and protein in substantial excess of maintenance requirements. From a nutritional standpoint, elk appear to be able to obtain high quality diets in a variety of habitat types. We conclude that animal condition at the end of summer is probably variable from year to year. 3. Examine Dietary Relations of Elk, Bighorn ahd Mule Deer and assess the effects of those relationships on the Carrying Capacity of each Species in RMNP. Nutritional evaluation of the major forage plants of these ungulate is currently in progress. in the diet of each �-213- DIET COMPOSITION Grass and Grass Grasslike 64 Grasslike 64 0/0 22 1977 Figure 1. and 0/0 0/0 1978 Summer forage class composition of tame elk diet's on alpinesubalpine habitats, Colorado during 1977 and 1978. �Figure 2. 18 Crude protein content of summer diets of tame elk on alpine-subalpine summer range, Colorado during 1977 and 1978. Numeral 1 indicates a value for 1977. Numeral 22 indicates a value for 1978. Regression equation for 1977 is Y = 18.31 - 0.09X, R = 0.91, != 0.00001, SE of estimate = 0.87, for 1978 Y = 13.5 + O.llX - 0.002X2, R2 0.93, P = 0.000001, SE of es t Lmatie = 0.55. I I III 197711~ 16~ II~ z m ba:: 0- 1978 14~ ~2 22 I N •..... -I:'- ~ I W 0 ::::> a::: 12 o ~ 0 10 8 ~ July I Au gust I September I �64 DIET IVDDM 60 12 2 II 56 1977, 2 :E o o > 52 2 2 2 ~ 48 ~ ------ ~978 ~ 2 2 2 "" I N ",. I-' VI I " 2 ~2 2 44 2 2 I I 40r July I Figure 3. II Aug ust I September I In vitro digestible dry matter (IVDDM) of summer diets of tame elk on alpine-subalpine summer range, Colorado, during 1977 and 1978. Numeral 1 indicates a value for 1977. Numeral 2 indicates a value for 1978. Regression equation for 1977 is Y = 50.7 + 0.34X 0.005X2, (R2 = 0.58, E. = 0.0000002, SE of estimate = 4.66); for 1978 Y = 53.1 - 74/x, R2 = 0.14, P = 0.35, SE of estimate = 3.7. �-216- LITERATURE CITED Anderson, A. E., D. E. Medin, and D. C. Bowden. 1972. Indices of carcass fat in a Colorado mule deer population. J. Wildl. Manage. 36:579-594. Baker, D. L., and N. T. Hobbs. 1978. Systems Modeling Big Game Pop.: Simulations of the carrying capacity of the Rocky Mt. National Park elk winter range. Colo. Div. Wildl. Fed. Aid W-38-R-33, WP17, Job 2. Prog. Rep., Game Res. Rep. July, Part 2. 169-216. Bear, G. D., and R. A. Green. 1980. Elk population and ecological Colo. Div. Wildl. Game Res. Rept. July: In Press. studies. Blair, R. M., H. L. Short, and E. A. Epps, Jr. 1977. Seasonal nutrient yield and digestibility of deer forage from a young pine plantation. J. Wildl. Manage. 41(4):667-676. Bobeck, B. size. 1977. Summer food as the factor limiting roe deer population Nature 268:47-49. Caughley, G. 1976. Wildlife management and the dynamics of ungulate populations. p. 183-224 in T. H. Coaker ed. Applied Biology Vol. 1. Academic Press. N. Y. Caughley, G. 1979. What is this thing called carrying capacity? p. 2-8 in M. S. Boyce and L. D. Hayden-Wing eds. North American Elk: Ecology, Behavior, and Management. Univ. of Wyoming. 294 p. Coe, M. J., D. H. Cumming, and J. Phillipson. 1976. Biomass and production of large African herbivores in relation to rainfall and primary production. Oecologia 22:341-354. Craighead, J. J., F. C. Craighead, R. L. Ruff and B. W. O'Gara. 1973. Home ranges and activity patterns of nonmigratory elk of the Madison drainage herd as determined by biotelemetry. Wildl. Mono. 33. 50 pp. Dean, R. E., E. T. Thorne, and I. J. Yorgason. mountain elk. J. Mammal. 57:186-189. Gates, C., and R. J. Hudson. 1978. Acta Theriologica. 23:365-370. 1976. Weights Energy costs of locomotion Geis, A. F. 1950. The food consumption and relative various winter diets fed to elk under controlled Thesis. Montana State Univ. 68 pp. of rocky in Wapiti. digestibility of conditions. M.S. Hobbs, N. T. 1979. Winter diet quality and nutritional status of elk in the upper montane zone, Colorado. Ph.D. Thesis. Colo. State Univ., Ft. Collins. 131 pp. Hobbs, N. T., and D. L. Baker. 1979. Systems Hodeling Big Game Pop.: Simulation of the carrying capacity of the Rocky Mt. National Park elk winter range. Colo. Div. Wildl. Fed. Aid W-126-R-2, WP3, Job 1, Prog. Rep., Game Res. Rep. July, Part 2, 231-372. �i>. .< -217- Hobbs, N. T., D. L. Baker, J. E. Ellis, and D. W. Swift. 1980. Botanical composition and nutritional quality of elk winter diets in the upper montane zone, Colorado. J. Wildl. Manage. 44. Hungerford, C. R. 1948. The food consumption and weight response of elk under winter conditions. M.S. Thesis. Univ. of Idaho. 60 pp. Johnston, A., 1. M. Bezeau, and S. Smoliak. 1968. Chemical composition and in vitro digestibility of alpine tundra plants. J. Wildl. Manage. 32(4) :773-777. Maloiy, G. M. 0., R. N. B. Koy, E. O. Goodall, and J. H. Topps. 1970. Digestion and nitrogen metabolism in sheep and red deer given large and small amounts of water and protein. 24:843-855. Mautz, W. W., H. Silver, and H. H. Hayes. 1974. bility of winter deer browse from proximate Zoo1. 52: (11):1201-1205. Predicting the digesticomposition. Can. J. Mautz, W. W., H. Silver, J. B. Holter, H. H. Hayes, and W. E. Urban. 1976. Digestibility and related data for seven northern deer browse species. J. Wildl. Manage. 40:630-638. Mautz, W. W. 1978. Nutrition and carrying capacity. p. 321-348 in J. 1. -Schm.Ld t and D. L. Gilbert eds. Big Game of North America. Stackpole. Harrisburg, Pa. 17105. McEwan, E. H. 1975. cervids compared The adaptive significance of the growth patterns in with other ungulate species. Zool. Zh. 54:1221-1232. Mentis, M. T. 1976. Stocking rates and carrying capacities for ungulates on African rangelands. S. Afr. J. Wildl. Res. 7:89-96. Mentis, M. T., and R. R. Duke. 1976. Carrying capacities of natural veld in natal for large wild herbivores. S. Afr. J. Wildl. Res. 6:65-74. Milchunas, D. G., M. I. Dyer, O. C. Wallmo, and D. E. Johnson. 1978.· In vivo 1 in vitro relationships of Colorado mule deer forages. Colorado Div. Wildl. Special Rep. 43. 44 pp. Moen, A. N. cisco. 1973. Wildlife 458 pp. Ecology. H. H. Freeman and Co., San Fran- Moir, R. J. 1961. A note on the relationship between the digestible dry matter and the digestible energy content of ruminant diets. Aust. J. Agr. Anim. Husb. 1:24-26. Mould, E. D. 1980. Asepcts of elk (Cervus canadensis nelsoni) nutrition and associated analytical procedures. Ph.D. Thesis. Wash. St. Univ., Pullman. 72 pp. �-218- Mould, E. D., and C. T. Robbins. 1981. J. Wildl. Manage. (Submitted). Olmstead,G. E. 1977. Mountain National 141 pp. Nitrogen metabolism in elk. The effect of large herbivores on aspen in Rocky Park. Ph.D. Thesis. Univ. of Colo., Boulder. Pieper, R. D., C. H. Herkel, D. D. Dwyer, and R. E. Banner. 1974. Management implications of herbage weight changes on native rangelands. J. Soil and Water Cons. 29(5):227-229. Rittenhouse, L. R., C. L. Streeter, and D. C. Clanton. 1971. Estimation, digestible energy from digestible dry and organic matter in diets of grazing cattle. J. Range Manage. 24:73-75. Robbins, C. T. 1973. The biological basis for the determination of carrying capacity. Ph.D. Thesis. Cornell Univ., Ithaca. 239 pp. Robbins, C. T., R. L. Prior, and J. T. Reid. 1974. white-tailed deer. J. Anim. Sci. 38:871-876. Robbins, C. T., Y. Cohen, and B. B. Davit. 1979. elk calves. J. Wildl. Manage. 43:445-453. Body composition Energy expenditure of by Ruggerio, L. and J. B. Whelan. 1976. A comparison of in vitro and in vivo food digestibility by white-tailed deer. J. Range Manage. 29:82-83. Simpson, A. M. 1976. A study of the energy metabolism and seasonal cycles of captive red deer. Ph.D. Thesis, Univ. of Aberdeen, Aberdeen, Scotland. 190 pp. Smith, G. E. 1971. Energy metabolism and metabolism of the volatile fatty acids. p. 543-562. In D. C. Church ed., The Digestive Physiology of Ruminants. Vol. 2.--Nutrition. COlvallis, Ore. 801 pp. Swick, R. W., and N. J. Benevenga. 1977. Labile protein protein turnover. J. Dairy Sci. 60:505-515. reserves and Swift, D. M., J. E. Ellis, and N. T. Hobbs. 1980. Nitrogen and energy requirements of North American cervids in winter - a simulation study. Proceedings of the Second Reindeer and Caribou Symposium. In press. Thompson, C. G., J. B. Holte, H. H. Hoyes, H. Silver, and W. E. Urban. 1973. Nutrition of white-tailed deer. I. Energy requirements of fawns. J. Wildl. Manage. 37:301-311. Van Soest, P. J. The chemical basis for the nutritive evaluation of forages. Pages V1-V19. In Proc. of Natl. Conf. on Forage Qual. Eval. and Utile 1969. Nebr. Cent. for Contino Educ., Lincoln. �-219- Wallmo, O. C., L. H. Carpenter, W. L. Regelin, R. B. Gill. and D. L. Baker. 1977. Evaluation of deer habitat on a nutritional basis. J. Range Manage. 30:122-127. Weins, J. A. 1977. On competition Scientist. 65:590-597. in variable environments. White, T. C. R. 1978. The importance of relative animal ecology. Oecologia. 73:71-86. ~-. Prepared by ~~--------~----------------Dan L. Baker Wildlife Researcher C Am. shortage of food in �Table 6. Nutritional composition of summer diets of tame elk in alpine-subalpine Colorado during July-September 1977 and 1978. ._------ADFE.! CWC~/ Habitat X Year SE X X - Willow Park krummholz ecotone Alpine tundra X SE --.------.--~ .... in .._.---- CRUDE PROTETN LIGNIN SE habitats ~------[VDD~/ SE X -. SE ------ 1977 52.6 1.9 29.1 0.9 8.3 0.5 l'L4 0.9 48.8 3.8 1978 48.4 1.2 25.6 0.6 7.45 0.7 13.0 0.9 .51.0 3.4 1977 51. 9 1.7 29.6 1.0 7.1 0.8 13.4 1.3 50.2 3.8 1978 45.5 1.2 24.7 0.4 7.3 0.8 13.2 1.0 49.5 1.6 1977 51.2 2.9 27.2 1.0 7.0 0.9 13.4 1.3 49.7 3.8 1978 42.7 1.6 23.3 0.5 n.n 0.7 13.7 0.7 ')1.9 2 . '-') ~! Cell wall constituents E./ Acid detergent fiber ~/ In vitro digestible dry matter I N N 0 I �-221- JOB PROGRESS State of Colorado Project No. W-126-R-3 ------- July, REPORT Big Game Investigations Work Plan No. 3 Elk Investigations Job No. 2 Elk Population Period Covered: July 1, 1979 Personnel: 1980 and Ecology Studies through June 30, 1980 G. D. Bear, R. A. Green, other contributors in Appendix B are listed ABSTRACT One hundred nine tv-eight elk (91 cows, 65 calves, and 42 ~ulls) were captured during 1979-80 winter period. Trapping success was 8n% Seasonal distribution and within season movements were determined frow frequent relocations of 37 radio collared elk. Elk tagged with radic collars during winter 1978-79 at Masonville summered near Stormy Park in nor t he rr, Rocky Mountain National Park then returned to the Masonville where they spent the 1979-80 winter. Elk radio collared at Buttonrock Reservoir near ~vons in winter 1978-79 summered west of Wild Basin in RMNP then returned to the Buttonrock Reservoir area in winter 1979-80. Elk radio-colla~ed on east slope winter ranges in RMNP summered along the Colorado River (Kawuneeche Valley), Speciemen Mountain, and headwaters areas Fall River and Forest Canyon. Population estimates projected from a weighted mean of 4 capture-recapture estimate replicates indicated 1,575 + 66 elk winter on east slope RMNP winter ranges. Aerial classifications of population structure indicated ratios of 42 calves:100 cows and S6 bulls: 100 cows. Ground classifications revealed ratios of 31 calves: 100 cows and 37 bulls: 100 cows. Known calf mortality during the measurement period was: l3 percent to predators, 13 percent to hunting mortality, 17 percent to apparent malnutrition, and 4 percent to unknown causes. Known adUlt mortality was: 12 percent to hunting mortality and 3 percent to apparent malnutrition. Telemetry accuracy tests indicated that elk were located within 20.5 ± 4.8 ha. During summer elk habitat use was predominately (59.8-9%) in the krumholz types and spruce-fir types (34.8%). Winter range habitat use was dominated by the coniferous tree (22.7%), ponderosa pine-shrub (30.9%), and wet meadow (21.0%) types. Intensity of feeding activity declined from November through February while resting activity increased. ��-223- ELK POPULATION AND ECOLOGICAL STUDIES George D. Bear and *Ronald A. Green P. N. OBJECTIVES 1. Develop techniques population levels. to more accurately 2. Define natality 3. Determine seasonal movements, daily activity patterns, and habitat preferences of elk in Rocky Mountain National Park and adjacent seasonal ranges. and mortality problems and precisely of selected estimate elk elk populations. SEGMENT OBJECTIVES 1. Capture 2. Trap and mark up to 200 elk on the winter 3. Monitor telemetry collared elk to determine mortality rates and probable causes, seasonal and daily activities, and habitat selection and preferences. 4. Conduct and mark 20 elk calves with mortality aerial censuses and production METHODS Trapping collars. range. surveys. AND MATERIALS and Marking Winter Range Elk were captured with portable elk-sized Clover Traps. Six trap sites were located on the winter range within Rcoky Mountain National Park, and two trap sites were outside the Park (Figure 1). The trap sites and the respectiv:e number of traps at each.location were as follows: Beaver Meadows Entrance Station (3), Morraine Park (5), Hallowell Park (2), Beaver MeadowsWest (4), Little Horseshoe Park (5), Horseshoe Park (2), McGregor Ranch (5), and Crocker Ranch (5). Traps were baited with 50 lb block of livestock salt and alfalfa hay. * Ronald A. Green is a graduate student enrolled at Colorado working under contract to DOW to study elk activity patterns to habitat selection; Habitat Selection and Activity Section State University and relate them of this report. �Creek 1....-, L..._, I Th. -!~rNeed MeCir.·gar 1:1 les I Mt. I r-:National 'h' ~' o "'or, •• h • fJ/J;. -, '----, ..•..•• L.. ---, ..__ r: .bLk~ "'·'''...Castle ~-~,,'1 Mt. I I I N N L_ --, .l:- r;""-....J I I "'"- .•..••..•..••. T'~ , Pros pee t s:»: ..~,,, Mt "'< Gia n t trae k ""'_.,' Mt. -"" Rams'" '~Horn "v Figure 1. _J ••• .J I ••,I". Deer A.••.' Mt. - __.r Elk trapping stations and the number of traps at each station. . �-225- During the last work segment elk captured in the traps first were tranquilized before handling; however, in this segment's trapping the elk were physically restrained for handling. This latter technique requires a minimum of five persons in the team. One person restrained the head of the elk by encircling the neck with his arm and blindfolding the elk, while the other team members secured the elk's feet with half-inch ropes. The elk's feet are pulled to one side of the trap and the animal is laid on his side for processing. It is believed this latter technique of handling elk was bette~ than tranquilizing the animal, because the elk was processed and released, much more quickly (approximately 5 minutes versus 30-45 minutes for tranquilized elk). Therefore elk are cleared from the traps in a shorter period of time. However, more persons are needed to physically restrain the animals. Each elk was marked with large (2 inch) plastic livestock eartags. The tags were orange with black numerals on them for individual identification. Duplicate tags were placed in each ear of each elk. Eight elk (4 cows and 4 bulls) were fitted with telemetry activity to study elk activities as they relate to habitat selection. collars In cooperation with Dr. Robert McLean (Center for Disease Control, Fort Collins, Colorado) blood and ticks were collected from the captured elk when feasible. Blood samples were extracted from the jugular vein, using a hypodermic syringe and needle. Blood samples and ticks are being anzlyed for incidence of Colorado tick fever and several other viruses. The blood samples are also being analyzed for brucellosis, leptospirosis, vibriosis, and black tongue. Marking of Elk Calves Elk calves were located and captured either by searching calving grounds on foot or from a helicopter. Calves during the first few days after birth generally avoided danger by lying down and remaining motionless. Then as they got older they tended to jump up and run if they suspected they had been detected. Therefore, only calves 1-3 days old could be caught by a person on foot. By using a helicopter a researcher could capture calves until they were 20-25 days old. When a calf was first observed the pilot separated it from the cow or other adults. Then the helicopter hovered 20-30 feet directly over the calf, this would generally stimulate it to try to hide. Once the calf laid down, the helicopter would land nearby permitting a person(s) to get out. With older calves it was necessary for the helicopter to return to a position above the calf to divert its attention away from a capturer until he could get close enough to catch the calf. The calf was blindfolded and placed in a large cotton bag. It was sexed, weighed (to the nearest pound), and the following measurements were recorded (to the nearest centimeter): total body length, shoulder height, and girth. Age of the calf was estimated based on hoof coloration, total body coordination, body size, and other characteristics of general physical appearance. A telemetry collar was then placed on its neck. Calf telemetry collars consisted of the telemetry unit made by Telonics attached to a two-inch (5 cm) collar. The collar was made from a two-inch (5 cm) cotton firehose with the telemetry unit attached to it with pop-rivets, and the ribbon antennae inserted inside the hose for protection. Overall �-226- diameter of the collar was reduced by folding the collar and fastening each fold with elastic, thus forming an expanding collar that would expand as the calf grew. Each of these collars contained a mortality sensor as in the telemetry units placed on larger animals trapped last winter. Normal pulse rate is 550 ms, and 850 ms in the mortality mode. The mortality switch is activated if the elk does not move its head (or collar) within a five-hour period. Each calf was located and moving. Locating Collared 3-4 times a week to determine Elk for Distribution if it was still alive and Mortality Radio-collared elk were located 1-3 times a week (depending on accessibility) for the distribution and mortality segments of this study. Majority of the monitoring work was done from a vehicle while the elk were on the winter range; foot travel or aircraft surveys were employed more extensively as the elk migrated to summer ranges. A monopole whip antennae was used when searching an area with a vehicle or aircraft. Once a signal was received, one of the more directional antennaes (loop H-beam, 3-element beam) was used to locate the animal. The latter antennaes were more accurate, but also larger; thus, the type of antennae used depended on the situation or condition of use. The loop was used in the cockpit of aircraft. The H-beam was used when backpacking into remote areas was required. The 3-e1ementbeam was used from the vehicle or on short excursions when locating a dead animal. Habitat Triangulation Selection and Activity Accuracy The technique of triangulation was used to locate a radio-collared elk. This procedure involved the determination of the angular bearings from 2 different preselected points to a radio-collared elk. A 7-element yagi directional antenna was used to obtain the reading. An elk's location was then estimated by plotting the location of the intersection point of these 2 bearing lines on an aerial photo. Preliminary tests indicated that the error in the location of a radio-collar was approximately 12 ha (Bear and Green 1979). Additional testing was continued and completed in February 1980 using identical methods used in the preliminary test. Sampling Design A stratified random sampling design was used to obtain estimates of both elk habitat selection and activity patterns. Initially 28 elk were radio-collared during the months January-March 1979. An additional 14 elk were fixed with radio collars during January and February 1980. Eleven of the collars have activity switches (Bear and Green 1979). A small number (4-10) of these elk were then sampled each month. The major criterion for selecting a particular elk was the.ir accessibility to �-227- monitoring from roadways. An attempt was made to include as many different elk with activity collars in the sample as possible. An effort was made to sample the same individual elk each month, however changes in spatial location by different individual elk throughout the course of the year made this nearly impossible. Initially 3 days each month were selected as the sampling time each month. Again changes in spatial locations of different radiocollared elk relative to each other required that during some months more days be sampled to monitor the same number of elk. When the elk remained in relatively large groups, 3-5 elk could be monitored during the same sampling day. During those times when the elk were splitting into smaller groups or were widely distributed over their range (spring and summer) often only I or 2 elk could be monitored during the same sampling day, thence requiring more sampling days to sample a given number of elk. The days sampled each month were chosen by a systematic random draw. Those days were scheduling conflicts prevented fieldwork were thrown out and a randon draw of the remaining days was made. Each day sampled was stratified into 4 sampling intervals: 1) 2 hours before and after sunrise; 2) 2 hours before and after sunset; 3) the daylight hours between the sunrise and sunset periods and 4) the night hours between the sunset and sunrise periods. The principal advantages of this design are: 1) breaks a 24-hour day into workable parts for one person but still allows estimates of habitat selection and activity for a 24-hour period. 2) Separate estimates of habitat selection and activity patterns be obtained for each stratum without additional sampling. 3) Still permits comparisons of habitat during different times of the day. selection and activity can. patterns Data Analysis Both habitat selection and activity data will be analyzed using a east squares analysis of variance to partition out differences due to individual elk, sex, time of day, months and interactions of these factors. This will allow independent measures of the relationships between habitat selection or activity and each factor without the confounding effects of the other factors. Analysis of the relationships between weather conditions (wind, temperature) and activity or habitat selection will be done with regression analysis with weather conditions as the independent variable. Both wind speed and temperature will be classified into 5 unit increments for analysis. Population Surveys Population surveys on the summer ranges were limited primarily to open alpine basins and the large grassy or willow parks along the streams in the spruce-fir forest. Helicopter surveys covered the entire summer range in Rocky Mountain National Park; and were conducted in the early morning �-228- hours. The following data were recorded for each elk observed; location, sex, and age (adult or calf). The ground surveys were less extensive and were limited primarily to the large alpine basins. Also, during ground surveys an effort was made to classify the number of yearling elk (cows and bulls) observed, and identify marked (eartagged or collared) individuals. Winter range of elk population in the Estes Valley were determined by monitoring movements of telemetry-collared and eartagged animals during this and previous segments. The entire winter range was covered during aerial surveys (Figure 2). These surveys were conducted with a helicopter during the early morning hours. The survey was divided into workable segments using natural land features, such as rivers, ridges or roads; then strip-surveyed while flying at approximately 300-500 feet above the land surface. Elk were classified only as being marked with eartags or unmarked. They were tallied only when it could be positively determined that the elk was or was not eartagged. Survey work was scheduled to be conducted at weekly intervals immediately following the trapping and marking program; however, poor weather conditions and unavailability of helicopters precluded this prescibed sampling scheme and flights were made whenever possible. The following formula population estimate: from Seber (1973) was used to compute a weighed (n N = N n = l n2= m2= l + 1) (n 2 + 1) m2 + 1 estimated population total number of elk marked number of elk observed in survey number of marked elk observed in survey The following formula was used to compute variance or standard error among counts. (n - m ) (n - m ) l 2 2 2 V (N) = (N + 1) (m + 1) (m + 2) 2 2 RESULTS AND DISCUSSION Population Distribution Five elk marked with telemetry collars on Green Ridge, near Masonville during the last work segment summered in the northern section of Rocky Mountain National Park (Bear and Green 1979). Then in the fall (September-October) they returned to Green Ridge. During the October elk hunting season they moved to the Storm Mountain Area. Two of these collared elk were harvested by hunters in November; one in the Storm Mountain area and the other near Estes Park (Devil's Gulch). Another collar ceased to function, thus that �I N N 1.0 I North PARK Corter lake . . ~ . . N ~cale: , , Figure 2. " , Portion , "" of elk winter range (shaded area) surveyed during census flights. f I in. = I mi. �-230- elk is unaccounted for. The remaining two elk returned to the Green Ridge winter area until June, 1980 when they again moved westward into the Storm Mountain area. The four radio-collared elk marked previously at Buttonrock Reservoir (Bear and Green 1979) migrated to a summer range in the southern portion of Rocky Mountain National Park (Bear and Green 1979). One collar ceased to function. The other three elk migrated back to the Big Elk Meadows area last fall, then to the B~ttonrock Reservoir area later in the winter. The cow wearing the non-functional collar was harvested by a hunter during December. The other elk returned to the alpine range in Rocky Mountain National Park in June, 1980. Bear and Green (1979) reported that the 28 elk collared on the winter range in Rocky Mountain National Park moved between winter concentration sites in Morraine Park, Beaver Meadows, and Horseshoe Park (Bear and Green 1979). In late May and early June they migrated to summer ranges along the Colorado River drainage, and alpine ranges at the head of ForeRt Canyon, Poudre River-Specimen Mt. and Roaring River areas (Fig. Appendix A). In the fall (September-October) these migration patterns were reversed, and the elk returned to winter ranges in Horseshoe Park, Beaver Meadows, and Morraine Park. There was some movement of elk eastward along Fall River into the Estes Valley with the first heavy snow storms in November, however, they generally moved westerly back into the Park as the snows melted on the south-facing slopes of Fall River canyon, and hunting pressure outside the Park increased. Then in late December-January-February heavy snow accumulation caused a large portion of the elk in Fall River drainage to move out into the Estes Valley and onto Crosier Mountain, as far east as the Drake Fish Hatchery. There was a large concentration of elk in Morraine Park and the extreme easterly portion of Beaver Meadows and Deer Mountain. Very few elk were found around the trapping stations in west Beaver Meadows, Little Horseshoe, and Horseshoe Parks in late January and early February. It was April and early May before elk wintering in easterly portions of Estes Valley and Crosier Mountain moved back to the winter ranges in Rocky Mountain Park. By late May and early June the radio collared elk were moving to alpine ranges along the Continental Divide and along the Colorado River. Four telemetry collars had been placed on elk trapped on the Crocker Ranch in February, 1980. In May and June three of these marked elk migrated south to the Twin Sisters area while the fourth migrated to the Colorado River area. Several elk ear-tagged on the Crocker Ranch were observed in Horseshoe Park during June. Population Estimation One hundred ninety-eight elk (91 cows, 65 calves, and 42 bulls) were captured and marked during the period December 18, 1979 - February 27, 1980. Six of these elk were recaptures marked during the previous winter. Another six recaptured elk had been tagged as calves the previous spring. In addition 35 elk first tagged during the 1979-80 winter were captured more than one time. The overall trapping success was 86%. Six elk (5 mature bulls and one female calf) died during the trapping operation. �-231- Four population surveys were flown during March and April (Table 1). Flight conditions on all flights were good with a poor background (spotty snow cover). Efforts to obtain a helicopter when the ground had a solid snow cover proved fruitless. The weighted average population estimate for the four flights was 1575 (± 66) elk. Table 1. Aerial elk surveys on the Estes Valley winter Total Counted Date Number Marked Elk Observed grounds. Population Estimate March 20* 395 57 1,344 March 27 602 71 1,649 April 5* 596 70 1,656 April 9 799 95 1,641 * Weather prohibited surveying the alpine segment of census area. Two ground-surveys were attempted in between the aerial surveys. A total of 491 elk were observed March 24-25 with 63 tagged animals for annual overall population estimate of 1,528 elk. On April 1, 453 elk were observed with 41 tagged animals for an overall population estimate of 2,166 elk. Area covered on the ground was limited to the large open areas and roadsides. Therefore, the data seemed to be more variable than the data collected on the aerial surveys. However,. these sample sizes are extremely small. The orange ear-tags were very visible. Comparison between the number of tagged elk observed in the larger herds in Beaver Meadows and Morraine Park was nearly identical for two aerial and two ground tallies; however, the total number of elk counted on the aerial surveys (198 and 201 animals) was 16% lower than from the ground (234 and 246 animals). Since these surveys were not made at the same time, and the sample sizes are very small, they are likely to be valid only for general comparisons. Aerial counts on the smaller group appeared to be very accurate; discrepancies between the aerial and ground counts were apparent only for those elk in very large herds (200-250 elk). Mortality Mortality data is limited to information on telemetry collars elk and incidental observations on the winter range. Twenty-five calves were captured and collared in the spring of 1979, however, one collar was non-functional, so 24 were followed throughout the year. Three of the calves were killed by predators (coyote, bobcat, and mountain lion); three were harvested by hunters in the fall, four lost their collars when captured in the elk traps, three pulled their collars off in livestock fences, four died of what appeared to be malnutrition in late winter, one died of unknown causes, and six are still active (Table 2). �-232- Table 2. Status of elk calves radio-collared in 1979. Date Collared Capture Location 610 6-18 Little Horseshoe Park Still active - moved to summer range on Specimen Mountain then back to winter range Horseshoe Park - Beaver Meadows 620 6-22 Headwaters of Poudre River Remained on Poudre River drainage for summer, then moved to winter range on Crocker Ranch, 2-26 collar came off in trap, put on eartags 11192 630 6-28 Specimen Mountain Remained in Specimen Mountain area for the summer, no signal since 10-22, May 1980 on Holzworth Homestead upper Colorado River 660 6-28 Ypsilon 670 6-26 Headwaters of Poudre River Remained in Poudre River area for the summer, then to winter range on Devil's Gulch-McGregor Ranch; killed by lion in Black Canyon, March 680 6-4 Beaver Meadows Killed 6-8 in Horseshoe bobcat 680 6-15 Beaver Meadows Killed 6-25 in Beaver Meadows coyote 680 6-28 Mummy Mountain Remained in Mummy Mountain area for summer, then on winter range in Horseshoe Park-Deer Mountain, mortality signal on Green Ridge June, 1980 690 6-28 Specimen Mountain Remained in Specimen Mountain area for summer, then to winter range in Horseshoe Park-Beaver Meadows, died of malnutrition on Deer Mountain, April 700 6-26 Headwaters of Poudre River Remained in Poudre River-Forest Canyon area for summer; then wintered in Horseshoe Park-Beaver Meadows area; shot by hunter in Dry Gulch area, December Collar Number Remarks L. Remained in Ypsilon Lake basin for summer, then on Horseshoe Park winter range, pulled collar off in fence on Devil's Gulch, April Park by by -------------------------------------------------------------------------- �-233- Table 2. Status of elk calves radio-collared in 1979. (Continued). Date Collared Capture Location 710 6-28 Specimen Mountain Still active - remained in Specimen Mountain area for summer; then to Crosier Mountain in winter 720 6-28 Headwaters of Poudre River Remained in Poudre River area for summer; then Horseshoe Park - Beaver Meadows winter range; collar came off in trap, Horsehsoe Park 1-17; eartag #85 now 730 6-26 Head of Forest Canyon Remained in Forest Canyon for summer; then on winter range in Beaver Meadows; lost collar in fence east of "1ary's Lake, April 740 6-19 Beaver Meadows Remained in Forest Canyon (hean-end) for summer; then on winter range in Beaver Meadows; May 1, Hallowe.11 Park, suffering from malnutrition-shot by Park Ranger 750 6-22 Specimen Mountain Remained in Specimen Mountain area for summer; went to Ho~seshoe Park winter range; shot by hunte~ December 760 6-26 Specimen Mountain Remained in Poudre-Forest Canyon area for the summer; went to Dry Gulch Crocker Ranch winter range; collar came off in trap, Crocker Ranch, 2-27; now eartag 11195 770 6-6 Beaver Meadows Remained in Forest Canyon area (head-end) for summer; no signal since 10-26 Little Horseshoe Park Remained in Forest Canyon area (head-end) for summer; went to Devil's Gulch winter range; lost collar in fence on Devil's Gulch, April Collar Number 780 Remarks 790 6-28 Headwaters Poudre River Still active - remained in Poudre River area for summer; went to winter range in Horseshoe Park - Beavers Meadows Crosier Mountain 800 6-26 Specimen Mountain Still Active - Lost contact for summer; found on Bighorn Mountain in fall; then moved to winter range on NE Crosier Mountain -------------------------------------------~--------------------------------- �-234- Table 2. Status of elk calves radio-collared in 1979. (Continued). Date Collared Capture Location 810 6-26 Specimen Mountain Still active - remained in Poudre River area for summer; went to winter range in Horseshoe Park NE Crosier Mountain 820 6-28 Head of Forest Canyon Remained on summer range in Forest Canyon; went to winter range in Beaver Meadows; collar came off in trap, January, Beaver Meadows; now eartag #64, found dead by Park Headquarters, June, 1980 830 6-28 Flatiron Mountain (NE side) Remained in area at head of Hague Creek during summer; in fall moved to headwaters of North Fork Big Thompson River; shot by hunter, west of Drake, 'October 840 5-31 Beaver Meadows Remained in Iceburg Pass area (Fall River during summer; moved to winter range in Beaver Meadows; collar removed in trap 1-29, Little Horseshoe now eartag If 140 Collar Number Remarks Aerial and ground surveys indicate a low calf:cow ratio for elk in Rocky Mountain National Park. However, there is a discrepancy between data collected on the two surveys (Tables 4 and 5). The ground-surveys combined for July and August indicated a ratio of 36 calves: 100 adult cows. When yearling cows are combined with adult cows (as on aerial surveys) the over-all ratio is 31: 100 (Table 5). The calf:cow ratio for the aerial survey was 42: 100 (Table 4). There isn't any apparent reason for this discrepancy, except differences in sample sizes or misclassification on the aerial surveys. The ratio of yearlings:cows is only slightly lower than the calf:cow ratios; which may be an indication of high calf survival after the first few weeks of life. Of the 34 adult elk with active radio collars four were harvested by hunters and one died of malnutrition in early spring. This latter elk was first ear tagged as an adult cow in 1965, than she was fitted with a radio-collar last year. The incidental elk mortalities documented on the winter range in late winter and early spring indicated heavy losses in this calf age class (Table 3). Of the thirteen recorded mortalities eleven were calves and two adult cows. Starvation or malnutrition appeared to be the major cause of these mortalities. �-235- Table 3. Known elk mortality during this work segment, July 1, 1979June 30, 1980. Animal Date Remarks Adult Cow October RC #350, shot by hunter near Grand Lake Adult Cow November RC #900, shot by hunter near Estes Park (Devil's Gulch) Adult Cow November RC #750 shot by hunter on Storm Hountain (Seam Rock) Adult Cow December RC 11280, shot by Hunter Parachute Hill Calf December RC #75, shot by hunter, location unknown Calf February 11 ET #105, Fran Marcoux found lying along side a fence (Ranger Road, NE Deer Mt., Estes Park) and collected it; it was very weak, pelage rough, back bone very prominent; necropsy; extensive hemorrahaging subcutaneously in upper rt. foreleg, no fat on mesantary or kidney Calf March 1 ET #158, Fran Marcoux found it near Mary's Lake; broken leg, destroyed it Calf March 14 VanSlyke, Park Ranger, reported killed by coyotes, Morraine Park, RMNP Calf March 26 RC #67, found in Black Canyon (RMNP); remains buried on ledge in cliff; appeared to be lion Calf April 3 Wagner, Park Ranger, reported killed by dogs in campground ~ mile east of RMNP headquarters Calf April 15 RC #69, dead 1 mile NW of RMNP headquarters, bone marrow was red and gelatinous, prob. malnutrition Calf April 28May 2 ET #92 Park Ranger reported; no cause of death Calf April 28May 2 RC #74, Park Ranger found by road in Hallowell Park; very weak and poor condition thus destroyed it; bone marrow was red and gelatinous; prob. malnutrition Calf April 29 Park Ranger found by Eagle Cliff; bone marrow red and gelatinous, poor body condition; probably malnutrition Calf April 28May 2 ET #144, reported by Park Rangers; no cause of death �-236- Table 3. Known elk mortality during this work segment, July 1, 1979June 30, 1980. (Continued). Animal Date Remarks Adult Cow May 2 RC #100, ~ mile east of RMNP Headquarters, Wagner, Park Ranger, reported it was in poor condition, prob. malnutrition; this was a very old cow (approx. 18-20 year old), first tagged in 1965 Adult Cow Hay 2 ET #46 McLoren, Park Ranger, found dead Calf June ET #64, dead on ridge ~ mile north of Park Headquarters, dead for several weeks Table 4. Month July Ground-classification of elk during July..,..August, 1979. Location Bulls Hague Creek 41 12 3 6 Poudre River 32 12 5 2 Specimen Mountain 62 25 29 1 7 Mummy Mt. 74 25 21 6 5 Ypsilon Basin 43 12 9 2 5 257 86 67 17 22 Hague Creek 15 7 5 4 10 Poudre River 13 3 2 Specimen Mountain 118 53 38 9 50 Chapping Creek 40 17 6 7 12 186 73 46 16 85 TOTAL Calves/lOa cows:33 Yearlings/lOa cows:33 August Cows Elk Observed Yearlings Bulls Calves Cows 5 10 Mummy Mountain* Ypsilon Basin* TOTAL Calves/adult 100 cows: 39:100 Yearlings/adult 100 cows: 33:100 *Elk were down in the timber. Calves/lOa cows: Bulls/lOa cows: 31:100 37:100 �-237- Table 5. Helicopter survey of elk on the summer range in Rocky Mountain National Park, September 9, 1979. Location Total Cows 1 17 7 (1) 1 1 48 21 (1) 7 14 6 30 21 (3) 7 1 1 1 12 8 Fall River 26 Fall River Forest Canyon 14 6 (1) 5 1 14 7 4 3 19 10 (1) 5 1 3 23 10 (1) 3 9 1 4 2 (1) 1 (1) 1 10 4 4 2 15 8 (1) 5 2 5 3 1 8 (1) 2 1 26 7 8 12 6 1 5 16 10 5 1 8 3 3 2 42 16 5 21 15 8 5 2 15 8 5 (1) 2 5 2 15 10 9 Wild Basin 11 Mummy Mt. 38 Dunraven Chapin Creek 11 11 Long Meadow Hague Creek 2 5 13 13 5 5 11 11 6 6 29 20 9 (2) 4 4 Poudre River Bulls Xature Yrlg. 3 Stormy Park Mt Ida Calves 29 18 10 1 20 13 4 3 13 16 9 (1) 10 4 4 2 --------------------------------------------------------------------------- 1 �-238- Table 5. Helicopter survey of elk on the summer range in Rocky Mountain National Park, September 9, 1979. (Continued). Location Cows Total Specimen Mountain Calves 21 (1) 38 Bulls Mature Yrlg. 13 3 1 Willow Creek 8 5 2 2 1 Specimen Mountain 8 3 1 3 1 127 65 19 (1) 40 3 739 TOTAL Calves/100 cows:42:100 373 187 158 Bull/IOO cows:56:100 21 Blood and ticks were collected from 102 of the elk captured in the traps. Analyses of these samples are pending. Twenty-nine calves (16 females, 13 males) were captured in the spring of 1980 (Table 6). Thirteen of these were captured during ground-searches and 16 were captured with the aid of a helicopter. Four of the calves died shortly after tagging, they were frozen and transported to the CSU Diagnostic Laboratory at Fort Collins for necropsy. Table 6. 1980. Elk calves collared in Rocky Mountain National Park, May-July, Collar Number Date Collared 01 02 03 05 06 * 07 07 08 * 09 09 10 ~'c*11 11 12 * 13 13 14 15 16 * 17 17 5-27 6-2 6-3 6-3 6-4 6-4 6-18 6-5 6-10 7-1 6-10 6-12 7-1 6-12 6-13 7-1 6-19 6-19 6-19 6-19 7-1 Sex F F F M F M M F F M F M F M M F F F F M F Weight (lbs) 47 44 32 38 35 37 34 ~·7 35 86 52 45 54 58 30 58 47 56 45 43 68 Shoulder Ht (cm) 66 70 72 70 71 70 70 73 73 77 75 74 83 79 68 77 76 79 71 72 73 Girth (cm) 62 66 62 63 61 61 60 66 61 86 76 65 71 72 59 72 68 69 62 64 72 Total Estimated Lg. (cm) Age (days) 103 103 93 95 102 96 99 112 103 127 122 108 116 126 102 113 113 124 106 105 133 5 7 3 3 2 3 2 7 3 20+ 14 5-7 14+ 10-14+ 1 14+ 7-10 10-14 5 5 14-20 �-239- 4) Although the method has limitations and has been evaluated as such, it is my opinion the method represents a very usable technique that exceeds any other existing technique currently available to obtain the same type of information. 5) The potential of the telemetry equipment to be used in such a technique has largely yet to be seen and offers an attractive area for future developments. Habitat Selection - May 1979-April Summer Range - July - October 1980 1979 Estimates of habitat selection were obtained for each of the 4 months that the elk occupied their summer ranges. Due to the ease and accessibility to monitoring of the radio-collared elk in Forest Canyon, all estimates of summer habitat selection were made in this region. Thekrummholz type (KRHZ) was the most widely utilized habitat type of the 4 types present (Table 7). Not until late summer and early fall (October) did use of the krummholz region decline substantially. As a general trend, habitat use appeared to shift slightly from the higher elevation types to greater use of the lower elevational types (SPFR and WLPK) as summer advanced. Elk use of the krumholz type was greatest during the month of July (69.2%) while usage of spruce-fir types was the lowest in July. The only usage of the alpine type (4.1%) during the summer months was also recorded during July. Although this may represent an under estimation of total alpine usage by elk in Rocky Mountain National Park, it may represent a very realistic estimation of alpine use in the Forest Canyon region since alpine types comprise a very small portion of that particular summer range. Elk were visually observed using alpine types in other parts of the summer range (Specimen Mountain and Trail Ridge Road) as late as October. Willowpark types (WLPK) comprised a relatively small proportion of the total habitat usage on the summer range. Like the alpine type, the willow-park type also occupied relatively small proportion of the total summer range. Considerable differences were found in habitat selection between different times of the day (Tables 8-11). Habitat use during the daylight hours was divided nearly in half between the k.rummhoLz and spruce-fir types with use shifting toward the spruce-fir types in the latter portion of summer (Table 8). During the evening use shifted more heavily toward the krummholz type(Table 9). Use of the krummholztypes increased through the night and remained high through the morning hours (Tables 10 and 11). As the summer months waned, the general trend was for the use of the krummholz types to decline and the use of spruce-fir areas to increase during all periods of the day. Winter Range: May-June 1979, November 1979-April 1980 habitat use on the winter range was dominated by 3 types: 1) coniferous (CONF) (22.7%), 2) ponderosa-pine shrub (PPSH) (30.9%) and 3) wet. meadow (WTMP) (21.0%) (Table 12). Use of the other 4 types contributed less than 10% each to the total habitat usage. No definitive month to month trend in the use of various habitat types could be demonstrated. It appeared that the elk were in general, selecting habitat types relatively independent of months. The situation appeared the same even when the data were summarized by month for different times of the day (Table 13-16). �-240- Table 6. 1980. Elk calves collared Collar Number Date Collared Sex 18 19 20 22 62 65 66 76 6-24 6-27 6-24 6-24 6-24 6-24 7-1 7-1 M M F M M F M F in Rocky Mountain Weight (lbs) National Shoulder Ht (cm) Girth (cm) 88 80 80 76 76 80 94 76 69 72 63 68 63 69 91 69 56 54 40 50 46 63 85 55 Park, May-July, Total Lg. (cm) 127 118 109 108 100 115 138 120 Estimated Age (days) 10-14 14+ 5 10-14 5-7 20 20+ 10-14 * Calf died - collar reused **Calf lost collar - collar reused Habitat Triangulation Selection and Activity Accuracy An additional 64 radio collars locations were sampled in February to estimate the error in locating a radio-collared elk using triangulation methods. This brought the total sample size to 74 locations, including the 10 sample locations from the previous year. The estimated mean location error with 95% confidence limits was 20.5 (± 4.8) ha. This represents an increase in error of 8.6 ha over that previously reported for the preliminary testing in April 1979 (Bear and Green 1979). Although 20.5 ha represents a relatively large error, it may be somewhat deceiving though in evaluating the overall accuracy of the telemetry equipment. A frequency distribution of the errors for each sample location indicated they are strongly skewed to the right suggesting that a small number of very erroneous locations may be accounting for a larger proportion of the total error. This was indeed found to be the case. Elimination of the 12 most erroneous locations (16% of the sample) reduced the mean location error from 20.5 ha to 13 ha. Thus 16% of the sample was contributing 37% of the location error. When a series of elk locations are plotted on aerial photographs, quite often largely erroneous locations can be identified and eliminated. Thus the effective mean location error may be somewhat smaller depending on a person's ability to identify grossly inaccurate locations. Conclusions drawn from the testing were: 1) Locating elk using the triangulation method does have its limitations. 2) The method as evaluated is best suited to those species their environment (habitats) relatively general. 3) Further comprehensive research is needed to identify all possible sources of error in the method and to give a more thorough evaluation of the technique to guide future equipment developments. that define �-241- Table 7. Mean elk habitat selection percentages by month for summer range, July-October 1979, Rocky Mountain National Park. Habitat Type July August KRHZl 69.2 25.4 SPFR 2 WLPK3 > ALPN4 1 4.1 > September October 65.7 68.1 36.3 59.8 33.6 28.2 52.0 34.8 1 3.7 10.7 3.8 0.0 0.0 0.0 1.0 X WLPK3 - willow park type KRHZ1 - krumholz type spruce-fir type ALPN4 - alpine type Table 8. Mean elk habitat selection percentages by month for daylight sampling period, summer range, July-October 1979, RMNP. Habitat Type July August KRHZ 55.6 SPFR WLPK ALPN > September October 49.7 49.6 37.5 48.1 43.7 50.3 48.1 61. 1 50.8 1 0.0 2.3 1.4 1.1 0.0 0.0 0.0 0.0 0.0 X Table 9. Mean elk habitat selection percentages by month for sunset sampling period, summer range, July-October 1979, RMNP. Habitat Type July August September October KRHZ 79.3 56.2 50.2 29.5 53.8 SPFR 20.7 39.8 49.8 70.5 45.2 WLKP 0.0 3.9 0.0 0.0 ALPN 0.0 0.0 0.0 0.0 X > 1 0.0 �-242- Table 10. Mean elk habitat selection percentages by month for night sampling period, summer range, July-October 1979, RMNP. Habitat Type July August September KRHZ 72.1 84.3 73.5 32.1 65.5 SPFR 8.6 15.7 17.3 42.9 21.2 WLPK 0.0 0.0 9.2 25.0 8.5 ALPN 19.3 0.0 0.0 0.0 4.8 October X Table 11. Mean elk habitat selection percentages by month for sunrise sampling period, summer range, July-October 1979, RMNP. Habitat Type July August September October KRHZ 95.9 84.7 97.7 50.0 82.1 SPFR 4.1 15.3 2.3 43.2 16.2 WLPK 0.0 0.0 0.0 6.9 1.7 ALPN 0.0 0.0 0.0 0.0 0.0 X �Table 12. Mean elk habitat selection percentages by month for winter range, May-June 1979, November 1979-April 1980, Rocky Mountain National Range. Habitat Type November December CONFl 13.9 23.6 16.2 33.9 PPSH2 49.8 20.2 50.0 PPGS3 0.0 29.3 ASPN4 1.4 WTMD5 January February March April May June 12.9 9.0 34.2 37.7 22.7 16.7 35.5 70.7 4.2 1 30.9 10.1 0.0 0.0 0.0 11.2 8.9 7.4 1.9 0.0 3.2 0.0 0.0 16.6 25.0 6.0 25.3 9.5 13.7 32.8 41.9 20.3 11.0 13.4 21.0 WILW6 6.8 3.7 6.7 13.5 8.9 0.0 12.4 9.5 7.6 GRSL7 1.4 13.8 4.4 0.0 0.0 0.0 10.3 5.3 4.4 > X I N .j::-o w 1 CONF PPSH 2 - coniferous pine - ponderosa pine-shrub PPGS3 - pon~erosa pine-grassland 4 ASPN - aspen WTMD WILW 5 6 7 GRSL - wet meadow I - grassland - willow Table 13. Mean elk habitat selection percentages by month for daylight sampling period, winter range, MayJune 1979, November 1979-April 1980, RMNP. March April May June X 76.9 34.1 14.9 64.0 60.6 52.5 57.1 16.7 60.7 85.1 0.0 1 34.0 0.0 0.0 0.0 0.0 0.0 6.8 8.5 1.9 0.0 1.7 0.0 5.1 0.0 0.6 21. 7 24.5 6.6 WTMD 0.0 0.0 4.8 0.0 0.0 0.0 0.0 2.1 WILW 12.1 0.0 1.6 1.3 5.2 0.0 0.0 0.0 2.5 GRSL 0.0 0.0 1.6 0.0 0.0 0.0 6.3 3.8 1.5 Habitat Type November December January CONF 50.0 84.4 34.9 PPSH 37.9 13.9 PPGS 0.0 ASPN February > > 1 �Table 14. Mean elk habitat selection percentages by month for sunset sampling period, winter range, MayJune 1979, November 1979-April 1980, RMNP. Habitat Type November December January February March April May June X CONF 6.9 3.0 14.3 30.0 2.1 4.2 12.5 29.9 12.9 PPSH 66.7 26.6 57.1 16.7 33.3 78.6 0.0 0.0 34.9 PPGS 0.0 0.0 10.1 0.0 0.0 0.0 20.9 27.1 7.3 ASPN 0.0 0.0 0.0 0.0 0.0 0.0 20.8 22.0 5.3 WTMD 5.5 3.7 0.0 13.3 37.5 17.3 29.2 14.7 15.1 WILW 20.8 0.0 0.0 40.0 27.1 0.0 16.7 4.2 13.6 GRSL 0.0 66.7 0.0 0.0 0.0 0.0 0.0 2.1 8.6 I N Table 15. Mean elk habitat selection percentage by month for night sampling period, winter range, May-June 1979, November 1979-April 1980, RMNP. Habitat Type November December January February March April May June X CONF 0.0 1.7 5.2 13.9 0.0 0.0 0.0 l3.5 4.3 PPSH 51.0 16.7 42.6 16.7 15.8 54.6 9.9 0.0 25.9 ·pPGS 0.0 61.9 10.4 0.0 0.0 0.0 0.0 0.0 9.0 3.1 0.0 2.8 0.0 0.0 6.7 36.9 6.2 ASPN > 1 WTMD 43.5 11. 1 25.4 50.0 82.9 45.4 23.3 22.0 37.9 WILW 1.6 4.8 11.7 16.7 1.3 0.0 26.7 25.7 11. 1 GRSL 3.4 0.0 0.0 0.0 33.7 1.9 5.6 > 1 4.8 ~ ~ I �Table 16. Mean elk habitat selection percentage by month for sunrise sampling period, winter range, May-June 1979, November 1979-April 1980, RMNP. Habitat Type November December January CONF 1.4 20.4 19.0 PPSH 47.3 12.8 PPGS 0.0 ASPN February March April May June 12.5 10.4 13.9 22.1 12.8 14.1 48.3 16.7 31.3 53.9 11. 1 0.0 27.7 11.9 4.8 0.0 0.0 0.0 27.8 2.8 5.9 6.9 1.4 0.0 4.2 0.0 0.0 11.1 14.7 4.8 WTMD 37.4 23.7 8.8 66.7 44.3 32.2 5.5 32.5 31.4 wnw 6.9 9.5 7.1 0.0 14.1 0.0 22.2 20.5 10.0 GRSL 0.0 14.2 11.9 0.0 0.0 0.0 0.0 16.7 5.3 X I N ~ VI I �-246- When winter habitat selection was averaged across the 8 months and monthly differences ignored definite changes emerged in habitat selection throughout the day. Habitat use during the daylight hours was almost exclusively restricted to the coniferous (CONF) and ponderosa-pine shrub (PPSH) types (Table 7). During the evening hours habitat use shifted primarily to the wet meadow (\VYMD). willow (WILW) and ponderosa-pine shrub. The night hours were spent primarily in the same 3 types with more use of the wet meadow (37.9%) and slightly less use of the ponderosa pine-shrub (25.9%) types. Habitat use in the morning hours was very similar to that during the night. In summary. winter habitat use by elk in Rocky Mountain National Park was dominated by the coniferous. ponderosa pine-shrub and wet-meadow types. Habitat appeared to be selected rather independently of months but there was a consistent change in the habitat types selected during different times of the day throughout the winter. Activity Patterns-May 1979-April Winter Range - November-April 1980 1980 Definite trends in activity patterns were established across both months and time of day (Figure 3-8). An inverse relationship between time spent grazing and time spent resting existed across the 8 months the elk occupied the winter range (Figure 3). Grazing activity remained relatively high at the beginning of winter (November) but slowly declined as the winter months advanced. As grazing time declined to a low of 37.5% in February. resting activity proportionally increased to 52.1%. Moving activity remained relatively the same each month. During March the trend reversed and time spent in grazing activity slowly increased with proportional declines in resting. By May the activity patterns had returned to those observed at the beginning of winter. One hypothesis to explain this relationship is that the elk shift their energy strategies over the winter from one of energy intake to one of energy conservation. Throughout the winter months. pronounced changes in activity patterns were found during different times of the day (Figure 4). The elk exhibited a highly crepuscular and nocturnal grazing behavior and a diurnal resting pattern throughout the winter (Figures 5-8). The daylight hours represented the time of least activity. Averaged across the 8 months spent on the winter range. resting comprised 61.2% of ·the elk's time during the day while only 26% of the day consisted of grazing (Figure;4). For the most part, grazing during the day was minimal throughout the winter (NovemberApril) while 65~75% of the daylight hours consisted of resting (Figure 5). Not until May and June did feeding comprise a major portion of the daylight hours. The evening hours around sunset were without a doubt the most active hours of the day. Grazi~g activity averaged nearly 82% during this time (Figure 4). Resting activity generally comprised less than 10% of the evening hours (Figure 6). Although the elk were approximately 30% less active during the night than evening. grazing and moving still comprised a greater �i: l/;t~:~] ->- ~ff:1; I~I I~ • I~I I~, t- ~~ ~~ ••••• C',,) = ••••• - I AUG SEPT I I I W////A W/////l IW////hl f<"''''';,.~,J fi~5';'iT»jf V//////,I W///////.I V/////fil ~ ~ ~ ~~ ~ ~ ~ ~ ~ A. JULY OCT GRAZING Figure 3. I N RESTING o MOVING Percent elk activity by month, May 1979-April 1980, Rocky Mountain National Park. "-I I �10.9 >- -I- > IU C ~lmlll E'iIf/; .. ;i.ili, '. ~ ~ ~ ~ I N .j:-- IZ ~~,. '- > ••• C;' ~ ~ ~ co ~ I ••••• U &II: ••••• A. SUNSET grazing NIGHT resting SUNRISE C~ 24-HOUR :'PRD moving ., Figure 4. Elk activity by day periods averaged across the 8 months spent on the winter range (November-June), RMNP. �z z ,. ,. ,. 0 ~ -t 0 ~ :=1 ~ .. ~~ ~-~1~~~~ t""S>"l .1-*::4;:*:1 n:;~,:lii~ ~l:/;~J r~:j~[;lr~~:~i~ ~ I ~ N ~ \0 I JULY AUG SEPT GRAZING Figure 5. JUN OCT RESTING D MOVING Mean elk activity percentages by months for the daylight sampling period May 1979April 1980, Rocky Mountain National Park. �z z ~ ~ 0 :. -t :. 0 !i :. r~I""A r""". ."""". ..""" ...• ....""",~ .. """,. ."""" ... .""","'. I N VI a I JULY AUG SEPT OCT GRAZING Figure 6. RESTING o JUN MOVING· Mean elk activity percentages by months for the sunset sampling period, May 1979April 1980, Rocky Mountain National Park. �~I I~ ~I I~ I N V1 •... I JULY AUG SEPT OCT GRAZING DEC NOV ".' r.' '5j~1 RESTING L."'~,,"1, ..'.'}\ l:~t~i1-:"!'L~~!:(-~J . . ~~ii ..•:«:;\~~ Figure 7. JAN FEB MAR D APR MAY JUN MOVING Mean elk activity percentages by months for the night sampling period, May 1979-April 1980, Rocky Mountain National Park. �> -- (///////A I\";:·-"~';' ~A.l:l ,~I ~. - I~ •••• > '=' 1=,. :DI -t -t ~~ ~ I I AUG SEPT :DI, 1///////. 1.//////.11 V///////6 li·",.~tl".")!1 I////////JI V///////I ••• ////////1 •.•///////.1 I I ~ ~ NOV DEC ~ E;[}:~~l ~ ~ ~j;J ~ W = U w a. JULY OCT GRAZING Figure 8. I N U'1 RESTING l] MOVING Mean elk activity percentages by months for the sunrise sampling period, May 1979April 1980, Rocky Mountain National Park. N I �-253- portion of the night hours (Figure 4). As the winter progressed, nocturnal grazing and movements slowly declined while resting activity increased but resting still never comprised more than 50% of the night hours even during February, the month of least activity (Figure 7). Movement activity during the night displayed a pronounced decreasing trend across the winter months. Movements during the night in November consisted of nearly 21% of the time. By April, movements comprised less than 4% of the night hours (Figure 7). In the latter months of winter, the elk would often move into one area during late evening or early night and remain there until morning and simply relegate their activity to either grazing or resting. Activity (grazing and moving) increased slightly in the morning but not appreciably. Grazing activity comprised 58.1% of the morning hours while 31.2% of the time was spent in resting. Although the elk often grazed intensively in the morning it was usually of much less duration than that observed in the evening. Grazing and resting in the morning showed marked changes across the winter. From November to February, grazing in the morning declined over 50% while resting increased by a similar percentage (Figure 8). In March the trend reversed and by June nearly 65% of the morning hours were spent grazing. Although definite changes in activity were found between months, a very consistent model of daily activity patterns and movements emerged over winter that did not change a great deal month to month. The daylight hours were generally spent resting in either the coniferous or denser ponderosa pineshrub habitat types with only occassional movements and grazing. During the hardest winter months it was not uncommon for the elk to rest for 4-5 hours without moving. In late afternoon or early evening, the elk would move from their resting areas to the more open habitat types (wet meadow, willow, grassland and ponderosa pine-shrub) and begin an intensive grazing period that generally lasted until several hours after sunset. As the night hours advanced, the intensive grazing shifted to a periodic grazerest cycle which generally lasted until early morning. During this time the elk would remain in the open habitat types with grazing and resting occurring together in the same areas. During the night the elk would quite often use areas near the roads without hesitation for both grazing and resting. Through the morning, the periodic graze-rest cycle shifted to a short intensive grazing period which coincided with movements toward either the coniferous or denser ponderosa pine-shrub types. Once into these types, the daylight resting period started again. The only real monthly differences observed in this daily pattern was the absolute time spent in each activity during each of these time spans. An increase or decrease in the average daily time spent in a particular activity usually corresponded to an increase or decrease in that activity during all times of the day. These changes in average daily activities were not attributable only to large changes during one particular time of day but rather similar changes in activities throughout a 24-hour period. Summer Range:July-October 1979 Activity patterns on the summer ranges were very similar to those observed during May and June. Grazing remained the dominate activity (Figure 3). �-254- Differences in activities with respect to time of day were much less pronounced than those on the winter range. Grazing was much more evenly distributed throughout the day (Figures 5-8). Only small differences were found in activities between the evening, night and morning hours. Resting was greatest through the daylight hours but still only averaged 36.5% during this time. Logistical problems prevented estimation of activity patterns for September and October so no indication of activity patterns trends across the summer could be obtained. LITERATURE CITED Bear, G. D., and R. A. Green. 1979. Elk population and ecological studies. Colo. Div. Wildl. Fed. Aid W-126-R-2, Work Plan 3, Job 2. Prog. Rept., Game Res. Rept. July 1979, pp. 373-402. Clover, M. R. 1956. 42: 199-201. Seber, G. A. F. parameters. Prepared by Single gate trap for deer. Calif. Fish and Game. 1973. The estimation of animal abundance Hafner Press. New York, NY 506pp. 7/ ::.,.;;=~~_ ~. <Y Georg€:Bear Wildlife Researcher Ronald A. Green Graduate Student ~t1./~' C and related �-255- APPENDIX A �lIllie -, I Y.)IOws~ f"a,n.:hlld -!' \, ~" . =. \ •• oIJ'\ . 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'. \.": ' " ..,~ I �-303- APPENDIX B �-304- ELK TRAPPING ESTES PARK VALLEY Summary Duration: December 18, 1979 - February 27, 1980 Trap Sites: Location 1. 2. 3. 4. 5. 6. 7. 8. No. of Clover Traps Beaver Meadows Entrance Station (BME): at jct. of Bear Lake Rd. and Deer Ridge Rd., north side of road in trees 3 Morraine Park (MP): in Morraine Park Campground on Loop A, meadow area near sf.te 92 5 Hollowell Park (HoI): on bench 100 yds north of turn-around 2 Beaver Meadows (BMW): approximately 100-400 yds west of turn-around at west end, in timber 4 Little Horseshoe Park (LHS): on ridge 100 yds NE of barn and park at base of that ridge, approximately ~ mile NE of barn 5 Horseshoe Park (HS): on bench 50 yds north of road at east end 2 McGregor Ranch (McG): in mouth of Black Canyon, along creek bottom, 100 yds west of Park Boundary 5 Crocker Ranch (Cr): just east of yellow house along edge of meadow (southeast base of Mt. Olympus) 5 Marking of elk: Orange eartags (maxi Al f kex livestock plastic tags) with black numerals. A few radio-collars were placed on elk - green or white in color, except one red (#600). Trapping Crew: Rocky Mountain National Park DOW George Bear, Paul Neil, Barry VanSant, Bruce Gill �-305- Trapping Crew (Continued) RMNP Dave Essex George Wagner Jack Gartner Burt McLaren Lorin Casebeer Charles Logan James Wilson Bob Seibert Jim Protto Larry Van Slyke Ron Maitland Dwight Hamil ton Bob Gutherie Doug Bueler Skip Betts Dave Stevens CDC Robert McLean McGregor - Crocker Ranches RMNP George Wagner Burt HcLar-en Jack Gartner Lorin Casebeer Charles Logan Bob Gutherie Doug Bueler I'm,} George Bear Barry VanSant Fran Marcoux Paul Neil Courtney Crawford Ron Oehlkers Frank Rinella Roger Lowry Ely Wagner Mike Babler Elk Trapped-Marked: RMNP McGregor Crocker Cows Yearling Adult Total 15 59 74 0 2 2 1 14 15 Calves Males Female Total 27 28 55 1 0 1 4 5 9 Bulls Yearling Adult Total 15 21 36 0 4 4 2 0 2 165 7 26 Totals Trapping Success: 86% Recaptures: First tagged a year ago 6 First tagged this year 35 Calves collared last spring 6 �-306- Blood Samples: Blood (B) 102 Ticks (T) Mortality: Six elk (5 large bulls, I female calf) died in the trapping operation. The calf dislocated its back therefore was destroyed. Three of the large bulls literally destroyed the traps, became entangled in the netting and twisted their heads around to where they suffocated. The other two large bulls were in very poor body condition and apparently could not withstand the stress of being caught in the trap. �-307- Elk trapped and marked in the Estes Park Valley December 18, 1979-February 27, 1980. Ear Tag Number Date Sex Age Trap Location 1 12/18 F Adult BME 2 12/18 F Calf BMW 3 12/18 M Yrlg LHS 4 12/18 F Adult LHS 5 12/18 F Yrlg LHS 6 12/19 F Yrlg BME 7 12/19 F Adult BME 8 12/19 M Calf MP q 12/19 F Adult MP 10 12/19 M Calf MP 11 12/19 F Adult MP 12 12/19 F Adult HoI 13 12/19 F Yrlg BMW 14 12/20 M Yrlg LHS 15 12/20 M Yrlg LHS 16 12/19 F Adult MP 17 12/20 F Yrlg LHS 18 12/21 F Adult BME 19 12/21 F Calf BME 20 12/21 F Adult BME 21 12/21 F Calf MP 22 12/21 F Yrlg MP 23 12/21 M Yrlg MP 24 12/21 F Yrlg MP 25 12/21 F Adult BMW 26 12/21 F Adult BMW 27 1/3 M Calf BME 28 1/3 M Calf BME 29 1/3 F Adult MP 30 1/3 F Calf MP 31 1/3 F Yrlg MP Yellow E.T. 1118 32 1/3 M Adult MP R.C. 1145 Remarks E.T. left only Broken antler - RC#75 Right E.T. only Yellow E.T. 116 ------------------------------------------------------------------------------------ �-308Cont'd Ear Tag Number Trap Location Date Sex Age 33 1/3 M Yrlg MP 34 1/3 F Calf HoI 35 1/3 M Calf HoI 36 1/4 M Yrlg BMW 37 1/4 M Adult BMW 38 1/9 F Yrlg LHS (B) 39 1/9 F Adult LHS (B) 40 1/9 M Adult LHS 41 1/9 M Adult LHS 42 1/9 M Yrlg LHS 43 1/10 M Calf BME 44 1/10 F Calf BME 45 1/10 M Calf BME 46 1/10 F Adult MP (B) 47 1/10 F Calf MP (B) 48 1/10 F Adult MP 49 1/10 F Adult MP (B) 50 1/10 M Adult BMW (B) 51 1/11 F Adult BME (B) 52 1/11 M Adult BME (B) 53 1/11 M Adult LHS (B) 54 1/11 F Calf LHS 55 1/15 M Adult BME 56 1/15 F Adult BME (B) 57 1/15 F Calf MP (B) 58 1/15 F Calf MP RC 1179 59 f/15 F Adult MP Yellow E.T. 1119 by trap (B) 60 1/15 F Adult LHS (B) 61 -1/15 F Adult LHS (B) 62 1/15 F Calf HS (B) 63 1/15 M Calf HS (B) 64 1/15 M Calf LHS 65 1/15 F Yrlg LHS 66 . 1/16 M Calf BME Remarks RC 1140 RC 1141 (B) (B) (B) (B) RC 1118 (B) ---------------------------------~---------------~------------------------------- �-309Cont'd Date Sex Age Trap Location 67 1/16 F Yrlg BME (B) 68 1/16 M Adult BMW (B) 69 1/16 F Adult BMW (B) 70 1/16 M Calf BMW (B) 71 1/16 M Yrlg LHS (B) 72 1/16 F Adult LHS (B) 73 1/16 M Yrlg LHS (B) 74 1/16 F Calf HS (B) 75 1/16 F Adult HS (B) ;6 1/17 M Adult BME (B) 77 1/17 F Adult BME (B) 78 1/17 F Calf MP (B) 79 1/17 M Calf MP (B) 80 1/17 F Adult BMW (B) 81 1/17 M Adult BMW (B) 82 1/17 M Calf HS (B) 83 1/17 F Yrlg HS (B) 84 1/17 F Calf HS (B) 85 1/17 M Calf HS 86 1/18 M Yrlg MP (B) 87 1/18 F Adult MP (B) 88 1/18 M Adult MP 89 1/18 M Adult BMW 90 1/18 M .Adult BMW 91 1/18 M Adult LHS 92 1/18 M Calf LHS (B) 93 1/18 F Adult LHS (B) 94 1/18 F Adult LHS (B) 95 1/22 F Calf BME 96 1/22 M Calf BME (B) 97 1/22 F Adult BME (B) 98 1/22 M Adult BMW (B) Eartag Number Remarks RC 1172 RC 11300 (B) (B) (B) Right E.T. Only (B) --------------------------------------------------------------------------------- �-310- Cont'd Eartag Number Trap Location Date Sex 99 1/22 F Adult LHS 100 1/22 F Calf LHS (B) 101 1/22 F Adult HS (B) 102 1/22 F Adult HS (B) 103 1/23 M Yrlg BME (B) 104 1/23 F Yrlg BME (B) 105 1/23 F Calf MP 106 1/23 F Adult MP (B) 107 1/23 F Adult MP (B) 108 1/23 F Adult LHS (B) 109 1/23 F Adult LHS (B) 110 1/24 F Calf BME (B) 1/24 F Calf BME 112 1/24 F Yrlg MP 113 1/24 F Calf MP 114 1/24 F Adult MP 115 1/24 F Yrlg MP 116 1/24 M Yrlg MP 117 1/24 M Adult BMW 118 1/24 M Calf LHS (T-B) 119 1/24 F Adult LHS (B) 120 1/25 F Adult BME (B) 121 1/25 F Calf BME (T-B) 122 1/25 M Calf BME (T) 123 1/25 M Calf BMW (B) 124 1/25 M Yrlg BMW (T) 125 1/25 M Adult BMW 126 1/25 F Adult MP (B) 127 1/25 M Calf MP (T-B) 128 1/25 M Adult MP (T-B) 129 1/25 F Adult MP (B) 130 1/25 F Adult MP (T-B) 131 1/25 M Adult HS 111 , Age Remarks Right E.T. Only Died 2/11 (B) (B) (T-B) RC 11120 (B) ---------------------------------------------------------------------------------- �-311- Cont'd Eartag Number Date Sex 132 1/25 F Adult LHS (T-B) 133 1/25 F Adult LHS (B) 134 1/25 F Adult LHS (T-B) 135 1/29 M Yrlg BME 136 1/29 F Adult MP 137 1/29 M Calf MP 138 1/29 F Adult MP 139 1/29 F Yrlg LHS 140 1/29 F Calf LHS 141 1/29 F Adult LHS 142 1/29 F Adult LHS Left E.T. only 143 1/30 F Adult BME R.C. #220 144 1/30 M Calf BME 145 1/30 F Calf MP 146 1/30 .F Calf MP 147 1/30 M Adult MP 148 1/30 M Yrlg LHS 149 1/30 F Adult LHS 150 1/31 M Calf BME 151 1/31 M Calf MP 152 1/31 F Adult LHS RC 1144 153 1/31 F Adult LHS RC 1142 154 1/31 F Adult HS 155 1/31 M Calf HS 156 2/7 F Adult BME 157 2/7 M Adult BME 158 2/7 F Calf MP (T-B) 159 2/7 M Calf MP (T-B) 160 2/7 F Adult LHS Left E.T. Only (T-B) 161 2/8 F Calf BME (T-B) 162 2/8 F Adult MP (B) 163 2/8 M Calf MP (T-B) 164 2/8 F Calf MP (T) 165 2/8 F Calf MP (T) Age Trap Location Remarks RC 1135 RC #3 (R) RC 1143 (T) ------------------------------------------------------------------------------------- �-312Cont'd Eartag Number Date Sex 166 2/13 M Calf Cr (T-B) 167 2/13 M Calf. MeG (T-B) 168 2/13 M Adult MeG 169 2/13 F Adult MeG (T) 170 2/13 F Adult MeG (T-B) 171 2/14 F Calf Cr (T-B) 172 2/14 F Adult Cr RC 1/900 (T-B) 173 2/14 M Adult MeG RC 11390 (B) 174 2/14 M Adult MeG Very 19. bull 175 2/15 F Adult Cr RC 11240 176 2/15 M Adult MeG 177 2/19 F Calf Cr 178 2/19 M Yrlg Cr Rt. antler broke (B) 179 2/19 F Yrlg Cr RC 11600 180 2/20 M Yrlg Cr 181 2/20 M Calf Cr 182 2/20 F' Adult Cr 183 2/20 F Calf Cr (T-B) 184 2/20 F Adult Cr (T-B) 185 2/20 F Adult Cr 186 2/20 F Adult Cr 187 2/21 F· Adult Cr RC 11210 188 2/21 F Adult Cr RC 1/310 189 2/26 F Adult Cr "(T-B) 190 2/26 F Adult Cr (T-B) 191 2/26 F Adult Cr Hole in ventral surface of snout (shot?) (T) 192 2/26 M Calf Cr RC 1/62 - Came off in trap (T-B) 193 2/27 F Adult .Cr 194 2/27 M Calf Cr (T) 195 2/27 F Calf Cr (T) 196 2/27 F Adult Cr (T-B) 197 2/27 F Calf Cr (T-B) 198 2/28 F Adult Cr (T) Age Trap Location Remarks (B) (B) (T-B) (T-B) �-313- Recaptures in Traps - 1979-80 Tags Date Eartag Number Trap Location 1/11 4 HS 1/15 17 MP 1/15 8 MP 1/15 54 LHS 1/18 64 BMW 1/18 14 LHS 1/18 Unk. LHS 1/18 Unk. LHS 45 BMW 1/24 22 LHS 1/24 8 BMW 1/24 64 MP 1/25 51 BMW 1/25 110 LHS 1/31 144 LHS 1/31 43 MP 1/31 2 MP 2/1 92 MP 2/1 79 MP 2/1 43 MP 2/1 116 LHS 2/1 61 LHS 2/1 63 LHS 2/1 99 HS 2/7 Unk. MP 2/7 27 BME 2/8 43 MP 2/8 66 MP 2/14 141 McG 2/19 166 Cr 2/20 47 McG 2/21 183 Cr 2/26 166 Cr 2/27 166 Cr 2/28 166 Cr 1/23 RC ��July, 1980 -315- JOB PROGRESS REPORT State of Colorado ----~~~~~------------ Project No. W-126-R-3 --~~~~~----------- Work Plan No. 4 ------~------------- Job No. 1 Big Game Investigations Bighorn Sheep Investigations Prescribed Burning to Improve and Enlarge Bighorn Sheep Ranges Period Covered: Personnel: July 1, 1979 through June 30, 1980 Colorado State University, U.S. Forest Service, and Larimer County Sheriffs Department personnel; Dr. N. T. Hobbs and T. N. Woodard. ABSTRACT Prescribed burning was conducted on 29-30 September 1979. problems encountered during burning are discussed. Objectives and ��-317- PRESCRIBED BURNING TO IMPROVE AND ENLARGE BIGHORN SHEEP RANGES Thomas N. Woodard P. N. OBJECTIVES 1. To test the hypothesis that -pr.escr Lbed burning of understory vegetation improves the production and quality of bighorn sheep forage in a ponderosa pine-Douglas fir timber zone. 2. To test the hypothesis that bighorn sheep use of burned areas increases compared to adjacent unburned areas. SEGMENT OBJECTIVES 1. Complete analysis of all pretreatment data including plant cover, herbage yield, forage quality, and bighorn sheep food habits and treatment preference. 2. Coordinate pre-burn, burn, and post-burn activities with U.S. Forest Service and Colorado State University Fire Science personnel. METHODS AND MATERIALS Methods and materials were described by Woodard (1979a) and Woodard (1979b). Dr. N. Thompson Hobbs, currently principal investigator for this project, will complete a revised study plan during the next segment and report changes in methodology in future reports. STUDY AREA LOCATION The study area location was described by Woodard (1979a). RESULTS AND DISCUSSION Prescribed burning was conducted on 29-30 September 1979. Appendix I is a report by Dr. Philip N. Omi, Colorado State University, Department of Forest and Wood Sciences, which discusses objectives and elucidates problems encountered during burning. LITERATURE CITED Woodard~ T. N. 1979a. Prescribed burning to improve and enlarge bighorn sheep ranges. Colo. Div. Wildl. Game Res. Rep. January: 45-67. Woodard, T. N. 1979b. Prescribed burning to improve and enlarge bighorn sheep ranges. Colo. Div. Wildl. Game Res. Rep. July, Part Two:403-418. �-318- APPENDIX I �-319- SUMMARY REPORT TO COLORADO DIVISION OF WILDLIFE PRESCRIBED FIRE FOR WILDLIFE HABITAT Philip N. Omi, Principal Investigator February 27, 1980 �-320- This report summarizes work completed in cooperation with the Colorado Division of Wildlife in the planning and implementation of a series of prescribed burns in Wintersteen Park near Rustic, Colorado. This report augments previous summaries by Barrows and Yancik (1978), written in contemplation of the burns originally scheduled for 1978, and by Omi (1979), describing revisions to the original burning plan resulting from the postponement of burning operations until fall, 1979. The objectives of the study described in this report were: 1. To determing grass fuel loadings and the total grass and woody fuel loadings at the prescribed fire sites; 2. To test prescriptions for prescribed fires and revise as necessary; 3. To determine the behavior of each prescribed fire; and 4. To determine immediate effects of prescribed fires on fuels and vegetation. The work initiated to fulfill the above objectives reached culmination with the completion of the prescribed burns on September 29-30, 1979. This report sequentially discusses the degree to which objectives were fulfilled, and elucidates some of the problems encountered. Implications for the project "Prescribed burning to improve aridenlarge bighorn sheep ranges" (Colorado Division of Wildlife Project No. W-41-R) are also discussed. Objective 1: Grass and total fuel loadings at the prescribed fire sites. The herbaceous, shrub and litter fuels were resampled in summer, 1978 for comparison with the previous year's inventory. The average loadings (tons/ac) measured in the two sample years are presented in Table 1. While subject to sampling error and measured by two different persons, the inventory figures suggest that the relatively we.t precipitation year preceding the resampling effort produced the greatest impact in the live and dead shrub fuels in the grass type (Figure 1, plots 3-1, 3-2, and 3-3). The anticipated impacts on fire behavior characteristics were not appreciably different between two years. The computerized fire models used to compare the two years focus on contributions to fire spread on the ground surface and therefore proved insensitive to the higher shrub loadings in the grass type. Objective 2: Prescription tests and revisions to burning plan. The original prescriptions for the different vegetation types were revised due to: 1) inconsistencies in fuel moisture, windspeed, and relative humidity ranges specified in the original prescriptions; and 2) potentially adverse synoptic weather conditions historically associated with the original burning period. Ultimately, the coordination between the four agencies involved (Colorado Division of Wildlife, Colorado State University, U.S. Forest Service, and Larimer County Sheriff's Department) proved as difficult as selecting the proper meteorological "window" for the desired burn. Abnormally high rainfall during July-August, 1979 delayed curing of the fine herbaceous fuels and necessitated deviation from the planned burning date. �<, =r <> -e<><><> <. <>-><> <>0 <> 0<><> o < • <><>..0-<> <> -<>I <>C>o<> <> -e-, <><> <> 0 C> <> <>1 0<> -e<> 0 <> -<> C> <>C> <> <>. <> <><> <> <> \ <> <><> 0 <> <> • <> \ <><> H~ 0<> <><><> <><><> <> C> <><> <> <><><> <> ~: C> <> <> \ C><> <> <> <. C> <> ~ ¢ <> <> <> <>. <> <> ~-{ '~-z....:...J. _ <> C> __. .--0 .--0 <> C>o <> <> <> <> <> -<> <> < <> -<> <> <><>-<><><> -< C> <> <> <>c><>c>0._· < <><><>c><>-..<>o-<> -e...•.... ., 0<> <>-c>C><><>_ <> • <> <> .<> <> .<> <> <> <> " <><><> ~ <> ~ <><>. 0 .<> c:> c:> <> <> <> c> C> <><>"1. <>c::>c><><>c:>C> .<><><>:., <><><><><><><> \ C>e>C><>' <>_<><><> <> \\<> <> <> <> '. <> <> <> <> <> <> <>. C> ~<> <> <> <>-oC> ~<> <> _. 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"::> • <> <> <> <> oC><> <><><><>0)'0'<><>00<><><><> <> <> <> <> <> pc><> <> <> <> <> <> <> ,<> <> <> .<> <> <> <> <> <> <> <> <> <>E-<> <> <> eo.<> <> _ <> <> <>. <> <> <> eo- <> .C><> <> eo- <> C> <>_<><> <><><> <> <> <> ~ -::<> <>-_ <>-c>e><> ~ <><><> <> < <> <> <> <> <> <> <> <> <> <> <> <> <> -< C> <> .<> C> <> <>-<> <> <> ' <>.<> <> <> <> <> <> <> <> <> <> <><><> <> <> <> <> .<> <> <><>< • <> _.- _. '::l.; ooz = ·U"; 1 -<><> <> -e<> <> <><> .c><><> \.0 <> <> <> '<=><>C> ~<><> • C> <> \<> <> 0-0-<><> = :iTI"JS l ~-Z N \ L-Z -1l£- �-322- Table l. Average grass and total grass and wrJody fuel Lc ad Lnga in 1977 and 1978 (tons/ac). ------- Bunchgrass ShruG P. Pine 19771/ 1978 1977U 1978 : 9 771/ Li.ve .1297 .1873 ;1017 .0632 .101 j .0610 Dead .3026 .4356 .2372 .1481 .2'~62 .1416 14.8890 8.9589 8.1928 1978 Gr a s s ---- 2/ 3.2628 Total Grass and Woody-- l/From 2/ Barrows and Y~ncik fJ.2386 9.4099 (1978). - Includes dead down woody, woody inventory estimates loads in both years). litter, dead and livE: shrub fuels (dead down from 1977 were used in computing total fuel �-323- As the time for burning approached, intense rainstorms and snow twice forced last-minute cancellations of burning operations (September 7 and September 15). The prolonged absence of key USFS personnel, while assisting with the California fire situation, forestalled another opportunity (September 22). The use of 15 CSU Fire Science students as fire behavior monitors caused some difficulties, as weekends relieved them of classroom obligations, but didn't always coincide with suitable burning conditions. At the same time, Larimer County depended upon CSU students to beef-up its weekend fire control forces. Public involvement efforts, including affected property owners and local radio and news media, were coordinated by the USFS. The USFS also developed a detailed burning plan which stressed the scientific nature of the proposed burning operations. These plans were orally presented, along with CSU's original recommendation for a two-day burning effort, in discussions and meetings on September 4 and September 12. The apparent thoroughness of our planning efforts doesn't dispel my belief that the actual implementation of our plans proved unsatisfactory. Objective 3: Fire behavior determinations. Prescribed fire monitor teams recorded fire behavior characteristics in the burn "treatment" plots of the three vegetation types presented in Figure 1. These teams monitored rates of spread, flame lengths, flame heights, soil time-temperature profiles, and energy release at the representative thermocouple locations depicted in Figure 2. The figure also identifies the location of preburn inventory transects and weather recordings. Fire behavior characteristics observed by the monitor teams are summarized in Table 2. Future research efforts at CSU will include the correlation of fire behavior measurements in each vegetation type with post-fire plant community descriptors, for comparison with the preburn vegetation and with paired unburned control plots. As expected the differential fire behaviors observed reflect the variations in fuels, weather, and topography among the different treatment plots. In addition, the starting time for burning operations was delayed by almost two hours and subsequently affected fire behavior late in the day: The incomplete burn in grass plots 3-1 and 3-2 is directly attributable to attempting to burn when the plots were nearly in late afternoon shadows. In retrospect, the ignitions of all grass plots should have been deferred until the following day, as soon as it was known that all burning could not have been completed in one day. This determination probably could have been made after firing of the ponderosa pine plots, had we not been committed to a burning plan which called for one day completion of all plots. This modification to CSU's recommended two-day burning operation resulted from the U.S. Forest Service's concerns for the availability of line-holding crew personnel. Again in retrospect, the eventual availability of a skeleton crew to finish burning operations on the second day (September 30) indicates that the commitment to completing all firing in one day perhaps was unnecessary. However, personnel limitations may have accounted for the one major control problem encountered on September 30, at approximately 2:00 PM. The extra burned area (approximated in Figure 3) occurred during the firing of shrub plot 2-2, while the tanker-truck responsible for control in the area was refilling its water tanks at the nearest water source. Preliminary analysis of weather records from the nearest weather station �-324- Table 2. Average fire behavior characteristics in the three vegetation types, Wintersteen Park prescribed burns (standard errors parenthesized). Vegetation Ponderosa type/type of fire Rate of spread (ft/min) Flame length!/ (ft) 2/ Energy Release(Kcal) pine Head fire .09 (.03) 1.83 (.33) 14.37 (1.23) Flank fire .01 (.01) 4.17 (2.92) 17.46 (7.02) .03 (.001) 4.13 (.52) 31. 98 (8.84) Head fire .11 (.03) 1.7 (.51) 7.8 (.31) Flank fire .17 (.03) Shrub Back fire Grass 1/Fireline intensity by the formula (I) may be computed I l/Derived from·water-can = 5.76 F2•174 analogs. directly no measurements from flame length (F) �F\V Figure 2. Map of stud! area depicting sampling lccations. 13 B Plots 2.tr: c<;si r.a'ts-d by If'::geta t ~on type -'12 I icat ion ;:ur:1bertreatment us i:1 "'''e':~ ~, 0' ''; ni"l cede: ~~~~; : ~,,, , ~" ""l ..-t 0 . 1r: .. ::: 200 r:. 13U~ e.g. plot 2 3 8 is the third burn replication in the shrub vece cat icn type. 23U~ B Belt vica~nr::r" k~: rl::ac;:i';s F lO-hr fJel sticks Recording ~~~:~cr :tatic~ . Ther:i.C~:';..i~12 ~C~.::~:,:~ \~: , -Sf'.Jss t r··:~:~~2::~ -·"--~;1,"ub t i"Z::i3·~':~ �N 1 SCALE: Figure 3. Approximate burned area, Wintersteen Park prescribed burning project , .FINAL BURN 1 in. = 200 ft. �-327- (located in plot 3,-2U, Figure 2) indicates the simultaneous occurrence increasing easterly windsl/as the fire jumped its control line. of Objective 4: Immediate fire effects. The post-fire inventory intended to document fuel reductions and immediate post-fire effects. Reductions in fuel loading are an indicator of total energy release in the larger fuels, provided information on rate of spread and heat of combustion are available. The completion of the burn at the end of September did not allow sufficient time for all post-fire resampling tasks. Water can analogs provide one indication of energy release, while the rate of energy release can be inferred from fiLeline intensity and flame length. These data are presented in Table 2. The inventory of post-fire fuels and burned areas and a survey of short-term fire effects is the highest priority of the 1980 field season. Upon completion" inventory summaries will be forwarded to the Colorado Division of Wildlife. CONCLUSIONS The Wintersteen Park burning operations are completed and the Colorado Division of Wildlife may now proceed with grazing trials and tests of hypotheses concerning bighorn sheep preferences in burned and unburned plots. The frustration of the many postponements, patchiness of the burn (especially in the grass plots), and failure to confine the fire to its predetermined boundaries should not be allowed to overshadow the overriding significance of this cooperative research effort. At least the burns are done and the Division and CSU may proceed with their postfire analysis of bighorn sheep preferences and plant succession on burned vs. unburned plots respectively. In spite of the problems identified with the burning operations, the Wintersteen burns provide a valuable learning experience for the several agencies involved. The remarks in this report are intended to provide constructive criticism of the planning and implementation of the Wintersteen burns and hopefully will facilitate improved cooperative efforts and exchanges of ideas. LITERATURE CITED Barrows, J. S., and R. F. Yancik. 1978. Prescribed fire planning and analysis for wildlife habitat. Report to Colo. Div. of Wildlife. Colo. State Univ., Dept. of For. and Wood Sci. Omi, P. N. 1979. Wintersteen Park Updated Prescribed Burning Plan. Report to Colo. Div. of Wildlife. Colo. State Univ., Dept. of For. and Wood Sci. 1The wind records of the weather station are based on wind counts over a given time period and actually mask instantaneous gusts. Prepared by ;(;ZdYV14 s /tl tJ{)-(j~ Thomas N. Woodard Wildlife Researcher C ��-329- JOB PROGRESS State of July, 1980 REPORT Colorado __ --~~~~---------------- Big Game Investigations Pro j ec t No. __;,,;W_-;;;:..1;;;:..2,.;;..6-....;R;;.:.-....;3:;__ _ Work Plan No. 4 __ __;,,;------------------- Job No. 2 of Bighorn Period Seasonal Sheep Investigations Dietary Preferences Sheep Covered: Personnel: Bighorn July 1, 1979 through June 30, 1980 T. N. Woodard, T. V. Dailey, R. B. Gill, M. L. Stevens, D. L. Baker, Dr. N. T. Hobbs, K. Pals, D. F. Reed, D. Butler, B. Van Sant, J. Ritchie, C. A. ~veinland, University of Colorado Mountain Research Station personnel. ABSTRACT Grazing trials with 3 tame bighorn sheep were conducted in the alpine vegetation zone during summer 1979 and winter 1979-80. Graminoids, forbs, and browse comprised 56, 38, and 6 percent, respectively, of the over winter diet and 72, 27, and 0.4 percent of the over summer diet. Crude protein content of diets was lowest in January and March (6.14%) and highest in early August (17.57%). . .. ��-331- SEASONAL DIETARY PREFERENCES BIGHORN SHEEP OF Thomas N. Woodard P. N. OBJECTIVES 1. To describe and quantify bighorn sheep. seasonal diet selection 2. To evaluate nutritional forages. 3. To test the hypothesis that forage yields in seasonal bighorn sheep habitats can be quantified within acceptable tolerance limits with available fiscal resources. 4. To quantify bighorn potential of Rocky Mountain and digestibility of bighorn sheep sheep forage yields within seasonal habitats. SEGMENT OBJECTIVES 1. Conduct bighorn sheep grazing trials in selected seasonal 2. Begin nutritional 3. Determine sample size requirements for estimating a clip and weigh sampling technique. analyses of bighorn habitat types. sheep forages. herbage yields with HETHODS AND HATERIALS Procedures used during this segment were similar to those described Woodard (1979). by RESULTS AND DISCUSSION Bighorn Sheep Food Habits In early August forbs were the most important dietary component with Polygonum bistortoides and Trifolium sp. being especially important (Table 1). Graminoids and browse were chosen less frequently with Agropyron scribneri showing the greatest use (Table 1). Late August diets were similar to those of early August (Table 1). During late September the amount of graminoids in the diet doubled, with Agropyron scribneri accounting for most of the increase (Table 1). Forbs still dominated the diet with Trifolium parryi being most important. �-332- Table 1. Percentage composition of bighorn sheep diet across 3 animals in alpine tundra type during summer 1979. Entries are mean percentages with 90 percent confidence intervals for plant species comprising 2 percent. or more of the total diet during anyone month. Species are ranked in order of importance across all months. Species Early August Percent in Diet Late August September All MonthJ-' Polygonum bistortoides 32.7 + 24.3 25.6 + 15.4 4.8 + 5.9 19.1 + 12.8 Trifolium das;YEh;yllum 23.3 + 23.9 22.6 + 34.1 9.9 + 11.6 17.9 + 22.8 Trifolium parryi 111.5 + 17.2 + 5.3 21.0 + 34.1 17.0 + 14.9 10.2 + 14.6 25.0 + 31.6 15.9 + 20.9 1.6 Agropyron. scribneri 8.8 + 11.1 CamEanula. rotundifolia 2.8 + 5.7 4.2 + 6.3 5.5 + 9.1 4.3 + 6.6 Carex spp , 3.9 + 1.8 4.1 + 4.9 4.3 + 3.6 4.1 + 2.9 Mertinsia sp. 2.3 + 5.7 7.3 3.7 + 5.0 3.1 + 5.3 Trifolium nanum --- 5.2 + 3~6 2.9 .± 3.7 + 7.6 0.6 + 0.8 3.0 + 2.7 Poa spp , 1.6+ 1.1 .1.0+ 0.6 3.6 + 3.2 2.3 + 1.0 Trisetum sEicatum 0.6 + 1.0 0.8-+ 0.9 3.2 + 3.1 1.7+ 1.6 Forbs 83.0 + 13.4 80.0 + 21.2 59.3 + 39.2 72.4 + 26.8 Grasses 16.0 + 13.4 .19.7 + 21.4 40.5 + 39.7 27.2 + 27.0 Shrubs 0.9 + Total Bites ,!/calcuiated with computed 1.4 ~0,629 percentages 0.3 + 0.4 12,547 0.2 + 0.5 15,730 0.4 + 38,906 by animal across all months. Across the summer-fall period forbs were the most frequently chosen forage class (Table 1). Important forage species include Po1ygonum bistortoides, Trifolium dasYEhyllum, Trifolium parryi, and the grass Agropyron scribner!. During November graminoids were the most frequently chosen forage class with Ca1amagrostis Eurpurescens being especially important (Table 2). The forb CamEanu1a rotundifo1ia was also important, and accounted for almost 5{)% of the forb consumption. 0.6 �-333- In January graminoids again dominated the diet with Calamagrostis purpurescens and Agropyron scribneri being especially important (Table 2). Campanula rotundifolia was again important. During March forbs dominated the diet with Trifolium dasyphyllum being • the most important forb (Table 2). The graminoid Calamagrostis purpurescens was the most frequently chosen forage species. Across all 3 winter months graminoids were taken most frequently with Calamagrostis purpurescens being especially important (Table 2). Table 2. Percentage composition of bighorn sheep diet across 3 animals in alpine tundra type during winter 1979-80. Entries are mean percentages with 90% confidence intervals for plant species comprlslng 2% or more of the diet during anyone month. Species are ranked in order of importance across all months. Percent in Diet -:N-o-v-e-m-b-e-r-----J-a-n..::.u-=a-=r-=y==-..=::.....::-=-=M~a:-r-c-h---- Species Calamagrostis purpurescens 34.8 + 39.1 49.0 + 34.2 Campanula rotundifolia 17.5 + 23.3 11.2 + 15.6 Carex rupestris 8.4+ Agropyron scribneri 2.2 Trifolium dasyphyllum 2.2 + 3.1 Salix planifolia (Leaves) 9.0 + 6.6 Polygonum 3.7 + 4.9 bistortoides Arenaria fendleri 5.1 10.0 + 1.9 + 1.7 5.5 + 9.7 5.0 + 3.9 3.9 + 3.7 + 7.6 2.1 + 2.0 6.8 + 12.2 3.8 + 7.2 2.9 + 2.5 3.3 + 0.9 0.4 + 0.4 2.2 + 2.1 + 2.3 2.1 + 2.1 1.8 + 2.4 3.4 2.2 2.7 Senecio canis 0.7 + 2.0 arvense + 2.8 10.2 + 2.1 + Cerastium 3.9 + 2.4 2.1 Kobresia myosuroides 0.5 7.6 + 7.3 1.9 + 0.2 + 11.9 + 15.1 5.6 2.3 alpina + 3.3 7.9 + 4.6 5.5 7.8 + 4.2 + OreoxiE 36.2 + 30.8 12.3 + 11.8 Salix planifolia (Stems) Sedum lanceolatum 5.3 25.3 + 17.0 4.5 + 4.5 0.6 + 1.1 5.1 + 0.1 + 0.2 5.7 + 15.9 1.6 0.9+ 1.7 2.1 + 2.1 1.1+ 1.0 0.5 + 0.8 2.3 + 3.5 0.8 + 0.9 1.4 7.1 + 4.4 Grasses 50.7 + 30.8 72.4 + 23.5 45.1 + 31.3 56.1 + 29.3 Forbs 36.0 + 27.7 25.4 + 25.1 52.4 + 30.7 37.6 + 27.3 Shrubs 13.3 + Total Bites l'Calculated 4.0 9,305 with computed percentages 2.2 + 9,279 5.2 2.5 + 1.3 8,353 by animal across all months. 6.3 + 26,937 2.1 �-334- Generally, forbs in the diet declined from late summer to January and then inclined from January to March (Fig. 1). This may reflect forb unavailability in winter rather than a decline in actual preference for forbs. Percent gr,pminoids in bighorn diets exhibited opposing trends to dietary forb trends, i.e. graminoids were lowest in August diets, highest in January diets, and 'declined from January to March. Browse composition of diets was minor all months sampled except November when percent of browse in the bighorn diets' increased to 13 percent (Fig. 1). Forage Quality Plant species listed in Tables land 2 have been collected, and are currently being analyzed for chemical constituents and in vitro digestibility. Nitrogen analysis for winter 1978-79 and summer 1979 has been completed. Dietary crude protein peaked in early August at about 17.6% and reached lows of about 6% in January and March (Fig. 2). Forage Yields Forage yield measurements were not attempted during this period. For further iItformation, refer to previous forage yield results section in Woodard (1979). LITERATURE CITED Woodard, T. N. 1979. Seasonal'dietary preference of bighorn Colo. Div., Wildl. Game Res. Rept. July, Part 2:419-430. Prepared by _;_1r.L:......~__:_:______:__:./l/..:...... LJ_' _O__;;D Thomas N. Woodard Wildlife Researcher ~..::....b-<4~,--' C sheep. �-335- ..• •• ,..... ••• Aug. 1 Figure 1. • • •••• Shrubs • •••••• Dec. 1 eo· • Mar. 1 Tame Rocky Mountain bighorn sheep diets summarized by forage categories for the period August 1979-March 1980 from alpine vegetation zones at Niwot Ridge, Colorado. �-336- 25 M~d Nov. Figure 2. Jan. Mar Aug~ Sept. Mean crude protein of diets of 3 bighorn sheep grazing alpine zones of Colorado during winter 1978-79 and summer 1979. �July, 1980 -337- JOB PROGRESS REPORT State of Colorado ----~~~~~---------- Project No. l-J-126-R-3 Big Game Investigations Work Plan No. 5 ----~-------------- Pronghorn Investigations Pronghorn Population Job No. 1 Period Covered: Personnel: Study July 1, 1979 through June 30, 1980 Marc Elkins, Gordon East, John Ellenberger, Lynn Sexton, Paul Neil, Beth Williams, Bruce McCloskey, and District Wildlife Managers in his Area. Appreciation is extended to David Peterson of the Utah Division of Wildlife Resources for demonstrating the X-ray equipment and to Jennine Regas of Animark Inc. for demonstrating the ultra-sound machine, and to Rob Deblinger and Jim Ellis at the Natural Resources Ecology Laboratory. ABSTRACT A series of six helicopter flights were made over Unit 56 between 22 June and 18 September. The average buck:doe:fawn ratio for the six flights was 41:100:55. Fawn to doe ratios exhibited a gradual increase throughout the summer. Group composition shifted from predominantly single does in early summer to doe:fawn groups by mid-summer and buck-doe-fawn groups by late summer. Buck-doe associations indicate that rutting behaviour begins by mid-summer. A portable X-ray machine and an A-scan ultra-sound machine were used on three mule deer and three pronghorn females. The ultra-sound machine detected 9 of 11 fetuses for 82 percent accuracy and the X-ray detected 8 of 9 fetuses for 89 percent accuracy. Both detected pregnancy in all cases. �-338- PRONGHORN POPULATION STUDY Thomas M. Pojar This was the first year this project was active, therefore much of the work was exploratory in nature. Work was continued on the literature search relating to pronghorn nutrition, reproduction, and genetics. More specific study plans for various aspects of the overall study and are currently in preparation. P. N. OBJECTIVE To determine southeastern causes for low fawn:doe ratios of pronghorn Colorado. populations in SEGMENT OBJECTIVES 1. Test the hypothesis that measurements of annual pronghorn production (i.e. classification counts) are inaccurate and tend to underestimate real productivity. 2. Test the reliability 3. Relate incidence of ultra-sound of diseases scanning in detecting to statewide pronghorn fetal rate. productivity. METHODS AND MATERIALS Herd Structure All herd structure estimates were made from a helicopter flown at 60-80 knots at an altitude about 30 to 40 meters. Three different types of helicopters were used because of limitations of the contractor. The three types are: Hughes 500, Hiller-Soloy, and Bell 47G3. Most flights began about I hour after sunrise and were completed by mid-day. Every third north-south mile line served as the transect line. When a group of pronghorn was spotted they were pursued as closely as possible (usually 10 to 20 meters) for classification. The information was recorded on a portable tape recorder and later transcribed. Pregnancy Testing. Tame captured animals maintained at the DOW foothills facilities were used in these tests. Three mature pronghorn does and three mule deer does were available for testing. They were first physically restrained then given an injection of M-99 and Rompen for immobilization. An A-scan ultra-sound machine produced for pregnancy detection goats and a portable X-ray machine were used on each animal. in sheep and �-339- RESULTS AND DISCUSSION Herd Structure Estimates Six helicopter flights were made over Unit 56 between 22 June and 18 September and one flight was made over Unit 3 on 9 September. Using each group observed as a sampling unit the confidence interval for the buck:doe and fawn:doe ratios were calculated according to Cochran (1963:67) (Table 1). Count number 2 on 6 July was not begun until 1055 because of fog and clouds. The late start and weather conditions may have been a factor in the low number of animals seen that day. Count number 6 on 18 September was begun at 0914 which was about an hour late because of mechanical problems with the helicopter. However, a relatively large sample was still obtained on that count. With the exception of count number 2, which is based on a small sample, the fawn to doe ratio generally increased throughout the summer (Fig. 1). The trend in fawn:doe ratios does not exhibit the reported double peak referred to in Pojar (.1979). Kitchen (1974:81) observed that "Fawns spent up to 90 percent_ of their first 2.5-3.5 weeks of life lying hidden in vegetation". If this is the case, then it would be expected that the number of fawns seen would increase as the summer progressed, assuming that the fawns would remain hidden with their antipredator behavior at the approach of the helicopter. Twenty-three does were observed unaccompanied by other does during. the first count (22 June). Seventeen of them (73.9 percent) did not have fawns with them. According to Prenzlow et al. (1968) the peak of fawning should have been past, therefore it could be assumed that fawns were there, but still exhibiting antipredator hiding behavior rather instead of fleeing with the doe. On the last count (18 September), 14 does were observed unaccompanied by other does, 5 of which (35.7 percent) were without fawns. The group composition exhibited a decided shift during the summer. The early count (22 June) was dominated (47.6 percent) by doe groups, while the mid-summer count (24 July) had predominantly (47.6 percent) doe-fawn groups, and the last count (18 September) was composed mostly of buck-doe-fawn groups (59.0 percent) (Table 2). The percent of buck-doe-fawn groups more than doubled in counts 3 (24 July) and 4 (8 August) compared to the first 2 .counts. The increase in mixed groups indicates the onset of the rutting season (Neivergelt 1974), which supports Kitchen's (1974) contention that rutting behavior in the pronghorn begins in early summer. One flight was made by NW Region management personnel over Unit 3 on 9 September to estimate herd structure. The fawn to doe ratio of 83:100 is 30 percent greater than a comparable count from Unit 56, on 18 September. All speculation is tentative, however, until statistical analyses have been performed to test for significant differences. Detection of Pregnancy and Fetal Rate A portable X-ray machine and an A-scan ultra-sound machine were used on three mule deer and three pronghorn females in an attempt to detect pregnancy and number of fetuses. These procedures were done on 1 May which is late in pregnancy for both species. The results of the tests are as follows: �-340- Table 1. Herd structure estimates based on helicopter No. of Animals Observed Bucks Fawns Total Does flights over Unit 56. 90% Confidence Interval B:D F:D Count Number Date 1 6-22 33 56 25 114 59:100:45 9 to 127 31 to 59 2 7-6 11 29 20 60 38:100:69 5 to 70 31 to 107 3 7-24 9 63 31 103 14:100:49 6 to 22 35 to 64 4 8-8 30 87 42 157 34:100:48 10 to 59 34 to 62 5 8-22 61 122 69 252 50:100:56 28 to 72 44 to 69 6 9-18 38 88 56 182 43:100:64 29 to 57 51 to 76 182 445 243 870 41:100:55 Total Table 2. Composition of groups observed Ratio from helicopter flights over Unit 56. * 1 Group No. B,D,F, 2 Count Number 3 4 No. No. % % 18 40.0 23 59.0 36.1 18 40.0 6 15.4 4 11.1 1 2.2 3 7.7 0.0 1 2.8 3 6.7 3 7.7 47.6 11 30.6 5 11.1 4 10.2 % 3 7.1 1 5.3 5 23.8 7 19.4 B 8 19.0 5 26.3 4 19.0 13 D 20 47.6 5 26.3 2 9.5 B,D 1 2.4 2 10.5 0 D,F 10 23.8 6 31.6 10 See Table 1 for data that corresponds 6 No. No. * 5 No. % % with count numbers. % �-341- Predictions Animal No. Ultra-Sound X-Ray' Results Twins Twins Twins Twins Twins Twins Pronghorn 223 224 225 Twins Tw:ins Single Mule deer 13 146 ORF Twins Pregnant Twins * ** Could detect pregnancy No Picture Single Single ** * Twins Single" Twins but number of fetuses was not obvious. The X-ray picture was taken too far forward on the doe or else the second fetus may have been detected. LITERATURE Cochran, W. G. 1963. New York. 413pp. Sampling CITED techniques. Kitchen, D. W. 1974. Social behavior Wildl. Monograph No. 38. 96pp. John Wiley & Sons, Inc. and ecology of the pronghorns. Nievergelt, B. 1974. A comparison of rutting behaviour and grouping in the Ethiopian and Alpine Ibex. In: V. Geist and F. Walther (eds.). The Behaviour of Ungulates and its Relation to Management. International Union for Conservation of Nature Publications New Series No. 24:324-340. Pojar, T. M. 1979. Pronghorn population study. Job Prog. Rept. pp. 451477. In Colo. Div. Wildl. Game Res. Rept. Part Two pp. 231-502. Prenzlow, E. J., D. L. Gilbert, and F. A. Glover. 1968. Some behavior patterns of the pronghorn. Colorado Department of Game, Fish, and Parks. Division of Game Research. Special Report No. 17. 16pp. Prepared ~2t/ Thomas M. Poja Wildlife Researche~ ~ C �80 70 60 (/J Q) 0 Q 50 0 0 .-! 1-1 40 Q) p., I W .p. N I (/J ~ ~ 30 20 10 0 I 224 60 103 157 252 22 JUN 6 JUL 24 JUL 8 AUG 22 AUG 182 (Sample size) 18 SEP Date of Count Figure. Serial fawn:doe ratios obtained from helicopter flights over Unit 56 duri~g the summer of 1979. �July, 1980 -343- JOB PROGRESS REPORT State of Colorado ----~~~~~-------Big Game Investigations W-126-R-3 Project No. Work Plan No. 6 Rocky Mountain Job No. 1 Seasonal Dietary Forage Quality of Rocky Mountain Period Covered: Personnel: Goat Investigatio~s Selection and Goats July 1, 1979 through June 30, 1980 T. V. Dailey, T. N. Woodard, R. B. Gill, J. E. Ellis, M. L. Stevens, K. Pals, N. T. Hobbs, S. J. Dailey, D. L. Baker, D. Butler, B. VanSant, R. A. Binford, D. F. Reed, L. Adams, T. J. Spezze, D. Pskowski, D. Collins, University of Colorado Mountain Research Station Personnel, J. Ritchie, C. Weinland. ABSTRACT Three mountain goats obtained during the previous year were utilized in grazing trials during the summer and fall of 1979. All 7 mountain goats were used in grazing trials during the winter of 1979-1980. Across During winter protein August all months forbs were the most frequently chosen forage class. summer forbs accounted for a large portion of the diet. In the amount of graminoids and browse in the diet increased. Crude content of diets was lowest in January (6.48%) and highest in early (18.12%). ��-345- SEASONAL DIETARY SELECTION AND FORAGE QUALITY OF ROCKY MOUNTAIN GOATS T. V. Dailey, and T. N. Woodard P. N. OBJECTIVES 1. To describe and quantify Mountain goats. the seasonal diet selection 2. To evaluate the chemical constitution selected by Rocky Mountain goats. of Rocky and digestibility of forage SEGMENT OBJECTIVES 1. Obtain, rear and train 3-5 additional habit study. 2. Conduct mountain types. 3. Begin nutritional mountain goat kids for food goat grazing trials in selected analyses of mountain seasonal habitat goat forages. METHODS AND MATERIALS Capture, Rearing and Training of Mountain Goat Kids Four mountain goat kids were captured from free-ranging nannies on Sheep Mountain the Collegiate Range of Colorado during June-July 1979. One kid was captured by hand, and 3 were taken in clover traps. Rearing and training procedures previously outlined (Dailey and Woodard 1979) were followed for this second group of kids. Study Area Selection The procedures for this section were described Woodard (1979). previously in Dailey and The summer study area is about 3700 m in elevation with level terrain bordered by steep north and south facing slopes, and boulder fields. Grazing Trials The procedures for this section were covered in Dailey and Woodard (1979). During summer of 1979 forage selection of 3 yearling mountain goats was quantified at the Niwot Ridge summer study area. During winter of 1979-80, �-346- 6-7 mountain goats (4 kids and 3 yearlings) were utilized for grazing trials at the Niwot Ridge \vinter study area. Animals were transported to and from winter holding pens via over-snow machine and most often by following the researcher on foot. Forage Forage nutritional Woodard (1979). evaluation Evaluation procedures RESULTS were outlined in Dailey and AND DISCUSSION Diet Composition In early August forbs were by far the most important dietary component with Trifolium sp., and Polygonum bistortoides being especially prominent (Table 1). Graminoids and browse were selected considerably less frequently. Table 1. Percentage composition of mountain goat tundra type during summer 1979. Entries are mean confidence intervals for plant species comprising diet during anyone month. Species are ranked in months. diet across 3 animals in alpine percentages with 90 percent 2 percent or more of the total order of importance across all Percent in Diet -E-a-r-l-y-A-u-g-u-s-t---L-a-t-e~A:'=u"';:g;"::u=s;"::t;'_:::=""":::";;;S;"::e;"::p Species o21 P o b ~- 21.2 + 31.5 Trda 14.9 + 20.4 Trpa 17.0 + 16.6 9.2 + Mesp 22.8+13.5 9.8 + 13.8 2.3 16.3+41.8 Trna 4.7 + 3.9 16.4 + 13.6 12.4 + 7.5 12.2 + 12.9 10.2 + 8.2 8.8 + 6.6 11.9 + 8.0 13.0 + 6.3 6.2.+ 2.5 9.6 + 2.8 6.6 + 22.6 3.6 + 8.2 8.6 + 20.8 Caro 2.4 + 6.4 9.6 + 12.7 4.6 + 6.5 6.3 + 7.8 Phse 3.0 + 5.7 2.9 + 8.3 +17.2 4.6 + 7.9 Agsc 0.3 0.4 8.6 + 6.0 3.0 + 2.7 Podi 2.4 + 4.3 2.6 + 3.0 + 2.0 2.6 + 2.6 Hepa 0.0 0.7 7.1 + 1.9 2.6 + 0.8 Casp 1.6+ 1.2 4.0 + 2.6 1.8 + 1.4 2.5 + Forbs 98.0 + 1.6 Graminoids 0.9 Browse 1.1 II with computed ]J Species codes: Pobi Trda Trpa Mesp Trna Caro = = percentages Polygonum Trifolium Trifolium Mertensia Trifolium Campanula 6.0 2.1 93.5 + 8.6 83.9 + 9.8 91.8 + 2.1 2.8 + 5.9 ·13.8 +10.5 5.8 + 4.8 3.7 + 4.3 2.3 + 3.7 16,879 15,090 Total Bites - Calculated + 2.5 3.0 by animal bistortoides dasyphyllum parryi sp. namiun rotundifolia 15,481 2.4 + 2.7 47,450 across all months. Phacelia sericeae Phse Agsc = Agropyron scribneri Podi = Potentilla diversifolia Hepa = Heuchera parvifolia Castilleja sp. Casp �-347- Diet composition in late August was similar to early August. Campanula rotundifolia (a forb) was consumed more frequently August (Table 1). However, in late During September forbs were again very important when Trifolium sp., Phacelia sericea, and Heucheraparvifolia were frequently accepted (Table 1). Graminoids increased in importance due largely to consumption of Agropyron scribneri (Table 1). Across the summer period forbs were the most frequently chosen forage class (Table 1). Trifolium sp., Polygonum bistortoides, and Mertensia sp. were especially important. During November, graminoids and forbs were taken with approximately equal frequency. Important graminoids included Carex rupestris, Calamagrostis purpurescens,and Kobresia myosuroides (Table 2). The most frequently chosen forb was Campanula rotundifolia. Browse was taken less frequently than forbs and graminoids; however, Salix sp. (leaves) were the most important individual dietary component in November (Table 2). In January forbs were the most frequently chosen forage class with Campanula rotundifolia, Arenaria obtusiloba, Paronychia pulvinata and Trifolium dasyphyllum being most important (Table 2). Graminoids were chosen slightly less frequently with ..Carex rupestris, Calamagrostis purpurescens, and Kobresia myosuroides being important (Table 2). The amount of browse in the diets was very low. In March forbs were taken slightly more frequently than graminoids. Small quantities of a large variety of forb species made up this forage class with no individual species being especially prominent in the diets. The graminoid, Carex rupestris was especially important in the diets relative to all other forage species (Table 2). Similarly, the browse Salix sp. (leaves) was chosen quite frequently. Across all 3 winter months forbs were taken more frequently than graminoids. Browse consumption was highly variable (Table 2), probably because Salix sp. was unavailable in January because of deep snows. ----The change in diet composition Figure 1. across all months sampled is depicted in Forage Quality Evaluation Plant species listed in Tables 1 and 2 have been collected, and are currently being analyzed for chemical constituents and in vitro digestibility. Nitrogen analysis for winter 1978-79 and summer 1979 has been completed. Dietary crude protein peaked in early August at about 18%, and reached a low of about 6.5% in January (Figure 2). LITERATURE CITED Dailey, T. V., and T. N. Woodard. 1979. Seasonal dietary selection and forage quality of Rocky Mountain goats. Job Prog. Rept., Proj. No. W-126-R-2, Work Plan 6, Job I, Bighorn sheep and mountain goat investigations. Colo. Div. Wildl. lOpp. �-348- Table 2. Percentage composition of mountain goat diet across 6-7 animals in alpine tundra type during winter 1979-80. Entries are mean percentages with 90% confidence intervals for plant species compr1s1ng 2% or more of the diet during anyone month. Species are ranked in order of importance across all months. Species November Percentage January in Diet ~arch All Months!.! Carex.rupestris 17.6 + 11.8 16.6 + 8.7 Salix planifolia (leaves) 21.1 + 13.7 0.2 + 0.3 9.6 + 6.4 12.0 + 6.9 Campanul8. rotundifolia 13.3 + 5.7 10.9 + 5.5 4.2 + 2.0 9.9 + 4.0 Calamagrostis purpurescens 8.1 + 5.2 14.6 + 8.9 1.7 + 0.9 7.6 + 3.1 Kobresia myosuroides 6.7 + 5.9 10.0 + 6.4 1.1 + 0.9 5.6 Trifolium 2.1 + 1.5 6.2 + 2.3 5.9 + 3.7 4.9 + 1.2 0.9 + 0.8 9.0 + 7.4 3.6 + 1.3 4.9 + 3.3 5.2 + 4.0 3.8 + 1.9 6.2 + 3.8 4.3 + 2.5 0.4 + 0.6 7.0 + 6.3 2.0 + 1.6 3.4 + 2.8 fendleri 2.6 + 1.8 4.3 + 2.5 2.8 + 2.7 3.3 + species 2.6 + 1.5 1.7+ 1.5 2.1+ 1.3 2.4 + 0.9 Senecio canis 1.0+ 1.3 0.9 + 1.5 3.9 + 6.3 2.0 + 2.4 Solidago nana . 0.5 + 0.7 o 4.8 + 5.5 2.0 + 2.4 Oreoxis alpina 1.4+ 1.3 1.4 + 0.8 2.1+ 1.4 1.9+ 0.7 Erigeron 2.1 + 0.8 1.9 + 0.9 1.4+ 1.2 1.8 + 0.8 3.2 + 0.2 + 0.4 0.5 + 0.4 1.7+ 1.6 Arenaria dasyphyllum obtusiloba Geum rossii Paronychia Arenaria pulvinata Artemisia simplex Polemonium viscosum 3.4 36.2 + 16.5 21.0 + 11.9 + 4.3 1.4 Grasses 38.9 + 21.9 44.2 + 17.5 41.6 + 17.2 37.2 + 18.7 Forbs 38.4 + 10.1 54.4 + 17.8 47.3 + 12.2 49.6 + 11.8 Shrubs 22.7 + 13.8 11.1 + 13.2 + Total Bites 1/ 19,322 1.3+ 1.0 19,825 6.7 18,743 7.1 57,890 - Calculated with computed percentages by animal across all months. One animal included in calculations for November and March was deleted from all months calculations because of its overall small bite sample size. This accounts for some apparent discrepancies in means. �-349- 100 '. • ". .80 . 60 _ /._----_: . ·40 •••••. Graminoids I / / / 20 • / , . •/ / .: --if· ••• Aug. 1 -~ Figure 1. 0 ...•..... 0 .0 00 Oct. 1 .. .. .''2;\\",u,!~• • • ••• • • ••• Dec. Feb. Mar. 1 1 1 Diet by forage category of three~ocky Mountain goats during the period August 1979 through March 1980 from alpine vegetation zones at Niwot Ridge, Colorado. �-350- 20 / 15 / / d °ri III / oI.J / 0 $-I p.. III "Cl 10 - ::s u • ~ , / ~.~. $-I / I 5 Mid Nov. Figure 2. Mid Jan. Mid Mar. Mid May Mid July Mean crude protein content of diets of 3 mountain goats grazing alpine zones of Colorado during winter 1978-79 and summer 1979. Mid Sept. �July, 1980 -351- JOB PROGRESS REPORT State of Colorado ----~~~~~--------- Project No. Work Plan No. 6 ----~--------~- Job No. 2 Period Covered: Personnel: Big Game Investigations W-126-R-3 November Rocky Mountain Goat Investigations Rocky Mountain Goat Ecology Study 12, 1979 - June 30, 1980 Dale F. Reed ABSTRACT Literature of mountain goat (Oreamnos americanus) ecology with emphasis on dispersal, colonization, and interspecific competition was reviewed. Additionally, a wide spectrum of literature regarding "competitive exclusion" was reviewed. Detailed study plans outlining objectives, procedures, and study sites, and an environmental assessment of study activities, were prepared. ��-353- ROCKY MOUNTAIN ROCKY MOUNTAIN GOAT INVESTIGATIONS: GOAT ECOLOGY STUDY Dale F. Reed P. N. OBJECTIVE Prepare a detailed, long range Program Narrative describing specific hypotheses and/or objectives, procedures, schedules, costs, etc., and prepare a thorough Environmental Assessment describing anticipated environmental impacts of the proposed research. SEGMENT OBJECTIVES 1. Review literature of Mountain Goat (Oreamnos americanus) ecology with emphasis on dispersal, colonization, and interspecific competition. 2. Review: a wide spectrum exclusion". 3. Prepare detailed study plans outlining and study sites. of literature 4. Prepare an environmental assessment METHODS regarding "competitive objectives, procedures, of study activities. AND MATERIALS Methods involved literature search and study, much of which was done at Colorado State University Library, Colorado University Library, and Denver Public Library. Numerous reprints were requested directly from selected authors. RESULTS AND DISCUSSION For Objective No. land 2 see Appendix For Objective No. 3 see Appendix B. For Objective No. 4 see Appendix C. i ~-' Prepared by .0; if' JL (i1--Le ----7 -----; J r (\& :' .../.' ~ Dale F. Reed Wildlife Researcher C I I / A. �-354- APPENDIX A �-355- A review of mountain goat literature reveals there has been little more than speculation published concerning the problem of mountain goat-bighorn sheep competition. Klein (1953) reported a behavioral intolerance between mountain goats and bighorn sheep in Alaska. Although limited, his observations suggested that goats were indifferent to the presence of sheep, but sheep exhibit avoidance towards goats. While studying goats in the Collegiate peaks of Colorado, Hibbs (1965) observed goats and sheep grazing together on one occasion and agonistic behavior of goats toward sheep on another. However, he and others (Hibbs et al. 1969) considered mountain goat-bighorn sheep competition minimal except when both species occupied common winter ranges at lower (subalpine or montane) elevations. The unstated implication was that as mountain goat and bighorn sheep populations increased in density on common winter ranges, competition could become problematic. Geist (1971a) also minimized the potential for mountain goat-bighorn sheep competition because his observations indicated they selected distinct and separate niches within common habitats. But he emphasized " ...sheep could suffer if they depended exclusively on cliff terrain as a feeding ground in late winter and if the absence of alpine fir forced goats to feed extensively on grasses and herbs" (p. 272). In both of these studies the conclusions were based upon observational evidence collected over a short time interval. They did not study the density-dependent and environmentally dependent nature of competition over an array of density and environmental conditions. Competition could vary in intensity as species densities and winter serverity (principally snow dynamics) increase. Until at least a portion of the mountain goat-bighorn sheep competition question is resolved, there will not be sufficient information upon which to base informed management decisions. This program proposes to examine the nature of bighorn sheep-mountain goat competition and management strategies for coping with the problem. Unfortunately, the question of competition involves a multiplicity of ecological phenomena such as population regulation and community structure, and is at best complex. Competition or the "competitive exclusion principle" (sometimes referred to as Gause's, Volterra-Gause, or Lotka-Volterra principle) as developed over many years (Darwin 1859, Volterra 1926, Lotka 1932, Gause 1934, Elton 1946) is one element in a system of ecological thought (Hardin 1960). However, ecological thought has not lead to a concensus on the definition of competition or the exclusion principle. No single definition seems to please all workers (Jaeger 1974, Lidicker 1979). Considering all possible interspecific relationships, competition as defined by Odum (1953:170) is any interaction between two or more species populations which adversely affects their growth and survival. Jaeger (1974) ventured that competition occurs when two species jointly utilize a vital resource that is in short supply and one of the species eventually eliminates the other from the habitat where their distributions overlap. Rhetorical differences aside, it is certain that species interact with each other and their resources in numerous and complex ways leading to coexistence, survival through exclusion, or extinction over geological time (Jaeger 1974). It is equally certain that sympatric mountain goats and bighorn sheep fit this same paradigm. �-356- Generally, as with other ecological processes, competition can not be tested directly. Studies have grouped competition into two main categories: exploitation competition (Ricklefs 1973) and interference competition (Jaeger 1974, Levine 1976). Exploitation competition occurs when an individual or species is better adapted to obtain a limited resource, thus securing a larger portion of it. Interference competition occurs when an individual or species inhibits another species access to a resource. Both categories are considered. . Exploitation competition has been measured based on the concept of niche overlap (Levins 1968, MacArthur 1972, May 1974, 1975, Vandermeer 1972). How much overlap in utilization of a resource is necessary before exclusion occurs is a challenging question. Two species which overlap by 10% will interact differently than those that overlap by 80%. A number of different methods have been used to measure and analyze the common utilization of an array of resources by two species. Levins (1968), Vandermeer (1972), May (1975), Levine (1976) used competition coefficeints and matrices. Schoener (1974), Shannon et al. (1975), Hudson (1976, 1977) and Singer (1979) have used various methods to evaluate resource partitioning or division, habitat partitioning or utilization, and spatial distribution. Hudson (1976, 1977) concludes that the results of his study are simply descriptive of habitat selection and resource partitioning, and that they are difficult to interpret in terms of competitive processes. Contrary to the usual interpretation, resource overlap can be used as evidence either for or against the existence of competition (Sale 1974). Definitive conclusions can only be drawn when it is demonstrated that distribution and habitat preferences are altered in the presence of another species and this change is consequential to productivity (Hudson 1977). To resolve this problem it may be necessary to demonstrate that one species limits another's use to a limited resource. This could be demonstrated most directly by monitoring changes tollowing the planned or fortuitous removal of one of the species. Interference competition is usually measured by dil:l~ctobservation techniques (Geist 1971b, Kramer 1973, Altmann 1974). For interference competition to occur there must be some form of intraspecific communicatory mechanism, such as defense of space, that has been extended to interspecific interactions (Miller 1967). Considering the extent of intraspecific dominance and agonistic behavior exhibited by mountain goats (Chadwick 1977, Dane 1977), it is plausible that such extensions of behavior readily occur. Also, it is possible that agonistic behavior may be exhibited even before a resource becomes limited; thus becoming effective immediately and therefore having a high exclusion rate (Jaeger 1974) compared to exploitation competition (low exclusion rate potentially occurring over many generations). Jaeger (1974) emphasizes that competitive exclusion is a continuum of dynamic processes rather than a discrete event, and that it may result in (1) one species replacing a second species throughout the latter's entire geographic range forcing the latter to extinction, (2) one species excluding a second from part of the latter's range effecting some form of allopatry, often contiguous, and (3) two species remaining sympatric with the competitively superior species forcing a less efficient or aggressive species to abandon limited resources. Based on the period (1948-present; albeit brief in evolutionary terms) of potential mountain goat-bighorn sheep interactions �-357- in Colorado and observations on these species, it would appear that some degree of (2) and (3) could be occurring between mountain goats and bighorn sheep on sympatric ranges. Related to the potential of interacting and competing mountain goats and bighorn sheep is the dispersal of introduced mountain goats. Disper.sal-rate studies by Caughley (1970), Clarke (1971), and Davidson (1973) of populations of thar, red deer, and sika deer introduced into New Zealand indicate that dispersal rates for these three ungulate species varied by species, with sika deer dispersing slowest (1 mi/yr), followed by thar (3 mi/yr), and red deer (7 mi/yr). Caughley (1970) proposed two models to account for observed dispersal rates of thar: one was density-dependent, while the other was a model of random dispersal. His data suggested that thar dispersal rates were most closely approximated by the random or "innate" dispersal model. It would be valuable to conduct "goodness of fit" tests between observed mountain goat dispersal rates and predicted dispersal rates from Caughley's (1970) two models, because if the predicted pattern closely fit either of the two predictive models it would allow managers to predict the rate of spread of mountain goats from new introduction sites and adjust management strategies accordingly. Based on the general areas discussed several major hypotheses and associated corollary hypotheses can be formulated concerning the potential problem of mountain goat-bighorn sheep competition. 1. 2. Exploitative competition between sympatric mountain goat and bighorn sheep populations results from some degree of niche overlap. a. Habitat, especially winter habitat, overlap by sympatric mountain goat and bighorn sheep populations is related to similar habitat preferences (species graze same area). b. Diets of mountain goats and bighorn sheep overlap within habitat types sufficiently to warrant concern that forage demands exceed supply (species graze same vegetation). c. Biotic and abiotic resource (e.g. aspect, minerals, slope [percent], terrain, vegetation [cover and forage], and snow characteristics) partitioning by sympatric mountain goats and bighorn sheep is related to different anatomical and behavioral adaptations. d. Spatial distribution of mountain goats can be predicted and the relative importance of the numerous variables which affect it can be quantified. e. Seasonal movements of mountain goats are different than those of bighorn sheep in areas where at least some of their habitat overlaps. Interference competition between sympatric mountain goats and bighorn sheep results from interactions of species-specific adaptive behaviors and social environments. Such competition tends to influence mountain goat and bighorn sheep distribution and acts to regulate population density through differential vulnerability. �-358- 3. a. The frequencies of mountain goat-bighorn sheep (interspecific) agonistic encounters vary according to season, sex and age class, and other environmental variables. b. The frequencies of intraspecific agonistic encounters according to mountain goat sex and age class. c. Dominance is exhibited by one species when socialized mountain goats and bighorn sheep are maintained in a limited spatial setting. d. Avoidance or threat is exhibited by mountain goats when a socialized bighorn sheep is introduced into their area of activity. e. Avoidance or threat is exhibited by bighorn sheep when a socialized mountain goat is introduced into their area of activity. f. Avoidance, threat, or investigative behavior is exhibited by mountain goats when a bighorn sheep simulation is placed into their area of activity. g. Avoidance, threat, or investigative behavior is exhibited by bighorn sheep when a mountain goat simulation is placed into their area of activity. Dispersal and colonization sheep habitat or potential vary by translocated mountain goats affect bighorn bighorn sheep habitat to a limited degree. a. Given that a sufficient number of mountain goats are studied and that a sufficient number of years are available for study, mountain goat dispersal rates can be predicted. b. Mountain goat dispersal rates, like those suggested for thar in New Zealand (Caughley 1970), are more closely approximated by Caughley's (1970) "random" dispersal model than his "densitydependent" model (test "goodness of fit"). LITERATURE Altmann, J. 1974. Observational Behaviour 49:227-267. Caughley, G. 1970. Liberation, thar (Hemitragus jemlahicus) CITED study of behavior: sampling methods. dispersal, and distribution of Himalayan in New Zealand. N. Z. J. Sci. 13:220-239. Chadwick, D •.H. 1977. The influence of mountain goat social relationships on population size and distribution. Proc. Int. Mountain Goat Symp. 1:74-91. Clarke, C. M. H. 1971.- Liberations and dispersal of red deer in northern South Island districts. N. Z. J. For. Sci. 1(2):194-207. Dane, B. 1977. Mountain goat social behavior: social play structure and "play" behavior as affected by dominance. Proc. Int. Mountain Goat Symp. 1:92-106. �-359- Davidson, M. M. 1973. Characteristics, liberation, and dispersal of sika (Cervus nippon) in New Zealand. N. Z. J. For. Sci. 3(2):153-180. Darwin, G. 1859. On the origin of species by means of natural selection, or the preservation of favoured races in the struggle for life. John Murray, London. 469pp. Elton, C. 1946. Competition J. Anim. Ecol. 15:54-68. and the structure of ecological communities. Gause, G. F. 1934. The struggle for existence. Baltimore. 163pp. Williams and Wilkins, Geist, V. 1971a. Mountain sheep: a study in behavior and evolution. of Chicago Press, Chicago and London. 383pp. Univ. Geist, V. 1971b. A behavioural approach to the management of Wild ungulates. Pages 413-424 in E. Duffey and A. S. Watts. eds. The scientific management of animal and plant communities for conservation. Eleventh Symp. British Ecol. Soc., Blackwell Scient. PubIs., Oxford. 652pp. Hardin, G. 1960. The competitive exclusion principle. Science 131:1292-1297. Hibbs, L. D., F. A. Glover, and D. L. Gilbert. 1969. The mountain goat in Colorado. Trans. N. Amer. Wildl. and Natur. Resour. Conf. 34:409-418. Hibbs, L. D. 1965. The mountain goat of Colorado. Colorado State Univ., Fort Collins. 152pp. Unpubl. M.S. Thesis. Hudson, R. J. 1976. Resource division within a community of large herbivores. Nat. Can. 103(3):153-167. Hudson, R. J. 1977. Habitat utilization and resource partitioning by wild ruminants: multivariate analysis of nominally-scaled attribute data. Northwest Sci. 51 (2): 101-ll0. Jaeger, R. G. 1974. Competitive exclusion: comments on survival and extinction of species. BioScience 24(1):33-39. Klein, D. R. 1953. A reconnaissance study of the mountain goat in Alaska. M.S. Thesis. Univ. of Alaska, Fairbanks. 121pp. Kramer, A. 1973. deer species. Interspecific behavior and dispersion of two sympatric J. Wildl. Manage. 37(3):288-300. Levine, S. H. 1976. 110(976):903-910. Competitive interactions in ecosystems. Levins, R. 1968. Evolution in changing environments. University Press, Princeton, NJ. 120pp. Lidicker, W. systems. z. 1979. A clarification BioScience 29(8):475-477. of interactions Am. Nat. Princeton in ecological Lotka, A. J. 1932. The growth of mixed populations, two species competing for a common food supply. J. Wash. Acad. Sci. 22:461-469. �-360MacArthur, R. M. 1972. May, R. M. 1974. 5:297-232. Geographic ecology. Harper and Row. On the theory of niche overlap. May, R. M. 1975. Some notes on estimating Ecology 56:737-741. Ricklefs, R. 1973. Ecology. the competition matrix. of ecology. Adv. Ecol. W. B. Saunders Co., Chiron, Newton, Massachusetts. 861pp. Sale, P. F. 1974. Overlap in resource use and interspecific Oecologia 17:245-256. Schoener, T. W. 1974. Science 185:27-39. 269pp. Theor. Pop. BioI. Miller, R. S. 1967. >:Pattern and process in competition. Res. 4:1-74. Odum, E. P. 1953. Fundamentals Philadelphia. 384pp. New York. Resource partitioning in ecological competition. communities. Singer, F. J. 1979. Habitat partitioning and wildlife relationships of cervids in Glacier National Park, Montana. J. Wildl. Manage. 43:437~444. Shannon, N. H., R. J. Hudson, V. C. Brink, and W. D. Kitts. 1975. Determinants of spatial distribution of Rocky Mountain bighorn sheep. J. Wildl. Manage. 39(2):387-401. Vandermeer, J. H. 1972. Niche theory. Annu. Rev. Ecol. Syst. 3:107-132. Volterra, V. 1926. Vartatzioni e fluttuazioni del numero d' individui in specie animali conviventi. Mem. R. Accad. Lincer ser. 6:1-36. �-361- APPENDIX B �-362- STUDY PLAN Habitat Utilization and Resource Partitioning of Mountain Goats and Bighorn Sheep THE PROBLEM Mountain goats (Oreamnos americanus) not being indigenous to Colorado, were first introduced into the State in 1948. Several additional releases have been made since then, including a 1961 release in the Mt. Evans area. This area was within the range of an indigenous bighorn sheep (avis canadensis) population. The problem is that as these ungulates occupy this range and their populations fluctuate, it is speculated that at some threshold density of both species, "competition" will occur resulting in a competitive advantage for mountain goats. It is plausible that this has already occurred, and that any sympatric mountain goat and bighorn sheep populations have increased or decreased accordingly. The question of "competition" or competitive exclusiona involves a multiplicity of ecological phenomena such as population regulation and community structure. It can not be tested directly. This initial study proposes to test several hypotheses concerning habitat utilization and resource partitioning. Other study plans, such as those concerning "experimentally perturbed populations" or "removal experiments" may be added later. LITERATURE REVIEW There is a substantial volume of literature concerning concepts of species' exploitative interactions and methods to evaluate them. Such interactions have been measured based on the concept of niche overlap (Levins 1968, MacArthur 1972, May 1974, 1975, Vandermeer 1972). How much overlap in utilization of a resource is necessary before exclusion occurs is a challenging question. Two species which overlap 10% will interact differently then those that overlap by 80%. A number of different methods have been used to measure and analyze the common utilization of an array of resources by two species. Levins (1968), Vandermeer (1972), May (1975), Levine (1976) used competition coefficients and matrices. Berry and Marble (1968), Dacey (1971), Schoener (1974a, 1974b), Shannon et a1. (1975), Hudson (1976, 1977), McFetridge (1977), Harrington (1978), Ford and Krumme (1979), Krzysik (1979), Murai et al. (1979), and Singer (1979) have used various methods to evaluate resource allocation, division, partitioning, or use; habitat partitioning or utilization; ecological segregation; or spatial distribution. Hudson (1976, 1977) concludes that the results of his study are simply descript.ive of habitat selection and resource partitioning, and that they are difficult to interpret in terms of competitive processes. Contrary to the usual interpretation, resource overlap can be used as evidence either for or against the existence of competition (Sale 1974). Definitive conclusions can ~en t\lO species jointly utilize a vital resource that is in short supply ... one of the species will eventually eliminate the other from the habitat where their distributions overlap (Jaeger 1974). �-363- only be drawn when it is demonstrated that distribution and habitat preferences are altered in the presence of another species and this change is consequential to productivity (Hudson 1977). To resolve this problem it may be necessary to demonstrate that one species limits another's use of a limited resource. This could be demonstrated most directly by monitoring changes following the planned or fortuitous removal of one of the species. As it is imperative to understand that definitive conclusions concerning competitive processes can not be drawn merely from habit preferences/ utilization and resource partitioning studies, it is equally imperative to understand that there is a need for bas~c ecological information upon which to base a "crucial experiment" (Platt 1964). This study will proceed on the basis that at least a modicum of ecological information on mountain goat and bighorn sheep habitat and resource use must be gathered before experiments can be fully designed. OBJECTIVES To determine the types of habitats utilizeci and the amount of resource partitioning by sympatric mountain goats and bighorn sheep ,~ ~he Mt. Evans alpine tundra, krummholz ecotone, and suba.ip Lne regions. WORKING HYPOTHESES Several working hy~otheses can be formulated concerning the objective. 1. Habitats, especially winter habitats, of mountain goats and bighorn sheep overlap to the extent that the species populations are not in equilibrium (species graze same area, use same escape terrain, and use same shelter/cover). 2. Diets of mountain goats and bighorn sheep overlap within habitat types to the extent that forage demands exceed supply ~species graze same vegetation and use same foraging strategies). 3. Biotic and abiotic resource use (i.e. aspect, minerals, slope [percent], terrain, vegetation [cover and forage] and snow characteristics) is partitioned by sympatric mountain goats and bighorn sheep. 4. Seasonal movements of mountain goats are different than those of bighorn sheep in areas where at least some of their habitats overlap. The second working hypothesis, although central to the overall question of competition, has been addressed at least in part by Work Plan 4, Job 2 and Work Plan 6, Job I (Seasonal Dietary Preferences of Bighorn Sheep and Seasonal Dietary Selection and Forage Quality of Rocky Mountain Goats, respectively). Hence, this study will test hypotheses 1, 3, and 4. �-364- METHODS Mountain goats and bighorn sheep will be trapped in some of the areas previously used as saltlicks (Baumann undated) and in other mineral lick areas that may not be documented. An attempt will be made to cover as much of each lick area as possible with the traps. One to several clover traps will be used. They will be baited with salt, a grain/pellet feed preparation, apple pomace, or alfalfa depending on seasonal response. The traps will be set only when project personnel are in the area so that any banding and tagging, collections, and measurements can be accomplished promptly. The various sex and age classes will be banded or ear tagged as follows: Both speciesb Adult females - telemetry (148 and 172 MHz) or non-telemetry collars Adult males All yearlings - ear tags, possibly telemetry collars and males- non-telemetry some collars Kids and lambs - ear tags The telemetry collars to be used initially will be for locating and eventual sight confirmation of the animals. Later, telemetry activit~ (tip switch) collars will be used in conjunction with a remote, portable telemetry data site to collect sample periods of continuous movements and activity (head up and head down). Mountain goat and bighorn sheep habitat utilization or preference will be studied by observing banded and tagged individuals of both species. Any instances of aggressive or dominance behavior between individuals of the two species will be noted and described. Observations of marked individuals will initially encompass all four seasons. If possible, observations will be replicated, over time, within seasons, and at different densities. Ground and aerial "utilization" censuses will be conducted on a regular basis (2-3 times per month, weekly, or more often for selected areas or seasons) to detect changes in habitat utilization. Since much of the terrain is relatively inaccessible during the critical season, a technique employing a helicopter to survey either predetermined routes and/or randomly located quadrats may be necessary. If aerial "utilization" censuses are likely to provide reliable estimates of habitat utilization or preferences, a sampling scheme similar to those reported by Gill (1969) and Kufeld et al. (1980) may be devised. Preliminary information will be necessary before such a scheme can be fully planned. b All collars and tags will be individually numbered or color coded for purposes of identification. Non-telemetry collars for mountain goats and bighorn sheep will be black collars with white numbers, letters, or symbols, and white collars with black numbers, letters, or symbols, respectively. �-365- For purposes of preliminary work, the area to be considered will be all the alpine and any of the subalpine that has sympatric utilization within a 100 km2 area (10 x 10 km square, UTM right 41-50, up 78-87) (Fig. 1) which includes the Mt. Evans shelter house on the east, Hells Hole and Bierstadt on the west, Goliath Peak on the north, and Rosalie Peak on the south. The sample unit to be chosen will be the square km (1000 m2) or next lower metric scale (100 m2). Sub-sampling could even be done at the 10 m2 scale in high density or high utilization strata. Each 1.0 km2 quadrat can be designated by the 1000 m UTM grid lines drawn from the UTM tick marks of 1:24,000 or 1:62,500 topographjc maps. For example the 1.0 km2 quadrat that includes Lincoln Lake (Fig. 1) is designated 48-85. Unless preliminary information indicates a different course of action, a rectangular area (~ 3 x 6 km) ,which includes Mt. Spalding, Mt. Warren, Rogers Peak, and Lincoln Lake will be used for the ground census routes. During either the ground or aerial utilization censuses, data will be recorded on sex and age age, location (UTM), elevation, aspect, slope, snow, terrain, vegetation, and activity of each ungulate species (Fig. 2). ANALYSIS OF RESULTS The fundamental question asked in this study is whether there is a difference in habitat utilization between mountain goats and bighorn sheep and also whether they partition use of resources. Part of this question is how differences in habitat utilization or preference vary in time and space. A meaningful measurement of the variation may be a preference rating for each of the attributes studied. However, a number of analytical procedures have been used, some of which are as follows: Bonferroni z statistic used with x2(Neu et al. 1974) Coefficent of association (C7) Dice (1945), Cole (1949), Singer Multiple classification analysis Hudson (1977) Multiple discriminant analysis Hudson (1976), Matthews (1979) Multiple nominal scale analysis Hudson (1977) Multiple regression and partial correlation Hudson (1977) Utilization coefficient May (1975) (1979) Some of these analytical procedures have greater prec1s1on than the measurements of the attributes which they were intended to describe (Hobbs, personal communication). These procedures should be carefully examined before application is made to the preliminary data sets collected in this study. Other kinds of analyses or treatments of data may also be useful. Home range data based upon either polygon, ellipse (Koeppl et al. 1975), or grid cell may be analyzed and displayed by procedures discussed by White (personal communication). Possibly a computer generated x and y plot of locations (UTM) over time could be placed on video tape and used for building concepts and as part of the reported findings. �-366- APPLICATION OF RESULTS Results of this study will be most important in the planning of future studies to determine if a system can be developed to estimate the intensity and nature of competitive interactions. PUBLICATION Journal of Wildlife Management or Journal of Range Manag.ement will likely be a suitable outlet for the results of this study. Final data analysis will be done following the fifth winter's observations, and should be ready for publication by the fall of that year. PERSONNEL AND SCHEDULE Personnel Dale F. Reed Principle Investigator Wildlife Technician Schedule Activity Fiscal Years Establish census routes and telemeter and mark selected animals 1980-81 Estimate mountain goat and bighorn sheep habitat preferences on sympatric ranges 1981-85 LITERATURE CITED Baumann, T. G. Undated. Final report for the 1978 Mt. Evans bighorn sheep and mountain goat study. 38(56)pp. Typescript. Berry, B. J. L., and D. F. Marble. 1968. Hall, Engelwood Cliffs, NJ 512pp. Cole, L. C. 1949. The measurement Ecology 30(4):411-424. Spatial Analysis. of interspecific Prentice- association. Dacey, M. F. 1971. Regularity in spatial distributions: A stochastic model of the imperfect central place plane. Pages 287-309 in G. P. Patil, E. C. Pielou, and W. E. Waters, eds. Statistical Ecology. Vol. I. Spatial Patterns and Statistical Distributions. Univ. Park, PA. Penn. State Univ. Press. 582pp. Dice, L. R. 1945. Measures of the amount of ecological between species. Ecology 26(3):297-302. association �-367- Ford, R., and D. Krumme. 1979. Theor. BioI. 76:125-155. The analysis of space use patterns. Gill, R. B. 1969. A quadrat count system for estimating Colo. Div. Game, Fish and Parks Game Inf. Leafl. 76. J. game population. 2pp. Harrington, F. A. 1978. Ecological segregation of ungulates in alpine and subalpine communities. Ph.D. dissertation, Colo. State Univ. 164pp. Hudson, R. J. 1976. Resource division within a community of large herbivores. Nat. Can. 103(3): 153-167. Hudson, R. J. 1977. Habitat utilization and resource partitioning by wild ruminants: multivariate analysis of nominally-scaled attribute data. Northwest Sci. 51(2):101-110. Jaeger, R. G. extinction by 1974. Competitive exclusion: comments on survival and of species. BioScience 24(1):33-39. Koeppl, J. W., N. A. Slade, and R. S. Hoffmann. 1975. A bivariate home range model with possible application to ethological data analysis. J. Mammal. 56(1):81-90. Krzysik, A. J. 1979. Resource allocation, coexistence, and niche structure of a streambank salamander community. Ecological Monographs 49(2):173-194. Kufeld, R. C., J. H. Olterman, and D. C. Bowden. 1980. A helicopter quadrat census for mule deer on Uncompaghre Plateau, Colorado. J. Wildl. Manage. 44(3):632-639. Levine, S. H. 1976. 110(976):903-910. Competitive interactions in ecosystems. Levins, R. 1968. Evolution in changing environments. University Press, Princeton, NJ. 120pp. MacArthur, York. R. M. 1972. 269pp. Geographical ecology. Am. Nat. Princeton Harper and Row, New Matthews, J. A. 1979. A study of the variability of some successional and climax plant assemblage-types using multiple discriminant analysis. J. Ecology 67(1):255-272. May, R. M. 1974. 5:297-332. On the theory of niche overlap. May, R. M. 1975. Some notes on estimating Ecology 56 (3):737-741. Theor. Pop. BioI. the competition matrix. McFetridge, R. J. 1977. Strategy of resource use by mountain Alberta. M.S. Thesis. Univ. of Alberta. 148pp. goats in Murai, M., W. A. Thompson, and W. G. Wellington. 1979. A simple computer model of animal spacing. Res. Pop. Ecol. 20(2):165-178. �-368- Neu, C. W., C. R. Byers, and J. M. Peek. 1974. A technique for analysis of utilization-availability data. J. Wildl. Manage. 38(3):541-545. Platt, J. R. 1964. Strong inference. Sale, P. F. 1974. Overlap Oecologia 17:245-256. in resource Science 146(3642):347-353. use and interspecific competition. Schoener, T. W. 1974a. The compression hypothesis and temporal partitioning. Proc. Natl. Acad. Sci. 71:4169-4172. Schoener, T. W. 1974b. Resource Science 185(4145):27-39. partitioning in ecological resource communities. Singer, F. J. 1979. Habitat partitioning and wildlife relationships of cervids in Glacier National Park, Montana. J. Wildl. Manage. 43(2):437-444. Shannon, N. H., R. J. Hudson, V. C. Brink, and W. D. Kitts. 1975. Determinants of spatial 4istribution of Rocky Mountain bighorn sheep. J. Wildl. Manage. 39(2):387-401. Vandermeer, J. H. 1972. Niche theory. Annu. Rev. Ecol. Syst. 3: 107-132. �-369- Figure 1. 1:62,500 topographic map of the Mt. Evans study area showing the UTM 1000 m (1.0 km) grid lines. A 100 m grid scale is shown on the lower left. o 2 4 6 8 1 :62.500 ��-371- APPENDIX C �-372- COLORADO FEDERAL AID PROJECT W-126-R ENVIRONMENTAL ASSESSMENT Bighorn Sheep and Rocky Mountain Ecology Studies: Goat Bighorn Sheep and Rocky Mountain Competition Study Goat Program Leader R. Bruce Gill, Wildlife Research Leader Supervisor Harold M. Swope, Wildlife Manager/Research for Colorado Department of Natural Division of Wildlife Active Dates: Prepared November by: Resources 12, 1979-June ~\(~~ R. Bruce Gill and Dale F. Reed Fort Collins, Colorado 30, 1985 :;.J_ " VcJL 1- U ~-r?u, - . [ �-373- Table of Contents Page I. II. III. Proposed Action A. Habitat Utilization or preference and resource partitioning B. Dispersal Rates 375 375 The Environment A. Abiotic Environment B. Biotic Environment 1. Plants 2. Birds 3. Mammals 377 377 377 377 379 379 Environmental 380 IV. Mitigating V. Unavoidable VI. 376 Impacts Measures Adverse 380 Impacts 380 Local Short-term Use of Man's Environment vs. the Maintenance and Enhancement of Long-term Productivity 380 VII. Irreversible and Irretrievable 380 VIII. Alternatives to Proposed Action 381 IX. Consultation with Others 381 X. Literature Cited Commitment of Resources 381 �-374- SUMMARY This study proposes to investigate competition between bighorn sheep and Rocky Mountain goats within sympatric seasonal ranges. The investigation will center on several phases: habitat utilization or preference and resource partitioning; dispersal rates of translocated mountain goats; and others associated with estimating the intensity and nature of competitive interactions. This environmental assessment addresses only the habitat utilization and resource partitioning phase. Additional assessments will be prepared for the other two phases prior to their beginning dates. The studies will be located in 2 major environments: alpine tundra climax region and subalpine forest climax region. Major biotic and abiotic environmental impacts will be the removal of vegetation by trampling and eating soil at the saltlick areas used for trapping and vegetational disturbance with temporary weather and observation facilities. None of these impacts will be large nor permanent and are judged to be non-significant. A significant positive impact is expected to result to the human environment as a consequence of increased knowledge of bighorn sheep-mountain goat competitive interactions. Improved management for recreational aesthetics and sport hunting should result. �-375- I. Proposed Action The state of knowledge regarding the biology of bighorn sheep and Rocky Mountain goats is deficient and precludes meaningful management of the species. If populations of these animals in Colorado are to be maintained and perhaps increased for present and future generations, much ecological information must be assimilated. One aspect of sheep and goat ecology which is not well understood is competitive interactions. Habitat utilization of both species is a first step in understanding the relationship between these animals and ·their use of available resources. A second step is to determine if a system can be developed to estimate the intensity and nature of competitive interactions. I This research program, scheduled to continue through June 1985, will address bighorn sheep and mountain goat competition and mountain goat dispersal. The specific objectives of the research program are: 1. Describe mountain goat and bighorn sheep seasonal habitat utilization or preferences and resource partitioning within sympatric seasonal ranges and develop hypotheses concerning the intensity and nature of competitive interactions. 2. Measure dispersal rates of translocated mountain goat populations and test the "goodness of fit" between observed rates of dispersal and Caughley's (1970) predictive models of dispersal rates. Habitat Utilization and Resource Partitioning--Mountain goats and bighorn sheep will be trapped in some of the areas previously used as saltlicks, and in other mineral lick areas that may not be documented. An attempt will be made to cover as much of each lick area as possible with the traps. One to several clover traps will be used. They will be baited with salt, a grain/pellet feed preparation, apple pomace, or alfalfa depending on seasonal response. The traps will be set only when project personnel are in the area so that any banding and tagging, collections, and measurements can be accomplished promptly. The various sex and age classes will be banded or ear tagged as follows: Both speciesb Adult females - telemetry (148 and 172 MHz) or non-telemetry collars Adult males All yearlings and males- - ear tags, possibly telemetry collars non-telemetry some collars Kids and lambs - ear tags b All collars and tags will be individually numbered or color coded for purpose$ of identification. Non-telemetry collars for mountain goats and bighorn sheep will be black collars with white numbers, letters, or symbols, and white collars with black numbers, letters, or symbols, respectively. �-376- The telemetry collars to be used initially will be for locating and eventual sight confirmation of the animals. Later, telemetry activity (tip switch) collars will be used in conjunction with a remote, portable telemetry data site to collect sample periods of continuous movements and activity (head up and head down). Mountain goat and bighorn sheep habitat utilization or preference will be studied by observing banded and tagged individuals of both species. Any instances of aggressive or dominance behavior between individuals of the two species will be noted and described. Observations of marked individuals will initially encompass all four seasons. If possible, observations will be replicated, over time, within seasons, and at different densities. Ground and aerial "utilization" censuses will be conducted on a regular basis (2-3 times per month, weekly, or more often for selected areas or seasons) to detect changes in habitat utilization. Since much of the terrain is relatively inaccessible during the critical season, a technique employing a helicopter to survey either predetermined routes and/or randomly located quadrats may be necessary. If aerial "utilization" censuses are likely to provide reliable estimates of habitat utilization or preferences, a sampling scheme may be devised. Preliminary information will be necessary before such a scheme can be fully planned. For purposes of preliminary work, the area to be considered will be all the alpine and any of the subalpine that has sympatric utilization within a 100 km2 area (10 x 10 km square, UTM right 41-50, up 78-87) which includes the Mt. Evans shelter house on the east, Hells Hole and Mt. Bierstadt on the west, Goliath Peak on the north, and Rosalie Peak on the south. The sample unit to be chosen will be the square km (1000 m2) or next lower metric scale (100 m2). Sub-sampling could even be done at the 10 m2 scale in high density or high utilization strata. Each 1.0 km2 quadrat can be designated by the 1000 m UTM grid lines drawn from the UTM tick marks of 1:24,000 or 1:62,500 topographic maps. For example the 1.0 km2 quadrat that includes Lincoln Lake will be designated 48-85. Unless preliminary information indicates a different course of action, a rectangular area (tV 3 x 6 km) which includes Mt. Spalding, Mt. Warren, Rogers Peak, and Lincoln Lake will be used for the ground census routes. During either the ground or aerial utilization censuses, data will be recorded on sex and age, location (UTM), elevation, aspect, slope, snow, terrain, vegetation, and activity of each ungulate species. Dispersal Rates--Dispersal rates of experimentally translocated mountain goats will be measured by releasing mountain goats into area(s) without existing mountain goat populations within the release area or within a broad expanse (40~50 km) of contiguous habitat. Systematic relocation efforts will be conducted and the sex, age, (young, yearling, or adult) and location of all observed mountain goats will be recorded over several consecutive years (up to 10 years). Dispersal rates will be estimated according to the method described by Caughley (1970) for thar in New Zealand. �-377- These investigations will occur in several separate stages. Habitat utilization or preference studies will commence in FY 1980-81 and conclude in FY 1981-85. Dispersal rates of experimentally translocated mountain goats will be recorded commencing in FY 1981-82 through FY 1990-91. This assessment will include only those activities concerned with habitat utilization and preference investigations. Separate assessments will be prepared for the dispersal rate study and other studies as they are developed. II. The Environment A. Abiotic Environment Location and Topography The Mt. Evans study area is located in the Front Range of the Colorado Rocky Mountains about 58 km west-southwest of the City of Denver. The elevation of Mt. Evans is 4,348 m and is surrounded by 6 mountain peaks over 4,000 m in elevation (Rogers Peak, Mt. Warren, Gray Wolf Mtn., Mt. Spalding, Mt. Bierstadt, Epaulet Mtn.) located within a radius of 4 km. Numerous glacial cirques occur at the origin of major drainages also within 4 km of Mt. Evans. The alpine topography is composed of gently rolling to steep slopes, rocky ridges, and rock slides. Geology and Soils The Mt. Evans study area is of granitic orlgln. The geology of the Front Range is discussed in detail by Lovering and Goddard (1950). Soils in the Collegiate Range are derived from 4 general classes of rocks, 1) pre-Cambrian crystalline, 2) Paleozoic sediments, 3) Tertiary igneous, and 4) Quarternary deposits. Most rocks are pre-Cambrian and consist of metamorphosed quartzite, limestone, and schists and gneisses of sedimentary origin (Hibbs,1965). Pleistocene glaciation contributed greatly to the geological formation of this area. Heather The following weather parameters change, as stated, with increasing elevation on Mt. Evans; 1) Precipitation increases, 2) average annual solor radiation totals do not change, 3) mean duration of frost free season decreases, 4) diurnal temperature range decreases, and 5) mean wind speed increases. Temperatures as low as -29°C and wind speeds of up to 90 mph have been recorded on Mt. Evans (Streeter 1969). The nearest weather monitoring facilities are located 15 km to the northeast. B. Biotic Environment Plants The Mt. Evans area supports a large continuous area of alpine vegetation. Vegetation of the area is typical of the alpine and subalpine zones �-378- (xarr 1961). The vegetation in the Mt. Evans area has been classified into 7 types by Streeter (l969) as follows: Rock Type Characterized by cliffs, outcrops, and stabilized and unstabilized slide areas. Rock covered 75% to 100% of the area. Slopes ranged from 350 to 900• Vegetative cover averaged less than 25%*. Dominant vegetation species were Trifolium nanun, Geum rossi, members of the Cyperaceae, Silene ~is, and~aria obtusiloba. Plant density was "low" with distribution "scattered". Sedge,.. TrifoliumRock Type Characterized by rock cover of 50% to 100% with numerous rock slides, and usually associated with cirque-basin slopes of 450 to 600• Trifolium spp. covered 5% to 50% of the area, while Geum rossi and Festuca spp. each covered 5% to 25%. Plant density was "scattered" or in "patches". CushionFellfield Type Found on exposed ridge and mountaintops above 12,500 ft. elevation. Rock covered 50% to 75% of the area and slope angle varied from 50 to 200• Dominant vegetation species were Trifoli~~ nanum, Silene acaulis, Arenaria obtusiloba, Paronychia sessiflora, and Claytonia megarhiza. Plant density was "continuous" with distribution "scattered" or in "patches". Wet-meadow Type This type was found only in the Mt. Evans study area and was not utilized by bighorn sheep. It was characterized by standing or running water or bog areas. Slope was less than 100• Major vegetation of the type was either Salix spp. (5% to 50% cover), Carex spp. (5% to 25% cover), and Deschampsia caespitosa (5% to 25% cover), or Carex spp. (50% to 75% cover) and Salix spp. (25% cover). Plant density was "continuous-"-.-- Sedge-GrassMeadow Type Rock cover was 5% to 50% and slope angle varied from 50 to 350• Members of the Cyperaceae covered 5% to 25% of the area. Poa spp., Festuca spp. and Agropyron scribneri each covered less than 5% to 25% of the area while Trisetum spicatum, Deschampsia caespitosa and Calamagrostis purpurescens each covered less than 5% of the area. Dominant forbs were Geum rossi, Trifolium spp. and Arenaria spp. Plant density was "continuous". Tree-Shrub Grass Type Scattered outcrops and boulder-fields accounted for the majority of the 25% to 50% rock cover of this type. Slope angle varied from 300 to 600• Trees covered 5% to 50% of the type and were various associations of Populus tremuloides, Pinus ponderosa, R_. contorta, Picea engelmanni and Pseutosuga menzesii. Shrub cover varied from 5% to 50% *- Vegetative cover is expressed as percent of the vegetated the classification, not including rock cover. area in �-379- Tree-Shrub Grass Type (Cont'd) Old-Burn Type with Cercocarpus montanus, Ribes spp. and Jamesia americana as the dominant species. Grasses covered 50% to 75% of the type with Bouteloua gracilis, Muhlenbergia montana, Festuca spp.and Calamagrostis purpurescens as dominants. Artemisia frigida, the major forb, covered 5% to 25% of the type. Plant density was "interrupted" to "continuous". Rock covered 25% to 75% of the type and slope angle varied from 250 to 450• Populus tremuloides, Pinus contorta and Picea engelmanii covered less than 25% of the type. The dominant grasses, in order of decreasing percent cover, were Bromus anomalus, Poa spp., Trisetum spicatum, Calamgrostis purpurescens, De?champsia caespitosa and Festuca spp. Plant density was "discontinuous" or in "patches". Birds Frequently observed in the alpine climax region of the Colorado Front Range are: marsh hawk, red-tailed hawk, golden eagle, prairie falcon, white-tailed ptarmigan, horned lark, common raven, water pipit, browncapped rosy finch, and white-crowned sparrow. Species common to the montane climax region are: sparrow hawk, mourning dove, great horned owl, mountain bluebird, broad-tailed hummingbird, red-shafted flicker, yellow-bellied sapsucker, williamson's sapsucker, traill's flycatcher, western flycatcher, western wood pewee, violet green-swallow, barn swallow, black-billed magpie, black-capped chickadee, house wren, robin, Townsend's solitaire, starling, Virginia's warbler, Audubon's warbler, redwing, Brewer's blackbird, black-headed grosbeak, Cassin's finch, red crossbill, Green-tailed towhee, savannah sparrow, vesper sparrow, chipping sparrow, song sparrow. Species common to the subalpine climax region are: mallard, green-winged and blue-winged teal, Goshawk, Cooper's hawk, peregrine falcon, blue grouse, Rivoli's hummingbird, belted kingfisher, downy woodpecker, hairy woodpecker, Hammond's flycatcher, gray jay, Clark's nutcracker, Stellar's jay, mountain chickadee, white-breasted nuthatch brown creeper, dipper, Swainson's thrush, ruby-crowned kinglet, warbling viero, yellow warbler, Wilson's warbler, western tanager, pine grosbeak, gray-crowned rosy finch, pine siskin, dark-eyed junco, gray-headed junco, and Lincoln sparrow. Mammals Many mammals are so ubiquitous in their distributIon that they do not fit well into community classifications. Migratory species such as mule deer, elk, and various predatory species fall into this category. Realizing this, the following lists of species common to the various life zones does not imply they are restricted to those zones. Alpine climax region: Coyote, red fox, long-tailed weasel, badger, yellow-bellied marmot, bushy-tailed woodrat, heather vole, white-tailed jackrabbit, pika, Rocky Mountain bighorn sheep and Rocky Mountain goats. �-380- Subalpine climax region: Water shrew, hoary bat, marten, Colorado chipmunk, red squirrel, northern pocket gopher, beaver, long-tailed vole, and snowshoe rabbit. III. Environmental Impacts Approximately 50 individual animals of each species (mountain goats and bighorn sheep) will be captured and marked with either ear tags, individually marked collars, or radio telemetry collars. Although maximum care will be taken possible trapping injuries could occur to some animals. There will be adverse impact on the vegetation located at the trapsites. This will be caused by trampling and eating the soil ~t the saltlick areas used for trapping. An area of less than 25 m2 will be disturbed for each of up to 5 trapping sites. An additional, although slight, disturbance to vegetation will occur as a result of temporary weather and observation facilities. A beneficial impact of this project will be the knowledge gained regarding bighorn sheep and goat ecology. Basic ecological knowledge relevant to habitat utilization will result in a better understanding of habitat needs and serve as a portion of the biological information needed to manage the species. IV. Mitigating Measures Adverse impacts resulting from trapping 50 animals of each species will be minimized by removing the clover traps after trapping is completed. Weather and observation facilities will be removed upon completion of the project. V. Unavoidable Adverse Impacts Disturbance of small areas of vegetation and possible injuries to trapped animals will be the only unavoidable adverse impacts. VI. Local Short-term Use of Man's Environment Enhancement of Long-term Productivity VS the Maintenance and Despite the susceptibility of alpine vegetation to disturbance (Billings 1973) most of the impacts related to this project are not expected to persist for more than 2 years following conclusion of field work. Longterm productivity for bighorn sheep and Rocky Mountain goats will be enhanced through improved management with ecological knowledge gained. VII. Irreversible and Irretrievable Commitment of Resources One irretrievable commitment of resources will be the removal of any seriously injured animals from the herd. This will have minimal measurable effect on the long-term productivity of the herd. The only other irretrievable commitment of resources will be the removal of vegetation at the trapsites. There will be no irreversible commitment of resources. �-381- VIII. Alternative to Proposed Action One alternative to this proposed research is to not conduct it and have no environmental impact. Without this and other continuing research efforts, bighorn sheep populations specifically, will most likely maintain their current status of declining trends. Another alternative is to proceed with the study as outlined and incur the minimal impacts delineated previously. The knowledge to be gained will improve man's understanding of bighorn sheep and mountain goat interrelationships with the environment. IX. Consultation with Others u.s. Forest Service representatives and scientists in the Big Game Research Section, Colorado Division of Wildlife reviewed the proposal. X. Literature Cited Billings, W. D. 1973. Arctic and alpine vegetations: similarities, differences, and susceptibility to disturbance. BioSci. 23(12): 697-704. Caughley, G. 1970. Liberation, dispersal, Himalayan thar (Hemitragus jemlahicus) J. Sci. 13:220-239. and distribution in New Zealand. of N. Z. Hibbs, L. D. 1965. The mountain goat of Colorado. Unpubl. M.S. Thesis, Colorado State Univ., Fort Collins. 152pp. Lovering, T. S., and E. N. Goddard. of the Front Range, Colorado. 223. 319pp. 1950. Geology and ore deposits Geol. Surv. Prof. Paper No. u.s. Marr, J. W. 1961. Ecosystems of the east slope of the Front Range in Colorado. Univ. Colo. Studies. Series in Biol. 8:1-134. Streeter, R. G. 1969. Demography of two Rocky Mountain bighorn sheep populations in Colorado. Ph.D. Dissertation, Colorado State University, Fort Collins. 96pp. ��July, 1980 -383- JOB PROGRESS State of Colorado Proj ect No. W-126-R-3 REPORT Big Game Investigations Work Plan No. 7 Black Bear Investigations Job No. 1 Black Bear Investigations Period Covered: Personnel: July 1, 1979 through June 30, 1980 A. Anderson, T. Beck, G. Bock, B. Boydstun, R. Danvir, B. Gill, L. Green, M. Haroldson, D. Kenvin, D. Miller, D. Sizemore, L. Stevens, M. Taylor; Colorado Division of Wildlife ABSTRACT A total of 21 individual bears were caught between 29 May-20 September 1979. Sex of captured bears was 12 females and 9 males. The katamine hydrochloridexylazine hydrochloride drug mixture worked exceptionally well with no drug related problems during handling of bears. The study area was enlarged to include areas of heavy :use by bears during the early fall berry season. Location of dens via aerial tracking was difficult and ground tracking was hampered by terrain and avalanche danger. ��-385- BLACK BEAR INVESTIGATIONS Thomas D. I. Beck P. N. OBJECTIVES 1. Develop techniques bear populations. to accurately and precisely estimate black 2. Determine habitat preferences 3. Describe black bear population dynamics sufficiently of various harvest and habitat manipulations. of selected black bear populations. to allow analysis SEGMENT OBJECTIVES Same as P. N. Objectives METHODS AND MATERIALS Study Area Delineation The Black Mesa-Crystal Creek area was initially selected as the general study area. Final delineation of study area will be determined after radio-tracking provides data on seasonal movements. Capture and Marking Capture efforts lasted from 29 May to 20 September 1979. All captures were made using Aldrich spring-activated snares and basic techniques described by Flowers (1977). All snare sets were out of sight from roads and maintained trails and were checked daily. All sets were marked with signs warning people to stay clear of both snares and snared bears. Snared bears were immobilized with a combination of ketamine hydrochloride and xylazine hydrochloride. The ketamine kydrochloride (100 mg/cc) was freeze-dried and concentrated to 200 mg/cc and rehydrated with xylazine hydrochloride (100 mg/cc) ina2:1 ratio (200 mg ketamine - 100 mg xylazine/cc). Drug was administered with use of a 6-foot jab pole. Dosage used, time to immobilization, and duration of immobilization were recorded. Bears were tattooed, ear-tagged with calf-size Ritchey rubber ear tags, and some were instrumented with a Telonics radio transmitter collar. Numerous physical measurements were taken and a premolar removed for subsequent aging by cementum annuli counts. Habitat Selection Data on habitat selection was of insufficient quantity to warrant analysis during this segment. Den site locations were found in mid-winter using aerial radio-tracking. �-386- RESULTS AND DISCUSSION Study Area Selection The initial study area was expanded to the north when all radio-collared bears (7) moved into the Smith Fork drainage in early September. These movements coincided with the ripening of chokecherries (Prunus virginiana) and serviceberries (Amelanchier sp.). The specific drainages added are the Clear Fork Creek and the Smith Fork River with all its tributaries. The area closed to bear hunting includes all the added areas except the roadless regions of the Smith Fork drainage nort.h of the river. A detailed delineation of study area will be completed after the 1980 field season and will appear in the next annual report. Capture and Marking The data reported are for the 1979 field season (basically calendar year 1979). Reporting of results on a biological year basis rather than fiscal year basis is the most reasonable approach and will be followed throughout duration of the project. Twenty-four black bear captures were made during the 1979 field season of which 21 were initial captures and 3 were recaptures. Nine of the captured bears were males and 12 were females (Table 1). Ages of bears were estimated from tooth eruption and wear and cementum annuli counts. There appears to be some serious problems in laboratory techniques in cementum annuli counting. Therefore, analysis of age data will not be attempted until the problems are resolved. Two radio-collared bears were legally shot by hunters outside of the closed area. A dosage rate of 3.0 mg/1b was found to be very satisfactory for ketamine/xylazine. No drug-related problems in handling bears were encountered and the compound has been very adequate for black bears. More detailed analyses of reactions to the drug and physical measurements will not be conducted until a larger numer of bears has been handled. Habitat Selection Data on den site location was severely hampered by the absence of the principal investigator after 30 September (on leave without pay). However, four den sites were generally located from aerial radio-tracking. Two sites were inacessib1e in March 1980 because of avalanche danger, one could not be found on the ground, and one was located. The located den was at the end of a tunnel at least 13 m long. A detailed description of this den will be obtained during the 1980 field season after emergence of the denned bear. Although data are limited, the general movement of bears during SeptemberOctober was very restricted as all the radio-collared bears stayed in the berry producing areas. The beginning of deer season (mid-October) sent all bears moving throughout the study area although berries were still abundant in the oak brush zones. Keeping up with the bears was difficult during the hunting seasons as daily movements were often quite large. Intensive radio-tracking is planned for this period in 1980 as the hunting seasons continue through the probable denning periods. �-387- Table 1. Colorado. Sex and weight of captured black bear, Black Mesa Study Area, Capture Date Sex Weight (kg) Remarks 5-29-79 M 141 Killed by hunter 7-13-79 6-13-79 F 70 6-13-79 M 68 6-17-79 F 11 Cub of the year 7-01-79 F 13 Cub of the year 7-18-79 F 27 7-19-79 F 27 7-20-79 F 48 , 7-25-79 M 120 7-27-79 F 75 7-30-79 M 107 8-01-79 F 43 8-01-79 F 68 8-02-79 M 41 8-11-79 M 84 8-27-79 M 52 9-10-79 F 59 9-11-79 M 70 9-14-79 F 107 9-17-79 F 50 9-20-79 M 48 Lost radio collar Killed by hunter 10-22-79 �-388- Nearly all of the 1979 field season was devoted to learning the area and capture efforts. Capture efforts will continue to be the major field effort in 1980. LITERATURE CITED Flowers, R. 1977. The art and technique Forest Protection Assoc., 37p. Prepared by .._J~j)j Thomas D. I. Beck Wildlife Researcher gtC~ C of snaring bears. Washington �July, 1980 -389- JOB PROGRESS REPORT State of Colorado --~~~--~--------- Project No. W-126-R-3 Work Plan No. -=8 Job No. 1 Period Covered: Personnel: Big Game investigations _ Mountain Lion Investigations Mountain Lion Population Dynamics July 1, 1979 through June 30, 1980 Allen E. Anderson, Lynn Sexton, Craig Albright ABSTRACT A review of the published and unpublished literature on the mountain lion was continued with about 900-1,000 references now read, catalogued and crossreferenced under 31 subject headings. Analyses of quantitative data on the biology of the mountain lion was the major activity. A list of the resultant 17 completed tables and 8 tables and 3 figures in progress is presented along with a working outline for the critical synthesis of literature now underway. ��-391- MOUNTAIN LION POPULATION DYNAMICS Allen E. Anderson P. N. OBJECTIVE Develop improved mountain lion inventory procedures and ~xpand on knowledge of mountain lion population dynamics and predator-prey relationships. Specific objectives are: 1. Estimate density and population 2. Develop an improved method which is also economically 3. Assess mortality rates of young mountain maternal-filial bond separation. 4. Assess 5. Estimate the impact of hunting dynamics. 6. Measure the inter-relationships large ungulate populations. dispersal size of selected mountain to reliably estimate mountain feasible to apply. and subsequent lion populations. lion densities lions after the period of fate of sub-adult and removal mountain on mountain between mountain lions. lion population lion populations and SEGMENT OBJECTIVE 1. Preparation of a detailed study plan describing the specific research strategies which will be employed to meet the previously stated specific objectives. This includes the selection of specific study site(s}. METHODS AND MATERIALS A comprehensive review of the published and unpublished mountain lion was undertaken as the first step. literature on the RESULTS AND DISCUSSION About 900-1,000 references pertaining to the mountain lion have been read, catalogued and cross-referenced under 31 subject headings. In perusing this material, it was apparent that very few biological parameters of the mountain lion have been quantified. It was also apparent that undocumented statements are common even in the most recent literature. This is particularly true of the presumed interactions between mountain lions and their prey species. For example, Russell (1978:213) repeats one of the most common unsupported assertions in the literature as follows: �-392- "Adult mule deer are taken by lions in greater proportion than they occur in their populations. This may be because the preferred habitat of adult bucks, to a greater extent than deer of other sex and age classes, closely coincides with that of mountain lions. The other, smaller deer are more abundant, however, and no doubt are taken in greater total numbers". No author is cited but only one (Hornocker 1970) of several studies, has reported a statistically significant higher proportion of adult bucks than adult does killed by mountain lions. The spatial juxtaposition of mature bucks and mountain lions is an untested hypothesis. In no other body of literature I've encountered, is it harder to separate fact from fiction. Thus, it appeared that, as part of both the planning process and as an aid to other researchers; the projected comprehensive review would be most useful as a critical synthesis; assembling, analyses, and a critical evaluation of all available quantitative data on the biology of the mountain lion. In view of the currently chaotic and unanalyzed body of information such a synthesis is essential before testable hypotheses can be formulated and a productive research effort initiated. A list of 17 completed tables and 8 tables in progress is given in Table I and a tentative outline of the critical synthesis of literature in Table 2. Current projections place the completion of a rough draft of this synthesis during the spring of 1981. As another part of the planning process, I visited 8 individuals during the spring of 1980 who are or have studied the mountain lion in New Mexico, Arizona, California, Nevada, Idaho, and Montana. I became acquainted with much unpublished information and learned something about problems to be expected in the field. LITERATURE CITED Goldman, E. A. 1946. Classification of the races of the puma. Pp. 177301, Part II in Young, S. P., and E. A. Goldman. The puma, mysterious American cats. The American Wildl. Inst., Washington, D.C. 358pp. Hornocker, M. 1970. An analysis of mountain lion predation upon mule deer and elk in the Idaho Primative Area. Wildl. Monographs. 21:1-39. Nowak, R. M. 1976. The cougar in the United States and Canada. U.S. Fish arid Wildl. Serv., Washington D. C. and New York Zoological Soc. 190pp. (Processed) • Russell, K. R. 1978. Mountain lion. Pp. 207-225. In Schmidt, J. L., and D. L. Gilbert. (Eds.) Big game of North America; ecology and management: Wildl. Management Inst., and Stackpole Books, Harrisburg, PA. XV + 494pp. Prepared by tilt.:gv £_~ tideA-d.ezJ Allen E. Anderson Wildlife Researcher C �-393- Table 1. Tentative titles of 25 tables and 3 figures to be included in the critical synthesis of literature on the puma. The scope of the information, and in some instances; the analytical procedures are given for the completed tables. Completed Tables Body or carcass weights (kg) of puma (Felis concolor) ssp. believed to be about 25 months or older. 22 references, N = 181 males, 164 females, of 6 subspecies. Changes in body weight (kg) of wild puma kittens prior to and after they had become independent of their mother. 3 references, N = 22 Sex ratios among North American puma. The hypothesis being'tested is whether the observed frequency of males is in accordance with the Mendelian expectation of a 1:1 sex ratio. 21 references, about 80 single classification chi square tests performed on samples totaling over 9,000 puma from bounty hunters, state and federal trappers, sport hunters, and three research studies. Measurements of the body or carcass of puma (Felis concolor) ssp. believed to be about 25 months of age and older. 13 references. Seven statistically described measurements of 211 males and 212 females of 3 subspecies. Comparisons of sub specific differences among 3 body or carcass measurements in 3 subspecies of puma. 13 references, 3 subspecies compared by ANOV for one measurement, 2 subspecies compared by ~ test for 3 measurements. An approximate record of change in numbers of puma killed within states and provinces, 1910-74, based on the compilation of Nowak (1976:154-166). Minimum total kill of puma; 66,665. A list of food items reported to be eaten by puma in North and South America. 10 references. Ranked major and minor food items identified from both stomachs and droppings of puma and estimates of sample size requirements (at + 10% of true value at P - 0.05) calculated. 12 references Parasites and diseases reported for puma. 47 references. Definitive, intermediate hosts, life cycle, major location in puma, pathology, incidence, and relationship to humans. Diseases and pathological 13 references conditions reported for puma. �-394- 2 home ranges (km ) of 23 males and 36 females puma in 7 states. 12 references. Mean, SD, of home ranges calculated where N > 3. Estimated Chi square analysis of buck:doe and fawn:doe ratios of deer killed by puma among 5 research studies. 5 references, in 5 states, N = 158 deer Comparison of buck:doe ratios of mule deer killed by puma with buck:doe ratios sampled in the living populations 2 references, analyzed by chi square contingency tables Comparisons of fawn:doe ratios of mule deer killed by puma with fawn:doe ratios sampled in the living populations. 2 references, analyzed by chi square contingency tables Comparison of subspecific differences in the mean body or carcass weight of North American puma. 22 references, 6 subspecies, ANOV and comparisons of means by the Studentized Range procedure. Tables in Progress Genetical characteristics of puma. Estimates of puma numbers 10 references on various Breeding dates in North American Litter size in North American study locales in North America. puma. puma. Weight of organs and glands in puma. Blood values in the puma. Comparison of 13 cranial measurements among 20 subspecies concolor) as tabulated by Goldman (1946:269-276). Chronology of dentition replacement Figures Distribution of puma (Felix in puma. in Progress of puma in North America. Regressions of body weight on body length in subspecies about 25 months of age and older. Growth in 3 young captive puma. puma believed to �-395- Table 2. puma. Working outline of the critical synthesis of literature on the Introduction I. Animal 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. II. Movements and Activity 1. 2. 3. III. Hunting Accidental Sex and age structure of the kill Population Characteristics 1. 2. 3. 4. VI. Qualitative Quantitative Nutrition Mortality 1. 2. 3. V. Home range Activity patterns Dispersal Food L 2. 3. IV. Taxonomy and distribution Chromosomes Cranial characteristics Dentition Weights and measurements Blood and organs Reproduction and productivity Growth and longevity Parasites and diseases Behavior Population structure Density Survival simulations Guesses about population trends by state Managemen t 1. 2. Outline by states Colorado procedures as one example Summary �-396- APPENDIX I PROGRAM NARRATIVE State: Colorado Project Title: Work Plan Title: Job Title: Big Game Investigations Mountain Lion Investigations Mountain Lion Population Dynamics Project No. Work Plan No. Job No. W-126-R 8 ---1 NEED Research on mountain lions (Felis concolor) increased dramatically during the decade of the 1970's. Nearly every western state initiated some kind of mountain lion investigation (Christensen and Fischer 1976). Some of this impetus stemmed from the perception of a western-wide decline in mule deer numbers during the late 60's and early 70's (Phelps 1976). Still another concern was that wildlife preservation groups were pressuring for a cessation of mountain lion hunting seasons (Hornocker 1972) while livestock interests were requesting increased harvests to cont ro l, predation on domestic livestock herds (Christensen and Fischer 1976). Most of the research of the 70's was directed towards inventory, population dynamics, and determination of home ranges (Russell 1978). Little research was directed towards the question of exploitation effects (Christensen and Fischer 1976) and even less was directed towards predator-prey interactions (Hornocker 1972, 1976). Hornocker (1972:401) summed up the first problem as follows: "There has never been an objective appraisal of the effects of long-term killing on predator populations, in spite of the fact that millions of dollars have been spent." Hornocker (1976:107) "Any discussion of the influence or effect of mountain lion predation on prey populations becomes, of necessity, a discussion of predator-prey relationships. The reason is obvious -- there are practically no data available on lion-prey interactions. Lions are known to kill deer wherever the two species occur together but this merely establishes the fact. The effect of this killing on deer numbers -- the really meaningful and important aspect to consider has scarcely been looked at in an objective way." The historic approach to evaluating the impacts of hunting on mountain lion populations has been to study an unhunted population (Hornocker 1969; Seidensticker et ale 1973) and then compare population dynamics, behavior, and movements of this "natural population" to a proximate and presumably similar hunted population (Christensen and Fischer 1976:38). This method has several limitations. First, studies of the two comparative populations have rarely been conducted within the same time frame. Secondly, levels of hunting removal are difficult to control especially regarding target sex and age categories. Thirdly, observed differ~nces between the two populations with respect to sex and age structure, territoriality, natality, mortality and density are interpreted as treatment effects. But the experimental design is inappropriate to do more than formulate these "effects" as hypotheses. �-397- A more productive approach would be to study the dynamics of an unhunted population for a pretreatment period then remove designated numbers of mountain lions from specified sex and age categories and measure population responses. This approach has been used with considerable success to study the effects of removal on populations of blue grouse (Bendell et al. 1972; Zwickel 1972, 1980; Zwickel and Bendell 1967; and Zwickel et al. 1977) and tits (Cedarholm and Ekman 1976); studies of the effects of predators on prey populations require simultaneous measurements of predator and prey population dynamics. The paucity of such studies in the literature attests to both the difficulty and the expense of this approach (Sinclair 1974; Theberge et al. 1978); Hornocker 1970; Kolenosky 1972; Shaw 1977). Hornocker (1976:107) commenting on the effects of mountain lion predation on mule deer populations remarked "We can't critically review the literature because there isn't any." OBJECTIVES 1. Investigate seasonal and diet activities, territoriality, population dynamics, habitat use, food habits and bioenergetics of an unhunted mountain lion population. 2. Remove designated numbers of mountain lions from specified sex and age classes and measure individual mountain lion and mountain lion population responses to removal. 3. Simultaneously monitor population dynamics of mountain lions and ungulate prey species to study the effects of mountain lion predation upon ungulate population dynamics. EXPECTED RESULTS AND BENEFITS The successful accomplishment of the objective of this reaearch should be central to the realistic attainment of the DOW strategic plan goals for mountain lion and should permit a more objective evaluation of alternative strategies for goal attainment. APPROACH Objective 1: A study area will be chosen where mountain lions currently are unhunted or where a long-term hunting closure (10 years) can be effected. All mountain lions encountered on the study area will be captured with the use of hounds and chemical immobilization equipment. Each mountain lion will be marked with ear and lip tatoos and radio transmitters equipped with mortality sensing device using a 6 hr delay in changes in pulse period. Drugs used will be ketamine hydrochloride and xylazine hydrochloride (Ramsden et el. 1976). Telonics, Inc. radio transmitters and receiving equipment will be used aerially and on the ground to measure seasonal and diet activities, territoriality and habitat use. Habitat preference will be. evaluated according to the method of Johnson (1980). �-398- The diet of mountain lion will be estimated qualitatively by identification of items in droppings picked up in the course of other field work. It may be possible to use a laboratory method of potentially positive identification of predator scat by species and perhaps by sex (Major et al. 1980). Kutilek and Clinite (n.d.) are studying its possible application to mountain lion. Food intake rates and energy consumption will be estimated for a period of one year using captive lions. We hope to use an experimental design with a sufficient number of lions to measure within and between variation in both sexes, 3 age groups « 1 month, < 13-30 months, > 30 months) utilizing 3 diets (mule deer, porcupine, and mule deer-porcupine combined). Except for the report by Golley et al. (1965) on energy balance in bobcats, there are few data on the bioenergetics of large North American carnivores which would be useful in the design of this experiment (Gessaman 1973). Objective 2: During the 6th year of study, those adult mountain lions which displayed activity and movement patterns judged most representative of their sex and age group over a minimum 2-year period will be recaptured and removed from the study area to a release site in comparable habitat hundreds of miles away. The activity and movements of the remaining individuals and the removals will be radiotracked at the same level of intensity as before. But telemetric surveillance of the transplanted mountain lions will be limited to one year. In addition, all mountain lions captured on the study area will be captured, radiocollared and tracked at 1 to 5 day intervals for four years. Inferences on changes presumably effected by this removal on the seasonal and diet activity, movements, sociability, home ranges, numbers, reproduction, mortality, and survival will be on the basis of qualitative and quantitative comparisons prior to and following removal. Estimated age and physiological and morphological measurements will be made on each captured lion with emphasis on those variables most likely to reflect social stress and perhaps treatment effect. The specific variables will be selected after additional literature search and consultation. Objective 3: During the 6 to 10th year of study, deer and elk will be censused during late winter on their winter ranges. The census will be in two places: (1) total winter range area, (2) confined to the home ranges of 2 randomly selected male and 2 randomly selected female mountain lion. The specific methodology used will depend on the vegetative and topographic characteristics of the study area and include either the helicopter quadrat census (Kufeld et al. 1980) or pellet group counts on sample units (Anderson 1980). Both methods use permanent plots in a stratified, random sampling design. �SCHEDULE Fiscal Objectives Segment 1,2 3,4 5 5 Segments 5-14 1 1 1 1 1 1 1 1 1 1 1 2 3 1 3 1 Activity July I· 1 Oct 1 ···T r 1 Jan Mar r --T· 1 4 I II r--l --_ Review of literature Selection/examination of study area Capture-radiocollaring of 2-3 mtn. lion Monitoring radio-collared lions Habitat Delineation Monitoring radio-collared lions Capture &radio-collared lions (3 hunters) Bioenergetics investigation (graduate student) J une 198081 -+ -+ -+ 198182 --------------+ I W 1.0 1.0 I Habitat Delineation Monitoring radio-collared lions Capfure & radio-collared lions (3 hunters) Bioenergetics Investigation (graduate student) 198283 Habitat Delineation Monitoring radio-collared lions Capture and radio-collar lions (1 hunter) Removal of selected mtn. lions (l hunter) Deer Census (Total Area) Monitoring radio-collared lion Installing quadrat census for deer within the home ranges of selected mtn lions Capture & radiocollar mtn. lions (1 hunter) Y ear -+ 198386 -+ 198687 -+ -+ -r -+ �Objectives 3 3 1 Activity Deer Census (Total Area) Deer Census (Lion home ranges) Capture & radio-collar lions (1 hunter) 1 Monitoring radio-collared lions . 3 3 Deer Census (Total Area) Deer Census (Lion home ranges) Monitoring radio-collared lions 1 , , , , , r=r: T July Oct Jan Mar H June T T ---,-- -, Fiscal Year ~ ~ 1987'90 ------------------~---------------------+ ~ - 199091 ----------------~------------------~--~-----+ I ~ o o I �-401- PERSONNEL Annual Person Days 205 156 267 210 Total 838 A. E. Anderson Professional Lion Hunter Wildlife Tech IA Graduate Student Estimated Annual Costs (01) (21) (28) (31) Personal Services Operating Supplies and Services Travel Expenses Capital Expenditures (Equipment) $ 76,610.00 37,630.00 8,745.00 2,300.00 Total GEOGRAPHIC Tentative: $125,285.00 LOCATION East side, Uncompahgre Plateau (GMU 62) RELATED FEDERAL PROJECTS None Known. LITERATURE CITED Anderson, A. E. 1980. Experimental deer inventory. In Game Research Report, July 1980, Part 1. Colorado Div. Wildlife-(In Press). Bendell, J. F., D. G. King, and D. H. Mossop. 1972. Removal and repopulation of blue grouse in declining population. J. Wildl. Manage. 36:1153-1165. Cedarholm, G., and J. Ekman. 1976. A removal experiment on crested tit Parus cristatus and willow tit Parus montanus in the breeding season. Ornis Scand. 7:207-213. Christensen, G. C., and R. J. Fischer (eds.). 1976. Transactions of the mountain lion workshop of the western states and western Canada. U.S. Fish and Wildl. Serv., Region 1. Portland, OR. 213pp. Gessaman, J. A. 1973. Methods of estimating the energy costs of free existance. Pp. 3-31. In Gessaman, J. A. (ed.). Ecological energetics of homeotherms: a view compatible with ecological modeling. Utah State Univ. Press Monograph Series 20:1-155. Golley, F. B., G. A. Petrides, E. L. Rauber, and J. H. Jenkins. 1965. Food intake and assimulation by bobcats under laboratory conditions. J. Wildl. Manage. 29(3):442-447. �-402- Hornocker, M. 1976. The possible influence of the mountain lion on mule deer populations. Pp. 107-109. In G. W. Workman, and J. B. Low (eds.). Mule deer decline in the West. A symposium. Utah State Univ. College Natur. Resources and Utah Agr. Exp. Stn. Logan, UT 84322. 134pp. Hornocker, M. G. 1972. Predator ecology and management - what now? J. Wildl. Manage. 36(2):401-404. Hornocker, H. G. 1970. An analysis of mountain lion predation upon deer and elk in the Idaho Primitive Area. Wildl. Monogr. 21:1-39. Hornocker, H. G. 1969. Winter territoriality in mountain lions. Wildl. Manage. 33(3):457-464. J. Johnson, D. H. 1980. The comparison of usage and availability measurements for evaluating resource preference. Ecology 61(1):65-71. Kolenosky, G. B. 1972. Wolf predation on wintering deer in east-central Ontario. J. Wildl. Manage. 36(2):357-369. Kufeld, R. C., J. H. alterman, and D. C. Bowden. 1980. A helicopter quadrat census for mule deer on Uncompaghre Plateau, Colorado. J. Wildl. Manage. 44(3):632-639. Kutilek, M. J., and E. W. Clinite. n.d. Species and self identification of large carnivores from fecal material with special reference to the mountain lion (Felis concolor): a research proposal. Dept. BioI. Sci., San Jose State Univ., San Jose, California. 6pp. (Typed). Major, M., M. K. Johnson, W. S. Davis, and T. F. Kellog. 1980. Identifying scats by recovery of bile acids. J. Wildl. Manage. 44(1):290-293. Phelps, J. E. 1976. Introduction to conference. p. 1. In G. W. Workman, and J. B. Low (eds.). Mule deer decline in the West: A symposium. Utah State Univ. College Natur. Resources and Utah Agr. Exp. Stn. Logan, UT 84322. 134pp. Ramsden, R. 0., P. F. Coppin, and D. H. Johnston. 1976. Clinical observations on the use of ketamine hydrochloride in wild carnivores. J. Wildl. Diseases 12:221-225. Russell, K. R. 1978. Mountain lion. Pp. 207-225. In J. L. Schmidt, and D. L. Gilbert (eds.). Big game of North America. Ecology and Management. Stackpole Books. Harrisburg, PA 17105. 494pp. Seidensticker, J. C., IV, M. G. Hornocker, W. V. Wiles, and J. P. Messick. 1973. Mountain lion social organization in the Idaho Primitive Area. Wildl. Monogr. 35:1-60. Shaw, H. G. 1977. Impact of mountain lion on mule deer and cattle in northwestern Arizona, Pp. 17-32. In R. L. Phillips and C. Jonkel, eds. Proc. of the 1975 Pred. Symp., Univ. Montana, Missoula. 268pp. �-403- Sinclair, A. R. E. 1974. The natural regulation of buffalo in East Africa. III. Population trends and mortality. Wildl. J. 12:185-200. Theberge, J. L., S. M. Oesenbrug, and D. L. Pimlott. seasonal variation in food of wolves, Algonquin Canad. Field - Natur. 92(1):91-93. populations E. Afr. 1978. Site and Park, Ontario. Zwickel, F. C. 1980. Surplus yearlings and the regulation density in blue grouse. Can. J. Zool. 58:896-905. of breeding ZWickel, F. C., J. A. Redfield, and J. Kristensen. 1977. Demography, behavior, and genetics of a colonizing population of blue grouse. Can. J. Zool. 55:1948-1957. Zwickel, F. C. increasing 1972. Removal and repopulation of blue grouse in an population. J. Wildl. Manage. 36:1141-1152. Zwickel, F. C., and J. F. Bendell. 1967. Early mortality and the regulations of numbers in blue grouse. Can. J. Zool. 45:817-851. � https://cpw.cvlcollections.org/files/original/9a12a335db7dfef2bd367412b7c67f99.pdf 32b4a751a48b3bff6835be8795c14047 PDF Text Text 1 JOB FINAL REPORT State of COLORADO Proj ect No. -'-S_E_-...:;..3_-_3 Work Plan No. 1 ----------------------- Job Title Colorado Period Covered: Personnel: _ Squawfish Endangered Job No. Investigations 4 ----------------------------- Propagation July 1, 1979 to February Wildlife Study 28, 1981 Gary Barta, Walter Graul, Charles Haynes, Robin Knox, David Langlois, Tom Lytle, Keith Murray, Clee Sealing, Stead and John Torres. Ken ABSTRACT A Colorado squawfish hatchery feasibility study was initiated at the Rifle Falls State Fish Hatchery near Rifle, Colorado. An engineering plan was prepared to accomodate research and production of 100,000 young fish annually. The hatchery plan includes: a hatching house to research spawn taking techniques, egg incubation, and fry rearing; two half-acre ponds and one quarter-acre pond for rearing fingerlings and overwintering brood fish, respectively; four concrete raceways for additional fingerling rearing space; a pollution abatement pond to restore water quality; and, a quarantine raceway to hold wild brood fish. �2 COLORADO SQUAWFISH Charles Haynes, PROPAGATION David Langlois, PROGRAM NARRATIVE STUDY Tom Lytle OBJECTIVES To design Colorado squawfish propagation facilities for construction at the Rifle Falls fish hatchery capable of producing 100,000 juvenile Colorado squawfish yearly; and, to begin construction of such facilities by June 30, 1979. INTRODUCTION With the knowledge that Colorado squawfish can be successfully propagated in a hatchery (Toney, 1979; Hamman, 1980) the State of Colorado decided to investigate and refine production technology. Spawn-taking techniques, handling of brood fish, and rearing methods for progeny are some of the research unknowns which must be answered to maximize production of fingerling squawfish on an annual basis. It was also decided that a facility devoted. totally to Colorado squawfish production and research would enhance the chances of La rge=sca'le success. The Rifle Falls State Fish Hatchery became a candidate site for a feasibility study based upon the following criteria: 1) it appeared to have land and water available to develop the facility; 2) it is adjacent to the historical habitat of t~e Colorado squawfish, i.e., the Colorado River and, 3) the existing hatchery and regional staff had ShO'Vllan interest in the culture of endangered species. While hatchery propagation alone will not result in the declassification of the species, such a source of young fish will aid the recovery effort in several ways. Fish of a uniform size and age are needed for bioassay work, including salinity, thermal tolerance, pesticide, and heavy metal toxicity studies. This knowledge would supplement (not replace) habitat requirement studies using wild fish. Taxonomic work with larval squawfish would facilitate field identification of wild fish, making it easier to monitor annual reproduction. Life history studies could also be conducted using hatchery fish both in the hatchery environment and in historic habitat. Squawfish·are needed to investigate competition and predation with other fishes, in addition to migration, growth, and survival. Such uses of hatchery - reared squawfish demonstrate the value of the hatchery product on the squawfish recovery effort EVALUATION AND DESIGN On August 2, 1978 personnel from the Division of Wildlife and the U.S. Fish and Wildlife Service met at the Rifle Falls hatchery to tour the unit, evaluated potential construction sites, and discuss the operation of the squawfish facility. Three possible sites were discussed and it was decided that the Engineering Section should design a construction plan. �3 The regional office made the engineering request. As a result, an on-site inspection was completed by the engineering section and three alternative hatchery plans were prepared for review during the winter months. On March 5, 1979 the Division of Wildlife and the U.S. Fish and Wildlife Service met again to review the three engineering designs and select the best one for further work. The details of the hatching house and pond layout were also reviewed. A copy of the preliminary design of the hatching house and the entire squawfish facility are included (Appendix A). Final approval is pending. The hatching house hopefully will provide an environment conducive to survival and spawning of larvae and juveniles. It includes several features requiring advanced fish culture and engineering technology. As water enters the hatching house from the settling pond or the existing trout hatchery source, it will be passed through an ultraviolet filter to kill potential pathogens. A water pump will push water through the system at a flow up to 225 gallons per minute. Because the incoming water will seldom be warmer than 550F (12.80C) a water heater will be used to increase water temperature to approximately 70 to 750 F (21.1 to 23.90C). By recirculating the heated water at a rate of 90 percent recirculated and 10 percent fresh water, it will be possible to significantly reduce heating costs. The energy source will either be electrical, propane or solar powered but in any case will have a back-up power source. Particulate material will be removed when the water passes through a settling basin. Additionally, the water will- pass through a denitrifying filter to reduce toxic ammonia waste products. Aerators will be used to restore dissolved oxygen to saturation and eliminate possible gas bubble disease caused by supersaturated nitrogen. Holding facilities include two indoor raceways for brood stock spawning and six small troughs to incubate eggs and rear fry. A small storage area is also included. Production raceways and ponds, .the quarantine facility, and the broodstock overwintering pond will be located outdoors. In the raceways, it will· be possible to rear fingerling squawf Lsh and conduct diet, growth and survival experiments. Similar experiments can be conducted in the ponds for comparison with raceway data. A quarantine facility will be used to isolate wild brood fish as they are captured and delivered to the hatchery. Disease inspection will be made in this raceway. A brood stock overwintering pond will be used to produce forage for the brood fish, hold the brood fish during non-spawning months, and research brood stock maintenance techniques. In July of 1979, regional personnel visited the Fish Hatchery, Willow Beach, Arizona to inspect and discuss propagation techniques. A synopsis obtained during this tour is attached (Appendix Willow Beach National this squawfish facility of culture parameters B). �4 The letting of bids and contracting of construction work were not accomplished by June 30, 1979. A request by the Colorado Legislature that squawfish not be propagated at the Rifle facility has placed an indeterminant delay on construction, which may proceed when, and if, this situation is resolved. LITERATURE CITED Toney, D.P. 1974. Observations on the propagation and rearing of two endangered fish species in a hatchery environment. Proc. West. Assoc. State Game and Fish Corom. 54: 252-259. Hamman, Roger L. 1980. Spawning and Culture of Colorado squawfish, humpback chub, and bony tail chub during 1980 at Willow Beach National Fish Hatchery. Paper presented at Colorado River Fisheries Project Workshop; Salt Lake City, Utah; November 20-21, 1980. 8pp. Prepared by ~L Charles M. Haynes Nongame Researcher �5 APPENDIX A: PRELIMINARY ENGINEERING DESIGN Note: The attached figures CA-I and A-Z) represent generalized schematics of the detailed engineering designs. The latter are on file with the Engineering Section, Colorado Division of Wildlife, Denver and are available upon request. Key to Figure A-I A B C D E F G Diversion structure, trash rack, and gate Settling pond Rifle Creek Hatchery building Production raceways Outdoor rearing ponds Clarifier Key to Figure A-Z A 6-foot double door B Generator room C = Storage room Settling basin D E = Settling basin with denitrifying box F Sand filter Boiler G H Ultra-violet filter I Aeration box J = Brood fish raceways K Egg incubation and rearing troughs Space heater L �8 A ==1~1 ... F Figure A-I ENGINEERING SCHEMATIC: COLORADO SQUAWFISH PROPAGATION FACILITIES, RIFLE FALLS STATE FISH HATCHERY, GARFIELD CO. , COLORADO (Modified from preliminory hatchery layout by Gary Barta, Engineering Section, Colorado Division of Wildlife, 6/5/79) ... �I I I I I I I $ : I I I I I I I $ I- e 4 I J B I I E ® D A r III~II , -- - 0 W~ 0 I I I I I 0 _-- --L~ , G I I I I I I ..hi')~ ~-~' ~ r ThJ~~··v ® K I $ $ $ $ f 8> ~O ENGINEERINGSCHEMATIC: COLORADO SQUAWFISH PROPAGATION FACILITIES, HATCHERY BUILDING (Modified from preliminary hatchery layout by Gory Barto, Engineering Section, Colorado Division of Wildlife, 6/5/79) 0 0 o C -...J �8 APPENDIX B: PROPAGATION GUIDELINES The following propagation guidelines are based on squawfish propagation experience gained at Willow Beach National Fish Hatchery, Arizona (Toney, 1974; Hamman, 1980). Additional information was provided by Tom Lytle (SW Region) and Keith Murray (Rifle Falls Hatchery), Colorado Division of Wildlife. Brood Stock Maintenance 1. Minimum holding water temperature should be no less than 60 to 65 degrees F (approximately 15 to 18 degrees C), while the recommended optimum temperature is 70 to 75 degrees C (21 to 24 degrees C) (D. Toney, personal communication). 2. Maximum holding-water temperature for brood stock has not been determined. An 8 inch (approximately 20 cm) specimen was maintained in an aquarium at water temperatures in excess of 90 degrees F (32.2 degrees F) with no observable stress. 3. Live rainbow trout fingerlings appear to be suitable as squawfish food. Toney (1974) observed that adult squawfish prefer crippled trout (heads pinched) and will not utilize commercial trout pellets. Feeding at Willow Beach was not conducted according to a rigid schedule; rather, brood fish were fed all that they would consume three to four times per week. Seven adult squawfish have been observed to consume as many as 160 three-inch trout fingerlings at one feeding. 4. Squawfish at'Willow Beach grew 3 inches (approximately 7.6 cm) per year (D. Toney, personal communication). ~.Jeightgain for six adults averaged 1.4 pounds (approximately 635 grams) per year (Toney, Disease and Parasites 1. Ich (Ichthyophthirus sp.) has been successfully treated at Willow Beach with a combination of formalin (25 mg per liter) and malachite green (0.05 mg per liter). Control was unsuccessful with a 1/1000 solution of potassium permanganate. 2. Bacterial diseases have been controlled by holding brood stock in a stagnation treatment of Furacin (10 ppm) for 7 hours or longer. Should Willow Beach stock be transported to Colorado, it is recommended that they be shipped in water containing 5 ppm active Furacin (D. Toney, personal communication). Spawning 1. Age at maturity has been determined for females (Hamman, 1980). as age V for males and age VI 1974). �9 2. Hormone (carp pituitary) injections followed by hand stripping has been used to induce ovulation (Hannnan, 1980). Rate and dose of pituitary extract which induces ovulation has been highly variable and fertility has been low. This would be a research item. 3. Spawning by wild adults has been accomplished at Willow Beach without benefit of hormone injections and/or hand stripping (Toney, 1974; Hamman, 1980). The following parameters were observed: a) Water temperature at egg deposition (June, 1974) approximately 72 degrees F (22 degrees C) after holding the spawners for one month at approximately 70 to 73 degrees F (21-23 degrees C) (Toney, 1974) Hamman (1980) observed spawning at the same facility at 68 degrees F (20 degrees C). b) A 95% fertility hormone-induced c) Spawning was accomplished in one of two raceways (Figure B-1). Substrate consisted of 8-10 inch rocks overlain by rocks of decreasing size until the top layer was composed of 3/4-1 inch material. The screen at the head of the raceway was 3/4 inch plywood with several 3/4-1 inch holes drilled below the level of the substrate. The bottom 12 inches were open, permitting water to flow under the platform and upwell through the rock substrate. The rock spawning area was covered with 6 inches (approximately 15 cm) of water (Toney, 1974). d) In 1979, the same raceway was utilized; however, spawning occurred in somewhat deeper water i.e. 8-22 inches (20-55 cm) of water (Hamman , 1980) . resulted spawning from natural spawning in 1980, while resulted in 80% fertility. Culture 1. Following spawning, Willow Beach personnel have employed two methodologies for hatching and culture. Toney (1974) removed eggs and rocks with attached eggs into a hatchery building where eggs were hatched in raceways with incubator trays of the type frequently utilized in salmonid culture. Hamman (persona.l connnunication) has permitted hatching to occur in the spawning raceways, afterwhich the larvae « 10 rom) are seined and transferred to hatching raceways. Although comperable data are unavailable; however, the second approach could be useful particularly when spawning is staggered and it is desirable that the spawning substrate not be disturbed. This would be a research item. 2. Hamman (1980) filled rearing raceways in advance of spawning and treated them with small amounts of inorganic and organic fertilizers to induce phytoplankton growth. Zooplankton appeared approximately 10 days later and served as the diet of the recently hatched fry. A starter trout prep~ration was added when fry reached 20-25 rom in length. �2t" PIPE BRINGING IN RIVER WATER AS NECESSARY \ -fa PLYWOOD WITH 15-20 III HOLES DRILLED BELOW LEVEL OF GRAVEL til- 1974 •... o FLOW SPAWNING SUBSTRATE 3" X 12" REDWOOD PLANKS Figure B-1 DiAGRAM OF SPAWNING AREA ~ R/T\CEWAY {J, (Willow Beach National Fish Hatchery ~Willow Beach, Arizona) �11 3. Jar culture of eggs will likely be unsuccessful, because fry tend to be lost in the overflow due to their small size (D. Toney, personal communication). Miscellaneous 1. Raceways at Willow Beach are concrete. All squawfish operations occur in either the raceways or hatchery house facilities. Ponds are not utilized. 2. Fry do not appear to school appreciably. 3. Estimated 4. The Rifle Hatchery facility would receive domestic River stock), lot number 4 DYAWB. Cannibalism is not apparent. time to bring fish to 9 inch (23 em) size is three years. progeny (Yampa �12 JOB PROGRESS State of Project COLORADO No. Job Title: Job No. Wildlife Investigation 1 ~~----------------------- II --~------------------ Nesting Performance Period Covered: Personnel: Endangered SE-3-3 Work Plan No. REPORT of Peregrine July 1, 1979 - February Falcons in Colorado 28, 1981 J. Enderson, Colorado College; E. Bauer, D. Berger, G. Craig, R. Meese, J. Patz, B. Pendleton and J. Rucks, Colorado Division of Wildlife; J. Hogan and S. Petersburg, National Park Service. ABSTRACT In 1980, peregrine falcon nest site occupancy continued to decline. Five adult pairs produced eggs and fledged young, primarily through augmentation. Shell thickness of eggs collected in 1980 remained constant with averages observed over the past seven years which is approximately 16 percent thinner than pre DDT era eggs. Organochlorine residues in eggs collected in 1979 did not vary significantly from residues observed in previous years. Organochlorine residues of potential prey collected in the vicinity of eyrie sites were compared. Killdeer, Brewer's blackbird, violet green, tree and cliff swallows and white-throated swifts possessed levels of DDE elevated sufficiently to be hazardous to peregrines. �13 NESTING PERFORMANCE OF PEREGRINE FALCONS IN COLORADO Gerald R. Craig and James H. Enderson P. N. OBJECTIVE The objective of this study is to annually monitor the breeding numbers and reproduction of Colorado peregrine falcons in an effort to document further population declines as well as eventually record the population's response to various recovery·efforts which are implemented. Additionally, health of the population may be monitored indirectly by analyzing pesticide residue levels in the falcons' eggs and principal prey they feed upon. Information obtained from these investigations will be made available through annual reports to the Rocky Mountain/Southwest Peregrine Falcon Recovery Team as well as cooperating agencies to aid in evaluation of various recovery efforts. SEGMENT OBJECTIVES 1. Annually document the number of breeding and their productivity in Colorado. 2. Annually survey potential habitats to locate previously unknown nesting peregrines and document their reproductive success. 3. Annually document egg shell thinning of nesting peregrines in Colorado. 4. Collect samples of principal avian prey species utilized by nesting peregrines and analyze the samples for pesticide residues. 5. Compile data and submit reports to appropriate state and federal personnel and the Rocky Mountain/Southwest Peregrine Falcon Recovery Team for use in evaluating recovery efforts. Note - These objectives corre~pond in the approved American Peregrine Southwest Population). pairs of nesting and pesticide peregrines residues in eggs to tasks 1113., 1114., 212., and 221 Falcon Recovery Plan (Rocky Mountain/ PROCEDURES 1a. Visit all nest sites throughout Colorado which have been occupied within the past three years and observe them from a distance with spotting scopes and binoculars to establish the presence of breeding adults. All occupied sites will be revisited periodically throughout the nesting season to document reproductive success. lb. Prior to, or immediately after hatching of the eggs, nest sites will be visited and eggshell fragments and addled eggs will be collected �14 for pesticide analysis. Eggshells will be measured for a thickness index according to standardized methods. Egg contents will be shipped to the Fish and Wildlife Research Laboratory at Patuxent for analysis. lc. Successful nests will be visited prior to fledging of the young and they will be banded and color marked. These sites will be kept under surveillance to determine actual fledging success. 2. Favorable habitats will be surveyed for potential nesting sites. The surveys will be conducted from the ground, but where necessary a helicopter may be used in remote regions. When new pairs are located they will be surveyed as outlined in la, lb and lc. 3. Ten to twenty individuals of each of 5 principal avian prey species will be collected from representative hunting areas utilized by breeding peregrines. This will be done at two sites annually. The individuals of each species will be combined into a single sample for pesticide residue analysis. The analysis will be accomplished by the Fish and Wildlife Service Laboratory at Patuxent or an independent laboratory recognized by the Patuxent Laboratory. 4. Compile data and submit reports to appropriate state and federal personnel and the Rocky Mountain/Southwest Peregrine Falcon Recovery Team. METHODS AND MATERIALS Nesting Investigations. Methods been described in the Procedures for nesting investigations enumerated above. have been Pesticide Analysis of Prey. Adult individuals of potential prey species within 8.3 km (5 mi) of active eyries were collected. The species were selected on the basis of high prevalence, and both migratory, non-migratory, herbivorous and insectivorous forms are included. Specimens were taken by shotgun and stored frozen. After thawing, the birds were weighed, plucked, and·the feet, beak and large and small intestine removed. The remainder was finely chopped, weighed and a sample taken for pooling. Species pooled samples were frozen in acetonewashed foil and submitted for analysis. Analytical procedures followed by Raltech Scientific Services, Inc. (formerly WARF Institute, Inc.,) at Madison, Wisconsin were as follows: Each sample is allowed to thaw in the refrigerator overnight. A 10 gram aliquot is weighed into a 250 ml beaker and mixed with a spatual with 150 grams of sodium sulfate. At the same time an aliquot is weighed into a preweighed 100 ml beaker for moisture determination and placed in a 400C oven for 2 weeks. After 2 weeks, weigh the beaker �15 and dry sample back and calculate percent of moisture. The 10 gram portion is allowed to dry overnight and is transferred to a 43 x 123 Whatman extraction thimble and plugged with glass wool. Place in a large soxhlet extraction aparatus and extracted with 50:50 Ethyl Ether: Petroleum Ether for 8 hours. Sample is removed and taken down just to dryness on a steam bath. Sample is brought to 25 ml with 25% Toluene in Ethyl Acetate. A 5 ml aliquot is transferred to Gel Permeation apparatus. The settings'are 28 mins discard, 14 mins collect and 2 mins wash. The sample is taken down on a flash evaporator just to dryness and made to 10 ml for injection. Injection: Hewlett Packard 5710A with Ni 63 Electron capture detector, with allta injector and hooked to a Hewlett Packard integration computor model 3352C. Column: 1.5% OV-17 x 1.95% QF-1 on 80/100 G.C.Q. 300oC; DDT Retention 200oC; Detector Temperature: 9.8 mins; Injection Temperature: 250oC; Carrier: Methane Flow 31 ml/min. Column Temperature: time of approximately 95% argon; 5% Lipid Determination: From the 25 ml volumetric containing the sample a 5 ml aliquot is pipetted into a preweighed 2 dram vial. These are placed into a 400C oven for 2 days, desiccated, weighed and percent lipid calculated. Pesticide Analysis of Peregrine Eggs. Infertile and unhatched eggs which are encountered in the wild, as well as those wild eggs which are removed for incubation in captivity and subsequently fail to hatch were analyzed for pesticide residues. Pesticide analysis of eggs was accomplished by the u.s Fish and Wildlife Service Research Center at Patuxent. In an effort to maintain uniformity in results, Patuxent should continue to perform the analysis. Samples were prepared for shipment by placing the egg contents in foil wrapped glass jars which had been rinsed twice in laboratory grade acetone~ The samples were then frozen and shipped in dry ice to the laboratory. Analysis techniques are the same as those described in Cromartie et al. 1975. Pesticide Monitoring J. 9:11-14. Since the eggs have undergone dehydration during incubation, residue values were corrected to fresh weight values assuming 85% moisture contents. RESULTS AND DISCUSSION Eyrie Occupancy. Table 1 includes available data on the historical use of 34 peregrine eyries in Colorado through 1980.· The presence of adult pairs, mixed pairs, and lone adults is shown. Data on the number of young fledged are incomplete. In 1980 there were seven adult pairs (BK, BU, PX, HR, GW, SB, CE) assuming that G~..;r and BU are distinct. There were three mixed pairs involving adult males and yearling females (SI, CR, WC), and three eyries with lone adults in the vicinity (RW, FX, HS). In all, thirteen eyries were occupied by at least one peregrine. An obvious trend toward abandonment continues. Only two of ten eyries �16 Table 1. Occupancy and productivity of Colorado peregrine eyries, 1972-1980 Year 1972 1973 1974 1975 1976 1977 1978 1979 1980 Eyries visited 15 23 24 26 27 31 32 33 34 Occupied 11 12 9 8 8 12 11 12 13 Adult pairs 8 11 7 6 5 11 7 6 8 Immature 0 0 0 0 2 0 2 2 3 3 1 2 1 1 1 2 3 2 1 5 2 4 6 5 4 5 eyries pairsl/ Lone adults Successful pairs Total young fledged No. of young augmented 2/0 11/0 5/0 6/4 11/5 16/11 12/8 Young Fledged/adult pair.~/ 0.18 1/67 0.83 1/20 1.00 2.29 2.00 2.00 Young Fledged/ successful pair 2.00 2.20 2.50 1.50 1.83 3.20 3.00 3.20 16/16 Percent of sites occupied~/ 73% 52% 38% 31% 30% 39% 34% 36% 38% Percent of sites w/ adult pairs 53% 49% 29% 27% 26% 35% 22% 18% 24% 1/ 2/ 1/ At least one member of the pair was in juvenile plumage. 1.25 young fledged per pair is considered normal reproduction. 80%-90% of the eyrie sites should normally be occupied in any particular year. with adult pairs in 1973 had adult pairs in 1980. Of nine eyries with adult pairs newly found during 1973-77, only three had adult pairs in 1980 (PX, HH, GW). Productivity of Nesting Peregrines. Five of the eight adult pairs produced eggs in 1980 and all succeeded in rearing and fledging young. Augmentation activities which were undertaken at four of the sites obscured actual production of the pairs since their eggs were removed and incubated artificially. �17 Table 2 summarizes the assumed natural reproduction had the pairs not received manipulation while Table 3 provides actual production as a result of manipulation activities. Although four young were seen on the nest ledge of site PX, only three of the young were observed on subsequent visits. At site HH, two of the brood of four were found dead on the talus slope below the eyrie. It is probable they fell to their deaths. Site BU was discovered late in the breeding season after 3 young had fledged, thus the number of eggs produced or young hatched could not be determined. Eggshell Thickness. Eggshell measurements were obtained from 11 wild eggs hatched in 1980 at the Peregrine Fund facilities in Fort Collins. Ten whole, unhatched eggs were sent to Patuxent for pesticide analysis and the shells were not returned in time for this report. The 1980 mean of 0.303 mm (with membranes) is very near the 1973-79 mean of 0.302 mm (n - 141). Thus, no change is apparent among samples collected since 1973. Table 2. Assumed Site Manipulation HH SB PX CN BU_Y Total Eggs Young Young Young Young Young Young Yes Yes Yes Yes No natural production !I No. of Eggs which would have been produced 4 4 4 5 3+'!:__/ 20 per laying pair hatched per laying pair per brood fledged for all pairs fledged per adult pair fledged per laying pair fledged per successful pair of managed and unmanaged No. of young which would have hatched 4 3 4 4 3+'!:__/ 18 4.00 3.60 3.60 1.36 2.14 3.00 3.00 II Natural reproduction is estimated had not been manipulated. II Pair was located after they had flying young-egg so assume a minimum of 3 eggs. for managed sites No. of Young which would have fledged 2 3 3 4 3 15 sites as though they production is unknown �18 Table 3. Site Actual production Manipulation of managed if of Eggs Produced and unmanaged II of Eggs Hatched sites. Young Returned to Pair If of Young Fledged HH Yes 4 SB Yes 4 PX Yes 8 eN Yes 4 BU No 3+ Total 23 Eggs per laying pair Young hatched per laying pair Young per brood Young fledged for all pairs Young fledged per adult pair Young fledged per laying pair Young fledged per successful pair 0 3 7 1 3+ 14 4.60 2.80 3.20 1.45 2.29 3.20 3.20 1/ so no young were returned. Site received no manipulation, 2 4 3 4 3 16 4 4 4 4 NAY 16 Organochlorine Residues in Egg Contents. As previously mentioned, ten wild eggs obtained in 1980 have not yet been analyzed at Patuxant and cannot be reported at this time. Results from eight eggs obtained in 1979 average 27.6 ppm DDE and 1.8 ppm PCB. A sample of 20 eggs or clutches obtained in 1973-78 averaged 21.2 ppm. The higher 1979 average may be accounted for by the unusually high (63 ppm) residues in one egg obtained from eyrie CR. When this value is deleted, there appears to be no difference in contamination in 1979 and former years. In previous years, (1977, 1978) a few eggs were found to contain as little as 7 ppm DDE. In 1979, the majority of the eggs had 16-32 ppm with none below that range and only one above it. Organochlorine Residues in Prey. Seven individuals each of 13 prey species were collected at six eyrie sites. Analyses were made under auspices of R. De Weese, U.S. Fish and Wildlife Service. Table 4 summarizes the data and compares them with results on ten species collected in previous years. In all ten cases, the 1980 levels are lower than earlier determination, and most of these are substantial decreases. The 1980 pesticide levels found in several species are noteworthy: 1) The starling, red-winged blackbird, and hairy woodpecker less than 0.1 ppm DDE, very low levels for insectivores. 2) The robin and white-throated swift were reported 0.2 ppm DDE, even though our earlier collections included pools with as much as 1.95 ppm DDE. samples have to have less than for both species �19 3) The Steller's jay pool had no detectable pool from New Mexico (LV) had 0.51 ppm. 4) One species not previously collected, the hermit thrush, contained 0.65 ppm DDE, far less than might be expected for a migrant insectivore. 5) Brewer's blackbird and the tree swallow, both bearing enough DDE to be a major source to peregrine, show levels one-third or less of those found in earlier years. 6) The robin was collected at CE in 1977 and 1980; DDE was about 11 times higher in the earlier pool. Table 4 Organochlorine residues DDE «0.04 in prey collected in 1980. Residues No. of Eyrie individuals Species ppm); a 1978 DDE1 (ppm, wet wt.) Total Orgchl.3 PCBs2 Solitary vireo CR 7 0.51 (-1.39) ND 0.56 Starling CR 7 0.06 (-0.39) ND 0.06 CR 7 1.10 (-4.9) 1.44 blackbird CR 7 0.06 (-0.43) ND 0.13 cowbird CR 7 0.37 (-0.83) ND 0.42 Hairy woodpecker FX 7 0.07 ND 0.07 Steller's jay FX 6 ND (-0.51) ND ND thrush FX 7 0.65 ND ND Tree swallow CE 7 11.1 (-21.7) 0.45 12.96 Robin CE 7 0.17 (-0.35) ND 0.17 PP 7 0.30 ND 0.30 Brewer's blackbird Red-winged Brown-headed Hermit Warbling Vireo 0.24 White-throated Swift RG 7 0.17 (-1.33) 0.17 0.42 Yellow-rumped warbler WC 7 0.68 (-0.29) 0.27 0.95 1 2 3 (difference from mean of 1977-79 pool(s) limit of detection - 0.10 ppm limit of detection = 0.04 ppm ) �20 Homogenates of individuals of four species collected 1977-79 and found to have high pool residue values were run separately in 1980 (Table 5). In three of four cases the mean of DDE values for the individual homogenates does not correspond closely to the earlier pool result. The mean for individuals is higher than the pool value for the robin, about the same for the killdeer and Brewer's, and substantially lower for the tree swallow. General agreement would be expected between the mean of individual analyses and the pool result because near-equal amounts usually 109, of individual homogenates were combined in the pool. Furthermore, there should be close agreement for percent fat and percent moisture values between individual homogenate means and the corresponding pool value (Table 5). There is close agreement in the case of moisture but the pool lipid values are consistently and substantially higher, especially in the case of the robin. It is clear that the individual homogenates did not loose moisture in frozen storage and that considerable error is apparent in lipid determination. However, since residues are reported on a wet weight basis, this error cannot explain the discrepancies between the average of individual and pool results. In any case, the discrepancies point out that pesicide determinations in different years by the same laboratory should be viewed with caution because of inevitable variation between technicians and techniques. The outstanding conclusion to be drawn from comparison of individual and pool results is the great variation in level of DDE contaminat'on between individuals within species. In the robin the variation is about 85-fold; two of the seven robins had low levels, one had a very high 11 ppm. The variation was seven-fold, 77-fold, and 14-fold in the Brewer's blackbird, killdeer, and tree swallow, respectively. Table 6 summarized all data on DDE and PCBS gathered from Colorado and New Mexico prey species 1977-80. Generally, birds with less than about 0.5 ppm DDE can be considered minor sources of DDE; those over about 1.0 ppm as contribution to significant shell-thinning in peregrines were they a major part of a female peregrine's diet. On this basis the following species are probably insignificant sources region-wide: pigmy nuthatch, mountain chickadee, red crossbill, nighthawk, Clark's nutcracker, mourning dove, western tanager, pinon and Steller's jays, flicker, the vast majority of robins, mountain and western bluebirds, Townsend's solitaire, blackheaded grosbeak, pine siskin, starling, and hairy woodpecker. The clearly hazardous forms include killdeer, Brewer's blackbird, violet-green, tree, and cliff swallows, white-throated swift, Say's phoebe, western wood pewee, and probably solitary vireo. In a few years, collections of such species as swifts, swallows, blackbirds, meadowlarks and robins should be made to ascertain possible changes in DDE levels. �21 Table 5. Organochlorine residues among individual prey compared with pool results Species Eyrie Year Robin CE 0.90 1977 (sample lost 0.13 11.0 0.35 2.21 0.52 MEANS POOL Brewer's blackbird CM 1978 MEANS POOL Killdeer HS 1978 MEANS POOL Tree swallow HS MEANS POOL 1979 DDE PCBs ND at lab) ND ND ND ND ND Residues (ppm, wet wt.) Total % lipid Orgchl. % H2O 0.90 2.8 69.7 0.13 12.19 0.42 3.51 0.57 4.3 5.2 3.9 4.1 5.6 68.3 67.2 69.2 70.4 65.7 2.95 4.3 6.2 68.4 69.6 2.52 2.07 0.12 22.3 9.56 34.4 4.82 5.93 8.89 7.17 0.17 0.16 0.12 ND 0.24 ND ND 26.8 9.84 34.58 4.93 6.32 9.12 7.33 3.7 4.3 4.0 5.0 6.2 6.2 6.4 69.2 69.5 70.2 66.7 66.5 68.2 67.7 13.3 16.7 0.1 0.21 14.13 5.1 6.0 68.3 67.6 12.6 35.7 5.03 21.0 1.97 153.0 9.93 0.14 ND ND ND 0.50 ND ND 14.66 37.13 5.03 21.57 2.59 153.75 13.97 8.7 10.(\ 4.4 6.1 7.8 4.2 4.9 65.4 58.9 68.5 66.4 67.1 69.6 69.0 34.2 31.7 0.09 0.31 35.5 6.7 7.9 66.4 66.0 27.7 17.2 55.1 16.7 4.03 4.25 6.38 ND ND 0.88 ND ND 0.42 0.58 27.95 17.43 56.64 17.63 4.03 4.82 7.06 8.1 10.1 10.3 4.7 4.3 8.2 4.9 63.0 61.9 65.3 65.4 68.6 64.9 67.9 18.8 32.8 0.27 0.42 19.37 7.2 9.3 65.3 66.1 �22 Table 6. Chlorinated Hydrocarbon Residues in Potential Peregrine Prey, 1977-1980 Species 1. Killdeer 2. Brewer's blk-bird 3. Violet-grn swallow 4. Cliff swallow 5. Tree swallow 6. Wht-throat swift 7. Meadowlark 8. Rd.-wg. blackbird 9. 10. 11. 12. 13. 14. 15. w. kingbird Say's phoebe W. wood pewee W. flycatcher Pigmy nuthatch Mtn. Chickadee Solitary vireo 16. Red crossbill 17. Nighthawk 18. Blk-throated gry. warbler Individ. Per pool 10 7 8 7 7 7 7 7 7 7 7 7 7 7 7 9 7 7 7 10 7 8 7 7 7 7 7 5 7 7 7 7 7 7 7 7 7 8 8 7 7 8 10 8 7 Eyrie, year HS, PR, HS, CE, CM, WE, PR, HS, CR, CR, CM, PX, CA, WE, PR, HS, HS, CE, CR, HS, PX, HS, FX, RG, PX, WE, HS, CE, CM, FX, MS, CR, CM, HS, HS, LA, LV, LA, LV, CA, CR, CA, HS, HS, 1978 1979 1979 1977 1978 1979 1979 1979 1980 1977 1978 1978 19781 1979 1979 1978 1979 1980 1977 1978 1978 1979 1979 1980 1978 1979 1979 1977 1978 1979 1979 1980 1978 1979 1979 19781 19781 19781 19781 19781 1980 19781 1978 1979 PX, 1978 Percent moisture fat 66.0 67.7 65.5 68.4 67.6 68.7 69.7 67.0 67.5 67.1 64.0 66.9 63.0 65.3 65.4 58.7 66.1 62.9 64.3 62.6 63.4 66.9 64.2 57.9 67.2 70.4 68.9 68.8 68.5 69.6 69.5 66.3 65.4 69.5 67.6 66.9 68.2 68.6 70.1 68.1 67.8 64.4 65.2 60.4 7.91 5.57 8.12 4.25 5.99 3.35 3.57 7.28 4.70 5.50 13.2 14.4 11.1 9.05 8.24 17.0 9.31 8.45 5.3 11.7 11.6 7.44 10.5 17.4 4.20 3.03 4.03 2.5 4.3 2.68 3.19 4.6 5.77 6.03 6.74 6.78 6.05 5.30 3.55 4.46 2.94 8.84 8.87 17.0 31.7 10.0 16.9 0.84 16.7 9.05 1.47 1.92 1.10 5.81 8.33 4.82 7.26 0.96 8.5 1.98 32.8 11.1 1.75 1.16 1.62 1.95 1.04 0.17 1.81 0.31 0.47 1.28 0.45 0.17 0.043 0.06 1.78 0.42 2.03 1.23 0.51 0.11 0.20 1.93 0.51 0.02 0.50 0.19 0.31 <0.01 <0.01 <0.10 0.21 <0.01 <0.01 <0.01 0.24 0.31 1.46 0.21 0.33 0.47 0.54 0.12 0.42 0.45 0.12 0.11 0.27 <0.01 0.76 0.17 0.21 <0.01 <0.01 <0.01 0.08 0.15 <0.01 ND 0.24 <0.01 0.01 0.15 0.18 0.13 0.10 0.10 ND 0.11 <0.005 <0.01 64.2 4.59 0.54 <0.005 % ppm (wet basis) PCB DDE �23 Table 6 (cont'd) Individ. Per pool Species 19. Y-rump. warbler 20. C. nutcracker 21. M. dove 22. W. tanager 23. Pinon jay 24. Steller's jay 25. Flicker 26. Robin 27. Mtn. bluebird 28. W. bluebird 29. T. Solitaire 30. Blk-hd. grosbeak 31. Brn-hd cowbird 32. Pine siskin 33. Starling 34. Hairy woodpkr. 35. Hermit thrush 7 6 7 7 7 7 7 7 7 7 7 7 6 7 7 7 7 7 9 7 7 7 7 7 11 6 7 7 7 7 7 7 7 7 7 7 7 7 Eyrie, year CE, LV, WC, CR, CA, CR, PX, LA, CA, CS, PX, LV, FX, CE, CR, FX, CE, LV, HS, PR, CS, FX, HS, CE, HS, LA, WE, CS, LA, 1977 19781 1980 1977 19781 1977 1978 1 1978 19781 1979 1978 19781 1980 1977 1977 1979 1977 19781 1978 1979 1979 1979 1979 1980 1978 19781 1979 1979 19781 PR, 1979 'HE, 1979 FX, 1979 CR, 1980 CS, 1979 PR, 1979 CR, 1980 FX, 1980 FX, 1980 1 New Mexico Prepared by: C. R. ~ Gerald R. Craiit Wildlife Researcher C Percent moisture 67.9 68.1 65.2 67.8 68.3 70.4 67.7 66.7 68.3 70.7 66.7 69.6 70.4 70.6 69.3 69.6 69.6 67.9 67.5 69.1 70.1 69.4 71.7 67.0 67.9 67.5 69.4 69.4 69.4 69.0 67.6 68.5 66.2 67.5 70.3 69.0 67.5 67.2 fat ppm (wet basis) DDE PCB 3.98 4.67 6.1 4.54 3.95 3.50 3.90 4.83 4.87 4.13 7.18 3.41 3.1 2.6 4.21 3.83 6.21 4.12 5.19 3.85 3.11 3.98 1.02 2.03 5.25 5.14 4.57 4.60 4.48 2.91 4.70 4.69 4.7 4.35 3.62 4.01 4.4 3.6 0.82 1.11 0.68 0.03 0.04 0.08 0.42 0.14 0.25 0.45 0.12 0.51 ND 0.06 0.04 0.087 2.07 0.49 0.10 0.13 0.18 0.15 0.52 0.17 0.096 0.09 0.26 0.30 0.08 .038 0.80 1.61 0.37 0.081 0.45 0.06 0.07 0.65 % <0.10 0.18 0.27 <0.10 <0.10 <0.10 <0.005 <0.10 <0.10 0.13 0.51 <0.10 ND <0.10 <0.10 0.13 0.12 <0.10 <0.005 <0.01 <0.01 0.14 <0.01 ND <0.005 <0.10 <0.01 <0.01 <0.10 <0.01 <0.01 <0.01 ND <0.01 <0.01 ND ND ND �24 JOB PROGRESS State of COLORADO Project No. ~S~E_-~3_-~3 Work Plan No. Job Title: _ II Physical Period Covered: Endangered Wildlife Job No. 2 and Biological Peregrine Personnel: REPORT Analysis Nesting July 1, 1979 - February Investigations of Colorado Habitat 28, 1981 J.E. Enderson, Colorado College; E. Bauer, M. Berman, B. Busby, G.Craig, M. McWhorter, J. Patz, B. Pendleton and J. Rucks, Colorado Division of Wildlife. ABSTRACT Physical characteristics and biological parameters have been collected for 32 peregrine falcon eyries. The 1980 season was devoted to completing photographic records of four sites. �25 PHYSICAL AND BIOLOGICAL ANALYSIS James H. Enderson OF COLORADO PEREGRINE NESTING HABITAT and Gerald R. Craig P. N. OBJECTIVE The primary objectives of this study are to examine physical and biological features of peregrine eyrie sites in an effort to eventually delineate specific factors which favor occupancy by peregrines. This information will aid in delineating those habitats which should be protected or enhanced for future occupancy by the falcons. SEGMENT OBJECTIVES 1. Establish physical and biological parameters which are uniform at all eyrie sites currently or historically occupied by peregrine falcons. 2. Delineate those human activities by nesting peregrine falcons. 3. When no alternative method exists, adults at specific eyrie sites may be trapped after their young have hatched and radio packages will be affixed to them to monitor their movements and hunting ranges. 4. Assemble the data and prepare designating new and potential land managers. Note - These objectives correspond American Peregrine Falcon Recovery and disturbances which are tolerated a report of the results eyrie sites by wildlife for use in agencies and to tasks 1111, and 1112, of the approved Plan (Rocky Mountain/Southwest Population). METHODS AND MATERIALS The investigation conformed to the following broad procedures. Procedures 1a. Annually, eight known historic and currently occupied peregrine eyrie sites in Colorado will be visited and the following physical aspects will be recorded: topography, geology, elevation, snow depth and precipitation, mean temperature, soil, presence and distance to water, and cliff characteristics. In addition, a series of photographs will be taken of each site. Each year, eight new sites will be surveyed until all known sites have been studied. lb. The vegetative types of the habitat within a distance of 15 miles of eight nesting cliffs will be cataloged whenever possible, appropriate Forest Service vegetative maps will be utilized and meadows and other potential hunting areas will be located on maps. �26 1c. Through use of standardized census techniques, avian prey diversity and abundance will be calculated for the months of April, May, June and July. This will be done for at least two sites annually. Censuses will be run once monthly in two localities for each habitat type represented. 2. Human activities, land use practices and audio and visual disturbances will be noted at each site and within an area of 15 miles of each site. If breeding pairs are present, they should be observed to note their reactions to potential disturbances. 3. At one or two sites annually, adult peregrines may be trapped and equipped with radio packages. This will be accomplished after the young have hatched and will extend for a period of approximately 3 weeks. The adults will be monitored as they embark upon hunting forays and locations will be triangulated primarily from the ground. Periodically, hunting adults will be tracked using a fixed-wing-aircraft to follow the radio signals. All locations of radio marked falcons will be recorded on appropriate topographic maps of the region. 4. Compile data forms and photographs taken during 1, 2 and 3 into notebooks. Prepare and submit reports of the results to appropriate state personnel, federal agencies and the Rocky Mountain/Southwest Peregrine Falcon Recovery Team. Physical Analysis Physical analysis of nesting cliffs and the immediate vicinity was accomplished by visiting the site and making a visual inspection of the area. Photographs were taken of the nest cliff and panoramas were compiled of the surrounding area from vantage points at the top of the cliffs. Vegetative information was also recorded during the visits and was further augmented from timber maps of the region. Elevational information was obtained from topographic maps of the region. Prey Abundance Line-transect counts of bird populations were taken in June and July at the seven eyries under study. Transects 805 m (0.5 mi) long were established within 8 km of each eyrie to quantify bird populations within the hunting range of resident peregrines. Transects were usually set in the most prevalent plant communities in each region and in the nearby riparian habitat where possible. Bird counts were made between 0630 and 1000 hrs by an observer walking the transect in 45-60 minutes, and then returning after a 15 minute pause. Species and numbers of individuals were recorded for all seen or heard, and unidentified birds were classed in three size groups. Individuals of each species for the "round t r Lp" on the transect were divided by two to yield the average number of individuals seen on the 805 m �27 transect on a given day. These daily means were in turn averaged to give the mean number of individuals of each species for each transect in the study period. Each species was classified in one of three subjective categories of vulnerability to peregrine predation, based on the relative amount of time each species was seen to fly in the open at least 15 m from the ground or other cover. "A" category includes forms often seen away from cover, e.g. Clark's nut cr-ackers ' "B" includes species often seen in cover but given to occasional long flights in the open, e.g. robins and flickers' "C" category includes species that seem infrequently subject to peregrine attack, e.g. rufous sided towhees. Large species such as buteos, ravens, and large waterfowl were given a "not applicable" (N.A.) rating. RESULTS AND DISCUSSION Physical characteristics of 32 peregrine falcon eyries and biological parameters relating to known territories have been collected and are being compiled into a final report. Panoramas of cliff faces at four eyries (PX, HH, SB and RG) were processed in the 1980 field season which completes the photo file for nearly all sites. During 1981, nest ledges will be marked on the photographs to record historical use of ledges. Prepared by \S Gerald R. Craig Wildlife Researcher C �28 JOB PROGRESS State of COLORADO Project No. SE-3-3 ----~~---------------- Work Plan No. Job Title: II ---------------------- Reintroduction Period Covered: Personnel: REPORT Endangered Job No. and Augmentation July 1, 1979 - February Wildlife Investigations 3 ~--------------------------- of Peregrine Falcon Production 28, 1981 J. Enderson, Colorado College; M. Elder, J. Hogan, S. Petersburg and D. Stevens, National Park Service; E. Bauer, D. Berger, M. Berman, G. Craig, R. Meese, J. Patz, and B. Pendleton, Colorado Division of Wildlife; W. Burnham, W. Heinrich and J. Hoolihan, The Peregrine Fund, Inc. and E. Freienmuth. ABSTRACT Four wild peregrine eyries were manipulated to increase fledging success. One pair was recycled to move them to a more accessible ledge and another site was altered to reduce the possibility of young falling from the ledge. Four hack sites were activated of which 3 successfully released a total of 11 young peregrines. �29 REINTRODUCTION AND AUGMENTATION OF PEREGRINE FALCON PRODUCTION Gerald R. Craig P. N. OBJECTIVE The objective of this study is to augment natural reproduction effort to sustain the wild peregrine population. in an SEGMENT OBJECTIVES 1. Augment natural production of peregrines by various techniques. 2. Monitor the results of the efforts, compile data and submit reports to appropriate state and federal agencies and the Rocky Mountain/ Southwest Peregrine Falcon Recovery Team. Note - These objectives correspond to jobs 222., 3133., 321., 322., 331., 332., and 3211, in the approved American Peregrine Falcon Recovery Plan (Rocky Mountain/Southwest Population). METHODS Augmentation AND MATERIALS efforts will be undertaken according to the following methods: 1a. Breeding pairs of peregrines will be observed to determine dates of initiation of egg laying. Within a week to 10 days after completion of the full clutch of eggs, the eyrie will be visited and all the eggs will be removed and artificially incubated. Approximately two weeks after removal of the eggs, the pair will recycle and lay a second clutch. The second clutch mayor may not be replaced with dummy eggs which the adults will be permitted to incubate. Dummy eggs should be substituted in situations where there may be concern about the adults' ability to incubate the eggs without breaking them. After a suitable period (if dummy eggs were substituted), the site will be revisited and the dummy eggs will be replaced with chicks from the eggs which were incubated and hatched in captivity. If the adults were permitted to hatch their own eggs, they will be permitted to continue to rear and fledge the young. Since there will be a 28 day difference in the age between the young produced from the first clutch and those produced from the second clutch, the young from the first clutch will be placed in other wild eyries containing similarly aged broods. Several representatives may also be retained for captive propagation purposes if similarly aged broods cannot be located. lb. As in 1a., initiation eyrie site dummy eggs breeding pairs will be kept under surveillance to determine of egg laying. Shortly after completion of the clutch, the will be visited and all the eggs removed and replaced with which the adults will be permitted to incubate. Since the �30 wild eggs usually are thin-shelled they will be artificially incubated to avoid their being accidentally crushed by the adult. After several weeks, the site will be revisited and young peregrines will be exchanged for the artificial eggs. Up to four young may be placed at each site to assure the maximum number of young are fledged. 1c. Captive produced young may be released at unoccupied or potential sites without the benefit of protection or care from adults through the technique of "hacking." Young falcons of three to four weeks of age will be placed on a suitable ledge at a potential reintroduction cliff site. They will then be cared for and fed by human attendants until they are flying and capable of feeding themselves. In this manner, the young falcons will return to the site at which they were reared and hopefully breed. This approach requires constant attendance and observation in order to protect the vulnerable young and insure they have sufficient food while they are in the eyrie. Because of this, the first two techniques will receive priority attention. If there are no additional adult breeding pairs as required by approaches 1 and 2, then young will be placed into the wild using this technique. RESULTS AND DISCUSSION Manipulation Efforts In 1980, the primary emphasis was placed on augmentation of wild pairs. Due to sufficient captive production, it was not necessary to supplement production by recycling wild peregrines. Three (sites SB, HH, and CN) of 5 pairs which produced eggs were augmented, one pair (PX) was recycled and one pair (BU) was discovered late in the season after they successfully fledged young. Table 1 presents the assumed natural reproduction of the sites had they not received manipulation. Difficulties are encountered when attempting to describe probable reproduction since the act of manipulation obscures the results. The estimates were developed by starting with the total number of eggs produced in the first nesting effort, then subtracting those eggs known to be infertile 'or inviable when they were removed, and finally subtracting those young which died after they were placed in the nest. This is misleading since egg mortality in the wild is probably higher due to abnormal water loss and breakaged associated with thin shells. Nestling mortality is also incorrect since young are placed in nests at 18 to 21 days of age. Undoubtedly some mortality would occur in the wild if the adults hatched and reared them to that age. Although we are aware of these biases, no attempt has been made to correct for them at this point. Thus, the assumed natural production is probably an overestimate. Table 2 presents the results of actual manipulation efforts. In comparing Table 1 and Table 2 it appears that manipulation only succeeded in producing �31 Table 1. Site !I Assumed natural production Manipulation of managed No. of Eggs which would have been produced HH Yes 4 SB Yes 4 PX 4 Yes CN Yes 5 3+2) No BUY Total 20 Eggs per laying pair Young hatched per laying pair Young per brood Young fledged for all pairs Young fledged per adult pair Young fledged per laying pair Young fledged per successful pair and unmanaged No. of Young which would have hatched sites. No. of Young which would have fledged 2 3 3 4 3 4 4 4 3+']) 18 3 15 4.00 3.60 3.60 1.36 2.14 3.00 3.00 11 Natural reproduction is estimated had not been manipulated. for managed sites or though they 11 Pair was located after they had flying young-egg production is unknown so assume a minimum Table 2. Site of 3 eggs. Actual production Manipulation of managed if of Eggs Produced and unmanaged if of Eggs Hatched sites. Young Returned to Pair if of Young Fledged 4 4 4 4 2 4 3 HH Yes 4 SB Yes 4 PX Yes 8 CN Yes 4 BU No 3+ Total 23 Eggs per laying pair Young hatched per laying pair Young per brood Young fledged for all pairs Young fledged per adult pair Young fledged per laying pair Young fledged per successful pair 0 3 II so no young were returned. Site received no manipulation, 7 1 3+ 14 NtJ./ 16 4.60 2.80 3.20 1.45 2.29 3.20 3.20 4 3 16 �32 1 young more than would have occurred naturally. This comparison is probably not reliable due to the above stated biases, as well as the extremely low number of nests available for manipulation. One site (PX) was recycled in order to relocate the pair to a more accessible nest ledge. This also served to delay the pair so that similar aged young were not required for all sites at the same time. The following sequence of activities occurred at the PX site: On April 30, a clutch of 4 eggs was removed from the nest ledge and not replaced with plastic replicas. The four eggs were transported to Fort Collins for artificial incubation. All eggs subsequently hatched. The pair relocated their nest and laid a second clutch approximately May 20. On May 29, the nest ledge was visited and the second clutch of 4 eggs was removed and replaced with a brood of 4 young 3 weeks of age. The adults immediately adopted and cared for the young. One of the four eggs was cracked and dented and not viable at the time of removal. The other 3 eggs were artificially incubated. In all, 5 young survived from the 7 eggs and were returned to the wild, primarily at hack sites. All 4 young survived at the wild site, but one disappeared shortly after fledging and was assumed dead. A second nest site (HH) was situated in an extremely hazardous locality and the pair should have been recycled in order to relocate them to a more secure nest ledge. However, the pair was discovered after they had completed their clutch and dates of egg laying could not be established. Efforts were taken to shore up and expand the existing nest site in order to reduce the chances of the young falling off the ledge. This was partially successful, but 2 of the 4 young did perish. It is assumed they fell from the site since their remains were located on the talus slope below the nest. Reintroduction Efforts Four hack sites were activated in 1980. The hack site in Rocky Mountain National Park successfully released young for the third successive year. A lone adult male released in 1978 continued to frequent the release site, but did not succeed in attracting a mate. Four females were released at the site in 1980 in an effort tri increase the opportunity to establish a pair. A second hack site was developed in Dinosaur National Monument and proceeded to the stage where five young were placed in the hack box. The night before the hack box was to be opened, great horned owls moved into the area.. In order to avoid almost certain predation by the owls, the young were removed and hack efforts discontinued. Perin's Peak near Durango, Colorado was the site of another successful hack program in which four young were released. This was the first year the site was in operation. A fourtbruck site was established in Rio Grande National Forest and 4 young were released. Golden eagle attacks became an almost daily occurrence �33 and two of the young became separated from the others and never did return to the hack site. They did come to a feeding board and one successfully reached independence, the other apparently was killed by the eagles. The two young which remained at ,the hack box survived to independence. In all, 17 young peregrines were placed at hack sites, and 11 survived independence. Prepared by: ~. Q. ~ Gerald R. Craig \ Wildlife Researcher C 12 were released �34 JOB PROGRESS REPORT State of COLORADO --~~~~~--------------- Project No. ~S~E~-~3~-_3~ _ Endangered Wildlife Investigations Work Plan No. ~I~I~ _ Job No. Job Title: Peregrine Falcon Captive Maintenance Period Covered: Personnel: 4 July 1, 1979- February 28, 1981 W. Burnham, D. Konkel, W. Heinrich, J. Hoolihan, J. Trotta, C. Sandfort, The Peregrine Fund Inc., G. Craig, Colorado Division of Wildlife. ABSTRACT In 1980, 35 adult anatum peregrine falcons were maintained at 1424 N.E. Frontage Road, Fort Collins, Colorado. They were fed daily and maintained in healthy condition. No losses or injury were incurred during the period. All falcons were compatible when held in pairs and no excessive aggression was observed by any individual. They were fed a diet of quail and five to six week old cockerels on a daily basis. The security of the facility was not violated and no unusual events occurred. �35 PEREGRINE William FALCON CAPTIVE MAINTENANCE Burnham and Gerald R. Craig P. N. OBJECTIVE The objective of this program falcons in captivity. is to maintain 35 adult anatum peregrine SEGMENT OBJECTIVES 1. Annually contract with The Peregrine Fund, Inc. to maintain adult anatum peregrine falcons in captivity. 2. Prepare an annual report of maintenance contract. efforts associated 35 with the PROCEDURES 1. Make daily observations of adults, monitoring for compatibility, stress, behavior abnormalities, physical difficulties, and any unusual actions. 2. Provide freshly thawed, whole Coturnix quail and five to six week old cockerels on a daily basis in quantities so that falcons will be maintained in prime condition. 3. Periodically, depending upon the weather, bath and drinking will be changed so fresh water is constantly available. 4. Periodically good hygenic 5. When falcon chambers are to be cleaned, each falcon should be caught and examined for physical difficulties. If talons or the beak are overgrown they should be trimmed to appropriate length. 6. The falcons will be maintained at a location where good security possible and disturbance can be minimized. pans clean chambers where falcons will be held, maintaining conditions. METHODS Daily Observations. Methods in the procedures enumerated is AND MATERIALS for daily observations above. have been described Falcon Food. Quail were raised until eight weeks of age, killed with CO2, cooled, and quick frozen for later thawing and feeding. Egg laying adult Coturnix quail were maintained in a Petersime battery brooder where eggs �36 were collected daily. The egg laying quail were fed and watered daily and were maintained on a 16 hour light photo period. Eggs were collected and held at 550 C. in a humid air egg storage room until there were enough to set in the incubator. The eggs were fumigated with potassium permanganate and Formalin before setting. Standard hygenic procedures were followed for the incubator and hatcher to prevent disease and maximize the hatch. Chicks were removed from the hatcher and transferred to brooder facilities where they were held on wood shavings with overhead radiant heat until three weeks of age. The auail were then moved onto wire where they remained until collection and freezing. The chickens were reared under similar conditions, however, they were never moved from shavings to wire. All food itemswere reared on nonmedicated feed. Drinking Water. }1ethods for changing drinking and bathing water have been described in Procedure 3 above. Clean Chambers. Chambers were cleaned after first removing the adult falcons. All perches were wire hrushed to remove fecal material, feathers, and prey remains, then they were washed with a high presure hose. When the perches had dried they were sprayed with Nolvasan disinfectant. Nesting ledges were raked and in some cases the pea gravel nesting substrate was changed. Carpeted areas were brushed and washed. The total ledge was then washed and disinfected. All bars, panels, glass, lights, perches, nest ledges, and supports were examined and renailed or screwed if loose. One-way glass windows were washed and light bulbs checked. The floor was then raked several times and all prey remains, fecal material, and feathers were removed. The floor was then washed and disinfected before refilling. The chamber was allowed to dry for 24 hours before falcons were returned. Examination of falcons. When falcons were caught they were "hooded". The talons were immediately dulled by clipping the points. This was done so the feet were not punctured, which increases chances for infection and bumble foot. The talons were clipped just shorter than normal length. The beak was examined and if longer than what; is considered normal it was clipped back with large nail clippers. A small fine file was then used to smooth and' shape the beak. Security. Falcons were held in chambers with barred fronts with external wire covering. The chambers were within a steel sheeted post frame barn. Each chamber had a seperate entrance door which securely locked and the doors are locked on an inner hallway with large doors that were securely locked when workers were not in the building. All barns were surrounded by an open grass area within a seven fOQt chain link fence which had barbed wire on top. Listening devices were located in the hallway of the post frame barns so monitoring was possible at night. Guard dogs were maintained in the compound to prevent intrusion. �37 RESULTS AND DISCUSSION In 1980, 35 adult anatum peregrine falcons were maintained at 1424 N.E. Frontage Road, Fort Collins, Colorado. They were fed daily and maintained in a good healthy condition. No losses or injuries were incurred during the period. All falcons were compatible when held in pairs. No excessive aggression was observed by any individual. They were fed a diet of quail and five to six week old cockerels on a daily basis. The security of the facility was not violated. No unusual events occurred. Prepared by: (?.. R. G~ Gerald R. Craig Wildlife Researcher C �38 JOB PROGRESS REPORT State of COLORADO ~~~~~----------------_ Pro j ect No . ..::.S.=E_-..::.3_-..::.3 _ Endangered Wildlife Hork Plan No. II Endangered Job No. 6 Job Title: Development Birds of a Preservation of Prairie Period Covered: Personnel: 1 July 1979 - 28 February Program Investigations for Three Species Grouse 1981 R. Calderon, L.O. Cordova, L. Crooks, B. Fittante, W.D. Graul D. Homan, R. Kahn, C. Loeffler, G.C. Miller, T.E. Olson, J. Slater, L. Stelter, C. ~rJagner,T. ~lashington, D. Weyerman. ABSTRACT Colorado's prairie grouse were studied between July 1979 and February 1981. Numbers of known lesser prairie chicken leks have generally increased since 1959. A sharp-tailed grouse was documented on the Division of Wildlife's South Platte Management Area, although the presence of a breeding population has not been verified. Greater prairie chicken numbers are unknown, but they have been located in areas from which they had not been reported previously. Most greater prairie chickens in Colorado have been documented from sandy soil sites in sandsage-bluestem prairie vegetation. The Tamarack restoration area approximates some characteristics of occupied prairie grouse range in Colorado, but needs to be seeded with additional tall grasses. Preservation of prairie grouse in Colorado appears to depend upon maintenance of insular populations. �39 DEVELOPMENT OF A PRESERVATION PROGRAM FOR THREE SPECIES OF PRAIRIE GROUSE Gary C. Miller P.N. OBJECTIVES This program is designed to ascertain current population levels and distributions of all 3 species of prairie grouse in Colorado, and to formulate recovery plans. SEGMENT OBJECTIVES 1. Determine the present population levels and distributions three species of prairie grouse in Colorado. 2. Identify key habitat nesting areas. 3. Rehabilitate 3,755 acres of historic Greater Prairie Chicken habitat by 1985 on DOW's South Platte Wildlife Management area to a condition capable of sustaining a restored population of three hundred birds. use areas--booming grounds, winter of all concentrations, INTRODUCTION Three species of grouse (Order: Galliformes, Family: Tetraonidae) once ranged over most of Colorado's eastern plains. Each has undergone severe range reduction since the early to mid-1900's. In each case, this range reduction has accompanied changes in land use (Evans 1964, Rogers 1969, Hoffman 1973). The Colorado Wildlife Commission has classified greater prairie chickens and plains sharp-tailed grouse as endangered species, and lesser prairie chickens as a threatened species in Colorado. This report presents preliminary results of work to update previous information and identifies those activities showing promise for attaining a more secure status for Colorado's prairie grouse. STUDY AREAS Lesser pralrle chickens were studied in Baca County in southeastern Colorado, primarily on lands administered by the USDA - Forest Service, Comanche National Grassland. Occupied range was characterized by well drained, sandy soils with a flat to gently rolling topography. Vegetation consisted mostly of perennial grasses such as blue grama, (Bouteloua gracilis), buffalo grass (Buchloe dactyloides), side oats grama (Bouteloua curtipendula), and three-awn grass (Aristida longiseta). Sand sagebrush (Artemisia filifolia) occurred in dense growth in disturbed areas. �40 Other vegetation, such as yucca (Yucca glauca), barrel cactus (Echinocactus sp.), prickly pear cactus (Opuntia sp.), and several varieties of annual weeds and forbs were present. An occasional cottonwood (Populus sp.) or hackberry tree (Celtis sp.) occurred along low lying moist areas. Greater prairie chickens were studied primarily in Yuma County, although some field activities extended into eastern Washington and southern Phillips counties. Vegetation studies were conducted on the Division of Wildlife's South Platte Wildlife Management Area in Logan County. METHODS In April and May, 1980, counts of lesser prairie chickens were made at leks, with each lek visited at least 3 times. Population information for greater prairie chickens was gathered using line transect sampling techniques (Burnham et al. 1980). Flock counts of greater prairie chickens were made from October through December in 1979 and 1980. Reports of prairie grouse in areas outside their known distributional limits were used to update distribution maps only when documented by a specimen or photograph. For the line transect sampling of greater pralrle chickens, a sample frame of 2920 possible transects C10 per mi2 in a 292 mi2 area known to contain prairie chickens), oriented north and south, was delineated. Transects to be sampled were selected randomly. Two observers on horseback rode 20m on either side of the transect dragging a rope between them to meet the assumption of detection of all birds on the transect. A trained pointing dog was also used, and generally was kept on alternating sides of each transect. Key habitat use areas were identified in conjunction with population surveys. Greater prairie chicken microhabitat data were collected at roosting sites from which birds were flushed during line transect sampling, following the 1m x 1m frame technique of Graul (1978). Recent and historical records of greater prairie chicken occurrence were compiled and classified with respect to the soil and vegetation type of the site. Vegetation data were collected from sites representing 6 combinations of vegetation type arid range management practices, following Graul's (1978) technique. Between April and July, 1980, vegetation was sampled on the portion of the Colorado Division of Wildlife's South Platte Wildlife Management Area south of highway 1-76. A sampling frame of 1515 100m x 100m plots, oriented in the cardinal directions was delineated, and 31 (2%) were randomly selected for sampling, surveyed, and marked with steel fence posts. Each plot was subdivided into 4 SOm x SOm (0.25 ha) quadrats and permanently marked. Vegetation sampling within each quadrat was conducted along 5 permanently marked SOm transects (randomly selected from a frame of 50 at 1m intervals) oriented east and west. �41 Vegetation was characterized using cover board (Jones 1968), density pole (Robel et al. 1970), and 1m x 1m frame (Graul 1978) techniques at 5m intervals along each transect, beginning 5m from the outside edge of the quadrat. Further measurements were obtained using a modification of the technique described by Parker and Harris (1959). A 3/4 inch loop was placed on the ground on the south edge of the 50m transect tape at O.Sm intervals beginning 0.5m from the outside edge of the quadrat. Each "hit" was recorded by species if the root crown was more than 50% within the loop. Plant material other than root crowns within the loop or the 3/4 inch column above the loop were classified as either current or residual (from previous yearst growth) litter if more than 50% of the surface within the loop was obstructed by vegetation. All other samples were classified as bare ground. RESULTS AND DISCUSSION Plains Sharp-tailed Grouse Six reports of sharp-tailed grouse on the South Platte Management Area were received during November and December, 1980. The first reported sighting, on 9 November, was of 2 birds. One bird was reported in each of 3 sightings on 12, 13, and 15 December. "Four or five" were reportedly seen on 14 December, and "14 or 15 sharp tails or prairie chickens" were reported for 23 December. Sharp tail presence was verified by photographs of a single bird taken by L. Crooks, District Hildlife Manager, in December. Lesser Prairie Chicken Most lesser prairie chickens in Colorado appear to be part of populations that range into Oklahoma and Kansas. Two somewhat; isolated populations appear to exist entirely within the state, separated from other known populations by 8 to 16km (Fig. 1). During April High counts ranged from County were and May of 1980, 17 active leks in Baca County were located. of cocks on all leks totaled 171 (Table 1). Birds per lek 1 to 19, with an average of 10 per lek. Leks in Prowers not counted in 1980. Four "new" leks were loc!ated this year (numbers 25, 26, 27, and 28). Leks 25 and 26 were in the vicinity of number 17, which was not active. Leks 27 and 28 were in the same general area as numbers 1 and 2, which have lost birds or have completely been abandoned recently. Lek number 2, used as a public viewing area for a number of years, has shown a decline in numbers each year since 1976 .. One male was seen on one occasion during the 1980 study. An effort will be made to attract birds back to this site in 1981, using recorded calls of male prairie chickens. If this is not successful, one of the other leks in the area may be selected for a public viewing area. �\ ~ u 1 /;h " 'J') ,; ( !Jj c )'-1, .. "~ I .._ .•.... _" II "'1 t ,,I}'! " .~. ·'' c.·'7,.·...... , '• ~ .____j I I I A'" . .~.:: '" t I -nsn ... _ 0 T . __ ." U-~ ,! E R 0 COMANC~ A ' < •• l~oon".. ,;" II. ,)~\,~" I, NATIONAL _, ~~., ,j~ ~)) ! .... _, ,,_ , , I(i"'p ! ~ .,",' . R I 5' /' N I >:1J· 0 1 - • ,;/ til <11 U) c <11 ~ .p.. N ~ Fig 1. e ••~b" Distribution of the lesser prairie chicken in Colorado 1980. c !I 10 l::r::t:.:.c::r.:::t:::.=.:..:: u.••, Itl ~fJ ...;.t-:=.:...:_ -; ._1.'.::-.::--_-. ".• I .:..;::~ "., !~(~ In~'I\ II 1111 I, ae z :__,_., �43 Table 1. Results of lesser prairie chicken lek surveys in Colorado, 1959-1980. TOTAL BIRDS (HIGH COUNTS) YEAR GROUNDS COUNTED 1959 2 11 1960 6 39 1961 11 84 1962 11 116 1963 10 125 1964 No Data Available 1965 No Data Available 1966 No Data Available 1967 1 1968 No Data Available 1969 No Data Available 1970 3 1971 3 1972 7 82 1973 7 65 1974 11 107 1975 13 151 1976 15 158 1977 17 178 1978 16 156 19 175 17 171 1/ 197~ 2/ 1980 - 6 42 . 1/ Prowers Co.= 4 leks, 48 birds; 1/ Counts made only in Baca Co. 37 Baca Co. 15 leks, 127 birds. �44 Counts in 1980 ;ndicated that of the known active leks in Baca County, nine were new ct increased in size, two remained stable, and eleven decreased in size or were abandoned since 1979. Most of the leks which showed a decline or were abandoned were the smaller and apparently less stable ones. From 1979 to 1980 the total number of known active leks in Baca County increased from 15 to 17, and high counts of males on all grounds totaled 171 in 1980, as compared to 127 in 1979. Six lek sites (8, 9, 10, 11, 16, and 19) have been inactive for two or more consecutive years. Greater Prairie Chicken Most of the area in Colorado known to contain greater pralrle chickens is contiguous with occupied range in Nebraska (Fig. 2). An apparently isolated population exists on the Washington-Yuma county line. The existence of a greater prairie chicken population near the Arikaree River in south-central Yuma County was confirmed with photographs of birds on a lek taken in 1979 by T.E.Olson, and breast feathers found at the same site in 1980. A young-of-the-year greater prairie chicken was found dead by D. Weyerman, District Wildlife Manager, near Hale in extreme southeastern Yuma County in October, 1980, adding credence to several reports of prairie chickens in that area in recent years. Data were tabulated on the soils and vegetation type of 95 sections of land on which prairie chicken have been documented since 1963 (Table 2). Most prairie chickens have been known from sandy soil sites in KUchler's (1964) sandsage-bluestem prairie vegetation type, although other soils and vegetation types do occur within the overall range of prairie chickens in Colorado. Encounters with pralrle chickens during line transect sampling were too few and variable for a meaningful population estimate, with 29 birds flushed in 7 encounters over 102 km of transect in 1979 and 4 birds flushed in 3 encounters over 64 km of transect in 1980. Data from 1979 indicated that sampling should be concentrated in tallgrass-sandsage pastures (Table 3). This was done in 1980, but early fall flocking, not seen in 1979, may have caused increased clustering and decreased detection probability. Flocks of prairie chicken flying to feed in croplands were counted in 1979 and 1980 (Table 4). Flocks observed at distances of 8 km or greater from each other were assumed to be different birds (Hamerstrom and Hamerstrom 1973). In 1979, no flocks were seen until 15 November, roughly the peak of the corn harvest and none were seen again until 30 November following the first major snowfall of the season. All flocks fed in picked corn fields. In 1980, flocks were first seen on 7 October, feeding on alfalfa. Flocks ranging in number from 24 to 112 were seen on alfalfa and green rye fields into the 3rd week of November. Flocks were not seen in corn fields until the 3rd week of November, although most corn had been harvested in the area by mid-October. �LEGEND --. ••• ~ Presumed historic western and southern limits (Aldrich and Duvall 1955) • Q 0> Distribution •• in 1962-1963 (Evans COL 0 R ADO l"li'fM{f{ h'{L" tOCA,"" 1964). Distribution * J( in 1981. Tamarack restoration c c "",,,, 1' .• ' ~ I. /1"'; area. AN rrA ;/'fJ"'~/OH S')"CI) -'---, "r.#IO I I ••.• . •• CA • • • •. ~"l.~ ... ---=A. ~ Irlf (J.,-i'S;:N LlHCO"t ••• { JA +:-- VI C"UCNNt /VI ° • 10 II. C,U 1'(1((1'0 <, • C~()I{,(y . .....__ S~CII"("'~ J •• ,. M" ""-<- PO(OI1!l f't.<C,..'T{ Fig 2. cur rt /l ~/I. I'~C"'O." [jt"" "-""''' Il_ tLlNU Past and present distribution I~ t,«A of greater prairie chickens -5nColorado, and Tamarack restoration area. �46 Table 2. Soils and potential vegetation of 95 sections of land on which prairie chickens have been documented since 1963. Number of Sections SOILS Sandy Loamy Clay POTENTIAL 90 4 1 VEGETATION Sands age - Bluestem Prairie Grama - Buffalograss Northern Floodplain forest Wheatgrass - Needlegrass Bluestem - Grama Prairie 85 6 4 o o �Table 3. Line transect characteristics and prairie chicken encounters by vegetation types in Yuma County, Colorado, 1979. PASTURE Tallgrass and sandsage 47.5 Percentage of transect length Shortgrass and Yucca 20.3 9.4 OTHER CROPS FALLOW 1.6 2.1 0 0 Abandoned Agricultural 3.9 15.2 5 0 1 0 1 26 - 2 - 1 Number of encounters Number of prairie chickens Tallgrass with sandsage control CORN .t:- -...J Table 4. Counts of greater prairie chickens flying into croplands, Oct - Dec, 1979 and 1980. Number of Observations -------- Number of areas Number of areas with birds Sum of high counts from each area -_-------------------------------------------- 1979 18 9 5 216 1980 39 13 5 255 �48 Roosting sites of greater prairie chickens, located during line transect sampling, were identified by the presence of forms with fresh droppings. In general, roost sites were characterized by higher tallgrass and sandsage densities and a lower percentage of bare ground than sites Sm from the roosts (Table S). Vegetation data collected from sites representing 6 combinations of vegetation type and range management practices showed that the Tamarack restoration area exhibited the densest residual vegetation (least amount of bare ground) of all sites (Table 6). The Tamarack had intermediate values of sandsage and tallgrass frequencies when compared with the other sites, which were in occupied prairie chicken range. This may have been due to the plant species composition of the areas. Common grasses on sites in occupied range included the bluestems, switchgrass (Panicum virgatum), and sand love grass (Eragrostis trichodes), which were rarely encountered on the Tamarack. SUMMARY AND RECOMMENDATIONS 1. No adequate technique for ascertaining prairie grouse densities over their full range in Colorado has been developed. Counts of numbers of leks are useful for monitoring distributions and may provide longterm indices of population trends (Cannon and Knopf in press), and should continue as management activities. Flock counts are useful for ascertaining distributions, but should not be relied upon for population monitoring unless a large number of concentration areas are monitored at one time. Line transect sampling is subject to many variables, but may have applicability in small, relatively homogenous vegetation types. 2. The Tamarack restoration area continues to be the only identified option for the preservation of greater prairie chickens and/or plains sharp-tailed grouse. Although the area approximates some conditions found in occupied grouse range, it is deficient in others, such as plant species composition. Since most attempts to reintroduce prairie grouse have failed in other areas (Kruse 1973), no reintroduction attempt should be made until conditions on the Tamarack more closely approximate those of occupied range. Bluestems, switchgrass, indiangrass (Sorghastrum nutans), and sand lovegrass should be seeded to the Tamarack grasslands before a reintroduction. Additionally, the Tamarack restoration area and surrounding grasslands should be inventoried intensively for possible leks, in light of the recent record of prairie grouse in the area. 3. Fragmentation of prairie grouse habitats is not unique to Colorado -it is characteristic of prairie grouse habitats throughout the United States (Crawford 1980, Klopatek et al. 1979, Miller and Graul 1980, Westemeier 1980). Preservation of prairie grouse in Colorado will depend upon maintenance of populations in relatively small, isolated patches of prairie. Research efforts should be directed toward ascertaining the size and distribution of patches, and the population characteristics needed to reintroduce, establish. and maintain prairie grouse populations with a known probability of persistence. Areas on the Comanche National Grassland and the Tamarack restoration area appear suited to this endeavor. �49 Table 5. Comparison of 1m x 1m frame analysis of greater pra~r~e chicken day roosts and sites Sm from roosts in the cardinal directions. Roost N=13 sm from roost N=s2 Sandsage Plants/m2 x SD 1.77 1.09 0.79 0.72 2.15 2.54 1.54 1.86 18 12 41 20 Tallgrass Plants/m 2 x SD Bare Ground % x SD �Table 6. Characteristics of sites selected as representative of 6 combinations of vegetation type and range management practices. Site II Sandsage - Tallgrass Moderate Intensive Moderate Grazing & Sandsage Grazing Grazing Control 5S(N=6) 6(N=6) 2(N=10) 3(N=15) Shrub Mean II/transect Range 7.0 2-12 9.5 5-15 17.4 0-28 Mean Height(cm) Range 47 35-59 61 50-8.2 42 38-47 Range Type Range Practice Moderate Grazing Bluestem Swale Moderate Grazing & Reseeded Ungrazed 2 seasons 4(N=10) 5N(N=6) 5.9 0-16 2.4 0-5 3.7 0-9 4.4 0-18 42 30-54 34 20-45 32 26-50 45 15-75 Tamarack (N=40) V'I 0 Tallgrass Mean II/transect Range 43.5 23-62 34.7 25-46 63.5 41-120 53.4 23-81 59.7 32-92 91.8 74-106 64.9 22-116 Mean Height(cm) Range 74 64-85 64 56-82 86 73-96 84 70-97 81 76-85 100 85-120 54 10-125 Bare Ground Mean % per transect Range 46 27-55 62 57-66 46 36-62 48 28-65 44 30-62 53 46-61 39 7-65 �51 LITERATURE CITED Burnham, K.P., D.R. Anderson, and J.L. Laake, 1980. Estimation of density from line transect sampling of biological populations. Wildlife Monograph 72. 202pp. Cannon, R.W. and F.L. Knopf, in press. Lek numbers as a trend index to prairie chicken populations. J. Wildl. Manage. Crawford, J.A. 1980. Status, problems, and research needs of the lesser prairie chicken. Pages 1-7 in P.A. Vohs and F.L. Knopf, eds. Proc. Prairie Grouse Symposium. Oklahoma State University Publishing and Printing, Stillwater, OK. Evans, K.E. 1964. Habitat evaluation Colorado. M.S. Thesis, Colorado of the greater pralrle chicken in State Univ., Ft. Collins. 98pp. Graul, W.D. 1978. A technique for evaluating greater prairie chicken habitat in Colorado. Colo. Div. Wildlife unpubl. rep. 3pp. Hamerstrom, F., and F. Hamerstrom. 1973. The prairie chicken in Wisconsin: highlights of a 22-year study of counts, behavior, movements, turnover, and habitat. Wis. Dep. Nat. Resour. Tech. Bull. 64. 52pp. Hoffman, D.M. 1963. The lesser prairie Manage. 27: 726-732. Jones, R.E. 1968. A board to measure J. Wildl. Manage. 32: 28-31. chicken in Colorado. cover used by prairie J. Wildl. grouse. Klopatek, J.M., R. J. Olson, C.J. Emerson, and J.L. Joness. 1979. Land-use conflicts with natural vegetation in the United States. Environ. Conserve 6(3): 191-199. Kruse, A.D. 1973. Prairie chicken restoration projects. Pages 40-46 in W.D. Svedarsky and T. Wolfe, eds. Proc. Conf. on the prairie chickeninMinnesota, Univ. of Minnesota, Crookston. KUchler, A.W. 1964. Potential natural vegetation of the conterminous United States. Am. Geogr. Soc. Spec. Publ. 36. 39pp. Miller, G.C., and W.D. Graul. 1980. Status of sharp-tailed grouse in North America. Pages 18-28 in P.A. Vohs and F.L. Knopf, ed~ Proc. Prairie Grouse Symposium. Oklahoma State University Publishing and Printing, Stillwater, OK. Parker, K.W., and R.W. Harris. 1959. The three-step method for measuring condition and trend of forest ranges; a resume of its history, development and use. Pages 55-69 in U.S. Forest Service techniques and methods of measuring understory vegetation. Southern and Southeast For. Exp. Sta. �52 Robel, R.J., J.N. Briggs, J.J. Cebula, N.J. Silvy, C.E. Viers, and P.G. Watt. 1970. Greater prairie chicken ranges, movements, and habitat usage in Kansas. J. Wildl. Manage. 34: 286-306. Rogers, G.E. 1969. The sharp-tailed grouse in Colorado. Colorado Game, Fish, and Parks Div. Tech. Publ. 23. 94pp. Westemeier, R.L. 1980. Greater prairie chicken status and management 1968-1979. Pages 8-17 in P.A. Vohs and F.L. Knopf, eds. Proc. Prairie Grouse Symposium. Oklahoma State University Publishing and Printing, Stillwater, OK. Prepared by GOJcv ,-c. Pl1 Gary C. Miller Wildlife Researcher 014 �53 JOB FINAL REPORT State of COLO~~O Project No. ~S~E~-~3_-~3 Work Plan No. III Endangered Job Title Mammals: Job No. Lynx and Wolverine Period Covered: Personnel: _ ~3 _ Verification 1 July 1978 through 30 June 1980 S. Bissell, R. Calderon, M. Foster, J. Garcia, W. Graul, J. Halfpenny, C. Loeffler, L. Lofgren, T. Lytle, L. Marlow, G.C. Miller, D. Nead, P. Shearwood, J. Torres, T. Valenta, T. Washington. ABSTRACT SurVeys were conducted to ascertain the past and present status of Canada lynx (Lynx canadensis) and wolverine (Gulo gulo luscus) in Colorado. Lynx continue to exist in at least 4 of the 8 counties of Colorado from which they have been verified. No irrefutable evidence of current wolverine existence was found, although circumstantial evidence of wolverines in several areas was obtained. Management of spruce (Picea sp.)-fir (Abies lasiocarpa) stands and snowshoe hare (Lepus americanus) should benefit lynx, and protection and management of ungulate winter ranges should benefit wolverines if present. �54 LYNX AND WOLVERINE VERIFICATION Gary C. Miller P. N. OBJECTIVES This program was designed to assess the past and present status of lynx and wolverine in Colorado. Information about the requirements of both species was sought, and threats to their continued existence were to be identified. SEGMENT 1. Ascertain 2. Verify 3. Develop levels. 4. Identify 5. Determine the historical status of lynx and wolverine the current existence techniques of lynx and wolverine in Colorado. in Colorado. to detect both species and estimate key habitat further OBJECTIVES population areas. research needs and suggest management guidelines. INTRODUCTION Lynx and wolverine are mammals associated with boreal and, in the case of wolverine, tundra areas. Both species reach their southernmost distributional limits in Colorado. Neither species has ever been considered abundant in the state. The Colorado Wildlife Commission has classified both species as state endangered. Detailed accounts of the study, including study areas and methods are given in Appendix A (History and Status of Canada Lynx in Colorado) and Appendix B (History and Status of T.volverine in Colorado) and will not be repeated here. A summary 'of the findings follows. SUMMARY Canada Lynx 1. Lynx have been verified in 8 counties of Colorado and reported without verification in 10 counties. The most recent specimens from Gunnison, Summit, Conejos and Montrose counties were taken prior to 1926; the most recent specimens from Eagle, Pitkin, Lake, and Clear Creek counties were taken between 1969 and 1974. 2. The elevational distribution of lynx in Colorado, appears to be above 2,730 m. past and present, �55 3. The distribution of lynx in Colorado is probably discontinuous and currently includes: Fryingpan River drainage upstream from Meredith in Eagle and Pitkin counties; Vail area in Eagle County; southeast of Leadville in Lake County; the Guanella Pass - Mount Evans area of Clear Creek County. Lynx may also exist in other areas of those 4 counties and also in Summit, Grand, and Park counties where spruce-fir associations occur and snowshoe hare are abundant. 4. Lynx appear to be members of a spruce-fir - snowshoe hare association. Lynx tracks found during the study were usually in or near spruce-fir stands with large boulder outcroppings nearby, and either on north facing slopes or on the sides of narrow north - facing valleys. No present or potential threats to lynx, their habitat, or their probable prey base (snowshoe hare) were identified during the study. 5. Lynx never were known to be abundant in Colorado, and no evidence of an historical decline in numbers was found. Their current status as a state endangered species makes the gathering of population and distribution data extremely difficult. Wolverine 1. Wolverine accounts exist for 20 Colorado counties, but the species has been verified only in Summit, Clear Creek, Ouray, and Montrose Gunnison counties, excluding imported wolverines known from Douglas and Pitkin - Gunnison counties. 2. A total of 271 reports of wolverine observations were evaluated during the study. No irrefutable evidence of a viable wolverine population was found during the study, although several reports provided some circumstantial evidence of wolverine existence in the state. 3. Circumstantial evidence accumulated during the study indicated wolverines in winter may be associated with ungulate concentrations. If so, protection and management of ungulate winter range could benefit wolverines, if they are present. 4. The chance of obtaining irrefutable evidence of a viable wolverine population is slight because of the difficulty in detecting the species and because their endangered status, under present Colorado statutes, makes it unlikely that authorities will be notified in the event a wolverine is taken even accidentally. Prepared by: ~~ C."rn''-~ Gary c:Mlller Wildlife Researcher �56 APPENDIX A HISTORY AND STATUS OF CANADA LYNX IN COLORADO James C. Halfpenny The Canada Lynx inhabits boreal snowshoe hare and Gary C. Miller (Lynx canadensis) is a 7 to 20 kg carnivore forests of. North America. (Lepus americanus) Lynx dependency that upon (Brand et al. 1976). as prey is well-known but rodents and birds are also taken. In the field, lynx may be differentiated (Lyn~ rufus) by a completely from the similar bobcat black tail tip on lynx as opposed of bobcats. belief, Lynx have 4 mammae; light coloration (Hall and Kelson Colorado 1959). has been reviewed confusion distributional because Contrary by Armstrong literature Historical accounts Division lynx from bobcats until lynx were first given protection in Colorado. species by the Colorado to prevent in the state, the Colorado Division conducted information to evaluate Wildlife of Wildlife of Wildlife 1970, when Commission. of endangered initiated a status of lynx in Colorado. was sought, and additional the species' are also In 1973, lynx were classified the extinction study in 1978 to assess the past and present All available and of lynx in since 1939 did not differentiate species for lynx. true lynx and large pale bobcats, known as "lynx." As part of its obligation features (1972), who also noted apparent records kept by the Colorado as a state endangered to common limits in Colorado The historical in early records between which were sometimes incomplete bobcats have 6. and ear tufts are not diagnostic Lynx reach their southern Utah to black only on the top of the tail current status. field work was In addition, �57 information about the species' requirements was collected to identify present and potential study was conducted under Federal Aid Endangered and used threats to its existence. The Species Project SE-3. STUDY AREAS The overall conducted study was statewide in scope. Field activities in Eagle, Pitkin, Lake, and Clear Creek counties. study sites were centered around Vail Pass, Berthoud Mount Evans, and the Fryingpan communities of the study sites included Abies lasiocarpa) contorta), River upstream with stands of spruce-fir, aspen communities are abundant (Populus tremuloides) (Costello 1964). Pass, Leadville, lodgepole pine (Pinus and open parks, and also alpine On the Fryingpan are present Vegetation (Picea sp. - River, snowshoe hare (Dolbeer and Clark 1975) and white-tailed (Lagopus leucurus) Specific from Meredith. spruce-fir were ptarmigan (Braun et al. 1976). METHODS A literature in Colorado. Over 3,000 requests were distributed outfitters, to District private People believed specimens review was conducted knowledgeable were sought in museum Between January specifically for information Wildlife individuals, Managers, about lynx sightings taxidermists, trappers, and federal and state agencies. about lynx were interviewed. and private Lynx collections. and April of 1979 and 1980, 30 days were spent searching for lynx and lynx tracks using, at various times, skis, snowmobiles, and hounds in the study sites above an elevation greater to identify all reports of lynx trained to cats. All known trails of 2400m were traversed. than 85mm wide, with a straddle Tracks less than 180mm and stride �58 less than 430mm which sank only shallowly into the snow (a subjective judgement based on snow condition) were classified as lynx tracks (Murie 1963). Where lynx tracks were encountered, the topography, vegetation, and abundance of snowshoe hare were described. RESULTS Lynx have been reported from 18 Colorado counties, dating from the 1870's to the present (Table 1). For 10 of these counties, however, Table 1. Colorado counties from which lynx have been verified or reported. Underlined sources are those which verified lynx occurrence. SOURCES Conejos Summit Carter colI. DMNH. Carter colI. DMNH; Warren (1942), Carter notes DMNH Burns colI. DMNH T.E. Jacques colI. 246371 USNM; Terrell (1971) Cary (1911); Terrell (1971); Shearwood pers. colI.; this study Purkat pers. colI. Cary (1911); Seton (1929); Terrell (1971); Colo. DOW colI.; this study Carter notes DMNH; Jefferson County Outdoor School; this study late 19th century late 19th century Allen (1874); Young (1958 Cary (1911); this study Cary (1911) Cary (1911) Cary (1911) Cary (1911) Warren (1906) Cary (1911) Cary (1911) Cary (1911) no no no no no no no no no no Montrose Gunnison Pitkin Lake Eagle Clear Creek :>-t H Z 0 C/) E-! P::: 0 p.., lJ;:l P::: DATE OF MOST RECENT SPECIMEN COUNTY Park Garfield Routt Jackson Rio Blanco Grand Custer San Juan La Plata Archuleta 1915 1925 1969 1969 1974 1972 specimen specimen specimen specimen specimen specimen specimen specimen specimen specimen �59 specimens are lacking and most sightings specimens from 4 counties taken prior to 1926. four counties (Gunnison, pre-date 1911. The most recent Summit, Conejos, Montrose) The most recent specimens were from the remaining (Eagle, Pitkin, Lake, and Clear Creek) were taken between 1969 and 1974. Five lynx specimens from Colorado, located during the study. at the Denver Museum previously One specimen unreported, in the Edwin Carter collection of Natural History (DMNH) was undated but taken in the late 19th century at Cumbres Pass, Conejos County. specimen and collected this information C. Lincoln. by Burns, 23 May 1915. bears the collection The collection locality in Montrose County, and the elevation been 2,400m (8,000 feet) or lower. Anton Purkat of Leadville he trapped in December, seen by Mr. Purkat The specimen unreported Creek County. apparently was the La Sal Mountains, at which it was taken must have possesses 1969 southeast a fully mounted of Leadville at that time led him to believe lynx which (Lake Co.~ 2 lynx were illegally School in Evergreen, trapped near Vail and the other, a fully mounted Pass, Clear headquarters, Denver. Another of the In 1973-74, One escaped is at the Colorado Division recent specimen, Terrell (1971), was shot by Pete Shearwood River Colorado. (Eagle Co.). specimen, Tracks 2 lynx were present. The fully mounted ·specimen is in the possession County Outdoor the Fryingpan tag with number and initials of Frederick In 1972, a lynx was trapped by Bill Buxton at Guanella Wildlife Another (skin only) at the DMNH was from the "La Salle Mountains, Colorado" Jefferson were reported by of Eagle, Colorado, (Pitkin Co.) and is presently of in 1969 on held by Mr. Shearwood. �60 Recent (1969 or later) lynx specimens from Colorado, then, number 4 and have been taken from 4 counties. Additional found during records the study. of lynx, for which no verification In Edwin Carter's were notes of a lynx from Breckenridge September taxidermy exists, were records (DMNH) in Summit Co. (female, dated 18 1878), and one from Soda Gulch in Clear Creek Co. (male, dated 30 January 1878). Carter's notes also contained records of 3 other lynx, taken in 1883, 1889, and 1897, but gave no location. Additional reports received during the study may have been lynx. Charles Co l.bv of Gilman, Colorado described tail of 3 animals He reported indicated Co. In the fall of 1969, Durbin McIlnay of Gilman; of 3,000m. Mr. McIlnay's description (Eagle Co.) of the animal it was a lynx. Further indications (1971) reported of lynx have been based on tracks. seeing lynx tracks in 1969 on the Roaring (Pitkin Co.). Three sets of tracks matching lynx tracks were found during (Eagle and Pitkin Counties) Chicago (1 in 1930, 2 in 1936). shot an animal on the south side of Vail Mountain at an elevation Aspen and a friend black tip on the the animals were taken at Golden Park on the upper Homestake Creek, Garfield Colorado trapped by himself a completely Creek drainage, at elevations above 2,730m Tracks were usually Fork above the description 1979 field work on the Fryingpan of River and 5 sets were found in 1980 on the West near Mount Evans All tracks identified Terrell (Clear Creek Co.). as lynx tracks during the study were found (9,000 ft.) and near large boulder associated with spruce-fir stands, facing slopes or on the sides of narrow north-facing outcroppings. either on north- valleys. �61 In addition to the lynx tracks on the West Chicago Creek drainage, 28 sets of snowshoe hare tracks were found while traversing of transects. stands Hare tracks occurred (mean inter-tree distance found in more open spruce-fir tree distance greater most commonly 1,234m in dense spruce-fir less than 5m); tracks were seldom or spruce-fir-aspen stands (mean inter- than 7m). DISCUSSION Failure to differentiate are the main reasons in Colorado evaluate an the efforts analysis reported by Cary an assessment is impossible. over time yields a Analyzing weak indication involved of lynx reports (1911). the numbers are: 1870-1889 1950-1969 If Caryts some parts of Colorado (1911) exceptional occurred periods (4); 1930-1949 (4); lynx as common in of lynx in Colorado may yield of their past and present status documentation in 10 other counties, 1972), in the state. counties, 3km north of the New Mexico border the southernmost by 20 year (Allen 1874, Coues 1879 cited in Armstrong have been verified :in at least 8 Colorado Pass, roughly 120 lynx effort is not considered, some early authors reported of the distribution indication For instance, information-gathering from Colorado (1); 1910-1929 of lynx since there is no way to 1890-1909 indicate intensive that lynx were ever numerous An analysis abundance those reports. for the period lynx and bobcat of lynx reported of abundance, of lynx reported Although it is doubtful the numbers in obtaining (5); 1890-1909 (6). in identifying of the historical -- but this number reflects however, a better and confusion (Fig. 1). as far south as Cumbres (the specimen of lynx in North America). but no verification Lynx exists. represents Lynx may have Published �62 o o 107 I 105 I COLORADO LYNX DISTRIBUT ON ~-RECENT o o SPECIMEN LYNX 50 ki lometers VERIFICATION PROGRAM A-TRACKS .-ACCEPTED .-MUSEUM 1>- REPORT SPECIM.EN REPORTED BY TERRELL D-COLORADO TRAPPER'S O-UTERATURE PRIOR SURVEY DENNEY 1975 TO '1912 SHADED AREA IS ABOVE 2.130 'MEHRS SOUTHERN LYNX DISTRIBUTION AND EXTRALIMITAL "-EXTRALIMITAL o 38O_ Fig. 1. Distribution of lynx in Colorado REPORTS REPORT �63 records Grand, of lynx in 7 of these 10 counties, San Juan, La Plata, Archuleta) summary, which was based largely Armstrong (1972) stated, caution." Reports " (Routt, Jackson, are solely from Cary's on observations of lynx in Garfield of the Colorado the published For the 8 counties Co. have come from 2 sources (Warren 1906). Division (1969), Eagle from which Conejos no judgement in Conejos, Monstrose, populations 1952) and Wyoming unoccupied (Long 1965). habitats in Conejos, years, neither County until similar probably Montrose, (1980). Lynx counties; probably lynx in these long have existed to their distributions Local extinctions Summit, 1971), Lake can be made about current to verify have occurred. Summit (Lake, Eagle, Pitkin, or Gunnison, in attempting the time (late 1800's), (1931) (Terrell In any case, lynx in Colorado discontinuous reported are: (1979), and Clear Creek At present, little effort was expended counties. in 1975 largely lynx have been verified, in at least 4 of these counties and Clear Creek). lynx occurrence A survey of trappers of Wildlife (1915), Gunnison (1979), Pitkin may be expected (Allen 1874, Young records. of the most recent known reports (late 1800' s), Montrose and, as ought to be regarded with a degree of 1958); Custer Co. from 1 source reflected (1911) by trappers (Cary 1911, this study); Park Co. from 2 sources by R.N. Denney Rio Blanco, in Utah and emigrations as (Durrant to Although lynx have not been and Gunnson counties were they known in Lake County until 1972 (in the latter case, however, for many 1969 or Clear Creek this study found evidence of lynx taken in 1878). The elevational appears distribution to be above 2,730m of lynx in Colorado, (9,000 ft.) (Fig. 1). past and present, Only 4 exceptions of 44 �64 records and reports elevations. While these exceptions to believe outside for which elevations lynx in Colorado the elevational 2, 700m) • are known occurred are notable, historically there is little reason existed zone of spruce-fir to any great extent forests (primarily Cary (1911) noted that lynx" •.. seldom ... wander 8,000 feet (2,440m author) even in the heaviest Colorado, the spruce-fir As presently the following known, areas: in Eagle and Pitkin counties; southeast of Leadville River drainage not be considered possible, Since verification is difficult all-inclusive. to obtain, The existence at least, in other areas of Pitkin, Creek counties fir associations Further and statutes (post-1969) from Meredith the area Pass - Mount or even substantive this listing should of lynx should be considered Eagle, Lake, and Clear spruce- occur and snowshoe hares are abundant. of lynx occurrence will be extremely part of the reasort for this is because intended evidence Since 1973, however, to protect the species. of lynx has resulted 30 days imprisonment (Title 33, Colorado it is unlikely of legislation from trapping lynx have been classified difficult. Nearly all recent or hunting. as endangered taking of lynx can result in a fine of one thousand Therefore, of includes and also in Summit, Grand, and Park counties where documentation Paradoxically, upstream in Lake County; and the Guanella of lynx occurrence In densities in Colorado Vail area in Eagle County; Evans area of Clear Creek County. to below (Dolbeer and Clark 1975). then, lynx distribution Fryingpan above snows of winter." zone is also the zone of highest the lynx's main prey, snowshoe hares evidence at lower Revised and the dollars and/or Statutes that even lynx taken accidentally 1973). will be �reported. Reports of observations may accrue, but field work specifically designed to locate lynx, even if based on reported observations, will be very time - consuming and expensive. On a positive note, however, is the fact that much of the area where lynx are now known or expected to exist is public land, administered by the USDA-Forest Service, and afforded the protection of the 1976 National Forest Management Act. No forseeable threats to lynx, their habitat, or their probable prey base (snowshoe hare) were identified during this study. LITERATURE CITED Allen, J.A. 1874. Notes on the mammals of portions of Kansas, Colorado, Wyoming, and Utah. Bull. Essex Inst., 6:43-66. Armstrong, D.A. 1972. Distribution of mammals in Colorado. Monogr., Mus. Nat. Hist., Univ. Kansas, 3:x+1-415. Brand, C.J., L.B. Keith, and C.A. Fisher. 1976. Lynx response to changing snowshoe hare densities in central Alberta. J. Wildl. Mgmt., 40:416-428. Braun, C., R.W. Hoffmann, C.E. Rogers. 1976. Wintering areas and winter ecology of the white-tailed Ptarmigan. Spec. Rpt. No. 38 Colorado Division of Wildlife. Cary, M. 1911. A biological survey of Colorado. N. Amer. Fauna, 33:1-256. Costello, D.F. 1964. Vegetation zones in Colorado. Pages iii-x in H.D. Harrington, Manual of the plants of Colorado. Swallow Press, Chicago. Coues, E. 1879. Notice of Mrs. Maxwell's exhibit of Colorado mammals. Pp. 217-225 in On the plains and among the peaks; or, How Mrs. Maxwell made her natural history collection. M.A. Dratt-Thompson. �66 Dolbeer, R.A. qnd W.R. Clark. 1975. Population ecology of snowshoe hares in the central Rocky Mountains. J. Wildl. Mgmt. 39: 535-549. Durrant, S.D. 1952. Mammals of Utah. Taxonomy and Distribution. Univ. Kansas Publ., Mus. Nat. Rist., 6:1-549. RaIl, E.R. and K.R. Kelson. 1959. The mammals of North America. Ronald Press Co., New York, 2:547-1083 +~. Long, C.A. 1965. The mammals of l-lyoming.Univ. Kansas Publ., Mus. Nat. Rist., 14:493-758. Murie, O. 1963. A field guide to animal tracks. Roughton Mifflin Co., Boston, xii + 1-375pp. Seton, E.T. 1929. Lives of game animals. Doubleday, Doran, and Co., Inc., Garden City, 1(1):1-337. Terrell, B. 1971. Lynx. Colorado Outdoors, 20(5): 19. Warren, E.R. 1906. Mammals of Colorado. Colorado College Publ., Gen. Ser. 19 (Sci. Ser. 46):225-274. ___ . 1942. The mammals of Colorado. Univ. Oklahoma Press, Norman. 330pp. Young, S,P. 1958. The bobcat of North America, its history, life habits, economic status and control, with a list of currently recognized subspecies. The Stackpole Co., Harrisburg and Wildlife Mgmt. Inst., Washington, xi + 1-193pp. �67 APPENDIX B HISTORY AND STATUS OF WOLVERINE IN COLORADO James C. Halfpenny The wolverine (Gulo gulo luscus) is a 14 to 27.5 kg mammal (Walker et al. 1975) of boreal and tundra regions (Wilson 1979). The largest members of the family Mustelidae, wolverines are carnivorous, and may feed on carrion as well as prey on living animals (Ognev 1935, Grinnell et al. 1937, Rausch 1959, Jackson 1961). Individuals range over large areas (Kratt 1960) and appear to be territorial -- males have territories as large as 2,000 km2 and female territories are 400-500 km2. Wolverines appear to be sol~tary during much of the year (Bee and Hall 1956). Historical records of wolverines in Colorado exist, but most are based on sight records (Cary 1911, Warren 1942, Spahn 1954, Field and Feltner 1974). Armstrong (1972) reviewed historical records and located just one specimen, taken near Idaho Springs in Clear Creek County (Denver Museum Natural History #2723). Although Wilson (1979) suggested wolverine numbers were increasing in Colorado, there never has been a population estimate upon which to base such a suggestion. Wolverines once were classified as fur-bearers in Colorado, but in 1965 the season was closed and the species was given complete protection. In 1973 the Colorado Wildlife Commission classified the wolverine as a state endangered species. In 1978 this study was initiated under Federal Assistance Project SE-3, to summarize the history of wolverines in the state and to accumulate information about their current status. James C. Halfpenny designed the study, coordinated field activities, and performed the data analysis. Field research activities were conducted by Halfpenny, C. Loeffler, W. Travnicek, D. Nead, M. Foster, and L. Lofgren. The assistance of the following individuals in various phases of the study is appreciated: R. Adams, B. Andrews, D. Armstrong, D. Collar, J. Fitzgerald, J. Garcia, J. Goodyear, D. Gore, L. Green, C. Hibler, T. Lytle, L. Marlow, S. Porter, M. Smith, T. Valenta, J. Wagner, and B. ~.Junder. METHODS Historical information was sought through a review of published literature and searches of museum material. Unpublished records of the Division of Wildlife were also inspected. Historical and current information was sought by a statewide "information request" campaign that employed slide programs, newspaper articles, and the distribution of over 3,000 "WANTED" posters (Fig. 1) to District Wildlife Managers, taxidermists, trappers, outfitters, private individuals, and state and federal agencies. �68 'j f..... mo INFORM W!NTE THE 0 OLV RINE, An Endangered Mammal 1111 Colorada The wolverine, largest member of tho weasel family, usually feeds on carrion. They often travol in pairs and have very large territories. APPEARANCE: Looks like a small bear but with a bushy tail; 30-40 inches in length including tail; weighs up to 40 Ibs.; dark brown with two yellowish stripes that merge above thetai!. hind PROBABLE LOCATIONS: Isolated high mountain areas of Colorado. CHARACT RISnCS: and bouncing gail. nr1 II possible, a good sighting should include: • a photo or drawing; • "photo 01 tracks with a scale included; o adrawingolthetracksortrail; o droppings; andlor • date, time and location. Widt:easurements front do not include PAW PRINTS claws I ,- sometimes stride only four ..I':.~ ,,-. toes show ,':0. ,;~ II,,' walking paltern ~ii~· YOUR ~di:i galloppaltem Information is al~ desired on all Colorado Lynx sightings. Moves with a bumping COOPERATION INGS OF THIS I?ROGRAM VERINE. MAMMAL AIMED AT REPORTING WILL SIGHT- STRENGTHEN AT PRESERVING WE NEED YOUR HELP! NOTE: This animal is endangered and is protected by law. If seen, please write or call the Colorado Division of Wildlife on Il8 possible. COLORADO DIVISION OF WILDLIFE-Nongame Section-P 6060 Broadway, Denver, Colorado 80216 •• 825-1192 Fig. 1. wolverines "Wanted" poster soliciting in Colorado. A THE WOL- information about �69 A report form was designed to standardize the evaluation of wolverine sighting reports (Fig. 2). A form was sent to all individuals reporting wolverine encounters. After the completed form was returned, the data were evaluated and rated from A to F in a standardized fashion, according to the criteria shown in Table 1. Hhen possible, all A,B, and Crated reports were followed up by personal interviews. Field study areas were selected on the basis of the observation reports (Table 2). In each study area, trails were searched for wolverine tracks using skis and snowmobiles. Baited hair snags -- hardware cloth cylinders with barbed wire interiors (Fig 3) _- were set to obtain hair samples (Hummel 1978). Reference slides of hair samples were prepared from collections at the Denver Museum of Natural History. Three keys were used to identify samples taken from the snags -- Brown (1942), Mayer (1952). and Moore et aI , (1974). RESULTS The literature review provided several hundred articles on wolverine. Reviewed articles were compiled and published as "A Bibliography of Mustelids. Part VI: wolverine" (Halfpenny et aI., 1979). The initial phases of the project were reported at the Colorado-Wyoming Academy of Science meetings (Halfpenny, et al., 1979). Historical Records Wolverine were never common in Colorado. Coues in 1879 remarked that "it is only of late years that the presence of this remarkable animal so far south has been known." Although many reports of encounters with wolverine in Colorado exist (Anon, 1946; Cary, 1911; Field and Feltner, 1974, Coues 1879; Grinnell, 1926; Halfpenny et a L, , 1979; Spahn, 1954; Smith, 1974; Warren, 1906, 1942) only 22 reports representing at least 25 animals were based on wolverine carcasses between 1871 and 1919 (Table 3). The last verified kill of a Coloradan wolverine occurred in 1919 (Grinnell, 1926). Skins or skulls of wolverine taken in Colorado are located in state museums (Table 4). Rocky Mountain National Park has kept detailed records on reported wolverine sightings for several years (Table 5). These have been summarized in an in-house report by David R. Stevens. Most notable is the sighting of 5 September 1974 where 60 visitors, two interpretive personnel, and several maintenance personnel watched a "wolverine" from the Fall River Pass visitor center. The sighting lasted 15 minutes. A long series of reports by Katherine Suhrbies proved to be unverifiable and subsequent happenings have suggested that her observations were not wolverine (D. Stevens, NPS, pers. comm.). No history of wolverine would be complete without including the history of wolverine raised at the Cheyenne Mountain Zoo, Colorado Springs. In February, 1964 the zoo received a pair of wolverine from Montana. �70 W 0 L V E R I N E 0 B S E R VAT ION REP 0 R T 10 #__ Please do not refer to a picture for filling out this form. How long observed: Minutes time of day: Date: _ Direction and Distance from nearest town: _ _______________________________ Legal Description: 'Sect.ton, County Townsh ip__ _ Range _ PM _ Topography and vegetation: _ Elevation: ft. Number of animals observed_,-__ --:c::--_ Was animal dead ? alive ? Were binoculars used? yes Distance from t-'-h-e--an-,"-'m-a-'1Number of observe-rs-Size: no -- (compare in your mind to pet dogs and cats) Height at shoulders: Tail 1ength : Weight: Body 1e·-n-g"'"'th:-:--------Characteristics: (describe size, shape and other features) Feet: _ Tail: _ Color patterns: _ _ Head: _ Body: _ (describe colors and position on each body part) Feet:, _ Head: _ Tail: _ Body: _ Chest: _ Behavior: (describe running pattern, reaction before and after the animal realized that it was being observed) Comments: (any other additional information) ------------------ On back side of this sheet, please sketch (no matter how crudely) the animal you saw and its color patterns. Observed by: Name _ Address, _ C ity-::--,;::--.,.-_-;-~S tate Zip _ Phone#: (Bus i ness) (Home )-c-----:---;-:-:-L"":-=O:--Please list other observers and their address and phone numbers 00 the back. Report taken by Date, _ Fig. 2. Form supplied information to all individuals about wolverine in Colorado. reporting sightings or other �71 Table 1. Rating system for Coloradan wolverine Rating Definition A Positive report - A report which includes proof e.g. hair, skin, photos, tracks, or skulls. B Probable report - A report which includes three or more key identifying characteristics e.g. side stripes, white chest markings, light colored claws, long bushy tail, loping movement or growling behavior; photos of secondary evidence (tracks, scat). C Possible report - A report containing one or two key identifying characters. D Weak report - The evidence is not available to rate as an "F" but the report lacks any key identifying characters. F Non-wolverine a wolverine. INC. Incomplete - The information provided was inadequate determine the identity of the mammal. Table 2. Wolverine Dates 1 Feb-30 April 15 Jan-30 April 1980 5 Feb-27 March and Criteria report - Evidence indicates the animal was not Identification as another mammal may be possible. 1980 to field study areas. Area 1979 reports. County Red Canyon American Lakes, Middle Fork Michigan R. Chair Mountain Ranch, Crystal Ri.ve r Coffee Pot Road Number Hair Snags 40 total Independence Mt. Canadian River S. Fork, Michigan River Red Canyon West of East Rifle Creek West and Main Elk Creeks Coffee Pot Road Jackson Jackson Jackson Jackson Garfield Garfield Chalk Creek Chaffee 72 total 12 �72 Fig. 3. Hair snags--hardware cloth cylinders with barbed wire interiors-- were used in an attempt to obtain evidence of wolverine occurrence. �Table 3. Reports of wolverine killed in Colorado . __ __ ._----------------------------------------- .. Source Cary 1911 Cary 1911 Cary 1911 Cary 1911 Cary 1911 Cary 1911 Cary 1911 Cary 1911 Cary 1911 Cary 1911 Cary 1911 Cary 1911 Cary 1911 Coues 1879 Grinnell 1926 Grinnell 1926 Warren 1906 Warren 1942 CSU Museum DMNH 112723 DMNH 1183 Logan Allen . Individual Fred Selah Alpert and Co. T.J. McKenna Wood Galloway R.N. Wheeler Allen (J.A.?) C.A. Cooper C.A. Cooper Carter Collection Maxwell Collection L.B. Crawford L.B. Crawford E.L. Chesnut Smith W. Weston This study O. Proffett B. Allen A. Williams This study R. Goecher & Location Date Notes Owl Mt, SE North Park Ranch Creek W. Slope Williams Fork Grand River Tin Cup Mine, Union Park, Gunnison Co. Antelope Park, Mineral Co. E. Base Pagosa Pk Calico Mt., West of Rico Silver Picket Mine, Mt. Wilson Huerfano Rvr Head Montgomery Gore Pass, between Middle & Egeria Pks Trappers Lk, Garfield Breckenridge, Summit Co Near Boulder Los Pinos Creek, SE Silverton Between Meeker & Cgaig Irwin, Gunnison Co. San Miguel Co. Camp Bird Mine, Ouray Co. Vicinity of Idaho Spgs Pass Creek, Summit Co. Wildlife Creek 1903 1903 1903 1883 n.d. 1905 1905 1895 n.d. 1871 1883 1889 n.d. n.d. 1919 1918 1890 n.d. 1913 n.d. 1876 1915-20 trapped killed trapped killed killed E. fork Cimarron, Montrose-Gunnison post1877 1966 skull found in 1977 Probable zoo escapee Co. Pine Creek, Rampart Range, Douglas Co. DMNH = Denver Museum of Natural History CSU = Colorado State University skin killed killed skin capture 4 skins -...J V-l captured taken caught skin skin/skull killed �Table 4. Known wolverine specimens from Colorado Museum or Collection Identification Number Location UNC #3 UNC 114 Unknown possible 1906 from Colorado Unknown possible from Colorado Purchased McFadden Purchased McFadden DMNH 1185 Pass Creek, Summit Co. Vicinity of Idaho Springs 20 Sep 1876 E. Carter Collection Camp Bird Mine, Ouray, Co. 1913 112723 CSU' #4503 UNC DMNH CSU Date Comments from from W. Weston Collection is the University of Northern Colorado, Greeley is the Denver Museum of Natural History, Denver is Colorado State University, Ft. Collins Two unidentified wolverine skins are present··in the CSU collection. Wolverine skins taken outside Colorado were located at the Liar's Lair, Rand, Co., and Rustic Pine Motel, Rustic, Co. Table 5. Summary of Wolverine reports in Rocky Mountain National Park. Records were taken from a special report by David R. Stevens, Research biologist (1974). Date Number of Animals Fall River Pass Wild Basin Lake-of-the-Clouds, W side of Never Summer Mts Fall River Pass area Hidden Valley Lost Lake Mirror Lake 1953 1954 1954 1957 1962 1962 1964 1965 1973 Sep 1974 Oct 1974 1976 Aug 1976 1976 Location 5 1 1 Bear Lake Fall River Pass Fall River Pass Leon Creek Bridge, Weld Basin Tundra Curves Rimfire Site Willow Basin Comments Several Reports 1 report 1 report 1 report 1 report 1 report I-Neal Guse, Ranger Possible Tracks 2 adults 3 young, Park Visitor 70 observers Pete Armington Jack Seybold Vis tor-Glenn Dent Visitor �75 In March of 1964 the female escaped through the bars. The next day Terry Schmidt of Colorado Springs killed her on v.!estLas Vegas. During April the zoo replaced the female and in Feb. 1965, the two male young were born. On Jan. 21, 1966 the parents escaped due to keeper error. Tracks from the zoo led northwest towards the Rampart Range (Cliff Meyers, superintendent, pers. comm.). February 18, 1966 Roy Goecher, Littleton, who was hunting along Pine Creek Rampart Range (Douglas Co.) encountered and killed a wolverine. Herman Schultz (D01.J)confirmed that the female was very old based on the canine teeth which were worn nearly to the gums. The worn teeth may also have been due to gnawing on bars of a cage. In all likelihood the animal shot was the female that escaped from the zoo. There has not been an accounting for the other wolverine. One of the young born at the zoo was sold in February of 1969 and a female arrived in May of 1970. Since that time two litters have been born at the zoo but both were destroyed by the female. Bill Agron, Director, was most helpful in reconstructing events at the zoo. In July, 1977 Mr. Al Williams, assistant principal of Delta High School, Delta, Colorado, found a skull on the east fork of the Cimarron drainage (Montrose and Gunnison Co.) while leading students on a hike. He made sketches of the skull. Later, students burned the skull, and only a few fragments were saved. Steven Bissell of Colorado DOW positively identified the fragments as wolverine, and judged the age to be from 10 to "as old as 100 years." The fragments are being held at the Delta High School. Two wolverine, a male and a female, were released by private individuals during the filming of "Rocky Mountain Reunion." The director, Mark Stouffer (Aspen) filmed the movie in conjunction with the Colorado Division of Wildlife. In the movie John Denver was involved in several Division projects which included releasing extirpated or endangered species into Colorado. However the wolverine release was not a Division project. The wolverine, which were imported, wild, from Canada, were released two kilometers due east of Castle Peak (Pitkin--Gunnison Co. line) on 26 October 1978. When the tracks of the wolverine were last seen they were traveling together. Since the release, three wolverine sightings have been reported that may relate to these animals. The reports were from Willis Lake (Lake Co.), Independence Pass (Lake Co.), and Grizzly Creek (Pitkin Co.) all of which are within 35 km of the release site. The report at Willis Lake (Mrs. George Rowe, Leadville) was of a mother and two young. In March of 1979, Robert Kay (Vernal, Utah) shot a wolverine. According to Kay the wolverine was killed two miles west of the Colorado-Utah border just off U.S. 40. The wolverine, which Mr. Kay retained, was a male, weighed 14.5 kg (32 lbs), and was reproductively active. Of particular interest is the greasewood desert habitat where the wolverine is supposed to have been taken. The location of the kill is far from any mountainous habitat and certainly would represent a somewhat unusual �76 occurrence. Since wolverine may roam over 32 km per day, the animal may have had part of its range in Colorado. Observation Reports A total of 271 reports of wolverine observations were evaluated during the study. Of these, 188 were received during the study -- the remainde:r;were from reports gathered by R. Zaccagnini (DOW) in 1974 and other reports periodically received by the DOW. Three reports could not be rated. The remaining 268 reports were rated as follows: A - 3(1.1%); B - 19(7.1%); C - 38(14.2%); D - 60(22.3%); F - 105(39.1%); Incomplete43 (16%) . The A - rated reports were those from the Williams skull, the Goecher kill, and the Kay kill in Utah. Analysis of wolverine reports revealed several trends. Most (87%) observers reported one wolverine per sighting and observations were usually made by one (54%) or two (26%) people (Table 6). Most sightings were for a brief interval (35% less than one minute) and at a short distance (57% less than 50m) (Table 7). The majority (61%) of the reports occurred at elevations of 2,700 to 3,600m (Table 8). Most reports (88%) occurred during the months May through October (Table 9). It would be expected that during these months more people are in the backcountry. Reports were nearly equally distributed between mornings (41.4%) and afternoons (39.3%) while a smaller percentage (19.7) of reports occurred during the evening or at night. As would be expected from the elevational distribution most reports occurred within the conifer zone. Ten of 11 reports of observations from November through April contained enough information for a detailed analysis (Table 10). Some important trends were obvious. Wolverine reports occurred at low elevations during the winter and as winter progressed, reports came in from lower elevations. During the winter, reports were usually associated with oakbrush, sagebrush, or fields at the lower montane ecotone. The locations of reports are closely associated with winter ungulate ranges. Some of the reported observations deserve· special mention. On 7 July 1979 Mr. Dick (C.R.) Wegner (Bayfield) photographed a series of tracks west of Dollar Lake (La Plata, Co.), which appear to be wolverine tracks. Mr. Wegner felt that the tracks had been made early the morning he took the photos. Judging from the cover of snow algae in the photo, the tracks were not melted out and therefore the photo represented a very good indication of wolverine in that area. Dr. P. Wright who had dealt with many wolverine in the field, has tentatively concurred on this judgement (pers. comm.). Mr. George Lanum (Denver) shot three photos of an animal crossing a snowfield near Trinchera Peak during June 1978. By studying the progression of slides, one receives the impression that the animal may be traveling with a lope. The tail passes far below the level of the back and then above the back. The general shape of the body is suggestive of a wolverine as was Mr. Lanum's description. However the animal is silhouetted and no color patterns are evident. �77 Table 6. Frequency of the number of observers per wolverine report and the frequency of animals observed per report. Includes only reports rated at A,B,C, or D. Number Percentage of observers Percentage of wolverine per observation 1 2 3 4 5 TOTALS (n=106) 53.7 26.4 6.6 12.3 .9 99.9 (n=108) 87.0 9.3 1.9 .9 .9 100.0 Table 7. Frequency of observational distance and lengths of observation. Includes only reports rated at A,B,C, or D. Distance (m) 0-25 26-50 51-100 101-200 201-400 Percentage of Sightings (n=88) Length of Observation (min) 39.8 17.0 22.7 11.4 9.1 < ~ ~1 >1-5 >5-10 > 10 Percentage of Observations (n=92) 19.6 15.2 31.5 10.9 12.0 Table 8. Frequency of occurence of wolverine reports at different elevations. Includes only reports rated at A,B,C, or D. Elevation (m) 1,500-1,799 1,800-2,099 2,100-2,399 2,400-2,699 2,700-2,999 3,000-3,299 3,300-3,599 3,600-3,899 3,900TOTAL Elevation '(ft) 5,000- 5,999 6,000- 6,999 7,000- 7,999 8,000- 8,999 9,000- 9,999 10,000-10,999 11 ,000-11 ,999 12,000-12,999 13,000 Percentage of reports (n=79) 1.3 5.1 11.4 11.4 19.0 20.3 21.5 7.6 2.5 100.1 �78 Table 9. Frequency of occurance of wolverine Includes only reports rated as A,B,C, or D. reports by months. Percentage reports Number of reports Honth 1 January February Harch April May June July August September October November December 1 0 2 10 15 21 17 9 10 6 1 1.1 1.1 0.0 2.2 10.8 16.1 22.5 18.3 9.7 10.8 6.5 1.1 TOTAL 93 100.3 Table 10. Analysis of reports November through Apr il. occurring Honth November November November November January Harch April April April Spring Elevation m (ft) 2,200 3,000 2,000 2,660 Low 1,880 1,880 1,700 1,970 2,100 during Habitat (7,300) (10,000) (6,600) (8,800) Oakbrush Pine and Spruce Pasture Dense Timber (6,200) (6,200) (5,700) (6,500) (7,000) Oakbrush Pasture fields Scrub, Pine Sage and Rabbit Brush of the time period of Location 24 km S Steamboat 19 km S Edwards 10 km W Steamboat 16 km Topanas 24 N Durango 19 km N Rifle N New Castle 5 km N Rifle 13 km S Meeker SW Hesperus Rating C C D C C B C B C C �79 During the second week in June 1978 Mr. and Mrs. Robert Zaas (Houston, Texas) photographed 4 animals west of Lake City. Again the animals were too distant to be positively identified. However the drawing and descriptions by the Zaas' could have been wolverine. On 16 March 1979 Don Roberts, Gary Guggenberger, and Ron St. Pierre (DOW) watched an animal for three minutes at 25 m. The men were very sure of their wolverine identification and described a wolverine when interviewed. The sighting was south of the Rifle Fish Hatchery. This canyon has been the location of other very credible sightings. (L. Kontour, Rifle). On 24 June 1979 Kurt Keskimaki watched what he described as a wolverine for four minutes at 11 m. Mr. Keskimaki was bear hunting and the animal took his bait from in front of his blind. The location was north of Parshall, (Grand, Co.). This was one of two reports by bear hunters of wolverine coming to their bait in 1979. Three behaviors are mentioned in several of the wo Lve r i.ne reports. Many of the summer reports are of wolverine chasing or hunting marmots. Three reports refer to wolverine watching a marmot while the tail was moved in a circular motion. In four instances wolverine were reported to have approached (charged?) the observer. In two situations the animal in question was accompanied by young. For 33 of the F - rated (non-wolverine) reports it was possible to identify the mistaken animal. Badgers (Taxidea taxus) (34.9%) and marmots (Marmota flaviventris) (36.4%) were most often mistaken for wolverine. Other species mistaken for wolverine included marten (Martes americana) (18.2%), weasel (Mustela sp.) (3.0%), and porcupine (Erethizon dorsatum) (3.0%). Hair Snags No hair obtained from the hair snags were identified as that of wolverine. Three hair samples could not be identified to species. Identified hair samples collected included: marten, weasel, coyote (Canis latrans), bobcat (Lynx rufus), striped skunk (Mephitis mephitis), spotted skunk (Spilogale putorius), raccoon (Procyon lotor), and chickaree (Tamiasciurus hudsonicus). No hair of any kind was collected at the Chalk Creek study area. In 1979, the 40 hair snags were checked approximately 1 month after they were set out and baited. Most of the bait was gone by that time. In 1980, therefore, the snags were generally checked and rebaited, if necessary, at 10 day intervals (the Chalk Creek snags were checked less frequently). DISCUSSION Wolverine were present in Colorado at least until 1919, when the last known documentation of a wild specimen occurred in the state. Since �80 that time, a skull has been recovered, the age of which may be 10 to 100 years old, one animal believed to have been a zoo escapee has been killed in Colorado (1966), and another animal has been killed in Utah, reportedly 2 miles west of the Colorado-Utah border (1979). In addition, 2 wolverines were released on the Pitkin-Gunnison County line (1978). Despite a number of credible reports, the study did not verify the existence of a viable wolverine population in Colorado. However, names of several areas show up repeatedly in the reports, suggesting areas where wolverine are more likely to be found. During the colder months these areas would be NW Colorado, especially SE of Meeker, N of Rifle, and Nand E of Durango. In the warmer months key areas may be the Flat Tops, NW Rocky Mountain National Park, and the San Juan Mountain complex. Wolf Creek Pass has a number of reports year around. Other areas also may have wolverine, but they are not reported as often. The San Isabel National Forest south of La Veta and the Sangre de Cristo Land Grant to the west are of interest due to the one set of photos from that area. If we assume that at least some of wolverine reports received by the project are true, we may hypothesize about the ecology of wolverine in Colorado today. The wolverine usually travels alone, although some may be reproducing as evidenced by reports of young. The wolverine is very secretive and most observers surprise the animal, obtaining but a short glimpse (less than 1 minute) at a short distance (less than SOm). During the warmer months, May through October, the wolverine occurs primarily at higher elevations from the upper montane to the tundra regions. A portion of their diet may consist of marmot during the summer. S-tarting around November, at least some wolverine may start an elevational migration to the lower limits of treeline or their movements within their ranges are great enough to reach low elevations. These migrations probably follow wintering ungulate herds and may go down into the oakbrushsage brush zone. The distribution of winter reports suggests that the animals may favor major south facing slopes or drainages. Wolverines may continue to exist in Colorado. The chances of obtaining irrefutable proof of a viable population are slight, however, because of difficulty in detection and-because their endangered status makes it unlikely that authorities will be notified in the event a wolverine is accidentally taken. If a wolverine population does exist, it appears that management of ungulate winter range will also benefit wolverines in winter and, conversely, the loss of such winter range could have detrimental effects upon wolverine. No human activities potentially detrimental to wolverine in summer ranges were identified during this study. LITERATURE Anonymous, 1946. The wolverine Comments, 9 (1) :22. CITED in Colorado. Colorado Conservation Armstrong, D.M. 1972. Distribution of Mammals in Colorado, Kansas University. Museum of Natural History. Monograph No.3. 416pp. �81 Bee, J.W. and E.R. Hall. 1956. Mammals of northern Alaska on the Arctic slope, Misc. Publ., Univ. Kans., Mus. Nat. Hist. 8: 1~309. Brown, F.M. 1942. The microscopy of mammalian hairs for anthropologists. Amer. Phil. Soc., Philadelphia. Proc., 85(3): 250-274. Coues, E. 1879. Notice of Mrs. Maxwell's exhibit of Colorado mammals. Pp. 217~225, in: M.A. Cartt-Thompson, On the Plains and Among the Peaks: or, How Mrs. Maxwell Made Her Natural History Collection. Claxton, Remsen, and Haffelfinger, Philadelphia. 237pp. Cary, M. 1911. 1-256. A biological Field, R.J. and G. Feltner. Grinnell, G.B. 1926. survey of Colorado. 1974. Wolverine. N. Am. Fauna, no. 33: Colo. Outdoors. Some habits of the wolverine. 23(2): 1-6. J. Mamm. 7(1): 30~34 • --- ., J. Dixon, and J.M. Zensdale. 1937. Fur-bearing mammals of California: their natural history, systematics, status and relations to man. Univ. Calif. Press, Berkely. 2: 251-256. Halfpenny, J.C., D. Nead, S.J. Bissell, and R.J. Aulerich. 1979. A bibliography of mustelids, part VI: wolverine. Michigan Agric. Exp. Sta. 9214: 1-91. Jackson, H.H.T. 1961. Mammals Madison, xiii + 504pp. of Wisconsin. Univ. Wisconsin Press, Krott, P. 1960. Ways of the wolverine. This animal inhabits a vast area to which it is peculiarly adapted. Nat. Hist. 69(2): 16-29. Mayer, W.V. 1952. The hair of California mammals with keys to the dorsal guard hairs of California mammals. Amer. MidI. Nat., 48: 480-512. Moore, T.D., L.E. Spence, L.E. Dugnolle, W.G. Hepworth. 1974. Identification of the dorsal guard hairs of some mammals of Wyoming. Wyoming Game and Fish Dept. Bull. 14. "VJyomingGame and Fish Dept. Cheyenne. 177pp. ° Ognev, S.I. 1935. Mammals of U.S.S.R. and adjacent countries. Vol. III. Carnivora (Fissepedia and Pinnepedia). Transl. from Russian by Isreasel Program for Scientific Translations, Jerusalem, 1962. 641pp. Rausch, R. 1959. Studies on the helminth fauna of Alaska. XXXVI: Parasites of the wolverine, Gtilo gulo, L., with observations on the biology of Taenia twitchelli, Schwartz 1924. Journal of Parasitology 45(5): 465-484. Smith, B. 1974. Spahn, J.J. 1954. Outside, insight Wolverine. 1974. Colo. Div. Wildl. News Release. Colo. Cons. 3(2); 1-3. �82 Walker, E.P., et al. 1975. Mammals of the world. 3rd ed. (J.L. Paradisco ed.). The Johns Hopkins Univ. Press, Baltimore. Warren, E.R. 1906. Mammals of Colorado. Colo. College Publ., Gen. Ser. 19 (Sci. Ser., 46): 225-274 . --- . 1942. The mammals of Colorado. Univ. Okla. Press. 330pp. Wilson, D.E. 1979. Wolverine (Gulo gulo). In Game, Pest, and Commercial Mammals of North America. In prep. �83 JOB PROGRESS REPORT State of COLORADO Project No. FW-22-R -------------------------- Work Plan No. Job Title 1 ------------------------ Job No. 3 Population Surveys of Selected Bird and Mammal Species in Colorado Period Covered: Personnel: Nongame Investigations 1 July 1979 through 31 December 1980 W. Graul, M. Moulton, J. Freeman, C.'Reckling, E. McGrath, S. Vallejos -- Colorado Division of Wildlife. ABSTRACT From 1 July 1979 through 31 December 1980 intensive field studies were conducted on the following species or species groups: Great Blue Heron, Greater Sandhill Crane, bats, and eastern prairie mammals. Based on this work, tentative distributional data were compiled for the mentioned species or species groups. Data on habitat associations, productivity, and nesting requirements were collected for some of the species. �84 POPULATION SURVEYS OF SELECTED BIRD AND MAMMAL SPECIES IN COLORADO Walter D. Graul P.N. OBJECTIVES 1. Determine statewide distributions, population levels, and population trends for select bird and mammal species; those with relatively restricted habitat requirements. 2. For the same species, determine species - habitat associations. SEGMENT OBJECTIVES 1. Determine statewide distributions bird and mammal species. and habitat associations for select INTRODUCTION Historically, wildlife management has been orientated toward single species. This approach is not feasible for the broad spectrum of nongame species. For instance, there are 347 species of birds and 73 species of mammals classified as nongame in Colorado. An alternative approach, termed a species-ecosystem approach, has be n implemented for management of all nongame species in Colorado. In fact, this approach now constitutes the foundation of the Division of Wildlife's management program for those nongame species not classified as threatened or endangered. The method basically consists of determining: (1) what species in a community have the most restricted habitat requirements, and (2) what are the habitat requirements of these restricted species. The latter habitat requirements can then be used to develop management guidelines for broad habitat types that will insure the preservation of a wide spectrum of species -not just a few. Based upon preliminary data collected by the Division's nongame program, it is possible to determine what bird and mammal species in Colorado should receive top priority in terms of applying the species-ecosystem approach. Namely, these are the species that in terms of distribution in Colorado are relatively restricted while not being peripheral. Furthermore, those species whose distributions are unknown need to be investigated immediately. The species addressed in this study were chosen according to the preceding criteria. METHODS AND MATERIALS All the intensive field studies were conducted by temporary procedures of each major segment will be described herein. employees. The �85 Nesting success data for the Greater Sandhill Crane (Grus canadensis tabida) were obtained and compiled by Candy Reckling from 1 July-21 July 1979. Previously, the main trend indicator for the nesting success of Colorado's breeding crane population had been based on spring and fall staging ground counts. These data were compiled. As a check on the accuracy of these counts, Colorado's main nesting population in California Park, Routt County (22 miles north of Hayden) was examined in July to determine production. Nests had been located in June by a U.S. Forest Service employee. In the 1979 portion of the Great Blue Heron (Ardea herodias) study, Stan Vallejos and Candy Reckling compiled and analyzed data pertinent to the complete 1979 nesting season. These data were presented in the last Job Progress Report. Elizabeth McGrath conducted the 1980 field season work on the Great Blue Heron from 22 March through 22 September. Her work consisted of two segments. First, she verified the location and status of heron nesting colonies reported after the 1979 field season. The second segment consisted of making detailed field observations at 4 of the largest heronries in Colorado: Chatfield Reservoir, Boulder Creek, Lonetree Reservoir, and Fossil Creek Reservoir. The primary function of the latter observations was to ascertain the population levels for the 4 sites throughout the breeding season. Associated with this, it was possible to examine the relationship between heron activity and human activity in an ancillary manner. All observations were made with a 15x - 45x spotting scope from distances of 300 m or greater. From 22 March 8 May observations were made at Chatfield Reservoir and Boulder Creek. From 9 May on, observations were made at all 4 sites. Throughout the study the observation days were alternated between the sites in a standardized, rotating manner. On a given day, observations were made either from sunrise to midday, or from midday to sunset. Small mammals were collected in July, August, and September of 1979 in Las Animas, Yuma, and Baca Counties. An equal number of snap traps and live traps was set in each of nine sites in an equal number of stations in parallel lines. A total of 10,600 trap/nights was run. Each animal that was trapped was removed from the area after it was labelled according to its trap-station number. Trap lines were operated for 4 consecutive days and nights in each area in Las Animas County, 20 consecutive days and nights in Yuma County, and 10 consecutive days and nights in Baca County. Vegetation was analyzed with the step-point method (Evans and Love 1957), in the Yuma and Baca county sites. In the Yuma County sites 400 "po In t s" were taken (10 points of each of the 40 trap stations). In the Baca County sites 300 "points" were taken (6 points of each of 50 stations). All points were taken along an imaginary line perpendicular to the trap line. Trapping stations were used as starting points. In the Yuma County sites five points were taken at 25 cm. intervals on both sides of each station. In the Baca County sites, three points were taken at 50 cm. intervals on both sides of each station. These points were categorized according to �86 whether or not they intercepted a basal crown of a plant. An (X) was recorded if a basal crown was intercepted, and a (0) was recorded if a basal crown was not intercepted. Plants were described either as "grass" or "nongrasstl• The "nongrass" category includes broad-leaf forbs, shrubs, cactus, and cactus-like plants. In the event that a basal crown was not intercepted, it was recorded whether the point intercepted bare ground (symbolized by "bg") or "mulch" (symbolized by m) . "Mulch" included any dead vegetation and cow pies. The points were summarized in terms of vegetative composition (percent grass vs. percent non-grass), and basal cover (perfect basal cover of grasses, non-grasses, mulch and bare-ground). Microhabitats of select species were described by summarizing the vegetative points for all stations where a given species was trapped. Vegetative composition and basal cover for the Baca and Yuma County sites are listed in table 1 of the mammal segment. Microhabitats of select species are recorded in table 2 of the mammal segment. After reviewing the scientific literature and contacting professional mammalogists, areas were selected to be trapped for bats. Most of these areas had never been investigated or had not been investigated in the recent past. Some areas which had been studied previously were used to validate some important recorded distributions. At each area, bats were trapped by placing three to five mist nests over isolated ponds or metal tanks or along streams. Trapping was done thoughout the night for one to three nights in each area. For each individual captured, species, sex, reproductive conditions, time of capture, weight, and length of forearm were recorded. During three field seasons more than 140 nights (approximately 500 netnights) of netting have been completed. In addition to trapping by mist netting during the night, old buildings, cellars, mines, bridges, rock crevices and caves were searched during the day for bats. �87 RESULTS AND DISCUSSION Greater Sandhill Crane Segment Staging ground counts for the only two known major crane staging areas in Colorado are shown in Figures 1 and 2. The Morgan Bottom area is northeast of Hayden, along the Yampa River. The Hayden Power Plant site is about one mile south of the Morgan Bottom area. It serves as a secondary spring staging area for the Morgan Bottom cranes and is used during springs with major snow cover. The Elk River staging area is on the Selby property and is located about two miles north of the confluence of the Elk and Yampa Rivers. All staging areas are located in Routt County. They are located south of the main breeding range of the Colorado nesting population. The most significant known nesting population of the Greater Sandhill Crane in Colorado is located in California Park. A Colorado Division of Wildlife study yielded 11 nests both in 1973 and 1974 in this park. In 1977, 9 nests were located. In 1979, only 8 nests were found. Based on the number of eggs laid, the hatching success in 1976 was 30.4% and in 1977 it was 33% (Bieniasz, 1978). In 1979, the hatching success was 38%. In comparable studies in other states, Drewien (1973) reported a 78% hatching success for the Greyts Lake National Wildlife Refuge population in Idaho, and Littlefield and Ryder (1968) reported a 42.7% hatching success for the cranes nesting at Malheur National Wildlife Refuge in Oregon. Both of the latter populations are located in extensive marshes, whereas the Colorado breeding population is characterized by nests associated with willow-lined drainages in mountain parks. The Colorado nesting crane population is centered in a portion of the Routt National Forest that can be expected to be utilized by more and more people during the crane nesting season (in conjunction with the anticipated human population increase related to energy development in northwestern Colorado). Compounding the human population increase problem (for the cranes) is increased accessibility to the crane nesting areas. For example, the road through California Park is being upgraded considerably in relation to a new timber sale; previously, it was 4-wheel drive only into July. As a consequence of the preceding trends, it is necessary to closely monitor the status of the Colorado crane nesting population. The problem is how to do this monitoring most efficiently. One possibility is to just monitor spring and/or fall staging ground populations. Another is to monitor the nesting success of a selected population such as California Park. Another alternative is to conduct complete nesting and fledging success surveys for the total crane nesting population. Obviously, the first two alternatives would be preferable if they reflect the status of the overall nesting populations. Consequently, an analysis of the staging ground and California Park nesting data is useful. �350 Maximum Spring Count Maximum Fall Count I I I 300 I I I I I r. \ \ \ \ \ \ \ \.--~-- / / / 250 " 200 CJ) Q) g H U I 4-1 0 H \ ./ / 150 Q) ~ ::l Z LOO 50 ....1.-__ ~._._ -L. -.. ...1..... -L-._--f-------L., 19h(' __ --'-.. . _.L._. --.i.. , ____...L__ __ .. ....l-. __ w. __ .1.. " 197n 196'5 19T5-~' Ypi:lrs ~ .i ~I, \', ir ar» .:::,'ig"jl)~' r ounr :.OIJllt'O ~organ BPi film-Hayden Power Pl ant. ~----J1980 00 00 �350 Maximum Maximum Spring Count Fall Count 300 250 200 Ul <I) ~ ~ u 00 ~ 1.0 0 ~ <I) 150 / ~;:l z / / ; " \ \ \ \~ I 100 / •... _ _J I 50 ~ , I .I 1975 Years Figure 2. Crane Staging Ground Counts: Elk River --''-__ -'-__ -'-__ -'-__ OJ 1980 �A comparison of the two staging ground data sets (Fig. 1 and 2) is enlightening. Note that between 1976 and 1979 the spring count data for Morgan Bottom indicate a gradual increase, whereas the Elk River data for the same period indicate a decrease, followed bv a slight increase. Similarly, the fall staging ground counts for the two areas do not follow a parallel pattern. A comparison of spring and fall counts for a given staging ground is also interesting. Note that there is no positive correlation between fall and spring counts for either of the staging areas. The final comparison that is of potential value is the relationship between the California Park nesting data and the staging ground data for Morgan Bottom (immediately south of California Park). This comparison would be useful if crane nesting success in California Park is indicative of the overall nesting success in given vears. There is no correlation between the number of birds staging in the spring and the subsequent number of nesting birds in California Park. Likewise, there is no apparent correlation between hatching success in California Park and subsequent fall staging populations on Morgan Bottom. None of the above comparisons, therefore, provide any hope for a shortcut method of monitoring the nesting crane population on an annual basis. There are several possible reasons for this. First, it appears that spring staging ground counts are heavily influenced by weather. Namely, in springs with heavy snow cover, the cranes seem to stay on the staging grounds longer, and consequentlv reach higher maximum counts. Also, thpre may well be movement between the two major staging areas. The fall staging ground counts undoubtably do not just reflect the local nesting population. BirdS banded at Grey's Lake have been observed on the Morgan Bottom staging area. Finallv, it is entirely possible that nesting success in California Park is not reflective of the overall nesting success fer the Routt Forest crane population. At this point, some tentative recommendations can be made. Apparently, the major value of the staging ground counts is to monitor Long-term population changes and these changes may well be more indicative (If the overall Rockv Mountain crane nesting population than the local Colorant) nesting population. Also, it appears that the only way to monitor the nesting success and overall population status of the Colorado crane nesting population is through intensive nest surveys. Prior work has illustrated that nests over a wide area can best be located through helicopter surveys. Finally, in view of the decreasing number vf nests in California Park and the relatively low hatching success for this area, it appears that a long-term monitoring program is warranted. Literature Cited Bieniasz, K.A. 1978. Biology of Greater Sandhill Cranes in Routt M.S. Thesis, Univ. of Northern Colo., Greeley, 71p. Drewien, R.C. 1973. Ecology of Rocky Mountain Greater Ph.D. Thesis, Univ. of Idaho, Moscow, l06p. Sandhill County, Cranes. Colo. �91 Littlefield, C.D. and R.A. Ryder. 1968. Breeding biology of the Greater Sandhill Crane on the Malheur National Wildlife Refuge, Oregon. Trans. N. Amer. Wildl. Nat. Res. Conf., 33:444-454. Great Blue Heron Segment As of 1979, 38 active and 21 inactive Great Blue Heron nesting colonies had been located. In 1980, three new active and two new inactive heronries were documented. The 41 active and 23 inactive heronries are detailed in the updated versions of Appendices I and II. The remainder of this segment report will address detailed at four of the most significant heronries in Colorado. Description data collected of Sites The Boulder Creek heronry is located between Hwy. 287 and 95th St., TIN, R69W, Sec. 16, SE ~ in Boulder County. The property, Boulder Valley Farm, is currently being leased as a commercial cattle operation. The heron colony lies on the north bank of Boulder Creek, approximately 1.1 km southeast of the ranch buildings, in a grove of cottonwoods (Populus spp.). Use of land adjacent to the colony is limited to grazing cattle and occasional ranch-related traffic. Access to the heronry is restricted and the property is posted private. Infrequently, trespassers were observed in the vicinity of the heronry. The owner of the property is currently negotiating with the Division of Wildlife to have the Boulder Creek heronry designated a natural area. This colony has been in existence for 30-35 years and appears to be in stable condition (Graul, 1979). The exact number of active nests could not be determined without disruption to the colony, but 62 active Great Blue Heron nests were observed with the use of a spotting scope. A final nest count after the birds had left the heronry revealed the presence of 127 large platform nests assumed to be Great Blue Herons, but it was not possible to determine how many of these nests were active during the 1980 breeding season. In addition, other species present within the colony included: Night Herons (Nycticorax nycticorax), Great Egret (Casmerodius and Great Horned Owl (Bubo virginianus). Black-crowned albus) The Chatfield Reservoir heronry is located 8 km south of Littleton, T6S, R69W, Sec 12 in Jefferson County. Chatfield Reservoir, and the adjacent lands, are used as a recreational area for metropolitan Denver. Human activities on the lake occur in designated areas and include: boating, fishing and waterskiing. An overlook on the southeast shore approximately 350 m from the heronry was constructed by the Colorado Division of Parks for observation of the heronry. The heronry, situated in a grove of cottonwoods, was originally located along the banks of the South Platte River. In 1979, this area was inundated when the �92 Army Corp of Engineers raised the water level on the reservoir. The heronry is now completely surrounded by water. The Division of Parks has established a "buffer zone" approximately 138 m around the heronry with the use of buoys. All boating is prohibited inside this zone until September 1 to eliminate disturbance to the birds during the breeding season. Trespassers were observed inside the buoy line throughout the summer. This colony has been in existence for 80 years and the number of nesting herons has increased 8-fold in the last 9 years CH. Kingery, pers. comm.). The exact number of active nests could not be determined without disruption of the colony, but 68 active Great Blue Heron nests were observed with the use of a spotting scope from the overlook. A final nest count after the birds had left the heronry revealed the presence of 117 platform nests. In 1979 and 1980, Double-crested Cormorants CPhalacrocorax auritus) began nesting in the colony. The cormorants appear to be in competition with the herons for nest platforms (Ryder et al. 1979). A nest of Great Horned ~vls was also in the colony. Fossil Creek heronry is located approximately 14.4 km southeast of Fort Collins, T6N, R68W, Sec. 9, SW ~ in Larimer County. The heronry is situated on the northwest shore of Fossil Creek Reservoir. The reservoir and a "buffer zone" of shoreline is presently privately owned. Use of the lake is limited to members and consists primarily of motorboating and waterskiing. To the north and west of the heronry, land use is agricultural. to the trees from this direction is not regulated. Access Activity at Fossil Creek included both land and water-related disturbances. Motorboats and waterskiers continually passed the southeast portion of the heronry. Numerous people on foot, motorcycle, horseback and motor vehicle entered the heronry from the northwest. Fossil Creek seemed especially susceptible to nature watchers and photographers. The heronry has been in existence for at least 10 years (Graul, 1979) and appears to be decreasing. The actual heronry is u-shaped in appearance and follows the contours of the shoreline. Observations of the season revealed the after the birds had Black-crowned Night heronry with a spotting scope during the breeding presence of 80 active nests. A final nest count left the colony yielded 85 nests. In addition, Herons also occupied this heronry. Lone Tree heronry is located 3.2 km northwest of Berthoud, T4N, R69W, At Lone Tree the heronry is divided Sec. 9, NE ~ in Larimer County. into two separate colonies. The original site is located in a grove of cottonwoods on the east shore of Lone Tree Reservoir. The front of this grove faces west onto the reservoir; the north and south edges of the grove border agricultural land; and the rear (east) of the grove is directly in contact with a causeway separating Lone Tree from \Velch Reservoir. Most nests were concentrated at the rear (east) of the grove but a few nests were observed throughout the grove. Reports (C. Cummings, pers. comm.) indicate that a major disturbance (2 boys shooting herons) �93 to the original site caused the birds to abandon this grove in the mid1970's and move 0.5 km east to a small group of cottonwoods on the south shore of Welch Reservoir. Trees at the new site are situated between the shoreline of Welch Reservoir to the north and an agricultural field to the south. The east shore of Lone Tree, which contains the original heronry, is private property. However, it receives considerable traffic from the public beach and campsite directly across the lake. Activity at the original site consisted of boating, horseback riding, motorcycling and pedestrian traffic. New site activity consisted primarily of farm equipment and occasional boat traffic. During colony; counted active the 1980 breeding season, some herons began to reoccupy the original the majority continued to nest at the new site, where 22 nests were with the use of a spotting scope. It was not possible to count nests at the original site without disturbing the herons. Final nest counts made at the original and new sites after the nesting season revealed 25 and 22 nests, respectively. Black-crowned Night Herons were observed determined if they were nesting. at the old site but it was not Human Activity ~ll human intrusions were quantified during the study as to the type of activity. The locations of the intrusions were recorded via two zones: Zone I -- directly under the nest trees; Zone II -- within 100 m of nest trees (at Chatfield this zone ranged from 100 m - 138 m in accordance with an established busy line). The reactions of herons to the various human intrusions were recorded as follows: (1) minimal or no response, (2) local response - temporary abandonment of nests in the area proximal to the intrusion, and (3) general response temporary abandonment of nests throughout a heronry. The number of human intrusions per observer hour per month for each of the four study sites is presented in Table 1. The type of intrusion and its location (zone) in relation to the heronry is also presented. Neither the Chatfield Reservoir nor the Boulder Creek heronries experienced any human intrusions during 38.0 hours of observation for the month of March. During the month of April, Chatfield Reservoir heronry had a higher ratio of intrusions than Boulder Creek heronry R = 0.2364 and = 0.0448, respectively. = number of human intrusions per hour of observation). R (R In the month of May, there was an increase in the number of intrusions at Boulder (R = 0.1260). Lone Tree (R = 1.934) and Fossil Creek (R = 0.400) heronries had the highest intrusion rates for May. The Chatfield heronry experienced the lowest rate of intrusions = 0.0880) of the four sites in May, undoubtably because the area was closed to boating in early May because of high water levels. (R �94 Table 1. The Number. and Types of Human Intrusions per Observer Hour by Month for Each of the Four Study Sites. BOULDER CREEK HERONRY Zone1 TYPE OF HUMAN INTRUSION LAND motor vehicle2 foot motorcycle horseback WATER unmotorized motorized Total hours observation/month Total human intrusions/hr/month March I II o o o o o o o o 15.5 0.0 April I II o o .0224 .0224 o o o o May I II o .0315 .0315 .0315 .0315 o o o June I II July I II o o o o o o o .0465 o .1395 o .0930 o o .057 .057 44.5 31.75 17.5 21.5 0.0448 0.1260 0.114 0.2790 June I II July I II CHATFIELD RESERVOIR HERONRY March I II April Zone TYPE OF HUMAN INTRUSION LAND motor vehicle foot motorcycle horseback WATER unmotorized motorized o o o o Total hours observation/month Total human intrusions/hr/month o o 22.5 0.0 I II .1182 .1182 May I II .044 o o .044 o o 0 .0588 50.75 22.5 17.0 0.2364 0.088 0.0588 .0889 .2222 .1333 .0444 22.5 0.4889 �95 Cont. Table 1 FOSSIL CREEK HERONRY March I II Zone TYPE OF HUMAN INTRUSION LAND motor vehicle foot motorcycle horseback wATER unmotorized motorized NO DATA Total hours observation/month Total human intrusions/hr/month LONE TREE RESERVOIR 1 April I II May I II NO DATA 0 0 0 0 0 0 0 0 0 0 0 .400 I t June I II July I II 0 .030 .470 0 .030 0 0 0 0 0 0 .220 0 0 0 0 0 0 0 0 .0667 0 0 0 12.5 31.75 15.0 0.400 0.760 0.0667 May I II June I II July I II HERONRY March I II lone TYPE OF HUMAN rNTRUSION LAND motor vehicle foot motorcycle horseback WATER unmotorized motorized NO DATA April I II NO DATA , I I Total hours observation/month Total human intrusions/hr/month 1 .0394 .0394 .7896 .1974 .0394 0 .1579 0 .1974 .0790 0 0 0 0 0 0 0 .0656 0 .0714 0 0 0 0 0 0 0 0 0 .2764 .0656 .0328 .1184 .1311 0 .0714 25.33 30.5 14.0 1.934 0.2951 0.1/.29 II Zone indicates location of the intrusion; Zone I is directly under the trees; Zone II is within 100 m of the trees at Fossil Creek, Lone Tree and Boulder Creek: and within 138 m at Chatfield Reservoir. 2/ The motor vehicle category includes trucks, cars and farm machinery. �96 In June, the rate of intrusions at Boulder heronry remained stable = 0.114), while the rate of intrusions at Chatfield heronry continued to decline = 0.0588). The rate of intrusions at Fossil Creek heronry increased to its highest level for the summer (R = 0.760). Lone Tree heronry experienced a decline ell = 0.2951) in the rate of intrusions but remained second highest for the month of June. (R (R In July, both Boulder Creek and Chatfield heronries demonstrated an increased intrusion rate, = 0.2790 and = 0.4889, respectively. Both Fossil Creek = 0.0667) and Lone Tree = 0.1429) experienced drop in the rate of intrusions. (R R R (R a A comparison of the estimated ratios of human intrusions per hour of observation on weekends verses weekdays for each of the four sites is presented in Table 2. There was no difference at the 0.1 level of significance in the number of human intrusions per observer hour on weekends versus weekdays at Boulder and Chatfield heronries for the months of March and April. During the months of May through July, the number of intrusions at Boulder on weekdays was significantly higher at the 10% (0.1) level than the number of intrusions at Chatfield. A comparison of the May through July estimated ratios of weekend verses weekday intrusions between Boulder and the heronries at Fossil Creek and Lone Tree demonstrated no significant difference at the 10% (0.1) level. A comparison of the ratio for Boulder on weekends with Lone Tree on Memorial Day weekend demonstrated a significant difference in the number of intrusions at the 1.0% (0.01) level. A comparison of Chatfield with Fossil Creek and Lone Tree also demonstrated no significant difference in the number of intrusions on weekends or weekdays at the 10% (0.1) level. In addition, a comparison of Fossil Creek with Lone Tree demonstrated no difference at the 10% (0.1) level of significance in the number of intrusions on weekends or weekdays. The estimated ratio of human intrusions per hour entire observation period (Ma~ch - July) for the compared to the ratio of intrusions during May Fossil Creek and Lone Tree is presented in Table of observation for the Boulder Creek heronry July for the Chatfield, 3. Again, the ratio of intrusions on weekdays at Boulder was significantly higher at the 10% (0.1) level than Chatfield. The ratio of intrusions on weekends at Fossil Creek and Lone Tree was also significantly higher at the 10% (0.1) level than the ratio on weekends at Boulder. There was no difference at the 10% (0.1) level of significance in the number of intrusions on weekdays between the Boulder Creek, Fossil Creek and Lone Tree heronries. �97 Table 2. A Comparison of the Estimated Ratios of Human Intrusions per Hour of Observation on Weekends Verses Weekdays for Each of the Four Sites. Boulder WEEKENDS LEVEL OF SIGNIFICANCE VERSES: Chatfield (March-April) Chatfield (May-July) Fossil Creek (May-July) Lone Tree (May-July) Lone Tree (Memorial Day) -0.0563 ~ 0.2078-0.0588 ~ 0.3543 -0.3293 ~ 0.4333-0.1284 ~ 0.9391 -0.2449 ~ 0.800-0.1284 ~ 1.5881 -0.1407 ~0.3099-0.1284 > 0.5037 0.3046 ~ 2.256-0.1284 ~ 3.9506 Boulder HEEKDAYS NS NS NS NS 0.01 LEVEL OF SIGNIFICANCE VERSES: Chatfield (Harch-April) Chatfield (May-July) Fossil Creek (May-July) Lone Tree (May-July) -0.0557 -0.2901 -0.3484 -0.504 ~ 0.1151-0.0 ~ 0.2859 ~ 0.0312-0.1813 ~ -0.0101 ~ 0.2727-0.1813 ~ 0.5312 > 0.3372-0.1813 > 0.8158 - Chatfield WEEKENDS NS 0.1 NS NS LEVEL OF SIGNIFICANCE VERSES: Fossil Creek (May-July) Lone Tree (May-July) -0.4208 -0.5744 > 0.800-0.4333 ~ 0.3099-0.4333 > - 1.1542 ~ 0.3276 NS NS LEVEL OF SIGNIFICANCE Chatfield WEEKDAYS VERSES: Fossil Creek (May-July) Lone Tree (May-July) -0.1965 -0.3403 ~ 0.2727-0.0312 ~ 0.3372-0.0312 ~0.6781 ~0.9523 Fossil Creek WEEKENDS NS NS LEVEL OF SIGNIFICANCE VERSES: Lone Tree (May-July) -1.1691 ~ 0.3099-0.800 ~ 0.1889 NS LEVEL OF SIGNIFICANCE Fossil Creek WEEKDAYS VERSES: Lone Tree (May-July) -0.8932 ~ 0.2727-0.3372 ~ 0.7642 NS �98 Table 3. A Comparison of the Estimated Ratio of Human Intrusions per Hour of Observation for the Entire Observation Period (March-July) for Boulder Heronry and the May-July Observation Period for Chatfield, Fossil Creek and Lone Tree. Boulder WEEKENDS LEVEL OF SIGNIFICANCE VERSES: Chatfield (May-July) -0.0851 Fossil Creek (May-July) 0.0594 > 0.800-0.0807 > 1.3792 0.1 Lone Tree (May-July) 0.0331 > 0.3099-0.0807 > 0.4253 001 Lone Tree (Memorial Day) 0.4263 0.01 > 0.4333-0.0807 2.256-0.0807 > > > 0.7903 3.9243 Boulder WEEKDAYS NS LEVEL OF SIGNIFICANCE VERSES: Chatfield (May-July) -0.2908 > 0.0312-0.1813 > -0.0094 0.1 Fossil Creek (May-July) -0.2822 > 0.2727-0.1232 > 0.5812 NS Lone Tree (May-July) -0.4375 > 0.8655 NS > 0.3372-0.1232 �99 The difference in the number of intrusions on weekends at Boulder Creek compared with the number of intrusions on Memorial Day weekend at Lone Tree was significant at the 1.0% (0.01) level. Of the total human intrusions within 100 m (138 m at Chatfield) at the four heronries, 56.7% resulted in minimal or no response; 39.1% resulted in a localized response causing birds in the vicinity of the intrusion to temporarily abandon their nests; and 4.3% caused a generalized temporary abandonment of the heronry (Table 4). At Boulder Creek heronry, which experienced the least number of intrusions (n = 14) per total observer hour, 50.0% of the responses in Zone II resulted in minimal or no response (Table 5). Local response to Zone II intrusions (28.57%) was higher than Zone I intrusions (14.29%). The Boulder heronry also experienced the only Zone II general response (7.14%). Of all observed intrusions which occurred at Chatfield heronry (n = 26), 88.4% resulted in minimal or no response. The remaining 11.6% of the responses at Chatfield were in the local category. The Chatfield heronry experienced no general response to intrusions (Table 5). At the Fossil Creek heronry (n = 30),56.67% of the responses were in the local category; followed by 36.7% in the minimal or no response category. Fossil Creek had two Zone I general responses (6.66%), when birds throughout the heronry temporarily abandoned their nests (Table 5). The Lone Tree heronry had the greatest number of intrusions (n = 60) per total hours of observation. Of these total observed intrusions, 51.68% elicited minimal or no response; 45.01% elicited a local response and 3.33% caused the general response (Table 5). The total number of observed human intrusions by month for the four study sites is presented in Table 6A. Human intrusions were separated into the two zones and the three possible categories of response. There were no intrusions during the hours of observation for either zones I or II in the month of March. In April, only 2 intrusions occurred in Zone I. The highest levels of intrusions in Zone I occurred during the months of May (n = 34) and June (n'= 24). In Zone II, May received the most intrusions (n = 26) while April (n = 12), June (n = 12) and July (n = 14) had lower but similar levels of disturbance. Table 6B presents the percentage of each category of response for the months of observation. There was an equal percentage(50.0%) of minimal and local responses for April. In May, this value shifted as local responses (64.7%) increased and general responses (5.9%) were observed for the first time. Conversely, the percentage of minimal or no responses declined (29.4%). In June, this trend continued as local (70.8%) and general (8.3%) responses reached their season highs, while minimal or no response reached its season low (20.8%). In July, there were no observed local or general responses to human intrusion. The minimal or no response category contained 100% of the responses. �Table 4. A summary of the Percentage of Total Responses at the Four Study Sites to Human Intrusion by Zone and Response Category. Res)2onse Zone Local Minimal General I II I II I II -- 50.0 14.29 28.57 -- 7.14 26 .. 92 61.52 7,69 3.85 --- 36.67 46.67 10.00 6.66 Lone Tree 25,01 26.67 40.01 5.00 3.33 -- % of Total 12.98 43,72 27.2 2.5 1.8 SITE Boulder Chatfield Fossil Creek 11.9 Response/Zone % Total Response/Category 56.7% 39.1% 4.3% •.... 0 0 �101 Table 5. The Percentage of Response of Herons to Human Intrusions in Zone I and II at each of the Four Study Sites. BOULDER CREEK HERONRY n = 14 Local2 Minimall II Response Zone I II 7.14% 14.29 o 7.14 21.43 21.1~3 o o o o 14.29 28.57 I General 3 I II TYPE OF HUMAN INTRUSION LAND o o o o Motor Vehicle Foot Motorcycle Horseback 7.14 14.29 o o o o o 7.14 o o WATER Unmotorized Motorized TOTAL %/Zone/Type of Response 50.00 CHATFIELD RESERVOIR HERONRY Response Zone n = 7.14 26 I Minimal II I Local II General I II TYPE OF INTRUSION LAND Motor Vehicle Foot Motorcycle Horseback WATER Unmotorized Motorized 11.54 15.38 30.76 30.76 7.69 TOTAL %/Zone/Type of Response 26.92 61.52 7.69 o o 3.85 3.85 o o o o �102 Fig. 5 Continued. n = 30 FOSSIL CREEK HERONRY Response Zone Minimal I II Local I II General I II TYPE OF INTRUSION LAND Motor Vehicle Foot Motorcycle Horseback 3.33 46.67 3.33 3.33 WATER Unmotorized Motorized 36.67 TOTAL %/Zone/Type of Response LONE TREE HERONRY 36.67 n = 6.67 46.67 10.00 6.66 60 Response Zone Minimal I II Local I II General I TYPE OF INTRUSION LAND Motor Vehicle Foot Motorcycle Horseback 11.67 1.67 6.67 1.67 21.67 1.67 6.67 6.67 6.67 15.0 3.33 5.0 3.33 1.67 25.01 26.67 40.01 5.00 3.33 3.33 \.JATER Unmotorized Motorized TOTAL %/Zone/Type of Response 1 2 3 3.33 Minimal or no response to intrusion-(birds did not leave the nest). Local response-(birds in immediate vicinity of intruder flew from nests). General Response-(Birds throughout the heronry left nests). II �]()3 Table 6A. The Total Number of Observed Human Intrusions the Four Study Sites by Zone and Response Category. by Month at ZONE --.-.I RESPONSE March April May June July Minimal 1 I) 10 5 6 Local 0 1 22 17 0 General 0 o 2 2 0 lONE II ._-_.--- _-_ March _______~ril RESPONSE --_ .. __ May June July Minimal 0 9 22 7 13 Local 0 2 4 5 1 General 0 1 0 0 0 Tab]~ 6B. The Total Percentage of Each Response by Month for Zones I and II. ZONt. I RESPONSE March ----------____:.:~= April Minimal ')0.0% May June July 29.4% 20.8% 100% I.oca1 o ')0.0% 64.7% 70_8% o Genera] o o 5.9% 8.3% o ___ II Zone ._._ . June July 84.6% 58.3% 92.9% 15.4% 41.7% 7.1% March April May Minimal 0 75.0% Local 0 16.7% General 0 8.3% RESPONSE .. _ --"-_ .. - --_._" 0 0 0 �104 In the month of April, the only Zone II general response (8.3%) was observed. In April (75.0%) and May (84.6%) the greatest percentage of responses fell in the minimal category, In June, the number of local responses increased from 15.4% to 41.7%. In July, the minimal category contained 92.9% of the responses. The percentage of human intrusions by foot, horseback, motorcycle or motor vehicle that yielded a local or general response at Boulder, Fossil Creek and Lone Tree heronries is presented in Figure 1. At Boulder heronry, both Zone I intrusions (n = 2) elicited a local or general response and 71.43% of the intrusions (n = 7) in Zone II elicited these responses. At Fossil Creek heronry, there was a major difference between the number of intrusions in Zone I (n = 16) and Zone II (n = 3). However both zones yielded 100% local or general responses. At Lone Tree heronry, Zone I experienced the highest number of intrusions (n = 21) of the three sites and had the lowest percentage (75.0%) of local or general responses to these intrusions. In Zone II, only 28.6% of the intrusions (n = 6) provoked a local or general response. The numbers of visitors to Chatfield Reservoir Recreational Area from March 1 through August 31 in 1979 and 1980 were analyzed. The total estimated people use of the Park in 1980 (n = 590,416) increased slightly over the 1979 season (n = 576,574). The months of March$ April and August demonstrated an increased people use in 1980 as compared to 1979. Visitation in May, June and July was lowered for the 1980 season, again because of the area being closed to boating in May and June. Total estimated visitation at Chatfield was separated into weekend (Saturday, Sunday and Holidays) versus weekday (Monday-Friday) usage for the March 1 through August 31, 1980 season. In March and April the primary people usage of the Park occurred on weekends (n = 28.224 and 48,989 respectively). In May and June, usage on weekends Cn = 10,198 and 17,055 respectively) and weekdays Cn = 28,069 and 20,803, respectively) declined. In July and August, there was little difference between weekend (n = 76,082 and 82,334 respectively) verses weekday (n = 85,063 and 78,207 respectively) usage. Overview Discussion Because of their location and adjacent land-use patterns, each of the four heronries possessed its own types and levels of human disturbance. The Boulder Creek heronry experienced the lowest level of intrusions and it is only accessible by land and had controlled access (private property). The Chatfield heronry was only accessible by water with use during the nesting season restricted within 100 m - 138 m of the actual heronry. The heronries at Lone Tree Reservoir and Fossil Creek Reservoir were accessible by both land and water and access to both was not well regulated. In March, the herons first arrived and began pair-formation Neither the Chatfield nor the Boulder heronries experienced human intrusions during this month. and courtship. any observed �105 - 100% ~% UJ en z: 0 a... en UJ 0:.- 80% _J - ~ UJ z: UJ (.!;' - 70% 0:: 0 _J « S •......• « •......• (!) -:3 >z UJ en z: 0 en Em ., I I Cl z: « •......• 50% en UJ z: 0 N ! llO% I z: ::> 0:: ~ •......• ?JJk f-- z: ~ .:c LL 20% 0 UJ ~ I- z: UJ 10%- U 0:: UJ CL ZONE I II Boulder I II Fossil Creek I II Lone Tree TOTAL Nl..MBER OF HUMAN INTRus IONS Figure 1. The percentage of human intrusions by foot, motorcycle, horseback or motor vehicle which yielded a local or general response at Boulder Creek, Fossil Creek and Lone Tree. �106 In April, the herons incubation. At this occurred in spite of of Parks prohibiting began nest building followed by egg-laying and time, intrusions into the heronry at Chatfield the buoy line and regulations of the Colorado Division entry into the heronry. On May 4, 1980 young herons were first observed at Chatfield. By the end of the month, young were observed at all four study sites. Generally, while the chicks were small, one adult remained at the nest with the young while the other adult obtained food. On May 8, 1980 Chatfield was closed to all boating and remained so until June 28, 1980; during this period, the herons were relatively free of all human disturbance. In contrast, the Lone Tree heronry experienced its highest level of human intrusions per hour of observation in late May. This peak occurred over Memorial Day weekend and included a variety of activities. By June, the young were generally observed to be alone on the nests as both adults sought food for them. The number of intrusions at Boulder remained stable and low. At Fossil Creek, these intrusions increased as nature watchers, photographers and asparagus hunters used the area directly under nest trees. Lone Tree did not experience the degree of human disturbance that it had in May. By July, most young herons had left their nests. The heronries were virtually empty and thus no longer provided a subject for photographers and nature watchers. At Lone Tree, the water under the old site had receded making access by boat impossible. In addition, vegetation on the causeway had obtained heights of ~ 1.0 m, making travel across it difficult. The response of herons to human intrusions varied in degree of intensity and often depended upon the location of the intrusion. Generally, a Zone I intrusion elicited a higher level of response than a Zone II intrusion. For all four sites, 56.7% of all human intrusions elicited minimal or no response from the birds. Only 4.3% of all intrusions provoked a general temporary abandonment of the heronry. Intrusions which occurred in Zone II elicited weaker responses than Zone I intrusions. An intrusion which occurred in Zone II might not be perceived by herons throughout the heronry thus eliciting a less intense response. A general response was usually the result of unusually loud (eg. shooting, motorcycles) or highly visible disturbances which were perceptible throughout the heronry. The Boulder heronry received the least number of observed intrusions, (n = 14) of which 50.0% elicited minimal or no response. These were generally farm-related disturbances involving horseback, motorcycles and trucks and the herons may have habituated to them (Grubb 1978). �107 Chatfield received only boat-related disturbances of which 88.4% had minimal or no response for Zones I and II. Intrusion by boats at Chatfield elicited 7.69% local flights from nests in Zone I and only 3.85% in Zone II. Herons at Chatfield never experienced a general temporary abandonment of their nests as a result of intrusion. Most boating intrusions into the heronry were for the purpose of fishing and were quiet and slow-moving. The louder, faster motorboats were not permitted in the general vicinity of the heronry. The greatest percentage of response (46.67%) at Fossil Creek occurred in Zone I when herons flew from their nests as people walked under nest trees. Herons at Fossil Creek experienced a high level of Zone II motorboat and waterskier traffic. These disturbances were generally ignored {36.67% minimal response), and elicited the local flight of birds on only one observed occasion (6.67%). Grubb (1978) states that Great Blue Herons may habituate to noise from outboard motors while Parker (1980) observed disturbance of herons by excessive canoe traffic. Lone Tree received the highest number of intrusions (n = 60), but 51.68% fell into the minimal or no response category. Most minimal disturbance occurred when boats paddled under the trees. Again, herons were less tolerant of people walking, riding horseback or motorcycles under the nest trees. The response of the herons to intrusions as the breeding season progressed. was not constant, but altered Intrusions in Zone I elicited a 1:ligherpercentage of flight (local or general) responses as the season progressed, increasing until the young left the nests in July. In Zone II, the birds appeared Zone I throughout the breeding of minimal or no responses. less sensitive to intrusions than in season as evidenced by the high percentage The number of human intrusions by boat, motorcycle, horseback and motor vehicle which yielded a flight (local or general) response at Boulder, Fossil Creek and Lone Tree were compared to determine if the herons could be habituating to these intrusions. Intrusions into Zone I elicited a 100% flight response at Boulder (n = 2) and Fossil Creek (n = 16) while Lone Tree (n = 21) experienced only a 75.0% flight response. Intrusions into Zone II at Boulder (n = 7) and Fossil Creek (n = 3) elicited a 71.4% and a 100% flight response, respectively. At Lone tree (n = 6), Zone II intrusions provoked only a 28.6% flight response. Although the sample sizes were too small to prove any significant difference, it may be that the birds at Lone Tree are not as responsive to human intrusions because of their higher human activity level. In summary, previous evidence has indicated as the Great Blue Heron, are more sensitive that colonial birds, such to human intrusion early �108 in the breeding season (Dusi 1979, Palmer 1976 and Parker 1980). At this time, disturbance could cause adults to abandon nests and newly laid eggs. Yet in this study, the birds remained sensitive to human intrusions throughout the breeding season, well into the month of June. The most severe response to intrusion that was observed involved the temporary flight of birds throughout the heronry from their nests. However, this was still a less drastic response than the permanent, complete abandonment of the heronry. Information supplied by the Division of Parks revealed that human use of Chatfield Reservoir increased throughout the summer of 1979 and appears it would have done likewise in 1980 had the Park remained open to boating. As the number of people using the Park increased so did the number of intrusions into the heronry. It appears that herons may be able to tolerate more human disturbance than had been previously expected. It is also possible that heronhuman interactions occur less frequently because of temporal differences in the activity partern of each. Early in the breeding season, the birds were relatively disturbance-free. Human use of Chatifield in March and April was concentrated on weekends leaving the remainder of the week with few disturbances. High levels of human intrusions did not occur until after the young had hatched and were actively being fed by the adults. At this late stage, the adults may be less likely to abandon their young. The daily activity pattern of the herons generally did not coincide with human recreational activity patterns. While the birds were most active in early morning and late afternoon, human use of the area appeared to peak around midday. Literature Cited Dusi, J.L. 1979. Heron colony effects on man, Proc. Colonial Group, 1979; Vol. 3: 143-144. Waterbird Graul, W.D. 1979. Population Surveys of Selected bird and mammal species in Colorado. Colo. Div. W~ldl. Job Progress Rept., Proj. FW-22-R, 26pp. Grubb, M.M. 1978. Effects of increased noise levels on nesting herons egrets. Proc. Colonial Waterbird Group, 1978: 49-54. Palmer, R.S. 1976. Handbook of North American Press New Haven. 567pp. and Birds, Vol I. Yale Univ. Parker, J. 1980. Great Blue Heron in Northwest Montana: nesting habitat use and the effects of human disturbance. M.S. Thesis, University of Montana. 82pp. Ryder, R.A., W.D. Graul and G.C. Miller. 1979. Status, distribution and movements of Ciconiiformes in Colorado. Proc. Colonial Waterbirds Group, 1979: Vol 3: 49-58. �109 Small Mammal Segment The objective of this segment was to clarify habitat affinities of the following select species: Perognathu8 flavesceus, Perognathus flavus, Perognathus hispidus, Reithrodontomys montanus, and Permyscus leucopus. STUDY AREAS Site Localities: Las Animas County: Mesa de Maya; T33S, R56W. Yuma County: Bonny Reservoir; T5S, R43W. Ungrazed Sand Sage: Section 21 Ungrazed Shortgrass: Section 27 Grazed Shortgrass: Section 10 Grazed Sand Sage: Section 14 Little Bluestem: Section 22 Baca County: Ungrazed Site: 4!zmiles south Vilas T31S, R45W Sec. 25. Grazed Site: 6 miles south, 9!z miles east Campo T35S, R44W Sec. 5. The Little Bluestem were all cultivated community of Yuma County, and the two Baca County sites at one time. The ungrazed site in Baca County was reseeded in 1958 with sideoats grama, sand dropseed, blue grama, crested wheat grass, and sand love grass. The grazed site in Baca County was reseeded in 1960 and in 1965 with sand dropseed and sideoats grama (U.S. Forest Service, unpublished data). It is unknown when the little Bluestem community was reseeded, but long-time residents maintain that this site was under cultivation in the "dust-bowl" days. The remaining Yuma County sites have not been cultivated during the last 30 years if they indeed ever were. Characteristic Plant Species: 1. Mesa de Maya grassland 1. Western wheatgrass (Agropyron smithii) 2. Blue grama (Bouteloua gracilis) 2. Mesa de Maya forest (1. Pinus-Juniperus-Quercus) 3. Yuma County -Ungrazed Sand Sage 1. Sand sage - (Artemisia filifolia) 2. Needle-and-thread - (Stipa comata) -Grazed Sand Sage 1. Sand sage (Artemisia filifolia) 2. Blue grama (Boutelous gracilis) 3. Sand dropseed (Sporobolus cryptandrus) �110 -Ungrazed Shortgrass 1. Snakeweed - (Xanthocephalum sarrothrae) 2. Blue grama - ~uteloua gracilis) -Grazed Shortgrass 1. Blue grama (Bouteloua gracilis) 2. Buffalo grass (Buchloe dactyloides) -Little Bluestem 1. Little bluestem (Schizachyrium scoparium) 2. Side oats grama (Boutelouacurtipei1dtifa") 4. Baca County -Ungrazed 1. Sideoats grama - ~uteloua curtipendula) 2. Big bluestem - ~dropogon gerardii) -Grazed 1. Sideoats grama (Bouteloua curtipendula) 2. Sandsage (Artemisia filifolia) Discussion Perognathus flavesceus - The ecology of this species in Colorado is poorly understood. Armstrong (1972) described typical habitat for it. In this study P. flavesceus was captured most frequently in the ungrazed sand sage site (7.05 per 1000 TIN). This habitat type was characterized by low basal cover of grass, and low basal cover of bare ground. This supports the findings of Maxwell and Brown (1968). Perognathus flavus - In this study P. flavus was trapped most frequently in the grazed shortgrass site. This site had a high percentage of basal cover of non-grasses « 1%). Maxwell and Brown (1968) found a similar habitat affinity for this species in Wyoming. Perognathus hispidus - This species occurred in all but three sites during this study. However, Perognathus hispidus was captured in each of these three areas in previous years. In terms of microhabitat affinities P. hispidus showed a similarity to Perognathus flavus. This is in contrast to the findings of Maxwell and Brown (1968) who found a greater similarity between Perognathus hispidus and Perognathus flavesceus. In as much as the preliminary data showed ~. nispidus to occupy the ungrazed sand sage site in 1978·, it seems possible that the absence of this species from this site in 1979 may reflect a weakness in my trapping techniques, or a gross change in population size. In either case a large sample of ~. hispidus may result in a closer similarity between Po hispidus and P. flavesceus in terms of microhabitat. Reithrodontomys montanus - This species is an inhabitant of grassy environments (Hill and Hibbard, 1943; Armstrong, 1972). Maxwell and Brown (1968) captured this species with greatest frequency in a Bouteloua:-stipa community. This species was captured most frequently in short grass communities before this study. In 1977, six individuals were captured in the area that was used in this study for an ungrazed shortgrass site. In May 1979, a single specimen of R. montanus was collected in a similar exclosure·of ungrazed shortgrass. �111 However, in this study despite a 10,600 trap/night effort only six more individuals were collected. Of these six, three were taken in the ungrazed Baca county site. This site might be the most grossly disturbed site in this study. It has a history of "blow-outs" and cultivation. (See site descriptions for details on reseeding). Because of the small sample size herein, microhabitat be viewed in preliminary terms. affinities should only Peromyscus leu copus - This species inhabits the woodlands of eastern North America, but it also occupies brush communities in southeastern Colorado (Armstrong 1972). During this study, ten individuals of this species were collected in the ungrazed site in Baca County. Another individual was collected on Mesa de Maya. The ungrazed Baca County site was characterized by a tall-grass element (Andropogon). Grant and Birney (1979) reported ~. leucopus from three sites in North American grassland. Of these three, two are tall grass areas, whereas the third was a midgrass area. Literature Cited Armstrong, D.M. 1972. Distribution of mammals in Colorado. Mus. Nat. Hist. Univ. Kansas No.3 pp. i--x+1-415. Grant, W.E. and E.C. Birney, 1979. Small mammal community North American Grasslands. Jour. Mamm. 60:23-36. Monogr. structure in Hill, J.E. and C.W. Hibbard. 1943. Ecological differentiation between two harvest mice (Reithrodontomys) in western Kansas. Jour. Mamm., 24:22-25. Maxwell, M.H. and L.N. Brown. 1968. Ecological distribution of rodents on the High Plains of eastern Wyoming. Southwestern Nat., 13:143-158. Report Michael Statewide Segment Prepared by: P. Moulton Bat Study Segment This project was designed to document distributional patterns and habitat associations of bat species in Colorado. Additionally, this project provided the opportunity to gather basic ecological data (food habits, reproductive patterns, relative abundance, sex ratios, and parasites) on several Chiropteran species which previously have been the subject of little or no research in Colorado. The project was divided into three phases: 1) the documentation of distributional patterns of all Chiropteran species, 2) the status (number, sex ratios, breeding) of a colony of the Brazilian free-tailed bat, Tadarida brasiliensis, and 3) the detailed analysis of habitat associations of a select group of species (Myotis volans, ~.- evotis, !1_. californicus, !1_. leibii, Eptesicus fuscus, Lasiurus cinereus, and Antrozous pallidus) in northwest Colorado. �112 During the Chiroptean breeding season of 1978, 1979, and 1980, more than 630 individuals of 14 species (Table 1) were taken; more than 300 have been prepared as standard scientific specimens. Two species (Lasiurus borealis and Tadarida macrotis) of rare occurrence in Colorado (Armstrong, 1972) were not captured. Unfortunately, three species (Myotis velifer, Plecotus phyllotus, and Euderma maculatum - an endangered species) of probable occurrence (Armstrong, 1972) were not found. Range extensions of four species will be discussed below. Figure 1 indicates areas trapped. Myotis yumanensis (Fig. 2) previously know from only the most northwestern and southwestern counties in western Colorado was taken in several locations extending along the western border of the state. Yuma Myotis should be regarded as occurring along the entire Western Slope at lower elevations. Note that Figures 2-5 show previously described ranges (shaded). The maps are from Armstrong (1972). Myotis californicus (Fig. 3) was captured in northern and southern Moffat County, and in Rio Blanco County. These data extend its range to cover lower elevations along the entire Western Slope. Pipistrellus hesperus, the western pipistrelle, previously known to occur along the western border as far north as the Colorado River Valley was captured northward along the White River in Rio Blanco County (Fig. 4). The collection of Antrozous pallidus, the pallid bat, in southern Moffat County and in Rio Blanco County (Fig. 5) also indicates that it occupies lower elevations along the entire Western Slope. A colony of 5,.000Tadarida brasiliensis, the Brazilian free-tailed bat, in the San Luis Valley, Saguache County, was reported previously. This colony now numbers between 100,000-250,000 bats. Juveniles and lactating females were captured in 1979. Juveniles, lactating females, and females wf th embryos were taken in 1980. This is the fartherest north breeding colony of th Ls species east of the continental divide. Populations of this species have been declining dramatically elsewhere. Data show that pesticide poisoning has contributed to this decline (Geluso et al., 1976). Due to the importance of any breeding colony of this species, we have sent fecal material and bodies to Patuxent Wildlife Research Center for pesticide analysis. We are awaiting the results at the current time. Myotis thysanodes, the Fringed t~otis, was captured only twice; one in Rio Blanco County (two individuals) and once in Mesa County (one individual). It seems as suspected by Armstrong (1972) "that M. thysanodes is not especially connon at the edge of its range."No major range extensions for other Coloradan species have been documented. As part of this Freeman, J. and Portions of the Wildlife Society study, the following publication has been completed: S. Bissell, 1979. Bats. Colorado Outdoors, 28(2): 1-5. data will be presented to the Colorado Chapter of the and to the Colorado-Wyoming Academy of Science in 1981. �113 Table 1: Number of individuals Species caught per species. No. Caught Myotis lucifugus 6 Myotis yumanensis 21 Myotis evotis Myotis thysanodes 113 2 Species No. Caught Pipistrellus Eptesicus hesperus fuscus Lasionycteris noctivagans Lasiurus cinereus Myotis volans 65 Plecotus townsendii Myotis californicus/leibii 65 Antrozous pallidus Tadarida brasiliensis TOTAL 22 59 54 48 8 53 121 637 �-../'---:";,.7), FdFFAr------ ---- ---- I /, ~ . -r ._-. ''',RIO 8LANCQ'-":::::" L r- __ ~! ~\ Cllulr' ~WAl~'t: / ...• .".l "0_ _ Y<lmpa •._, --" /" g • hr I ., , I 'M , w'!Y • ._ .-00. novm , A~NOUFi . , I } _._._~ 'MoNTEZ'fJii-ii II ~ ,o ~r,IIo"".l'u (.,."'s,, 0 J • ,., ~,.--), •.....~ ~r- i r ., '1 .)4 "ON ~ ._. ). I. ."... I / \ rr,"~.'!YI4'.I • .,,' ,{ Figure 1. ' 1 • It;' -" L_ ~,.. L ."".;1, r.J I' .~,._. ( ,_ _j''Wf'''' ._._. °8 - " O(jr:u}f:lplul/N"~ · \ .' 1.ALAMOSA "," I• ""'''0' OR •••• olh"p,,/,I, "" .. ~:: Pam•• . ~_______ ° r~O.' . ' ON•• '''''m''' $/,., r,a,.} _ ••••. ~ •. _____ .. 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"""./ I ~ 1I,,-, O!'" / '-'L /~ "'••. 1"'''' •..• _ " ../ ._ r _/._ "-' ..../ ~ • 1 '\ .~./ , ."• ~--i .' ~Wo - / ,/ 0"'" ' 1 ,,•• - .-. 1 \.@S!£VEPCWFI,/,··_·c,;;,,,hsw-.,.,,,,wt_._.-1 ~.. ' - 1 . \ ,_. /" .-" ( orad., • Rvsbo .S ;{ .s. , I Y __.- .- ._._._ ."6",,J.,_._. ,-:.._. . ,'?;iI A:;r<· I '.- ,-" ,~-~ I !17_.A..•.'.••J.••.•,,_. ivNCOCN $,ml, .,~. '\ F<'k{y 0 1 ._,- • - COLORAOO Sill/1M L. o -.~' _. ° I _....., ' ._ "' , /' 1' ,,00'/' w"/o." ","",... '£C"'-,--'''"'".;'''}''. PIO"'" 1• o)S,I, oN_' i N,.,JI 0 .,' ._._. • C~'PP"@ 'l _J w. ,(."'.' _.-",X,lIso", .., • _ 0 ' ••••• r" 1/oi'J - •~ •.••• _._.-. o· 0M f"''' .•••.••l: -'60~" --.. •• ~". I 0"""0 ~ ••• ". . F'~ 6 t""f'" ••_ -"I ,•• • h. \ ' I ~,' ._,!!!{onoo~"'" oeM/O' _'... j' -·T.·-,.-n,opr.·-·-·\ ElNORTe I _M' 1~~, ••.•1 \,' s.•• ,,,,·, " .._. . . f-i'>'>"'o ._! -=._._ ...l-._. L ' - _- osrvsh" , ~~' ~." _.;.r.... .-.~ .~.. ~~ e .1 _. _-. . ._ ••• ." "WO'_' ,FREMONT' -~-'I __ ..RloaiiA"-·-· ,""'" <'.-t'-'L' NDE P...... / - P'~·"'ff.ol"( ,,;).ryo \ w R Am/omlo: . L --'-'r-:l _L • . ~""""iw'-' 0'9 JPHi .• ~", U7fiJA}"ELB •• ~ 7P /~ error<. ._._j' I -' rolo "II,a,,,,O ) ( I~~'m L ...." 0 ,'~ / v " ~ M'N"~ '" ."""~N 1(0',1 " PlA1'" ° ' ,,,,._. -, ~" ~ • ......._ ......,.\. m"e\ I' OW'O Nalh . 1 +" J'~' ' B"I"... -- ~."'I.,. " 'l I .~ ~"'t7(1 ______.. /. 1 • I .'_woo. ,0 ;. 0'"" ",NV,. co"' "m•• . !! 1 '~M'?'-" ~"fH;'-" i • . ~L> ra l.~oo. <. T'-. (' '. _._ • r..,vpO I ! • ) .~.'~ .,;. ~A J od$woc' 1 -~, ~~. '''"''''-''''"~''_-'_'l' .-. _... ,.- . o·,/d·.. -~ "''') CAm•• oc~ I ~ ..' a"""il",L... '. J•• , - ,...~ --J.1 " . " "SSUR6, I••.• oKet u6U1'9 ._L. .'''HrOr • ~ .';';;11. ~ J"7mM''''ry" ..". c•• .-,. "_'. l1 0" ,. .,,,,,Vo ,,,,w.•)~.,,,. '.'."~' fit,,,-,,,i(, ,_ I .' -_ • ., N/i?lh'@LEA4Y1LI.FI,.)I""W'_oo -_ .,~ ••i <s: 4 '7,.. 'or. \ ( J' ,D ".'I 1"',"<eNE'- ._._L./< ,- 1-....~~G/6 • c • "-, "''''''''O@TFtwnG.>.''l'' ~I ___ _,..;x_" j,-- <.~ _, - _', " _. .' ~.~ / '\ !<H',,* " 04."',;" . . !. .....,,_ '\" / ©am".,."," -i' «"oct \film' """"00 I ~_/ cr.wr",,,,, , f ',••.. '-. N '81)![.'J,ERSON • MM -_ "O!: ASPEII 1. /fillf5/""'· ""£CK~"OG£ •• ., ~!IT ',--'''' ~.. -' .,,' . I ..' ORr" , • :?rr: )_ f' if o.l("m~" .~. "I • _ '-'_'"'1 • O;q~ES e ._. _ • ,'0 ~O'L_L. _•.~f, • •• ".., 0OUI oNvd. , ,.J ~ ., ~l r _. , ••••••• __ DElTA J' . ON'PPOSE _. O{/.~' /" . , e(~ " ._. I r ._.... ~ .-- - ~ :f"'"~'',,_.' Jl;7;)C.v;;;-.,v:;;, r ,···;::c ") \ ( !IG=Nd...-0",,61."'! .r"" DELTA ,/ • ,r t f "'''S" . ./,,/ Wh,/,w,l" ._ ~._ ao<~.' oIir"';r':plon . ",o>o",_·_j·· -- _.. oH",d",M P_oo , ••••• mo.:.;~'.I .". \...{"'" - ~. _'.. .'00' -- .,' «oe (..... 1'."' . ,s,mu/o "'- " 1 -_. SI.1'ham --, ~ II tiL·' • • 08"H"Shom '01. ..... .-- r..." d~, ~ __,-, i! ,,:-. .. o",'"v";, •._ ...- " ~.. oJ,d/wal,,\ ,,\0,..' .' ll,dChRi OM.h"o' Md .. ,'-. , .oo I'armonl 08"99 Avllo ••----~-- c,..,~~ ""0 1 1 1 ,,_Mo -- • !~"'._~ .,,,",, OLLINS /' ,r"'Ms~ ..... ,.,.an's,ttlA"fi"f', , _ ~ If 00 L I "I (irandl r oJIi/O .1\.........'--' . '_ v. • <, '00'_ " -'rna". .."..-, P.. "'OM ~ a.:= r""PPI6ur 1GRANO.~.~~ I, .SO",","",·'_' 8'''~::_ I 0""",,'1. ,. mo/(,~ .1 ~.' ' \_, S~kl "'.. • _. ~_., a,!:H'tJR J w",' o: ,__. .'Am _._, 1 ..:;. .'__ 0 tnJ .~. ' oOo"r ,HM'" ••••• I 'LOGAN oK"flinan _ . ' ' d",,,rJ '''Sl' "'" '>-- \. ","0 ,_ r' • ~" ------- 06ro' "'"mo" ",\" ,;:, ~' ,\" \_~' s.)f//;1a0Ar lP" •_ oG""''''J "..., ~ ob,o'oT" " • __ .__ .__.__ 7 .~ ~ \7ARiMER"-------'/ "<, ~ OYI~";n-:;;;;;i;'·-1 WElO--o Herero,,>, cc orr \'-r O(j/~".)'rl!.r· - - ------o.d'~' ~ "' , OJ" If "- /(gmi"ono , ' ~ •• ,I ,I MAP OF COLORA"O li?o""liTr-----r--JACKSON'\. \'ocolvmh,:y\ ~_ _~ N.,.,,,,,,,, , .W_ ••• _ .. ""- or,I",h,,,. •.• .• I' SPRINGfiElD ,.i ,.,..... . I'1i·••.• - 0 - 4rull,1I •'" .- .M, oCampo 0 I �115 107 10' Ii ! I, ; i T~ 107 I()' Figure 2. Distribution (.) of Myotis yumanensis in Colorado. 3. californicus in l lOT 103 Figure 4. Distribution (.) of Pipistrellus hesperus in Colorado. 107 10' 105 !C3 "" Figure 5. Distribution (8) of Antrozous pallidus in Colorado. lOT �116 Currently, we are analysing food habits, reproductive patterns, and activity periods of bats in various habitats throughout the state. We have cooperated with the Denver Museum of National History by providing scientific and educational material for the development of a permanent exhibit on bats at the Museum. Literature Cited Armstrong, D.M. 1972 Distribution of mammals in Colorado. Mus. Nat. Hist. Univ. Kans. No.3, 415pp. Monogr. Geluso, K.N., J.S. Altenback, and D.E. Wilson. 1976 Bat mortality: pesticide poisoning and migratory stress. Science, 194: 184-186. This report segment prepared by: Jerry Freeman Overall report Prepared By: tI~ f). ~ Walter D. Graul Wildlife Research Leader �APPENDIX I ACTIVE GREAT BLUE HERON COLONIES IN COLORADO (1980) Location Years in Existence1 Number of Nests4 Tentative Status Information Source3 Decreasing* Carolyn Armstrong 1-5 Stable* Carolyn Armstrong 36-40 Other Nesting Species in Colony5 Adams County 1. Barr Lake (East) 15 mi. N.E. Denver TIS, R66W, Sec. 28, 20-25 Double-crested Cormorant SW~. 2. Barr Lake (West) 15 mi. N.E. Denver TIS, R66W, Sec. 27, SE~. 3. Horse Creek Reservoir 6 mi. S.E. Hudson, TIN, R64W, Sec. 31, SE~. 40 Double-crested Cormorant, Black-crowned Night •.... Heron •.... -...J Unknown Unknown D.O. 16-20 Double-crested Cormorant Black-crowned Night Heron, Great Egret Boulder County 4. Boulder Creek Between Hwy. 287 and 95th St., TIN, R69W, Sec. 16, SE~. 30-35 Stable D.O. 76-80 5. Panama Lake 6 mi. S.W. Longmont, T2N, R69W, Sec. 35 NE~. 40 Stable D.O. 1-5 Double-crested Cormorant �APPENDIX I CONTINUED Years in Existence1 Tentative Status Unknown Unknown D.O. 1-5 7. Colorado River 8.1 mi W. Rifle, T6S, R94W, Sec. 29, SW~. Unknown Stable D.O. 16-20 8. Colorado River at Grand Valley T7S, R96W, Sec. 13, SE~. Unknown Unknown 9. Colorado River at Grand Valley, T7S, R96W, Sec. 13, SE~. Unknown Increasing D.O. 11-15 Unknown Unknown D.O. 31-35 15-20 Stable)'( Location Information Source3 Number of Nests4 Other Nesting Species in Colony5 Eagle County 6. Gypsum T5S, R85W, Sec. 6, NE~. Garfield County 10. Colorado River 0.5 mi S.W. Silt, T6S, R92W, Sec 10 NW~. Howard Green 1-5 •.... 00 Grand County 11. Colorado River, 6 mi. E. Kremmling, TIN, R80W, Sec. 18, NE~. •.... Lloyd Palmer 11-15 �APPENDIX I CONTINUED Location Years in Existencel Tentative Status Information Source3 Number 0f Nests4 Other Nesting Species in ColonyS Gunnison County 12. Gunnison River 5.5 mi. SW Gunnison, T49N, RIW, Sec. 8, SW!t;. 11 Unknown D.O. Kevin Cook 21-25 Black-crowned Night Heron 6 Stable* John Wagner 1-5 Black-crowned Night Heron Jackson County 13. Walden Reservoir, 3 mi. W. Walden TI2N, R80W, Sec. 24, SE!t;. •.... •.... '.0 Jefferson County 14. Chatfield Reservoir 5 mi So. Littleton, T6S, R69W, Sec. 12. 80 Increasing* D.O. , Hugh Kingery 71 Kit Carson County 15. So. Republican Drainage 3 mi. S.E. Flagler 20-25 Stable D.O. 1•.. 5 Larimer County 16. Lone Tree Reservoir, 2 mi. NW Berthoud T4N, R69W, Sec. 9, NE!t;. 5 Decreasing D.O. Camille Cummings 41-45 Cormorant, Black-crowned Night Heron �APPENDIX I CONTINUED Location 17. Fossil Creek Res., So. Ft. Collins, T6N, R68W, Sec. 9, Years in Existence1 Tentative Status Information Source3 Number of Nests4 Other Nesting Species in Colony5 Black-Crowned Night Heron 10 Decreasing D.O. 61-65 5 Increasing D.O. 1-5 5 Increasing D.O. 1-5 SW-t; 18. Horseshoe Lake 1 mi. N.E. Loveland T6N, R68W, Sec. 31, SW-t;. 19. Timnath Reservoir 4 mi. N.E. Timnath, T7N, R68W, Sec. 24, SW!t:. •.... N 20. Wellington Res. #3 1 mi. N.E. Waverly, T9N, R68W, Sec. 18, SE~. 5-10 Stable Lee Franz 50 Stable D.O. 26-30 25-30 Stable D.O. 6-10 0 50-60 Logan County 21. Jumbo Reservoir 8 mi. W. Sedgwick, TIIN, R47W, Sec. 7, SE~. 22. Sterling Res. 14 mi. N.W. Sterling, T9N, R53\<],Sec. 1, NE~. Double-crested Cormorant �APPENDIX I CONTINUED Location Years in Existence1 Tentative Status Unknown Decreasing* Information Source3 Number of Nests4 Other Nesting Species in Colony5 Mesa County 23. Colorado River 4 mi. So. DeBeque, T9S, R97W, Sec. 18 SE~. D.O. 11-15 Moffat County 24. 13 mi. N.E. Craig, T7N, R89W, Sec. 3, SE!t;. 25. Green River !t;mi. S.E. Brown's Park National Wildl. Refuge 26. Yampa River 1 mi west of bridge that is north of Maybell. T7NR95W. 2 15 Increasing Stable* Chuck Woodward D.O. 1-5 11-15 Unknown Unknown Charles Haynes 2 Unknown Unknown D.O. Jennifer Slater 2 Prowers County 27. Two Buttes Creek 30 mi. So. Lamar, West end of State Mgmt. Area (Two Buttes). •.... N •.... ��APPENDIX I CONTINUED Location Years in 1 Existence Tentative Status Information Source3 Number of Nests4 10 Unknown D,O. 1-5 10 Stable D.O. 11-15 35. Empire Reservoir 3 mi. E. Masters, T3N, R61W, Sec. 1, NE!t;. 50 Stable 36. Franklin Lake 0.5 mi. S.E. Serverance, T6N, R67W, Sec. 1, NW!t;. 5 Increasing 33. Yampa River 5.25 mi. W. Steamboat Springs, T6N, R85W, Sec. 9, NW-t;. Other Nesting Species in Colony5 Washington County 34. Prewitt Reservoir T5N, R4W, Sec. 14, N~. Weld County 37. Johnstown 1.25 mi. N. Johnstown, T5N, R67W, Sec. 33, SW!t;. 38. Mil ton Res. 9 mi. W. Platteville T3N, R65W, Sec. 10, SE!t;. 60-65 Increasing 5-10 Decreasing Ron Zacagnini D.O. Brad Kirshner D.O. 41-45 •..... Double-crested Cormorant. Black- ~ crowned Night Heron 1-5 46-50 46-50 Double-crested Cormorant, Blackcrowned Night Heron �APPENDIX I CONTINUED Location 39. Riverside Res. 2 mi. N. Masters, T4N, R62W, Sec. 12, NE~. Years in Existence1 Tentative Status 10-15 Unknown 40. Barbour Ponds (New-1980) 300 mi E of 1-25 and due E of Barbour Ponds. T2N, R68W, Sec.2, Unknown Information Source3 D.O. Number of Nests4 1-5 Walter Graul Other Nesting Species in Colony5 Double-crested Cormorant 2 NW-t;. Yuma County 41. So'.Republican River 1.1 mi. E. Hale, T5S, R43W, Sec. 12, SE~. 5 Stable D.O. 1-5 1. The "years in existence" is based on interviews and/or questionnaires from people familar with the sites. 2. The asterisk indicates that the assigned category is based on detailed data (those without the asterisk are, therefore, tentative). 3. D.O. refers to direct field observation by personnel in this study. 4. This value was determined by direct observation and/or interviews, and/or questionnaires. 5. Same as 4. •..... N ~ �APPENDIX II INACTIVE GREAT BLUE HERON COLONIES IN COLORADO (1980) Location # of Nests Prior Probable Reason for Abandonment2 to Abandonment1 Information Source Boulder County 1. Wonderland Lake NW edge Boulder, TIN, R71W, Sec. 13. SE~. Inis Baker 1-5 Trees cut for housing development 2. East LyonsT3N, R70W, Sec. 20,N~. Inis Baker Unknown Commercial development •..... Delta County N V1 3. Gunnison River 4 mi. N. DeltaT1sS, R96W, Sec. 19, SW~. (Hamilton Property). Hal Burdick 6 Unknown 4. Gunnison River 1 mile downstream from Escalante Canyon Bridge. T1sS, R97W, Sec. 7. Dave Langlois 6 Unknown 5. Gunnison River 8 miles downstream from Escalante Canyon Bridge. T14S, R2E, Sec. 10. Dave Langlois I~ Unknown �APPENDIX II CONTINUED Location 6. Fruit Grower's Res. 1 mi. So. Eckert, T14S, R94W, Sec. 18, SW~. Information Source # of Nests Prior to Abandonment1 Probable Reason for Abandonment2 Merle Hodges 6-10 Unknown Howard Green 1-5 Highway Construction Ann Loughteridge 6-10 Corral construction Eagle County 7. 6 mi. W. Gypsum, T5S, R86W, Sec. 5, NE~. Garfield County 8. 2.5 mi. SE Carbondale T7S, R87W, Sec. 36, NW~. •..... N '" Jefferson County 9. Standley Lake 88th and 100th Ave. T2S, R69W, Sec. 20, SW~. Ann Abbott 1-5 Nests removed by people Larry Strode 6~10 Reservoir drained 1966, human activity at heronry. Eva Radenacher 1--5 Gravel pit and recreation area near the heronry. Kit Carson 10. SW Reservoir, 2 mi. SW Flagler Larimer County 11. Barbour Ponds 7 mi. E. Longmont T2N, R68W, Sec. 3, S!z. �APPENDIX II CONTINUED Location Information Source # of Nests Prior Probable Reasons for Abandonment2 to Abandonment 1 12. Bodecker Res. 3.25 mi. W. Campion, T5N, R69W, Sec. 20 NW~. Judy Sisler 1-5 Housing development nearby. 13. Boyd Lake 1 mi. NE Loveland, T6N, R68W, Sec. 32, NE~. Judy Sisler Unknown Housing development adjacent to heronry. 14. Terry Lake (Island) N. edge Ft. Collins, T8N, R69W, Sec. 26, SE~. Ronald Ryder Unknown Cutting down of trees 15. Terry Lake (Shore) T8N, R69W, Sec. 26, SE~. Rene Ellis 1-5 Unknown 16. Lonetree Res. 3 mi W. Campion, T4N, R69W, Sec. 9, NE~. Camille Cummings 75 Illegal shooting, recreational activity adjacent to area. Joe Gumber 1-5 Unknown •.... N '-I Mesa County 17. Colorado River 3.2 mi. So. DeBeque T9S, R97W, Sec. 8, SE~. �APPENDIX II CONTINUED Information Source II of Nests Prior to Abandonment1 Probable Reasons for Abandonment2 18. Colorado River 4 mi. So. Debeque T9S, R97W, Sec. 18, SE~. Joe Gumber 6-10 Highway Construction nearby. 19. Colorado River 2.25 mi. W. Fruita, TIN, R3W, Sec. 14, NE~. John Gray 21.•... 25 Highway constructed through heronry. 20. Colorado River 1.5 mi. W. Fruita, TIN, R3W, Sec. 13. SE~. John Gray 11-15 Unknown Location •..... N 00 Moffat County 21. Yampa River 1.5 mi. SW Craig, T6N, R91W, Sec. 11, NE~. Samuel Scanga 6-10 Unknown Dal Schaefer 6-10 Nesting trees cut with reservoir development, intensive boating. Charles Reichertt 6~10 Unknown Pueblo County 22. Arkansas River 4 mi. W. Pueblo, Pueblo Reservoir area. Rio Blanco County 23. White River 4 mi. E. Meeker, TIN, R93W, Sec. 19, SE~. �APPENDIX II CONTINUED ------------- ------------------ ---- -1. Size of heronries prior to abandonment was determined by interviews, questionnaires, and/or direct observation. 2. Probable reason for abandonment as determined by site inspection, interviews, and/or questionnaires. t-' N \0 �130 JOB PROGRESS REPORT State of COLORADO --~~~~~-------------- Project No.__~W_-~1~2~4_-~R~ Work Plan No. _ I --~------------------ Job Title Job No. ~1 _ Bald and Golden Eagle Nesting Studies Period Covered: Personnel: Raptor Investigations March 1,1979 through December 31,1980 Thomas Bohanon, Gerald Craig, Joe Frothingham, Cynthia Janik, David Langlois, Chuck Loeffler, Tom Lytle, l\layne Russell, Gordon Saville, Babette Tully, and Ted Washington, Colorado Division of Wildlife. ABSTRACT Nest site occupancy and productivity surveys of golden eagles were conducted in 1979 and 1980. In 1980, this job was reassigned to regional nongame biologists and results were presented on a regional basis. l~en taken on a statewide basis, the reproductive index was significantly lower in 1980. Lagomorphs accounted for 89 percent of the prey encountered at a sample of 25 productive golden eagle nests in 1979. In 1979, one bald eagle nest succeeded in fledging a single young and in 1980 two bald eagle nests produced one young each. �131 BALD AND GOLDEN EAGLE NESTING STUDIES Gerald R. Craig P. N. OBJECTIVES The objectives of this study are: (1) to estimate the breeding numbers and obtain production data of bald and golden eagles nesting in Colorado; (2) to identify important nesting areas and associated hunting habitats of bald and golden eagles in Colorado; and (3) to compile data and submit reports to associated state personnel and federal agencies for use in delineating and protecting eagle nest sites. SEGMENT OBJECTIVES 1a. Continue to locate and map nesting sites of golden and bald eagles throughout Colorado. Nest searches will be conducted with fixedwing aircraft and in some instances with helicopter. Additional field work will be done from the ground by vehicles. Photographs of nest sites will be taken and pertinent information recorded about physical features and the habitat. lb. Nesting areas will be stratified by such features as climate, elevation and habitat type. Sample areas then will be delineated which are representative of important nesting areas. The sample areas throughout the state will be flown with a fixed-wing aircraft in late April and early May to ascertain the number of active sites. The same sites again will be checked from the air in June to dete 1 ine the nesting success and productivity of the sample sites. The percent of active sites, percent of succesful pairs, and total production will be extrapolated for each region of the state. 2a. '~en nests are visited in la, details will be recorded as to characteristics of the nesting habitat. Nesting eagles also will be observed from a distance to locate and map key hunting areas. 2b. Radio transmitters may be 'attached to several young bald eagles and a sample of young golden eagles to follow their movements after they fledge. Color markers mayor may not be used to mark the eagles as well. 2c. A sample of sites will be selected and visited to determine the prey species present at the nest sites. This information will be valuable in assessing the prey composition and availability to eagles. 3. Analyze all data obtained and prepare a report of the findings. �132 METHODS AND MATERIALS The golden eagle nest surveys were conducted aerially, employing a Cessna 185 aircraft in the Northeast, Northwest, Southwest and Southeast regions of the state. Nests were located with the aid of descriptive materials, maps and photographs. Nests were inspected as the aircraft flew past at distances of 50-150 feet. As time permitted, accessible nests were inspected from the ground if inclement weather or scheduling difficulties precluded observation from the aircraft. No specific searches were made to locate new nests, but potential areas were inspected in route between known nests and any new nests which were encountered were recorded. Although considerable variability exists in hatching time, an effort was made to observe all nests on record by mid-May to determine the number of eggs or young produced before nest mortality began to take its toll. Active nests are defined as those observed to contain either an incubating adult, eggs, or young birds. Neither the presence of flying adults in the area of a barren nest, nor the presence of fresh nesting material on an unoccupied nest was considered evidence of activity, but did stimulate further search for an occupied alternate site nearby, and note was taken of these conditions. Observations included presence or absense of adults, condition of nest, age of the eaglets, and possible disturbance factors. New nests were plotted on 1.250,000 U.S.G.S. maps, described and photographed to aid in future location, and photographs and descriptive materials for old nests were added when lacking. Those nests which were determined to be active were rechecked in late May and the first 2~ weeks of June, depending on the age of the young birds, to determine the number of eaglets which would fledge. Young golden eagles fledge at 9-10 weeks of age, and birds over 7~-8 weeks were considered to have attained fledging age since mortality in the 8-10 week period is insignificant. RESULTS AND DISCUSSION Golden Eagle Nesting Results Table 1 summarizes golden eagle nest occupancy and productivity on a statewide basis for 1979. In 1980, responsibility for administration of this job was transferred to the regional nongame biologists and for future reference, will be published under project numbers, W-137-R, W-139-R and W-140-R. Tables 2, 3, 4, and 5 summarize nest occupancy and productivity for 1980 on a regional basis. Although the reproductive index was up in the northwest, it was lower than 1979 when considered on a statewide basis. Prey Remains in Golden Eagle Nests In 1979, 25 active golden eagle nests were visited to determine the prey utilized by the eagle. Table 6 summarizes the results of the survey. �133 Table 1. Statewide summary of golden eagle nesting productivity in Colorado, 1979. activity and 1. Nests on record. . . . . . • . . . . . . . . . . . . . .. 496 2. Nests not found . . . . . . . . . • . . . . • . . • . . .. 26 3. Nests not checked . . . • . . . . . . . . . . . . . . . .. 23 4. Nests checked. 5. New nests 6. Nests inactive. 7. Nests active. 8. Young produced. 9. Young fledged • . . . . . . . . . . . . . . . . . . . .. 444 found . . . . . . . . . . . . . • . . . . . • .. 26 . . . . . • . . . . . . . . . . . . . . .. 253 . . . • . . . . . . • . . . . . . . . . . .. 191 . . . . . . . . . . . . . . . . . . . . .. 248 . . . . . . . . . . . . . . . . . . . . . .. 218 10. Young produced per active 11. Young fledged 12. No. of known successful 13. Young produced per successful 14. Young fledged 15. Active nests per nests 16. Reproductive index. per active nest. . . . . . . . . . . . . .. 1. 30 nest 1.14 nests 154 per successful nest. . . . • . . . . . . .. .53 . . . . . . . . . . . •. 1.42 . . . . . . . . . . • . .. 0.43 nest checked. . • . . . . . . . . . . . . . . . . . . 61.06 �134 Table 2. Summary of golden eagle nesting activity and productivity Northwestern Colorado, 1980. in 1. Nests on record •••••••••••••.•...•.. ' 244 2. Nests not found • . • • • • • • • • • • • • • • • . . • • • 1 3. Nes ts not checked 4. Nes t checked. • • . . • . . . . • • . . • . • • . . • . .. 242 5. New nests found • . • . . • • • • • • • . . . . . . . . .' 38 6. Nests inactive. 7. Nests active •••.•••••••••.•••...••. , 114 8. Young produced .•..•••••.•.••.••••.•• ' 152 9. Young fledged ' 138 10. Young produced 11. Young fledged per active nest 12. No. of known successful 13. Young produced 14. Young fledged per successful 15. Active nest per nests checked. 16. Reproductive . . . • • • • • • • • • . . • . • • • • • • • • • • • • • • . • . • • • • • . . • •. ....•••••.•••.•••••.•• per active nest. 1 128 • • . • • • • • • • • • •• 1.33 ............... 1.21 nests 97 per successful nest. • • . . • • • . • • •• 1.57 nest • • • • • • • . . • • .• 1.42 • • • • . • • • • • . • •• index .•••••.••.•••.•..•••• 0.47 66.74 �135 Table 3. Summaryof golden eagle nesting Southwestern Colorado, 1980. activity and productivity in 1. Nests on record. . . . • . . . . . . . . . . . . . . . .. 176 2. Nests not found. . . . . . • . . . . . . . . • . . . . .. 17 3. Nests not checked . . . . . . . . . • . . • . . . • . . .. 14 4. Nests checked . . . . . . . . . . . . . . . . . . . . • .. 145 5. Newnests 6. Nests inactive. 7. Nests active. 8. Young produced. 9. Young fledged found . . . • . . . . . • . . . . . . . . • . .• 14 . . . . . . . . . . . • . . . . . . . . .. 106 . . . . • . . . . . • . . . • . . . . • . .. 39 . . . . . . . . . . . • . • . • • . . • .. 45 . . . . . . . . . . • . . . . . . . . • . .. 26 10. Young produced per active 11. Young fledged 12. No. of known successful 13. Young produced per successful 14. Young fledged 15. Active nests per nests 16. Reproductive index ..•..•••..•..•.•...•. per active nest 1.15 nest 0.67 nests per successful 21 nest. nest checked. • • . • • • . . . . .. 2.14 . . . • . . • . . . . .• 1.24 • • . . • • . • . . . . .. 0.27 33.48 �136 Table 4. Summary of golden eagle nesting activity Southeastern Colorado, 1980. and productivity in 1. Nests on record. • • • . . • • • • • • • • • • • • • • • •• 58 2• N es ts no t found • • • • • • • • • • • • • • • • • . • • • •• 0 3. Nests not checked • • • • . • • • • • . • • • • • • • • • •• 8 4. Nes ts checked . • • • • . • • • • • • • • • • • • • • • • •• 50 5. New nests found. 6. Nests inactive. 7. Nests active. 8. Young produced. 9. Young fledged 10. Young produced 11. Young fledged per active nest 12. No. of known successful 13. Young produced 14. Young fledged per successful 15. Active nests per nests checked. 16. Reproductive • • • • • • • • • • . • • • • . • • • • •• 0 • • . • • . • • • • • • • • • • . • . • • .. 31 • . . • • • • • • • • • • • • • • • . • • . •• 19 • • . • . • • • • • • . • • • • • • • . • •• 29 • . . • • . . . • • . • . • • • • • . • . • .• 26 per active nest. • • • . • . • • • • • • • •• 1.37 nests per successful 1.53 19 nest. nest. • • • • . • • • • • . •. 1.53 • • • • • • • • • • • •• 1.37 • • • • • • • • • . . • • • • index ••••.••••••••••••••••• 0.38 52.06 �137 Table 5. Summary of golden eagle nesting activity and productivity Northeastern Colorado, 1980. . . . . • • . • • . . • . . . . . • . .• in 1. Nests on record. 2. Nests not found • . . • . . • • • • • • • . • • . . • . . • 0 3. Nests not checked 0 4. Nests checked. 5. New nests found . . . . • • . • . . • . . . • . • . . • . • 6. Nests inactive. 7. Nests active. 8. Young produced. 9. Young fledged . . . • • • . . . . • • • • • • . . • . • • • . • • • • • . • • • . • • • • . . • .. 87 87 1 . • . . . • . • . • • • • . . . . . . • .. 49 . • • . . • . • • • . • . • • . . • . . • • . 36 • . . . • . . . . • • . • . • • • . . • •• 48 . . . . . . . • • . • • . • • . . • . . . .. 36 10. Young produced per active nest. • • . • • • • • . . • . •• 1.33 11. Young fledged per active nest • . • • • • • • . • . • • .. 1.00 12. No of known successful nests. 13. Young produced per successful nest. . • • . • • • • . • •. 1.85 14. Young fledged per successful nest . . • • . • . • . • • .. 1.38 15. Active nests per nests checked. 0.41 16. Reproductive 26 • . . • . . • • • . . . •. index. . . . . • . . . • . • • • . . . . • • . 75.85 �138 Table 6. Prey composition in 25 golden eagle nests, 1979 Prey Species If of Nests If of Items % of Nests % of Total Jackrabbits 25 289 100 87 Cottontails 5 6 20 2 Lagomorphs 25 295 100 89 Richardson~s grnd squirrel 5 7 20 2 Prairie dog 4 6 16 2 Marmot 3 7 12 2 12 20 48 6 5 11 20 3 All rodents Mule deer All mammals Birds 25 326 100 98 5 6 20 2 �139 Lagomorph remains consisted almost exclusively of lower legs and cottontail legs were distinguished from jackrabbit legs by size and color. The percent of lagomorph remains is slightly misleading in that each carcass potentially contributes four items (legs and feet) which tend to remain in the nest. Smaller rodents can be consumed whole and may not be represented at all. Remains of deer were usually lower legs of fawns, but 2 adult deer legs were found in one nest and an entire fawn in another. Avian prey remains consisted of black-billed magpie (2 items), 2 common flickers, and 2 unidentified. Bald Eagle Nesting Results In 1979, one pair of bald eagles in northwestern Colorado succeeded in rearing a single young from two which hatched. Shortly after the young hatched, the adult male disappeared and the female was forced to assume the duties of young rearing, foraging, and nest protection. The second historic nest in southwestern Colorado was occupied initially in 1979 by a lone adult but the adult was not successful in attracting a mate. An adult and juvenile were observed fishing along the river just north of Durango in July and although no nest was located it is probable one exists in the vicinity. In 1980, the historic site in La Plata county was vacant, but a first nesting attempt was recorded in Montezuma County. Two young were hatched and one successfully fledged from the site. The historic site in northwestern Colorado was occupied by an adult pair which reared and fledged a single young. Prepared by C. .(2.. ~ Gerald R. Cralg Wildlife Researcher C �140 JOB PROGRESS REPORT State of COLORADO Project No. \oJ-124-R ------~--------------- Raptor Investigations Work Plan No. I --------------------- Job Title: ~B~a=l~d~a~n~d~G~o~l~d~e~n~E=a~g~l=e=_W~in==t~e~r~P~oLP~u=l~a~t~i=o~n~S~u=rv~ _ Period Covered: Personnel: Job No.2 ---------------------------- March 1, 1979 through December 31, 1980. Gerald Craig, Joe Frothingham, David Langlois, Tom Lytle, Wayne Russell, and Ted Washington, Colorado Division of Wildlife. ABSTRACT Midwinter bald and golden eagle flights were conducted in census areas in northeastern and northwestern Colorado. The San Luis Valley study area was not censused due to weather and aircraft scheduling difficulties. �141 BALD AND GOLDEN EAGLE WINTER POPULATION SURVEYS Gerald R. Craig P.N. OBJECTIVES The objective of this study is to obtain winter population trend information for bald and golden eagles on selected wintering areas in Colorado. The information will be used to estimate wintering populations of bald and golden eagles throughout the state. SEGMENT OBJECTIVES 1. Aerial Counts of Wintering Golden Eagles: Once annually an aerial flight will be made on census areas in the San Luis Valley, northeastern and northwestern portions of the state. These census areas were established in 1972 and the procedures will be essentially the same. Random, north-south transects will be flown throughout each study area with a Cessna 185 aircraft and all eagles observed within ~ mile of each side of the transects will be counted and classified as adult or juvenile. Transects will be flown at speeds averaging 100-120 mph at altitudes of 100 to 300 feet, depending upon topographic features. From the transects, an area estimate will be obtained as to eagles per 100 square miles. Identical census areas are also flown in Wyoming, Idaho, Montana, North Dakota, New Mexico and Nevada by the U.S. Fish and Wildlife Service and cooperating agencies. Information forthcoming from these states are then collected and analyzed by the Fish and Wildlife Service to obtain population estimates for the West. Age ratio information which is obtained provides an indication of the previous breeding season's reproduction. 2. Aerial Counts of Wintering Bald Eagles: Since bald eagles tend to congregate primarily along river courses and impoundments, aerial flights will be made along major river courses throughout the state and a direct count will be made of all bald eagles observed. The eagles will be classified as to age in order to obtain information about reproduction. The flights will be made in January when the highest concentration of eagles are usually present. Flights will be made along the South Platte River, Yampa River, White River and Rio Grande River. It is proposed to expand the censuses to the Colorado River and possibly the Gunnison, Dolores and San Juan Rivers. 3. Compile data and prepare annual progress and final reports them to appropriate personnel and agencies. and submit �142 METHODS AND MATERIALS Midwinter golden eagle flights were conducted in census areas established in the San Luis Valley, northeastern and northwestern Colorado according to procedures described in Segment Objective 1. The San Luis Valley census area encompasses 2,500 square miles which is the majority of the valley lands. Transects account for a 10 percent sampling of the area. The northeastern census area which accounts for 3,000 square miles in Weld and Logan Counties is also sampled at 10 percent. The northwestern census area includes Moffat and Rio Blanco Counties and is the largest with 4,100 square miles. Linear transects account for 7 percent coverage of the area. Winter bald eagle counts are conducted in the San Luis Valley in conjunction with golden eagle flights mentioned above. Since bald eagles are generally distributed throughout the San Luis Valley, it is possible to obtain an effective area estimate of bald eagles during the same flight for golden eagles. Bald eagles are also censused in conjunction with golden eagle flights in northwestern Colorado. A direct count is made of the South Platte River between Fort Morgan and Greeley in Weld County upon completion of the golden eagle flight of northeastern Colorado. The course of tIle river is flown at altitudes of 100 to 200 feet and all eagles are counted along the river from Fort Morgan to Greeley. RESULTS AND DISCUSSION Results of the 1980 aerial flights for wintering golden eagles are presented in Table 1 and the winter bald eagle census results are summarized in Table 2. The number of golden eagles utilizing northeastern Colorado increased in 1980 to the highest since 1975 but decreased in the northwest. Weather conditions and aircraft scheduling difficulties precluded flights in the San Luis Valley. Responsibility for these flights have been transferred to the regional nongame biologists and will be administered and reported in the future under other federal aid projects. �143 Table 1. Golden eagle aerial census of Colorado, 1972-1980. Northeastern Colorado (10% sample of 3,000 sq. mi.) Date 1/24/1973 1/16/1974 1/22/1975 2/19/1976 1/13/1977 1/04/1978 1/24/1979 1/31/1980 - - - - - - Adults Juveniles Unknown 16 19 22 17 18 18 14 13 8 3 5 3 1 3 4 11 0 0 0 0 0 0 0 1 -- --- - - - --- Eagles per 100 sq. mi. - - - - - - 8.0 7.3 9.0 6.7 6.3 7.0 6.0 8.3 - - --- Est. of Total Eagles 240 220 270 200 190 210 180 250 - - -- - - - - Northwestern Colorado (7% sample of 4,100 sq. mi) Date 1/25 & 2/26,1972 1/23/1973 1/22 & 1/29,1974 1/29/1975 1/26/1976 1/19/1977 1/19/1978 2/13/1979 2/06/1980 Adults Juveniles Unknown Eagles per 100 sq. mi. Est. of Total Eagles 86 35 6 11 29 29 23 46 38 80 14 8 8 3 5 5 25 27 97 70 47 17 10 2 0 3 2 91.6 41.5 21.2 12.5 14.6 12.5 9.8 25.8 23.3 3,757 1,700 871 514 600 514 400 1,057 957 San Luis Valley, Colorado (10% sample of 2,500 sq. mi.) Date 1/29/1976 2/16/1977 1/14/1978 2/08/1979 1980 - - - Adults Juveniles Unknown Eagles per 100 sq. mi. 17 14 10 9 9 3 5 4 2 1 0 0 11.2 7.2 6.0 5.2 Est. of Total Eagles 280 180 150 130 - counts were not conducted this year - - - - - �Table 2. 1972-1980 Bald eagle aerial census of Colorado, South Platte River, Colorado Date (exact count) Juveniles Adults Unknown % Juveniles Total 1/24/1973 18 13 0 31 42% 1/16/1974 16 15 0 31 48% 1/22/1975 28 14 42 0 33% 2/19/1976 10 19 0 29 35% 1977 - - - - The South Platte River was not flown this year - - - - - 1/4/1978 17 8 0 25 68% 1979 - - - - - - - - -Counts were not conducted this year- - - - - - - 1980 - - - - - - - - -Counts were not conducted this year- - - - - - - - Northwestern Colorado Date Adults 1/25 & 2/26/72 1/23/1973 1/22 & 29/1974 1/29/1975 1/26/1976 1/19/1977 1/19/1978 2/13/1979 2/06/1980 i~ - - - - 1 7 1 4 8 3 Prepared - 4 0 0 2 0 was made between San Luis Valley, 1/29/1976 2/16/1977 1/14/1978 2/08/1979 1980 - - - * * *0 ,;': --- Date Juveniles * * No distinction -- (7% sample of 4,100 sq. mi.) - --- Colorado - - Total Eagles per 100 sq. mi. Est. of Total Eagles 35 4 5 1 11 0 4 10 3 12.2 1.4 1.7 0.4 3.8 0.4 1.4 3.5 1.0 500 57 71 14 157 14 57 143 43 on these flights adults and juveniles -- - - - - - - - - - Juveniles Total Eagles per 100 sq. mi. 28 12 18 31 12 6 8 8 40 18 26 39 16.0 7.2 10.4 15.6 -- - - - - - -- (10% sample of 2~500 sq. mi.) Adults - - - -Counts were not conducted by _ _",G"",:--,:, _,_p.:..>....:-. -=~~0J...Cl=-.;:;~ 'r--Gerald R. Crai~( Wildlife Researcher C Est. of Total Eagles this year- - - 400 180 260 390 �145 JOB PROGRESS REPORT State of COLORADO Raptor Investigations Proj ect No. _-"W_-.,::1;.::2;,_:4_-.:;,;R;__ _ Work Plan No: II _..::~-----------------_ Job Title --------------------------- Osprey Nesting Studies Period Covered: Personnel: Job No. 1 March 1, 1979 through December 31, 1980 Gerald Claassen, Gerald Craig, Frank Fiala, Marta McWhorter, Steve Porter, Wayne Russell, John Wagner, and John Ward, Colorado Division of Wildlife. ABSTRACT Although the number of active osprey nests increased in 1979 and 1980, recruitment declined to 0.33 and 0.11 young per active pair in 1979 and 1980 respectively which were the lowest reproductive figures encountered in recent years. Chlorinated hydrocarbon pesticide analysis of 7 nonviable eggs revealed concentrations below those generally considered to inhibit reproduction. These results are in conflict with approximately 22 percent shell thinning observed from egg shell fragments collected below nests. Human disturbance and high winds were factors which may have adversely influenced reproduction. �146 OSPREY NESTING INVESTIGATIONS Gerald R. Craig and Marta McWhorter P.N. OBJECTIVES The objectives of this study are: (1) locate previously unknown nesting ospreys and monitor productivity of all sites, (2) determine cause of reproductive failure and develop methods to restore normal reproduction, (3) determine nesting habitat requirements and implement protective measures to assure continued occupancy, and (4) compile data and prepare annual and final reports. SEGMENT OBJECTIVES lao Locate and map previously Colorado. unknown osprey nesting lb. All known nest sites will be visited reproductive success. 2a. Observe nesting pairs from concealment to determine any behavioral abnormalities, weather conditions, predation, or human disturbance which might impact reproduction. Sites in remote localities will be compared with those adjacent to public activities to determine responses to human visitation. 2b. All unhatched eggs and shell fragments encountered during nest visitations described in 1a. will be collected and submitted for pesticide analysis. Shell fragments will be measured and compared with pre DDT era eggs for possible thinning. 2c. Once cause of reproductive failure has been determined, steps will be taken to alleviate the problem. In the case of human disturbance, closures may be initiated and attempts will be made to relocate some pairs away from areas of high disturbance. Nest alteration activities may be undertaken to make them more secure if predation or weather related failures are established. Should pesticide contamination be the culprit, steps will be taken to maintain or augment reproduction through artificial hatching of eggs. 3. Habitat features such as key hunting areas, topography, vegetative type, climate, and geology will be recorded for each nest site in an attempt to establish habitat requirements. 4. Analyze annually sites throughout to establish data and prepare a report of the findings. �147 METHODS AND MATERIALS Known osprey nesting localities were visited in late April and early May determine the presence of breeding adults. Due to late spring snow storms in 1980, access to sites in Jackson and Larimer counties was limited and nests were not checked until late May. In 1980, a fixed-wing aircraft was used to survey all nests in Grand, Jackson and Larimer counties to determine occupancy, presence of eggs and locate potential nesting areas. In 1979, emphasis was placed on observing only those active sites located in Grand County on an intermittent basis due to man power restrictions. The paucity of nesting details for Larimer and Jackson counties were supplemented by District Wildlife Managers Steve Porter and John Wagner. In 1980, the addition of another observer and two volunteers permitted the Grand County sites to be observed at least bi-weekly throughout the season to document reproductive success, identify disturbance factors and make behavioral observations. All observation points were situated so as to avoid dis.turbance of the site. Such behavioral aspects included nest construction, mating displays, copulation, incubation, hunting, intraspecific and interspecific interactions, re-nest attempts and postreproductive failure activity. Probable cause of nesting failures were recorded. In 1979, nests were entered when incubation was underway and the number of eggs were recorded and measured •. In 1980, the nests were entered only after reproductive failure when incubation was prolonged beyond the normal hatch period. All unhatched eggs as well as shell fragments were collected for pesticide analysis and shell thickness measurement. Whole, unhatched eggs were·submitted to Raltech Scientific Services, Inc for chlorinated insecticide scan in accordance with procedures stanqardized by the U.S. Fish and Wildlife Service Wildlife Research Center at Patuxant, Maryland. Nestling osprey were banded at an appropriate age with U.S. Fish and Wildlife Service bands. Shell thickness measurements were obtained optically using a microscope equipped with a stage micrometer. This permitted the technician to avoid abnormal shell projections or voids and obtain a more accurate measurement of average shell thickness. RESULTS AND DISCUSSION Osprey productivity in Colorado remained almost nonexistant for those sites which were investigated through the 1980 nesting season (Table 1). Twelve sites were occupied of which nine were known to have laid and incubated eggs. Total production consisted of two young at one site in Jackson County which one survived. Cause of the demise of the second juvenile was unknown. Several other osprey sightings were reported during the. breeding season in 1980, suggesting the presence of additional pairs in Colorado. DOW personnel recorded a lone adult female fishing in the Colorado River near Parshall in early May and two adults were observed over Twin Lakes �Table 1. Colorado osprey reproduction~ 1973-1980 1973 1974 1975 1976 1977 1978 1979 1980 Total Total Nests Checked 3 7 12 13 15 9 10 17 86 Occupied Nests!/ 4 7 8 10 12 7 7 12 67 Active Nests.!l 2 5 4 7 5 6 6 9 44 Young Hatches-~/ 0 7 0 3 3 4 2 2 21 Young Fledged~./ 0 7 0 3 3 4 2 1 20 Nests Producing Young Recruitment21 0 3 0 2 2 3 2 1 13 0.00 1.40 0.00 0.43 0.60 0.67 0.33 0.11 0.46 !/ For th.eperiod 1973 - 1978 the number of active nests equal occupied nests since surveys were conducted late in the breeding cycle and those occupied sited which failed were not normally encountered. 2/ Due to late visitation of nests from 1973 to 1978, the nunilierof young actually hatched could not be documented, thus this estimate is identical to the number fledged, ]j No. Young Fledged (or of sufficient age to be presumed fledged) / No. Active Nests. ~ ~ ex> �149 near Leadville in early June. U. S., Forest Service biologists reported three ospreys hunting Haviland Lake, La Plata County in early August after an unsuccessful visit was made to relocate the pair which once occupied a nest at adjacent Electra Lake. Lone adults were also observed interacting with nesting pairs in Grand County. Due to personnel scheduling difficulties from 1975 through 1979, activity and reproduction of some nests in Larimer, Jackson and Grand Counties (Tables 2, 3 and 4) were uncertain and recruitment figures are biased toward the conservative (active sites which failed early in the season were not counted). In 1978 and 1979 few nests were visited in Jackson and Larimer Counties. Of 44 known active nests over the eight year period, 13 produced young. This resulted in an over-all recruitment rate of 0.46, representing levels of 1.17, 0.82, and .04 for Larimer, Jackson and Grand Counties respectively. Referring to the most reliable data obtained in 1974, 1975, 1976 and 1980 for Larimer and Jackson Counties and 1976, 1978, 1979 and 1980 for Grand County, a fluctuating, but declining trend in recruitment is observed. In 1980, recruitment slipped to an all time low of 0.11. Henny and Wight (1969) calculated that ospreys in New Jersey and New York must produce 0.95 to 1.30 young per breeding pair to maintain stable population levels. If such reproductive rates are applicable to western breeding pairs, then osprey reproduction in northern Colorado is insufficient to maintain itself. Several factors may be contributing to low productivity in the threecounty area. The Shadow Mountain - Lake Grandby complex, now delineated as the Arapaho National Recreation Area, receives extremely high visitor use from Memorial Day weekend through Labor Day. The sudden influx of fishermen, boaters and hikers coincides with early to mid-incubation period of nesting ospreys. Hikers and recreational vehicle access to nest areas is also facilitated at this time by the lowered water level of the reservoir. Because the highest concentration of nests occurred in this area, it was the most intensively studied in 1980 to ascertain effects of human pressure on reproduction. Alone, or in combination with other factors, human disturbance may result in ultimately flushing birds from nests. Repeated or prolonged exposure may cause slight temperature fluctuation of eggs, causing arrestment of embryonic development and eventual abandonment by adu l.t s, Two breeding pairs failed to continue incubating after prolonged human activity occurred within 100' of nests over Memorial Day weekend. In one case the female was kept from feeding or exchanging with male for 8 hours. Both adults grew exceedingly agitated by continual presence of nine fishermen, and trolling and speeding boats within 50-100' of nest. In the second instance, both adults were flushed from the nest on the two consecutive days by campers within 50~ of the site. They were kept from resuming incubation activity for a minimum of 10 and 77 minutes on the two occasions observed. Fishermen were observed within 400' of a third nest on Memorial Day after desertion by adults had already occurred. Since this was an area of easy access, it is indicative that fishermen may have frequented it earlier in the weekend. The following weekend a �150 Table 2. Larimer County osprey reproduction, 1973-1980 Site No LA-I LA-2 Status 1973 1974 1975 1976 1977 1978 1979 1980 OC AN ND UO AN UO OC TD 0 ND AN AN Production ?/?/O Status (-- ?/?/2 ND Not discovered ?/?/2 --) Production AN ND AN ?/?/2 ?/?/1 ?/O/O Total Active Sites 0 1 ND 0 2 1 1 1 Total Young Produced 0 2 ND 0 2 2 1 0 Recruitment }j 0.00 2.00 ND 0.00 1.00 2.00 1.00 0.00 Status Codes OC occupied breeding territory (not necessarily productive) UO unoccupied breeding territory AN active nest (known to have produced eggs) ND no data available TD tree or nest blown down Production Codes 3/2/1 = 3 eggs, 2 young hatched, 1 young fledged ? = unknown 1/ Young produced/active nest �151 Table 3. Jackson osprey reproduction. 1973-1980 Site No 1973 1974 1975 1976 1977 ND ND AN ?/3/3 OC ?/?/O AN ?/?/2 AN AN ?/?/1+ ?/?/1+ AN ?/?/2 OC ?/?/O JA-1 Status Production JA-2 Status Production AN ?/?/O JA-3 Status Production OC ?/?/O JA-4 Status Production JA-5 Status Production (Not discovered) ND ND UO JA-6 Status Production (Not discovered) OC OC JA-7 Status Production (---- UO TD AN OC ?/?/O 2+/0/0 Not discovered ----) 1978 1979 1980 UO IA (--- Site abandoned --- ) ND ND ND ND OC 0/0/0 AN ?/?/O ND ND OC ND A.."N 0/2/1 AN ?/?/O TD (Site abandoned) ND ND UO ND ND -Not discovered- UO UO --) AN ?/?/O Total Active Sites 1 2 0 2 3 1 a 2 Total Young Produced 0 5 a 2 1 0 0 1 Recrui tmen 0../ 0.00 2.50 0.00 1.00 0.33 0.00 0.00 0.50 Status Codes OC UO AN ND TD = occupied breeding territory (not necessarily productive) unoccupied breeding territory active nest (known to have produced eggs) no data available tree blown down Production Codes 3/2/1 = 3 eggs, 2 young hatched, 1 young fledged ? = unknown 1/ Young produced / active nest �152 Table 4. Grand County osprey reproduction 1973-1980. Site No. GR-1 Status Production GR-2 Status Production GR-3 Status Production GR-4_Y Status Production GR-5 Status Production GR-6 Status Production GR-7]) Status Production cs-al/ Status Production GR-9 Status Production GR-lOl/ Status Production GR-ll Status Production GR-12 Status Production GR-13 Status Production GR-14 Status Production GR-15 Status Production 1973 1974 1975 OC VO ?/O/O AN AN 2/0/0 1/0/0 AN AN 2/0/0 ?/O/O installed AN ?/O/O (Not discovered) AN 0 (Not discovered) OC ?/O/O installed UO ?/O/O installed VO Total Active Sites Total Young Produced RecruitmentJ:j ND ND ND ND OC 0 1977 1978 1979 VO AN ?/?/1 OC ?/O/O AN ?/O/O AN ?/O/O VO OC ?/O/O OC ?/?/O AN +/0/0 OC ?/O/O OC ?/O/O TD AN 3/0/0 AN 3/1/1 VO AN AN 3/0/0 2+/0/0 AN AN 3/0/0 2+/0/0 AN TD ?/O/O AN ND ND 3/0/0 (Abandoned) OC ?/O/O VO OC ?/O/O VO AN 3/0/0 VO OC installed ( DO Not discovered ) AN 1/0/0 ND OC ?/?/O TD ND ND TD (-Site abandoned-) ND (---------------Not discovered----------------) (---------------Not discovered----------------) (---------------Not discovered----------------) 1 0 0.,00 4 0 0.00 4 0 0.00 1 0 0.00 4 1 0.00 Status Codes OC DO AN ND TD AN 3/0/0 VO occupied breeding territory (not necessarily productive) unoccupied breeding territory active nest (known to have produced eggs) no data available tree or nest blown down Production Codes 3/2/1 = 3 eggs, 2 young hatched, 1 young fledged. + = at least 1 egg produced, possibly more. ? = unknown i/ Artificial nest 1/ Young produced / active nest AN 3/0/0 DO DO ND ND ND ND AN DO DO 3/0/0 (Site abandoned) (----------- Not discovered---------)DO 0 0 0.00 1980 1976 4 0 0.00 AN 2+/0/0 OC ?/O/O OC 0/0/0 AN 2+/0/0 6 0 0.00 �153 fourth nest appeared to fail under observation, but the area beneath the nest was obscurred from view. From the defensive attitude of adults at the nest and their behavior following desertion, it is suspected that hikers may have been in close proximity for at least 30 minutes. At the two remaining nests in the Shadow Mountain - Lake Granby complex, hikers, fishermen and trail rides within 20-100' flushed incubating birds on several occasions through June for periods of up to 8 minutes. In both cases eggs received prolonged incubation and were found to have addled at some point in embryonic development. Nests in North Park are situated by beaver ponds and are relatively isolated compared to those in Grand County. Two unproductive sites received occasional to moderate fishing pressure acknowledged by District Wildlife Managers and observation of recent footprints within 100' of nests. Ospreys in these more remote areas may be accustomed to very little human disturbance. They exhibited far less tolerance to observers and flushed more readily at longer distances than ospreys in the Shadow Mountain - Lake Granby area. Beaver Creek in Jackson County contains one active and two historic nests and features difficult access to adjacent beaver ponds. The pair breeding here has consistently showed highest production level in the three-county area over the past 8 years and was the only success story of 1980. Swenson (1979) suggested that the abrupt advent of heavy human activity in mid-June was a major factor in lower reproduction on Yellowstone Lake than on adjacent streams receiving continual, but light use through the nesting season. This implication may have application in similar situations existing in northern Colorado. High velocity wind storms appear to have severely influenced productivity in the past, especially after incubation was well in progress. Natural nests in snags have experienced the most destruction. In the nesting seasons of 1978 and 1979 wind blew down one nest tree and destroyed two nests. It is uncertain whether these had reproductively failed beforehand. Four other nest trees have been blown down off-season in the past 5 years. Nests on artificial platforms in live trees have withstood wind pressures to a greater extent. In 1980 incubating birds exhibited a close tenacity to nests during high wind periods and heavy hail ~nd snow storms. This behavior suggests ospreys are tolerant to harsh weather situations under normal conditions, possibly until severity causes actual demise of the nest. This year no nests were destroyed, but the highest velocity wind storm of the season occurred on Memorial Day weekend. In this instance high winds may have served to further antagonize already nervous birds and precipitated abandonment. In combination with human activity, winds may have expedited alterations of egg temperatures, and/or blown eggs from nests during flushing periods. District Wildlife Managers reported strong winds in North Park through the nesting season. One nest was partially blown apart, but after hatching should normally have occurred. Predation does not appear to be a primary factor in reproductive failure, but may have occurred after the fact at failed nests. All eggshell �154 fragments found contained little or no embryonic fluid. Two eggs left after partial clutches were taken had been cracked open with fluids partially removed. Ravens and gulls were noted in close proximity to nests, and ravens repeatedly harassed incubating, perched and flying ospreys. At one nest where incubation was extended, intra-specific aggression was observed several times during the latter 2 weeks of activity. At this point the nest had technically failed, but the pair was incubating as normal. From one to three intruding adults, presumably from failed nests in the area, flew close to, or hovered over the active nest. In one instance an osprey perched on the nest three times with male and female residents present. On all occasions of interference the active pair behaved nervously and attempted to drive off intruders. Pesticide contamination of adult breeding birds was also considered as a cause for reproductive failure. Although no eggs were collected in 1979, four nonviable eggs were taken from nests in Grand County in 1980 and were submitted for chemical analysis along with three eggs obtained in 1978. The DDE residue levels of the seven nonviable eggs averaged 4.22 ppm (range 1.99-6.89 ppm) DDE fresh weight (Table 5). This is be Low the concentrations encountered with declining populations in Connecticut which averaged 9.9 ppm and 8.9 ppm (~Jeimeyer et al. 1975). Although data are meager for the western United States, small samples of eggs from Wyoming (4 eggs) and Idaho (11 eggs) averaged 7.2 ppm and 8.5 ppm respectively (Swenson 1975; Johnson et al. 1975). Despite the elevated DDE levels, both populations were considered stable or increasing which was contrary to the trend experienced along the eastern seaboard. In any case, it would appear that the Colorado samples fall in the range of low to moderate DDE levels. Shell thickness measurements from 13 eggs collected in 1975, 77, 78 and 80 (Table 6) averaged 0.3955 rom (with membrane) which is 22 percent thinner than eggs collected prior to 1947 from the eastern U.S. (Anderson and Hickey 1972). Spitzer et al. (1972) observed that shell thickness indexes of osprey eggs were identical throughout the continent which rules out geographical variation in shell thickness and permits direct comparison of the Colorado eggs with results obtained elsewhere. Below normal production was observed in'populations averaging 15-18 percent thinning (Weimeyer et al. 1975) and thinning above 20 percent is generally considered severe. It is perplexing that extreme shell thinning was observed without associated high pesticide ~vels. One bias is that the shell fragments which were measured were encountered below nests indicating they may have broken during incubation. The intact, nonviable eggs encountered after incubation and subsequently removed for analysis would be expected to possess thicker shells in order to remain intact. Thus, pesticide contaminant levels should be lower in the intact eggs. At the time of this writing, the shells have not been returned from the pesticide laboratory so thickness measurements cannot be compared to pesticide levels or shell fragments in possession. �Table 5. Organochlorine insecticide residue analysis of osprey eggs collected in 1978 and 1980. Site No. Sample No. Year Collected Percent Moisture DDT Residues 1 (ppm corrected to fresh-Iwet weight) DDD DDE PCB Dieldrin GR-2 78-3 1978 78.8 0.16 0.67 6.89 2.88 ND.Y ND 10.60 GR-4 80-2 1980 80.3 ND 0.11 1.99 1.09 0.03 ND 3.22 GR-4 80-3 1980 79.8 ND 0.12 1.95 1.12 0.04 ND 3.23 GR-4 80-5 1980 83.1 ND 0.14 2.79 1.55 0.04 ND 4.56 GR-6 78-1 1978 78.7 ND 0.15 3.64 3.56 ND ND 7.35 GR-6 78-2 1978 80.1 ND 0.17 6.45 5.41 ND ND 12.03 GR-7 80-1 1980 78.0 ND 0.88 5.81 1.12 0.04 0.06 Mean Value 0.32 4.22 2.39 0.04 Range 0.11-0.88 1.99-6 .89 1.09-5.41 0.03-0.04 11 assumed wet weight of fresh egg at 85% 1/ none detected Endrin Total Residues 7.91 6.99 3.22-12.03 •.... VI VI �156 Table 6. Shell thickness Site No of Colorado Year Collected osprey eggs Average Shell Thickness (mm) Hith Membrane "I/O Membrane GR-2 1975 0.3874 0.2889 GR-3 1977 0~41373../ 0.3152 GR-1 1978 0.3940'1:} 0.2955 GR-1, l!/ 1980 0.3874 0.2889 GR-1,2 1980 0.3809 0.2824 GR-1,3 1980 0.3743 0.2758 GR-2,1 1980 0.3874'l:_/ 0.2889 GR-2,2 1980 0.3809 0.2824 GR-3,1 1980 0.4073'l:_/ 0.3088 GR-3,2 1980 0.440oY 0.3415 GR-3,3 1980 o . 4400'1:} 0.3415 GR-15,1 1980 0.39403../ 0.2955 GR-15,2 1980 0.3546 0.2561 0.3955 + .0146l! 0.2970 Average Thickness 1/ Last digit indicates 2/ Shell membrane measurements could not be taken either because the shell was not clean or it was separated from the membrane. In these cases, a membrane thickness of .0985 mm was assumed since this was the value invariably obtained when measurements were taken. 1/ Confidence egg number, not necessarily limit of 95% in order laid. �157 LITERATURE CITED Anderson, D.W. and J.J. Hickey. 1972. Eggshell changes in certain North American birds. Proc. XVth Int. Orithol. Congress 1972. pp. 514-540. Henny, C.J. and H.M. Wight. 1969. An endangered osprey population: estimates of mortality and production. Auk 86: 188-198. Johnson, D.R., W.E. Melquist, and G.J. Schroeder. 1975. DDT and PCB levels in Lake Coeur d' Alene, Idaho; osprey eggs. Bull, Environ. Contam. Toxicol. 13(4): 401-405. Spitzer, P.R. R.W. Risebrough, J.W. Grier and C.R. Sindelar, Jr. 1977. Egg shell thickness - pollutant relationships among North American ospreys. in Trans. of North Am. Osprey Res. Conf., College of William and Mary. Williamsburg VA, 10-12 Feb. 1972. J.C. Ogden (ed.) Proceed and Trans. Nattl Park Servo No.2: 13-19. Swenson, J.E. 1975. Ecology of the bald eagle and osprey in Yellowstone National Park. M.S. Thesis, Montana State Univ., Bozeman 146pp. Swenson, J.E. 1979. Factors affecting status and reproduction of ospreys in Yellowstone National Park. J. ~.;rild1. Manage. 43 (3): 595-601. Wiemeyer, S.N. Spitzer, P.R., Krantz, W.C., Lamont, T.G., and Cromartie, E. 1975. Effects of environmental pollutants on Connecticut and Maryland ospreys. J. Wildl. Manage. 39(1): 124-139. Prepared by _G=..,.:....,.. -"-Q-=-.___:::~=:-=:.::...'__,.. Gerald R. Craig \ Wildlife Researcher C _ �158 JOB PROGRESS REPORT State of COLORADO Project No. ~W_-~1~2~4_-R=Work Plan No. Raptor Investigations III Job ~~~---------------- Job Title: No: I Prairie Falcon Nesting Studies Period Covered: Personnel: _ March 1, 1979 through December 31, 1980 Gerald Craig, James Enote, Brandon Grebence, James McKinley, and Richard Williams, Colorado Division of Wildlife. Abstract Prairie falcon nest occupancy and reproductive data were summarized for the 1979 and 1980 breeding seasons. The lower fledging success in 1980 may, in part, be due to unseasonally late spring snow storms which occurred at egg laying. �159 PRAIRIE FALCON NESTING STUDIES Gerald R. Craig and Brandon Grebence P.N. OBJECTIVES 1. Document the breeding range and estimate the number of breeding pairs of prairie falcons in Colorado. 2. Obtain production data at selected nesting areas throughout the state and estimate total production of prairie falcons. 3. Delineate nesting habitat requirements of prairie falcons. 4. Document movements and mortality of prairie falcon throughout Colorado. SEGMENT OBJECTIVES 1. Locate and map all known nesting sites of prairie falcons throughout Colorado. From data obtained through approach 3, extrapolate the number of breeding pairs potentially occupying similar habitat types throughout the state. 2. Establish study areas in select habitats which represent important nesting areas. Study areas will be selected that represent shortgrass prairie, foothills and mountain nesting populations and productivity will be determined and compared between areas. 3. Physical and biological parameters of each nest site which is visited will be recorded on appropriate field forms and analyzed to establish those features which favor occupancy by prairie falcons. The field information will be gathered in conjunction with. approach 1. 4. When nests are visited to determine productivity, the young will be banded with Fish and Wildlife Service lock-on bands and their movements will subsequently be traced through reports filed with the Office of Migratory Bi~d Management. Should the occasion permit, transmitters will be placed on several breeding adults to determine extent of hunting ranges. Radio transmitters will also be placed upon young at fledging and their movements and activities will be monitored. 5. Compile data and prepare annual and final reports. METHODS AND MATERIALS Known nest sites were visited during courtship and early incubation to determine the presence of breeding adults. Invariably all sites could not be visited early in the season and some only received visitation once �160 after the young were produced. Of those sites which were checked, a sample of active nests was selected and an attempt was made to return to each of the sample sites and determine reproductive success. Young were banded with U.S. Fish and Wildlife Service bands when nests were visited to determine brood size, locate prey remains, and record other information about the nest. In addition, the following information was recorded on field forms: elevation, topography, geology of the nest cliff, major vegetative communities, potential hunting areas, distance and direction to disturbance factors, location of nest site, height from nest to top and bottom of the cliff, dimension of the nest ledge, and presence of other raptors in the vicinity. RESULTS AND DISCUSSION For purposes of summarizing the data, the state was divided into 6 geographic areas. Area 1 is the northwest, Area 2 is the northeast, Areas 3 and 4 represent the east-central, Area 5 is the southeast, and Area 6 represents southwestern Colorado. Tables 1 and 2 present site occupancy and productivity for the 1979 and 1980 breeding seasons. The primary effort was directed toward obtaining physical and vegetative parameters of known nest sites. Table 1. Prairie falcon reproductive success in Colorado, 1979 No. Sites Checked No. Occupied Sites % of Sites Occupied Total No. of Pairs No. of Successful Pairs Total Young Produced Young Produced per Successful Pair Young Produced per Total Pair Young Produced per Occupied Site Total Young Fledged Young Fledged per Successful Pair Young Fledged per Total Pair Young Fledged per Occupied Site Fledgling Male to Female Ratio Area 1 Area 2 Area 3 Area 4 Area 5 Area 6 Total 16 8 50.0 8 3 17 5.67 50 42 84.0 41 17 77 4.53 32 18 56.3 13 8 34 4.25 16 5 32.0 4 3 7 2.33 27 10 37.0 6 3 9 3.0 21 11 52.4 5 3 10 3.33 162 94 58.0 77 37 154 4.16 2.13 1.88 2.61 1.75 1.50 2.00 2.00 2.13 1.83 1.89 1.40 0.90 0.91 1.64 15 5.00 61 3.59 30 3.75 7 2.33 9 3.00 5 1.67 127 3.43 1.88 1.49 2.31 1.75 1.50 1.00 1.65 1.88 1.45 1.67 1.40 0.9 0.45 1.35 1.67 5.0 2.5 0.67 0.88 �161 Table 2. Prairie falcon reproductive success in Colorado, 1980 No Sites Checked No. Occupied Sites % of Sites Occupied Total No. of Pairs No. of Successful Pairs Total Young Produced Young Produced per Successful Pair Young Produced per Total Pair Young Produced per Occupied Site Total Young Fledged Young Fledged per Successful Pair Young Fledged per Total Pair Young Fledged per Occupied Site Fledgling Male to Female Ratio 1/ Area 1 Area '})j Area 3 Area 4 Area 5 Area 6 Total 22 15 68.2 13 6 24 4.00 28 9 32.1 8 8 22 2.75 13 5 38.5 2 1 4 4.00 26 6 23.1 5 1 6 6.00 16 6 37.5 4 2 7 3.50 105 41 39.0 32 18 63 3.50 1.85 2.75 2.00 1.20 1.75 1.97 1.60 2.44 0.80 1.00 1.17 1.62 24 4.00 22 2.75 4 4.00 3 3.00 7 3.50 60 3.33 1.85 2.44 2.00 0.60 1.75 1.87 1.60 2.44 0.80 0.50 1.17 1.54 0.44 0.80 3.00 0.50 1.33 0.74 Data not yet available from National Audubon Society. Prepared by ~~~.~R~. __~~~~ _ Gerald R. Craig Wildlife Researcher C � https://cpw.cvlcollections.org/files/original/c7907bc9375948477dcfc8e68d2e8527.pdf f7519c21d5186a50a753eddce1d48083 PDF Text Text 1 April JOB PROGRESS State of Project REPORT Colorado ---------------------No. Game Bird Survey W-37-R-34 ------ Work Plan No. 1 ---------------- Job Title: Evaluation in Relation Period 1981 Covered: Personnel: Job No. of Nesting ------------------ 22 Cover Preferences of Pheasants to Wheat Farming Methods 1 April 1980 through 31 March Kirk Snyder, David Palmer, Martin Colorado Division of Wildlife. 1981 Staab, and Warren Snyder, ABSTRACT Approximately 2 feet of snow, which fell in late March 1980 and which persisted for over a week in early April delayed the onset of hen ringnecked pheasant (Phasianus colchiaus) spring dispersal and nesting compared to 1979. This delay, accompanied by sparse, short wheat stubble and excellent early growth of green wheat in late April and May, resulted in most known first nests being placed in green wheat.. Consequently, spring stubble plowing which was later than usual .(occurring primarily during the. 2nd half of May) had little negative impact on 1980 pheasant production. Thirteen of 18 known first nests placed in green wheat in Mayor early June were successful yielding large broods. This excellent early hatch was primarily responsible for the increased fall population. Most hens, whose initial nests were destroyed by predators abandoned renesting efforts in early July because wheat harvest crushed nests or buried them under straw. Extremely hot, dry weather, from late June through July, also affected renesting activities. However, 2 hens successfully renested and completed known 3rd nesting attempts with their clutches hatching in late August. Hens with broods made pronounced movements toward green irrigated row crops or tall annual weed cover soon after hatching. Data are presented concerning cover type availability, use of cover types by hen pheasants, clutch size, nest placement, grasshopper densities, and other environmental comparisons with preceding years. The chemical fallow treatment of wheat stubble in August 1979 was not effective in suppressing weeds in 1980. Failure of hens to nest in wheat stubble made treatment failure irrelevant since evaluations could not be made. Mild, dry fall-winter conditions permitted excellent pheasant survival. Percent harvest of the fall population of male pheasants was low. Forty-two hens were trapped, using nightlighting techniques in March 1981, equipped with solar powered transmitters, and released for subsequent monitoring. ��3 EVALUATION OF NESTING COVER PREFERENCES OF PHEASANTS IN RELATION TO WHEAT FARMING METHODS Warren D. Snyder Ring-necked pheasants are important game birds in Colorado and, in most years, are the most heavily sought after resident game bird. While pheasants occur in suitable habitat throughout Colorado, this species is most widespread in eastern Colorado. Densities over large areas are highest in the wheatlands of northeastern Colorado, an area that is intensively farmed for wheat production. More recently, use of center-pivot irrigation systems has resulted in increased acreage being planted to corn and pinto beans. With changing cropping systems and higher costs for fuel, land and machinery, farmers have indicated interest in chemical treatments and minimum tillage systems that could reduce their operating costs. This report covers the 2nd year of a 3 year study designed to examine pheasant nesting preferences and the potential of chemical treatment and minimum tillage systems to impact pheasant nesting. P. N. OBJECTIVES 1. To document the relative importance of wheat stubble, green wheat, and other vegetative cover for (1) pheasant nest site selection, and (2) successful production of young in the wheatlands of northeast Colorado. Other variables pertinent to understanding the basic ecology of the nesting pheasant in the Tablelands include determination of primary limiting factors to reproduction and brood survival. 2. Upon documentation that pheasants use wheat stubble extensively for initial spring nesting and that adequate sample sizes can be obtained in the chemical fallow treated fields and their controls, a second 'primary objective will be to determine if the chemical fallowmini-till farming method will increase pheasant nesting success when compared to conventional summer fallow methods. SEGMENT OBJECTIVES 1. Monitor a. b. c. d. e. pheasant production activities in relation to the following: Nesting cover selection for first, second and subsequent attempts. Determine clutch size, nesting fate, use of vegetation and litter and recycle time to subsequent renest when applicable. Relate nesting activities to farming methods and activities including time of stubble tillage and wheat harvest. Record predation and predator influence on nesting and brood activities. Monitor brood rearing activities in relation to cover types and weather. �4 2. Monitor environmental pheasants including: a. b. c. d. e. f. g. h. i. variables which directly and indirectly impact Precipitation amount and pattern and obtain temperature data. Compare soil moisture conditions when appropriate. Record period and peak of wheat stubble tillage. Obtain height-density measurements of residual cover. Obtain height-density measurements of green wheat. Sample grasshopper densities in selected covers. Sample the green vegetation in chemical fallow and untreated stubbles. Compare the amount of straw residue on chemical fallowed-minitilled fields with that on conventionally farmed fields. Record dates of various field activities. 3. Trap and equip a m1n1mum of 30 hens with transmitters monitoring of hens in late winter, 1981. 4. Analyze second year findings. 5. Prepare an annual progress and initiate report. METHODS AND MATERIALS Reference materials is made to Snyder (1979, 1980) for a summary of methods used in this study. and RESULTS AND DISCUSSION Environmental Precipitation Measurements and Temperature Sixteen inches of precipitation were recorded in the Sand Draw vicinity in 1980, slightly below the long-term average and below the amount received in 1979 (Table 1). Significant precipitation was not received from mid-August 1980 through 28 January 1981, and top soils were extremely dry through the fall - early winter period. The most significant weather in 1980 was a snowstorm on 27-29 March which yielded over 3 inches of precipitation at Holyoke and a similar amount at the Sand Draw. Snow accumulated to depths approximating 2 feet on the average and considerable drifting occurred, covering most herbaceous food and cover. Winds accompanying the snow were moderate rather than gale force and little direct pheasant mortality was noted. Pheasants abandoned plum (Prunus spp.) thickets in search of food and were exposed and vulnerable to avian predation with several hens being taken. �5 Table 1. 1978-80.a Precipitation (inches) recorded in the Sand Draw vicinity in Month 1978 Year 1979 Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec 0.45 0.43 0.07 0.74 2.78 1.37 1.63 1.25 0.11 0.50 0.43 0.75 0.64 0.09 2.14 1.45 2.44 3.81 2.32 1.77 0.65 1.00 1.60 Tr 0.90 0.73 3.20 1.80 2.79 2.12 2.72 1.53 Tr 0.10 Tr 0.31 0.29 0.86 1.53 3.29 3.83 2.82 1.88 1.64 1.13 0.46 0.36 10.51 17.91 16.00 18.40 Totals 1980 Long-term o.ri b aprecipitation for winter months was obtained Commerce records in nearby towns. b Long-term mean precipitation for Holyoke, mean from u.s. Dep. Colorado. Snow remained on the ground through approximately the first 10 days of April 1980 with accompanying colder than average temperatures. Daytime maximums averaged 47.9 F for the first 2 weeks of April in contrast to an average of 72.3 F for the remainder of the month (U.S. Dep. Commerce 1980). Rapid temperature increases in late April, accompanied by abundant soil moisture, stimulated rapid growth of previously well established green wheat. Hot, relatively dry conditions began in the area on 23 June 1980 and persisted through the remainder of June, nearly all of July, and part of August. Rapid ripening of wheat followed by rapid wheat harvest resulted. A localized thunderstorm, containing considerable small hail, occurred on 10 May. It reduced the height, density, and yield of several wheat fields immediately south and southeast of the Sand Draw property. This thunderstorm occurred too early in the spring to have had a major impact on pheasant nesting efforts or broods. �6 Vegetation Height and Density Residual cover. -- Height and height-density (H-D) measurements (Kirsch, unpubl. rep., U.S. Fish and Wildl. Serv., Jamestown, N.D. 1977) showed that wheat stubble and other residual herbaceous vegetation provided poor early spring nesting and protective cover (Table 2). The stubble remaining from the poor 1979 wheat crop was short and open, and was partially lodged after the heavy, wet snow in March. Fields on the Sand Draw property, which contained little wheat stubble and dense cheatgrass (Bromus tectorum) offerred no significant cover after the snow (Table 2). The late March snow lodged residual annual and perennial cover on the property and in roadsides throughout the region. However, most of this cover was of marginal value for nesting pheasants prior to the snowstorm. Table 2. Comparative height density ments of wheat stubble, green wheat, within or near the San Draw Property Date measured H-D a index Height Mid-Apr Mid-Apr Mid-Apr Late Janc near 0 0.27 0.33 0.65 7.43 11.53 10.42 17.24 Sand Draw Sand Draw Sand Draw Late Jan Late Jan 20 May 0.28 1.58 0.98 7.22 32.50 9.40 Sand Draw 20 May 1.41 10.35 Sand Draw Sand Draw Eight fields on and near Sand Draw property 2 Jun 2 Jun 22 Apr 1 May 10 May 20 Mayd 2 Jund 2.04 0.91 near 0 1.12 2.73 3.60 7.48 12.58 11.84 5.00 10.19 16.79 19.21 33.72 Location Cover type Wheat stubble Alfalfa/cheatgrass (burned in 1979) Unburned alfalfa/ cheatgrass Weed patch (Kochia) Alfalfa/cheatgrass (burned spring, 1980) Unburned alfalfa/ cheatgrass 1-yr-old alfalfasweet clover Mixed perennial grass Green wheat Sand Draw Two adjacent Other nearby Sand Draw aThe H-D index is expressed are presented in inches. b Two stubble roosting. cResidual late March. d Excludes mid-May. (H-D) indices and height measureand other vegetation obtained in 1980. fieldsb fields in decimeters and the height measurements fields heavily used for winter-early cover was subsequently measurements obtained severely spring feeding and lodged by heavy snow in on wheat fields damaged by hail in �7 New growth. -- Green wheat used subsoil moisture accumulated in 1979, along with moisture from the March snow and had excellent growth starting in late April 1980 (Table 2). Height density (H-D) indices obtained in 1980 contrasted markedly to those obtained in 1979 (Fig. 1). The growth in 1980 compared favorably with growth of wheat in the 1960's (Snyder 1980). Fields on and south of the Sand Draw Property that were treated with herbicides in summer 1978, showed no residual herbicide effects. They yielded the tallest and most dense wheat among the 8 fields measured. Cheatgrass was not a problem in Sand Draw Property fields in 1980, as it was in 1979, in part due to the excellent stands of wheat established in September 1979, and possibly because of deep plowing at the onset of the previous summer. Limited H-D and height samples obtained in stands of old alfalfa, a 1979 seeded alfalfa-sweet clover stand, and perennial native grass, indicated that none of these was comparable to the H-D and height samples obtained in green wheat in late May and early June (Table 2). Acreages of these cover types were small and no nesting use by radiomarked hens was documented. H-D and height comparisons. -- A close correlation (~ = 0.975) between height and the height-density (H-D) method (Robel et al. 1970, Kirsch 1977) was obtained in green wheat during 1979 and 1980 (Fig. 2). The H-D index is considered a better indicator of the concealment value of cover for ground nesting birds than height alone. Vegetation Types Approximate acres and percentages of vegetative types within the 9-section area, including and surrounding the Sand Draw Property during early spring 1980 varied (Table 3). There were few differences in acreage of cover types available between March 1980 and March 1979 (Snyder 1980). Corn stubble was not present in March 1980 and there were several hundred more acres of millet stubble. The latter, however, is of little value to wintering, feeding, or nesting pheasants. A decrease in shortgrass and mixed-grass prairie occurred on the Sand Draw Property within Section 19 where reseeding efforts resulted in forb dominated grass-alfalfa stands. Part of this acreage was converted to annual forbs, primarily sunflowers (Helianthus spp.), by disturbance tillage. A narrow tree row was partially removed along the west edge of the southwest 1/4, Section 18 in late winter 1980. All trees were also removed in the southwest corner of Section 16 just outside the 9-section extensive study area. Cover types present in late July and August 1980 varied (Table 4). The acreage of wheat stubble increased whereas millet and millet stubble decreased over the preceding year. The wheat stubble was generally of better height and density than in 1979. Irrigated sugar beets replaced beans in the field immediately north of Sand Draw (Fig. 3). Both row crops, while offering late summer brood cover, result in bare ground over winter. �8 8 I I -../ fgl 7 It fJl 6 #/ ~ ~ ~ ~ e' ~ I ! ~7 5 >t f'j ~ {Il z 4 ~ I ) ~ ~ .!2 / / fal :c f! / 3 L. 2 ~ 1 ~ ~ / ~ .".,. •••• ~ 0 30 20 APR _20 10 MAY 30 JUN Fig. 1. Comparison of 1979 and 1980 wheat growth as indicated by heightdensity measurements in fields on and near the Sand Draw Property. �9 45 40 35 30 U) ~ 25 ~ 20 ~ ~ 15 10 r •.0.975 5 Y = o 1 2 3 4 -1.69 5 HEIGHT-DENS ITY INDEX + 0.247 ~ 678 (DECIMETERS> Fig. 2. Correlation of height and the height-density (H-D) index for 1979 and 1980 wheat growth measurements. 9 �Table 3. Acres per section of cover types present on the 9-section extensive study area in March 1980. Cover type 13 18 17 24 Section 19 20 25 30 29 Totals % of total Wheat stubble 194.6 103.5 62.0 358.8 172.6 59.6 100.1 270.5 368.9 1,690.6 29.4 Green wheat 270.3 200.2 566.4 238.5 291.5 269.3 363.0 313.6 167.1 2,679.9 46.6 Millet stubble 179.4 10.0 38.5 175.1 37.3 95.0 535.3 9.3 139.9 2.4 386.1 6.7 5,431.8 94.4 Sorghum stubble 130.0 Plowed ground 131.2 Total cropland 644.3 Shortgrass or mixed grass 564.9 638.4 464.6 9.4 130.0 124.9 634.0 625.3 593.5 7.0 26.8 171.7 3.0 5.3 5.2 14.5 0.2 75.0 4.0 Forbs and grass 0.8 Trees 3.0 4.0 0.4 63.5 3.0 2.0 0.8 1.6 0.5 1.8 15.8 1.3 2.0 15.7 640.8 636.2 1.0 Shrubs Total acreage 635.8 62.9 Occupied farmyard Roadsides 0.5 645.1 635.8 644.4 638.8 631.0 4.4 3.7 79.0 1.4 1.2 2.3 3.0 15.2 0.3 0.2 2.5 22.6 0.4 17.7 0.3 639.0 634.7 637.7 5,752.5 ,_. 0 �Table 4. Acres per section of cover types present on the 9-section extensive study area in late July and August 1980. Cover type 13 18 17 24 Section 19 20 25 30 29 Totals % of total Summer fallow 342.0 233.5 72.0 397.1 176.6 234.7 121.1 278.0 463.9 2,318.9 40.3 Wheat stubble 270.3 141.4 566.4 238.7 177 .4 269.3 346.0 315.5 167.1 2,492.1 43.3 168.9 2.9 65.6 1.1 2.0 0.1 254.6 4.4 131.2 2.3 5,433.3 94.4 Mulched stubble Millet 58.8 110.1 33.6 32.0 Sorghums 2.0 Corn, irrigated 130.0 Sugar beets, irrigated Total cropland 131.2 644.3 Shortgrass and mixed grass 564.9 638.4 466.1 634.0 75.0 4.0 Forbs and grass 0.8 Trees 3.0 4.0 0.4 62.0 3.0 2.0 0.8 1.6 0.5 1.8 15.8 1.3 2.0 15.7 640.8 636.2 1.0 Shrubs Total acreage 635.8 62.9 Occupied farmyard Roadsides 124.6 645.1 635.8 644.4 638.8 625.3 593.5 631.0 7.0 26.8 171.7 3.0 5.3 5.2 14.5 0.2 4.4 3.7 77.5 1.4 1.2 2.3 3.0 15.2 0.3 0.2 2.5 22.6 0.4 17.7 0.3 639.0 634.7 637.7 5,752.5 tt- �12 2 1 5 4 91b .__.....-----~-~ l1S. / ,,/ S 9 10 15 22 26 30 29 27 3S-Pc, .:: CE 35 36 31 2 1 6 ~IVOT 33 1\~.:;:GA'rED FIT 34 5 Fig. 3. Location of unsuccessful pheasant nests (0), successful nests (e) and movements to brood areas ~) in the Sand Draw vicinity during summer 1980. D �13 Roadsides within the extensive study area and within the region were nearly all farmed to the shoulder. Those remaining unfarmed were dominated by cheatgrass and were of little value to nesting pheasants. One farmer chemically sterilized roadsides eliminating all cover along the east side of the northeast 1/4, Section 20. This practice was 'repeat ed along the edges of several other sections in the vicinity. The Sedgwick County road maintenance crew sprayed roadsides between sections 20-29, 19-30 and 24-25 (Fig. 3) with atrazine in summer 1980. The impact of this treatment on 1981 roadside cover remains uncertain although no great impact is anticipated. Progress of Spring Stubble Plowing Initial spring tillage of wheat stubble was later than average in 1979, but in 1980 was the latest since initiation of data collection in 1963 (Fig. 4). Part of the delay was due to wet field conditions, but priorities in preparing and planting irrigated crops, and rising fuel costs also may have affected start of spring tillage. Stubble tillage within the 9-section study area progressed at about the same rate as tillage along the more extensive transect route (Snyder 1980). Wheat stubble fields contained poor cover in spring 1980 and little nest placement occurred there, consequently few nests were destroyed by spring tillage. Wheat Harvest Rapid wheat growth (Fig. 1) in combination with warm weather in late June ripened the 1980 wheat crop so that harvest began on 5 July in the study region. Harvest progressed rapidly with peak activity occurring from 8-15 July. Harvest was approximately one week or more ahead of that in 1979. Yields were good to excellent. Grasshopper Density A slight reduction in grasshopper density occurred in 1980 compared to 1979 (Table 5). Annual forbs and perennial grasses continued to contain high densities (Table 6) and considerable damage was inflicted to sorghum food-cover plantings on the Sand Draw property. Table 5. type. Vegetation Comparison of 1979 and 1980 grasshopper type Sunflowers, weed strip Roadside, cheatgrass and forbs Perennial mixed grasses Uncut ripe wheat densities Average 1979 35.5 39.3 14.5 3.0 by vegetation 2 density/yard 1980 14.5-31. 0 4.2-11.8 11.5-14.2 1.0 �100 Q 80 ! w ~ 60 co ::> ~ U) r.r.. 0 ~ 40 w u Il: W ll. l J/ / •....• _/ RI/ .l:"- 20 o __ ~ ~ __ ~a10 17 APR Fig. 4. 24 __~ ~ 1 '___~~ 8 15 MAY 22 __~ 29 '___ 5 JUN Progression of spring stubble plowing in the study region, 1963-68, 1970-76, and 1978-80. �15 Table 6. Property, Date Grasshopper densities Summer 1980. Location Vegetation obtained on and near the Sand Draw tlpe Cover height(ft) Grasshoppers/ yard2 Forbs 13 Jul 13 Aug NE~ S19 NE!t.S19 Sunflowers/cheatgrass Dock/sunflowers 1.5 - 4.0 14.5 2.5 31.0 Roadside 15 15 31 31 Jul Jul Jul Jul SW~ SW~ N~ NW~ S21 S17 S13 S28 Sunflowers/cheatgrass Cheatgrass Cheatgrass/sunflowers/Kochia Sunflowers/cheatgrass/Kochia 4.2 11.1 1.5 5.2 2.5 U.8 Reseeded pasture Mixed native grass Ungrazed shortgrass 0.5 - 1.0 l3.S U.S 14.2 Uncut ripe wheat Irrigated beans 3.0 - 4.0 1.0 1.0 0.4 Sorghum 2.0 - 4.0 35.0 Grass and grass/forb 16 Jul 13 Aug 13 Aug NW~ S30 NE~ S19 SE!t.S19 1.5 0.5 Cropland 13 Jul 15 Jul l3 Aug NE~ NW~ S~ NW~ S19 S32 S3S S19 food plot Pheasant Trapping and Transmitter Hen Monitoring Attachment Wheat stubble fields used by roosting hens thawed in mid-February 1980, became muddy, and did not freeze enough to permit access for nightlighting from pickups. Two local farmers, one with an all-terrainvehicle, assisted K. Snyder in trapping 10 hens on 11-12 March. A small tractor with lights and pulling a trailer containing the generator, was used to capture 24 additional hens on 18-19 March. The tractor was again used for field access to obtain 7 replacement hens on 20 April. The solar-powered single stage transmitters, weighing approximately 15-18 gms, were harnessed in a back-pack to the hens so the solar panels could receive direct sunlight. Flat, ~-inch wide nylon-elastic material was used for harnesses. In 1979, the harness straps came over the front of the wings, fastened together in the center of the breast, and then came up behind each wing to the rear of the transmitter. This permitted the transmitter to ride too far forward on the hen and feathers at the base of the neck frequently covered part of the solar panels causing inconsistent transmitter operation. Harnesses were modified in 1980 to come directly around under each wing. This �16 permitted the transmitters to ride further back and dramatically improved signal operation and ease of locating hens~ Some reduction in flight capability, primarily in slower take-off, was noted, especially in 1980. However, hens seldom fly during the spring-summer nesting season. One additional hen, #151-361, whose single stage capacitor-type transmitter signal had been lost shortly after release in 1979, was recaptured near Sand Draw by K. Snyder in spring 1980. The hen was fitted with a new harness, and released on 16 April. Her signal was again lost several days after release in 1980. Transmitter Efficiency The single stage solar-powered transmitters, containing ani-cad battery for temporary power storage, worked effectively most of the time. Several transmitters from mortalities were recovered in both 1979 and 1980 even though the transmitters were upside down. Enough light reflected onto the panels permitted operation at least part of the day. The transmitters became least effective during incubation in dense, tall stands of green wheat from late May to late June. However, persistent monitoring usually verified hen location sometime during the day. The dampening effect of tall vegetation on signal transmission distance also made monitoring difficult in late spring and early summer and when hens were in tall corn. Six transmitters purchased in 1979 contained capacitors instead of ni-cad batteries. These capacitor-type units have not produced signals consistent enough to permit effective monitoring, staking of nests, and collection of other pertinent information in most instances. Four of the six were lost during the last 2 years whereas 18 of 24 ni-cad equipped solar transmitters purchased in 1979 are still functional. Nine of 10 transmitters obtained in 1980 were recovered for future use, although 2 were returned for repair. Predation and Mortality Most hen pheasant mortality in the vicinity of Sand Draw occurred in April (Table 7). Seventeen of 29 known mortalities (58.6%) in springsummer 1979 and 1980 occurred in April. Predation by resident great-horned owls (Bubo virginianus) was suspected in a majority of the April losses. Hens were especially vulnerable in 1980 when snow covered all herbaceous cover during the first part of the month. In addition, the wheat stubble was short and open, and provided little protective cover at a time when they were joining harems. At the same time resident great-horned owls had increased food demands to feed their rapidly growing young. Information obtained during the last 2 years strongly supports the findings of Petersen (1979) in Wisconsin. He found that great-horned owls and red-tailed hawks (Buteo jamaicensis) reduced and held pheasant populations below carrying capacity with peak predation occurring in April. Weigand (1980), working with gray partridges (Perdix perdix) in Montana, found that the primary �Tahle 7. lien 020111 020112 040111 040112 060111 060112 080111 0801/2 102111 102112 118 140111 140112 160 180 200 850 863 887 913 919 935 962 971 013 032 052 087111 087112 107 146 163 186 202 258 309 337 341 361 387 439 448 Statistics Location tra~l'cd 19SW 19SI~ 19S\~ 19SW 19SW 19SW 19SW 19SW 19SW 19SW 19SW 19SW 24SE 19SW 19SW 19SW 19SW 19SW 19SW 19SW 19SW 19S\-1 19SW 19SH 19SW 19SW 19SW 19SW 24 SE 19SI. 19SW 24SE 24SE 24SE 24SE 24SE 24SE 24SE 24SE 24SE 24SE 24SE for radio-marked Date released 18 20 18 20 12 20 12 20 12 20 12 12 20 12 12 18 12 12 12 18 18 18 18 18 18 18 18 18 20 18 18 19 19 19 19 19 19 19 16 19 19 19 Mar Apr Har Apr Mar Apr Har Apr Mar Apr Mar Mar Apr Mar Mar Mar Mar Ha r Mar Mar ~jar Mar Mar Mar Mar Mar Mar Mar Apr Mar Mar Mar Mar Mar Mar Mar Mar Mar Apr Mar Mar Mar Projected age Juv Juv Juv Juv Juv Ad Juv Juv Ad Juv Juv Ad Ad Juv Ad Juv Juv Juv Juv Ad Juv Juv Ad Juv Juv Juv Ad Juv Ad Juv Juv Juv Ad Ad Ad Juv Juv Juv Ad Ad Ad Ad pheasant hens, spring and summer 1980. I~eight Known nest attempts (gms) 800 800 840 810 860 900 780 900 770 880 890 830 930 850 920 820 860 800 850 950 860 770 830 830 785 800 860 820 870 895 920 830 910 820 1050 780 780 860 880 850 910 900 0 1 0 1 0 2 0 1 Suspected attempts 1 a 1 1 0 1 1 Combined total Success No. young hatched 1 Yes 14 1 Yes 10 3 No 1 Yes 8 1 1 Yes 7 1 1 Yes Yes 11 6+ a 0 I 3 1 0 1 1 3 2 Yes No Yes No 1 Yes 2 1 1 1 No Yes 11 Yes 12 2 1 2 Yes Yes No 5 15 2 3 Yes Yes 11 5 1 2 Yes No 10 I 1 1 9+ 11 10 a 2 1 1 1 a 1 1 2 1 a 2 3 0 1 ,2 0 0 2 0 a 2 . , No lien fate (if known) ttor t a l.Lt y Retrapped l'ortality r.etrapped Mortality Retrapped Nortality Mortality Mortality Unknown . Retrapped Nortality Retrapped Retrapped Unknown Mortality Retrapped Hortality Retrapped Retrapped Unknown Retrapped Hortality Retrapped Retrapped Mortality Retrapped Mortality Retrapped Retrapped Unknown Mortality Retrapped Retrapped Mortality Retrapped Retrapped Unknown Unknown Unknown Unknown Mortality and released and released and r.eleased and released and released and released and released and released and released and released and released and released and released and released and released and released and released and released and released Approximate date 4-9 Apr 18 Aug 9-15 Apr 20 Aug 4-9 Apr 14 Aug 20-31 Mar 1-4 Aug 20-31 Mar 3 Jul 18 Aug 4-9 Apr 14 Aug 14 Aug 19 Apr 26-27 Apr 9 Oct 26 May 20 Oct 26 Aug 20 Apr 14 Aug 22 Apr 18 Aug 4 Nov Late May 20 Aug 20 Apr 9 Oct 20 Oct 3 Jul 20 Apr 26 Aug 9 Oct 4 Nov 24 Aug 18 Aug Late Apr 17 Apr 11 Aug 20 Mar Late Apr T~ee of mortalit~ Avian predation Predation suspected Avian predation Avian predljltion Unknown predation Avian predation (signal lost) Avian predation (signal lost) Unknown predation Unknown predation Unknown predation Unknown predation Unknown Avian predation (signal lost) Avian predation Unknown predation (signal (signal (signal (signal Unknown lost) lost) lost) lost) predation .- -..J �18 limiting cover in the same in spring factor appeared to be the quantity and quality of protective spring. Spring was the period of major mortality. He observed factors as found in the Sand Draw vicinity, i.e. minimal cover combined with an influx of raptors. All mortalities discovered by 20 April 1980 were hens trapped immediately south of the Sand Draw Property in Section 19 (Fig. 5). Hens trapped and resident in Section 24 southwest of Sand Draw sustained less and later predation. The impact of the transmitters on hen survival was unknown, however several carcasses of hens and males not marked with radio transmitters were found during both springs. Several mortalities found in shrubby cover in late April 1980 were possibly killed by Cooper's hawks (Occipiter cooperii). Coyotes (Canis latrans) were infrequent visitors to Sand Draw in late winter and no evidence of foxes was noted. Weasels (Mus tela spp.) feral house cats and dogs, prairie falcons (Falcomexicanus) and other predators were present. Striped skunks (Mephitis mephitis) and badgers (Taxidea taxus) were resident on and near Sand Draw but were not believed responsible for direct hen mortality. Some hens may have died because of weather or harness related stress, age, or other factors. These birds may then have been eaten by predators. Fourteen (41.2%) of the 34 hens trapped and instrumented in March died in spring. Contact was lost with 4 other hens, however, equipment failure, emigration, or mortality could have been responsible. Signals were lost from 2 hens in early July and 1 in August. One hen lost in early July was believed to be incubating at the time and predation was suspected. Equipment failure was the apparent reason for the other July signal loss. An accipiter was flushed from the partially consumed carcass of hen 102 on 7 January 1981. The nonworking transmitter was still intact. One hen with a capacitor type transmitter was relocated near Sand Draw in February 1981 after contact had been lost in August 1980. Spring Dispersal Significant spring dispersal away from the vicinity of Sand Draw occurred in 1980 and movements were more pronounced than in 1979. Thirteen hens showed no long-distance movements or made short shifts into green wheat, in late April - early May. Thirteen other hens had movements of usually at least 1 mile and often several miles (Table 8). Dispersal of hens peaked in late April with most movements to the east, northeast or southeast (Fig. 5). Seven hens moved outside the boundaries of the 9-section extensive study area. Other hens, whose signals were lost, may have dispersed equal or even greater distances. Searches for radiomarked hens were conducted for several miles away from Sand Draw in all directions but chances of locating hens diminished in proportion to the distance they moved and the height of green wheat. Increased dispersal made locating all hens more time consuming and difficult. �19 2 1 6 3 01 • 146 11 12 14 13 10 16 23 15 22 26 25 35 36 31 2 1 6 5 28 27 33 34 4 3 Fig. 5. Dispersal of hens moving extensive distances away from Sand Draw in Spring 1980. Known or suspected nests (.); successful nests ~. �20 Table 8. 1980. Timing and relative Period of dispersal distance of hen dispersal during spring Number of hens moving Long distances Short distancesa Totals Apr 2- 9 10-16 17-23 24-30 4 2 1 1 9 2 1 1 13 2 6 13 2 19 May 1- 7 Totals aShort distance movements appeared to be into suitable nesting cover rather than pronounced dispersals. Spring dispersal in 1979 (Snyder 1980) was much less pronounced than in 1980 and was characterized by shorter dispersal distances. Most movements occurred prior to or around 10-13 April 1979, approximately 2 weeks earlier than in 1980. Reason for the more pronounced dispersal in 1980 could have resulted from poorer nesting conditions, higher hen densities, or other variables. Nesting Activities Nest Initiation. -- The delayed dispersal in 1980 was followed by delayed initiation of nesting activity (Table 9). Nesting began in mid-to late April in 1979, but the earliest known attempts in 1980 were in early May and most activity appeared to begin in mid-May. The late March snow and subsequent early April snow cover possibly caused considerable stress to the hens and may have been a factor in delaying dispersal and onset of nesting. Table 9. Initiation pheasants, 1979-80.a of known first nesting attempts of ring-necked 1979 Interval 15-30 Apr 1-15 May 16-31 May > 1 June Totals 1980 N % N % 3 9 3 1 16 18.75 56.25 18.75 6.25 0 9 9 3 21 0.0 42.9 42.9 14.2 aThese were the first located nests per hen. Other prior nesting attempts may have occurred for some hens but were not confirmed. �21 Cover Type Selection for Nesting. -- Although one or more hens may have started nests in wheat stubble in 1980, these nests were not verified, and were destroyed before incubation started. Quality of the wheat stubble was poor (Table 2). In contrast, excellent stands of green wheat made rapid growth in late April and May (Fig. 1, Table 2) and attracted hens so that most known first nesting attempts occurred there (Table 10). Renesting was initiated in May and June and was concentrated in green wheat, the only significant nesting cover available. Fence rows were known sites for two nesting attempts and one suspected attempt. Wheat stubble was used for late summer nest efforts after completion of wheat harvest (Table 10). Five of 26 nests (19.2%) in green wheat in 1980 were in turnrows where overlapping drill furrows placed wheat clumps more closely together than normal. Three other nests were in narrower than average drill rows. Three hens nesting in Section 24, all used a field drilled with 10-inch row spacing. Two nearby and readily accessible fields of equal size containing 14-inch row spacings were not used. These data indicate nest site selection in locations with higher than normal densities of green wheat clumps. However, other hens selected nest sites in green wheat which were exposed and open. Cheatgrass, if present in green wheat, was frequently selected for nest placement. Nest Fate in Relation to Cover Type and Location. -- Delayed stubble tillage (Fig. 4) would have destroyed many incubated nests if hens had nested in wheat stubble. Except for one abandoned nest (reason for abandonment unknown), all nests placed in green wheat that were not destroyed by predators hatched yielding large clutches (Table 10). These successful nests provided most of the excellent production observed in the vicinity in 1980. A few renest efforts hatched prior to wheat harvest, but those that didn't were destroyed or covered with straw resulting in hen abandonment. Hen 087 nested in an open weed-dominated turnrow in ripe wheat just north of Sand Draw. Due to her location in the corner of the triangular field, both the combine and a tractor and sweep plow missed her nest while she was incubating. This was the only hen to successfully complete a nest in a field subjected to wheat harvest activity. In contrast, hen 887 hatched a clutch from her 2nd known nest on 11 July only to have a combine wheel crush the young before they were old enough to leave the nest. As in 1979, the Sand Draw property continued to be a poor production site and no nests of radio-marked hens were known to have been successful there in 1980. Resident striped skunks and badgers were suspected of extensive, repetitious predation to nests of 5 monitored hens whose activities also centered in the northeast part of the Property. Several nests were suspected but could not be verified before they were assumed destroyed. Only 2 of the 5 hens brought off young near the property. These were relatively late hatches lending evidence that persistent nest predation probably occurred on and near the Sand Draw Property. �Table 10. Data on known nesting attempts and fates, radio-marked pheasant hens, 1980. Nesta number Hen Layingh onset Cover type association Incubation onset ·Termination date Days incubated Clutch size Nest fate Number chicks 0-020 Green wheat 11 May Jun 23 Jun Full term 16 Hatched 0-040 Green wheat 4 Jun 16 Jun 9 Jul Full term iO Hatched 5 0-060 Fence row mid-May 2 Jun 9 Jul link Pred.dest. 14 Green wheat-cheatgrass 11 Jul 21 Jul 22 Jul 6 Abandon o o 0-080 Green wheat 30 May 9 Jun 2 Jul Full term 8 Hatched 8 0-118 Green wheat 17 May 4 Jun 27 Jun Full term 14 Hatched 7 11 2 0-140 Green wheat 15 May 30 May 21 Jun Full term 12 Hatched 0-160 Green wheat late May early Jun early Jun Full term unk Hatched 6+ 0-850 Green wheat 10 May 22 May 15 Jun Full term 10 Hatched 9 Late May Unknown unk Pred .dest. Crushed o o 11 Green wheat mid-May late May 2 Green wheat 6 Jun 16 Jun 9 Jun 3 0-887 23 7+ Wheat stubble 18 Jul 1 Aug 23 Aug Full term 11 Hatched 0-913 Green wheat 14 Jun 22 Jun 11 Jul 19 6 Abandon o 0-935 Green wheat 16 May 29 May 21 Jun Full term 10 Hatched 10 Green wheat 21 May 4 Jun 15 Jun 11 11 Pred.dest. Green wheat-cheatgrass late Jun early Jul 11 Jul 4 or 5 7 Pred.dest. 0-971 2 o o 1-013 Green wheat 25 May 8 Jun 30 Jun Full term 11 Hatched 11 1-052 Green wheat 18 May 4 Jun 27 Jun Full term 14 Hatched 12 1-087 Weeds-ripe wheat Jul 11 Jul 3 Aug Full term 8 Hatched 5 Full term 15 1-107 Green wheat 3 May 1-146 Green wheat 4-8 May 2 Fence row (weeds) Green wheat 1-186 Transmitter functioned improperly Young killed in nest by combine wheel Combine flushed at wheat harvest Believe predator destroyed night before wheat harvest Nest missed by combine & sweep plow (in corner) 16 Jun 15 Hatched 11 Jun 17-20 14 Pred.dest. mid-Jun late Jun Unknown Unknown unk Pred.dest. 7 May 24 May 16 Jun Full term 13 Hatched 11 11 Aug 18 Aug 18 Aug 6 Abandon Hen flushed from nest Nest partially crushed by combine wheel o o Green wheat 14 May 28 May 8 Jun 1111 11 Abandoned Green wheat 14 Jun 25 Jun 11 Jul· 1616 9 Destroyed o 3 0-1 -- Wheat stubble 21 Jul 31 Jul 22 Aug Full term 8 Hatched 5 1-309 Green wheat 14 May 27 May 19 Jun Full term 11 Hatched 10 1-387 Green wheat mid-May late May 11 Jun 14+ unk Pred.dest. o Green wheat 25 Jun 6 Jul 13 Jul 7 9 Abandon 9 2 3 of 10 died at hatch 22 May 2 1-202 Wheat stubble Flushed by combine 21-26 May o o 2 Comments Hen signal lost Chicks may have died when young. Hen renested Unknown Flushed by combine and nest covered under straw 22 aFirst, 2nd or 3rd confirmed nest. Other previous nests could have been established for which information was not available. bprojected dates based on clutch size and a laying rate of 1.3 days/egg. N N �23 Nest Success Only 16 hens from the 20 April population of 34 -- less than one-half -were known to have successfully brought off broods in 1980 totaling approximately 150 chicks (Table 10). One of these broods wa s probably lost at an early age because the hen (186) attempted to renest in August. Five hens that survived to late summer were unsuccessful nesters and apparently discontinued nesting activities. In 1979, only 1 hen was unsuccessful. Seven other hens were known to have died between 20 April and early summer and did not have a chance to complete nests. Signals were lost from 4 hens in spring and in early July from 2 other unsuccessful hens. One transmitter whose signal was lost in spring was recovered in November from an apparent early fall predation. This suggests that dispersal was the reason for signal loss. Wheat harvest, which flushed hens from nests, crushed nests or covered them with straw, was the primary factor in several hens being unsuccessful. Extremely hot, dry weather prevailed during and after wheat harvest, which may have stymied renest efforts. However, 2 hens renested in late July and completed successful late summer clutches. Clutch size of 14 nests started in May 1980 averaged more eggs per nest (12.14) than in subsequent months. Four June and 4 July nests averaged 8.50 and 8.25 eggs per nest, respectively. These findings are similar to those obtained in 1979. Hatchability of eggs was apparently higher among earlier nests. Hatchability averaged 87% for May initiated nests, 85% for June nests, and 75% for July nests based on combined 1979-80 samples of 18, 5.and 5 nests, respectively. These data did not show any marked difference in.hatchability between the 2 years. Young per successful nest averaged 9.6 in 1980 compared to 8.4 in 1979. Hen Use of Vegetation Types Hen use of vegetation types varied seasonally in 1980 (Fig. 6, 7). Monitoring often did not represent early morning and late evening feeding periods so the data only partially represent hen use of available habitats. The high use of wheat stubble and woody cover in April rapidly gave way to high use of green wheat through May and June (Fig. 6). Use of annual forbs increased markedly after mid-June with corn and other row crops becoming increasingly important in July, August, and September. Hen occurrence in each vegetation type was examined by activity periods (Fig. 7). These data indicate rapid movement into green wheat during laying and incubation with subsequent movement to weedy habitats and green row crops for brood rearing. Reduced use of wheat stubble in spring and later summer 1980 was documented in contrast to 1979. Relative importance of cover types for use in relation to availability can be made (Tables 3, 4; Figs. 6, 7). Direct comparisons cannot be made because several hens resided outside the 9-section extensive study area. �24 100 90 80 GREEN WHEAT 70 60 40 30 20 10 ~ N .1 OJ APR \D r-f It"I 1 1 N N M MAY JUN .., r-f r-f I 0'\ r-f 1 r-f N JUL Fig. 6. Use of cover types by radio-marked hen pheasants intervals from early April through early September 1980. AUG It"I N 0'\ J, 1 N or-t N SEF at biweekly �25 100 90 :::::::::::::::: ":::::::::::::: 80 Corn & Sorghum Green Wheat 50 40 30 20 & 10 Stubble 0 !:i! U1 0:: I:il {!)a. ~~ 0::0 a. til Fig. 7. habitats gel {!) ~ )-i .:( o tl )-i 0:: a. ~ <...:a ~ ~ ~ u Z H el § I:il~ :X:1:il ~~ :x:r-t Percent occurrence of radio-marked during activity period in 1980. 0:: ~ ~ N r-t 0:: I:il til 0 ~a ~~ <""1< ~~ ~.:( u~ I:ilO O::Z hen pheasants in different �26 Hen Brood Activities Use of Vegetation Types. -- Hens, after successfully hatching a clutch, usually remained in green or ripening wheat for only a few days before moving to cover apparently more suited for brood rearing (Fig. 3). The fact that hen 186, who subsequently renested, remained in green and ripening wheat away from weeds, grass, or row crops for an extended period, lends support to the hypothesis that her brood may have died at an early age. Hens nesting in fields near the Sand Draw Property usually moved into the forb-grass-shrub complex there to rear broods. This area contained abundant grasshoppers preferred by young chicks (Table 6). The disturbance tillage strips dominated by sunflowers, the old and reseeded alfalfa and grass-alfalfa stands, and the sorghum food-cover plantings received high use by hens with broods in 1979 and 1980. Hens with broods residing away from Sand Draw moved to roadsides and weedy edge areas, to grass-forb pastures, and to irrigated row crops (corn, sugar beets, and beans). Their activities were concentrated primarily along the edge of large fields. Movements into irrigated fields placed hens and broods into close association with aerially applied insecticides and fungicides throughout July and August. Several, if not all corn fields, containing radio-marked hens, were sprayed with insecticide mixtures. Fungicides were used on both sugar beet and pinto bean fields. No direct mortality of radio-marked hens was detected and the fate of broods in corn could not be determined. Moderately high numbers of pheasants remained in these fields until crops were harvested in September and October. This pattern of henbrood movement to irrigated fields, which were subsequently spr ayed .with insecticides,occurs through most of eastern Colorado's pri~ary pheasant range. Movements in Relation to Water. -- Water was available in all irrigated fields and from windmill overflow on Sand Draw. There was a definite movement of hens with broods toward these areas (Fig. 3). However, whether these movements were toward green vegetation and increased insect abundance, free water, or both, could not be determined. Brood Size and Mortality. -- Limited information was obtained concerning brood size and brood reduction with time. Young chicks were difficult to flush and little confidence is placed in the accuracy of counts of flushed broods. Two or more hens with broods were often seen together and some mixing of chicks among broods may have occurred. Consequently, flushes of hens and broods were minimized rather than subject the hens and young to excessive harassment with possible increased mortality and movements. �27 Evaluation of the Impact of Minimum Tillage Summer Fallowing Information concerning hen pheasant use of chemically treated stubble was not obtained in 1980. No nests, if they occurred, were documented in wheat stubble because of the excellent stands of green wheat available for nesting. Wheat stubble in spring was short, open, partially lodged by heavy spring snow and some fields were partially dominated by cheatgrass which did not provide nesting cover. In addition the atrazine-2,4-D herbicide mixture did not surpress spring germination of cheatgrass on the treated fields. Therefore, the leasee was instructed to till the treated fields in early May rather than to let a new crop of cheatgrass mature and produce seed by late June. Herbicide treatments were applied to 50 acres of better quality wheat stubble in mid-August 1980 by the leasee and a commercial applicator. However, little significant rain was subsequently received to place it into the topsoil. Its effectiveness in supressing spring weed growth will not be determined until spring 1981. Fall-Winter 1980-81 Conditions Mild, dry weather permitted harvesting of all row crops earlier than normal in early fall 1980. Most corn fields were disced and/or plowed prior to- onset of the 8 November 1980 pheasant season. Pheasant numbers were above average in the Sand Draw vicinity, but lack of significant snow permitted birds to remain in large stubble fields throughout the fall and winter. Abundant, moderately tall wheat stubble and the learned ability of the male pheasants to avoid hunters with season progression made them difficult to find or approach after the first 2 weekends of the hunting season. Only one snow, in early March 1981 was sufficient to fill the stubble and force pheasants to seek other cover. No significant weather related mortality occurred and an abundant, well dispersed breeding population was present going into spring 1981. 1981 Trapping Activities Nightlight trapping and instrumenting of hens with transmitters was initiated on 2 March 1981 and halted 3 March by deep snow. Trapping of 42 hens was resumed and completed on 19-20 March. LITERATURE Petersen, L. 1979. in southeastern 63pp. CITED Ecology of great-horned owls and red-tailed hawks Wisconsin. Wis. Dep. Nat. Resour. Tech. Bull. Ill. �28 Robel, R. J., J. N. Briggs, A. D. Dayton, and L. C. Hulbert. 1970. Relationships between visual obstruction measur ement s and weight of grassland vegetation. J. Range Manage. 23:295-297. Snyder, W. D. 1979. Evaluation of nesting cover preferences of pheasants in relation to wheat farming methods. Colo. Div. Wildl. Prog. Rep., Fed. Aid Proj. W-37-R-32, Work Plan 1, Job 22. Pp. 1-10. 1980. Evaluation of nesting cover preferences of pheasants in relation to wheat farming methods. Colo. Div. Wildl. Prog. Rep., Fed. Aid Proj. W-37-R-33, Work Plan 1, Job 22. Pp. 1-40. u.S. Department of Commerce. National Oceanic and Atmospheric Administration. 1980. Climatological Data Annual Summary, Colorado. 85. No. 13. Prepared by ~~//~"_,~ __·__~~~__.~·rf_·'~,~~-~~·~~·~~=·-~=·~------Warren D. Snyder r tft.?) Wildlife Researcher C· �29 April 1981 JOB FINAL REPORT State of Colorado Project No. W-37-R-34 ._--- - - ----, --- 3 Work Plan No. Job Title: Game Bird Survey Vulnerability Job No. and Population .---- 11 Characteristics of Sage Grouse in Moffat County Period Covered: Personnel: 1 April 1978 through 31 May 1981 Jack Baker, Mike Bauman, Tom Beck, Kevin Berner, Clait Braun, Marty Bray, Charles Brown, Jack Corey, Larry Crooks, Jim Dingman, John Ellis, Steve Emmons, Howard Funk, Ken Giesen, Larry Green, Jim Haskins, Dave Hoart, Don Hoffman, Richard Hoffman, Laura Spess Jackson, Ken Kendall, Tom Lines, Debra Martin, Russ Mason, Jim Miller, Sue McElderry, Brent Renfrow, Dave Roche, Ed Rodriquez, Glen Smith, Trish Sweanor, John Testa, Lou Vidakovich, Duane Wagner, Nancy Watkins, Sue Wielgopolan, Perry Will, and Charles lA)'oodward, Colorado Division of Wildlife. ABSTRACT Population characteristics, harvest statistics and vulnerability to hunting were investigated for sage grouse (Centrocercus urophasianus) in Moffat and western Routt counties, Colorado from 1978 through spring 1981. Counts of male sage grouse on leks increased in 1979 and decreased in 1980 and 1981. Brood counts were discontinued after 1978 as no correlation could be found with percent chicks in the fall harvest. During the 3-year period, 2,824 sage grouse were banded; 506 in 1978; 1,250 in 1979: and 1,068 in 1980. This included 1,059 chicks, 476 yearling males, 934 adult males, 177 yearling females and 178 adult females. Trapping of adults and yearlings was most successful using spotlights and long-handled nets while most chicks were captured using cannon nets and walk-in traps. Three check stations and 16 volunteer wing collection stations were used to collect harvest data and wings. Birds harvested per hunter varied and was highest (2.3) in 1978 and lowest (1.4) in 1980. Longer sage grouse seasons increased number of hunters and hunter days but did not noticeably influence total birds harvested. Most sage grouse hunters originated from the Eastern Slope with the greatest increase in the 3-year period being in the number of hunters from the Denver metropolitan area. Age and sex structure of the harvest varied among years with good production in 1978 and only average production in 1979 and 1980. The estimated annual mortality rate for adult males was 50.3%; that for adult females was 41.5%. Nesting success and production varied by area and year with adult hens being consistently more successful than yearlings. A high percentage of the adult and yearling hens ovulated each year. Total harvest fluctuated between �30 5,185 and 7,840 sage grouse and was not affected by season length or bag and possession limits. Hunters responded to more l.iberal seasons by hunting more frequently with substantial pressure occurring late in the season. Small Game Management Units 16, 18, 20 and 14 in that order provided the bulk of the hunting opportunity and sage grouse harvest. Direct recovery rates of all sage grouse were low (7.0-10.9%) but chicks were more vulnerable to hunting than were adults and yearlings. The direct recovery rates of chicks markedly increased from 1978 to 1980. RECOMMENDATIONS 1. The number of active sage grouse leks and their locations annually ascertained. This is a management function. should be 2. Check stations should be annually operated at Cedar Mountain, Dinosaur and Maybell on the opening weekend of the hunting season to obtain wings from hunter harvested sage grouse. This is a management function. 3. Wing barrels should be placed at the following locations to obtain wings from hunter harvested grouse: California Park Road, M.C. Road 2, M.C. Road. 101, Colorado 318 at Maybell, National Park Road at Dinosaur, Junction of M.C. 3 and 7, Lay Creek north of U.S. 40, Lay Creek south of U.S. 40, Maybell City Park, M.C. Road 3, and M.C. Road 4. Wings should be removed on Monday morning following the first weekend, Friday before the second weekend and on Monday following the second weekend when the wing barrels are removed. Wings should be stored and frozen in plastic bags with each bag being labeled with date and. location. This is a management function. 4. Studies of direct harvest rates of chick sage grouse on Cold Spring Nountain should continue as part of an intensive study of movements and recruitment to leks. This is a research function. �31 VULNERABILITY AND POPULATION CHARACTERISTICS OF SAGE GROUSE IN MOFFAT COUNTY Clait E. Braun Sage grouse are hunted annually in Colorado, with season lengths typically of 3 days and bag possession limits of 2 and 4 until recent years. Intensive bandings of adult and yearling sage grouse along with experimental hunting seasons in North Park, Jackson County, Colorado have indicated that only about 10% of the fall population of these segments is annually harvested. Data from this area indicate that yearling males are most vulnerable, with adult males and hens of both age classes having lower rates of exploitation. Knowledge concerning population characteristics and harvest rates are important if maximum allowable recreational opportunity through hunting is to be achieved. If turnover rates are moderately high (40-60%) and recovery rates (and by relationship, vulnerability rates) are low, conservative seasons and bag limits have little merit. This is the final report of a 3-year study to investigate population characteristics and vulnerability of sage grouse in Moffat County and adjacent areas in western Routt County, Colorado to hunting. The period covered was from mid-March 1978 through late May 1981. P. N. OBJECTIVES 1. Estimate young of the year vulnerability to adults and yearlings of each sex. 2. Provide reliable estimates 3. Provide reliable data on characteristics 4. Estimate to hunting of harvest rates in relationship (recovery rates). of the harvest. turnover and survival rates. METHODS AND MATERIALS Counts of male and female sage grouse on leks were made in early morning from late March through late May following procedures described by Rogers (1964) and Braun and Beck (1976). Intensive efforts to locate new leks were made only in 1978. Sage grouse were located and trapped at night from late March through late May in 1979 and 1980 where they roosted along trails, and on and near leks. During mid-July to early September, sage grouse were located and trapped where they roosted at night along trails and throughout the early morning and evening where they concentrated in meadows. Methods used included spotlighting and capture with long-handled nets, wire drift fences with a cannon net at the end of the fence, bumper mounted cannon nets, and modified lily pad traps (Lacher and Lacher 1964, Guillion 1965, Braun and Beck 1976). Birds captured were weighed, measured, classified as to age and sex (Eng 1955, Beck et al. 1975) and banded with serially-numbered aluminum leg bands (size 16 for males, 14 for females) and plastic �32 bandettes County). color-coded for year and location (5 different areas in Moffat Sage grouse hunters in Moffat County in 1978-80 were required to have in their possession while hunting a free numbered permit. Permits, unlimited in number, were available from Division of Wildlife offices in Craig, Denver, Fort Collins, Colorado Springs, Grand Junction, Meeker, and Montrose, all license agents in the Moffat County area, and all project and management personnel working in the area. Questionnaires were sent to all permittees immediately following the end of the sage grouse hunting season in Moffat County. One follow-up letter and questionnaire were sent in mid-October to all nonrespondents to the initial mailing. Check stations were operated at Cedar Mountain (junction of M. C. roads 3 and 7), Dinosaur (junction of National Park Service road and U.S. Highway 40) and Maybell (Colorado Highway 318 at U.S. Highway 40). Each station was operated on Saturday and Sunday of the first 2 weekends (except Maybell which was not open during the 2nd Saturday) of the sage grouse hunting seasons in 1978-80. Each station was operated from about 0800 to 1900 MDT depending upon traffic load. Data obtained per party were: county of origin, number of hunters, hours hunted (total of all hunters in each party), birds observed, birds bagged, birds lost, number of banded birds and location where each was harvested, areas hunted, and information on previous sage grouse hunting experience. One wing was obtained from 89 to 95% of the birds checked (1979: 1,692 of 1,898; 1980: 1,218 of 1,284). Ovaries of adult and yearlings were classified as to recent ovulation status and a sample was collected each year for laboratory examination. Sex of a sample of juvenile sage grouse each year was ascertained by gonadal inspection. These data were recorded on tags with wings being individually marked with corresponding information concerning actual sex of that bird. Wings were frozen and stored for later analysis. In addition to the 3 check stations, field checks of hunters were made on the opening weekend and 2nd Saturday at Cold Spring Mountain (1979-80) and wing barrels and signs (Hoffman and Braun 1975) were placed at 16 locations in Moffat and western Routt counties, including the 3 check station sites. Volunteer wing collection stations were in operation the entire season at all sites each year. Wings were collected following each weekend and week and frozen for later analysis. Collected wings were thawed and classified as to age (chicks, yearlings, adults) and sex following procedures outlined by Eng (1955) and Beck et al. (1975). Hatching dates were calculated for chicks of the year using data from Pyrah (1963). Ovaries collected were stored in AFA (alcohol-formalin-acetic acid) and later examined for presence of ovulated follicles as described by Kabat et al. (1948). �33 DESCRIPTION OF AREA Moffat County is in extreme northwestern Colorado, bordered by Hyoming on the north, Utah on the west, Rio Blanco County on the south and Routt County on the east. The major drainages include the Green River which flows north to south through western Moffat County, the Yampa River which flows east to west in the southern part of the county to its intersection with the Green River, approximately 3 miles (4.8 km) from the Utah border, and the Little Snake River which flows northeast to southwest through the middle of the county where it joins the Yampa River in Lily Park. The climate in Moffat County is semiarid with 8-20 inches (20.3-50.8 em) of precipitation annually. Precipitation occurs mainly in late summer to early winter. Mean annual precipitation for 6 weather stations was 12.99 inches (33.0 em). The annual mean temperature for 5 stations was 43.3 F (6.3 C) during years of record. in Moffat County varies from approximately 4,600 to 11,045 feet (1,402-3,367 m). The county seat, Craig, located in eastern Moffat County, is 6,240 feet (1,902 m) above sea level. Elevation Sagebrush ranges comprise approximately 60% (2,841 mi2; 7,358.2 km2) of the total land area of Hoffat County. Nonsagebrush ranges consisting of pinonjuniper, rough and rocky mountain shrub, aspen, or spruce-fir types comprise most of the remaining 40% (1,902 mi 2; 4,926.2 km2-) of the total land area (Hoffman 1979) . RESULTS AND DISCUSSION Counts of Sage Grouse on Leks Counts of sage grouse on known leks in Moffat and western Routt counties were made from late March through late May each year from 1978 through 1981. All data for 1978 and 1979 were presented earlier (Braun and Hoffman 1979, 1980). Data for 1980 (Table 1) and 1981 (Table 2) and summaries for all years (Tables 3, 4, 5) are presented in this report. Average number of male sage grouse counted on leks increased in 1978 and 1979 and decreased in 1980 and 1981 (Table 4). When all available data are considered (1969-81, Table 5), it is apparent that average number of males on leks decreased from 1969 through 1972, increased in 1973, decreased to a low in 1975 and 1976, increased to a high in 1979 and decreased in 1980 and 1981. Reasons for the fluctuations in average number of males counted on leks are not readily apparent as they occurred even with more intensive efforts in 1976-81 (number of counts per lek) which resulted in more leks being located (increase from 17 in 1975 to 72 in 1981 (Table 5). Beck and Braun (1981) presented data from North Park, Colorado which documented daily fluctuations of 25-100% in number of male sage grouse on leks when the leks were intensively surveyed (3-7 days per week). It is quite obvious that lek counts of male sage grouse vary daily, weekly, and markedly throughout the display season. In addition to daily and weekly changes in behavior patterns, sage grouse leks in Moffat County vary in elevation from 1,400 to over 3,300 m. Consequently, timing of breeding activities differs from area to area within the county and between "early" and "late" snow years. �34 Table 1. Peak lek counts of sage grouse, Moffat No. of counts Lek Males County, Colorado, Date(s) Females 1980. Date(s) Blue Mountain Bear Creek Escalante Haslim Cow Camp Karren Ranch Sixteen Road State Line Thirteen Mile Twenty-five Marker 7 7 6 4 7 5 4 1 36 45 65 61 44 27 2 18 19 9 9 13 13 13 13 7 May May May May May May May May 5 2 5 4 2 3 0 0 19 May 15,20 May 9,13 May 8,13 May 21 May 13 May All dates 7 May 1 1 0 0 46 45 27 May 23 Apr 2 14 2 2 2 0 3 2 5 5 3 3 2 0 1 4 3 29 21 4 May 21 Apr 25 May 0 24 3 All dates 21 Apr 25 May 57 2 8 5 1 27 0 25 Apr 30 Apr 8 Apr 19 Apr 11 May 25 Apr All dates Cold Spring Beaver Basin Gee Flats Goodman Draw Summit Spring 27 May 23 Apr East of Baggs Highway Cowboy Reservoir Eighty Road Elk Mountain Elkhead Creek Fan Rock Fly Creek Four Mile Creek 1 Four Mile Creek 2 Four Mile Creek 3 Mule Creek Sage Creek Slater Park TWenty-nine Road 1 Twenty-nine Road 2 Northcentral 0 92 25 Apr 30 Apr 19 Apr 29 Apr 29 Apr 25 Apr All dates 2 Apr 24 Apr 0 45 2 Apr 21 Apr 9 8 19 16 16 16 19 7 East Cottonwood Gulch Mud Spring Draw Pole Gulch Timberlake 1 Timberlake 2 Timberlake 3 Timberlake 4 Timberlake 5 West Timberlake 2 Northcentral 160 15 26 36 81 68 0 0 1 3 2 8 4 4 2 2 1 77 31 149 38 37 34 28 9 9 19 16 16 16 19 19 Apr May Apr Apr Apr Apr Apr Apr 0 8 21 52 12 42 11 1 3 6 3 3 3 3 55 25 28 26 25 59 8 22 22 23 1 22 May Apr May Apr May Apr 10 11 2 1 0 23 Apr Apr Apr Apr Apr Apr Apr May North Big Hole Butte Big Hole Gulch Cox Ranch Dressler Gulch 1 Dressler Gulch 2 Scandinavian Gulch 8 May 14 Apr 22 May 23 Apr All dates 22 Apr �35 Table 1. (continued) No. of counts Lek Northcentral North Date(s) Females Date(s) 8 May 22 Apr 8 May 0 7 0 8 May 14,22 Apr 8 May (cont.) Seven Uile Reservoir Thornburg Gulch Twenty-one Road West Timberlake 1 Northcentral Males 1 3 1 0 0 40 0 2 3 3 3 2 2 3 1 4 2 2 3 0 25 48 27 37 123 35 34 53 0 6 37 All dates 18 Apr 15 Apr 23 Apr 6 May 27 Apr 22 Apr 17 Apr 10 Apr All dates 15 Apr 17 Apr 0 3 13 13 2 17 3 4 5 0 0 13 All dates 18 Apr 15 Apr 23 Apr 6 May 27 Apr 15,22 Apr 17 Apr 10 Apr All dates All dates 17 Apr 6 3 3 3 4 0 2 5 5 2 4 3 3 2 4 70 83 31 50 79 17 Apr 10 May 4,17 Apr 8 Apr 8 Apr 24 4 59 9 20 8,17 Apr 10 May 4 Apr 8 Apr 8 Apr 0 62 9 0 32 45 28 44 27 All dates 28 Apr 28 Apr All dates 28 Apr 25 Apr 20 May 28 Apr 17 Apr 0 54 0 0 3 3 3 3 6 All dates 8 Apr All dates All dates 10 May 25 Apr 20 May 17 Apr 17 Apr 3 4 1 4 1 3 2 2 3 40 32 2 21 13 39 54 22 28 South Big Gulch 2 Big Gulch 3 Bord Gulch Grassie Reservoir Greasewood Lay Creek North Fork Big Gulch 2 Sand Creek Spring Creek 1 Spring Creek 2 Spring Creek 3 Upper Nineteen Road South of U.S. 40 Axial Basin Boxelder Gulch 1 Boxelder Gulch 2 Deception Creek Dry Lake 2 Duffy Mountain Horse Gulch Juniper 1 Juniper 2 Morgan Gulch 1 Morgan Gulch 2 Morgan Gulch 3 Round Bottom Temple Gulch Yellow Jacket Sunbeam West - North Coffee Pot Spring Cross Mtn. 1 Cross Mtn. 2 Cross Mtn. 3 East Sand Wash Lone Tree Powder Wash Hill Snake River West Thornburg Well 21 24 11 11 16 18 21 16 18 May Apr Apr Apr Apr Apr May Apr Apr 2 7 0 6 0 19 3 1 2 21 11 11 11 16 3 21 16 18 May Npr Apr Apr Apr Apr May Apr Apr �36 Table 2. Peak lek counts of sage grouse, Moffat No. of counts Lek County, Colorado, Females 1981. Males Date(s) Date(s) 4 3 4 3 3 3 3 3 18 55 83 54 0 68 0 22 13 Apr 7 May 13 Apr 7 May All dates 20 Apr All dates 6 Apr 6 6 5 5 36 26 8 12 27 Apr,1 May 27 Apr 28 Apr 21 Apr 4 6 0 1 7 5 4 3 4 4 6 6 6 4 2 0 3 3 22 53 5 24 85 12 6 22 27 48 4 5 May 23 Apr 20 Apr 29 Apr 27 Apr,5 May 6 Apr 2 Apr 12 Apr 27 Apr 5 May 7 Apr 14 21 8 8 21 5 0 8 15 12 2 2 Apr 16 Apr 16 Apr 16 Apr 6 Apr 19 Apr All dates 12 Apr 12 Apr 6 Apr 7 Apr 0 52 All dates 15 Apr 0 11 All dates 8 Apr 1 5 5 4 6 5 4 4 4 25 36 52 12 95 14 28 0 47 8 Apr 1,20 Apr 14 Apr 31 Mar 11 Apr 11 Apr 8 May All dates 11 Apr 0 9 56 5 82 2 29 0 10 Only date 20 Apr 2 Apr 31 Mar 31 Mar 11 Apr 31 Mar All dates 11 Apr 4 6 5 5 5 6 19 19 6 18 7 27 6 16 0 8 4 39 8,14 Apr 8 Apr All dates 7 May 7 May 9 Apr Blue Mountain Bear Creek Escalante Haslin Cow Camp Karren Ranch Sixteen Road State Line Thirteen Mile Twenty-five Marker 8 17 72 24 0 39 0 12 13 Apr 6 Apr 13 Apr 6 Apr All dates 13 Apr All dates 6 Apr Cold Spring Beaver Basin Gee Flats Goodman Draw Summit Spring 4 May 4 May All dates 28 Apr,6 May East of Baggs Highway Cowboy Reservoir Eighty Road Elk Mountain Elkhead Creek Fan Rock Fly Creek Four Mile Creek 1 Four Mile Creek 2 Four Mile Creek 3 Mule Creek Sage Creek Slater Park TWenty-nine Road 1 Twenty-nine Road 2 Northcentral East Cottonwood Gulch Mud Spring Draw Pole Gulch Timberlake 1 Timberlake 2 Timberlake 3 Timberlake 4 Timberlake 5 West Timberlake 2 Northcentral North Big Hole Butte Big Hole Gulch Cox Ranch Dressler Gulch 1 Dressler Gulch 2 Scandinavian Gulch 14 7 10 29 29 21 Apr May Apr Apr Apr Apr �37 Table 2. (continued) No. of counts Lek Northcentral North Date(s) 3 6 4 5 0 23 20 1 All dates 11 May 23 Apr 14 Apr 0 1 0 0 All dates 11,14 Apr All dates All dates 5 3 5 3 1 6 4 4 4 4 2 22 0 20 41 54 17 28 33 15 9 26 14 Apr All dates 29 Apr 10 Apr 10 Apr 13 Apr 10 Apr 10 Apr 10 Apr 16 Apr 8 Apr 16 0 13 26 20 8 10 4 1 2 6 1 Apr All dates 1 Apr 10 Apr 10 Apr 1 Apr 10 Apr 10 Apr 10 Apr 16 Apr 8 Apr 1 3 3 6 6 1 0 4 4 3 4 3 4 5 5 54 72 16 25 108 5 28 14 1 1 45 0 7 8 13 1,6 6 Only 15 6 0 8 13 4 13 13 26 Mar 9 Apr All dates 8 Apr 8 Apr 15 Apr 26 Mar 7 Apr 4 3 2 3 4 4 4 4 5 20 22 15 22 5 53 16 8 20 5 3 3 0 0 20 30 0 7 31 Mar 29 Mar 16 Apr All dates All dates 29 Mar 31 Mar All dates 29 Mar Females Date(s) (cont. ) Seven Mile Reservoir Thornburg Gulch Twenty-one Road West Timberlake 1 Northcentral Males South Big Gulch 3 Bord Gulch Grassie Reservoir Greasewood Gulch Lay Creek North Fork Big Gulch 2 Sand Creek Spring Creek 1 Spring Creek 2 Spring Creek 3 Upper Nineteen Road South of U.S. 40 Axial Basin Boxelder Gulch 1 Boxelder Gulch 2 Deception Creek Dry Lake 2 Duffy Mountain Horse Gulch Juniper 1 Juniper 2 Morgan Gulch 1 Morgan Gulch 2 Morgan Gulch 3 Round Bottom Temple Gulch Yellow Jacket 62 18 2 21 26 12 35 24 7 21 15 13 6 11 Apr Apr May Apr Apr Apr 26 Mar 9 Apr 7 Apr 13 Apr 15 Apr 15,27 Apr 26 Mar 7 Apr Apr Apr Apr Apr Apr date Sunbeam West - North Coffee Pot Spring Cross Mtn. 1 Cross Mtn. 2 Cross Mtn. 3 East Sand Wash Lone Tree Powder Wash Hill Snake River West Thornburg Well 24 29 16 29 25 29 13 13 5 Apr Mar Apr Mar Apr Mar Apr Apr Apr �38 Table 3. Lek Peak lek counts of sage grouse, Moffat County, Colorado, 1976-8l. a 1976 Blue Mountain Bear Creek Escalante Haslim Cow Camp Karren Ranch 26 Sixteen Road State Line Twenty-Five Marker (1980) Cold Spring Beaver Basin Gee Flats Goodman Draw Summit Spring(1977) East of Baggs Highway Cowboy Reservoir(1978) Eighty Road(1976) Elk Mountain Elk Head Creek(1977) Fan Rock 113 Fly Creek(1977) Four Mile Creek 1(1977) Four Mile Creek 2(1977) Four Mile Creek 3 (1977) Mule Creek (1980) Sage Creek Slater Park(1977) Twenty-nine Road 1 Twenty-nine Road 2(1977) Northcentral East Cottonwood Gulch(1977) Mud Spring Draw(1978) Pole Gulch(1977) Timberlake 1 29 Timberlake 2 60 Timberlake 3(1977) Timberlake 4 Timberlake 5(1980) West Timberlake 2(1977) Northcentral North Big Hole Butte(1976) 17 Big Hole Gulch 19 Conway Spring(1977) Cox Ranch 20 Dressler Gulch 1(1976) 36 Dressler Gulch 2 Scandinavian Gulch(1977) Seven Mile Res.(1977) Thornburg Gulch(1978) Twenty-one Road West Timberlake 1(1978) 1977 1978 62 40 65 92 101 64 13 19 153 55 42 18 11 20 12 116 8 34 27 40 36 18 136 36 4 62 8 1979 1980 1981 64 112 97 80 77 92 36 45 65 61 44 27 20 18 55 83 54 0 68 22 46 45 36 26 8 12 3 29 21 41 160 15 26 36 81 68 0 22 53 5 24 85 12 6 22 27 48 4 0 153 0 92 0 52 25 36 52 12 95 14 28 0 47 6 18 37 14 12 152 41 46 85 128 37 55 43 21 38 76 87 53 66 20 17 93 99 55 104 108 43 5 47 111 1 77 31 149 38 37 34 28 34 34 3 16 27 37 37 61 42 55 25 19 19 20 25 45 14 42 7 17 23 33 18 68 5 40 26 3 28 26 25 59 0 40 0 6 18 7 27 0 23 20 1 �39 Table 3. Lek (continued) a 1976 Northcentral South Big Gulch 1 Big Gulch·3 Bord Gulch Grassie Reservoir Greasewood Gulch Lay Creek North Fork Big Gulch 2 Sand Creek Spring Creek 1 Spring Creek 2 Spring Creek 3(1977) Upper Nineteen Road South of U.S. 40 Axial Basin Boxelder Gulch 1(1979) Boxelder Gulch 2(1978) Deception Creek Dry Lake 2 Duffy Mountain(1977) Horse Gulch(1977) Juniper 1 Juniper 2(1978) Morgan Gulch 1(1977) Morgan Gulch 2(1977) Morgan Gulch 3(1978) Round Bottom Temple Gulch(1978) Yellow Jacket(1976) 1979 1980 1981 8 42 34 27 125 76 45 46 20 11 50 25 48 27 37 127 35 34 53 0 6 37 22 0 20 41 54 17 28 33 15 9 26 34 44 44 17 56 14 35 45 15 4 43 24 50 32 72 71 62 43 117 70 83 31 50 79 54 72 16 25 108 5 45 8 2 43 78 43 57 38 0 62 9 0 32 45 28 44 27 62 18 2 21 26 12 35 24 26 25 13 29 31 5 16 22 55 12 36 28 24 69 8 32 24 27 17 30 29 68 76 23 35 40 32 2 21 13 39 54 22 28 20 22 15 22 5 53 16 8 20 1977 1978 32 21 30 32 31 23 6 52 34 22 37 22 20 57 37 25 24 31 16 33 24 42 12 30 28 58 60 14 57 22 35 27 4 31 3 32 24 8 39 55 36 18 54 Sunbeam West-North Coffee Pot Spring(1977) Cross Mtn. 1 18 Cross Mtn. 2 Cross Mtn. 3 b East Sand Wash(1978) Lone Tree(1977) 26 Powder Wash Hill Snake River West Thornburg Well(1977) a( ) year of location. b Sand Wash Road abandoned in 1978; East Sand Wash became established in 1978. �Table 4. Summary, peak lek counts of male sage grouse, tfuffat County, Colorado, 1976-81. 1976 Area Blue Mountain 1977 1978 x x N -x 26.0 1 64.8 4 49.3 4 67.0 Cold Spring Mountain N 1980 1979 .- N x N 37.3 8 50.0 6 1 45.5 2 20.5 4 x N 87.0 6 4 6.0 N 1981 x 113.0 1 35.9 10 50.3 4 66.1 11 53.1 10 30.0 12 Northcentral East 44.5 2 28.2 6 55.3 7 78.8 8 49.4 8 38.6 8 Northcentral North 23.0 4 24.7 4 23.5 8 31.7 10 36.9 7 15.6 9 Northcentral south 25.8 8 26.8 9 31.2 9 44.0 11 42.5 10 26.5 10 South of U.S. 40 32.7 6 34.0 11 32.6 14 52.2 13 46.7 12 34.3 14 Sunbeam West-North 22.0 2 25.6 8 28.3 8 36.6 9 27.9 9 20.1 9 31.9 24 33.5 59 37.2 54 53.4 69 42.6 66 29.4 72 East of Baggs Highway All areas ~ 0 �41 Table 5. Colorado, Trends in peak lek 1969-81. counts of male sage grouse, Moffat Number of leks counted Year 29 28 25 27 22 22 17 27 59 54 69 66 72 1969 1970 1971 1972 1973 1974 1975 a 1976 1977 1978 1979 1980 1981 alnitiation of research investigation County, Average number of males per lek 74.6 54.4 52.0 38.4 45.8 41.4 31.7 31.9 33.5 37.2 53.4 42.6 29.4 on sage grouse in Moffat County. Fluctuations in counts of male sage grouse on leks should have some relationship to nesting success and production of young the previous year as measured by percent young in the fall harvest. For example, percent young in the fall harvest was high in 1976 and lek counts of males should have increased in 1977 (actual increase from 31.9 to 33.5 males/lek). Percent young in the fall harvest in 1977 was the lowest in the 1976-80 interval and the number of males on leks should have decreased in 1978. However, the average number of males counted increased in 1978 (37.2 vs. 33.5) over 1977. Examination of all harvest data do not support any direct relationship between nesting success and resulting production and number of males counted on leks the next spring. Data on number of female sage grouse counted on leks were variable, primarily because most leks were counted after the peak of hen attendance. The meager data available indicate marked differences between areas within Moffat County and between years in timing of hen attendance. For example, peak attendance of hens on leks in 1980 on Blue Mountain occurred in early May while attendance of hens occurred in late March in southern Moffat County and early April in the northcentral portion of the county (Table 1). The available data suggest there is no value in hen counts outside of knowing when to expect hatching to occur. Brood Counts Systematic counts of sage grouse along established routes in Moffat County were discontinued after the 1978 field season (Braun and Hoffman 1979). Brood routes were discontinued after analysis of available data revealed no relationship between any statistic (chicks per hen, young per adult, birds per mile) and percent young in the fall harvest (Table 6). However, all distinct broods seen each summer (primarily in July) were tabulated. �42 Brood count data (Table 6) should be used with caution as brood size frequently increased in August. For example, 195 distinct broods averaged 4.8 chicks each in July 1978 while 36 distinct broods in August 1978 averaged 6.8 chicks. The increase in average number of chicks per brood in August was a function of brood shuffling and formation of gang broods. This left some successful hens without any chicks and other hens with more chicks than the average clutch size. During August 1978, 9 distinct broods were seen containing from 9 to 16 chicks. The average clutch size for sage grouse is 7-8 eggs. Table 6. Sage grouse brood data, Moffat No. of broodsa Year 1976 1977 1978 1979 1980 Chicks per hen 17 111 165 231 42 5.3 4.8 4.7 5.1 5.0 aAII broods seen in summer. bVehicle transects County, Colorado, 1976-80. Young per adultb Birds her mile Percent young in fall harvestC 2.4 1.6 1.2 No data No data 1.4 1.3 2.9 No data No data 63.6 43.0 68.7 55.0 56.4 only. cWoarig survey. Capture and Banding Sage grouse trapping and banding in Moffat County was conducted from 13 July through 29 August 1978, 27 March through 30 May and 18 July through 30 August 1979, and 25 March through 2 September 1980. Total sage grouse banded was 506 in 1978, 1,250 in 1979, and 1,068 in 1980 (Tables 7-9). Table 7. Jul Aug Totals Moffat County, Colorado sage grouse bandings, 1978. Chicks Males Yearlings 2+ Chicks 70 147 6 19 8 16 80 124 8 5 3 20 217 25 24 204 13 23 Total for year 506. Females Yearlings 2+ �43 Table 8. Moffat County, Colorado 2+ 0 191 97 13 246 179 1 52 9 2 25 8 288 438 62 35 37 102 4 18 4 58 119 6 23 15 28 139 22 17 177 29 43 139 310 455 177 91 78 Females Yearlings 2+ Mar Apr May Subtotals Subtotals Totals Total for year Table 9. Moffat Mar Apr May Totals Females Yearlings 2+ Males Yearlings sage grouse bandings, 2+ Chicks 1980. 2 99 26 13 324 110 0 27 10 2 32 8 127 447 37 42 98 48 7 0 9 2 4 5 3 119 42 7 18 14 2 17 15 5 153 11 12 168 34 37 153 138 459 168 71 79 Subtotals Subtotals Chicks 1,250. County, Colorado Chicks Jul Aug Sep 1979. Males Yearlings Chicks Jul Aug sage grouse bandings, Total for year 1,068. Trapping effort in spring 1979 and 1980 was relatively uniform (2, 2-person crews for about the same length of time) but more birds were trapped in Spring 1979 than in 1980. This was attributed to less wary birds in 1979 as no trapping had been done in the spring in Moffat County prior to this time. Also, there was a higher spring population in 1979 than in 1980. Summer trapping effort was also essentially equal during each year (1, 2-person crew for about the same length of time) but more birds were captured in 1978 than in 1979 and 1980. This difference was possibly the result of less wary birds in 1978 but more likely was the result of a higher density of birds in areas trapped in 1978 than in 1979 and 1980. �44 Although Moffat County was subdivided into 6 zones for sage grouse banding, trapping pressure and success were not evenly distributed. Chick sage grouse proved especially difficult to locate in sufficient concentrations for effective trapping except on Cold Spring Mountain and along Fortification Creek (Table 10). Trapping pressure and success within each zone were not uniform in spring 1979 and 1980 even though efforts were made to trap all leks (Table 11). These data illustrate the differences between zones and leks in 1979-80. In summer, trapping efforts were concentrated on Cold Spring Mountain, along Fortification Creek and near Stuntz Reservoir on Blue Mountain. All birds banded in spring were captured with long-handled nets at night. In summer, more birds were captured in the morning and evening with cannon nets and walk-in lily pad traps than at night with long-handled nets. For example, in 1978, 61 captures were made in 5 stationary cannon net attempts (~ = 12.2, range 6-20), there were 202 captures in 28 trap days using lily pad traps = 7.2, maximum in 1 trap = 33), 197 captures in 37 attempts with bu;per mounted cannon nets (x = 5.3, range 0-23), while spotlighting at night produced 105 captures in 96.4 hours (! = 1.1 birds/hour). Spotlighting was nonselective for age or sex of grouse captured although when a choice was possible, netters were instructed to select for hens first and then males in the spring and chicks first, then hens, and males last in summer. Cannon nets and lily pad traps were selective for chicks. Trapping was not representative of the sage grouse population during any season. However, it would appear that sex classes of chicks were captured in about the same ratio they occurred in the population (comparison of trapping and harvest data of chicks). ex Sage grouse had not been banded in Hoffat County prior to 1978. In 1979, 7 sage grouse were recaptured that had been banded in 1978. One chick female banded on 1 August 1978 along Fortification Creek was recaptured on 30 April 1979 between Four Mile Creek 1 and 2 leks, northeast of the original banding site. During summer, 6 additional birds banded in 1978 were recaptured (3 banded as chick females, 2 as chick males, 1 yearling male). All were correctly classified to sex when initially banded. Also, all ·6 recaptures of summer banded birds were in the immediate area (Cold Spring Mountain) where they had been initially banded. Recaptures of sage grouse banded in 1978 (1 in spring, 4 in summer) and 1979 (23 in spring, 2 in summer) were much more extensive in 1980. Of interest is the lack of recaptures of birds banded in summer 1979. This suggests that birds banded in summer 1979 survived poorly. One male banded as a chick in summer 1978 along Fortification Creek was recaptured as an adult male on Timberlake 4 lek, northwest of the initial banding location. Of 22 males banded in spring 1979 on leks that were recaptured in spring 1980, 6 changed locations (Four Mile Creek 1 to Four Mile Creek 2 [2 birds], Timberlake 3 to near Timberlake 2, Timberlake 4 to Four Mile Creek 3, Timberlake 4 to Timberlake 2 [2 birds]). The only hen recaptured in 1980 that was banded in 1979 was recaptured on the lek where initially banded. Only 2 of 13 recaptures (all males) of birds banded in 1980 in the same trapping season changed locations (Four Mile Creek 2 to Four Mile Creek 1, Juniper 2 to Four Mile Creek 2). The large distance involved in the 2nd movement, age class of bird (adult), and days involved (18 to 29 April), �Table 10. Sage grouse banded by area, -Moffat County, Colorado, Males Yearlings Chicks 1978-80. Adults Females Yearlings Chicks Adults 1978 1979 1980 1978 1979 1980 1978 1979 1980 1978 1979 1980 1978 1979 1980 1978 1979 1980 0 9 1 0 47 8 0 103 38 0 18 1 0 3 2 0 8 5 198 123 124 22 28 13 19 44 38 191 153 101 11 31 26 19 39 31 19 0 28 0 53 29 4 75 89 13 0 66 2 18 16 4 15 12 Northcentral 0 7 0 3 114 55 1 120 162 0 6 0 0 19 18 0 5 18 South of U.S. 40 0 0 0 0 34 19 0 72 87 0 0 0 0 14 7 0 8 13 Sunbeam West and North 0 0 0 0 35 14 0 39 45 0 0 0 0 8 2 0 2 0 Area Blue Mountain Cold Spring Hountain East of Baggs Highway +:- V1 �46 Table II. Sage grouse bandings Spring 1979-80. by area and lek, Moffat County, Colorado, Males Area and lek Yearlings 1980 1979 Adults 1979 1980 Females Yearlings Adults 1980 1980 1979 1979 Blue Mountain Bear Creek Escalante Haslim Cow Camp Karren Ranch M.C.16,Marker 25.2 State Line 3 17 17 4 1 0 2 3 2 1 0 14 35 26 7 19 11 9 8 6 12 4 0 0 0 0 0 0 0 1 1 1 0 1 1 0 0 1 0 2 1 2 0 0 Cold Spring Mountain Beaver Basin Gee Flats Goodman Draw 8 4 1 2 0 19 12 15 13 1 3 1 0 3 0 1 0 1 1 0 East of Baggs Highway Eighty Road Fan Rock Fly Creek Four Mile Creek 1 Four Mile Creek 2 Four Mile Creek 3 Mule Creek Twenty-nine Road Sunbeam West and North Cross Mountain 1 Cross Mountain 2 Cross Mountain 3 East Sand Wash Lone Tree Powder Wash Snake River West Thornburg Well South of u.S. 40 Axial Basin Boxelder Gulch 1 Boxelder Gulch 2 Deception Creek Dry Lake 2 Duffy Mountain Juniper 1 Juniper 2 Morgan Gulch 2 Morgan Gulch 3 Round Bottom Temple Gulch 8 9 27 9 2 0 4 3 0 1 10 5 5 1 6 34 13 6 22 5 1 7 4 20 0 9 9 0 3 9 1 2 7 3 0 3 1 0 4 0 1 1 3 0 3 3 2 2 0 4 40 2 2 6 11 14 9 2 1 3 5 9 0 0 9 10 12 3 8 8 1 11 25 14 5 8 0 20 12 0 13 6 8 4 5 23 0 10 6 6 18 1 0 4 1 1 0 0 0 0 0 3 1 5 6 0 0 1 3 4 0 1 5 0 3 0 1 0 5 0 0 0 0 1 0 0 1 0 2 1 1 0 2 0 0 0 2 0 2 0 0 0 0 0 0 1 0 1 0 1 0 3 2 2 0 0 0 0 0 0 0 0 0 0 0 1 0 2 0 5 5 0 �47 Table II. (continued) Females Males Area and lek Northcentral Big Hole Butte Bord Gulch Cox Ranch Dressler Gulch 1 Grassie Reservoir Greasewood Gulch Lay Creek Mud Spring North Fork Big Gulch 2 Pole Gulch Scandinavian Gulch Spring Creek 3 Timberlake 1 Timberlake 2 Timberlake 3 Timberlake 4 Timberlake 5 Thornburg Gulch Upper Nineteen Road West Timberlake 2 Yearlings 1979 1980 2 0 4 0 0 3 0 0 3 Adults 1980 1979 4 10 0 6 4 5 9 2 7 1 5 2 2 12 3 1 6 1 2 3 3 5 0 0 0 0 0 5 5 8 2 27 3 3 0 43 9 4 6 1 Yearlings 1980 1979 24 6 3 2 25 10 1 4 6 1 1 19 11 5 37 10 9 7 10 8 0 0 2 0 Adults 1980 1979 0 0 0 0 0 0 1 0 0 9 1 0 0 6 2 0 1 0 0 0 3 0 0 5 0 3 1 1 0 0 0 0 0 0 0 1 0 0 0 3 0 0 1 0 0 0 4 2 2 6 1 2 0 1 0 �48 suggest misreading of the band number of the bird recaptured. The 6 recaptures in summer 1980 of birds banded in 1978 (4; chick female, adult female, 2 chick males) and 1979 (chick female, adult male) were all in the area of initial banding (1 along Fortification Creek, 5 on Cold Spring Mountain). Hunting Season Data Collection The sage grouse season in Moffat County (Units 14, 16, 18, 20, 26) opened on the 2nd Saturday of September each year from 1978 to 1980 (Table 12). Season length in most of Morfat County increased from 7 days in 1977 to 9 in 1978, 16 in 1979 and 25 in 1980. Data were collected each year through use of volunteer wing collections stations, check stations, field checks of hunters and a questionnaire survey. Table 12. Sage grouse hunting seasons and bag and possession Moffat County, Colorado, 1978-80. Year Opening .da t e Closing date 1978 1979 1980 9 Sep 8 Sep 13 Sep 17 Sep 16b,23 Sepc 21 Sepd,7 Octe a In the aggregate with sharp-tailed Length(days) Daily 9 9b,16c 9d,25e 3 3 3 grouse (Pedioecetes limits, a Limits Possession 6 6 6 phasianellus). bU·nlts 14 , 26 . cUnits 16, 18, 20. dUnit 26. eUnits 14, 16, 18, 20. Check Stations Check station operation was identical each year in the 3-year period. Hunter contacts and total number of birds examined was greatest at Cedar Mountain where hunters from the northcentral part of Moffat County were checked. Smaller but significant numbers of hunters were contacted at Dinosaur (Blue Mountain hunters) and Maybell (primarily hunters from Cold Spring Mountain). Number of hunters contacted increased each year; this was most notable at Cedar Mountain. At Dinosaur, number of hunters contacted decreased while at Maybell, they slightly increased. Number of birds harvested and birds per hunter increased in 1979 from 1978 levels and decreased in 1980. Number of sage grouse reported seen decreased from 1978 to 1980 (Table 13). These data indicate that the sage grouse population decreased from 1979 to 1980. It is probable that the increase in hunter numbers was the result of more liberal seasons, greater publicity, word of mouth by hunters of their hunting success, and an increasing human population in the Moffat County area. �49 Table 13. Sage grouse harvest statistics, Moffat 12.7 15.3 12.7 8.1 6.6 7.4 1.4 2.1 1.2 435 357 247 12.5 11.0 17.6 6.9 9.6 6.4 2.2 2.2 1.4 6,707 4,227 3,746 404 442 340 6.0 10.5 9.1 7.6 6.2 6.3 3.6 3.4 2.3 15,295 14,639 10,621 1,490 1,898 1,284 9.7 13.0 12.1 7.6 7.1 7.0 1.9 2.3 1.4 No. birds observed Cedar Mountain 1978 1979 1980 459 534 604 5,114 7,175 5,468 651 1,099 697 Dinosaur 1978 1979 1980 199 162 174 3,474 3,237 1,407 Maybell 1978 1979 1980 112 130 149 770 826 927 Hunter efficiency No. birds harvested Crippling loss a (%) Area and year a 1978-80. Birds per hunter No. hunters checked Combined 1978 1979 1980 County, Colorado, (%) areas Only those statistics collected at check stations. Small game management units in Moffat County were subdivided into harvest zones to examine hunter pressure and harvest by area (Fig. 1). Data available from check stations, while biased because of their location, illustrate that hunter pressure and harvest were not uniformly distributed in Moffat County (Table 14). Table 14. Hunter pressure, harvest, Moffat County, Colorado, 1978-80.a Harvest zone 14A 16A 16B 16C 18A 18B 20A 20B 26A aCheck Total hunters, % 1978 1979 1980 22.7 37.5 2.5 9.0 2.3 26.0 0.6 19.4 46.6 2.3 9.1 2.1 19.8 0.1 station data only. 0.3 34.8 33.8 0.9 8.9 2.8 18.5 and hunter success by harvest Total harvest, % 1978 1979 1980 16.4 28.5 3.0 22.0 0.9 29.2 0.4 21.1 38.2 1.8 17.0 2.5 18.9 0.1 0.5 32.3 24.2 1.9 20.3 1.6 19.2 zone, Birds per hunter 1978 1979 1980 1.4 1.5 2.3 4.8 0.8 2.2 1.4 2.5 1.9 1.8 4.3 2.8 2.2 2.0 2.0 1.3 1.0 3.0 3.1 0.8 1.4 �~ "·z - 50 ~ z :::>0 00 Q<t a: 1-'3 Lt8 LL 0 ~ tCIJ UJ > W 0 �51 These data indicate that hunter pressure was low in zones 14A, 16C, 20B and 26A~ This is probably true for zones 16C, 20B and 26A but not for 14A. Harvest generally paralleled hunter pressure in all zones but 18A (Cold Spring Mountain). Data presented (Table 14) suggest a decline in hunter pressure in harvest zone 20A (Blue Mountain). Total harvest in this zone decreased from 1978 to 1979 but remained similar in 1979 and 1980. It is also apparent that percent of the total harvest almost doubled in harvest zone 16A from 1978 to 1980. Hunter pressure has also increased in this zone, probably because it is readily accessible from Craig, more liberal seasons, an increasing human population in the Moffat County area, word of mouth by successful hunters, and increased publicity. Sage Grouse Hunter Experience Most hunters contacted at check stations each year were asked whether not they normally hunted sage grouse in Moffat County (Table 15). Table 15. Previous sage grouse hunting experience Moffat County, Colorado, 1978-80. Normally hunt in Moffat County N % Year 1978 1979 1980 502 538 679 69.1 70.8 75.9 Normally hunt elsewhere N % 81 52 52 11.1 6.8 6.8 or of sage grouse hunters, First time sage grouse hunter % N 144 170 164 19.8 22.4 18.3 Data for the 3 years indicate that the percent of hunters that normally hunt sage grouse in Moffat County increased from 1978 to 1980 while the percent that normally hunt sage grouse elsewhere decreased. The percent of first-time sage grouse hunters remained relatively constant. It would appear that more liberal sage grouse seasons in Moffat County attracted some additional hunters in 1978 but did not in 1979 and 1980. This was most evident in 1980 when the season was liberalized to 25 days in most of the county. Overall, data in Table 15 are remarkably similar to data collected on sage grouse hunter experience in North Park, Colorado from 1973 to 1980. Hunter Origin Origin of hunters contacted at check stations was ascertained from hunter vehicle license plates each year (Table 16). The percent of the total hunters from Moffat County decreased from 1978 to 1980 indicating that the increase in total hunter numbers (Table 13) was not the result of more hunters from the Moffat County area. Instead, the increase in hunters was probably the result of more hunters from the Denver metropolitan area (from 21.3 to 25.7% of the total from 1978 to 1980). All other points of origin remained relatively stable or decreased (Mesa County) during the 3-year period. These data suggest that the more liberal seasons, increased publicity and words of mouth about hunter success, most affected people living in the Denver metropolitan area. �52 Table 16. Hunter origin, Moffat parties, 1978-80. County, Colorado 1978 County Moffat Mesa Rio Blanco Denver Metro. Area Jefferson Adams Arapahoe Denver Douglas Boulder Garfield Routt All others Totals N sage grouse hunting 1980 1979 % N % N % 129 46 41 (74) 32 18 11 11 2 12 11 6 28 37.2 13.7 11.8 (21.3) 9.2 5.2 3.2 3.2 0.6 3.4 3.2 1.7 8.1 III 45 33 (76) 32 14 11 19 0 13 8 17 55 31.1 12.6 9.2 (21.2) 8.9 3.9 3.1 5.3 0.0 3.6 2.2 4.8 15.4 122 35 49 (103) 34 22 19 25 3 14 11 15 52 30.4 8.7 12.2 (25.7) 8.5 5.5 4.8 6.2 0.8 3.5 2.7 3.7 13.0 347 100.0 358 100.0 401 100.0 Wing Receipts Sage grouse wings were collected at check stations, volunteer wing barrels placed at 16 locations, field checks, mail-in wing envelopes, and brought in to a CDOW office. Data collected in all 3 years were remarkably similar (Table 17) indicating that hunter pressure and success (as a percent of the total) remained constant by area each year. These data indicate that harvest of sage grouse is minimal south of U.S. Highway 40. Eastern Moffat County (including western Routt County) accounts for about 10-11% of the harvest while the Cold Spring Mountain area accounts for 16-20%. The only. noteworthy change during the 3-year period was the marked decrease (22.0 to 12.5%) in percent of total wings received from the Blue Mountain area. Data available on origin of sage grouse wings do not suggest there should be separate regulations for areas east of the Baggs Highway or south of U.S. Highway 40. It is also doubtful that Unit 26 should have differing regulations from other areas within Moffat County. Time of Harvest All wings received were identified to harvest period (Table 18). Number of wings received varied from 2,326 (9 day season), to 3,424 (16 day season) and 2,769 (25 day season). Thus more liberal seasons did not necessarily result in more sage grouse being harvested. Instead, number of sage grouse harvested reflected environmental conditions prior to and during the hunting season, and the size of the population available. It does appear that a 25 day season will increase the total harvest over a 16 day season by about 5-6%. All data available indicate that hunters respond to longer seasons by spreading their effort over a longer period. This does not necessarily result in a higher harvest rate but it should increase total recreational days. �53 Table 17. Origin of sage grouse wings, Moffat County, Number Area 1978 South of U.S. 40 Axial Deception Creek Juniper Springs Percent of total 17 20 20 Dinosaur (Blue Mountain) Check Station Wing barrel Wolf Creek Road Percent of total Northcentral Cedar Mountain Check Station Cedar Mountain wing barrel Lay Creek Maybell City Park M.C. Road 3 M.C. Road 4 Miscellaneous North Fork Big Gulch Percent of total a Includes western 17 36 20 2.6 34 9 29 41 1 9 44 58 198 88 10.9 93 136 16 10.7 16.3 378 168 ll2 19.2 330 144 78 19.9 417 93 21 22.0 323 ll5 15 13.2 236 83 27 12.5 558 160 84 54 134 83 98 989 297 57 130 143 ll7 96 664 322 105 197 98 95 7 II o 48.7 53.8 54.3 99 69 10.6 Maybell (Cold Spring Mountain) Check Station Wing barrel Cold Spring Mtn. Field Checks Percent of total Routt County. of wings received 1979 1980 25 47 27 2.9 2.4 East of Baggs Highwaya California .Park Road Elkhead Road M. C. Road 2 M.C. Road 101 Miscellaneous Percent of total Colorado, 1978-80. 337 58 22 �54 Table 18. Time distribution Colorado, 1978-80. of sage grouse wings received, % N First weekend First week Second weekend Second week Third weekend Third week Final 4 days Totals 1,745 224 357 Season Season Season Season 2,326 Age and Sex Structure % N 75.0 9.6 15.4 closed closed closed closed 100.0 County, 1980 1979 1978 Period Moffat % N 2,291 66.9 9.1 313 462 13.5 2.2 75 8.3 283 Season closed Season closed 100.0 3,424 1,786 271 274 63 221 23 131 2,769 64.5 9.8 9.9 2.3 8.0 0.8 4.7 100.0 of the Harvest Most wings received each year were classified to age and sex. Data for 1978-80 were included with those from 1976-77, the only other years available (Table 19). The data for 1978-80 were further subdivided by area (Table 20). Data presented in Table 20 indicate that marked differences occurred in production of young and survival of young to the yearling age class among areas and among years. Overall, production of young decreased in 1979 and 1980 from 1978. Production of young was poor in 1977 which resulted in a low percent of yearlings in the 1978 population. Percent yearlings increased in 1979 (good production in 1978) and decreased in 1980. It is expected that percent yearlings in the 1981 harvest will be similar to that recorded in 1980. Obviously, this age classification is a good indicator of the previous year's production. These data also indicate that the 1981 spring sage grouse population should have leveled off or decreased. This actually occurred, although the decrease was greater than expected (Tables 2, 3, 4). Table 20. Age structure Colorado, 1978-80. a of the sage grouse harvest Area 1978 b Moffat County East Northcentral Cold Spring Mountain Blue Mountain Moffat County South All areas 77 .8 71.8 62.3 62.1 70.2 68.7 a b Immatures 1979 1980 57.5 57.2 54.1 45.0 56.5 55.0 50.8 52.9 61.4 68.8 54.8 56.4 1978 by area, Moffat Yearlings 1979 1980 8.7 5.0 16.7 17.0 5.3 9.9 27.8 28.0 21.7 31.6 25.3 27.2 15.6 16.8 12.3 11. 0 9.6 14.9 1978 Adults 1979 1980 13 .5 23.2 21. 0 20.9 24.5 21.4 14.7 14.8 24.2 23.4 18.2 17 .8 33.6 30.3 26.3 20.2 35.6 28.7 In percent. Includes western 1980=58 wings. Routt County, 1978=45 wings, County, 1979=104 wings, �Table 19. Age and sex composition of the sage grouse harvest, Moffat and western Routt counties, Colorado, 1976-80. Males Immatures Females Totals Males Year N 1976 203 42.6 273 57.4 476 63.6 41 1977 121 39.3 187 60.7 308 43.0 1978 825 49.6 837 50.4 1,662 1979 871 46.3 1,011 53.7 1980 705 45.1 857 54.9 5-year avg. % 46.3 N % 53.7 % % Yearlings Females Totals N % N % 35.7 74 64.3 ll5 15.4 77 45.8 91 54.2 168 68.7 77 32.1 163 67.9 1,882 55.0 386 41.5 545 1,562 56.4 154 37.4 258 N 58.5 N 39.4 Adults Females Males % N 25 15.9 l32 23.4 89 36.9 240 9.9 141 58.5 931 27.2 62.2 412 14.9 60.6 18.5 N Totals Sample size N % 84.1 157 21.0 748 152 63.1 241 33.6 717 27.3 375 72.7 516 21.4 2,418 183 30.0 428 70.0 6ll 17 .8 3,424 289 36.4 506 63.6 795 28.7 - 31.3 % 68.7 23.0 2,769 VI VI �56 Composite data by age and sex for the entire Moffat County area for 1976-80 (Table 19) document that: 1. The sex ratio at hatching approximates 1:1 with some differential survival favoring females occurring before chicks are 3-4 months of age. 2. Differential survival favoring females occurs in the older age classes and is most pronounced in adults (68.7:31.3). 3. There are 2 hens to every male in the spring breeding population (5 year average = 65.1% females:34.9% males of all adults and yearlings). 4. Production of young was excellent and 1980 and poor in 1977. 5. Recruitment of yearlings was poor in 1978 following in 1977 and good in 1977 and 1979. 6. Recruitment (overwinter survival) was more than sufficient to maintain population stability in 1977 and 1979 but was not in 1978, 1976, and 1980. 7. The population should have increased decreased in 1976, 1978 and 1980. Survival and Mortality in 1978 and 1976, average in 1979 the poor production in spring 1977 and 1979, and Rates Estimated turnover of adult male and female sage grouse in Moffat County was estimated from the age composition of the adult and yearling components of the harvest each year (Tables 21, 22). Since a sage grouse population can only increase, decrease, or remain stable, the percent yearlings of the combined yearling and adult segment of the harvest should provide an indication of the annual turnover rate of adult sage grouse in the population. Table 21. Estimated annual County, Colorado, 1976-80.a turnover of adult male sage grouse, Moffat Percent in harvest Yearlings Year Adults 1976 1977 1978 1979 1980 37.9 53.6 64.7 32.2 65.2 62.1 46.4 35.3 67.8 34.8 5-year average 49.7 50.3 Estimated annual mortality for adult males in a stable population aDate from wing collections only. N 66 166 218 569 443 50.3%. �57 Table 22. Estimated annual turnover of adult female sage grouse, Moffat County, Colorado, 1976-80. Percent in harvest Yearlings Year Ada-lts 1976 1977 1978 1979 1980 64.1 62.6 69.7 44.0 66.2 35.9 37.4 30.3 56.0 33.8 5-year average 58.5 41.5 Estimated a annual mortality for adult females in a stable population N 206 243 538 973 764 41. 5%. . Data from wing collections only. Percent yearlings of the adult and yearling cohorts in the fall harvest varied annually for both males and females and averaged 50.3 and 41.5%, respectively. Thus, annual mortality in a stable population would be 50.3% for adult males and 41.5% for adult females. Data from 1980 (and 1978) suggest a decreasing population as apparent mortality rates should be higher in increasing populations. This is because these mortality estimates are based on the percent yearlings in the fall population. If the percent yearlings are low, the population is decreasing because of low production of young the previous year and/or poor overwinter survival of young. Data from 1976 and 1979 indicate that the male population increased in those years. The same is true for females in 1979. Data for females in 1976 and 1977 are less indicative. The 5-year average mortality estimate of 50.3% for males appears reasonable while the 5-year average of 41.5% for females appears somewhat high. Nesting Success and Production Each year, primary feather molts of adult and yearling hens were recorded and compared to primary feather molt patterns of males of the same age class harvested in the same time interval (Tables 23, 24). Estimated nesting success varied by area within Moffat County with Cold Spring Mountain having the highest success each year (Table 23). The data also indicate that nesting success of yearling females was consistently less than adult females. Less than one-half of the yearling hens were normally successful whereas at least one-half of the adult hens are successful nesters in most years (Tables 23, 24). Overall, nesting success in 1980 was lower than in 1979 although the chicks per hen computations were similar (1.9 vs. 2.0). These data support the conclusion that overall production in 1980 was only average. It would appear that percent yearlings in the 1981 spring population (and fall harvest) will be similar to or less than that measured in 1980. Obviously the Moffat County sage grouse population is not increasing at the present. The available data strongly document the relationships of nesting success ��59 and production of young to the size of the spring population in the succeeding year and the fall harvest in that year. The factors of nesting success and production of young are most important in determining population size. Outside of control of grazing and prevention of habitat loss, there is little that can be done over large areas to positively impact sage grouse populations without expensive programs to increase habitat quality and quantity. Table 24. Estimated sage grouse nesting County, Colorado, 1976-80. Year No. hens examined 1976 1977 1978 1979 1980 206 243 538 973 764 Hatching Dates success and production, Percent successful Yearlings Adults All hens 35.1 14.9 50.3 46.1 33.7 55.7 39.6 65.3 58.4 49.4 48.3 30.0 60.8 51.5 44.1 Chicks/hen 2.3 1.2 3.1 1.9 2~O Moffat Percent chicks in harvest 63.6 43.0 68.7 55.0 56.4 Ages and hatching dates were calculated for 1,662 chick sage grouse in 1978 (Table 25), 1,882 in 1979 (Table 26), and 1,557 in 1980 (Table 27). Hatching was earliest in 1978 and latest in 1980. Hatching started as early as the week of 4-10 May and was complete by the week of "20-26 July. The hatching peak occurred between 1 and 28 June, varying with year. Chicks hatching after 1 July were probably the result of renesting. In 1978 and 1979, renesting accounted for 10-13% of all chicks produced while in 1980, as many as 17-18% of the chicks resulted from renesting. It is apparent that climatic variables in late March and early April greatly affect initiation of breeding and nesting events. For example, 1980 initially appeared to be an early nesting year in all areas within Moffat County. However, frequent rain and snowstorms in April appear to have been major contributors to the lateness of the hatch and the lower nesting success in 1980. Ovarian Analysis A sample of ovaries was collected from hunter harvested adult and yearling female sage grouse each year and analyzed following procedures described by Meyer et al. (1947), Kabat et al. (1948) and Buss et al. (1951). Additional ovaries were classified in situ at check stations by C. E. Braun and K. M. Giesen. All data were pooled (Table 28). Only 4 (2 yearlings, 2 adults) ovaries were classified as not ovulating during the 3 years. The available data indicate that an exceedingly high percentage of female sage grouse ovulate and probably lay eggs each year with no differences (f > 0.05) between adults and yearlings. �Table 25. Estimated sage grouse hatching dates, Moffat County, Colorado 1978. East N.C. 4-10 May Cold Spring Males Blue Mtn. South 2.1 Totals East N.C. Females Cold Blue Spring Mtn. 0.4 0.9 6.2 0.8 2.8 2.5 12.5 5.1 11.1 9.2 3.8 1.0 0.7 0.7 18-24 May 7.1 7.4 2.1 25-31 May 7.1 17.4 4.4 27.1 12.5 10.7 23.2 24.3 6.7 1- 7 Jun 33.3 34.5 29.8 28.3 31.2 32.0 18.5 24.8 8-14 Jun 16.2 16.6 18.4 15.7 25.0 19.4 21.3 15-21 Jun 18.1 8.9 9.9 14.5 6.3 11.5 22-28 Jun 5.1 7.3 6.4 9.6 6.3 11.1 4.5 16.3 3.0 6-12 Jul 0.0 2.0 13-19 Jul 1.0 0.5 20-26 Jul Sample sizes a In percent. 8.3 1.9 20.8 7.3 6.7 16.7 18.3 23.8 22.0 33.3 23.5 23.9 26.7 25.6 8.3 23.8 12.0 5.3 3.8 18.9 0.0 8.5 8.2 4.6 4.8 7.6 12.2 4.2 6.6 8.2 1.9 3.4 12.4 8.5 4.2 5.4 8.5 3.0 2.8 1.3 12.4 6.1 .4.2 3.9 1.4 0.6 0.9 0.5 2.8 .1.8 0.2 99 403 Totals 0.1 11-17 May 29 Jun-5 Jul South 0.7 0.1 141 166 16 825 108 436 105 164 24 837 (J'I 0 �Table 26. Estimated sage grouse hatching dates, Moffat County, Colorado, 1979.a .East 11-17 May N.C. Cold Spring Males Blue Mtn . South 0.9 Totals East N.C. Females Cold Blue Spring Mtn. South Totals 0.1 18-24 May 25-31 May 2.7 1.7 3.0 1- 7 Jun 15.2 12.2 6.7 8-14 Jun 27.8 24.1 0.3 1.6 0.5 4.4 2.0 3.9 3.1 2.6 6.1 2.9 4.1 21.7 10.9 11.8 6.6 7.3 9.1 6.6 12.7 16.5 26.0 21.6 39.2 36.0 17.3 12.1 33.3 30.1 28.5 24.2 18.6 43.5 26.4 23.5 24.4 25.7 32.7 30.3 25.6 16.1 23.8 33.3 38.1 25.6 9.8 19.4 23.0 32.7 12.1 20.3 29 Jun-5Jul 8.8 7.6 14.0 18.6 10.1 11.8 8.0 15.7 17.8 6.1 10.8 6-12 Jul 4.4 2.1 6.1 4.1 1.7 6.3 3.7 3.0 2.7 13-19 Jul 0.3 0.5 0.9 20-26 Jul 0.2 15-21 Jun 22-28 Jun Sample sizes a 112 In percent. 474 165 97 4.4 3.3 23 871 102 578 0.4 0.1 191 107 33 1,011 0\ •.... �Table 27. Estimated sage grouse hatching dates, Moffat County, Colorado, 1980.a Dates East N.C. Cold Spring 18-24 May 0.6 25-31 May 1.4 0.6 Males Blue Mtn. South Totals 0.1 10.0 1.1 Females Cold Blue Spring Mtn. East N.C. South Totals 2.5 0.2 0.6 0.7 2.3 1.1 2.2 10.0 1.8 0.5 1- 7 Jun 7.2 8.5 6.2 1.0 25.0 7.2 6.2 5.7 2.8 15.5 35.0 5.3 8-14 Jun 37.7 29.0 12.3 13.9 20.0 23.6 30.8 25.5 6.8 32.6 15.0 20.2 15-21 Jun 23.2 24.7 24.7 36,6 20.0 26.1 32.1 32.5 29.9 30.4 20.0 31.6 22-28 Jun 23.2 20.7 25.3 28.7 25.0 23.3 18.5 19.6 28.8 10.4 15.0 23.0 7.2 11.4 21.6 14.8 13.5 3.7 9.5 11.3 5.2 5.0 9.4 1.5 2.8 7.4 4.0 3.8 6.2 3.6 11.9 1.5 5.7 13-19 Jul 0.6 1.9 1.0 0.9 0.9 5.1 1.5 1.8 20-26 Jul 0.3 0.2 0.2 1.7 29 Jun-5 Jul 6-12 Jul Sample sizes a In percent. 69 352 162 101 20 704 81 440 177 0.7 135 20 853 a- N �63 Table 28. Age class Yearlings Adults Totals Percent Sage grouse ovarian analysis, Moffat County, Colorado, No. ovaries examined 1978 1979 1980 55 85 141 140 88 228 74 127 201 1978 1978-80. Percent ovulatory 1979 1980 98.2 98.8 100.0 100.0 98.6 99.2 98.6 100.0 99.0 Hunter Questionnaire All sage grouse hunters in Moffat County were required to obtain a free permit each year during the 1978-80 interval. This provided names and addresses for a questionnaire survey of each hunter following the hunting season. One followup questionnaire was sent to all nonrespondents to the initial survey letter after about 3-4 weeks. Averages calculated for respondents to the followup questionnaire were used to project for those permittees not responding to either survey. Data for 1980 are presented in Table 29 while a summary of all 3 years is presented in Table 30. Response to the survey varied from 76 to 78%. Number of permittees increased each year from 2,006 in 1978 to 3,477 in 1980. Based on hunters contacted at check stations only 8.4% did not have permits in 1978 (B = 770 hunters contacted), 7.1% (B = 826) did not have permits in 1979, while 8.4% = 927) did not have permits in 1980. Thus the percent noncompliance before hunting remained constant over the 3 years. Each year, only 73-76% of the total permittees hunted sage grouse in Moffat County (Table 30). Hunter success was variable (66 to 81%) but total hunter days increased markedly from 3,161 in 1978 to 5,348 in 1980. Total harvest including birds reported crippled and lost ~creased from 5,185 in 1978 to 7,840 in 1979 and decreased to 6,680 in 1980. Thus liberalized seasons resulted in steady increases in hunters afield and hunter days but did not result in yearly increases in birds harvested. The number of birds harvested was a function of the number of birds in the fall population and climatic variables immediately prior to and during the hunting season. (B Addresses of all permittees were tabulated by county of residence each year (Table 31). These data document a decrease in the percent of the total permittees that originated in Moffat County and an increase in the percent originating in the Denver metropolitan area. These were the only notable changes in the 3-year period. Overall, the percent of hunters originating from the Eastern Slope increased from 1978 to 1980 while the percent originating from the Western Slope decreased (Table 31). These data indicate that more liberal seasons attracted more Denver area hunters to Moffat County. This is surprising considering increasing costs of gasoline. �64 Table 29. Moffat data, 1980. County, Colorado Nl Number in sample Percent of total permittees Number Percent of hunters hunters Number of nonhunters Percent nonhunters Number hunters successful 2,096 60.3 1,609 76.9 487 23.2 1,112 sage grouse hunter questionnaire N2 601 17.3 403 67.1 198 32.9 245 Projected for z 2,697 77.6 2,012 74.6 685 25.4 1,357 780 22.4 523 67.1 257 32.9 318 Projected for 3,447 100.0 2,535 72.9 942 27.1 1,675 Percent hunters successful 69.1 60.8 67.4 60.8 66.1 Percent permittees successful 53.1 40.8 50.3 40.8 48.2 Number of hunter days Days per hunter Number of grouse bagged 3,482 2.2 4,193 820 2.0 901 4,302 2.1 5,094 1,046 2.0 1,151 5,348 2.1 6,245 Grouse per hunter 2.6 2.2 2.5 2.2 2.5 Grouse per successful hunter 3.8 3.7 3.8 3.7 3.7 Grouse per permittee 2.0 1.5 1.9 1.5 1.8 Number of birds lost Birds lost per hunter Total harvest Percent crippling loss 330 0.2 4,523 7.3 53 0.1 954 5.6 383 0.2 5,477 7.0 52 0.1 1,203 5.6 435 0.2 6,680 6.5 �65 Table 30. Moffat Total permits Percent County, Colorado, issued Number hunters Percent hunters successful 2,006 2,673 3,477 1,128 75.9 2,041 1,653 Number of grouse bagged Grouse per hunter Grouse per successful hunter Number of birds lost Birds lost per hunter 4,697 4,448 crippling loss Season length Bag and possession Percent chicks Percent yearlings limit in harvest in harvest 66.1 5,348 2.2 7,260 2.1 6,245 3.1 3.6 2.5 4.2 4.4 3.7 488 580 0.3 5,185 Total harvest 72.9 1,675 81.0 2.1 Days per hunter 77 .6 2,535 76.4 75.1 3,161 Number of hunter days Percent 1980 74.9 successful 1978-80. 1979 1,502 hunters data, 1978 78.1 response Total hunters Percent sage grouse questionnaire 435 0.3 7,840 9.4 0.2 6,680 7.4 a 9 16 3/6 3/6 6.5 25b 3/6 68.7 55.0 56.4 9.9 27.2 14.9 aUnits 16, 18 and 20; 9 days in Units 14 and 26. bU . nl.ts 14, 16, 18 and 20; 9 days in Unit 26. �66 Table 31. County a Origin of Moffat or area Moffat Jefferson Rio Blanco Mesa Denver Adams Boulder Routt Garfield Larimer El Paso Out of state b Denver metro area West Slope East Slope Total permits issued County, Colorado sage grouse permittees, Number of permits 1978 1979 issued 1980 Percent 1978 883 142 191 179 l39 32 59 88 45 26 18 45 395 1,434 527 2,006 1,024 467 238 320 257 201 138 134 133 78 65 79 957 2,053 1,342 3,474 44.0 7.1 9.5 8.9 6.9 1.6 2.9 4.4 2.2 1.3 0.9 2.2 19.7 71.5 26.3 919 253 220 217 169 141 112 135 78 68 53 66 642 1,666 942 2,674 1978-80. of total permits 1980 1979 34.4 9.5 8.2 8.1 6.3 5.3 4.2 5.0 2.9 2.5 2.0 2.5 24.0 62.3 35.2 29.5 13.4 6.9 9.2 7.4 5.8 4.0 3.9 3.8 2.2 1.9 2.3 27.5 59.1 38.6 aOnly those counties with about 2.0% or more of the total permittees. b Adams, Arapahoe, Denver, Douglas and Jefferson counties. Time period of hunting was available for 1,586 hunts that resulted in 3,891 sage grouse being harvested in 1978; 1,691 hunts and 4,875 grouse bagged in 1979; and 2,228 hunts and 4,992 grouse bagged in 1980 (Table 32). These data indicate that as season length increased, hunter pressure and total harvest decreased on the opening weekend of the season. This may be somewhat misleading as the total number of hunters actually increased in the 3-year period and there may have been no actual decrease in total hunters or harvest the opening weekend. However, it is quite evident that sage grouse hunters responded to liberalized seasons by extending their hunting effort with almost 10% of the total effort and 7%+ of the total harvest coming after the 3rd weekend in 1980. Number of hunters reporting achieving bag limits for 1 or more days varied each year (Table 33). Hunter success was highest in 1979 and lowest in 1980. Generally, less than 40% of the hunters responding reported achieving the bag limit of 3 birds per day. Thus, a reduction in bag limit from 3 to 2 birds per day would increase the number of hunters achieving the bag limit but would not measurably decrease the total number of birds harvested. �67 Table 32. Time period of hunting effort and harvest, Colorado, 1978-80. Time interval Percent 1978 First weekend First week Second we ekend Second week Third weekend Third week Fourth weekend Last 2 days 61.3 20.6 18.0 Closed Closed Closed Closed Closed total hunting trips 1980 1979 47.3 15.4 15.8 6.5 15.0 Closed Closed Closed Table 33. Number of hunters achieving County, Colorado 1978-80. Number of days limit achieved 1 2 3 4 5 6 7 8 9 1978 44.2 13.5 12.1 7.4 14.5 3.5 3.1 1.7 Hoffat County, Percent 1978 66.1 19.0 14.8 Closed Closed Closed Closed Closed 56.5 13.0 14.0 5.2 11.2 Closed Closed Closed 52.2 13.6 10.3 6.0 10.5 2.9 3.1 1.4 a bag limit of sage grouse, Moffat Number of hunters obtaining 1979 220 219 21 10 2 2 total harvest 1980 1979 229 331 27 16 2 3 1 bag limit 1980 208 172 10 12 1 2 1 The questionnaire was designed to examine harvest and hunter activity by Small game units and harvest zones (Fig. 1). Data for 1980 are presented in Table 34 while data for all 3 years are presented in Table 35. These data indicate that hunter pressure and harvest were not uniform by Small game unit or harvest zone. Unit 16 had the highest hunter pressure and harvest each year. This is the northcentral portion of the county centered around Great Divide. Unit 18, especially Cold Spring Mountain (18A), had from 16 to 20% of the hunters and 20-22% of the total harvest. The 3rd leading unit was 20, primarily the Blue Mountain area (20A). Unit 14 consistently had about 10-12% of the total hunters and 10-12% of the total harvest. Hunting pressure and harvest was neglible in Unit 26 (A), and harvest zone 20B and was low in harvest zone 16C. It is of interest that the data on hunter pressure and harvest were remarkably similar for each unit and harvest zone each year. �68 Table 34. Hunter activity and harvest harvest zones, Moffat County, Colorado by small game management 1980. units and Percent of total Unit (Harvest zone) No. of hunters Percent of total 14 (A) 16 (A) 251 415 576 180 5 255 183 21 4 152 27 35 47 a 2,151 11.7 19.3 26.8 8.4 0.2 603 863 1,338 384 n .s 8.5 1.0 0.1 7.1 1.3 1.6 . 2.2 828 338 41 4 468 78 60 78 ll.8 16.9 26.3 7.5 0.2 16.4 6.6 0.8 0.1 9.2 1.5 1.2 1.5 100.0 5,094 100.0 .(B) (C) (?) 18 (A) (B) (C) (1) 20 (A) (B) 26 (A) Unknown Totals a No. of birds harvested II Total does not equal 2,012 as some hunters hunted in more than 1 zone. Table 35. Hunter activity and harvest by small game management harvest zones, Moffat County, Colorado 1978-80. Unit (Harvest zone) 14 (A) 16 (A) (B) (C) Percent 1978 10.8 17.7 27.8 7.5 (?) 18 (A) (B) (C) of total hunters 1979 1980 10.9 18.7 26.5 8.7 0.4 9.9 6.2 1.0 n .s 11. 9 1.2 2.6 3.4 7.3 0.6 1.9 3.5 8.3 1.4 (?) 20 (A) (B) 26 (A) Unknown 11.7 19.3 26.8 8.4 0.2 ll.8 8.5 1.0 0.1 7.1 1.3 1.6 2.2 Percent 1978 9.8 15.5 27.0 5.4 units and of total harvest 1979 1980 13.3 5.9 1.4 10.1 17.3 27.8 6.6 0.4 14.3 6.5 1.2 15.5 1.3 2.1 2.8 10.5 0.5 1.1 3.7 n .s 16.9 26.3 7.5 0.2 16.4 6.6 0.8 0.1 9.2 1.5 1.2 1.5 Vulnerability During the 3-year study period 2,824 sage grouse were banded in Moffat County. This included 509 chick males, 476 yearling males, 934 adult males, 550 chick females, 177 yearling females and 178 adult hens. Through 1980, 316 recoveries from all sources had been reported (11.2%) (Table 36). �69 Table 36. 1978-80. Year Sage grouse banding and recovery No. banded 1978 data,a Moffat Number recovered, 1979 County, Color ada all sources 1980 Males - chicks 1978 1979 1980 20 217 139 153 8 17 3 1 30 1 18 0 9 8 1 28 0 7 39 8 17 6 1 27 3 6 0 6 6 - Yearlings 1978 1979 1980 25 313 138 4 - Adults 1978 1979 1980 24 451 459 5 Females 1978 1979 1980 204 178 168 - Chicks 16 - Yearlings 1978 1979 1980 13 92 71 1 - Adults 1978 1979 1980 23 76 79 2 1 5 a Two unknown age sage grouse (1 male and 1 female) banded recovered in 1979 are not included. 1 3 6 in 1978 and Direct or first year recovery rates were calculated for each age and sex class for each year using all recoveries (Table 37). Most of the recoveries were from hunting although each year a few recoveries were reported from birds found dead as a result of predation, hitting power lines or from collisions with vehicles. Direct recovery rates for all birds banded were 9.5% in 1978, 7.0% in 1979, and 10.9% in 1980. Direct recovery rates for females were lower (range 6.5-10.9%) than those for males (range 7.912.3%). This indicates that there is no hunter selection for hens (i.e. smaller birds) and if selection occurs, it is for males. The slight difference in vulnerability between males and females may be a function of the larger size (i.e., bigger targets) and apparent slower flight speed of males. �70 Table 37. 1978-80.a Sage grouse banding and recovery data, Moffat County, 1979 1980 Colorado 1978 Direct recovery N b banded rate N banded N recovered Direct recovery rateb Females Chicks Yearlings Adults Subtotals 168 71 79 318 27 6 6 39 16.1 8.5 7.5 12.3 178 93 76 347 9.6 7.5 6.6 8.4 204 13 23 240 7.8 7.7 8.7 7.9 Males Chicks Yearlings Adults Subtotals 153 138 459 750 30 8 39 77 19.6 5.8 8.5 10.3 139 313 451 903 ll.5 5.4 5.8 6.5 217 25 24 266 9.2 16.0 20.8 10.9 1,068 116 10.9 1,250 7.0 506 9.5 Age and sex class Totals aAll b N banded Direct recovery rateb recoveries .. In percent. Direct recovery rates of chicks of both sexes were higher than for yearlings or adults (Table 37). Thus chicks are more vulnerable to hunting than are adults and yearlings. These differences were most pronounced in 1980. Reasons for the marked increase in direct recovery rates of chick sage grouse from 1978 to 1980 are not known. However, most chicks were banded on Cold Spring Mountain where hunter success was high during the 3 years studied. Recoveries of sage grouse were examined by zone of banding for each age and sex class (Tables 38, 39). Bandings of chicks were inadequate, except at Cold Spring Mountain, for detailed analyses. Based on the limited data available, chicks appear to be more vulnerable than adults no matter where they were banded. However, the apparent differences could be a function of small sample sizes. Adequate samples of yearling and adult males were banded in all zones except Cold Spring Mountain in 1979 and 1980. The data indicate a low harvest rate of adult and yearling males (average for all areas = 5.8, 5.8 for yearlings and 6.2 and 8.5 for adults in 1979 and 1980, respectively). Direct recovery rates were lowest south of U.S. Highway 40. No apparent pattern for adult and yearling males is evident in the data available. Too few adult and yearling hens were banded each year for meaningful analysis. The available data do indicate a low direct recovery rate of hens when the data for all areas are pooled. �71 Table 38. Direct recovery County, Colorado 1978-80. Area Cold Spring Mountain rates of banded male sage grouse, Moffat 1978 11979 1980 1978 1+ 1979 1980 1978 2+ 1979 1980 10.6 12.2 17.7 13.6 7.1 15.4 21.1 13.6 18.4 2.9 0.0 1.4 2.3 5.7 13.8 0.0 1.3 7.9 7.0 1.8 100.0 8.3 8.0 6.4 12.5 7.8 15.8 2.9 0.0 7.7 8.9 5.8 5.8 6.2 8.5 South of U.S. 40 East of Baggs Highway 0.0 25.0 Northcentral 14.3 Blue Mountain 11.1 33.3 100.0 Sunbeam West and North All areas 9.7 12.2 Table 39. Direct recovery Colorado, 1978-80. Area Cold Spring Mountain 19.6 16.0 rates of banded female sage grouse, Moffat 11979 1980 1978 1+ 1979 1980 1978 2+ 1979 1980 7.4 8.4 15.8 8.3 9.7 3.8 5.3 7.7 12.9 7.1 0.0 0.0 0.0 5.6 6.3 0.0 8.3 10.5 16.7 0.0 5.6 0.0 50.0 25.0 0.0 0.0 0.0 0.0 6.5 8.5 14.3 16.7 Northcentral 16.7 Blue Mountain 11.1 0.0 0.0 Sunbeam West and North All areas County, 1978 South of U.S. 40 East of Baggs Highway 20.8 7.8 9.6 16. 1 7.7 25.0 8.7 6.5 7.6 The marked increase in direct recovery rates of chicks merits more attention. Presently, it does not appear the longer sage grouse seasons have affected harvest rates of all sage grouse, especially those rates for adult and yearling males and females. The overall harvest rates of 7 to 11% are well below the 30% level above which hunting may have an impact on sage grouse population levels. �72 LITERATURE CITED Beck, T. D. I., and C. E. Braun. 1981. The strutting ground count: variation, traditionalism, management needs. Proc. West. Assoc. Fish and Wildl. Agencies 60:558-566. , R. B. Gill, and C. E. Braun. 1975. Sex and age determination of sage grouse from wing characteristics. Game Inf. Leaflet 49 (revised). Colo. Div. Wildl. 4pp. ---- Braun, C. E., and T. D. I. Beck. 1976. Effects of sagebrush control on distribution and abundance of sage grouse. Colo. Div. Wildl. Final Rep., Fed. Aid Proj. W-37-R-29, Work Plan 3, Job 8a. pp. 21-84. , and D. M. Hoffman. 1979. Vulnerability and population characteristics of sage grouse in Moffat County. Colo. Div. Wildl. Job Progress Rep., Fed. Aid Proj. W-37-R-32, Work Plan 3, Job 11. pp. 163-199. ---- ___ ~' and 1980. Vulnerability and population characteristics of sage grouse in Moffat County. Colo. Div. Wildl., Job Progress Rep., Fed. Aid Proj. W-37-R-33, Work Plan 3, Job 11. pp. 205-242. Buss, I. 0., R. K. Meyer, and C. Kabat. 1951. Wisconsin pheasant reproduction studies based on ovulated follicle technique. J. Wildl. Manage. 15:32-46. Eng, R. L. 1955. A method for obtaining sage grouse age and sex ratios from wings. J. Wildl. Manage. 19:267-272. Gullion, G. \.J'. 1965. Improvements in methods for trapping and marking ruffed grouse. J. Wildl. Manage. 29:109-116. Hoffman, D. M. 1979. Investigations of the distribution and status of sagebrush and sage grouse in the Moffat County area. Colorado Div. Wildlife, Final Rep., Fed. Aid Proj. W-37-R-32, Work Plan 3, Job 10. pp. 95-162. Hoffman, R. W., and C. E. Braun. 1975. A volunteer wing collection tion. Game Inf. Leaflet 101. Colo. Div. Wildl. 3pp. sta- Kabat, C., I. O. Buss, and R. K. Meyer. 1948. The use of ovulated follicles in determining eggs laid by the ring-necked pheasant. J. Wildl. Manage. 12:399-416. Lacher, J. R., and D. D. Lacher. Wildl. Manage. 28:595-597. 1964. A mobile cannot net trap. J. �73 Meyer, R. K., C. Kabat, and I. O. Buss. 1947. Early involutionary changes in the post-ovulatory follicles in the ring-necked pheasant. J. Wildl. Manage. 11:43-49. Pyrah, D. G. 1963. Sage grouse investigations. Idaho Fish and Game Dep., Wildl. Rest. Div., Job Compl. Rep., Fed. Aid Proj. W-12S-R. 71pp. Rogers, G. E. 1964. Sage grouse investigations in Colorado. Dep. Game, Fish and Parks, Tech. Publ. 16. 132pp. Prepared by __~~~~~··~~~.~.~.,r~t~a~,~~~'i_'/ _ Clait E. Braun (i8j Wildlife Research Leader Colo. ��75 April JOB PROGRESS State of Colorado Project No. W-37-R _______ Work Plan No. 3 ------- Job No. Job Title: Potential Covered: Personnel: REPORT ..Game Bird Survey 12 ------- Impacts of Strip Mining on Sage Grouse Movements and Habitat Period 1981 Use 1 January - 31 December 1980 R. A. Ryder, Colorado State University; Clait Braun, Howard Funk, Ken Giesen, Steve Porter, Tom Schoenberg, John Wagner, Colorado Division of Wildlife. ABSTRACT Investigations concerning sage grouse (Centrocercus urophasianus) movements and habitat use in North Park, Colorado continued in 1980. During February-May 515 sage grouse were captured in North Park including 166 females and 300 males banded for the initial time. Peak lek attendance of males in 1980 occurred during the same week of April as in 1979. Peak lek attendance of hens in 1980 occurred 1 week later than in 1979. Peak lek counts of male sage grouse in the northeast quadrat decreased 15% from 1979. Denmark lek, however, showed a 5% increase. Thirteen male and 18 female sage grouse were radio-marked between 1 February and 29 June. Twenty of 29 (69%) transmitter packages were recovered after use on sage grouse. Wildlife Materials transmitters had significantly longer (P < 0.05) transmitter life (209 days) than AVM transmitters (136 days). There was no significant difference > 0.05) between male and female sage grouse daily winter movements. Daily movements averaged 1.6 and 1.5 km for males and females, resp~ctively. Winter flock break-up and dispersal to breeding areas occurred during the first 2 weeks of April with the onset of the spring thaw. Movements of both sexes from wintering areas in the northeast and southeast quadrats were to the northwest (3) and southwest (4) quadrats. One male remained in the southeast quadrat during the breeding season. The mean distance from the lek visited to 7 nests was 2.5 km arid ranged from 0.3 to 7.8 km. Movements of both sexes from breeding areas and nests to meadows along the Michigan and Canadian rivers occurred throughout June, primarily during the latter half of the month. Four of 5 radio-marked males and 5 of 6 radio-marked hens moved to the meadow nearest the lek attended or nest site, respectively. Of 10 nests located, 3 were successful (30%), 3 were abandoned (30%), and 4 were depredated (40%). Mean sagebrush (Artemisia tridentata) canopy cover was greatest at male feeding-loafing sites (43%) followed by nest sites (42%), broodrearing sites (35%), and female pre-incubation feeding-loafing sites (32%). Average sagebrush height was greater at nest sites (35 em) (R �76 than male feeding-loafing sites (33 cm). Average sagebrush height was identical (26 cm) at female pre-incubation feeding-loafing sites and brood-rearing sites. Significant differences (~ < 0.05) in organic matter were found between soils from male and female use sites, and between soils from male use sites and random sites. Significant dif·ferences in manganese were found between all sites (male, female, random). Sage grouse pellet transect counts revealed declines in number of droppings/day of 38% from summer 1979 to summer 1980, and 42% from winter 1979-80 to summer 1980. The 1980 average of 1.4 birds/hunter was 0.5 bird/hunter lower than 1979 but still 0.3 bird/hunter higher than the previous 6-year average (1974-79). Two radio-marked hens from 1980 and 1 radio-marked hen from 1979 were shot during the hunting season. Only 11% of the sage grouse harvested in North Park in 1980 were bagged in the northeast quadrat. Although 30% of all banded birds shot in 1980 were recovered in the northeast quadrat, the direct recovery rate (harvest rate) was only 12%. Chick production in 1980 apparently decreased from 1979 but there was good yearling and adult survival. Overall nesting success was the lowest on record (42.7%) and the percent young in the harvest (49.1%) and young per hen (1.4:1) were below average. Young per successful hen (3.3:1), however, remained above average for the 3rd consecutive year. �77 POTENTIAL IMPACTS OF STRIP MINING ON SAGE GROUSE MOVEMENTS AND HABITAT USE Thomas J. Schoenberg Sage grouse are widely distributed throughout western North American rangelands dominated by sagebrush. Their dependence on sagebrush for breeding, nesting, brood-rearing, and wintering activities has been extensively documented (Girard 1937, Rasmussen and Griner 1938, Patterson 1952, Klebenow 1969, Eng and Schladweiler 1972, Wallestad and pyrah 1974, Wallestad and Schladweiler 1974, Wallestad et al. 1975, Beck 1977). Elimination and alteration of sagebrush for agricultural, grazing, and other developments have reduced both numbers and distribution of sage grouse throughout their historic range (Patterson 1952, Aldrich 1963). With increasing demands for western surface-mined coal, a greater reduction in sage grouse habitat can be expected in Colorado and other western states. Although reclamation of mined areas is required by state and federal laws, current rehabilitation practices concentrate on introduced grasses and forbs. Because reestablishment of the sagebrush community through succession is a long-term process, mined areas will be lost as sage grouse habitat for decades. In the face of shrinking amounts of sagebrush rangeland throughout the West, management of sage grouse populations will require a more complete understanding and detailed description of the sage grouse - sagebrush relationship. The need to identify important seasonal ranges and specific habitat preferences of sage grouse is especially important in areas to be disturbed by mining or other developments. Existing and proposed coal mines in Jackson County, Colorado will impact historic breeding, nesting, brood-rearing, and winter habitats. The sage grouse population in Jackson County is locally migratory with major wintering concentrations in the principal mining area in the northeast quadrat (Be~k 1975). Consequently, habitat disturbance in the northeast will impact the entire population in Jackson County. P. N. OBJECTIVES The objectives of this study during the predevelopment period are to: 1. Ascertain seasonal movement patterns and habitat preferences male and female sage grouse within the area to be impacted. 2. Continue population monitoring on known sage grouse leks within the study area. Data from monitoring population trends in adjacent areas and throughout North Park continued by research and management personnel will be used for comparative purposes. 3. Prepare management plan for predicting, impacts on the sage grouse resource. reducing, of . and mitigating �78 4. Based upon findings, prepare plan for rehabilitation of developed areas that would be attractive to and effective for maintaining and enhancing sage grouse populations. Hypotheses being tested are: a. Movements of sage grouse within sex class. b. Habitat class. c. Movement patterns for each sex class vary during critical periods of the annual cycle (prebreeding, breeding, nesting and brood rearing). d. Habitat use varies and brood rearing) selection (preference the study area differ for each and need) differs seasonally (prebreeding, for each sex class. for each sex breeding, nesting, SEGMENT OBJECTIVES 1. Review available literature concerning niche description and measurement, habitat use by sage grouse, sage grouse movements, use of radiotelemetry to identify habitat selection by birds, and western mineland reclamation. 2. Sage grouse pellet transects in the northeast quadrat of North Park will be searched in late Mayor early June and late $eptember or early October depending upon snow cover. All droppings found will be counted and cleared from the transect. 3. Establish 10 new sage grouse pellet transects in the northwest quadrat north of Independence Mountain. Transects, 0.5 km long and 2.5 m wide, will be counted and cleared in late September or early October. 4. Continue intensive counts (2-3 per week) of all sage grouse present on each known lek (Perdiz, Raven, Denmark, Roth, Pronghorn) within the study area. Emphasis will be placed on leks likely to be directly impacted (Raven, Perdiz, Denmark). Counts will be in the 0430-0730 interval from late March to late May. Counts of sage grouse present on other leks (4/season of leks with more than 10 males) in North Park will be continued. 5. Continue banding samples of male and female sage grouse within the study area. Grouse will be located where they roost by spotlighting and will be trapped with long handled nets. Some trapping of hens with cannon nets on leks may be necessary. Birds trapped will be banded with serially numbered aluminum bands and color coded (location) bandettes. A sample of 20% of the high spring count on each lek is desired each year. Samples of 50 cocks banded in each of the other 3 quadrats within North Park are desired for comparative purposes. �79 6. Equip 15 male and 15 female sage grouse with radios to ascertain movement patterns and habitat use preferences within the study area. While year-round documentation of habitat use is desirable, emphasis will be placed on spring (March-June) and late winter (January-February). Movements will be mapped and vegetation composition, height and canopy coverage will be ascertained for adequate samples (present sample size unknown) of habitats utilized by season for necessary functions of sage grouse. 7. Harvest data will be obtained from check wing collection barrels. Check stations Muddy Pass, and Willow Creek Pass during end of the hunting season. A sample of 8. Number and location of marked birds harvested through field checks of hunters and voluntary 9. Age composition of the harvest will be determined through examination of wings collected during field checks and through use of wing collection barrels. 10. Compile data, analyze results, and prepare progress and/or completion reports. Suitable findings will be submitted to appropriate technical journals for publication consideration. Management recommendations will be included in the final report. DESCRIPTION stations and volunteer will be operated at Gould, at least the opening week500 wings is desired. will be obtained mail reporting. OF THE STUDY AREA The investigation was conducted in North Park, Jackson County, Colorado (Fig. 1). The study area was centered in the mining area of the northeast quarter of North Park 13 km east of Walden, although all areas of North Park were included depending on movements of radio-marked birds. Vegetation of the area is characterized by the sagebrush-grassland type with native and irrigated meadows along major stream courses. Detailed geographic, geologic, vegetational and climatic features of the area will be treated in the final report. There are 5 known leks in the study area and 2 additional leks north of the Canadian River. Studies of sage grouse movements and habitat selection during 1980 were concentrated primarily on birds radio-marked on Raven lek but also those marked on Perdiz and Denmark leks. �80 INDEPENDENCE MOUNTAIN /Vl/\A.. LAKE JOHN AREA • WALDEN ~~.,.., ~~8 SPRING CREEK AREA ON ~/() G~ 0",(. ~/() G~ ~ N ~ 0 I 10 20 30 I I I KILOMETERS • FT. COLLINS • DENVER COLORADO Fig. 1. Location 1979-80. of coal mining study area in North Park, Colorado, �81 METHODS AND MATERIALS Sage grouse were captured while roosting along roads and on leks through-out the study area. Birds were located from a truck or on foot using a spotlight and were captured with a long-handled hoop net (Braun and Beck "1976). Captured grouse were marked with serially numbered size 14 (females) and size 16 (males) aluminum leg bands and unnumbered plastic bandettes color-coded to year of capture. Sex and age (Eng 1955, Beck et al. 1975), weight, and primary molt were determined for all birds captured. During 1980, 12 males and 17 females were fitted with 15-21 g 164 MHz tail clip radio packages (Bray and Corner 1972) attached to the 2 central rectrices. In addition, a 17 g solar-powered radio-collar and a 26 g battery-powered radio-collar (Amstrup 1980) were placed on a female and male, respectively. Transmitter packages were obtained from AVM Instrument Company (Champaign, Ill.) and Wildlife Materials, Inc. (Carbondale, Ill.). Daily locations of radio-marked sage grouse were made at leks; feedingloafing sites, and nests using a portable receiver and 3-element yagi antenna. A snowmobile was used during winter months to locate birds in areas inaccessible by 4-wheel drive vehicle. All locations, determined by triangulation or by flushing birds, were plotted on 7.5 minute U.S. Geological Survey topographic maps. Weather conditions, physiographic features, and flock size and composition were recorded at each site where birds were flushed. Weather conditions included temperature, precipitation, wind direction, and a wind speed estimate recorded as none, none~light, light, light-moderate, moderate, moderate-strong, strong, or very strong. Slope and aspect were measured with an Abney level and compass, respectively. Snow depth and percent snow cover were measured during winter months. Vegetation measurements were made at leks, feeding-loafing sites, nests, and" randomly located sites using a modification of Canfield's (1941) line interception method. Two 10-m transects were measured at each site. Transects were oriented along north-south and east-west axes with the bird location as the center of the plot at feeding-loafing and nest sites. The approximate mating center served as the center of the plot on leks. Random vegetation measurement sites were along 20 randomly located 500 x 2.5 m sage grouse pellet transects. Four sites were randomly chosen along each transect. Vegetation parameters included percent canopy cover of foliated and unfoliated sagebrush, forbs, grasses, litter, and bare ground. All forb species were collected and identified following Harrington (1954). Additional measurements were made of sagebrush crown length and width, and live and dead height of the tallest branch on the largest plant in each clump intersected by the transect. Sagebrush density was measured under a 2-m wide band along both transects at each site. Winter vegetation measurements were restricted to sagebrush and included both exposed and actual measurements after snow had been cleared from the transects. Grass height as well as canopy cover was measured in meadow areas where it provided most of the cover. �82 Soil samples, 15 cm in depth, were collected at each site measured in 1979. From these, 60 samples were randomly selected for analysis; 20 each from male feeding-loafing sites; female feeding, nesting and brooding sites; and randomly selected sites. Chemical analyses of the samples were performed by the Colorado State University Soil Testing Laboratory. Determinations of the following 12 soil characteristics were obtained: pH, conductivity (soluble salts), lime, organic matter, nitrate nitrogen, phosphorous, potassium, zinc, iron, manganese, copper, and soil texture. Conductivity and pH were evaluated using a saturated soil paste (Richards 1954). The lime test is an estimate based on sulfuric acid effervescence of the soil. Percent organic matter was determined using a potassium dichromate solution (Graham 1959). Nitrate nitrogen was extracted with water and evaluated using the chromotropic acid method of nitrate analysis (West and Ramachandran 1966). Phosphorus, potassium and micronutrients (Zn, Fe, Cu, Mn) were simultaneously extracted using ammonium bicarbonate in diethylene triamine pentaacetic acid (NH4HC03-DTPA) solution (Soltanpour and Schwab 1977). Phosphorous was determined by colorimetric evaluation and potassium and micronutrients were analyzed by atomic absorption (Soltanpour et al. 1979). Soil texture estimates were based on tactile examination. Soil data were analyzed using the Statistical Package for the Social Sciences (SPSS) programs (Nie et al. 1975). Student's ~-test was used to compare variables between sites at the P = 0.05 level of significance. Techniques used in lek census and sage grouse pellet transect counts were described earlier (Schoenberg 1980~). Ten new transects.were placed in the northwest quadrat of North Park north and east of Independence Mountain and west of the North Platte River. Transects were randomly chosen from 103 possible points on a l_km2 grid map of the approximately 55 km2 area of sagebrush rangeland. Wing collections were made during the sage grouse hunting season (13-28 September) at 4 wing collection barrels and 3 check stations. The wing collection barrels were maintained throughout the entire 16 day season at Gould, State Line, Willow Creek Pass, and Muddy Pass. Check stations were operated at Gould and Willow Creek Pass on 13, 14 and 21 September, and at Muddy Pass on 14 September. Data collected from each party of hunters at check stations were given by Braun (1979) and included county of origin; number of hunters; hours hunted (total of all hunters); number of grouse observed, bagged, and lost; number of banded birds bagged and location where shot; and the area hunted. In addition, individual hunters were asked whether or not they normally hunted sage grouse in North Park or if they were first time sage grouse hunters. �83 RESULTS AND DISCUSSION Banding Sample Trapping was conducted in wintering areas of the northeast and southeast quadrats of North Park on 25 nights between 21 January and 10 April. During this period 173 sage grouse were captured and 168 grouse (53 males, 115 females) were newly banded. The excellent trapping success during this period was due to locating large wintering concentrations of sage grouse during March in the coal-mining area along J. C. roads 12 and 29 and in the Spring Creek - Owl Creek area along J. C. Road 25. Of 173 grouse captured, 128 (74%) were caught in the northeast quadrat. Trapping during the breeding season was conducted throughout North Park on 20 nights between 16 April and 22 May. During this period, 342 sage grouse were captured. Fifty-one females (29 adults, 22 juveniles) and 251 males (145 adults, 106 juveniles) were newly banded. Recaptures included 2 females and 38 males. The quota of 50 newly banded males/ quadrat was exceeded in all quadrats with 63 in the northeast, 68 in the northwest, 53 in the southeast, and 67 in the southwest. During February-May 515 sage grouse were captured in North Park. Of these, 224 (44%) were captured in the northeast quadrat. There were 166 females (94 adults, 72 juveniles) and 300 males (171 adults, 129 juveniles) newly banded. Recaptures included 4 females (3 adults, 1 juvenile) and 41 males (39 adults, 2 juveniles). Six leks in the study area were trapped between 16 April and 20 May (Table 1). Sixty-three males (44 adults, 19 juveniles) and 12 females (3 adults, 9 juveniles) were newly banded. Recaptures included 12 adult males. The desired goal of banding 20% of the high spring count of males on each lek was exceeded on all but Denmark lek where only 16% of the peak count of males was newly banded. Table 1. Numbers of sage grouse banded on 6 leks, northeast Park, Colorado, 1980. Males Juveniles Lek Totals Juveniles 17 5 7 12 23 11 7 67 16 138 21 1 2 4 6 10 23 19 44 63 Canuck Denmark Perdiz Prague Pronghorn Raven -Totals Females % HSpa 3 6 6 apercent Adults 9 North of high spring count. Adults 2 1 1 5 9 3 Totals % HSP 3 3 15 27 1 5 12 12 �84 Lek Census Seven leks in the study area were censused between 17 March and 28 May. Denmark, Perdiz, Pronghorn, Raven and Roth leks were censused 1-2 times/ week during April and May while Canuck and Prague leks were censused once/week during May only. Sage grouse were first observed on Perdiz on 16 April. Initial observations of grouse on leks were about 1 week later than in 1979. Dates of peak lek attendance of males occurred between 21 April and 22 May (Table 2) with the season's peak for all leks occurring during the week of 11-17 May (Fig. 2). Peak male attendance occurred during the same week of April in 1980 as in 1979. Peak hen attendance on Denmark, Perdiz, and Raven leks occurred between 16-25 April, with the season's peak occurring during the week of 20-26 April, 3 weeks prior to peak male attendance. Peak hen attendance in 1980 occurred 1 week later than in 1979. Peak hen attendance dates for Canuck and Prague leks were not available since these leks were not counted during April. Table 2. Lek Canuck Denmark Perdiz Prague a Pronghorn Raven Roth High spring lek counts, Males .18 144 8 34 0 43 1 northeast Date(s) 01 May 14 May 2, 21 May 09 May all dates 07 May 22 Apr aOne male and 4 females were observed trapped on 29 April. North Park, Colorado, Females 2 20 11 10 0 38 0 1980. Date 15 May 29 Apr 16 Apr 09 May all dates 25 Apr all dates on the lek when it was Peak lek counts of male sage grouse decreased from 1979 to 1980 (Table 3) on all leks in the study area except Denmark, where there was a 5% in.crease. The 1980 total of 248 males represents a 15% decline from 1979. The greatest decline (100%) occurred on Pronghorn lek. Although no bf rd s were observed when censusing the lek, 1 male and 4 females were found roosting on the lek when it was trapped on 29 April. Perdiz and Roth leks had 50% declines from 1979 while Raven, Prague and Canuck leks declined 31, 21 and 14%, respectively. Raven lek declined in 1980 for the 2nd consecutive year whereas the remaining leks either increased or remained stable in 1979 and then declined in 1980. �85 40 /. . ::.::: ~ -: tI.l § 30 H ./ ~ ~ o MALES p:: ~ ~ z ~ ~ • 20 ~ ~ / • / .// i-:. • '\ / 10 / '\ FEMALES ,, '\ '. - - - '. -_ •...... N •....• I \0 0\ \0 N I 0 N •....• I C""l •....• APR C""l 0 I •...... 0 •....• I ...;:t •....• I •....• •....• --a -cr N I 00 •....• N MAY Fig. 2. Sage grouse lek attendance on 7 leks, northeast North Park, Colorado, 1980. �86 Table 3. Trends in peak lek counts of male sage grouse, northeast North Park, Colorado, 1977-80. Year 1977 1978 1979 1980 Canuck Denmark Perdiz Prague Pronghorn Raven Roth 27 58 21 80 18 144 8 34 15 73 8 35 10 94 1 21 136 16 43 10 63 2 Totals 181 241 291 248 36 40 42 35 Lek Avg./lek aOne male and 4 females were observed on 29 April. Oa 43 1 Percent change 1979 to 1980 - 14 + 5 - 50 - 21 -100 - 31 - 50 - 15 on the lek when it was trapped Radiotelemetry Between 1 February and 29 June, 29 transmitters were fitted on 13 male (9 adults, 4 juveniles) and 18 female (11 adults, 7 juveniles) sage grouse. Four males and 9 females were followed during the winter and 9 each males and females were followed during the breeding season. Twenty of 29 (69%) transmitters were recovered. The recovery rates of AVM and Wildlife Materials (WM) transmitters were approximately equal. Ten of 15 AVM transmitters and 10 of 14 WM transmitters were recovered. Tail clip radio package recoveries were made after premature loss of the central rectrices (5), predation (4), recapture (4), rectrixmolt (4), hunter kill (1), and unknown mortality (1). The solar-powered radio-collar was recovered after it either had slipped off or had been pulled off by the hen. Transmitter life was calculated for AVM (Table 4) and WM (Table 5) transmitters. Average transmitter life and range were evaluated using only those radios which were recovered and monitored or those which shut off while the bird was being tracked. Transmitters on birds with which radio contact was lost could not be included since the birds may have moved out of the area while transmitters were still functioning. It was especially difficult to maintain radio contact with AVM transmitters since the maximum line-of-sight receiving distance was usually less than 5 km whereas WM transmitters could often be heard up to 15 km. Differences in maximum line-of-sight reception distances between AVM and WM transmitters can be largely attributed to differences in power output. WM transmitters contained 3.0 V lithium batteries whereas AVM transmitters contained 1.5 V mercury batteries. �87 Table 4. Colorado, Life of AVM transmitters 1980. Channel Pulse 56 47 58 42 60 60 47 67 1 2 2 3 3 4 5 5 Transmitter life(days)b 249 207 213 34 44 6 6 8 8 9 10 12 64 51 65 47 64 45 Avg. Range a 37 129 47 169 used on sage grouse Comments Radio package " " and monitored " " " " 1.4 V Hg batteries. Table 5. Life of Wildlife Materials in North Park, Colorado, 1980. Pulse 50 47 51 48 42 43 46 59 60 59 53 66 72 Avg. Range " " Lost radio contact after 212 days Radio package recovered and shut off Lost radio contact after 10 days " " " "121" Radio shut off when tracking bird Lost radio contact after 73 days " " " "17" Radio shut off when tracking bird Radio package recovered after hunter kill Radio shut off when tracking bird Radio package recovered and monitored Lost radio contact after 72 days bOnly those radios which were recovered tracking bird were included. 2 5 6 7 8 8 9 9 9 10 11 11 11 recovered " " 136 34-249 aSMl with Channel in North Park, Transmitter life(days)b 199+ 215+ 214+ 246 208+ 145 260+ 193+ 199+ or which shut off while transmittersa used on sage grouse Comments Radio package " " " " recovered " " and monitored " " " " Lost radio contact after 128 days Radio package recovered and monitored Lost radio contact after 11 days Radio package recovered and monitored " " " " " " " " " " " " " " " " " " " " Lost radio contact after 87 days 81 " " " " " 209 145-260+ aHLP-2750-LD with 3.0 V Li batteries. bOnly those radios which were recovered were included. +rndicates transmitters were returned to manufacturer replacement prior to shut off. for battery �88 (R Average transmitter life of WM radios was significantly greater < 0.05) and averaged 73 days longer than average transmitter life of AVM radios. Individual and average transmitter life of WM radios would likely have been even greater had not 7 transmitters been returned to the manufacturer for battery replacement before the old batteries had expired. Sage Grouse Movements Winter Preliminary data on daily movements and winter ranges were compiled for 1980 (Table 6). Daily movements of males averaged 0.1 km greater than daily movements of hens but there was no significant difference (R> 0.05). The greater range of daily movements by males was due to the difference in reception distance of the WM and AVM transmitters used on males and females, respectively. Males could easily be located daily even after long distance movements of 8-10 km whereas if a hen moved a long distance, it often took several days to relocate her. Table 6. Daily movements and winter ranges of 4 male and 7 female sage grouse, North Park, Colorado, 1980. Daily movements (km) Avg. Range Sex Males 57 1.6 0-10.9 Females 34 1.5 0.1- 5.3 Winter range (ha) 5,000-25,000 6,400-n,900 Winter ranges were also greater for males than females although both sexes used the same wintering concentration areas. Here again, the discrepancy is probably due to the shorter reception distance of AVM radios used on hens. Movements by hens, however, were much greater than those reported by Eng and Schladweiler (1972) in Montana where female winter ranges were between 1,059 and 3,142 ha (2,615 - 7,760 ac). There were 2 major wintering areas used by sage grouse in 1980. These areas had been outlined previously by Beck (1975) during the winters of 1973-74 and 1974-75. One area was in the northeast quadrat along J. C. Road 12 and north and east of the Kerr Coal Mine approximately 16 km east of Walden. The 2nd area was located in the southeast quadrat in the Spring Creek - Owl Creek area 15-20 km southeast of Walden. Although all birds were radio-marked in the northeast, they all eventually moved into the southeast area, a distance of approximately 12 km from J. C. Road 12, prior to dispersing to breeding areas throughout North Park. One bird, male 7149, moved back and forth between the 2 areas on 2 occasions. �89 To Breeding Areas Winter flock break-up and dispersal to breeding areas began during the first 2 weeks of April, coincident with the onset of the spring thaw. All radio-marked birds had moved out of the northeast during early April. Movements of 3 males from wintering areas were to the northwest (2) and southwest (1) quadrats. Adult male 8829 remained in the Spring Creek Owl Creek area where he had spent the last week of winter and attended Spring Creek 2 lek. Adult male 7149 returned to Hound lek north of Walden Reservoir where he was originally banded as a juvenile in 1978. Adult male 8802 moved to Bighorn lek north of Lake John, also in the northwest quad ra t . Juvenile male 8801 first moved to the area near Fish Hatchery lek in the southwest quadrat. He continued to move southwest and was last located near Cheyenne lek. Movements of 4 hens were to the northwest (1) and southwest (3) quadrats. Adult hen 3429 attended Boettcher Lake Junction lek on 25 April. Her nest was located on 14 May approximately 1 km west of the lek. Nests of adult females 3427, 3432, and 5032 were in the southwest quadrat although it is not known which leks they attended. The nest of hen 3427 was found on 14 May just west of Beaver Creek. Cheyenne is the nearest known lek to her nest site. Female 3432 was located on her nest on Pole Mountain on 15 May. Coalmont and Migan are the nearest known leks in that area. Hen 5032 nested approximately 1 km east of Ridge Road lek. Juvenile hen 3428 was last located on 17 April in the Spring Creek drainage in the southeast quadrat. Radio contact was lost after that date so it is not known whether she remained in that area to breed and nest. Breeding Season Three adult and 2 juvenile males from Raven lek, 1 adult and 1 juvenile male from Perdiz lek, and 2 adult males from Denmark lek were fitted with transmitters during April and May. All 6 adult males remained in the area surrounding the lek where they were captured throughout the breeding season. However, all 3 juveniles moved to another lek during the breeding season. Male 8731 moved from Raven to Denmark on 25 May and remained there the rest of the breeding season. Male 8981 also moved from Raven to Denmark on 25 May but returned to Raven by 28 May. Male 8982, trapped on Perdiz lek on 12 May, was on Raven lek on 14 May and'remained there throughout the breeding season. Distances and directions of daily movements from the lek by males to feeding-loafing sites have not been evaluated for 1980. Ten nests of 9 radio-marked and 1 unmarked hens were located in the study area. Seven of 9 radio-marked hens were captured on Raven (1 adult, 3 juveniles), Denmark (2 adults), and Perdiz (1 adult) leks. Distances from the lek to the 7 nest sites averaged 2.5 km and ranged from 0.3 to 7.8 km , Distances of nests from Raven lek averaged 1.85 km. Adult hen 5063 nested 4.7 km south of Raven lek, just east of the Wyoming Fuels Company coal mine. Juvenile females 5072, 5086, and 5087 nested 1.4 km northeast, 1.0 km southwest, and 0.3 km southwest of Raven lek, respectively. Distance and direction from Denmark lek to nests of adult �90 hens 5075 and 5088 here 1.4 km west and 7.8 km east, respectively. hen 5064 nested 0.9 km northwest of Perdiz lek. Adult Juvenile females 5089 and 5090 were trapped along roads near Denmark and Raven leks, respectively. Hen 5089 nested 2.5 km southwest of Denmark. Hen 5090 nested 3.7 km southwest of Raven. To Summer Areas Males began moving from breeding areas to meadows along the Michigan and Canadian rivers during June. Four of 5 males moved to the meadow area nearest the lek they attended during the breeding season. Adult male 8991, however, moved approximately 7.5 km from Raven lek to the Michigan River meadows instead of 2.8 km to the Canadian River meadows. Both of the other males from Raven lek (7016, 8981) and 1 male from Denmark lek (8731) moved to the Canadian River meadows between 13 and 23 June. Male 8879 moved to the Michigan River meadows from Perdiz lek between 5 and 20 June. Movements of radio-marked hens to meadow areas occurred during mid-late June, regardless of nesting success. Five of 6 hens moved to the meadow nearest their nest sites. However, yearling hen 5086, which nested just south of Raven lek, moved 7 km to the Sage Hen Creek drainage along the Michigan River meadows when the Canadian River meadows were only 3 km from the nest. Hen 5072, also nesting near Raven lek, and hens 5088 and 5089 from the Denmark lek area moved to the Canadian River meadows between 19 and 23 June. Two hens nesting in the Perdiz lek area moved to the Michigan River meadows. Adult hen 5063 and her brood moved approximately 4 km south from the nest site to the meadows by. 23 June, 15 days after hatching. Adult hen 5064 continued to feed in sagebrush habitat throughout most of June after her nest had been destroyed. She did not arrive at the Michigan River meadows until after 20 June. Summer Summer movements within meadows were restricted to relatively small areas along the Michigan and Canadian rivers. Two males and 1 unsuccessful hen were followed from late June through early August in the Canadian River meadows. All 3 birds remained within a 1 km stretch of the irrigated meadow area between Sherman and Yockum creeks, primarily on the north side of the Canadian River. Two males and 2 females were periodically located along the Michigan River meadows. Hen 5063 and her brood remained within a 1 km stretch of irrigated meadow just south of Bolton Draw. An unsuccessful hen, 5064, spent the summer along a 1-2 km stretch of irrigated meadow south of the Brocker Ranch headquarters. Both males restricted their movements to about a 1 km stretch of irrigated meadow northeast of the Owl Creek Ranch headquarters. �91 Nest Success Of 10 nests located in the study area, 3 were successful (30%), 3 were abandoned (30%), and 4 were depredated (40%). Three of 4 nest predations appeared to have been caused by ground squirrels (Spermophilus richardsonii) while the 4th was probably destroyed by a coyote (Canis latrans) or badger (Taxidea taxus). Two of 3 nest abandonments were caused by the investigator. Cause of the 3rd abandonment is unknown -but may also have been caused by the investigator since the hen was flushed from her nest during the first few days of incubation. There had been some disturbance at the nest after my first visit, however. When the nest was revisited, 1 egg had been moved about 60 cm from the nest but none had been destroyed. Although the egg was returned to the nest, the hen did not return. Hen 5087 abandoned her nest after laying 3 eggs when her transmitter package got caught in sagebrush at the nest and was pulled out with her central rectrices. Hen 5072 abandoned her nest after 22 days of incubation when she was recaptured on her nest and her transmitter package was replaced. Nests of 4 adult hens (3427, 3429, 3432, 5032) totaling 31 eggs were collected outside the study area on 16 May and taken to Fort Collins to be incubated by bantam hens. Twenty-seven eggs hatched between 8 and 10 June. Weight and primary feather growth measurements of each surviving chick were recorded at weekly intervals. Habitat Selection Winter Winter vegetation measurements were made at 53 male feeding-loafing sites, 50 female feeding-loafing sites, and 52 random sites. Although data from these sites have not yet been analyzed, it was quite apparent that exposed sagebrush canopy cover and height at sage grouse- feedingloafing sites were markedly greater than at random sites. Breeding Season Comparisons of sagebrush canopy cover and height were made for a limited number of male feeding-loafing sites, female pre-incubation feedingloafing sites, nests, and brood-rearing sites in 1979 and 1980 (Table 7). Twenty additional random sites were also measured in 1980. A complete analysis of all data will be presented in the final report. Table 7. Sagebrush canopy cover and average height at random and sage grouse use sites, northeast North Park, Colorado, 1979 and 1980. N Sitea 12. 3. 4. 5. Random Male FL Female PIFL Nest Brood a 1979 1980 60 59 35 5 12 16 12 5 8 2.=male feeding-loafing loafing site. Canopy cover(%) 1980 1979 26 33 29 27 23 site, 3.=female Height (cm) 1980 1979 43 32 42 35 pre-incubation 19 28 23 30 19 33 26 35 26 feeding- �92 Measurements of sagebrush canopy cover increased an average of 10% and sagebrush height increased an average of 5 cm at the 4 sites between 1979 and 1980. This increase in canopy cover and height of sagebrush may be the result of small sample sizes in 1980. Sagebrush canopy cover was greatest at male feeding-loafing sites during both years with 33% in 1979 and 43% in 1980. In contrast to 1979, however, nest sites had the 2nd highest canopy cover in 1980 (42%) followed by brood-rearing sites (35%), and female pre-incubation feedingloafing sites (32%). In 1979, female pre-incubation feeding-loafing sites had the 2nd highest canopy cover (29%) followed by nest sites (27%), and brood-rearing sites (23%). The ranking of average sagebrush height at these 4 sites was similar between years. The greatest average sagebrush height was found at nest sites in both years, 30 cm in 1979 and 35 cm in 1980. Male feedingloafing sites had the 2nd highest average sagebrush height in 1979 (28 cm) and 1980 (33 cm). In 1979, average sagebrush height at female pre-incubation feeding-loafing sites (23 cm) was greater than broodrearing sites (19 em) whereas in 1980, these values were identical (26 cin). Sununer Vegetation measurement data for sage grouse feeding-loafing sites in meadow areas have not yet been analyzed. These data will be presented in the final report. Soil Samples Soil samples collected in 1979 at male and female sage grouse use sites and random sites were analyzed. Mean values of 9 soil variables were compared among sites (Table 8). Significant differences (~ < 0.05) in manganese were found between all sites. Significant differences in organic matter were found between male and female use sites and between random sites and sites used by males. Levels of pH, copper and the mean vectors of all variables were also significantly different between random sites and sites used by males. The differences in organic matter are of greatest biological importance, Organic matter directly and indirectly influences nutrient availability to plants. A 1% difference in organic matter is significant in plant nutrition (Kononova 1966). When organic matter is broken down it provides a direct source of nutrients in available form. About 99% of the nitrogen in the soil is contained in the organic matter (Thompson and Troeh 1973). The most important contributions of organic matter are indirect, however. Breakdown of organic ma.tter by microbial activity improves soil structure so that nutrients and water are more readily available. Soils with high organic ma t t e.r have greatest microbial activity leading to increased production of C02 and organic acids. Organic acids lower soil pH. Cations of micronutrients (Zn, Fe, Cu, Mn) are more available to plants under acidic conditions (Sauchelli 1969). �93 Table B. Mean values of 9 soil variables at sites used by male and female sage grouse and random sites, North Park, Colorado, 1979. 'Variable pH Organic matter, % Nitrate nitrogen, ppm Phosphorus, ppm Potassium, ppm Zinc, ppm Iron, ppm Manganese, ppm Copper, ppm Male 6.34 4.01 4.05 9.65 290 3.53 35.75 21.94 3.0B Sites Female 6.50 3.04 3.70 7.20 339 2.63 36.Bl 15.77 2.45 Random 6.77 2.95 3.10 7.B5 269 2.22 23.90 11.IB 2.34 A pattern of high organic matter, low pH and high level of available micronutrients can be seen in the data (Table B). Therefore, the significant differences in manganese, copper, and pH are a reflection of the differences in organic matter. Levels of all nutrients are adequate for healthy growth of most agricultural crops (Sauchelli 1969). Little work has been done on minimum nutrient requirements of sagebrush although some work has been done on mineral cycling in sagebrush ecosystems (Mayland and Murray 1979). Soil organic matter content is positively correlated with plant growth (Kononova 1966). Structural differences in sagebrush at sites used by male and female sage grouse and random sites have been recognized in the study area (Schoenberg 19BOa). Males selected areas with greater average sagebrush canopy cover and height than females and females selected areas with greater average sagebrush canopy cover and height than found at random sites. This same pattern is reflected in higher (f < 0.05) percent organic matter in soils collected at male use sites than at female use sites and random sites, with only slightly greater percent organic matter at female use sites than random sites. Whereas soils at male use sites were significantly higher in organic matter than random sites, and soils collected at female use sites and random sites had approximately the same levels of organic matter, this does not mean that female sage grouse are not selecting sites. Females may be selecting sites similar to random sites on the average. Percent organic matter at sites used by females was less variable than at random sites (Fig. 3). Differences between sites may also be obscured by the fact that random sites include potential sage grouse use sites as well as nonuse sites. �94 7.0 6.0 5.0 p:: ~ E-+ E-+ 4.0 X ~ U H ~ c.!) p:: a E-+ z ~ u 3.0 p:: ~ p.. Range 2.0 (B_=20) (B_=20) (B_=20) 1.0 o Male Female RANDOM Fig. 3. Percent organic matter at sites used by male and female sage grouse and random sites, North Park, Colorado. 1979. �95 Pellet Transects Twenty sage grouse pellet transects in the northeast quadrat were counted and cleared between 25 and 29 June and again on 11 and 12 October. The number of droppings/day (# dps/day) was calculated for summer 1979, -winter 1979-80, and summer 1980 (Table 9). Comparisons of number of droppings per day rather than total number of droppings are preferable since pellet counts were not made at equal intervals. Between winter 1979-80 and summer 1980, number of droppings per day decreased 42%. If pellet count data are used alone, they would indicate that 1980 overwinter and breeding season use of the northeast quadrat of North Park was 1.7 times greater (0.65/0.38 = 1.71) than 1980 summer use. It is known from Beck's work (1977) and radiotelemetry studies (Schoenberg 1980b) that grouse from all quadrats of North Park migrate to the northeast quadrat during winters with heavy snowfall. This occurs because there is lower snowfall and more available sagebrush in the northeast than in other quadrats. Withou~ long-term pellet count data and better understanding of variables (topography, snow cover and depth, sagebrush cover and height, habitat disturbance) affecting sage grouse use of sagebrush between seasons, it is not possible to determine whether or not this is an accurate estimate of the change in intensity of :use by grouse between seasons. The actual difference may be greater or less than 1.7 times. Other factors which bias the observed difference between seasons include placement of transects (random) and time period over which the count is made. Comparisons using all 20 transects may not be appropriate since some transects within the northeast quadrat are located in areas which receive greater winter use than others. Conversely, some transects may not be representative of summer use intensity. Inclusion of 1980 breeding season droppings with 1979-80 winter droppings may also bias the difference. Overwinter and breeding season droppings were indistinguishable in late June when transects were cleared. It would be expected that inclusion of breeding season droppings with overwinter droppings would lower the average number of droppings per day since there are fewer grouse in the northeast quadrat during the breeding season. Grouse which winter in the northeast quadrat return to breeding areas with the onset of the spring thaw (Schoenberg 1980b). Summer dropping accumulations in 1980 were 38% less than in 1979. Comparison of the number of droppings per day indicates 1.6 times greater (0.61/0.38 = 1.61) summer use in 1979 than in 1980. However, this difference may be exaggerated. Since the 1979 winter count was made in early June and the 1980 winter count was made in late June, the 1980 summer count does not include a 3-week period of breeding season use. This period is disproportionately more-important than the remaining summer months in terms of sage grouse use of sagebrush uplands. Since sage grouse moved from sagebrush uplands to meadow areas during late June in 1980 (Schoenberg 1980b), there was little summer use of sagebrush habitat where pellet transects are located. The actual difference between summers is probably less than the observed factor of 1.6. �Table 9. Number of sage grouse droppings/day encountered on 20 transects, northeast North Park, Colorado, Summer 1979, Winter 1979-80, and Summer 1980. Summer 1979 Winter 1979-80 ! dps/ Transect ! dps ! days day ! dps ! days 3 4 48 57 60 74 78 81 83 88 99 115 129 136 166 174 179 194 196 197 0 0 2 3 182 51 72 114 121 14 81 7 242 106 57 0 661 4 58 17 143 143 151 151 143 142 144 144 144 154 144 143 150 149 143 143 143 149 149 149 0 0 0.01 0.02 1.27 0.36 0.50 0.79 0.84 0.09 0.56 0.05 1.61 0.71 0.40 0 4.62 0.03 0.39 0.11 8 499 183 26 17 7 540 125 144 14 793 118 6 229 69 0 100 52 249 20 242 242 242 242 246 246 246 245 245 246 245 245 245 246 242 242 242 245 245 245 Totals Avg. 1,792 Summer 1980 ! dps/ 0.61 ! dps/ Summersummer day ! dps ! days day 0.03 2.06 0.76 0.11 0.07 0.03 2.20 0.51 0.59 0.06 3.24 0.48 0.02 0.93 0.29 0 0.41 0.21 1.02 0.08 3 26 154 6 107 8 63 72 2 6 87 26 12 55 132 0 5 108 108 108 108 105 105 105 104 104 105 104 105 105 105 108 108 108 106 106 106 0.03 0.24 1.43 0.06 1.02 0.08 0.60 0.69 0.02 0.06 0.84 0.25 0.11 0.52 1.22 0 0.05 0.05 0.21 0.12 + 100 100 142 200 20 78 20 13 98 33 50 400 93 27 205 0 99 67 46 9 106 0.38 - 38 3,199 146 Change in dps/day(%) 5 22 13 + + + + + + + + + - Wintersummer 0 88 + 88 - 45 +1357 + 167 - 73 + 35 - 97 0 - 74 - 48 + 450 - 44 + 320 0 - 88 - 76 - 79 + 50 - 804 244 0.65 - 42 1.0 a- �97 Pellet transects in the northwest quadrat of North Park were established on 18 and 19 October, and counted and cleared on 1 November. The number of fresh droppings per transect varied from 0-228. No fresh droppings were found on 5 of the 10 transects. Five transects on which few or no droppings were found are within 1-2 km of the sagebrush-aspen ecotone east and north of Independence Mountain. No droppings were found on another transect located on heavily grazed private land. Harvest Data Sage grouse harvest data from Gould, Willow Creek Pass and Muddy Pass check stations were tabulated (Table 10). During 7 check~station days of operation, 567 hunters bagged 794 sage grouse in North Park. This average of 1.4 birds/hunter represents a 0.5 bird/hunter decrease from 1979. However, it is still 0.3 bird/hunter more than the previous 6-year average (1974-79) as reported by Braun (1979). Table 10. Sage grouse harvest Park, Colorado, 1974-80.a statistics from check stations, Year No. of hunters checked No. of birds observed No. of birds harvested 1974 1975 1976 1977 1978 1979 1980 730 738 595 353 350 521 567 6,062 5,735 3,393 3,303 4,922 6,910 5,000 785 551 459 385 480 982 794 aUnpublished data North No. of banded birds No. of birds! hunter 57 47 1.1 0.7 0.8 1.1 1.4 1.9 1.4 (Braun 1980). There were 47 banded birds reported at check stations representing 5.9% of the total bag. This is comparable to 1979 when banded birds accounted for 5.8% of the total bag. Bands from 2 radio-marked hens from 1980 (5063, 5086) and 1 radio-marked hen from 1979 (3401) were recovered during the hunting season. All 3 hens were shot by hunters. Hunter and Harvest Distribution Sage grouse hunter pressure and harvest as determined from check station data have been consistently low in the northeast quadrat of North Park (Table 11). From 1974 to 1980, an average of only 11% of all hunters surveyed hunted in the northeast and bagged an average of 13% of the harvest. The northwest and southwest quadrats have traditionally had high hunting pressure and harvests. With the opening of the Arapahoe National Wildlife Refuge (ANWR) to sage grouse hunting in 1978, the harvest in the southwest quadrat increased substantially in 1978 and 1979. Although the 1980 sage grouse harvest decreased in the southwest, �98 it remained high in the ANWR area, where approximately one-third of the sage grouse harvested in North Park were bagged. The percent of the harvest in the northwest quadrat has remained relatively stable during 1978-80 but the percent of the hunters has decreased indicating a shift in hunter distribution to the other quadrats of North Park. Table 11. Sage grouse hunter and harvest Colorado, 1974-1980.a,b Northeast % Hunt. Harv. Year 1974 1975 1976 1977 1978 1979 1980 7-year avg. Southeast % Hunt. Harv. in North Park, Southwest % Hunt. Harv. 12 13 11 9 8 9 l3 14 14 11 15 15 9 11 35 42 44 46 49 35 29 38 45 36 37 28 30 33 18 14 11 8 6 13 14 13 6 6 8 3 9 13 35 30 34 36 38 44 44 35 32 47 40 53 51 43 11 l3 40 35 12 9 37 43 aUnpublished b Northwest % Hunt. Harv. distribution data (Braun 1980). Data from check stations only. Band Recoveries There were 26 band recoveries in the northeast in 1980 from hunter kills (20), predator kills (5), and unknown cause (1). This is the highest number of bands ever recovered in the northeast (Table 12). During the 1980 hunting season 30% (20 of 67) of all banded birds were shot in the northeast whereas in 1979, only 15% (11 of 72) were shot in the northeast. This may have occurred as a result of greater numbers of banded birds in the northeast quadrat compared to other quadrats. The northeast accounted for 33 and 44% of the banded sample in 1979 and 1980, respectively. However, it can be expected that many of these banded birds emigrated from the northeast quadrat after being banded since many were banded during February - early April in wintering concentration areas of the northeast. In 1980, only 27% of all birds banded during the breeding season were banded in the northeast quadrat. �99 Table 12. Sage grouse band recoveries Park, Colorado, 1973-80. in the northeast quadrat of North Year banded Total banded in North Park 1973 1974 1975 1976 1977 1978 1979 1980 288 191 421 379 540 258 357 466 2 0 0 0 0 3 0 0 0 0 0 0 1 0 3 0 0 0 0 3 4 0 0 0 0 3 2 6 0 0 0 0 2 2 9 13 2,900 2 0 3 0 4 7 11 26 Totals 1973 Year recovered in northeast quadrat 1974 1975 1976 1977 1978 1979 1980 The direct recovery rate provides a more realistic assessment of the harvest rate in the northeast quadrat in 1980. The direct recovery rate was 5% (10 of 206) for birds banded from February through May and 12% (10 of 83) for birds banded only during the breeding season. A 12% harvest rate is probably a more realistic estimate for the northeast due to emigration of birds banded in wintering areas during February-April. Check station hunter survey data (Table 11) indicate that 11% of the total harvest was from the northeast quadrat in 1980. Wing Analysis During the 1980 hunting season, 939 wings were collected in North Park. Examination of the age structure (Table 13) indicates that production was down 8.6% from 1979 but only 0.8% below the 7-year average. Yearling and adult survival increased from 1979. The percent of yearlings in the 1980 sample was the highest on record due to excellent production in 1979. Table 13. Age structure Colorado, 1974-80.a of the sage grouse harvest Immatures Year N 1974 1975 1976 1977 1978 1979 1980 350 212 210 290 385 624 461 7-year avg. aUnpublished in North Park, Yearlings % 50.1 42.0 42.3 45.9 53.2 57.7 49.1 N 138 III 117 123 143 256 250 49.9 data (Braun 1980). Adults % 19.8 22.0 23.5 19.5 19.8 23.7 26.6 22.4 N 210 182 170 219 196 201 228 % 30.1 36.0 34.2 34.6 27.1 18.6 24.3 27.7 �100 Wings from 328 hens (158 adults, 170 yearlings) harvested in 1980 were classified as to primary molt to estimate nesting success (Table 14). Estimated nesting success of adults and yearlings in 1980 was below the 7-year average. The overall success rate was the lowest ever recorded. Nevertheless, percent young in the harvest (49.1%) and young/hen (1.4:1) fell just slightly below the 7-year averages and young/successful hen (3.3:1) and birds/hunter (1.4) were above the 7-year averages. Table 14. Sage grouse nesting success and production success in North Park, Colorado, 1974-80.a Estimated nesting success Adults Yearlings Year (%) Overall % young in harvest Young per hen rates and hunter Young per successful hen Birds per hunter 1974 1975 1976 1977 1978 1979 1980 64.5 53.2 52.9 59.3 59.7 65.1 55.7 46.1 39.0 26.8 32.7 38.3 55.8 30.6 58.7 48.6 43.2 50.3 51.4 60.1 42.7 50.1 42.0 42.3 45.9 53.2 57.7 49.1 1.4: 1 1.1: 1 1.1: 1 1.2: 1 1.8: 1 2.2:1 1.4: 1 2.3:1 2.3:1 2.5:1 2.0:1 3.6:1 3.7:1 3.3:1 1.1 0.7 0.8 1.1 1.4 1.9 1.4 7-year avg. 58.6 38.5 50.7 49.9 1.5: 1 2.8:1 1.2 aUnpublished data (Braun 1980). LITERATURE CITED Aldrich, J. W. 1963. Geographic orientation of North American tetraonidae. J. Wildl. Manage. 27:529-545. Amstrup, S. C. 1980. A radio-collar Manage. 44:214-217. for game birds. Beck, T. D. I. 1975. Attributes of a wintering grouse, North Park, Colorado. M.S. Thesis. Fort Collins. 49pp. J. Wildl. population of sage Colo. State Univ. 1977. Sage grouse flock characteristics and habitat selection in winter. J. Wildl. Manage. 41:18-26. , R. B. Gill, and C. E. Braun. 1975. ----=of sage grouse from wing characteristics. Game Inf. Leafl. 49 (Revised). 4pp. Sex and age determination Colorado Div. Wildl. �101 Braun, C. E. 1979. Evaluation of the effects of changes in hunting regulations on sage grouse populations. Colorado Div. Wildl. Prog. Rep., Fed. Aid Proj. W-37-R-32, Work Plan 3, Job 9a. pp. 11-35. -~-- , and T. D. I. Beck. 1976. Effects of sagebrush control on distribution and abundance of sage grouse. Colorado Div. Wildl. Final Rep., Fed. Aid Proj. W-37-R, Work Plan 3, Job 8a. pp. 21-84. Bray, O. E., and G. W. Corner. 1972. A tail clip for attaching mitters to birds. J. Wildl. Manage. 36:640-642. Canfield, R. H. 1941. Application of the line interception sampling range vegetation. J. For. 39:388-394. trans- method in Eng, R. L. 1955. A method for obtaining sage grouse age and sex ratios from wings. J. Wildl. Manage. 19:267-272. , and P. Schladweiler. 1972. habitat use in central Montana. ---- Sage grouse winter movements and J. Wildl. Manage. 36:141-146. Girard, G. L. 1937. Life history, habits, and food of the sage grouse, Centrocercus urophasianus Bonaparte. Univ. Wyo. Publ. 3:1-56. Graham, E. R. testing. of soil 20pp. 1959. An explanation of theory and methods Univ. Missouri Agric. Exp. Sta. Bull. 734. Harrington, H. D. 1954. Manual of the plants of Colorado. Books, Inc. Denver, Colo. 666pp. Klebenow, D. A. 1969. Sage grouse nesting and brood habitat J. Wildl. Manage. 33:649-662. Kononova, M. M. 1966. Soil organic matter. Oxford, U.K. 544pp. Sage in Idaho. Pergamon Press Ltd., Mayland, H. F., and R. B. Murray. 1979. Mineral-cycling aspects within the sagebrush ecosystem. Pp. 62-73 in The Sagebrush Ecosystem: A Symposium. Utah State Univ. Logan. Nie, N. H., C. H. Hull, J. G. Jenkins, K. Steinbrenner, and D. H. Bent. 1975. Statistical package for the social sciences. McGraw-Hill Book Co. New York, N. Y. 675pp. Patterson, R. L. 1952. The sage grouse in Wyoming. Denver, Colo. 341pp. Sage Books, Inc. Rasmussen, D. I., and L. A. Griner. 1938. Life history and management studies of the sage grouse in Utah, with special reference to nesting and feeding habits. Trans. North Am. Wildl. Conf. 3:852-864. Richards, L. A., (ed.). 1954. Diagnosis and improvement alkali soils. U.S. Dep. Agric. Handb. 60. 160pp. of saline and �102 Sauchelli, V. 1969. Trace elements in agriculture. Reinhold Co. New York, N. Y. 248pp. Van Nostrand Schoenberg, T. J. 1980a. Potential impacts of strip mining on sage grouse movements and habitat use. Colorado Div. Wildl. Prog. Rep., Fed. Aid Proj. W-37-R, Work Plan 3, Job 12. pp. 243-264. 1980b. Potential North Park, Colorado. impacts of strip mining on sage grouse, Unpub. Prog. Rep. July. 9pp. Soltanpour, P. N., and A. P. Schwab. 1977. A new soil test for simultaneous extraction of macro- and micronutrients in alkaline soils. Commun. in Soil Sci. and Plant Anal. 8:195-207. ------- , S. N. Workman, and A. P. Schwab. 1979. Use of inductivelycoupled plasma spectrometry for the simultaneous determination of macro- and micronutrients in NH4HC03-DTPA extracts of soils. Soil Sci. Soc. Am. J. 43:75-78. Thompson, L. M., and F. R. Troeh. 1973. Soils and soil fertility. McGraw-Hill Book Co. New York, N. Y. 495pp. Wallestad, R. 0., and D. Pyrah. 1974. Movement and nesting of sage grouse hens in central Montana. J. Wildl. Manage. 38:630-633. ------- , and P. Schladweiler. tat selection 1974. Breeding season movements and habiof male sage grouse. J. Wildl. Manage. 38:634-637. J. G. Peterson, and R. L. Eng. 1975. Foods of adult sage grouse in central Montana. J. Wildl. Manage. 39:628-630. West, P. W., and T. P. Ramachandran. 1966. Spectrophotometric determination of nitrate using chromotropic acid. Anal. Chimica Acta. 350:317-324. Prepared b;~~~~~~_~~'~~~~-_?'~'/~'~~~~~'_'~~~~_~~~_~~~ Thomas J. Schoenberg Graduate Research Assistant Approved by . to. ~C_£~)~~=~~t __~b~',~£==~=iC.~~~<?_~~ Clait E. Braun (1~i Wildlife Research Leader _ �103 April 1981 JOB FINAL REPORT Colorado State of Project W-37-R-34 No. 9 Work Plan No. Job Title: Game Bird Survey ---- Population Job No. Dynamics 5 and Habitat Relationships of Blue Grouse Period Covered: Personnel: 1 April 1975 through 31 March 1981 L. Alexander, T. Beck, R. Binford, D. Benson, C. Bonney, J. Brache, C. Braun, B. Cade, L. Carpenter, A. Chappell, J. Claassen, R. Clippinger, J. Corey, D. Covic, B. Dupire, K. Duncan, D. Freddy, H. Funk, J. Gerrans, K. Giesen, W. Heicher, D. Hoart, R. Hoffman, J. Kautz, W. Larrick, D. Luce, B. McCloskey, S. McEllin, R. Oakleaf, J. alterman, L. Rottman, M. Sellitto, B. Sigler, M. Smith, L. Spess, S. Steinert, L. Strong, W. Woodard, Colorado Division of Wildlife. ABSTRACT Investigations concerning the effects of hunting on blue grouse (Dendragapus obscurus) populations, stability of breeding population levels, and habitat relationships of blue grouse were initiated in 1975 and continued through 1980 on 2 areas in northwestern Colorado. Data obtained from breeding and production surveys over the past 6 years indicate that the populations under investigation were stable and reproductively healthy. Ecological density of territorial males at Green Mountain and Eiby Creek ranged from 7.7 to 9.2 ha/male and 8.8 to 10.1 ha/male, respectively: The sex ratio in the breeding population . approximated 1:1. Evaluation of 8 nests located since 1975 indicated an average clutch of 6.0 eggs. Egg loss reduced the hatch by 33.3% (66.7% hatching success). Overall, eggs in 50% of the nests hatched successfully, 30% were partially successful, 10% were destroyed by predators, and 10% were deserted. Peak of hatch varied by as much as 3 weeks over the 6 years of study, but most eggs hatched within a 25 day period from 18 June to 12 July. Estimated nesting succ es s of females based on wing analyses was 69.3% (range 59.8-76.4%) for Middle Park and 65.7% (range 56.7-80.9%) for the Eagle area. Adult hens exhibited less variation and an overall higher success than yearlings. Brood survey!? resulted in complete counts of 310 broods from 1975 to 1980. Yearly average brood size ranged from 3.9 to 4.9 for Green Mountain and 3.4 to 5.5 for Eiby Creek. June to August mortality of chicks was much greater at Green Mountain (45.8%) than at Eiby Creek (26.4%). Production and mortality estimates indicate there are more chicks produced than necessary to replace natural losses in the breeding population. Consequently, �104 there were surplus birds available each fall and in most years the surplus exceeded 40%. Hunter activity and harvest was monitored throughout the study. Only a small portion (3.9%) of the fall population was removed by hunting compared to what could be safely harvested (22.5%) without adversely affecting the subsequent spring population. An operations plan for implementing a statewide wing collection program to monitor grouse populations for management purposes is presented. Examination of 137 crops collected at check stations resulted in identification of 45 genera of plants and 5 orders of insects in the fall diet of blue grouse. Vegetation maps were prepared for both study areas and characteristics of breeding, nesting, and brood habitat were described. �105 POPULATION DYNAMICS AND HABITAT RELATIONSHIPS OF BLUE GROUSE Richard W. Hoffman Blue grouse are the most widespread member of the grouse family in Colorado occurring in varying densities over more than 51,800 km2 of diverse habitats and terrain throughout the state (Rogers 1968). Furthermore, blue grouse rank first in population numbers and annual harvest among resident grouse species. Colorado has had a history of conservative seasons on this mountain grouse in spite of its abundance and wide distribution. This conservatism has resulted from: (1) lack of data upon which to base management recommendations, and (2) the misconception that hunting is a major mortality factor on blue grouse populations. Data presented in this report concern these problems and represent results of population studies initiated in 1975. P. N. OBJECTIVE Major objectives of this study are to: (1) increase the harvest of blue grouse in Colorado without harm to breeding populations in subsequent years, (2) identify differences in breeding densities due to differing habitat types, (3) document the stability of breeding densities over time, and (4) prepare a management plan for the design and implementation of a statewide wing survey using volunteer wing collection stations. METHODS AND MATERIALS The initial year of investigation was primarily devoted to selecting study areas and developing and testing new or existing techniques for censusing, capturing and marking blue grouse. Potential study areas were located by searching for concentrations of grouse. To assist in this endeavor, a blue grouse observation form was mailed to Colorado Division of Wildlife and U.S. Forest Service personnel working in mountainous portions of the state. Useful information was also obtained by personal communications with Division personnel and landowners. Areas where both breeding grouse and females accompanied by chicks were consistently observed were investigated to ascertain (1) density of grouse utilizing the area, (2) variations in structural and floristic characteristics within and between areas, (3) accessibility, (4) availability to hunters, and (5) hunter use. Grouse were located by systematic search with the aid of a pointing dog and by accoustical census. Accoustical census makes use of blue grouse behavior and involves playback of hen "cackle" or chick "distress" calls (Stirling and Bendell 1966). A portable, lightweight, battery powered, Norelco Model 150 Carry Corder utilizing cassettes was used for this purpose. Tape recorded calls were most effective in locating territorial males (spring, hen cackle call) and females with chicks less than 5 weeks old (summer, chick distress call). Systematic search of the habitat with the dog was the most successful method for finding hens and non-territorial males during the breeding season, hens with broods older than 5 weeks, unsuccessful hens, and fall flocks. �106 Once located, birds were first observed through 7 x 50 binoculars to count numbers present and ascertain sex. Attempts were made to capture all unmarked grouse. Several methods of capture were tested including: (1) noose poles (Zwickel and Bendell 1967a), (2) mist nets (Schladweilder and Mussehl 1969), (3) drive traps (Tomlinson 1963), (4) funnel traps (Henderson 1960), and (5) territorial traps (Bray et al. 1975). With the exception of noose poles, the other methods were considered impractical or ineffective for field use. However, a 4-ply, 10 cm black nylon mist net measuring 3 x 9 m was used throughout the study to capture incubating hens. This was less disruptive of the nest than trying to noose the hen while she was still sitting on the eggs. All grouse captured were weighed, aged (Braun 1971, Zwickel and Lance 1966, Redfield and Zwickel 1976), sexed (Braun 1971, Caswell 1954a) and banded. Pesola scales accurate to within ± 5 gm were used to weigh grouse less than 1000 gm; grouse exceeding 1000 gm were weighed with a pesola scale accurate to within + 10 gm. Captured grouse older than 6 weeks were individually marked with color combinations of anodized, aluminum, butt-end leg bands (size 14) following the banding scheme described by Gullion (1965). Birds less than 6 weeks old were marked with individually number pa tagium tags. A check station was operated in Middle Park by regional and research personnel along Colorado Highway 9 at the Prairie Point Campground from 1977 to 1979. In addition, volunteer wing collection stations (Hoffman 1975) were available to hunters each year at selected locations in Middle Park and Eagle County. A few hunters were contacted opportunistically in the field and some wings were obtained from the mail survey. The check station was operated opening weekend from about 1000 to 1800 MST, depending upon traffic load. All vehicles were stopped on the highway, but only hunters were directed to pull off at the check station. Data obtained per party included: county of origin, number of hunters, hours hunted, birds observed, birds bagged per hunter, area hunted, birds shot but not retrieved, and location where each bird was harvested. One wing was removed from each bird provided no wings had been deposited in barrels. Whenever a bird was missing 1 or both wings, the hunter was questioned as to what he did with the wings. Thus, the status of all wings was recorded as follows: (1) hunter disposed of wings, (2) hunter deposited wing in barrel, and (3) wing collected at check station. Sex by gonadal inspection was ascertained for immatures ·whenever possible. Ovaries were collected from adult and yearling hens when present. Whole body weights were obtained for all uneviserated grouse. Early fall food habits were studied by examlnlng the contents of crops collected during check station operations. All crops were labeled as to date, location, sex and age at time of collection and placed in frozen storage. Contents of each crop were later removed and representative samples of every food item were placed in small envelopes. Samples were subsequently dried for identification. Food samples were identified by aid of a disecting microscope, reference collections, plant and insect keys, and field comparisons. �107 Vegetative classification of the study areas was done in accordance with procedures outlined by Kuchler (1955) which are discussed in a later section entitled "Habitat Investigations". This method of vegetation mapping is essentially a derivation of Braun-Blanquet's (1951) system of floristic and physiognomic description of plant communities. Vegetation maps have been prepared elsewhere in Colorado using this method (Medin 1962, Braun 1969). STUDY AREAS Research was conducted on 2 study areas of differing habitat types in northwestern Colorado (Fig. 1). These areas are referred to as Green Mountain and Eiby Creek. Green Mountain is approximately 19 km south of Kremmling, Colorado in portions of Grand and Summit counties. Area investigated is in T2S, R80W, parts of sections 3, 10, 11, 14 and 15. While small portions of the area are under private ownership, the majority of this area is natural resource lands administered by the Bureau of Land Management and U.S. Forest Service. Topography of the area is irregular varying from steep slopes and ridges to gently rolling expanses hundreds of hectares in size. Cliffs and ledges are common. Maximum relief is about 330 m; elevations range from 2,539 to 2,868 m above sea level. No well delineated streams occur on the study area, where most runoff drains directly into the Blue River. Holt (1961) and Taggart (1962) discuss the geologic features of the lower Blue River and Mt. Powell areas which includes portions of the study area. Located 10 km north of Eagle, Colorado, the Eiby Creek study area is located in T3S, R84W, parts of sections 31 and 32, and T4S, R84S, parts of sections 4, 5 and 6. A small portion of the area is under jurisdiction of the Bureau of Land Management with over 95% of the area under private ownership. Topography is characterized by a flat valley floor that rises abruptly on the southwest and northwest sides. Cliffs and ledges occur along the valley rim. Above the rim, terrain quickly changes into a gently rolling upland. Maximum relief is about 400 m with elevations ranging from 2,500 to 2,900 m above sea level. Major drainage is to the south via Eiby Creek which flows into the Eagle River 10 km south of the study area. Numerous small streams occur throughout the study area and empty into Eiby Creek. Geology of this region has been described by Stauffer (1953). U.S. Weather Bureau precipitation and temperature data were available from Green Mountain Dam (elevation 2,360 m) approximately 1 km southwest of the study area. Weather records for Eiby Creek were from the Eagle County Airport (elevation 1980 m) about 12 km southwest of the study area. These data are summarized in Table 1. No wind measurements were available, but prevailing winds are primarily westerly with frequent high velocities in winter and spring. Eiby Creek supports slightly higher temperatures but lower annual precipitation than Green Mountain. Climate of both areas is typically continental with seasonal and sometimes daily variations in temperature, precipitation, and wind velocities. �MOFFAT ROUTT \JACKSON RIO BLANCO GARFIELD ..- o 00 • Fig. 1. Location of blue grouse study areas, northwestern Colorado. �109 Table 1. Temperature and precipitation, year averages from 1966 to 1976.a Temperature (OC) Month January February March April May June July August September October November December Mean Annual a - Eagle Precipitation (em) Eagle and Green Mountain, 10 Green Mountain Precipitation Temperature (OC) (em) 7.2 - 4.0 - 1.5 5.2 10.9 15.2 19.2 l3.9 12.9 3.7 - 1.0 - 8.0 2.4 1.2 1.9 2.1 1.1 2.7 3.6 1.8 2.6 3.2 1.2 2.7 - 7.1 - 6.7 - 1.9 3.3 9.3 13.3 16.8 16.2 11.7 2.6 - 1.3 - 7.0 2.9 2.2 3.8 3.6 3.2 4.4 3.8 3.3 3.2 3.8 2.9 3.0 4.9 26.5 4.1 40.1 Data from: u.S. Department of Commerce, Weather Climatological Data, Colorado Annual Summaries. Bureau. Green Mountain lies in the southwest corner of Middle Park, one of 4 major intermountain parks in Colorado. Unlike intermountain parks characterized by broad, rolling grass (South Park) or sagebrush (Artemisia spp.) (North Park) covered floors, Middle Park is mountainous and locally heavily forested; however, sagebrush types still predominate. The study area includes portions of both the sagebrush and Douglas-fir (Pseudotsuga menziesii) vegetation zones as described in Harrington (1964). Scattered to dense stands of Douglas-fir predominate above 2,700 m and extend downward to 2,600 m. Open areas below this elevation are vege-. tated primarily with big sagebrush (A. tridentata). Intermediate areas are best described as a mixed conifer-aspen (Populus tremuloides)-shrub association. This zonal pattern of vegetation is not distinct as extensions of various sagebrush types occur throughout the Douglas-fir zone creating a patchwork of vegetation types. Canopy coverage for the entire study area is about 40%. Seventy percent of the crown cover is Douglas-fir, 27% aspen, and 3% Rocky Mountain juniper (Juniperus virginiana) and mountain maple (Acer glabrum). Understory cover consists of a mosaic of shrub and grass-forb types. Snowberry (Symphoricarpos spp.), common juniper (J. communis), rose (Rosa acicularis), chokecherry (Prunus virginiana) and ser~iceberry (Amelanchier-8pp.) are the major understory shrubs; yarrow (Achillea lanulosa), pussy toes (Antennaria spp.), northern bedstraw (Galium boreale) and peavine (Lathyrus spp.) are the major forbs encountered in the understory, and sedge (Carex spp.) and bluegrass (Poa spp.) are the major grass and grasslike species. --- �110 Unforested areas constitute about 60% of the study area of which in excess of 90% is dominated by sagebrush. Other shrubs, forbs, and grasses (or grasslike) commonly found in association with sagebrush include: snowberry, rabbitbrush (Chrysothamnus spp.), serviceberry, rose, bitterbrush (Purshia tridentata), pussy toes, arrowlead balsamroot (Balsamorhiza sagittata), paintbrush (Castelleja spp.), sulphur flower (Eriogonum umbellatum), common lupine (Lupinus argenteus), bedstraw, sedge, wheatgrass (Agropyron spp.), Junegrass (Koleria cristata), needlegrass (Stipa spp.) and bluegrass. Plant nomenclature is according to Weber (1972) and Nelson (1969). Eiby Creek falls within a transition belt between the Pinyon-Juniper and Spruce-Fir zones and consists of a mosaic of mountain-shrub, aspen, sagebrush and riparian communities. Coniferous types are virtually absent within this belt, except in a few localized areas. Extreme variations in slope, aspect, soil type and moisture availability contribute to the complexity of vegetation on the study area. Open to dense stands of aspen are scattered throughout the area and occur in various growth forms from scrub thickets to over-mature, tall stands with a poorly developed shrub layer. The largest, continuous stand encompasses about 80 ha. Most stands range in size from 2 to 10 ha and occur in small patches that are separated by unforested areas alternately dominated by sagebrush, serviceberry, chokecherry, and snowberry. Thinleaf alder (Alnus tenuifolia), willow (Salix spp.), aspen, and redosier dogwood (Cornus stolonifera) form fingers of dense thickets along the stream courses. Approximately 45% of the area was classified into. forested types. Canopy cover was estimated at 32% of which 91% was aspen, 7% alder, and 2% balsam poplar (Populus balsamifera) and Rocky Mountain juniper. The dominant understory shrubs are snowberry, currant (Ribes spp.), chokecherry, serviceberry, and rose. Conspicuous herbs are yarrow, sweet cicely (Osmorhiza spp.), meadow rue (Thalictrum spp.), dandelion (Taraxacum officinale), peavine, northern bedstraw, and chickweed (Stellaria spp.). Grasses most commonly encountered are bluegrass, brome (Bromus spp.), and wheatgrass. The remainder of the study area (55%) is an assemblage of shrub types mainly dominated by sagebrush and secondarily by serviceberry and chokecherry. Rabbitbrush, snowberry and rose commonly grow in association with these types. Many of the forbs and grasses previously mentioned as being abundant in the forested types also frequently occur in unforested areas. Other notable forbs and grasses (or grasslike) include: arrowleaf balsamroot, common lupine, fleabane (Erigeron spp.), larkspur (Delphinium spp.), sulphur flowe~ geranium (Geranium fremontii), mint (Agastache spp.), violet (Viola spp.), bluebells (Mertensia spp.), sedge and Junegrass. �111 Several factors have markedly influenced the composition and structure of vegetation on both study areas. Between 1880 and 1890, the northern portion of the Green Mountain study area was set on fire by a local homesteader to create a source of firewood. Much of the area burned has reverted to sagebrush with charred logs, stumps, and patches of Douglasfir dotting the landscape. Around 1930 the Douglas-fir was selectively logged as a local source of ties for constructing the railroad. Few trees were removed, but enough to enhance development of the understory. Both study areas were subjected to extensive disturbance i.n the early 1960's when sagebrush dominated areas were sprayed with 2,4-D to enhance grass production for livestock. The degree of mortality in shrub species was not uniform throughout the sprayed areas, thus, vegetative composition and structure were altered more at some sites than others. Adequate time has since elapsed for the sagebrush to recover except in those areas where spraying was most effective. Dead or partially damaged sagebrush and serviceberry plants were still an important component of ground cover in such areas. Perhaps the greatest impact on the vegetation at Eiby Creek and to a lesser extent on Green Mountain has been excessive livestock grazing. Eiby Creek is still heavily grazed while Green Mountain receives only light summer use by cattle mostly at the lower elevations near available water. Regeneration of aspen on the Eiby Creek study area has been seriously impeded or virtually eliminated and in some places the aspen stands are deteriorating and reverting to a brushland or grassland type. Mule deer use both areas from late spring until early winter and elk are commonly found at Eiby Creek in spring and early winter. Besides grazing, no other use is presently made of either area. RESULTS AND DISCUSSION Breeding Survey Timing Blue grouse were first observed on the breeding range at both study areas in early April in all years with most early arrivals being males. Few females were present during the first 2-3 weeks of April; thus, male display was much curtailed, limited to the predawn hours, and associated more with territorial establishment than sexual responsiveness. The frequency and intensity of displays increased about the 3rd week of April which coincided with an increase in the number of hens observed on the study areas. Activities associated with breeding and density of breeding grouse peaked in early May. Timing of breeding events was similar in all years except 1977 when activities were advanced about a week. �112 Display by males was concentrated in the early morning period between 0430 and 0600 MST. Occasional display occurred after dusk, but was sporadic and involved fewer birds. Few males displayed after late June early July, although flutter flights were heard into early August. Displays Display of the male blue grouse consisted mainly of flutter flights, strutting and hooting. These displays have been described by several authors (Blackford 1963, Rogers 1968, Hjorth 1970, Stirling and Bendell 1970, Harju 1974). Flutter flight as used here is analogous to the flutter flight described by Harju (1974) and included either (1) jumping into the air with wings beating and turning a half circle before landing in full display, or (2) flight from ground to tree, or tree to tree with loud crackling wing beats at the termination of the flight. The full display or strutting position consisted of the tail being raised to the vertical and fanned to a complete half circle; wings extended slightly out and down so the wing tips dragged on the ground; eye combs and gular sacs expanded and engorged with blood so they became deep red; and the white rosette of cervical feathers around the gular sacs extended until forming a complete circle. Several variations of this display were observed, especially when the bird was hooting. During hooting the wings were often flat against the body. The white cervical feathers were seldom fully extended and the gular sacs were only partially visible. The tail was generally in the normal position and unspread, but at times it was raised and not fanned, raised and fanned, or fanned partially and not raised. Hooting was the 5 note type characteristic of the dusky race (~. ~. obscurus). Pulsations could be seen in the neck when the bird was hooting. Males selected open sites within their territories for display performance and could usually be located in the early morning at their favorite display spot. However, their display activities were not confined to these selected spots. When a hen was heard or seen, the male would frequently strut or fly in her direction. Males would sometimes display in trees, but usually only if they were disturbed on the ground or aroused while feeding or roosting in a tree. Flutter-flights were the most commonly heard and observed type of display. Flutter-flights, as noted by Caswell (1954b) and Harju (1974), had a chain reaction response that triggered flutter-flights from surrounding territorial males. It is believed that 1 function of the flutter-flight was to proclaim territoriality to other males. Flutterflights also served as a long distance display to attract hens to the territory and advertise the presence of a territorial male. There were some mornings when flutter-flights could be heard as far as 500 m. �113 Hooting was considered a close display designed to attract and stimulate the hen. Whenever a male was observed hooting there was usually a hen nearby. Males could be stimulated to hoot by the play-back of a tape recorded hen call at distances less than 50 m. At distances greater than 50 m males initially responded with a wing flutter. Often the male would move towards the source of the call in the strutting position stopping occasionally to perform wing flutters. As he approached closer, he stopped wing fluttering and started hooting. If disturbed, he would fly to a tree and continue hooting. Hooting was variable in audibility, but seldom could be heard over 30 m. Harju (1974) found no evidence to suggest that hooting served to proclaim territory and intimidate intruders or neighboring males. Communal display, described by Blackford (1958, 1963) and Harju (1974), was not observed on Green Mountain or at Eiby Creek. There were several instances when 5 or 6 males became concentrated in a small area where several hens were present. However, none of the males crossed territories of adjacent males to display collectively as a group. Little active display was noted by hens other than cackle calls and wing sounds. There were undoubtedly more displays performed by hens, but due to their secretive nature and large home ranges, fewer hens than males were encountered in the field. When disturbed they either sat motionless on the ground or in a tree, or else they flushed out of sight. Cackling was the main call produced by hens which stimulated males to flutter or hoot depending on how close they were to each other. This call was referred to as the "quaver cry" by Stirling and Bendell (1970) and was the chief call performed by hens in Wyoming (Harju 1974). A call resembling the precopulatory "whinny" described by Stirling and Bendell (1970) was heard only once and may be a less important part of dusky grouse display than the cackle call. Hens also produce a type of wing flutter when landing, which often initiated a response from nearby males. The wing flutter was not the same as that performed by males. Hens landed with a constant wing beat, while males terminated their flight with a more distinct, louder flapping. Territoriality Blue grouse males on the study areas were territorial and returned to the same territory year after year, although the boundary of the defended area occasionally changed. New territories were located in some years, but for the most part, the same territories were occupied each year. However, not all known territory locations were occupied by a male every year. A few yearling males were successful in establishing territories, but most yearling males captured and marked during the breeding season were classified as nonterritorial. Only 1 adult male, captured on Green Mountain, was not territorial. �114 Territory size was determined for 16 banded males identified at least 5 times each in 1 breeding season. Territory size ranged from 1.2 to 1.9 ha, with a mean of 1.5 ha , Harju (1974) reported a mean size of 0.7 ha (range 0.4-1.1) per territory in Wyoming. Bendell and Elliott (1967) found that the sooty grouse (.Q. ~. fuliginosus) occupied territories of 0.4-0.8 ha in a dense population and 1.6-2.8 ha in a sparse population. Martinka (1972) reported a mean territory size of 0.8 ha (range = 0.4-1.5) for blue grouse in MOntana, while Boag (1966) found that male blue grouse in Alberta occupied territories of 0.22-0.92 ha. Density Total number of territorial males recorded during the peak of breeding activities was used as an index to population size. Hens could not be accurately censused during the breeding season because of (1) greater daily movements, (2) more secretive habits, and (3) lower response rate to recorded calls. Since the study areas differed in terms of habitat type, ecological density rather than total males counted was used for comparison of densities. Any area on the breeding range where grouse were observed was considered habitable in the estimation of ecological density. For Eiby Creek and Green Mountain, the estimates were 291.6 ha (60.5% of total area) and 146.7 ha (80.9% of total area), respectively. The number of territorial males and ecological density of breeding males over a 6-year period at Green Mountain and 5-year period at Eiby Creek varied (Table 2). Breeding surveys were not conducted at Eiby Creek until 1976. Both populations have been relatively stable with gradual increases. A portion of the increase can be attributed to improved observer efficiency. Ecological density of males was slightly higher on Green Mountain than at Eiby Creek, but the difference was not significant (~. > 0.05). Table 2. Numbers at Green Mountain Area Size (ha) and ecological density of territorial and Eiby Creek, 1975-80. No. territorial males 1975 1976 1977 1978 1979 1980 male blue grouse Ecological density (ha/bird) 1975 1976 1977 1978 1979 1980 Green Mtn. 181.3 16 16 17 19 19 18 9.2 Eiby Creek 482.0 NDa 29 29 30 32 33 ND 9.2 8.6 7.7 7.7 8.1 10.1 10.1 9.7 9.1 8.8 aNo data. Number of hens on the breeding range could not be accurately ascertained from the breeding survey. It was possible to estimate the number of hens on the study areas just after most broods had hatched. At this time, hens were responsive to a tape-recorded chick distress call and were easily located. Data are only presented for Green Mountain �115 (Table 3) for 1976-80 because its smaller size and isolated nature from surrounding breeding populations made it easier to search and the hens observed were probably residents. Data from 1975 are excluded because less time was spent searching for broods than in other years. Table 3. Population estimates No. of females Year Succ 1976 1977 1978 1979 1980 13 17 15 9 12 Unsucc Totals 4 4 5 7 6 of blue grouse on Green Mountain, No. of males Nonterr Totals Terr 16 17 19 19 18 17 21 20 16 18 3 3 4 4 4 19 20 23 23 22 Male: Female 1976-80. Spring population 1.1: 1 1.1: 1 1.2: 1 1.4: 1 1.2: 1 36 41 43 39 40 Some hens and non territorial males undoubtedly escaped detection in all years, therefore, the total population estimate is minimal. The indicated sex ratio did not deviate from 1:1 (~ > 0.05). Data collected in other studies similarly suggest a 1:1 sex ratio in the spring population (Bendell et al. 1972, Zwickel 1972). Nesting Parameters, Hatching Dates and Production Natality Few nests were located during the study due to the difficulty in finding nesting hens and because no concerted effort was made to conduct nest searches on either study area. All nests were found while performing other duties. Twelve nests were located from 1975 to 1981. Clutch size and hatching success was determined for 10 nests (Table 4). The other 2 nests were found about 5-6 weeks after hatching and there were few egg shells remaining to determine the clutch size or fate of these nests. Sample size was too small for comparisons among years or' between age classes. Table 4. Natality 1975 to 1980. No. Nests 10 Clutch sizea Mean Range 6.0 4-8 rate of blue grouse based on 10 nests examined Total 54 Unhatched 6 No. of eggs DepreDedated serted 10 aBased on 8 nests for which total clutch 2 from Hatching success Hatched 36 (%) 66.7 size was ascertained. �116 Egg loss, including 1 nest that was deserted before laying was completed, reduced number of eggs hatching by 33.3%. Boag (1966) reported an approximate 18% egg loss of blue grouse in Alberta, whereas Zwickel (1975), reported what he considered a conservative estimate of 54% hatching success or a maximum estimate of 46% egg loss. Overall, eggs in 50% (5) of the nests examined hatched, 30% (3) of the nests were partially successful, 10% (1) were destroyed by predators and 10% (1) were deserted. Hatching Dates Time of hatching for blue grouse in the Eagle and Middle Park areas varied by year (Fig. 2). Time of hatch was similar between areas, so the data were grouped. All chicks were classified as to age following procedures outlined by Zwickel and Lance (1966). The estimated age was then adjusted for bias as described by Redfield and Zwickel (1976). Peak of hatch varied by as much as 3 weeks over the 6 years of study, but most chicks hatched within a 25 day period from 18 June to 12 July. Examination of the hatching curves show that no 2nd peak, indicative of renesting, was evident in any year. This does not imply renesting did not occur, but the contribution of renesting to production of young was small. Nesting Success Since hens with broods were easier to locate than those without, inflated estimates of nesting success were calculated from field observations. Consequently, analyses of the wing molt pattern of hunter harvested birds was used as the best estimate of nesting success (Braun 1971). The technique provides a minimum estimate as some successful hens will have completed their molt by mid- to late September and cannot be distinguished from unsuccessful hens. This problem is compounded during years when hens initiate nesting earlier, such as in 1977. Few yearlings were identifiable in 1977 and many adult hens were in the advanced stages of molt. Therefore, nesting success was probably better in 1977 than what the data indicate (Table 5). With the exception of 1979, combined nesting success of adult and yearling hens in Middle Park was relatively constant from year to year. Nesting success varied more within and between age classes than noted for combined success of both age classes. Adult hens exhibited less variation (66.7-80.9%) and an overall higher success (x = 73.3%) than yearlings (x = 57.1%, range 33.3-82.1%). Reasons for this difference are uncertain, but may be an artifact of sample size. Zwickel (1975) detected no significant difference in nesting success of yearling and adult hens. Overall, 59.8% of the adult and yearling female wings collected in Middle Park in 1979 were classified as from successful nesters. This represents the lowest estimate obtained from wing samples over the 6-year period and was attributed to (1) a late nesting season, and (2) adverse weather conditions during the nesting period. However, even under these less than optimal conditions, nesting success was still good. Based on field observations of successful (9) and unsuccessful (7) hens in 1979, estimated nesting success on Green Mountain was 56.2%. �~_17 70 1975 1976 1977 1978 1979 1980 60 (!) z 50 • • 0 0 .-----. 0-----0 K c ~ n I U ~ I40 • r- 1\ z w u I I a:: W 30 o, / I I I / 20 / / ,• / <, / / 10 <, <, "' / •I . <, <, I <, <, I 1-7 JUNE 8-14 15-21 22-28 29-5 WEEK OF HATCHING Fig. 2. Blue grouse hatching curves. 197."i-RO. __ "...• 6-12 JULY - "'" 13-19 20-26 �118 Table 5. Estimated nesting success Year Middle Park Estimated nesting successa YearTotals lingsb 1975 1976 1977 1978 1979 1980 80.3 (71) 68.3 (60) 68.5(143) 80.9(115) 66.7(150) 77.5(142) 33.3 82.1 68.7 65.2 42.4 59.2 Total 73.3(681) 57.1(219) aTotal (21) (28) (16) (46) (59) (49) Estimated Adults 1975-1980. Eagle nesting successa YearTotals lingsb 69.6 (92) 72.7 (88) 68.6(159) 76.4(161) 59.8(209) 72.7(191) NDc ND 57.1 (63) 8l.0 (58) 67.2 (64) 58.7 (70) ND ND c IS (4) 80.8 (26) 65.1 (43) 50.0 (20) ND ND 57.1 (63) 80.9 (84) 66.4(107) 56.7 (90) 69.3(900) 65.5(255) 66.3 (89) 65.7(344) sample of wings examined b (%), Middle Park and Eagle, in parentheses. Due to small sample sizes in some years, estimated may not be representative of this age class. nesting success cNo data. dInsufficient sample. Few wings were collected from the Eagle area in 1975 and 1976. In 1977, only 4 yearlings were identifiable so nesting success was based only on the sample of adult hens. From 1978 to 1980, nesting success of adults and yearlings was similar, although being slightly higher for·adults. This agrees with the data reported by Zwickel (1975), but not with the Middle Park data. However, yearly variations in nesting success had no known impacts on the population at either study area. There were always more chicks produced than necessary to replace natural losses in the breeding population. Adult Turnover Estimated turnover (based on the 5-year average from wing analyses) of adult male and female blue grouse was 23.9 and 27.5%, respectively (Table 6). These estimates are based on the assumption that in a stable population percent yearlings must equal annual loss of adults. These data suggest differential survival favoring males, but the difference is non-significant (P > 0.05). Zwickel (1965) banded and recensused 70 male and 68 female blue grouse on a breeding range on Vancouver Island. He found the mean annual death rate to be 27% for males and 38% for females. Bendell and Elliott (1967) reported estimates of 28 and 33% for male and female blue grouse, respectively. Both studies concluded that death rates did not vary significantly between sexes. �119 Yearly estimates varied more for males than females because of the compounded problem of separating yearling and adult males. Most likely adult males have a constant annual turnover rate as' shown by females (Table 6). Overall the estimated annual turnover rates (5-year averages) are probably lower than the actual turnover rates, again, as some year·lings were undoubtedly assigned to the adult age class. This problem was more pronounced for males than females and accounted for the lower turnover rates calculated from wing analyses compared to the estimates given by Zwickel (1965) and Bendell and Elliott (1967). Sample sizes were too small for similar calculations using the data collected from the Eagle area. Table 6. Park. Year a 1975 1976 1978 1979 1980 Estimated Males Yearlings Adults N 30 68 121 131 162 5-year average turnover of adult male and female blue grouse, Middle % 85.7 81.9 77 .1 61.5 87.6 N 5 15 36 82 23 76.1 Production and Juvenile Totals % 14.3 18.1 22.9 38.5 12.4 23.9 aData from 1977 excluded yearlings. Females Yearlings Adults 35 82 157 213 185 N 41 64 116 151 146 % 73.2 69.6 71.6 71.9 74.9 N 15 28 46 59 49 72.5 because of the unrepresentative % Totals 26.8 30.4 28.4 28.1 25.1 56 92 162 210 195 27.5 sample of Mortality Brood surveys conducted from mid-June to mid-August resulted in complete counts of 310 broods (Green Mountain = 151, Eiby Creek = 159) from 1975 to 1980 (Table 7). Broods were counted more than once, but not within the same month. Average brood size was similar between areas in 1977, 1978 and 1980. The apparently smaller brood size at Eiby Creek in 1976 was attributed to a larger sample of broods counted in late July - early August, whereas primarily early to mid-July counts were made on Green Mountain. All evidence indicates better production and survival of young at Eiby Creek in 1979 than at Green Mountain. Rogers (1968) found average brood sizes of 4.1, 2.6 and 3.9 in 3 years of study in western Colorado. �120 Table 7. Average brood size, range and number of broods observed monthly intervals, 1975-80.a Month 1975 Green Mountain 1976 1977 1978 1979 1980 1976 by Eiby Creek 1977 1978 1979 1980 Jun Mean Range N b ND ND ND 5.3 3-7 3 5.1 4-7 11 6.2 5-9 9 7.2 6-9 5 6.3 5-7 3 5.0 4-6 2 4.6 3-6 7 6.5 6-7 2 6.0 3-9 3 5.6 4-7 7 Jul Mean Range N 5.2 2-7 4 4.1 2-6 7 4.0 1-7 10 5.4 3-7 9 4.8 1-10 12 4.8 1-9 18 3.7 1-7 9 4.1 2-9 10 4.9 3-8 13 6.5 4-10 15 4.8 2-9 20 Aug Mean Range N 3.0 1-5 6 3.0 1 2.5 1-4 13 3.7 2-6 14 3.0 1-6 11 3.7 1-5 15 2.2 1-4 5 3.6 1-10 20 5.0 3-8 17 4.3 1-7 13 3.4 1-6 16 3.9 4.4 3.8 4.9 4.5 4.4 3.4 3.9 5.1 5.5 4.4 Yearly avg. aOnly distinct broods with full counts are included. bNo data due to late hatching year or no full counts obtained. Some doubt exists as to whether blue grouse broods can be accurately counted until after they are 4 weeks old (Boag 1966), consequently late June and early to mid-July counts may be underestimated. Brood counts in August may also be misleading due to shuffling of chicks between broods, formation of gang broods, and brood breakup and dispersal. Therefore, observed differences in brood size from June to August may not accurately reflect loss of chicks during this period (Table 8). The gradual attrition in brood size over the summer is the least reliable method'of evaluating chick mortality (Zwickel and Bendell 1967b), but was the only data available for determining losses in this study. Average annual summer mortality of chicks was 45.8% for Green Mountain and only 26.4% for Eiby Creek. Production and Breeding Populations Estimated total production in relation to breeding population levels varied by year and area (Table 9). Calculations are based on the following assumptions which are supported by data collected in this and other studies: (1) (2) (3) (4) A 1:1 sex ratio exists in the breeding population. Estimated percent nesting success is correct. Mean annual brood size and monthly estimated brood sizes represent the production of chicks on the study areas. Immigration equals emigration. �121 Table 8. Apparent mortality of juvenile blue grouse from late June to mid-August at 2 Colorado. study areas, 1975-80. Area and year Mean brood size Jun Aug Mortality Green Mountain 1975 1976 1977 1978 1979 1980 1975-80 a 5.2 5.3 5.1 6.2 7.2 6.3 5.9 3.0 3.0 2.5 3.7 3.0 3.7 3.2 42.3 43.4 51.0 40.3 58.3 41.3 45.8 Eiby Creek 1976 1977 1978 1979 1980 1976-80 5.0 4.6 6.5 6.0 5.6 5.3 2.2 3.6 5.0 4.3 3.4 3.9 56.0 21.7 23.1 28.3 39.3 26.4 aMean brood size for July as no full counts were obtained due to a late hatch. (%) in June Zwickel (1965) and Bendell and Elliott (1967) reported a mean annual death rate of approximately 30% for breeding populations of blue grouse on Vancouver Island, British Columbia. A similar estimate of 39% was obtained from reobservation of marked birds at Green Mountain and Eiby Creek. Assuming this estimate is constant from year to year, an estimate of total production that is needed for minimum replacement necessary to maintain a stable breeding population can be determined (Table 10). Production by mid-August rather than total production was used because mid-August figures more closely approximates production and survival of juveniles until fall. The replacement requirement then becomes an estimate of the fall to spring mortality of juveniles that should occur in a stable population. Both populations under investigation were essentially stable. It is apparent (Table 10) that production was more than adequate to replace natural losses in the breeding population. For Green Mountain, the average annual fall to spring loss of juveniles required to maintain the population from 1975 to 1980 was 52% (62% for Eiby Creek from 1977 to 1980). These data suggest there are surplus birds in the fall population and in most years this surplus exceeds 40%. �Table 9. Estimation of the annual production of young blue grouse on Green Mountain and Eiby Creek, 1975-80. • Total a Area and year population Nesting success (%) Green Mountain 1975 1976 1977 1978 1979 1980 32 32 34 38 38 36 69.6 72.7 68.6 76.4 59.8 72.7 br eed Lng No. hens with broods Average annual brood size Total production JunAug mortality Total production by midAug Total population by midAug Percent gain 11 12 12 14 11 13 3.9 4.4 3.8 4.9 4.5 4.4 43 53 46 69 49 57 42.3 43.4 51.0 40.3 58.3 41.3 25 30 22 41 21 33 57 62 56 79 59 69 44 48 39 52 36 48 N N Eiby Creek 1977 1978 1979 1980 58 60 64 66 57.1 80.9 66.4 56.7 aExcludes nonterritorial males. 17 24 21 19 3.9 5.1 5.5 4.4 66 122 115 84 56.0 23.1 28.3 39.3 29 94 83 51 87 154 147 117 33 61 56 44 �123 Table 10. Estimates of the fall to spring loss of juvenile necessary to maintain a stable breeding population. Area and year Total breeding population Annual mortality breeding . a popu 1 atl.on Total production by mid-Aug blue grouse Fall to spring mortal itt of juveniles (%) Green Mountain 1975 1976 1977 1978 1979 1980 32 32 34 38 38 36 12 12 13 15 15 14 25 30 22 41 21 33 52 60 41 63 29 58 58 60 64 66 23 23 25 26 29 94 83 51 21 75 70 49 Eiby Creek 1977 1978 1979 1980 aNumber of birds expected to be lost from the population a constant annual mortality rate of 39%. bMortality necessary to maintain assuming the population. Harvest Hunting Season The 1980 blue grouse season in Middle Park opened on 13 September and closed on 7 October. Blue grouse hunting was also permitted between 11 October and 11 November in conjunction with the deer (Odocoileus spp.) elk (Cervus elaphus), and combined deer-elk seasons. Season length was 52 days with daily bag and possession limits of 3 and 6 birds, respectively. Season structure, bag limits and length have varied from 1975 to 1980 (Table 11). Middle Park The check station was not an efficient or economical way of collecting wings and was established primarily to monitor hunter activity in Middle Park. Since this objective of the study was completed in 1979, the check station was not operated in 1980. Instead, hunters were contacted opportunistically in the field in an effort to increase wing samples from known sex juveniles. These contacts were made on Blue Ridge, Spring Creek, Gore Pass, and Chimney Rock, major harvest areas in Middle Park. Volunteer wing collection stations were also available to hunters in 1980 from 13 September to 7 October. No attempt was made to monitor the blue grouse harvest during the 1980 big game seasons as this objective was completed in 1979. �124 Table 11. Blue grouse hunting seasons, Colorado, 1975-80. Hunting season dates Year Season length (days) Bag limit Possession limit 1975 13 Sep - 5 Oct 23 3 6 1976 11 Sep - 10 Oct 11 Sep - 10 Oct and 16-26 Oct 10 Sep - 9 Oct 30 40 3 3 6 6 30 3 6 15 Oct - 15 Nov 27 3 6 30 27 30 27 25 27 3 3 3 3 3 3 6 6 6 6 6 6 1977 1978 1979 1980 9 14 8 13 13 11 Sep Oct Sep Oct Sep Oct - 8 Oct - 14 Nov - 7 Oct - 13 Nov - 7 Oct - 11 Nov Open areas Unit 80 and all units west of Interstate 25 except portions of Unit 52 Same as 1975 Unit 28 (Middle Park) only West of Interstate 25 and Unit 80 West of Interstate 25 and Unit 80 when open to deer or elk hunting Same as 1977 Same as 1977 Same as 1977 Same as 1977 Same as 1977 Same as 1977 From 1975 through 1980, 3,922 blue grouse wings were collected from Middle Park. Wing barrels accounted for 86.2% (3,379) of all wings collected (Table 12). Number, source and location of blue grouse wings collected in Middle Park from 1975 to 1980 varied by year (Tables 12, 13). Table 12. Number and source of blue grouse wings collected in Middle Park, 1975-80. Wing barrels N % Year 120(10)a 292(13) 501(15) 875(16) 839(14) 752(12) 1975 1976 1977 1978 1979 1980 Totals Mean 65.6 84.9 82.7 87.0 88.7 89.9 Check stations Mail survey N N % % 26 49 87 105 84 74 14.2 14.2 14.4 10.4 8.9 8.8 425 3,379 86.2 0 3 18 0 23 11 0 0.9 2.9 0 2.4 1.3 55 10.8 Miscellaneous N % 37 0 0 26 0 0 20.2 0 0 2.6 0.0 0.0 183 344 606 1,006 946 837 3,922 63 1.4 Totals 1.6 a Number in parentheses represents number of wing barrels used. �125 from wing Table 13. Location and number of blue grouse wings collected barrels, Middle Park, 1975-1980. No. of wings collected Station location City Reservoir Beaver Creek Williams Fork Corral Creek Troublesome Pinto Creek Lawson Ridge Spring Creek Blue Ridge Chimney Rock Gore Pass Trough Road Kremmling Rock Creek Willow Creek Cottonwood Pass Ute Pass 1980 NB NB 28 10 80 16 23 3 169 50 136 154 85 29 4 13 60 17 0 3 161 63 197 157 74 NB NB 162 36 74 183 147 NB NB NB NB NB 57 16 19. 62 8 20 NB 17 28 60 40 306 66 53 11 639 273 587 595 380 47 5 171 24 94 839 752 3,379 1976 1977 1978 8 1 0 21 13 9 0 30 38 3 19 1 43 2 0 3 55 23 95 32 17 8 0 53 18 5 2 62 63 85 69 74 14 5 NB NB NB NB NB NB NB NB 120 Totals 1979 1975 NB 4 NB NB NB NB NB 12 26 292 501 a 875 Totals 16 49 0 16 NB 52 aNo barrel at this location. During the 3 years (1977-79) of check station operations, 705 hunters with 862 blue grouse (1.2 birds/hunter, range 1.0-1.3) were contacted. These hunters reported observing 2,068 blue grouse. Of the 705 hunters checked, 49.2% (range 39.5-54.6%) were successful. Only 37 birds were reported wounded and not retrieved for an estimated crippling loss of 4.3% (range 3.1-5.6%) (Table 14). 'Eable 14. Blue grouse harvest Year No. hunters checked No. birds observed No. birds bagged 1977 1978 1979 228 222 255 480 758 830 226 294 342 statistics, Hunter efficiency % 47.1 38.8 41.2 Middle Park, Colorado, % Successful hunters, % 39.5 54.6 53.3 Crippling loss 1977-79. % Birds per hunter 4.0 3.1 5.6 1.0 1.3 1.3 �126 Status of wings from all birds examined at check stations from 1977 to 1979 was categorized as (1) hunter disposed of wing, (2) hunter deposited wing in volunteer collection station, or (3) wing collected at check station. Results were summarized for all birds harvested in areas where a wing station was available to the hunters and yielded a measure of .hunter participation and effectiveness of wing stations for sampling (Table 15). Over one-half (54.6%) of the wings were deposited, suggesting hunters were willing to cooperate. Still, some hunters were inconvenienced by the request and simply did not bother to deposit wings. Others indicated they would have cooperated, but were unaware of the stations upon entering their hunting area and disposed of the wings before leaving. Table 15. Status of wings from blue grouse examined Middle Park, Colorado, 1977-79. Status Wings Wings Wings Wings deposited, % not deposited, % disposed, % of unknown status, % at check station, 1977 1978 1979 60.7 18.9 17.4 3.0 50.6 20.3 28.3 0.8 52.6 22.8 16.4 8.2 1977-79 54.6 21.1 20.5 3.8 Distribution of the harvest within Small Game Management Unit 28 was not uniform (Table 16). Gore Pass was the leading harvest area closely followed by Spring Creek and Chimney Rock. There were 6 areas that consistently accounted for over 75% of the harvest each year since 1977. All other areas in Middle Park each had less than 5% of the harvest. Table 16. a 1977-80. Distribution of the blue grouse harvest within Middle Park, Percent of harvest 1979 1980 Area 1977 1978 Spring Creek Gore Pass Chimney Rock Piney Blue Ridge Corral Creek 12.2 14.1 16.0 15.5 12.7 9.1 18.8 17.2 14.0 9.7 7.4 8.0 19.2 18.7 23.5 8.8 7.5 7.1 20.9 22.6 10.0 18.7 8.8 6.0 22.7 23.1 20.1 15.5 8.7 9.9 79.6 75.1 84.8 87.0 100.0 Totals aTotals included. do not add up to 100% as only major harvest areas are 1977-80 �127 Distribution of wing collections according to time period is given in Table 17. The bulk of the harvest occurred opening weekend each year since 1975. Liberalization of the season spread the harvest and hunting pressure over a longer period of time. This effect was supported by data collected in 1977, 1978 and 1980, but not in 1979. During this interval, hunting pressure essentially remained the same, but declined on opening weekend as more hunters are hunting later in the season. Of the 3,922 wings collected from Middle Park since 1975, 3,820 were classified to age and sex (Table 18). Immatures were well represented (> 45%) in the harvest each year indicating production of young was good, being better in some years than others. Identified yearlings were poorly represented in the harvest samples in most years, except possibly 1979. This was not because of poor production and subsequent low recruitment, but was because few yearlings could be separated from the adult age class. This was due to time of the onset of primary molt and whether primaries X and IX were still present at the time of collection. If these feathers had already molted, as occured with many wi.ngs examined from 1975 to 1978 and in 1980, then yearlings could not be separated from adults and were classified as adult. The problem is compounded in "early" years (1977) (Table 18) and moreso for males than females. The first birds to initiate their primary molt are yearlings (most likely nonterritoria1 males) followed by adult males, lone females (brood1ess) and brood females. From 1975 to 1980 (excluding 1979), an average of 52% of the adult and yearling males and only 9% of the females had molted P X by the opening weekend. Consequently, fewer males than females we~e identifiable as yearlings. The sex ratio of adults and immatures approximated 1:1 in all years except 1978 when there were more (f < 0.05) immature males than females in the harvest sample. Reasons for the deviation in sex ratio are uncertain, but may be an artifact of sampling. Yearlings had a balanced sex ratio in only 2 (1978 and 1979) of 6 years. From 1975 to 1977 and in 1980, there were fewer (P < 0.05) males than females in the yearling segment of the harvest. This indicates that more yearling males than females were incorrectly assigned to the adult age class. The actual sex ratio of yearlings probably approximated 1:1. The percentage of juveniles in a population is often used as an indicator of year to year turnover. Data available (Table 18) suggest that about 56% (range 46-67%) of the fall population is comprised of juveniles. Assuming an even, sex ratio of adults (including yearlings), a theoretical autumn population of 100 birds would be composed of 56 juveniles, 22 adult and subadu1t males and 22 adult and subadu1t females. At an average annual adult (and subadu1t) mortality rate of 39%, only 17 juveniles would join the subsequent spring breeding population in a stable population. About 70% of the juveniles alive in fall would then be "surplus". This may represent an inflated estimate as intensive studies on Green Mountain have shown about 52% first-year mortality among juveniles that survive to autumn. The autumn to autumn loss of juveniles probably lies somewhere between 52 and 70% indicating that (1) annual mortality of juveniles is much higher than for adults, and (2) surplus birds are always present in the fall population. These are cornnon phenomenon characteristic of most upland game bird populations. �Table 17. a Time distribution of blue grouse wings collected, Middle Park, Colorado, 1975-80. 1975 Time period N 1st weekend 1st week 2nd weekend 2nd week 3rd weekend 3rd week 4th weekend 4th week 5th weekend Experimental season, 16-26 Oct. Deer season Elk season Combined season 53 9 20 0 8 11 19 Total - regular season 1976 % 44.2 7.4 16.7 0.0 6.7 9.2 15.8 1978 1980 % N 37.5 4.0 12.6 5.5 8.8 1.4 3.6 2.2 2.4 288 99 109 13 55 13 23 10 119 32.9 11.3 12.5 1.5 6.3 1.5 2.6 1.1 13.6 375 30 65 81 48 48 18 35 39 44.7 3.6 7.7 9.7 5.7 5.7 2.2 4.2 4.6 37 63 10 7.4 12.6 2.0 67 44 35 7.7 5.0 4.0 54 25 21 6.4 3.0 2.5 96.2 391 78.0 729 83.3 739 88.1 3.8 110 22.0 146 16.7 100 11.9 100.0 501 100.0 875 100.0 839 100.0 N 121 36 36 11 23 10 21 8 15 41.4 12.3 12.3 3.8 7.9 3.4 7.2 2.7 5.2 188 20 63 28 44 7 18 11 12 11 3.8 % 1979 N % % N % 270 78 168 22 98 25 91 35.9 10.4 22.4 2.9 13.0 3.3 12.1 t-' 120 100.0 Total - big game season Total - both seasons 1977 N 281 11b 120 100.0 292 aWing collections from barrels only. bExperimental season only. N 00 752 100.0 752 100.0 �Table 18. Age and sex composition of the blue grouse harvest, Middle Park, Colorado, 1975-80. Males Adults Females Totals Yearlings Females Males Year N % 1975 30 17.7 41 24.1 71 41.8 5 2.9 197.6 68 19.9 64 18.8 132 38.7 15 1977 133 22.1 144 23.9 277 45.9 1978 121 12.5 116 11.9 237 24.4 N - % N - % % - % 15 8.8 20 11.7 4.4 28 8.2 43 3 0.5 16 2.6 36 3.7 46 4.7 N - % Totals N - Immatures Females Males Totals % - % 36 21.2 43 25.3 79 46.5 12.6 74 21.7 92 27.0 166 48.7 19 3.2 164 27.2 143 23.7 307 50.9 82 8.4 364 37.4 290 29.8 654 67.2 N N - N N % .- N 1.0 1979 131 14.2 151 16 4 282 30.6 82 8.9 59 6.4 141 15.3 242 26.3 255 27.7 497 54.0 1980 162 19.9 146 18.0 308 37.9 23 2.8 49 6.0 72 8.8 233 28.7 200 24.6 433 53.3 6-year average 16.9 17.3 34.2 4.3 5.6 9.9 29.1 26.8 55.9 �130 Eagle A total of 417 blue grouse wings was collected from Eagle County (Unit 54) during the 1980 season. Six volunteer collection stations accounted for 98.6% (411 wings) of the wings obtained. Fewer stations were available to hunters in 1980 than in previous years; however, the 6 locations sampled have been recommended as permanent collection sites for Eagle County in anticipation that the Regions will assume this responsibility in 1981. The mail wing survey provided the only other source of wings (6 wings = 1.4%). Since 1977, wing barrels have been the primary means of procuring samples of wings from the Eagle area, contributing 95.8% (1,681) of the total sample (1,754). Number of wings collected from volunteer stations over the past 4 years has varied (Table 19). Some stations were eliminated while others were relocated to more productive sites. Table 19. Location area, 1977-80. Station location Squaw Creek Lake Creek Muddy Pass Milk Creek Coffee Pot Springs Red Sandstone Eiby Creek South Eiby Creek North Brush Creek Cabin Creek Gypsum Creek Wolcott Totals and number of wings collected 1977 1978 5 5 63 23 79 57 19 NB 27 NB 19 NB a NB NB 68 73 67 63 26 78 28 5 8 38 297 454 from wing barrels, No. wings collected 1980 1979 Eagle 1977-80 NB NB 83 78 69 52 14 59 30 8 44 82 NB NB 87 80 73 59 NB 52 NB NB NB 60 5 5 301 254 288 231 59 189 85 13 71 180 519 411 1,681 aNo barrel at this location. All but 12 of 417 wings collected in 1980 were classified to age and sex (Table 20). Immatures comprised over 50% of the harvest each year suggesting good production or differential vulnerability. Production of young varied from year to year. Few yearlings occurred in the harvest with the exception of the late nesting year of 1979. Therefore, in most years, the proportion of yearlings in harvest samples could not be used as an indicator of recruitment nor as an approximation of annual turnover in the adult segment of the population. �Table 20. Age and sex composition of the blue grouse harvest, Eagle area, 1977-80. Adults Females Males Year N 1977 87 25.8 1978 62 1979 1980 4-year avg. % Yearlings Females Males Totals Immatures Females Males % N 65 19.3 152 45.1 10 3.0 4 1.2 14 4.2 81 24.0 13.4 59 12.8 121 26.2 9 2.0 26 5.6 35 7.6 153 75 14.4 64 12.3 139 26.7 38 7.3 43 8.3 81 15.6 74 18.3 70 17.3 144 35.6 8 2.0 20 4.9 28 6.9 17.3 N Totals 15.0 % 32.3 N % 3.7 N % 5.4 N % 9.1 Totals % N % 90 26.7 171 50.7 33.1 153 33.1 306 66.2 153 29.4 147 28.3 300 57.7 118 29.1 115 28.4 233 57.5 N % 29.3 N 29.3 58.6 •....• w •....• �132 Sex ratio of adults and imrnatures approximated 1:1 in all years. Yearlings exhibited an unbalanced sex ratio favoring females in 1978 and 1980, but the ratio was not different (P > 0.05) from 1:1 in 1977 and 1979. There is no reason to suspect the sex ratio. of yearlings would be different from that of adults or imrnatures. The deviations detected in 1978 and 1980 undoubtedly occurred because more yearling males than females were incorrectly assigned to the adult age class. Data from Eagle are similar to findings from Middle Park. Both populations have similar attributes and all available evidence suggest that: 1. 2. 3. 4. 5. 6. 7. 8. 9. Harvest Overall, hunting pressure and harvest are light. Present harvest levels have no measurable impact on the subsequent spring breeding population. The sex ratio at hatch approximates 1:1. The sex ratio of the spring breeding population (adults and yearlings) approximates 1:1. Yearlings are poorly represented in fall harvest samples because many yearlings have already completed their primary molt by mid-September and cannot be distinguished from adults. The proportion of yearlings in the harvest sample is not a reliable measure of adult survival rates or recruitment of young birds into the breeding population. The problem of separating yearlings from adults is more pronounced for males than females and is compounded in "early" (1977) vs. "late" (1979) years. Production of young has varied from fair to excellent and has been more than adequate to replace natural losses in the breeding population. There are surplus birds in the fall population. Rate Direct evidence from banding data and indirect evidence based on the attributes of the populations under investigation indicate that present levels of harvest have no measurable impact on blue grouse populations. Available data suggest that even though there is a high and variable loss of grouse during their first year of life, production of young greatly exceeds the number necessary to replace natural losses in the breeding population. These "excess" birds represent a harvestable resource that can be removed without adversely affecting the subsequent spring breeding population. Few birds were banded, but the percentage of banded birds shot (3.9) indicate hunters removed a negligible portion of the fall population (Table 21). Bendell and Elliott (1967) reported approximately 5% of the hens and chicks banded on their study areas on Vancouver Island were shot each year, while few (0.7%) adult males were taken. Mussehl (1960) found that hunters removed 7 (1957) and 12% (1958) of the blue grouse banded in the Bridger Mountains, Montana, most of which were juveniles. Most chicks captured in this study were too young « 6 weeks) to carry leg bands, and were marked with only patagium tags. Some of these birds were probably harvested, but because of the less conspicuous nature of patagium tags, the hunters did not notice the birds were marked. No patagium tags were returned by hunters; a factor which undoubtedly contributed to the calculated low harvest rate of juveniles. �133 Table 21. Number of banded grouse available Middle Park and Eagle, 1976-80. Year 1976 1977 1978 1979 1980 Totals Males No. % Avail. Shot Females No. % Avail. Shot to and shot by hunters, Juveniles No. % Avail. Shot 6 22 15 17 16 0.0 0.0 6.7 5.9 12.5 13 17 33 32 50 0.0 0.0 3.0 9.4 4.0 5 14 18 58 69 76 5.3 145 4.1 164 0.0 20.8 5.5 0.0 1.4 3.1, Totals No. % Avail. Shot 24 53 66 W7 135 0.0 5.7 4.5 3.7 3.7 385 3.9 Hickey (1955) states that gallinaceous birds can safely withstand a hunting kill equivalent to about one-half their annual mortality rate. Based on a 6-year average for Green Mountain (Table 9), production contributes to about a 45% increase in the population. In a stable population there must be an annual loss from fall to fall of this amount. Therefore, according to Hickey (1955), the population can absorb a harvest of 22.5%. This represents a minimum estimate because the annual mortality of adults is not considered. The calculated yield (22.5%) greatly surpasses the estimated harvest rate (3.9%). These data support the conclusion that hunting has no measurable effects on blue grouse populations. The above data provided the justification to liberalize the statewide blue grouse season in 1977 (Table 10). Season length was increased from 30 to 57 days by allowing blue grouse hunting in conjunction with the deer, elk and combined, deer-elk seasons. A total of 1,500 big game hunters were surveyed at the Idaho Springs check station,during the deer (500), elk (500) and combined deer-elk (500) seasons in order to evaluate hunter participation and response to the longer grouse season. Hunters were contacted opportunistically at the check station. Only 1 hunter from each party was surveyed and only residents were included in the sample. Sampling periods included days at the beginning, middle and end of each season. ' From data presented in Table 22 it is apparent that few big gam~ hunters took advantage of the longer grouse seasons in 1977 (6.3%) and 1978 (5.5%). Based only on those hunters aware the grouse season was open, participation was 13.1% in 1977 and 9.8% in 1978. Even though participation was low, general response of hunters to the combined blue grouse - big game season was highly favorable in both years. The primary complaint of those hunters opposed to the season was their concern about the potential increase in number of hunters (big game + grouse hunters). However, less than 1 percent of all hunters contacted during the 1977 and 1978 big game seasons were strictly grouse hunters. More hunters were aware of the longer grouse season in 1978 (55.1%) than in 1977 (48.1%), but participants remained relatively constant. Therefore, only a small percentage of hunters were interested in hunting grouse during the big game season. �Table 22. Combined 1977 and 1978. blue grouse-big game season hunter questionnaire data, Idaho Springs, 1977 Deer Elk Colorado, 1978 Combined Total Deer Elk Combined Total No. in sample 500 500 500 1,500 500 500 500 1,500 No. big game hunters aware of season 253 260 210 723 259 285 283 827 % big game hunters aware of season 50.6 52.0 41.8 48.1 51.8 57.0 56.6 55.1 No. big game hunters hunting grouse 40 25 30 95 14 37 31 82 % big game hunters hunting grouse 8.0 % big game hunters aware of season that hunted grouse Total grouse observed % big game hunters observing grouse Grouse observed/big game hunter Total grouse harvested Grouse/big game hunter hunting grouse 5.0 6.0 6.3 2.8 7.4 6.2 5.5 w .f:- 15.8 693 9.6 2,102 14.3 620 13.1 3,415 5.4 1,384 13.0 2,053 10.9 1,455 9.8 4,892 24.8 41.8 18.5 28.4 32.4 35.6 37.8 35.0 1.4 4.2 1.2 2.3 2.7 4.1 2.9 3.2 13 28 12 53 17 28 32 77 0.3 1.1 0.4 0.6 1.2 0.7 1.0 0.9 52.0 52.2 52.0 52.1 60.0 57.0 62.0 59.7 % negative 13 .4 9.2 10.4 11.0 14.0 11.0 9.6 1l.5 % indifferent 34.6 38.6 37.6 36.9 26.0 32.0 28.4 28.8 % hunters favorable of combined blue grouse-big game season I-' �135 Due to good production, number of grouse observed increased substantially in 1978 (4,892) compared to 1977 (3,415). Consequently, hunters that took advantage of the longer season were more successful in 1978 (48.5% success, 2.3 grouse/successful hunter) than in 1977 (27.4% success, 2.0 grouse/successful hunter). As a result, fewer hunters 'killed more grouse in 1978 than in 1977. Based on wing collections from barrels in Middle Park, 22.0 and 16.7% of the total grouse harvest occurred during the big game seasons in 1977 and 1978, respectively. Application of Wing Collection Program The collection of blue grouse wings and their subsequent analyses has important practical implications for management. Wings can be efficiently and economically collected through the operation of volunteer collection stations; thus, the potential exists to implement a data collection program to obtain population and harvest trend information on blue grouse (and other upland game birds) for management purposes. Such a program has been successfully conducted as a research function over the past 6 years to provide continued support for liberal blue grouse seasons in Colorado. With the termination of these research efforts, it has been recommended that the wing survey be expanded statewide and continued by management personnel. An operations plan for the implementation of this program has been prepared and submitted to regional and staff personnel for their approval (Appendix A). FOOD HABIT STUDIES Contents from 137 crops collected at check stations provided data concerning the early fall food habits of blue grouse. Since only representative food items from each crop were saved, results are reported in percent frequency of occurrence. Gilfillan and Bezdek (1944) reported that "frequency of occurrence is an important index of the leading foods and affords a more accurate picture of food preferences than volumetric data". However, frequency data do not lend themselves to statistical analysis even though they may be a good index to preferred foods. Table 23 presents a complete list of plants, animals, and other items recorded in the crop analyses by age and sex classes. Represented in this table are 45 genera of plants and 5 orders of insects. Of the 45 genera, 21 had a frequency of occurrence greater than 5%. Fifteen crops were empty and excluded from the data. Vaccinium spp. leaves were the most common food item, being represented in 41% of the samples. The fruit and stems of Vaccinium spp. were also heavily used. Adult grouse used this plant 2-3 times more than juveniles., Vaccinium spp. has been found in the diet of blue grouse in other studies, but not at such high frequencies. This plant was found in only 3 and 9% of the crops examined by Boag (1963) in Washington and Knapp (1962) in northcentral Colorado, respectively. Members of the family Formicidae (ants) occurred in 32% of the samples. Adults and juveniles ate ants about equally as often, but females ate these insects more often than males. Ants were common in the diets of grouse in studies by Knapp (1962) and Boag (1963), occurring in 5 and 20% of the crops examined, respectively. �136 Table 23. Foods in 122 blue grouse crops collected Colorado, fall 1975-79. Food and plant part Vaccinium spp. leaves fruit stems Taraxacum spp. leaves flowers Eriogonum spp. leaves seeds Lathyrus spp. leaves Lupinus spp. leaves seeds Amelanchier spp. fruit Senecio spp. leaves flowers Ribes spp. fruit leaves Trifolium spp. leaves Antennaria rosea leaves seeds Picea engelmannii needles Poaceae leaves Juniperus communis fruit Erigeron spp. leaves seeds Achillea lanulosa leaves Fragaria spp. leaves fruit Symphoricarpos spp. fruit leaves Tragopogon dub ius fruit leaves Male in northwestern Frequency occurrence, % Adult Juvenile Female Male Female Totals 19 6 6 10 4 3 6 3 1 6 2 1 41 15 12 4 6 1 4 1 10 2 23 4 3 1 8 2 5 6 1 22 4 3 5 4 8 20 9 4 1 1 1 15 1 3 4 2 4 14 3 1 3 1 1 2 1 9 4 5 2 1 1 3 2 1 4 8 1 1 6 3 1 1 10 2 1 2 4 10 4 3 3 9 2 1 1 1 1 1 1 5 4 4 3 1 1 8 2 1 1 1 1 1 5 2 2 1 1 1 2 6 1 1 2 1 1 1 6 1 9 3 4 11 9 1 �l37 Table 23. (continued) Food and plant part Mentzelia spp , fruit leaves Oxyria digyna leaves Arctostaphylos _---uva-ursi seeds leaves Rosa spp. fruit leaves Pinus contorta needles Poa spp. leaves Prunus virginiana fruit Rubus spp. fruit Shepherdia canadensis leaves Carex spp. seeds leaves Draba .spp. fruit leaves Campanula spp. flowers leaves Ligusticum porteri leaves Pseudotsuga menziesii needles cones Acer spp. seeds Agoseris spp. leaves Alnus tenuifolia catkins Artemisia tridentata leaves Mahonia repens seeds Penstemon spp. leaves Populus angustifolia leaves Frequency occurrence, % Adult Juvenile Female Female Male Male Totals 1 1 1 1 2 5 1 1 3 1 1 6 1 1 1 1 1 4 1 1 1 1 2 4 1 1 1 1 4 1 1 1 4 1 1 2 4 1 1 1 1 4 2 1 4 1 1 1 2 1 1 1 1 1 1 2 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 �138 Table 23. (continued) Food and plant part Populus tremuloides leaves Potentilla spp. leaves Pterospora andromedea fruit Rumex spp. seeds Thermopsis divaricarpa leaves Vicia spp. leaves Unknown plant parts leaves seeds Insects Formicidae Orthoptera Hymenoptera Coleoptera Hemiptera Lepidoptera Unidentified Grit Feathers Mollusca Arachnida Chilopoda Frequency occurrence, % Juvenile Adult Male Female Male Female 1 Totals 1 1 1 1 1 1 1 1 1 1 1 1 1 1 13 1 12 3 6 3 4 36 6 6 9 6 3 1 2 2 10 3 3 2 1 1 2 12 1 3 32 1 1 1 1 11 3 12 1 3 8 1 1 1 1 1 7 6 4 4 1 6 43 6 4 1 1 Juvenile grouse tended to eat more Insecta, Mollusca and centipedes (Chilopods) than adults. This would be expected due to the higher energy and protein requirements of young grouse. The leaves of sulphur flower (Eriogonum spp.) were represented in 22% of the crops. Knapp (1962) and Haggstrom (1966) did not report this food item in their results. Boag (1963) found Eriogonum spp. in 23% of his crop sample. Dandelion (Taraxacum spp.) leaves were used by 23% of the birds in this study. Leaves of this plant were used by 37% of the birds in Boag's (1963) study, but were not mentioned in Knapp's (1962) or Haggstrom's (1966) studies. Conifer needles made up a significant part of the diet of blue grouse in Washington, Oregon, California, Idaho and Montana (Beer 1943, Stewart 1944). Although 4 species of conifer needles were found in the crops of blue grouse in Colorado, the highest frequency of occurrence was 10% for Engelmann spruce (Picea engelmannii). Adult grouse ate conifer needles more than juveniles. �139 There were major differences in the food contents of blue grouse crops examined in this study compared to the results reported by Haggstrom (1966) for birds collected in alpine areas of northcentral Colorado. Willow (Salix spp.) was the principle food item of alpine blue grouse, occurring in 31% of the crops. Willow was not found in any crop from 'this study. Other plants found in Haggstrom's (1966) study, but not recorded in this study include: Lychnis spp., Oxypblis fendleri, Saxifraga spp., Rananculus spp., Veronica spp. , Arenaria spp., Stellaria spp. and Cerastium spp. Many of these plants may have been used by grouse collected in this study, but were among the unidentified materials. However, these data indicate variatians in the foad habits af blue grouse accupying different habitat types. When reviewing the literature it becomes apparent that there are considerable differences in the diet of blue grouse according to locatian. For example, some food items which were in the diet af grouse in Washington, but were nat faund in this study, include: Larix occidentalis, Sambucus melanocarpa, Hieracium spp., Stellaria spp., Arceuthobium campylopadum, Ceanothus velutinus, Epilabium paniculatum and Polygonum spp. Many af these plants do. not occur in Calarada. Others occur in Colarada, but may be used during seasons ather than fall. Several plants were used in Calorada, but not in Washington. They include: Senecio spp., Juniperus communis, Oxyria digyna, Mentzelia spp., Ligusticum parteri, Draba spp. These dietary differences indicate variability in the diet af blue grouse. Blue grouse appear to be oppartunistic feeders and are able to. use whatever is available to. them, depending an locatian and time af year. However, they prefer succulent, herbaceous vegetatian, seeds, and fruits during the fall periad. Habitat Vegetatian Investigatians Type Mapping Aerial phatographs of the study areas were used as base maps. Befare going into. the field, the phatos were carefully examined and every area af vegetation appearing dissimilar from adjacent areas was delineated directly on the photo. The minimum area of vegetation bourtded by a line an the photo was appraximately 0.5 ha. No. types were amitted within the study area boundary. Upon delineation of all areas apparent on the map, each area was then field inspected. The first task was to examine the area fram end to end and across, abserving the vegetatian and field checking the boundaries shown on the pho t o , Salient features o f the area were recorded on prepared data sheets. Consecutive numbers were assigned to each area inspected and were entered both an the data sheet and the phatograph. Each area received its own number regardless of any similarities to previausly inspected areas. To better facilitate preparatian of the final map, a phatograph was taken af each vegetative unit. Number of the individual exposure was recorded along with the correspanding number af the unit where the photograph was taken. �140 The vegetative physiognomy and other physical features of the area were recorded including: slope gradient, exposure and character. The physiognomic classification (Table 24) reveals the appearance and structure of the plant community (vegetation unit), i.e., height and density of every item listed, and in addition, such special features as may be present. A detailed description of the classification scheme and its application is presented by Kuchler (1949). Slope gradient and exposure were measured with an Abney level and compass. After the physiognomic formula for an inspected area had been established, attention was focused on the floristic character of the vegetation. All species occurring on the area were recorded and each was assigned numerical values describing its coverage and sociability (Table 25). Cover is understood to mean the percentage of ground that would be covered if the full spread of the species were projected vertically to the ground." Sociability refers to the distribution of a species within the area of vegetation under consideration. Undoubtedly, some species for which only few specimens were present in a given type escaped detection. This was also true for early or late flowering species that peaked prior to or after the period of field work, which was designed to coincide with the peak of plant development (mid-Jun to mid-Jul). Preparation of the base map involved sorting the data forms, notes and photographs and rearranging the vegetation units into combinations encompassing similar floristic and physiognomic classifications. Woody plants including trees and shrubs were given primary emphasis during the first phase in the combining process. Units within these major vegetation types were then manipulated to produce categories based on similar floristic and structural characteristics. After assigning each unit to a major vegetation type and combining similar vegetation units, the final map was prepared from the base map, pooling similar units which were adjacent to each other. Similar units were then reassigned the same number to simplify the map. Total area mapped as planimetered from U.S.Geological Survey topographic maps was 181.3 ha for Green Mountain and 482.0 ha for Eiby Creek. Area encompassed by each vegetation type and unit was calculated by the weight method (Welch 1948; 85). Five major vegetation types and 26 vegetation units the Green Mountain study area (Figs. 3, 4) compared and 27 units delineated at Eiby Creek (Figs. 6, 7). tables (Appendix B) were compiled and are presented information describing the physiognomy, floristics, of each vegetation unit. were recognized on to 7 vegetation types Vegetative summary as supplementary and physiography Examination of the vegetation maps might indicate that the vegetation types and units were clearly distinct. While the major types and some units were distinct, most units were not, with the majority of units gradually intergrading into adjoining units. In some instances the transition was so gradual as to warrant establishment of a separate unit. When the transition was more abrupt, the boundary line was placed midway through the area of overlap. Thus, mixtures of adjacent units were �141 Table 24. Physiognomic classification of vegetation.a CAPITAL LETTERS: Herbaceous Woody Vegetation: B: D: E: N: 0: G: H: L: evergreen broadleaf deciduous broadleaf evergreen needleaf deciduous needleleaf without leaves Vegetation: graminoids forbs lichens and mosses SMALL LETTERS: Group I: Height: t: tall; m: medium tall; low; 1: s: z: Group II: c: i: p: r: b: Group III: e: j: k: q: a shrubs; dwarf shrubs; height of trees: height of herbaceous plants: of trees: of herbaceous plants: height of trees: height of herbaceous plants: height: continuous growth interrupted; plants usually do not touch plants scattered singly, or in groves or patches rare, yet conspicuous barren; vegetation largely or entirely absent Special Features: epiphytes lianas succulents cushion plants Floristic classification palms bamboos aquatic tree ferns and tuft plants of vegetation.a Sociability: cover very small Plentiful but less than 1/20 of the area Covering 1/2 - 1/4 of the Covering 1/4 - 1/2 of the Covering 1/2 - 3/4 of the Covering greater than 3/4 a u: v: w: y: (1955). + Very sparsely present; 2 3 4 5 1 m Maximum height Cover: 1 25 m 2 m 10-25 m !z-2 m 10 m !zm 1 m Density: From Kuchler Table 25. Minimum Minimum Height Height Maximum Maximum Minimum From Kuchler (1955). 1 2 3 4 5 area area area of the area Growing singly Grouped or tufted In small patches In extensive patches In great crowds �142 frequently present along their boundaries. "Islands" of dissimilar vegetation within a larger, individual unit were usually too small « 0.5 ha) for recognition, and consequently, were ignored unless they were of known importance in terms of grouse use. Characteristics of Breeding Areas Blue grouse males on the study areas were territorial and defended chosen territories throughout the breeding season (mid-April to early July). All territories located (GM = 22, EC = 36) from 1975 to 1979 were marked on aerial photos. Territories were subsequently plotted on the vegetation maps to facilitate their description in relation to vegetative features of the area (Fig. 5, 8). Delineation of territorial boundaries was limited to 9 territories on Green Mountain where the resident males were banded and observed 5 or more times within the same breeding season. Otherwise, only an approximate location of the territory was determined. No territory was located completely in any 1 vegetation unit as portions of at least 2 and up to 4 units were encompassed within the boundaries of all territories on both study areas. Breeding territories were primarily located where Pseudotsuga (GM) or Populus units (EC) intergraded into more open areas such as the Artemisia (GM and EC) and the Amelanchier, Symphoricarpos, and grass-forb units (EC). All territories at Eiby Creek were at least partially associated with the Populus units. Riparian units were not extensively used by territorial males except where the aspen extended away from the creek bottom and adjoined a more open unit. Douglas-fir units were the major vegetational components of 21 and 22 territories on Green Mountain, although aspen was frequently present in small amounts. The only territory without Douglas-fir was associated with an aspen-sagebrush mixture, and this territory was occupied for just 1 breeding season. Territorial males avoided dense stands of Douglas-fir except along the edges adjacent to more open units. Aspen was an important part of breeding habitat on Green Mountain as it was much less abundant in the overall tree cover than at sites where territorial males were observed. This was attributed to the fact that (1) territorial males were most frequently found while performing their displays, and (2) aspen occurred along the edges of coniferous stands and openings which were the same areas preferred by males for displaying. Features found in common among all territories examined included: (1) some form of tree cover, (2) shrub thickets, (3) edges, (4) open areas, and (5) some degree of openness in both the canopy and understory cover. The data do not imply that territorial males selected Douglas-fir and aspen over other species of trees for breeding habitat, but simply reflect the high incidence of Douglas-fir and aspen on the 2 study areas. What the data do suggest is that structural characteristics of the vegetation more than species composition are an important factor in breeding habitat selection. Within Colorado, and throughout its range, the blue grouse has been documented to breed in a variety of forest and mountain shrub types from the foothills to timberline with no apparent restrictions to any habitat type within this elevational range. �143 GREEN MOUNTAIN ,..~, ~ ," 1 ~~ ~~~ ~ ".r"'~ ( \ "" ~ ~ .. ...... '" '... Artemisia (77.l)a - mixed shrub D ...~~ m~ Pseudotsu ga - mixed shrub (43.7) [[JJIIJ] Pseudotsu ga-Populusmixed shrub (44.4) D Populus - mixed shrub (l3.2) § Carex-Agr opyronscattered shrub (2.9) [Z] aSurface area in hectares ~ N . 4~ SCALE ] I o I km Fig. 3. Major vegetation ~r ~ types, Green Mountain study area. �144 GM 1 Artemisia-mixed shrub-scattered Pseudotsuga GM 2 Artemisia-Symphoricarpos-mixed GM 3 Artemisia-Chrysothamnus-mixed GM 4 Artemisia-Amelanchier-mixed GM 5 Artemisia-mixed GM 6 Artemisia-Agropyron-Chrysothamnus GM 7 Dense Pseudotsuga-Symphoricarpos-mixed shrub GM 8 Dense Pseudotsuga~bareground-scattered shrub GM 9 Dense Pseudotsuga-Juniperus-mixed shrub-scattered Pseudotsuga shrub shrub-scattered shrub-rock-scattered Pseudotsuga Pseudotsuga shrub GM 10 Dense Pseudotsuga-Acer-mixed shrub GM 11 Semi-open Pseudotsuga-Symphoricarpos-mixed GM 12 Semi-open Pseudotsuga-mixed GM 13 Open Pseudotsuga-Artemisia-Symphoricarpos-mixed GM 14 Open Pseudotsuga-Prunus-mixed GM 15 Open Pseudotsuga-Artemisia-mixed shrub GM 16 Open Pseudotsuga-Juniperus-mixed shrub-rock GM 17 Semi-open GM 18 Open Pseudotsuga-Populus-mixed GM 19 Dense Pseudotsuga-Populus-mixed GM 20 Scattered GM 21 Low Populus-mixed GM 22 Medium Populus-mixed GM 23 Scrub Populus-mixed GM 24 Medium Populus-Pseudotsuga-Prunus-mixed GM 25 Medium Populus-Carex-Lathyrus-mixed GM 26 Carex-Agropyron-mixed shrub-rock shrub outcrop shrub shrub Pseudotsuga-Populus-Symphoricarpos-mixed shrub shrub shrub-rock Pseudotsuga-Populus-Artemisia-mixed shrub-rock shrub shrub shrub grass-scattered Fig. 4. Vegetation units, Green Mountain information presented in Appendix B). shrub shrub shrub study area (supplementary �145 GREEN MOUNTAIN N SCALE o I km Fig. 4. Vegetation units, Green Mountain information presented in Appendix B). study area (supplementary �GREEN MOUNTAIN Approximate location of breeding territories [[Ill Approximate boundary of breeding territories Brood concentration areas N SCALE Fig. 5. Major vegetation types and blue grouse use areas, Green Mountain. �147 EIBY CREEK Artemisia-mixed Populus-mixed shrub shrub Amelanchier-mixed (179.6) shrub Symphoricarpos-mixed Prunus-mixed (184.5)a shrub Forb-Grass-meadow shrub (10.6) (5.4) (11.7) Populus-Alnus/Alnus-Salix a N (45.8) SCALE riparian (43.4)!5{l o Surface area in hectares. Fig. 6. Major vegetation types, Eiby Creek study area. 1/2 Ikm �148 EC 1 Artemisia-Chrysothamnus-mixed EC 2 Artemisia-Chrysothamnus-Prunus-mixed EC 3 Artemisia-Cercocarpus-Amelanchier-mixed EC 4 Artemisia-Amelanchier-rock-mixed EC 5 Artemisia-Symphoricarpos-Chrysothamnus-mixed EC 6 Chrysothamnus-Symphoricarpos-Prunus-scattered EC 7 Artemisia-Amelanchier-mixed EC 8 Medium EC 9 Medium Populus-Symphoricarpos-Prunus-Amelanchier-rock shrub shrub shrub shrub shrub Artemisia shrub (sprayed area) Populus-Symphoricarpos-Ribes-mixed shrub EC 10 Medium Populus-Symphoricarpos-Prunus-Amelanchier-Ribes EC 11 Medium Populus-Symphoricarpos-Prunus-Ribes EC 12 Medium EC 13 Low Populus-Arnelanchier-Prunus-Symphoricarpos EC 14 Scrub Populus-Symphoricarpos-Prunus-Rosa EC 15 Open Populus-mixed EC 16 Open Populus-Symphoricarpos-mixed EC 17 Arnelanchier-Prunus-Symphoricarpos-mixed EC 18 Amelanchier-Prunus-mixed EC 19 Arnelanchier-Prunus-Symphoricarpos-scattered EC 20 Symphoricarpos-Prunus-mixed EC 21 Symphoricarpos-Chrysothamnus-mixed EC 22 Prunus-Symphoricarpos-mixed EC 23 Forb-Grass-scattered EC 24 Populus-Alnus-mixed EC 25 Alnus-Populus-Salix-Ribes-Cornus-riparian EC 26 Alnus-Salix-Ribes-riparian EC 27 Salix-Alnus-Cornus-riparian Populus-mixed (sprayed area) outcrop forb-grass shrub shrub shrub shrub Juniperus shrub shrub-scattered shrub-scattered Populus Populus shrub shrub-riparian Fig. 7. Vegetation units, Eiby Creek study area (supplementary information presented in Appendix B). �14 S' EIBY CREEK N + SCALE [ 0 Fig. 7. Vegetation units, Eiby Creek study area (supplementary presented in Appendix B). I 1/2 information I 1km �15) ElBY CREEK N Approximate location breeding territories Brood concentration Fig. 8. t of SCALE areas Major vegetation types and blue grouse use areas, [ I 0 1/2 Eiby Creek. I Ikm �151 Territorial males were seldom found more than 25 m from an opening and avoided areas with dense canopy or understory cover except along the edges. During early to mid-April when most ground cover was beneath the snow, the birds on Green Mountain spent the majority of time in trees using small, dense clumps of conifers in an otherwise open habitat. As phenological changes progressed and the birds reverted to ground dwelling habits, shrub thickets became important during resting and feeding activities while trees were primarily used as escape cover and for roosting. Conversely, the birds at Eiby Creek depended heavily upon shrub thickets for cover in early spring. The openness of the aspen stands afforded little cover from predators. Only in the early morning and late evening hours during feeding activities were the birds found in the aspen, unless disturbed while on the ground, they would then frequently fly into a nearby aspen. Some territorial males, both at Eiby Creek and Green Mountain, performed their displays considerable distances (750 m) from tree cover and therefore relied on nearby shrub thickets for cover. Areas characterized by an interrupted mixture of low growing vegetation, bare ground, stumps, logs and patches of tree and shrub cover were preferred over areas with continuous or homogeneous ground cover. Characteristics of Nest Sites Twelve nests were found on the study areas (GM = 11, EC = 1) from 1976 to 1979. Each nest site was described as to aspect, slope~ altitude and vegetative cover. All nests had some type of cover immediately above and surrounding the nest. The cover was mainly in the form of shrub clumps, but 2 nests were partially protected by a log while another was adjacent to the trunk of a Douglas-fir. Three nests were found under sagebrush, 3 beneath saplings of Douglas-fir, 1 at the base of a larger Douglas-fir (rv 6 m high), 2 in the center of a sagebrush-snowberry clump, 1 in a sagebrush-rabbitbrush clump (EC nest), 1 under a log partially covered by snowberry, and 1 in a serviceberry-sagebrush clump. Maximum distance from the nest to the nearest tree was 42 m. Excluding the nest under a large Douglas-fir, the height of vegetation immediately above the nest averaged 98 cm and ranged from 30 to 280 cm. General appearance of the habitat around the nest was open to semi-open. Of the 12 nests located, 6 were in transition zones (edges) between the following units: (1) dense Pseudotsuga-Symphoricarpos-mixed shrub and Artemisia-Amelanchier-mixed shrub-scattered Pseudotsuga (2 nests), (2) semi-open Pseudotsuga-Populus-Symphoricarpos-mixed shrub and Artemisia-Symphoricarpos-mixed shrub-scattered Pseudotsuga (2 nests), (3) semi-open Pseudotsuga-Populus-Symphoricarpos mixed shrub and Artemisia-mixed shrub-scattered Pseudotsuga (1 nest), and (4) semi-open Pseudotsuga-Populus-Symphoricarpos-mixed shrub and dense PseudotsugaSymphoricarpos-mixed shrub (1 nest). Two of the other 6 nests were in the Artemisia-Amelanchier-mixed shrub-scattered Pseudotsuga vegetation unit and 1 nest was found in each of the following 3 units: (1) ArtemisiaSymphoricarpos-mixed shrub-scattered Pseudotsuga, (2) scattered Pseudotsuga-Populus-Artemisia-mixed shrub-rock, and (3) Artemisia-mixed shrub-scattered Pseudotsuga. The 1 nest location at Eiby Creek was in the Artemisia-Chrysothamnus-mixed shrub vegetation unit. Nest sites were in the same vegetation units selected for breeding, but only �152 one-half the nests were located within the boundaries of a territory. Artemisia tridentata was associated with all nesting sites except 1, but this may be related only to the abundance of sagebrush on both study areas. However, hens apparently did select the more open units for nest sites and these same units were preferred brood use areas. All nests were between 2,500 and 2,740 m elevation. The sample size was inadequate to evaluate the importance of slope and aspect, but neither feature was believed to be a significant factor in nest site selection. Exposure of nests included all major compass directions except south. The majority of nests faced in northerly and easterly directions which were the primary exposures for the Green Mountain study area where most nests were found. Slope at the nest site ranged from 6 to 25%. Characteristics of Brood Areas Two hundred and fifty-seven observations of blue grouse broods were made on Green Mountain (B = 121) and at Eiby Creek (B = 136) from 1975 to 1979. At 1 time or another, broods were observed within most all vegetation units (Fig. 4, 7). However, there were definite concentration areas. Nearly 90% of all brood sightings on Green Mountain were in the open and semi-open habitats on the east face of the study area (Fig. 5). Broods tended to use more of the study area at Eiby Creek, possibly due to the greater availability of open areas and edges. Nevertheless, broods congregated in certain areas. These areas (Fig. 8), accounted for about 70% of all brood observations at Eiby Creek and together with the concentration sites on Green Mountain were the focal areas for evaluation of brood habitat. Broods used areas where vegetation had interspersions of plants of various life forms. Such areas were generally along the edges of brush or tree cover and provided a high degree of concealment. Forested areas with a dense understory or open areas with heavy herbaceous ground cover were avoided except along the edges. Almost invariably, any brood encountered in the field was originally observed on the ground. They seldom ventured more than 30 m from brush or tree cover. Shrub thickets and edges of forested areas were used for resting and for escape when disturbed. Broods were frequently found along stream courses or near other wet sites at Eiby Creek providing the vegetation was not too dense. However, there was no evidence of the necessity for open water near brood cover. Only 2 sources of open water occur on Green Mountain, and no broods were found at either location. Summer brood range partially overlapped breeding areas, but there tended to be a greater diversity of plant species, more ground cover (especially herbaceous), and a greater amount of open vegetation units in brood habitat than breeding habitat. Structural characteristics of the vegetation more than species composition appeared to be the critical factor in habitat selection by broods. Examination of the data (Fig. 5) indicates that a major portion of the brood range on Green Mountain was included within the semi-open Pseudotsuga-Populus-Symphoricarpos-mixed shrub vegetation unit. Even in this predominantly Douglas-fir unit, broods were most often found �153 near or in the many small clearings (Artemisia-Symphoricarpos-mixed shrub-scattered Pseudotsuga) dispersed throughout this unit. Characteristically, these openings, as with other forest edges on the study area, were often bordered by aspen; thus, the distribution of broods closely approximated the distribution of aspen on the study area. At lower elevations broods were found in transition areas between the coniferous and sagebrush types where aspen was commonly present. Vegetation units favored by broods in these areas included: scrub, low, and medium Populus-mixed shrub units, Artemisia-Chrysothamnus-mixed shrub, Artemisia-Amelanchier-mixed shrub-scattered Pseudotsuga, open PseudotsugaPopulus-mixed shrub, and open Pseudotsuga-Artemisia-Symphoricarpos-mixed shrub. There was a great deal of variation in brood habitat at Eiby Creek. Broods were found in and along the edges of aspen groves, on open sagebrush hillsides, along alder-willow dominated stream courses, and in dense patches of chokecherry and serviceberry. There was a clear preference for edges where open vegetation units occurred adjacent to aspen stands or patches of tall shrubs, i.e., serviceberry and chokecherry. Aspen was the dominant tree in brood cover, but there was no apparent preference for certain aspen units as long as the stands were bordered by open areas. During daylight hours, broods were most frequently found foraging on open hillsides covered with shrubs, while at night they roosted in aspen stands or patches of serviceberry and/or chokecherry. Sites used for roosting also served as resting and loafing cover during the day; when disturbed, the hen and chicks usually flew into the nearest aspen stand or shrub thicket. The Artemisia-Symphoricarpos-Chrysothamnus-mixed shrub was the single most important vegetation unit for brood use at Eiby Creek, especially where this unit bordered aspen. Other open units where broods were commonly located included: Artemisia-Chrysothamnus-Prunus-mixed shrub and Chrysothamnus-Symphoricarpos-Prunus-scattered Artemisia (sprayed area). Several smaller, localized areas were identified as brood concentration sites. One such area that was heavily used by grouse throughout the spring, summer, and fall consisted of a mixture of the Amelanchier-Prunus-mixed shrub, medium Populus-Symphoricarpos-Prunus Amelanchier-Ribes, and Artemisia-Cercocarpus-Amelanchier-mixed shrub vegetation units. Light to moderate grazing pressure had little impact on brood habitat at Eiby Creek. However, excessive grazing pressure virtually eliminated herbaceous cover and the grouse moved to less disturbed sites. Such influence was particularly noted in the forb-grass scattered shrub vegetation unit. Where herbaceous cover remained, broods were found in the same pastures with cattle, but usually on steep slopes or thickets less frequented by the cattle. �154 LITERATURE Beer, J. R. 1943. 7:32-44. CITED Food habits of the blue grouse. J. Wildl. Manage. Bendell, J. F., and P. W. Elliott. 1967. Behavior and the regulation of numbers in blue grouse. Can. Wildl. Servo Rep. Ser. 4. 76pp. ___ -:-'D. G. King, and D. H. Mossop. 1972. of blue grouse in a declining population. 1153-1165. Blackford, J. L. 1958. Territoriality lation of blue grouse in Montana. Removal and repopulation J. Wildl. Manage. 36: and breeding behavior of a popuCondor 60:145-158. 1963. Further observations on the breeding behavior blue grouse population in Montana. Condor 65:485-513. of a Boag, D. A. 1963. Significance of location, year, sex, and age to the autumn diet of blue grouse. J. Wildl. Manage. 27:555-562. 1966. Population attributes of blue grouse in southwestern Alberta.. Can. J. Zool. 44:799-814. Braun, C. E. 1969. Population dynamics, habitat, and movements of whitetailed ptarmigan in Colorado. Ph.D. Thesis. Colorado State Univ., Fort Collins. 189pp. 1971. Determination of blue grouse sex and age from wing characteristics. Colorado Div. Game, Fish and Parks. Game Infor. Leafl. 86. 4pp. Braun-Blanquet, J. 1951. Germany. pp. 58-66. Pflanzensoziologie. Springer-Verlag, Wien, Bray, O. E., J. L. Guarino, and W. C. Royall, Jr. 1975. A trap for capturing territorial male red-winged blackbirds. Western Bird Bander 50:4-7. Caswell, E. B. 1954a. Manage. 18: 139. A method for sexing blue grouse. J. Wildl. _---:-. 1954b. A preliminary study of the life history and ecology of blue grouse in west-central Idaho. M.S. Thesis. Univ. Idaho, Moscow. 105pp. Gilfillan, M. C., and H. Bezdik. 1944. Winter in Ohio. J. Wildl. Manage. 8:208-210. Gullion, G. W. 1965. Improvements in methods marking ruffed grouse. J. Wildl. Manage. Haggstrom, D. A. northcentral foods of the ruffed grouse for trapping and 29:109-116. 1966. Fall food habits of blue grouse Colorado alpine. Unpubl. rep. 9pp. in the �155 Harju, H. J. 1974. An analysis of some aspects of the ecology of Univ. Wyoming, Laramie. dusky grouse. Ph.D. Thesis. 142pp. Harrington, H. D. 1964. Manual of the plants of Colorado. Brooks, Denver, Colo. 666pp. Sage Henderson, V. B. 1960. A study of the blue grouse on summer range, northcentral Washington. M.S. Thesis. Washington State Univ., Pullman. 96pp. Hickey, J. J. 1955. Some American population research on gallinaceous birds. Pages 326-396 in A. Wolfson, ed. Recent studies in avian biology. Univ. Illinois Press, Urbana. 479pp. Hjorth, I. 1970. Reproductive behavior in Tetraonidae, reference to males. Viltrevy 7 :183-596. with special Hoffman, R. W., and C. E. Braun. 1975. A volunteer wing collection station. Colo. Div. Wildl. Game Infor. Leafl. 101. 3pp. Holt, H. E. 1961. Geology of the lower Blue River area, Summit and Grand counties, Colorado. Ph.D. Thesis. Colorado Univ., Boulder. 107pp. Knapp, D. B. 1962. September foods of blue grouse in north-central Colorado. Unpubl. rep. 6pp. Kuchler, A. W. 1949. A physiognomic classification Ann. Assoc. Amer. Geographers 39:201-210. of veget~tion. 1955. A comprehensive method of mapping vegetation. Assoc. Amer. Geographers 45:404-415. Martinka, R. R. territories 498-510. Ann. 1972. Structural characteristics of blue grouse in southwestern Montana. J. Wildl. Manage. 36: Medin, D. E. 1962. An ecological investigation of the Cache la Poudre deer herd. Colorado Dept. Game and Fish. Job Completion Rep. Fed. Aid Proj. W-I05-R. July 1962. pp. 187-204. Mussehl, T. W. 1960. Blue grouse production, tions in the Bridger Mountains, Montana. 24: 60-68 .. movements, and populaJ. Wildl. Manage. Nelson, R. A. 1969. Handbook of Rocky Mountain King, Tucson, Ariz. 331pp. plants. Dale Stuart Redfield, J. A., and F. C. Zwickel. 1976. Determining the age of young blue grouse: a correction for bias. J. Wildl. Manage. 40:349-351. �156 Rogers, G. E. 1968. Fish and Parks. The blue grouse in Colorado. Tech. Publ. 21. 63pp. Colo. Div. Game, Schladweiler, P., and T. W. Mussehl. 1969. Use of mist nets for J. Wildl. Manage. recapturing radio-equipped blue grouse. 33:443-444. Stauffer, J. E. 1953. County, Colorado. 62pp. Geology of an area west of Wolcott, Eagle M.S. Thesis. Univ. of Colorado, Boulder. Stirling, I., and J. F. Bendell. recorded calls of a female. ____ ~-' and Syesis 3:161-171. 1970. 1966. Census of blue grouse with J. Wildl. Manage. 30:184-187. The reproductive behavior of blue grouse. Taggart, J. N. 1962. Geology of the Mount Powell quadrangle, Ph.D. Thesis. Harvard Univ., Boston, Mass. 239pp. Tomlinson, R. E. 1963. A method for drive-trapping J. Wildl. Manage. 27:563-566. Weber, W. A. Boulder. 1972. Rocky Mountain 437pp. Welch, D. S. 1948. Limnological New York. 381pp. flora. methods. Colorado. dusky grouse. Colorado Assoc. Univ. Press, McGraw-Hill Book Co., Inc., Zwickel, F. C. 1965. Early mortality and numbers of blue grouse. Ph.D. Thesis. Univ. British Columbia, Vancouver. 153pp. 1972. Removal and repopulation of blue grouse in an increasing population. J. Wildl. Manage. 36: 1141-1152. 1975. Nesting parameters of blue grouse and their relevance to populations. Condor 77:423-430. , and ------grouse. J. F. Bendell. 1967a. A snare for capturing J. Wildl. Manage. 31:202-204. _______ , and of numbers 1967b. Early mortality and the regulation of blue grouse. Can. J. Zool. 45:817-851. ------ , and A. N. Lance. grouse. Prepared by: blue 1966. Determining J. Wildl. Manage. 30:712-717. the age of young blue �157 APPENDIX STATEWIDE Objective A GROUSE WING COLLECTIONS OPERATIONS PLAN and Goals The objective is to ~stablish a statewide wing collection program designed to obtain long-term population and harvest trend data on blue grouse and sage grouse for management purposes. The goal will be to collect 3,200 blue grouse and 2,000 sage grouse wings through the operation of volunteer collection stations at various locations in the state. Data should be assembled on a regional basis with the Small Game Management Section in Denver coordinating regional efforts. Specific Regional goals are given (see attached) and 56 locations are recommended as permanent sampling points: 11 for blue grouse, 24 for sage grouse and 21 for concurrent collection of wings from both species. The recommendations are by no means inclusive of the entire state, but represent areas where data are already available. For this reason, we cannot over-emphasize the need to. continue wing surveys in these areas. Specific goals for sage grouse are based on discrete populations rather than regional goals. Even so, there are no situations where the populations cross regional boundaries. Most sampling sites for blue grouse are confined to the northwest region where our research activities were centered. Here we took an area approach, setting specific quotas for defined geographical areas within the region. Because we lack knowledge related to hunter pressure, harvest and location of major hunting areas in the other regions, we were limited to establishing an overall realistic goal for each affected region without designating specific areas. Such knowledge will evenCually be obtained using wing barrels, but until then, we need to experiment with different sampling points leaving their actual selection to regional personnel. The idea of volunteer collection stations for obtaining anatomical parts from harvested game is not new nor is it inferred that their application be restricted to blue grouse and sage grouse. We certainly encourage the regions to explore other avenues of use and to develop objectives and goals for collecting wings from other species. The southwest region has already adapted their stations for collecting pheasant wings, while Mike Szymczak has used volunteer stations in North Park and' South Park for ascertaining species composition of the waterfowl harvest from wing samples. Expected Results From analyses of wings, data will be acquired concerning (1) age and sex composition of the harvest, (2) nesting success of breeding females, (3) hatching dates, (4) turnover rates, and (5) productivity. When collected consistently over a period of years, the data will also be useful in monitoring trends in both the population and harvest. Other information of significance to management that will be obtained from operation of wing stations include: (1) identification of major harvest areas, (2) evaluation of hunter success, and (3) assessment of the distribution of harvest over time. �158 Cost, Maintenance, Durability and Manpower Materials and labor for constructing 1 station presently cost about $50.00. Using scrap materials, less elaborate stations can be built at considerably lower cost. Most of the expense involves the construction of a quality sign. Our stations are still structurally sound after 6 years of use and have required only minor maintenance such as painting and rethreading the pipe. The signs are less durable and will require restencilling after 7-10 years of use. Most stations invariably sustain damage from bullet holes, but this has not affected their operation. Otherwise, damage from vandalism has been minor. Annual maintenance cost is estimated at about $4.60/station/year. No new or part-time employees will be necessary to conduct this project. However, it will be necessary to redirect some duties of DWM's and regional biologists to insure proper implementation. With all materials on hand, 1 person can construct a single station in about 2 hours. Setting up, checking and dismantling the stations can be partially accomplished in the course of performing other duties to minimize additional time, manpow~r, mileage and expense. One person should oversee the program in each region and coordinate activities with the other regions. The actual processing of wings and analysis and interpretation of data will probably require at least 4 working days. All interested personnel should be invited to participate. However, one person should be responsible for insuring wings are properly classified and all data accurately recorded for later analysis. Consistency in data collection and analysis is imperative for comparisons among areas, years and species. Total man-days and cost per region for the entire operation of this project will be less than 25 man-days and $5,000.00. Procedures 1. Incorporate information already available into one or several publications to include: (1) results of wing surveys conducted by research personnel, (2) procedures for conducting the survey, (3) techniques for classifying wings to age and sex, (4) methods of analysis and interpretation of data, and (5) design of a statewide survey. Towards this end, management reports have been prepared each year summarizing data derived from wing collections. Time, cost and manpower requirements plus step by step instructions for implementing a wing survey have been previously outlined to the regions via performance contracts. Techniques for separating sex and age classes of blue grouse and sage grouse have been published (Game Information Leaflets Numbers 49 and 86) along with information pertaining to the design and construction of wing barrels (Game Information Leaflet Number 101). In addition, we plan to produce several other written products which should further clarify the operation of wing stations, separation of age and sex classes, analysis of wing data, and interpretation of findings. One such product will be a "cookbook" type manual with procedures, recommendations and results of wing surveys conducted by research personnel. 2. Increase public awareness to enhance cooperation. It has been our experience that time is a big factor in terms of increasing public awareness. For instances, stations in Middle Park are now providing 3 times the number of wings they produced in 1975, not because of an �159 increase in hunters or harvest, but because of greater awareness and cooperation. Other ways to consider getting the word out include: news releases, hunter safety classes, Colorado Outdoors, local sportsmen's clubs, and Wildlife News. 3. Select locations for operation of wing stations. As previously noted and attached, we have compiled a list of specific locations for collecting wings. However, other areas should be identified. Attempts should be made to obtain a sample of wings from the entire region and to select major access routes in order to sample as large an area as possible. 4. Obtain materials and construct wing stations. Additional materials should be kept on hand for replacement of stations that are vandalized. We have 50 complete stations and approximately 20 additional barrels in Fort Collins, of which 35 stations and all barrels can be distributed to the regions to reduce costs. Furthermore, there are at least 25 stations in existence that have been independently operated by regional personnel in excess of the stations maintained by research. All total there should be somewhere in the neighborhood of 75 stations currently available for use. 5. Install stations 1-2 days prior to opening of the grouse season. Stations should be highly visible and readily accessible to the hunters. If possible, place stations near stop signs or junctions where traffic must stop or proceed slowly and where there is adequate space for vehicles to pull off the road. 6. Stations should remain in operation from opening weekend (2nd Saturday in September) through the last weekend in September. Our reasons for selecting this time period are twofold: (1) it avoids conflicts with other work assignments during the big game seasons, and (2) many birds have already completed their primary molt by early-October and consequently are not useable for calculation of nesting success or hatching dates. 7. All stations should be checked a m~n~mum of twice/week preferably on Friday afternoons and Monday mornings. Wings collected on Fridays are assumed to be from birds shot during the week, while wings collected on Mondays are assumed to be from birds harvested over the weekend. Knowledge concerning the approximate data of harvest is imperative for calculations of hatching dates and nesting success and for assessment of the distribution of harvest over time. 8. Record the date, station location and number of wings collected by species on a standardized form each time a volunteer station is checked. Wings collected from each station should be placed in a plastic bag, labeled as to date and location of collection, and stored in a freezer at a central location until they can be processed. 9. Conduct regional "wing bee" to classify wings and tabulate data. Research personnel will assist in processing wings and training personnel during the initial year of implementation. �160 10. Prepare management reports on results of wing surveys similar reports we have prepared over the past several years. to the 11. Assemble, compare and report findings from all regions into an annual report to be compiled by the Small Game Management Section and to function as a supplement to the annual small game harvest survey. �161 GROUSE WING COLLECTION I. Regional A. Goals Sage Grouse 1. NW Region a. b. c. d. e. 2. 3. North Park -- 500 wings 500 wings Gunnison Basin -- 500 wings Fruitland Mesa, if season is opened SE Region -- none, no sage grouse range NW Region a. b. c'. d. e. f. -- 1,700 Middle Park -- 500 Routt County -- 500 Eagle County -- 400 Piceance Basin -- 100 Moffat County -- 100 Grand Mesa -- 100 2. NE Region 500 wings 3. SW Region 500 wings 4. SE Region 500 wings Recommended A. 500 wings Blue Grouse 1. II. 1,000 wings Moffat and Western Routt counties -- 600 wings Eagle County -- 80 wings Middle Park -- 100 wings Yampa -- 140 wings Piceance Basin -- 80 wings SW Region a. b. 4. -- NE Region a. B. GOALS AND LOCATIONS Locations for Operating Wing Barrels Sage Grouse 1. NW Region -- 32 barrels a. Moffat and Western Routt counties -- 14 barrels 1. 2. 3. 4. 5. 6. Axial Deception Creek Juniper Springs M.C. Rd. 2 M. C. Rd. 101 California Park Rd. �162 Grouse Wing Collection 7. 8. 9. 10. 11. 12. 13. 14 . b. Milk Creek Eiby Creek Rd., No. end Wolcott Muddy Creek Pass Williams Fork Rd. Corral Creek Rd. Gore Pass, Jct. Colo. 134 and US 40 Pinto Creek Rd. Chimney Rock Rd. Spring Creek Rd. Williams Peak Rd. Yampa -- 4 barrels 1. 2. 3. 4. e. 4 barrels Middle Park -- 7 barrels 1. 2. 3. 4. 5. 6. 7. d. Maybell, Jct. US 40 & Colo. 318 Dinosaur, 1.6 km E. of Dinosaur on US 40 at junction with N.P.S. road Wolf Creek Rd. Cedar Mountain (Junction of M.C. roads 3 and 7) Maybell City Park Lay Creek M. C. Rd. 3 M. C. Rd. 4 Eagle County 1. 2. 3. 4. c. Goals and Locations Five Pines Mesa No. Five Pines Mesa So. DeGohanl Green Ridge Piceance 1. 2. 3. Basin -- 3 barrels Cathedral Bluffs Cow Creek Magnolia Creek 2. NE by at to 3. SW Region -- 14 barrels a. Region -- check stations and wing barrels will be operated research personnel in North Park for at least 3 more years which time recommendations will be developed for the region continue wing collections. Gunnison Basin -- 14 barrels 1. 2. 3. 4. 5. 6. 7. Blue Mesa Cochetopa at US 50 Doyleville Gold Basin Lost Canyon No. Parlin Flats Ohio Creek �163 Grouse Wing Collection 8. 9. 10. U. 12. 13. 14. b. B. Goals and Locations Sapinero at US 50 Sapinero at Colo. 149 Six Mile Lane So. Parlin Flats Waunita Hot Springs Willow Creek at Colo. 149 Woods Gulch Fruitland Mesa -- to be determined if season is opened. Blue Grouse -- note that 21 locations are recommended for the concurrent collection of blue grouse and sage grouse wings (not including North Park). Some other locations will occasionally produce blue grouse wings, but not in any significant numbers. 1. NW Region -- 27 barrels a. Moffat County -- 4 barrels 1. 2. 3. 4. b. Willow Creek, Jct. Colo. 125 & US 40 Corral Creek Gore Pass, Jct. Colo. 134 & US 40 Chimney Rock Trough Rd., Jct. with Colo. 9 Spring Creek Rd. Williams .Peak Rd. Routt County -- 6 barrels 1. 2. 3. 4. 5. 6. e. Red Sandstone Rd. Wolcott Muddy Creek Pass Milk Creek Eiby Creek Rd., No. end Coffee Pot Springs Middle Park -- 7 barrels 1. 2. 3. 4. 5. 6. 7. d. & Colo. 318 Eagle County -- 6 barrels 1. 2. 3. 4. 5. 6. c. M.C. Rd. 2 M.C. Rd. 101 Maybell, Jct. US 40 Dinosaur Elk River Rd. Buffalo Pass Rd. California Park Rd. Lynx Pass Dunkley Pass Green Ridge Piceance 1. 2. Basin -- 2 barrels Cow Creek Cathedral Bluffs �164 Grouse Wing Collection Goals and Locations f. Grand Mesa -- 2 barrels 1. 2. 2. & 330 NE Region -- 20 barrels -- additional areas to be selected by regional personnel a. North Park -- will be sampled by research personnel for at least another 3 years. b. Larimer County 1. c. Pingree Park (previously sampled) Other suggested locations: 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 3. Silt Jet. Colo. 65 Ted's Place, Jct. Colo. 14 Livermore Cherokee Park Masonville Drake E. side Rollins Pass Grant Squaw Pass, Jct. Colo. 103 Jet. Colo. 67 & 105 Rampart Range North & US 287 & 74 SW Region -- 20 barrels -- additional areas to be selected by regional personnel a. Gunnison Basin 1. 2. 3. 4. b. Gold Basin Ohio Creek Sapinero at Colo. 149 Blue Mesa Other suggested locations 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. Uncompahgre Plateau (at least 6 stations) Norwood Kebler Pass, E. and W. sides Almont - Taylor River Rd. Jet. Colo. 149 at Wagon Wheel Gap Jct. Colo. 92 and US 50 Mt. Crested Butte - Gothic Jet. Colo. 65 and 92 Dolores Telluride �165 Grouse Wing Collection 4. SE Region -- 20 barrels -- no areas previously areas to be selected by regional personnel a. Regional A. Sage Grouse -- IS barrels Blue Grouse -- 10 barrels Sage Grouse and Blue Grouse -- 17 barrels Sage Grouse -- No. Park research study Blue Grouse -- 20 barrels Sage Grouse and Blue Grouse -- No. Park research SW Region -- 34 barrels 1. 2. 3. D. Summary NE Region -- 20 barrels 1. 2. 3. C. Ophir Creek Gardner Cucharas Pass, both sides Apishapa Pass Bear Creek Cottonwood Pass Weston Pass Rampart Range South Buffalo Peaks Rd. Independence Pass Wolf Creek Pass East NW Region -- 42 barrels 1. 2. 3. B. sampled, all Suggested areas: 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. III. Goals and Locations Sage Grouse -- 10 barrels Blue Grouse -- 20 barrels Sage Grouse and Blue Grouse SE Region -- 20 barrels 1. 2. Sage Grouse Blue Groiise o barrels 20 barrels -- 4 barrels study �166 APPENDIX B �167 DESCRIPTIVE Legal Range: SUMMARY OF VEGETATION TYPE HAP. GREEN HQUNTAIN- BLUE GROUSE INVESTIGATIONS Description: T25. R80W, Sections 2, 3. 10. 11 County: SUClllllt 2500-2865 C1 Surface Area: ~ Aerial Photo No. U.S.F.S. Fl6CN 10-3-72 Elevational VEGETATIVECLASSIFICATION AND SUPPLEMENTARY FEATURES Classification GM-1 and Features !! Physiognomic BziDzlElp GM-2 GM-3 BdDziElp Exposure (Degrees Azimuth) 38-85 36-46 Gradient (Percent) 18-30 20-25 Area 14.7 (Hectares) Plant Species List 11 Woody - Trees: Acer glabrum Juniperus virginians Populus t reeukoddes Pseudotsuga menziesii Wood -Shr-ub s : Acer glabrum ADelanchier a1n1£o11a Arctostaphylos uva-ursi Artemisia cana Artemisia tridentata Ceanothus ve1utinus Cercocarpus montanus Chrysothamnus spp. Holodiscus dumosus Juniperus coeeunt s Juniperus virginiana Mahonia repens Pachystima myrsinites Populus tremuloides Prunus virgIniana Pseudotsuga cene tes i i Purshia tridentata Rhus r r t Iobat a Ribes spp. Rosa acicularis Rubus de1iciosus Sambucus racemosa Shepherdia canadensis Symphoricarpos oreopnrfus Tetradymia caneecens vacc rntue spp , Forbs: Achillea 1anu1osa Anemone spp. Antennaria spp. Aqui1egia caeru1en Arabis spp. Arnica spp . Artemisia frigida Artem1sia ludoviciana Aster spp. Astragalus spp. Balsa:norhiza sagittata Ca1ochortus gunnisonii Campanu1a rotundifo1ia CovYSocL' Asoigned according to BziO"szpElp BziDzpElp 350-90 160-180 5-25 25-27 10.0 Soc Cov 41.9 Soc Cov 180 10-12 3.8 Soc Cov UNIT NUMBER GM-7 GM-8 BzpGl1Dzp EmcliDzp Emc + + 2-3 1-3 3-4 1-4 3-4 + 1-2 1-2 1-2 1-2 1 1-2 + + 1 +-1 +-1 +-1 +-1 +-1 +-1 1-2 1 1 1-4 +-1 1 1 1-2 2 + +-1 1 2-3 1 28 20-32 315 20-90 17 18-25 25-35 17-22 18.3 .7 7.2 .7 4.0 2.9 2.2 + 3 1-2 1-3 2-3 1-2 +-1 + 1-2 1 +-1 1-3 COV Soc Cov +-1 4 1-4 1-4 1-2 1-3 +-1 +-1 + 2-3 1 1 1-2 1 1 1-2 1-3 2-3 1-2 1 1 + 2 +-1 1-3 1 1-2 2-3 +-1 1-2 1-2 +-1 +-1 1 2 + 1-2 1 1-2 2-3 1-2 +-1 +-1 +-1 1 2-3 1 +-1 +-1 1-3 1 + 1-2 +-1 1-2 1-2 +-1 1-3 +-1 1-2 +-1 1-3 1 +-1 +-1 +-1 1 1 +-1 +-1 +-1 +-1 1 + +-1 1-2 1 1 + + 1-3 1-2 1 1 1-3 1-2 + +-1 +-1 +-1 3 +-1 1-2 3-4 3 +-1 1-4 2 1-2 1-2 +-1 1-2 + 1 1-2 3 1 1-2 1-2 2-3 1-3 1 1-2 1-2 +-1 1-2 1 1 +-1 1-2 +-1 +-1 +-1 +-1 + +-1 1-2 +-1 +-1 1-3 1 3 +-1 1-2 1 +-1 +-1 1 +-1 1-2 1-2 1-2 1-2 1-2 2 1-2 +-1 1 +-1 +-1 1 1-2 1-2 1 1-2 +-1 + +-1 +-1 2-3 1-2 +-1 +-1 2-3 1 +-1 +-1 +-1 + 1 1 +-1 1-3 J +-1 +-1 + 1 1 +-1 +-1 3-4 2 1-2 +-1 +-1 +-1 4 Soc 1-2 +-1 +-1 1-2 1 1 Soc Cov 1-2 2 1-3 1-3 1 1 Soc Cov 1-2 1-3 + 3 1 1-2 1-2 Soc Cov 1 2 ' CM 13 ElpBslDzi 310 2-3 + + +-1 +-1 1 +-1 CM-12 EmlDzp 10-20 FLORISTIC CHARACTERISTICS Soc Cov Soc Cov Soc Cov Soc 3 1-3 cx-i i Em1Dzp 10 +-1 1-2 GM-lO EmlDlzp 125 1 +-1 GH-9 EmliDzpEzp 14-28 340-35 2.2 + Castelleja spp , Chaenactis doug1asii Chenopodium spp. Cirsium spp. cfesac rs columbiana Clematis hirsutissima Comandra. umbellata Crepis spp. Cryptantha spp. Cynogolcssum officinale Delphinium spp. . Descurainia spp , Dodecatheon pu1che1lum Ebilobium spp • Erigeron spp. Eriogonuc spp. Erysimum spp. Fragaria spp • Frasera speciosa Galium boreale Geranium fremontii Hap Lcpappus spp. Heuchera spp . !-telenium hocpes t f Helianthella quinquenervis Ipocopsis aggregata Lathyrus spp. Llnum lewisii Lf r he spe rmue spp. Lupinus argentius Hertensia spp. Oxytropis spp. Pens r eecn spp. Phacelia spp. Phlox app , Potentilla spp. Pricu1a spp. r'seccccveopeerus eont ecua Pulsatilla patens Saxifraga bronchialis Senecio spp. Smilacina spp , Solidago spp. Taraxacum officinale Tragopogon dubius Thalictrum spp. Viola spp. Gramlnoids: Agropyron spp. Bromus spp. Carex spp. Deschampsla caespitoss Festuca spp. Iris missouriensis Ko1eria gracilis Oryzopsis hymenoldes Ph1cum pratense Poa spp. Sitanion 10ngifolium Stipa app , 11 45-180 GH-5 5-17 4.5 Cov BziDzi VEGETATION GM-6 GM-4 + 2-3 1-3 1-2 1 1-2 +-1 +-1 +-1 +-1 1-2 +-1 1 +-1 +-1 + + +-1 +-1 +-1 1-2 +-1 1 +-1 1-2 +-1 +-1 1-2 1 1 +-1 1 3 1-2 1 +-1 +-1 +-1 1-2 +-1 +-1 1· +-1 +-1 + +-1 + 1-2 3 1-3 +-1 1-2 +-1 3 1 1 3 1 1 + 1-3 1 1-3 1 +-1 +-1 1-3 1 1-2 1-3 1 +-1 +-1 1-2 +-1 +-1 +-1 +-1 +-1 +-1 1 1-2 1-3 1 +-1 +-1 1-2 +-1 +-1 +-1 3 + +-1 +-1 + + + + 1 1-3 1-2 1 1 1 +-1 +-1 + 1-2 + 1-2 +-1 + +-1 1-3 2 1 1 +-1 + + + + 1 +-1 +-1 + +-1 1-2 + + 1 1 2 1-2 1 1-2 2 1-2 1 1-3 1 1 1 2 + + 1-2 +-1 1-3 +-1 2 +-1 +-1 +-1 + + + +-1 + + + +-1 + +-1 + + + + +-1 1 1-2 2 1 1-2 2 1-2 + 1-3 2 + 1-2 + ~ 1 1 1-2 +-1 + +-1 +-1 1-2 +-1 +-1 + +-1 +-1 +-1 +-1 +-1 +-1 + + +-1 1 +-1 1-2 1-2 +-1 1 +-1 1-2 +-1 2-3 1-3 1 1-3 +-1 2-3 1-2 1 1-3 1-2 + 1 1-2 +-1 1 +-1 1 1 +-1 +-1 +-1 +-1 + +-1 + 1 +-1 1-2 1 1-2 +-1 2 1 +-1 + +-1 +-1 + 1-2 +-1 +-1 + + + +-1 + +-1 +-1 2 1-2 +-1 ·t-l 1 +-1 +-1 +-1 + 1-3 1 1-3 1-2 +-1 1-2 1-3 1 1-3 1-2 2'-3 1-3 1-2 +-1 +-1 1-2 1-2 + 1-2 1-3 +-1 1-2 1-2 1-3 +-1 +-1 2-3 1 1 1-3 +-1 1 +-1 2-3 1-2 +-1 1 +-1 1-2 1-2 1 1 1-3 +-1 1-2 2 1 1-3 +-1 +-1 1-2 1 1 + +-1 1-2 1-2 +-1 1 1-2 1-2 Kuchler (1955). the phydosnoaic fonaulae reads left ee riabt with the IDOst conspicuous type placed at +-1 +-1 +-1 the 1 +-1 besinnlDS. 1 �168 Elevational DESCRIPTIVE SUMMARY OF VEGETATION TYPE HAP. eREEN MOUNTAINLegal Description: T2S. R80W. Sections 2, 3. 10, 11 Range: 2500-2865 CI Surface Area:....!!!..:.l... Aerhl VEGETATIVE CI...ASSIFICATION AND Classification Physlognou:ic and Features Exposure (Degrees Gradient (Percent) Aeca CH-16 ElpSzpDzp 35 90-190 65-120 17 12-25 35 .9 5.3 1.5 (acc ceees) Cov2/Soc2/ Cov Soc EIIIlpDszp Cov Soc EmizpOlzp SUPPLEMENTARYFEATURES VEGETATION UNIT NUMBER GH-19 GM-20 eM-IS GH-17 EmpDszp Azll11uth) Plant Species List3/ Weody - Trees: Acer glabrum Juniperus virginiana Populus tremuloides Pseudo t suge menzies1i ~ood I-Shrubs: Acer glabrum Amelanch1er alnifoUa Arctostaphylos uva-urs1 Artemisia cana Artemisia tridentata Ceanothus velutinus Cercocarpus montanus Chrysothamnus spp. Holodiscus dumosus Juniperus communis JuniperUS virginiana Mahonia repens Pachystima myrslnites Popu Lua tremuloides P'runus virginiana Pseudotsuga menziesii Pu r sb La t r rden c ar a . Rhcs trllobata Ribes spp . Rosa acicular is Rubus deliciosus Sambucus racemosa Shepherdla canadens!s Syephor-Lc a r pca oreophilus Tet radymia canescens vccc r nt ce spp. Forbs: Achillea lanulosa Anemone spp. Ant enna r La sp p , Aqul1egia caerulen Arabls spp , ArnIca spp . Artemisia f r LgLda A.rtemisia ludovlciana Aster spp. Astragalus spp. Balsaa:orhiza sagittata Calochortus gunnisonii Campanu1a rotundlEolia Castelleja spp. Chaenactis douglas!i Chenopodium spp. Cirsium spp. cteeet is columbiana c.teeac rs hf r suc Ls s Lea Comandra umbellata Crepis spp . Cryptantha spp. Cynogclossum off tc fnej.e Delphinium spp. Descurainia spp. Dodecatheon pulchellum Ebllobium s pp , Erigeron spp. Eriogonum spp. Erysimum spp , Fragaria spp. Frasera spec rosa Galium borea1e Geranium fremontii Haplopappus spp. Heuchera spp. Hc1enium hoopesli Hel1anchella qu1nquenerv!s Iporsop s I s aggregata Lathyrus spp , Lrnue lewisii Lithospermum sppLupinus argent1us Mertensia spp. Oxytropis spp. Penstemon spp . Phacel1a spp • Phlox spp. Potent ilia spp. Pr1mu1a spp. Pseudocymopterus ecneencs Pulsatilla patens Saxifraga bronchiali3 Senecio spp. . Scllacina spp • Solidago spp. Taraxacum officinale Tragopogon dubiuo Thal1ctrum epp , Viola spp. Graminoids: Agropyron app , GM-IS GH-14 1/ BLUE CROUSE INVESTIGATIONS County: Suc=it No. ~ F16CN 10-3-72 Photo EmiDlszpBzp EmiDlp DzpBzpEcpDlp 18-46 35-90 2-20 305-10 17-30 15-25 34 10-20 23.0 2.9 2.6 Cov Soc Cov 2 3-4 3 1-4 ·2 CH-22 GK-21 OlcszpHGlc 20-45 GK-23 GH-24 DmczpHGlc DlcziHGlc GM-25 DmpEmpElspDszp DIIIIGlpHlp GM-26 GlpHlpDsp 25-70 20 30-35 30-40 9-15 5-16 18 18-20 11-22 6-22 a. 9 4.3 .4 2.5 31 2 9 15.9 FLORISTIC CHARACTERISTICS Soc Cov Soc Cov Soc Cov Soc ccv Soc Cov Soc Cov Soc Cov 3-4 2-3 1-3 4-5 1 1 Soc ]22-72 Cov Soc 1-2 +-1 3 1-3 1-2 1-2 1-3 1-2 1-3 +-1 +-1 1-2 + 3 1-3 2-3 4 2-3 1-3 1-2 3 1-3 +-1 1-2 +-1 1-2 1-3 1-3 1-2 1-2 1-3 1-3 +-1 +-1 +-1 1-2 +-1 i +-1 +-1 +-1 1 1 1 1 2-3 +-1 +-1 +-1 1-2 -+-1 2-3 -+-1 +:1 -+-1 +-1 +-1 1 +-1 +-1 +-1 +-1 +-1 +-1 2-3 1 1 1-2 +-1 +-1 -+-1 2-3 1 2-3 1 1 1-2 + +-1 1-2 1 -+-1 +-1 +-1 +-1 2-3 +-1 +-1 1-2 +-1 1 +-1 +-1 +-1 2 1 1 1 2-3 3 1 1-2 +-1 +-1 +-1 +-1 2-3 1 1 1-3 + 1 2-3 1 1-2 2 +-1 1 1-3 1-2 1-2 1-2 -+-1 1-2 +-1 2-3 1 1-2 2-3 +-1 1-2 +-1 +-1 2-3 2-3 2-3 1 1 1 2 1-2 1 1-2 1-2 1-2 1 +-1 +-1 +-1 2 1-2 1 1 1 2-3 1-2 +-1 1-2 +-1 +-1 + 1-2 +-1 1 2 +-1 1 2 +-1 +-1 +-1 +-1 1-2 +-1 2-3 +-1 1-2 1 +-1 +-1 1-2 + +-1 2 2 1 2 2-3 1 +-1 1-3 +-1 + + + -+-1 -+-1 1-2 1 1-2 1 1-2 1-2 +-1 1-2 1 1 +-1 +-1 +-1 +-1 + +-1 1-2 1 + +-1 +-1 +-1 +-1 + +-1 +-1 +-1 +-1 +-1 + +-1 2 1-2 +-1 +-1 +-1 +-1 1 +-1 1-2 +-1 +-1 +-1 +-1 +-1 + +-1 +-1 +-1 +-1 +-1 + +-1 1-2 1-2 +-1 +-1 +-1 2 +-1 + +-1 1-3 2 1-3 +-1 +-1 +-1 1-2 3 1-3 3 +-1 +-1 +-1 1-3 1-2 +-1 3-0 +-1 3 1-2 1-2 +-1 1-2 2 1-2 +-1 +-1 1-2 1-2 +-1 +-1 +-1 1 +-1 +-1 3 1-2 1 +-1 1-2 +-1 +-1 +-1 +-1 + +-1 + +-1 + +-1 +-1 2 +-1 +-1 +-1 1-1 + +-1 + + +-1 +-1 +-1 +-1 1 1-2 +-1 1-2 3 + 1 +-1 +-1 +-1 1 1-2 3 1-2 2-3 1 +-1 -+-1 + + +-1 +-1 +-1 + 2-3 1 + 1-2 1-2 +-1 +-1 +-1 +-1 +-1 +-1 + + + 1-2 1-2 1 1-2 1-2 +-1 +-1 1 1-2 1-2 1 1-2 2 1 -+-1 + ,1 2-3 + +-1 -+-1 1-2 1 + + 1 1 1 2 +-1 +-1 +-1 1 1-2 1-2 +-1 + +-1 1-2 2 1 +-1 1 -+-1 2-3 +-1 + +-1 + +-1 1-3 1 1-3 +-1 +-1 1-2 +-1 +-1 +-1 +-1 3 1-2 +-1 +-1 +-1 1 1 1-2 +-1 + 1-2 �169 DESCRIPTIVE SUMMAltYOF VEGETATION TYPE HAP. lIlY ClI.!!;K-BLU!. CROUSE IKV!.STIGATIONS Legal ~.ct'1pt1on: T1S. R84W,Sec. 31 T4S, 184W.Sec. 4,5,6 County: !.agle !lev.tional Ranle: 2S00-Z896. Surhe • .\1:"': 482 VEGETATIVE ;~;:!!!~:~!!? Features and Exposure (DegreeD (Hectaru) .urn cu.sSIrICATION 140-180 IS-3D 32.1 " 5,] Co.,!' LhJ.1 spede. So~1 Cov Soc Cov Soc see 1-2 2-3 1-2 1 1-2 1 1-2 1 1-3 1-3 1 CS-Ufl-1l4 7-6-51 FUTUUS 5UPPI.DC!NTARY NUMB!R variable 5-28 EC-9 EC-6 EC-7 !C-B th:18zpHlcClc 21. BzpDnpHG Daoez1Hmpl1Cli '" 35-190 7-17 ·68.2 10-19 13.0 6-25 s,c 101.8 FLORISTIC Plant No.: BzpDziHG BdOlpdHC 25-30 12.4 Photo VEGETATION UNIT 8dDit1RG 25-40 20-]0 70-260 6-22 18.0 Gradient (Percent) Area az1D~hpHG BdDziHG .uilMath) Aerial ,,,"3 EC-2 I!N h.a o..cJ:ispHC 140-215 8-30 e.a OlAJIAcrElUSTlCS see see cev Soc Cov Soc cev Soc Woody-Treea: Alnus tCDuifoU. Juniperua virginiana Populua ball.lfcrll Populull trelllUlo1dea . Wood -Shruba: Acer glabnlll Alnua tenuifol1a Az:Ielanchh:r dnifolia Artemisia 1-3 tridentata Ccrcocarpua IDOntanuG Chrysotha.mnua Cornua app. 1-2 3 -+-1 2-3 1-3 3 2 +-1 2-3 1 '-3 '-3 '-3 1 , +-1 1-2 1 e~nia Juniperus virginiana iDvoluerata repena • "" PachystiDa lIIyt.initea Pentaphyllo1dcB flodbuDda Populus Ptunua delieioaua epp , "" H RollI!. ae1cI,I1&t111 Salix 2 1-2 2 1-2 1 Putl1hia ttidentlltll Ribcs app , H H 1-2 1-2 1-2 2-3 1-3 +-1 1-3 1-2 1 1 1 1 2-3 1-3 '-3 1-2 caneaceno ~I lanulosa 1-2 1-2 +-1 , +-1 app. Agoaeris AlliUJ:I glauca app , Angelico app. Antennaria +-1 Dpp. , +-l +-1 2-3 1 1 2-) 1-3 1-3 +-1 _I + 1-2 , +-1 z +-l Aqullcgla cllerulca Arabia app. Atel1arla 1 1-3 1-3 + +-1 tubrs Agastache 1-' 1-' 1-2 colWllbianum Actaea 1-2 2-3 1 1-2 Tctradymia Aconitu:a. 2-3 H Sambucus rael'!llO'& S)'mphor1ellrpoa oreophilus Achilles +-1 H ttClllulo1dcs vitginiana Rubus 1-' 1-' lI:olonif.ra Juniperua Lonicera Mahonia 3 '-3 +-1 2-3 1 +-1 2-3 +-1 congeal's ·Arnlca I 1 app. ArtCllliaia dracum:ulua Mtc..lsia frigida ArtCllliala A,atragalus ludovlciona app. 8.alsl1lllOrhlza Caloehortus Capselb +-1 +-l '-3 sagittata +-1 2-3 1-2 1-3 + + r-a 1-3 +-l + +-l +-l 3 gunnisonii burea-pasto\'la Cardaaine 1-' 1-' '-3 eordifoU..o. Caatillejs 2 app. Chenopodium Ciraium app. +-1 app. Clt:l:Lllti.eepp. COllinda +-l +-1 parviflora Cllloaia l1nellria Com:mdrll umbel lata ofUelnale npp. Deaeuralnla epp. Dodeelltheon pulc:hellulII Eplloblum Equiaetum , +-1 l-3 app. app. umbellat •.••• EryaimulI.llsper= fragarla .•.. , 1-' +-l 1-2 1-' a _1 1 1-2 1-3 +-l r-z +-l + +-1 1 1 a r-z '-3, 1 '-3 +-1 3 z +-1 + +-1 +-l +-l +-l 1-' '-3 1 1-2 , ,, '-3 +-l 1 1 + +-1 3 -+-1 2-3 +-1 1-2 1 +-1 fremont 1 Hack.elia floribunda Helenium hoopesii Heli.nthella 1-2 11 +-l +-1 1 + +-1 +-1 +-l + +-l quinquenervis HeraeleuQ +-1 2-3 3 npp. Ge\'llnium GeuQ spp. +-1 1 specioaa Calium +-l '-3 2 +-1 +-l +-l 3 r-a 1-3 1-3 '-3 1 1 + 2 +-l +-l t-a + +-l +-1 t-z 1 H + +-l ,, 1 3 ovaUa Fraser. + +-1 +-1 1-', upp , Erigeron Erlogonum '-3 '-3 + Cre?iaspp. Cynoglossum Delphinium , +-l 1 1 + lanatuill 1 ~l 2-3 +-1 ~l 1-2 1 +-1 +-1 Heueheraapp. lpol!lOpls Lappulll aggregllta redovakii Lathyrus Llgullticul!I +-1 IIpp. IIpp. +-1 + +-1 + +-1 lew1811 Linlllll Llthoaper"l!lUl!l. =ultiflonua Lo1lllltiw:aspp. w?inull argenteus He-ctcnaill app. ltemophlll1 brevi OomorhlZ11 8pp. Oxypol1s + +-l +-l +-l flora 2 1-3 1 , + +-1 1 + +-l fendlerl P~icu18r1s .•..+ , 1 1 +-l 2-3 app. Phaccl1a r-a l-3 1-' 1 1-2 +-l PhYIIDrl11 1-' ~I 2 + +-1 .pp. officlnale Thalietr\lll Thla.pi 1-' 1 + +-1 l-2 1 1 1 1 +-1 2-3 2 1 1-2 1 1-2 z +-l +-l +-1 1 ~1 1-2 +-1 , +-1 1-2 1-3 l-3 1-' 1-3 +-1 +-l 1-2 dublu. tenuipetal\lll .pp. Viguiera +-1 2 1-3 1-3 +-1 1 + 1-2 +-1 +-1 2-3 3 3 IlUltinora app. +-1 Zysadenu. Gr_1noida: eleg.ne Agropyron app. pp. +-1 1-2 1 1-' .pp. +-1 1-2 app. 1-2 1-2 +-1 1-3 1 2-3 +-1 1-' 1-2 1 2 1-3 2-3 1 1-' 1-3 1-' 1-2 ea •• p1to •• app. lracilta .•.. , 1-' +-l bulbo•• Phleum praten.1I: Poa epp , 1-' 1-3 1-3 +-1 + +-1 1-2 +-1 1-2 1 1 '-3 +-1 +-1 +-l 1-2 '-3 +-1 +-1 1-3 1-2 1-3 1-2 1-2 1-2 +-1 1-3 1 1-2 + +-1 1-' 1-'1 1 1-3 +-1 '-3 3 1 +-1 1 1 '-3 +-1 '-3 8pp. HordeUlli brachyantheru:ll Iria ai.nourien.ls ..Juacu. app. epp . 1-3 1-3 gl_rata oeaeha.p814 El)'"IDUn spp. Sti!!a , a spp. Veronf,ea Helic. 1-' dioica Veratl'l,lll J.oelll:ria +-1 app. Trifoliua Felltuca + +-1 1 .pp. Tugopogon Clyceria 3 1-3 +-1 app. Stellaria Urtica +-1 1-2 opp. Solidago ••.••• r-z +-1 +-l 1-' cllndUa '!.rax.ecu. Care:x + +-l +-1 +-1 lllciniau lanecalata ScaUaclna 61'0 1 1 app. Sidalcea DIlctylia 1 upp , Rudbeckia Scrophularla. Agroatia 2 2-3 1-' IIpp. Ranunculua Viola + +-l +-1 1 Potent1l1allpp. Senecio +-1 1 IIcopulorum Polygonum 1 , 1-2 1 vitul1hra Plaglobothrya +-l 1 1-' 3 spp. epp , +-1 +-l +-1 a 1 1 spp. Penotemon Phlox z 1-' 1 1 , , +-1 +-1 +-1 2 +-1 1-2 1-3 1 1 1-3 �170 DtsC1.1PTlVE lAS_l !lev.tlond SUMKAlY or 'I!G!TATION ns, o..criptlon: RanIa: 2500-2896. U4W. tTlE Sec. Surfac. IW'. ]1; EIIY T4S, Ar •• : VlClTAftVl CAlEX-Bt.U! IIBIlW. Sec. 482 ha ClOUSE County: Photo CLASSIFICATION" (Perc.nt) Gradient Mea se-n EO-IO o.c:.Pj6-."SS11Cl1 190-210 10-)0 12.7 24-40 65.0 (Hcctaru) EO-I] DlcapzlHC 180-220 10-)0 4.1 lC-I2 o.e.pziHC AInulI Soc! eav!' o.1nlfoU Artem1aia Juniperuo cosaunil Juniperu' virginiana .•.. , repena Pachyoti_ Popu11,lo tremu10idea .1 1-) 2-3 1-2 acicu1ario ).1 7-30 1.' 8.4 SYl=Ipnor 1carpos a-a )-4 ,-) ,-) a-a 1-' ...., Cov 2-) 1 +-1 2 ...., 1 Achillea , Agastache 1-4 1-3 1-2 1 1-2 2 +-1 I I 2-) 2 1-2 1-2 1-2 2-3 1 1-) I +-1 1-2 1-) 1-) ...., +-1 1-2 ...., .•.. , ...., +-1 1-2 +-1 1-2 81a"c. app. Angelico Opp. Antennar1:l spp. Aquilegia opp. Arnic. spp. a-a +-1 + caerulea Arabia Arenaria tongest. Artemisi. dracunculua ArtCI:I!si4 frigida Artel:lillttl l"dovi~lanA luIeragalu. Btl1oallOrhiza 1-' opp. sagittet. Calochortua ...., ....,+ , spp. Ciraiul:lOpp. CICQ4tis 1-3 ...., .•.. , + ...., ...., , +-1 1-2 I +-1 + +-1 + +-1 opp. CoIUnoia parviflora Collomitl l1neo.ris Comarnlrll umbellata +-1 ....,+ ...., ...., offic1D.1l1e Delphinium sp.,. tcscurolr:la app , Dodecathcon pulchellUli EpllobiUIIIIlPP· Equbetuc opp. -+-1 +-1 -+-1 ...., spp. ....,+ fremonel1 floribunda HelcnlulII boO)peo1i Heli.mthella ...., ...., ...., H +-1 ...., quinq"encrvio HeracleUI:I !polllOpis + +-1 redowakii L4thyrua spp. LtgUDticulIl Ilpp. ...., +-1 1-2 1-' I I , +-1 1-2 ...., ) l-2 +-1 1-3 +-1 .•.. , .•.. , .•.. , +-1 +-1 1 1-) +-1 +-1 H .•. +-1 2 2-) H ...., I ....,+ .•.. , .•.. , ...., ....,+ ...., ...., ...., ....,+ lewi9~i Llthollperllll,llll ml,lltiflorUlll Lom.:atiUIII opp. Lupinull Mertensia ...., argente"o IIpp. NCDOphlla breviflora OOlllOrhlza opp. Oxypolh z + ...., I fendled PedlCl,llariGspp. Pen9tcIlIon I I I I ...., 2-3 opp. Phaceliu +-1 1-2 H .•.. , +-1 +-1 ) 1-2 + +-1 1-3 ...., + Senecio Sid.lce. laoceolat. ...., .pp. Stell.ria opp. Tara14C\,lll officinale ThalictNIII ThllUllpi +-1 I app. H ...., 1 1-2 I +-1 1-2 1 I +-1 , tenuip app. .•.. , I .•.. , I 1-2 I 1 1-3 +-1 .•.. I 1 , 1-' ...., ...., 1-' , .pp. app. 1-' 1-2 1-2 1-) 1 I-J 1-' 810ller"'t. Dcllcha.psia 1-3 1-3 I +-1 -+-1 +-1 2-3 1-3 1 1 1 2 + +-1 +-1 + ...., 1-' app. spp. o.ctylil I 1 1-2 1 I ) ...., I +-1 1-2 1-2 1-3 I Graminolda: Agroltis 2-3 ...., .•.. , eleganl Agropyron +-1 1-2 I e tal\lll CNltifloro. Ipp. Zygadcnuo 2 +-1 H 1 opp. Viguiera + +-1 + TrllBoposon dubiull T1:1foli •••• "pp. Urrico. dioic. Veratrua Veronica 1 ...., , ,-) Ipp. Solidaao 2-3 I +-1 ...., 1-' app. candida SlIIllo.cin.ca 1-3 .•. +-1 acopulol'lla laciniato. Scrophuloria 2-3 I opp. epp. Rudbeck1a 2-3 2-3 .• 1 +-1 Potent1l1a Ranunculuo I +-1 z 1-2 ...., vitulHera Plaaiobothryo. Polygonu:lll app. I ...., H I app. Phyaaria + +-1 +-1 ...., I I ....,+ +. opp. caellpitoall EIYl=lua app. Featuca I 2-3 allugata Lappula BrOl:lUa I +-1 -+-1 H .•.. , l"Mtum app. geuchera 2 1-2 2 .•. I Geulll spp. Hllckelll!. I +-1 ...., umbellat"l:1 Fragaria ova119 Frllsero.speciosG GcranlW11 1 2 1 + I 2 1-2 Ery:!Ii.lllWil ;J.!)perum Gallum 1-3 .•. -+-1 1-2 I ...., El'igeronspp. El'iogonulll 2-3 _1 _1 +-1 + Cl'epbspp. CynogloslilulII +-1 I ,, epp , ChenopodiUIII 1-2 i-a buraa-pastor1. cordifolia C:ise1lleja +-1 +-1 gunnillon11 Capllella Cardam1ne 1-' +-1 1-2 1 1-3 ...., I + +-1 I +-1 1-2 3 1 2-3 1-2 I ...., I-l I 1-2 1-3 I 1-2 1-2 1-3 2-3 1-3 1 1-3 +-1 +-1 1-) opp. Glyceria app. HordeUIII .•.. , ~ac:hyantherUIII i-z Irismis!lourienois Juncua a-a spp. Koeleria + H H ) gracilis Melica ...., bulbosa Phleull Stipa 1-2 )-4 2 + .•.. , app. Agoaeria AlUuIII 1-' .•.. , ...., + rubra 1-' ...., 1-' ...., lonulolla Actaea Poa +-1 ...., 1-' Aconit\llllcohm.bi.= Carex 1-2 2-) 2 1 Soc 1-' ) orcophllua 1 2-) 1-2 2 a-a 1-' ...., 1-2 1-' Forbo: Vi~lo 32.0 +-1 eancncens TetradYl=lio Phlol!; 59-220 5-)7 s,>< '0' Soc a app. SG.QbucuorAcemoaa Linum Ee-III D.~pHC D·:!:r~PIP S-11 CHUAcnnlSTICS Cov Ru.'bua del1eiool,la S41b EC-17 !C-16 ~~~~g ao Soc 1-' tridentata app. Roua Ee-I) Dl~i&BC floribunda vifginiaD.ll Purahta Ribe:. NUMBER lIIyrainitea Pentaphylloidell Prunull 7-6-51 nATURES 1-' ...., ...., invo1uerat. Mahonu Cov 1-' I Cbryoothamnuo opp. Cornuo oto1onifera Lonicera 1-' z-a .• tridentata aontanull ceecceerpue Soc 3-4 tenuifolia Aac18m;hier Ccv bah GS-t.N2-1J\ Eo.14 nottISTIC Plant Speciea U ••tll Woody-Trees: Alnus tenuifoaa Junlpcrul virginian. Populul bala_iter. Populull trcau.loidea Wood -Shrub.: ker alabru. No.: AND SUPPLEMlNTARf VEGETATlON UNIT ClAOslflclltlon and Featur." Physlogno=aldl Exposure (Degrel!lI Ad.QUth) INV!STlCATIONS 4,5,6 Aedd pratense 1-' z spp. 1-) ...., I 1-2 1-2 spp. 11Assigned according 1/ Coverage and l/IIllPorta-r.t to) Kuchlo.r sociAbility plant apecies (19.5.5). the aaaSgned on the according buts of phyaiogoo.lc to coverage. Kuchler Plant 2-3 1-3 ...., I fGflIula reada ...., ...., left to l'ight with the (1955). no_nelature folJ.!IVlI Weber (1972). ...., , 2 1-3 H !IIOllt cOII,spimou$ type p Laced ...., , 2 I at 1-3 1-2 I 1 tbe beginning. 1-3 2 1-) 1-2 1 1 �171 DESCRIPTIVE Legal Elevat10nal SlDIQRY OF Description: Range: VEGETATION T1S. 2500-2896.. TYPE 1l84W. Sec. Surface MAP. 31; Area: !IBY T45, 482 CREEK-BLUE RB4W, Sec. ha CROUSE 4,S,6 Aeri.l VEGETATIVE CLASSIFICATION and Phyaiognomlc:,.!./ Exposure (negreee Gradient (Percent) ~p.. EC-I9 EC-20 DazpElrHC Dd.pRC Features ,., 20-30 (Hectares Dapzi1'llpHG 10-25 ,., 6-29 '.1 6.' FLORISTIC Plant Species So, L1BJ: 7-6-51 UNIT NUMBER DIal.18pzlH11pliCli 170-190 5-13 6-15 11.7 25.1 EC-27 EC-26 EC-25 EC-24 HlICI10zp 20-79 130-246 14. 6-2. !.ISle CS-Ul2-134 EC-23 EC-22 Ou~pHG 200-220 21. Azauth) EC-21 No.: AND SUPPLt:KENTARY FEATUR!S VEGETATION Classification INVESTIGATIONS County: Photo DupHGlc: DlpezpHG Dal1sz1HG ,-, 175-195 117 '-6 '.6 2lS ,., 6-' ~:)--- CHARACTERISTICS Cov Soe So, So, Cov Soc 2-3 2-3 Cov Soc: WOOdy-Trees: Alnus tcnllifo11a Juniperus vlrglnlana PopuluB baloamHera Populus 1-2 1-2 treauloldea 1-' 2-3 Wood -Shrubs: kef' gilibrum Alnuo , tenuifolia Aaelanchler alnifolia Artt=lsia 1-2 1 1 2-' 1-2 tridentata cerecceescs IIIOntanua 1-2 2-3 1-2 1-2 Cornua otolonlfera co-.nia +-1 repeno Pachyat1ao Populus , 1-2 1-2 1-2 +-1 floribunda 2-' H trcmuloldea 2-' 2-' Prunus virginians Purohl0 tridentata 2-3 ,-, 2-3 app. aclcularia Rubus 1 1-2 lIIyralnltea Pentaphylloides Ribes Rosa 1 1-3 .. virginiana invo]1,ICrat:a Hahonia 2 1-2 1-2 Juniperus Jl,mil'erus Lanicera 2-' 2-' 1-2 2-3 2-' spp , Chryaothamnus 2-'2 1 1-2 2 1-2 1-2 1-2 1 2 2-3 "-2 2-3 1-2 SYlllphoricarpoa Tetradymio 1-' oreophlluB caneacena 2-' 2-3 +-1 1-2 1 , 1 2-3 1-2 , dellclosua Salix spp. SOlllbucuaraceoQ(lsa 1-2 1-2 1 2 +-1 3 2 +-1 1 1-2 +-I +-I +-I H Forba: +-I Achillea lanulooa Aconitum c:olumblanulII Actaea 1-2 1-2 +-1 opp. AgollcJ;'io glauco AlliUIII +-1 1-2 +-1 +-1 +-I · spp • Antennaria Aqullegia opp. Artemisia drac:unc:ulua ArtClllo1a Artettiaia frigida ludovic:iana +-1 1-2 A!Jtrag.al»a spp. Rall,l8111Orhba sagittata Caloc:tlOrtuo 1-' 1-2 gunnioonl1 Capoella 1-3 1-2 1 . buroa-paotoris Card.!lllline Calltilleja cordifolia app. CiraiWl +-I +-1 +-1 app , ChenopodiuCl opp. Cle=latis Collinllia spp • p8rviflora Collomia lincario Coaaandra Wloollat.!l 1 +-1 2 +-1 +-1 1-2 1 2 +-1 +-1 . 1-2 1 +-1 +-I +-1 app. Cynoglossllll • +-1 1-2 +-I +-I app. Dcacur.inia opp. Dodecatheon pulchellua Epilobiull opp. EquisetuCi spp. +-I +-1 ulllbclbtum Erysimulll asperum Frllgorill oval +-1 +-I +-, +-1 1 1-2 +-1 1-2 . lanatulII aggregata redowalr.U Lathyrua Ligusticum Llnum app , spp , lewisii Llthospen:lum lIIultiflorum LomlltiuCl upp , Lupinull aegeneeue Mertensia spp. NemophUa brevifloro Osmorhizo spp. Oxypol1s 1 1 +-1 +-1 candida app. app. +-I +-1 +-1 +-I +-I 1-2 1 . 1 1-3 1-2 +-1 • +-1 +-1 2-3 1 +-1 +-1 +-1 1 1-3 2 1-) 1 1-2 1-3 1-2 1-2 +-I 2 app. +-1 elegana opp. 1-2 1-2 1-2 1 1 1-2 1-) 1 2-3 2 +-1 1-2 epp1 1-2 opp. 8loll('.r8ts caespitoa& app. spp. app. btachyantherUII is80urlenois app. +-1 +-1 grae11io Helie. bulboa& Phlellll +-1 1-2 1 1 • +-1 +-1 2 2-3 , 1-2 1 • 1 • +-I • 1 +-1 +-1 1 +-1 • 1 2 +-I 2-) +-1 2-3 +-1 +-1 1-3 1 1 1-2 1-3 1-2 2-' 1-2 app. !I Assigned according 1:.1 Coverage and to aociability plant epecfee Kuchler (1955). a •• t,gne.cf on the 1-2 1 +-1 the phyalogD01ll.ic according basis of to covera.e. 1 Kuchler Plant 2-3 2-3 • 1 2-3 2-3 +-1 1-2 1-2 1 fOl'lllUb readll , to right rith the 1-2 1 1 1-) 1 2 1-3 , 1 2-3 2-3 +-I , 1-' 1-) followa Weber (1972). con'sPi01oUS +-1 1 2'-3 2 1-2 2 2 1 .oat • +-1 2-3 type placed at the 2-3 1-3 +-1 2 2-,, +-1 2 1-3 +-1 2 1 left +-1 +-1 1-2 1-2 2 (1955). nOllM!nc1ature +-I +-1 2 +-1 1-2 1-2 1 H +-1 +-1 1 1 1-2 +-1 +-I 2 2 pr.tense app. l/Itiportsnt 1-2 1 +-I 1-2 Feutuc:a C1yceri. HordeUSI koeleria +-1 2 +-1 +-1 +-I 1 Viguiera 1:IU1tiflora Viola epp , E1}'IIUa +-I +-1 1 2-' Vel:Ooica Deschallpo:la 1 +-1 1 2 2 2 1-3 dubiuo app. tenulpetalUIII Agrootia Br~aapp. +-I +-I 1 1 dloica Zygsdenuo +-1 1-2 1-2 2 +-1 +-1 +-I +-I +-1 +-1 Veratnn. Daetylis 1 1-2 a app. Crallinoids: Agropyron +-I +-I 1-2 spp~ Urtica 1-2 1 spp. officil181e Tragopogon Trifol1U111. +-1 · +-I + +-I 1-2 1 2 Stellaria TaraXAcWII Thalictrllll 1-2 1 +-I app. Thlaapi +-I + +-I 1aoceo1ata Sailacins. Solidago Stip. 1 1-2 1 2 spp. Sida1cea Poa +-1 +-1 1 1-2 +-1 2 1-2 1 1-2 laciniata Scrophu1aria Senecio 2-) 1-2 +-I 1 +-1 spp. Rudbeclr.1a 1-2 +-1 +-I • +-1 +-1 vitulifera Ranunculua • +-1 +-I 1-2 1 2-3 • +-1 P1ag10bothrys scopu10rua Po1ygonull spp . Potentllla 1 +-1 1 · +-I +-I epp , Phyaar1a 1 +-1 +-1 + +-I 1 2 . spp. Phlox 1-3 1 +-1 +-1 +-I spp. spp. Phac:efia +-I +-I +-I +-1 +-1 1 fcndleri Pedieulal'iB senateeon +-1 +-I 2 +-I Heucheraapp. . Lappula 1-2 1 quinquenervl0 lpo:lopls +-1 1 f lor ibunda hoopclll1 Hcracleulll 1-2 +-I 1-2 2-3 is Geraniulll frClllOntii CeUIll app , Hclianthella , 1-2 +-1 FrllBcrllBpeciosa Gnlium spp. Hackelia Hclenium 1-2 2-' Erigeronspp. Eriogonulll +-1 +-1 2 1-2 ·· +-I officinale Delphinium 1-2 +-1 1 2-3 create Iri •• Juncus 1-2 +-1 app. caerulea Arabia app. Arenario congeata Arnica +-1 +-1 1-2 opp. Angelica Carex +-I +-1 1-2 ruhra Agllatac:he be.ginning. 1-2 2 ) 2-3 1 1 1-2 2 1-' ��173 April JOB PROGRESS St~te of Colorado Project .No. W-37-R-34 9 Work Plan No. Job Title: Response Period Covered: Personnel: J. B. C. D. 1981 REPORT Game Bird Survey Job No. 6 of Blue Grouse to Aspen Silvicultural Practices 1 April 1980 through 31 March 1981 Dickerson, M. Ward and F. Wild, u.S. Forest Service; Wilcox, Colorado State Forest Service; T. Beck, Braun, B. Cade, H. Funk, T. Hobbs, R. Hoffman and Miller, Colorado Division of Wildlife. ABSTRACT Location of a study area and development of a detailed study plan were priorities of this research project in 1980. Whereas a suitable study area has been located, efforts to obtain the desired level of treatment for development of a feasible study design have been unsuccessful. Lack of a market for quaking aspen (Populus tremuloides) wood and rising production costs have reduced the area that can be economically manipulated. Alternative means of obtaining the desired level of treatment are being considered. Implementation of this project is dependent upon the cooperation and a firm commitment of assistance from the u.S. Forest Service in conducting the necessary treatment and seeking an economical and practical means of increasing the acreage that can be manipulated. ��175 RESPONSE OF BLUE GROUSE TO ASPEN SILVICULTURAL PRACTICES Richard W. Hoffman Blue grouse (Dendragapus obscurus) are widely distributed in the mountainous areas of Colorado occupying varied habitats from shrublands to mature coniferous forests. This species is avidly pursued by sport hunters and the blue grouse is a major upland game resource in Colorado. Studies of blue grouse in Middle Park and near Eagle have focused on population dynamics and habitat use. These studies have identified the aspen type as being of major importance for blue grouse breeding and brood rearing activities. Aspen is a subclimax type and covers hundreds of thousands of hectares in Colorado and adjacent states of the Central Rocky Mountains. Management of aspen by public land management agencies will have tremendous impacts upon the blue grouse resource. The intent of this developing study is to identify blue grouse response to aspen management practices. To successfully accomplish this study, it will be necessary to have the cooperation and a firm commitment from public land management agencies, primarily the U.S. Forest Service. P. N. OBJECTIVE Prepare a detailed study plan describing specific hypotheses and/or objectives, procedures, schedules, costs, manpower requirements, etc., and locate a suitable study area for initiating field investigations. SEGMENT OBJECTIVES 1. Review literature pertinent to the subject of this study. 2. Visit potential 3. Prepare a detailed study plan outlining dures, and study sites. study sites and select a suitable study area. specific objectives, proce- METHODS AND MATERIALS Field personnel of the Colorado Division of Wildlife (CDOW), U.S. Forest Service (USFS) and Colorado State Forest Service (CSFS) were contacted regarding potential study areas. Field surveys were conducted in most recommended areas. Some time was spent in consultation with D. C. Bowden, Statistician, concerning design of appropriate sampling and analytical procedures necessary to reliably measure the response of a blue grouse population to aspen manipulation. Efforts were initiated to review and assemble literature pertinent to the subject matter of this study. �176 RESULTS AND DISCUSSION Study Area Selection Field personnel of the Colorado Division of Wildlife, U.S. Forest Service and Colorado State Forest Service were contacted for suggestions on study areas. General surveys were conducted on 5 potential study areas including Stoner Mesa, Groundhog Mesa, Terror Creek, Schaefer Creek, and Slater Creek. Four of the 5 areas are in south.western Colorado; 2 (Stoner and Groundhog mesas) in the San Juan National Forest and 2 (Terror and Schaefer creeks) in the Gunnison National Forest. Southwestern Colorado supports the largest, continuous stands of aspen in the state. Baker (1925) estimated that approximately 695,109 hectares of aspen cover types occur on national forest lands in Colorado of which 65% (451,821 ha) was in the southwest portion of the state. The only area investigated outside of southwestern Colorado was Slater Creek, Routt National Forest, in northwestern Colorado. There are few other areas where large scale aspen manipulation programs are currently in progress or where programs will be implemented in the near future. Most timber sales and wildlife habitat improvement projects involving aspen are done on such a small scale as to preclude any studies to measure impacts on wildlife populations. The problem stems from the lack of a commercial market for aspen wood accompanied by rising production costs which have reduced the area that can be economically manipulated. As a consequence, commercial operators are only interested in harvesting small areas and economic restraints have limited the amount of noncommercial timber that can be managed solely for wildlife habitat improvement. The Terror Creek management unit was considered the most promlslng area for conducting this study. Stoner Mesa, Groundhog Mesa and Slater Creek have already been subjected to aspen manipulation; consequently, no pretreatment evaluation could be performed on these areas. In addition, Stoner Mesa and Groundhog Mesa are not readily accessible for conducting breeding surveys in early spring. Access until late Mayor early June would be by snowmobile. Schaefer Creek fulfilled the study area criterion but the proposal to manage aspen in this area for watershed protection has not progressed past the district level planning stages. It is highly unlikely this project will be implemented in the near future. Terror Creek is located about 11 km north of Paonia, Colorado in portions of T125, R91W and T125, R92W, Delta County, Paonia Ranger District, Gunnison National Forest. Exact study area boundaries have not been delineated, but the area being considered is roughly bounded by Big Alder Creek on the north, the Overland ditch on the west, Cunningham Creek on the south, and the Steven's Gulch road on the west. This area encompasses approximately 2,995 hectares and is comprised almost entirely of the following cover types: aspen-tall shrub (2,066 ha), aspen-spruce fir (450 ha), aspen-tall forb (375 ha), and aspen-riparian (104 ha). More than one-half (58%) of the area has been classified as determinant stands (even-aged), most of which is in the mature and over-mature age classes with virtually no regeneration in the understory. �177 The drainage where treatment is contemplated ranges in elevation from 1,890 to 2,866 m. Access is from the Hubbard Creek and Steven's Gulch roads. Low standard roads exist along east Terror Creek, Cunningham Creek, Betty Park, and along the Curecanti-Hayden Powerline. Four domestic grazing allotments (3 cattle and 1 sheep) are within the proposed unit. USFS Proposed Treatment The Terror Creek Wildlife Habitat Improvement Unit was established during winter 1980. Management plans and an Environmental Assessment Report were prepared by USFS personnel. The plans dealt primarily with aspen and Gambel Oak (Quercus gambelii). Primary objectives for the project in aspen are to regulate the determinant aspen timber to sustainedyield and even distribution of age and size classes and to enhance wildlife habitat by diversification. Photo evaluation and stand mapping was completed before field work began in July 1980. Stage II inventory was completed on the unit during summer 1980. Standard timber information was gathered during this time along with a basic wildlife survey. Aspen cover types were classified as determinant (even-aged) and indeterminant (uneven-aged) and each category was further delineated into productive and nonproductive stands. Productive stands are defined as those capable of producing 1.4 cu m/ha/yr (20 cu ft/acre/yr) of sawtimber material and which are suitable for silvicultural practices. Only the productive, determinant aspen was designated for treatment. This included commercial (net volume/acre> 5,000 bd ft/acre) and noncommercial or cultural treatment (net volume/acre < 5,000 bd ft/acre). Since the indeterminant aspen supposedly would be self-perpetuating, it was not designated for treatment. Computed areas of aspen in each category varied (Fig. 1). ASPEN /2,995 , " / ha ~ Determinant 1,753 ha ' ! -, Unproductive 305 ha / Productite 1,448 ha \ Indeterminant 1,242 ha Unproductive 149 ha \ \ Commercial Harvest 924 ha >., Cultural Treatment 524 ha \ Total Treatment 1,448 ha // Fig. 1. Estimated areas of aspen classification Stage II Inventory. -. Productive 1,093 ha types based on USFS �178 A separate regulation schedule was established for cultural and commercial treatments. Both treatment types are based on a 60-year rotation wi t h 6-10 year cutting cycles (treatment periods). 1. Cultural Treatment -- There are 524 hectares designated for cultural treatment. Using a 60-year rotation, the average annual harvest would be 8.7 ha/yr or 87 ha/10 yrs. All stands designated for cultural treatment were prioritized according to defect percentage. At 1 end of the scale were stands with 0% defect and 0 net volume/ha which were basically seed saps or small poles. These stands were considered least in need of treatment and were placed in the far treatment periods (i.e. 5th and 6th). At the other extreme were stands with 100% defect and 0 net volume/ha which were severely decadent and in immediate need of treatment. These stands were placed in the first cutting period. The remaining stands wer e ranked according to defect percentage with the higher percentages rated as higher priority for treatment. Size of the areas were adjusted somewhat to maintain an even flow of cutting at about 87 ha/10 yrs. Aspen Cutting Plan Period Hectares to be treated 1 (First 10 years) 94 89 85 85 85 2 3 4 5 6 ~ Total 524 Since most of this volume will not be used except possibly as free fuel wood, no attempt will be made to remove the timber. The actual area treated in any 1 year is not binding as long as each unit of treatment is completed within a 10-year period. For instance, the 94 hectares designated for treatment in the first 10 years need not be divided equally among these years, but can be treated in any combination of ways including cutting the entire acreage in 1 year. The only stipulation is that the 94 ha be cut within the 10 years. 2. Commercial Treatments -- There are 924 hectares designated for commercial treatment. Based on a 60-year rotation, the average annual harvest would be 15.4 ha/yr or 154 ha/10 yrs. Since the volume in this treatment will be removed for commercial purposes, a transportation system must be developed. The area was divided into 6-10 year cutting periods. Priorities were established on the basis of present wildlife needs, expected transportation system needs, and distribution of treatment units to obtain desired habitat diversity. The area within period 1 would be the most accessible and receive the heaviest wildlife use. Periods 2-6 are increasingly less accessible and of decreasing importance to wildlife. Area to be treated within each period are: �179 Hectares Period 1 2 3 4 5 6 Total 183 135 145 154 151 156 924 For regulation to occur within the 60-year rotation, each 10 year unit must be treated in a 10 year period. However, the order in which each 10 year unit is treated is dynamic and can be changed according to current conditions. For example, if after the first 10 year-units (183 ha) are treated, there is a need to change priority area 5 to 2, this can be done without disrupting future age class distribution as long as the integrity of each 10 year-unit is maintained. Overcutting in any period would congest the age classes at 1 end of the spectrum and leave a deficiency at the opposite end. Undercutting would extend the time period for regulation beyond a rotation. Unlike the cultural treatments, commercial treatments are dependent upon market conditions and must be imposed on an annual basis to stay even with demand. Study Status and Recommendations Cooperation and a firm commitment of control over the treatment to be imposed are prerequisites to further development of a detailed study plan. No such commitment has been obtained. The USFS's proposal has not progressed beyond the planning stages and there is no assurance that the proposal is economically feasible considering the financial situation and the current unstable market for aspen wood. Whereas the proposal appears operational on paper, it's practicality is questionable as there are too many factors which could interrupt adherence to the cutting schedule; most notably, a continued lull in the aspen market and further restraints on spending by federal agencies. It is recommended that the USFS abandon their proposal, at least temporarily, and consider the possibility of a cooperative study to examine the impacts of aspen manipulation, specifically clear-cutting, on wildlife populations. This is based on the premise that before large scale aspen manipulation programs are implemented, controlled experiments should be conducted to document wildlife responses when aspen is disturbed. A suitable study area has been located but efforts to obtain the desired level of treatment for development of a feasible study plan have not been successful. Economic restraints created by the lack of a market for aspen wood have reduced the amount of acreage that commercial operators are willing to cut below that which any response from the blue grouse population can be accurately measured. It is therefore imperative for successful completion of this study to continue working with U.S. Forest Service personnel in seeking alternative means of increasing the amount of acreage that can be manipulated. The most practical alternative being considered is to use a combination of �180 services including (1) commercial timber sales to local operators, (2) service contracts for removal of unmarketable timber, and (3) a Colorado State Forest Service logging crew to cut the additional area necessary to attain the desired level of treatment. It is recommended that the planning phase of this project continue in an effort to meet specific research requirements because: 1. Aspen forests have extremely important wildlife values. 2. Forest managers have expressed interest in, and even requested input from, wildlife specialists in developing strategies for aspen management that are consistent with wildlife needs. 3. It is anticipated that silvicultural prescriptions for managing aspen will eventually become a standard practice throughout Colorado as manipulation of aspen forests is essential for regeneration and perpetuation of the species. 4. Large scale aspen manipulation programs can have tremendous ficial implications to wildlife populations. 5. Both big game and nongame researchers have expressed in conducting studies related to aspen manipulation; potential exists for a multispecies approach. bene- an interest thus, the In conjunction with the planning phase, study plots should be selected and field work initiated to collect pretreatment information. If by the end of 1 year there is no confirmed commitment to cut the required area, the project should be discontinued until a commitment can be obtained. Pretreatment information should provide further evidence to support the hypothesis that maintenance of diversity in size and age classes in aspen stands will enhance grouse populations. Hypotheses which have been developed a. Determinant, homogeneous aspen stands support low densities of territorial male blue grouse. b. Determinant, homogeneous aspen stands support low numbers of female blue grouse with broods. c. Determinant aspen stands supporting low densities of blue grouse can be manipulated to increase population densities. The approach 1. are: to be followed Continue reviewing blue grouse. in 1981-82 is: literature concerning aspen manipulation and �181 2. Continue cooperative efforts with the U. S. Forest Service and Colorado State Forest Service to obtain the desired level of treatment necessary to meet research requirements. Primary concerns to be considered are: (1) the present lack of a market for aspen to obtain maximum use of the harvestable resource, (2) capabilities of a commercial logging operation to complete the project within a specified time period, (3) economic feasibility of service contracts for removing noncommercial wood, (4) development of an additional transportation system, (5) adverse effects of excessive domestic grazing on treatment and control plots, and (6) economic feasibility of subsidizing logging operations. 3. Randomly select 3 pairs of study plots of approximately 81 ha/plot in determinant aspen stands designated for treatment. All plots will be selected on the basis of similar vegetative and structural characteristics with each pair of plots being most similar to 1 another than to other pairs. One plot from each pair will be randomly selected for treatment with the other plot designated as the control. Selection of study plots will be based on Stage II Timber Survey maps of the Terror Creek Unit depicting (1) aspen cover types (aspen-tall shrub, aspen-tall forb, aspen-riparian, and aspen-spruce-fir), (2) stand status (determinant or indeterminant) , (3) stand location, and (4) stand priority for treatment. 4. Layout the actual treatment to be imposed on each of 3 study plots including the number, size, shape and specific boundaries of each clearcut. Approximately 28 ha in 1-4 ha blocks will be assigned for treatment in each plot. Treatments will be designed to maintain homogeneity between plots. 5. Conduct pretreatment evaluation. a. Using established techniques, trap and individually mark as many blue grouse as possible on each study plot. Repeated observations of marked grouse will provide data for plotting of territories and ranges of individual birds. These data will be analyzed for (1) differences in territory size between study plots prior to treatment, (2) alterations of territorial boundaries resulting from treatment effects, and (3) differences in territory size between treated and untreated plots. b. Estimate the density study plot. Census using tape recorded territories located future reference. c. Estimate the number of individual broods using each study plot with census procedures as in 5b. Brood use areas and actual sightings will be plotted on aerial photos for future reference. of territorial male blue grouse on each procedures involve a systematic search calls and a trained pointing dog. All will be plotted on aerial photos for �182 d. Measure structural variables (slope, aspect, canopy coverage, number of existing openings> 0.5 ha, shrub. density, and amount of edge) in each study plot and correlate with the density of territorial males and broods estimated for each plot. These measurements are essential for evaluation of pretreatment conditions and to account for possible differences in densities between study plots prior to treatment. 6. Develop study plan for the implementation of post-treatment evaluation provided all research requirements are satisfied. As these study plans are developed they will be appended to and become part of the initial Program Narrative. 7. Compile data, analyze results, and prepare progress or completion reports. Recommendations will be formulated concerning the continuation of research efforts and management practices. LITERATURE CITED Baker, F. s. 1925. Aspen in the Central Rocky Mountain Dep. Agric. Dep. Bull. 1291. 46pp. Prepared by ~t{(YU2"V (~ckat<Z' a./ . Ri.chard W. Hoffmart Wildlife Researcher (jf!) C Region. U.s. �183 April 1981 JOB FINAL REPORT State of Project Colorado W-37-R-34 No. 13 Work Plan No. Job Title: Period Distribution Covered: Personnel: Game Bird Survey -----Job No. 7 and Status of Mountain Sharp-tailed Grouse 1 April 1979 through 31 March 1981 Mike Bauman, Clait Braun, Ken Giesen, Van Graham, Jim Hicks, Donald Hoffman, Robert Mangus, and Charles Woodward, Colorado Division of Wildlife. ABSTRACT First-hand sightings of mountain sharp-tailed grouse (Pedioecetes phasianellus columbianus) by Division of Wildlife personnel and members of bird clubs in Colorado in recent years have been limited. Data gathered in recent years by Colorado Division of Wildlife personnel is largely limited to counts of sharptails on a few dancing grounds in Routt and Moffat counties and wing survey data obtained through wing barrels and hunter check stations. A minimum of 123 mountain sharp-tailed grouse of both sexes was counted on 15 active dancing grounds in Routt and Moffat counties in 1979. A minimum of 24 sharptails on 5 active dancing grounds was counted in 1980. The average number of sharp tail males per active ground was 5.5 and 4.8 in 1979 and 1980, respectively . .Between 14 and 79 sharp tail wings have been collected annually in wing barrels and at hunter check stations in western Colorado since 1976; nearly all from the California Park area in Routt County. Analysis of these wings indicate rapid turnover in the adult segment of the population. Questionnaires circulated to field personnel in western Colorado indicated few recent confirmed sightings of mountain sharptails outside of Routt and eastern Moffat counties. Moderate populations exist in Rio Blanco County while remnant populations occur in Gunnison, San Miguel and Dolores counties. �184 Reported sightings in recent years, numbers of sharp tail wings collected annually in wing barrels during the open seasons, and findings from this investigation all indicate populations of this game bird are low on a statewide basis with populations being disp.ersed over extensive areas of seemingly suitable habitats. Highest present day populations are within Routt and Moffat counties where mountain shrub habitat is being lost through coal strip mining activities, agriculture, and homesite developments. It is expected that this trend of habitat loss will continue in the future because of the need for energy and the resulting increase in human population. RECOMMENDATIONS 1. Efforts should continue to locate leks, brood areas and winter use areas of sharptail grouse in western Colorado. 2. Counts of males and females on selected leks in Routt and Moffat counties should be conducted annually to determine the relationship between lek counts and population size. 3. Investigations of seasonal habitat requirements of the mountain sharp-tailed grouse in western Colorado should be initiated. 4. Investigations of important population parameters (age and sex structure, productivity, annual survival and mortality rates) should be initiated. 5. Hunter harvest in Routt and Moffat counties should be moni~ored using check stations, wing barrels, and hunter questionnaires. �185 DISTRIBUTION AND STATUS OF MOUNTAIN SHARP-TAILED GROUSE Kenneth M. Giesen Donald M. Hoffman Mountain sharp-tailed grouse have been hunted annually in Colorado since 1953 with season dates coinciding with those of sage grouse (Centrocercus urophasianus) and bag and possession limits in the aggregate with those of sage grouse. Little inventory work has been accomplished on mountain sharp-tailed grouse since 1962-65. This report includes all data collected during a study designed to gather and update available information on the current status of the mountain sharp-tailed grouse in Colorado. P. N. OBJECTIVE To increase the knowledge of the distribution grouse in Western Colorado. METHODS and status of sharp-tailed AND MATERIALS Literature searches were accomplished in libraries of Colorado State University and the Colorado Division of Wildlife Research Center in Fort Collins. A personal reference library was also used for this purpose. Questionnaires designed to obtain population information, sighting locations and names of knowledgeable individuals in regard to sharptails were circulated. Completed questionnaires were summarized and some follow-up interviews were conducted with individuals having first-hand knowledge of past or present distribution and status of sharp-tailed grouse in western Colorado. Visits were made to museums maintained by colleges, universities and the Denver Museum of Natural History. All mountain sharp-tailed grouse specimens were examined for location,and date of collection and name of the collector. Where personal contact could not be made with the museum curators, correspondence was initiated. Visits were made to offices of the Colorado Division of Wildlife in Denver, Fort Collins, Grand Junction and Montrose to obtain management and population information on sharp-tailed grouse in western Colorado. A spring census of dancing grounds in Routt and Moffat counties was conducted in 1979 and 1980 to count males and total birds on known dancing ground locations and locate new grounds whenever possible. Procedures used were similar to those described by Rogers (1969). �186 Attempts were made to locate and count sharp-tailed grouse broods along trails and roads and in potential habitats during summer 1979. Searches of potential winter habitats were conducted from mid-October 1979 through Mid-March 1980 to locate and count sharp-tailed grouse on wintering ranges. Locations of all known active dancing grounds, brood, winter and miscellaneous sightings were recorded on U.S. Geological Survey topographic maps for Routt and Moffat counties and on other suitable maps for other counties in western Colorado. Names of individuals securing the sighting or count, number of birds, and period of the year were recorded. RESULTS AND DISCUSSION Distribution and Status Information on historical and current distribution of the mountain sharptailed grouse in Colorado obtained from the literature, Museum specimens, a questionnaire, and field investigations has been compiled and summarized (Rogers 1969, Hoffman 1980). Although previous methods for population inventory of sharptails were inadequate, it is clearly apparent that the distribution and abundance of the mountain sharptails have declined in historic times (Miller and Graul 1980). The status of most remnant populations of mountain sharptails in Colorado is unknown as no systematic effort has been made to monitor these populations and their habitat. The known present distribution of the mountain sharptailed grouse in western Colorado (Fig. 1) was delineated from field investigations and follow-up of positive responses to the questionnaire. Undoubtedly many remnant populations are not included. Furthermore, little is known about size or status (increasing, decreasing, or stable) of the sharptail population in Routt and Moffat counties where they are apparently most abundant. Dancing Ground Counts Male and female sharp-tailed grouse were counted on 15 active leks in 1979 and 5 active leks in 1980. The low number of leks counted in 1980 was due to poor access in March and the retirement of the principal investigator (D. Hoffman). The average number of males per active lek in 1980 was the lowest recorded since 1964 (Table 1). The biological significance of sharp tail lek counts is poorly understood as little is known about patterns of lek attendance by male and female mountain sharptailed grouse. Observed annual variation in the average number of males per active lek may be due to count date, weather, variation between leks (not all leks were counted each year), personnel involved in counting and other factors. �MOFFAT RIO BLANCO MESA .•..... 0;) -._J MONTROSE SAN MIGUEL DOLOREb MONTEZUMA Fig. 1. The known present distribution of the mountain sharptailed grouse in western Colorado. �188 Table 1. Sharp-tailed grouse dancing ground counts Moffat counties, 1964-65 and 1977-80. Year No. active grounds counted No. of males 7 15 2 8 15 5 32 87 16 65 82 24 a 1964 1965a 1977 1978 1979 1980 a Data from Rogers in Routt and Average no. of males per active lek 4.6 5.8 8.0 8.1 5.5 4.8 (1969). Wing Collections Wings of hunter harvested sharp-tailed grouse were collected annually in Routt and Moffat counties since 1976, primarily through the use of voluntary wing collection stations (wing barrels). In 5 years 256 sharp tail wings were collected, of which 129 (50.4%) were from juveniles. Yearly samples of wings were small because wing barrels were primarily located to obtain sage grouse wings rather than wings from sharptails. Also, it, is possible that few wings were received because of low hunting pressure for sharptails in Routt and Moffat counties. Most sharp tail wings (178 of 242 wings collected since 1977) have come from a single wing barrel (California Park Road) although the exact location of harvest is unknown. Methods for ascertaining sex of mountain sharptails from wing characteristics are not available, thus limiting the amount of data obtained. Two age classes (juvenile and birds 1 year and older) are separable using wings and analysis of the wings indicate high production of chicks from 1977-80 (Table 2). Table 2. Age and sex composition of hunter harvested grouse in northwestern Colorado, 1976-80. Adultsa Year N 1976 1977 1978 1979 1980 10 47 13 32 25 Totals Juveniles % 71.4 65.3 46.4 40.5 39.7 127 5-year Average aAdults sharp-tailed and yearlings. N 4 25 15 47 38 % 28.6 34.7 53.6 59.5 60.3 129 49.6 Total Sample 14 72 28 79 63 256 50.4 �189 LITERATURE CITED Hoffman, D. 1980. Distribution and status of mountain sharp-tailed grouse. CoLo rado Div. Wildl. Job Prog. Rep., Fed. Aid Proj. W-37~R-33. April 1980. Pp. 325-347. . .Miller, G., and W. Graul. 1980. Status of sharp-tailed grouse in North America. Pp. 18-28 IN Proc. Prairie Grouse Symp. Oklahoma State Univ., Stillwater. Rogers, G. 1969. The sharp-tailed grouse in Colorado. Game, Fish and Parks, Tech. Publ. 23. 94pp. Prepared by ~ J1! ~ --~~--~~~-=~~~------------Kenneth M. Giesen Wildlife Researcher A Colorado Div. ��191 April 1981 JOB PROGRESS State of C_o_l_o_r_a_d_o _ Project No. W-37-R-34 Job No. 1 Upland Game Publications Job Title: Period Covered: Personnel: Game Bird Surve 22 Work Plan No. REPORT 1 April 1980 through 31 March 1981 T. D. I. Beck, C. E. Eraun, L. H. Carpenter, S. Emmons, K. M. Giesen, E. O. Hahn, T. A. May, B. E. Petersen, and T. J. Schoenberg, Colorado Division of Wildlife. ABSTRACT Publications accomplished under this job in Segment 33 are: Beck, T. D. I., and C. E. Braun. 1980. The strutting ground count: variation, traditionalism, management needs. Proc. West. Assoc. State Game and Fish Comm. 60:558-566. Braun, C. E. 1980. Alpine bird communities of western North America; implications for management and research. Pages 280-291 in R. M. DeGraff and N. G. Tilghman, compilers. Workshop Proc. Management of western forests and grasslands for nongame birds. U.S. Dep. Agric., For. Servo Gen. Tech. Rep. INT-86. Carpenter, L. H., and C. E. Braun. 1980. A new approach to the dilemma of federal cost-sharing and technical assistance programs that alter wildlife habitat. Proc. Annu. Summer Conf., Central Mtns. and Plains Sect., The Wildlife Society 25:Abstract. Emmons, S. R. Colorado. 1980. Lek attendance of male sage grouse in North Park, M.S. Thesis, Colorado State Univ., Fort Collins. 69pp. , and C. E. Braun. 1980. Breeding season ---selection of male sage grouse in North Park, movements Colorado. and habitat J. Colo.- Wyo. Acad. Sci. 12(1):36. Giesen, K. M., and C. E. Braun. 1980. Dispersal of juvenile whitetailed ptarmigan. Annu. Mtg. Am. Ornithol. Union. 89:Abstract. �192 Giesen, K. M., C. E. Braun, and T. A. May. 1980. Reproduction and nestsite selection by white-tailed ptarmigan in Colorado. Wilson Bull. 92:188-199. 1980. Hormonal induction Hahn, E. D., and C. E. Braun. pigmentation in ptarmigan. Auk 97:601-607. of feather Petersen, B. E. 1980. Breeding and nesting ecology of female sage grouse in North Park, Colorado. M.S. Thesis, Colorado State Univ., Fort Collins. 86pp. _______ , and C. E. Braun. 1980. Patterns of growth rates and weight gain by juvenile sage grouse. J. Colo.-Wyo. Acad. Sci. 12(1):43. Schoenberg, T. J., and C. E. Braun. 1980. and female sage grouse. J. Colo.-Wyo. Spring habitat use by male Acad , Sci. 12(1) :43-44. ______ ~' S. R. Emmons, and B.E. Petersen. 1980. Telemetry investigations of sage grouse in North Park, Colorado. Annu. Mtg. Am. Drnithol. Union 89:Abstract. P·repare d b y: /\. eJ?a,-C .L.) (;. -1-?ACt<.. •..• '-' Clait E. Braun (il].)' Wildlife ResearchvLeader � https://cpw.cvlcollections.org/files/original/de4e65da1634b7099d2c6b386706fb3b.pdf 00cf117e31f68039752ae126e2055fd8 PDF Text Text July 1981 1 JOB FINAL REPORT State of Colorado Project No. W-126-R-4 -------------------1 Work Plan No. Job Title: Multispecies Period Covered: Personnel: Big Game Investigations Job No. Investigations 2 - Deer and Elk Management Study July 1, 1980 through June 30, 1981 Regional Wildlife Biologist Staff members: John Ellenberger, John Gray, Jim Olterman, Mark Elkins, Jack Vayhinger, Gene Schoonveld, Jim Dennis, Area Supervisors and District Wildlife Managers; Big Game Supervisor, Bob Hernbrode; T. M. Pojar. ABSTRACT All four regions now have access to remote computer terminals in their r_egional offices. All regions also have Regional Inventory Biologists with expertise to operate the ONEPOP model. Full staffing of the Inventory Biologist positions was not accomplished until this past segment when the Southwest and Northeast Region filled these positions. Therefore, most effort was spent training and consulting with personnel from these two regions. All regions now have independent population modeling programs. Only minimal consulting and maintenance of computer files at CSU will be necessary from this point on for all regions to continue a viable modeling program. ��3 DEER AND ELK MANAGEMENT STUDY Thomas M. Pojar P. N. OBJECTIVE Devise and implement and continually improve a statewide big game management system. Provide technical and educational assistance to management personnel in implementing modeling programs and to train the Big Game Supervisor in the use of the ONEPOP model. RESULTS AND DISCUSSION For several years personnel of the Big Game Management project have worked closely with the Colorado Cooperative Wildlife Research Unit of Colorado State University and Colorado Division of Wildlife's (DOW) various management sections. The objective of these cooperative activities has been to develop, test and implement a big game population simulation modeling system to augment DOH's big game management program. Key personnel in all 4 DOW Regions have, been trained in the use of the system and remote computer terminals have been installed in the Denver Office and all 4 Regional offices. The operation of the entire system has, as of July 1, 1981, been transferred to these various management sections and the Big Game Management Study is terminated. Several publications (see References Section) are available that describe the development, validation and management use of the ONEPOP model. Anyone contemplating using this model should be familiar with these publications to ensure proper understanding of the limitations and legitimate use of the model. REFERENCES Model Documentation Gross, J. E., J. E. Roelle, and G. L. Hilliams. 1973. Program ONEPOP an information processor: A system modeling and communication project. Progress Report Colorado Cooperative Wildlife Research Unit, Colorado State Univ., Fort-Collins, Colo. 327p. Roelle, J. E. and J. M. Bartholow. 1977. User documentation: Program ONEPOP. 44 xerox pages. (Available from T. M. Pojar, Colo. Div. Wildl. Res. Center, Ft. Collins, Colo. 80526). Model Validation Bartholow, J. M. 1977. Fort Niobrara Refuge: Big game management M.S. Thesis, Colorado State Univ., Fort Collins, Colo. 224p. modeling. Roelle, J. E. 1977. Refuge management modeling: The National Bison Range. Ph.D. Dissertation. Colo. State Univ., Fort Collins, Colo. 3llp. �4 Williams, G. L., 1977. Simulation modeling Wildlife Refuge. Ph.D. Dissertation. Colo. Management of big game at Wichita Mountains Colorado State Univ., Fort Collins, use of Model Pojar, T. M. 1981. A management perspective of population modeling. In: Fowler, C., and T. Smith (eds.). Dynamics in Large Hammal Populations. John Wiley and Sons, New York.' (In press). --- , and D. Strickland (Eds'.). 1979. A workshop on the status and applicationof big game population modeling. Fort Collins, Colo. Jan. 16-17, 1979. 53p. (Available from the author, Colo. Div. Wildl. Res. Center, Fort .Co IlLns , Colo.). Salwasser; H. and T. M. Pojar. 1979. Simulation modeling of pronghorn populations. Mcnuscript submitted to: J. Yoakum (~d.), Pronghorn ment. Wildlife Management Institute. ;. ~ Prepared by /"1 i . { t".,~(!l' ./ Thomas M. Pojar Wildlife Researcher ' " Manage- �July 1981 5 JOB PROGRESS Colorado State of Project REPORT No. W-126-R~4~ Work Plan No. Job Title: Big Game Investigations Job No. 4 Multispecies Investigations - Animal and Pen Support Facilities for Deer/Elk Research Period Covered: Personnel: I _ July 1, 1980 through June 30, 1981 P. Neil, B. Gill, Dr. T. Spraker, S. Torbit, R. Parachini, C. Smith, D. Parkinsen, V. Jameson, J. Reeves, P. Smith, B. Van Santo ABSTRACT Modifications to the facility and minor construction projects were completed to facilitate on-going research projects and in preparation for up-corning programs. A study was conducted to examine and compare the endocrine system and thymicolymphatic systems of stressed (captive) and normal (free-ranging) pregnant mule deer and their fetuses in order to demonstrate that stress of the pregnant animal causes alterations within the fetal endocrine and thymicolymphatic systems. Preliminary results from 5 captive, stressed females suggest a moderate thymic atrophy and adrenal cortical hyperplasia in the adult deer and early thymic degeneration in two of eight fetuses. Intensive training of mule deer and elk for use in digestion cages occupied a major portion of the segment. The Animal and Pen Support Facility presently houses 21 mule deer, 7 elk, 10 pronghorn, 6 bighorn sheep and 7 Rocky Mountain goats. ��7 ANIMAL AND PEN SUPPORT FACILITIES FOR DEER-ELK RESEARCH Paul H. Neil P. N. OBJECTIVE To provide and maintain populations of captive big game animals facilities to support big game research programs. and pen SEGMENT OBJECTIVES 1. Continue to develop and maintain Foothills campus. 2. Coordinate rearing, training, and research activities with captive wild and tame big game animals for all big game research projects. 3. Integrate big game animal and physical plant facilities, resources, and fiscal resources under a single budget. METHODS a big game research facility at CSU human AND MATERIALS Materials were ordered to complete several modifications at the facility during the segment. Personnel from the Physical Plant metal shop at Colorado State University were enlisted to construct several modifications for the digestion cages. Cement work was completed under the cages and the collection holes under the cages were enlarged to accommodate larger collection bottles when working with elk in the cages. The Young-Adult Youth Conservation Corps of the Bureau of Reclamation was enlisted to assist with construction of a perimeter fence at the facility and other minor construction projects. Three yearling female elk and one 3-year-old bull elk were obtained from the Wildlife Disease Center at Colorado State University to supplement the DOW captive elk herd. With the exception of the bull, all were incorporated into an intensive training program for use in the digestion cages. Dr. Terry A. Spraker of the Colorado State University Diagnostic Lab used the DOW facilities to study the effects of stress on apparent immunocompetence of mule deer. Five pregnant wild mule deer does were trapped and placed in captivity at the facility for the purpose of evaluating the effects of stress on the endocrine system and thymicolymphatic system. Three of 10 antelope at the facility are being used for a Pronghorn Breeding Study described under W-126-R, WP 5, J 1. The remaining seven pronghorn are being maintained for nutritional studies scheduled for the winter of 1981-82. �8 RESULTS AND DISCUSSION All of the modifications and small construction projects conducted during the segment were for the purpose of preparing for up-coming digestion trials and to facilitate on-going research. The perimeter fence was constructed for dog control and security purposes. Five pregnant wild mule deer does were trapped during the last week of February and the first week of March 1981 and placed in the DOW facility. These 5 deer were kept in captivity for approximately 2 months for the purpose of evaluating the effects of stress on the endocrine system and thymicolymphatic system. This information is to be related to disease incidence and survival rates of fawns. The deer were euthenized the 3rd week of May 1981 and were necropsied. Parameters measured and examined grossly and histologically included the peripheral lymph nodes, spleen, kidney, brain, thymus, and adrenal glands of the adults and their fetuses. Preliminary results suggest there is a moderate thymic atrophy and adrenal cortical hyperplasia in the adult deer following two months of stress. Preliminary results from the necropsy of the 8 fetuses did show an early thymic degeneration in two animals. Further work is presently being done with these samples at the Veterinary Diagnostic Laboratory at Colorado State University and a complete analysis and report are pending. Two elk died of apparent "chronic wasting syndrome" during the segment which reduced the elk .research herd to 3 animals, a number insufficient for up-coming digestion trials. Three yearling female elk and one 3 year-old bull elk were obtained from the Wildlife Disease Genter at Colorado State University to supplement the herd. Training of mule deer and elk intensified during the later portion of the segment in preparation for digestion trials to be conducted during the winter of 1981-82. Details of this project are described under W-126-R, WP 3, J 3. The total research herd presently consists of 21 mule deer, 7 elk, 10 pronghorn, 6 bighorn sheep, and 7 Rocky Mountain goats. The "chronic wasting syndrome" is still a major problem among our captive mule deer and elk. The Pathology Department and Diagnostic Laboratory at Colorado State University are continuing their investigations as to the cause of this problem. Prepared by ~tf.-(\~( Paul H. Neil Wildlife Technician III �9 JOB PROGRESS State of REPORT Colorado --~~~~~---------- Deer-Elk Investigations Pro j ec t No. ;:.W_-,:1.:Z.,::::6_-,::.:R,-_.....:4 _ Work Plan No. Job Title: Period Job No. 5 Experimental Improvement of Oakbrush on Deer, Elk, and Cattle Ranges - Hightower Mountain Covered: Personnel: 1 July 1, 1980 through June 30, 1981. R. Kufeld ABSTRACT A lO-year post treatment evaluation of burning, spraying, and chaining was initiated during the last portion of the segment following procedures used in previous evaluations. This is the last evaluation that will be conducted on this study. ��11 EXPERIMENTAL IMPROVEMENT OF OAKBRUSH ON DEER, ELK, AND CATTLE RANGES - HIGHTOWER MOUNTAIN Roland C. Kufeld P. N. OBJECTIVE To determine the extent to which deer, elk, and cattle forage production, forage qualit~ and game use can be increased and maintained by chaining, sprayin~ and controlled burning of over-age Gambel oak winter game ranges. SEGMENT OBJECTIVES To determine the extent of vegetative composition and forage production changes, and extent of deer and elk use changes which have resulted from implementation of each habitat improvement method tested. Determine if cattle exhibit a preference toward certain habitat improvement units. METHODS AND MATERIALS Detailed Methods and Materials are presented by Kufeld (1978). A final report on this study will be presented next segment. LITERATURE CITED Kufeld, Roland C. 1978. Experimental improvement of oakbrush on deer, elk, and cattle - Hightower Mountain. Colo. Div. of Wildl. Game Res. Rep. July (3):335-411. ? Prepared by r i~eafoteic/ J?& r . ,~' Roland C. Kufeld Wildlife Researcher C '; i ��13 JOB PROGRESS State of REPORT Colorado --~~~~~---------- Deer-Elk Investigations Pro j ec t No. .:.:.W:.....-=1.::.2~6-_:R~-._4.:...._ _ Work Plan No. Job Title: Period Digestible Covered: Personnel: Job No. 1 Nutrient Content July 1, 1980 through R. Kufeld, 6 of Deer and Elk Winter Forage Plants June 30, 1981. M. Stevens ABSTRACT Chemical analyses and digestibility of serviceberry (Amelanchier alnifolia) and mountain mahogany (Cercocarpus montanus) was initiated during this segment. Lab results will be completed and a manuscript summarizing this work will be prepared next segment. A manuscript entitled "Winter Variation in Nutrient and Fiber Content and In vitro Digestibility of Gambel Oak (Quercus gambe1lii) and big Sagebrush (Artemisia tridentata) from Diversified Sites in Colorado" was published in the Journal of Range Management and the abstract is in Appendix A. ��15 DIGESTIBLE NUTRIENT CONTENT OF DEER AND ELK WINTER FORAGE PLANTS Roland C. Kufeld P. N. OBJECTIVE To estimate average nutrient content and digestibility values and the degree of variation within selected range forage plants during winter. SEGMENT OBJECTIVES To determine the degree of variation in nutrient content and digestibility of selected range forage plants during winter. METHODS AND MATERIALS Methods and Materials for this study have been described previously (Kufeld 1979). LITERATURE CITED Kufeld, Roland C. 1979. Digestible nutrient content of deer and elk winter forage plants. Colo. Div. Wild1. Game Res. Rep. July (1):47-51. Prepared by ~~J.,,-ciC 'K.JM Roland C. Kufeld Wildlife Researcher C ��17 APPENDIX A �18 WINTER VARIATION IN NUTRIENT AND FIBER CONTENT AND IN VITRO DIGESTIBILITY OF GAMBEL OAK (QUERCUS GAMBELLII) AND BIG SAGEBRUSH (ARTEMISIA TRIDENTATA) FROM DIVERSIFIED SITES IN COLORADO Roland C. Kufeld, Marilyn Stevens, and David C. Bowden ABSTRACT Nutrient and fiber content and in vitro digestible dry matter (IVDDM) were measured in Gambel oak (Quercus gambellii) and big sagebrush (Artemisia tridentata) samples collected during January from nine geographic areas distributed widely throughout the western half of Colorado, and representing three vegetation types. Coefficients of variation among areas were less than 10% in both species in dry matter content, IVDDM and most cell and cell wall components. Variation appears to be small enough to permit application of a suitably selected, constant value, which would reflect winter nutrient content, fiber content or digestibility of these species, regardless of where collected in Colorado, in surveys where winter nutritional status of big game rangelands is being estimated for management purposes. �July 1981 19 JOB PROGRESS State of Project REPORT Colorado ---------------------Big Game Investigations No. W-126-R-4 ------- 1 Hork Plan No. Job Title: Multispecies Period Covered: Personnel: Job No. Investigations 7 - Big Game Research Publications July 1, 1980 through June 30, 1981 Listed in citations in text ABSTRACT During the 1980-81 segment the Big Game Research Section had 13 publications published and 9 others accepted for publication. ��21 BIG GAME RESEARCH PUBLICATIONS R. Bruce Gill P. N. OBJECTIVE To publish the results of research conducted under the auspices of Federal Aid Project W-126-R in a variety of professional journals and other indexed publishing media to insure widespread dissemination and availability of this information to natural resource managers and ecological scientists. SEGMENT OBJECTIVES 1. Carpenter, L. H. Twenty-four hour activity patterns of mule deer at pasture. J. Wildl. }~nage. 2. Kautz, M. A., G. M. Van Dyne, L. H. Carpenter, and W. W. ~utz. Energy costs of some activities of mule deer fawns. J. Wildl. ~nage. 3. Torbit, S. C., L. H. Carpenter, D. M. Swift, and A. W. Alldredge. Body composition estimation in mule deer; a comparison of methods. J. Wildl. ~nage. 4. Torbit, S. C., L. H. Carpenter, D. M. Swift, and A. W. Alldredge. Depletion dynamics of mule deer energy reserves. J. Wildl. Manage. 5. Bartmann, R. M. An evaluation of winter diet selection by tame mule deer. J. Range Manage. 6. Bartmann, R. M., and L. H. Carpenter. Influence of prior foraging experience on tame mule deer forage selections. J. Wildl. Manage. 7. Bartmann, R. M. Distribution and movements of mule deer in the Piceance Basin, Colorado DOW Special Report. 8. Bartmann, R. M. Winter severity indices and mule deer winter mortality in the Piceance Basin, Colorado. J. Wildl. ~nage. 9. Bartmann, R. M. Tame mule deer forage selections and diet qualities on pinyon-juniper winter range in the Piceance Basin, Colorado. DOW Special Report. 10. Rutherford, W. H., and W. D. Snyder. Guidelines for wildlife habitat manipulation - a manual. DOW Division Report. 11. Milchunas, D. L. and D. L. Baker. In vitro digestion with respect to wild ruminant nutritional analyses. J. Wildl. Manage. 12. Hobbs, N. T., D. L. Baker, and R. B. Gill. Dietary overlap of elk~ deer, and bighorn sheep in Rocky Mountain ~ational Park. Ecological Honographs. �22 13. Hobbs, N. T., and D. L. Baker. Energy and nitrogen based estimates of elk winter range carrying capacity. J. Wildl. Manage. 14. Baker, D. L., and N. T. Hobbs. Nutritional quality of elk diets in alpine and ~ubalpine habitats, Rocky Mountain National Park, Colorado. J. Wildl. Manage. 15. Baker, D. L.,and N. T. Hobbs. Botanical composition and nutritional quality of elk winter diets in the upper montane zone, Colorado. J. Wildl. Manage. 16. Big Game Investigations Federal Aid Job Progress Reports. PUBLICATION PROGRESS 1. Carpenter, L. H. Twenty-four hour activity patterns of mule deer at pasture. J. Wildl. Manage. First draft of this manuscript is 3/4 completed. 2. Kautz, M. A., G. M. Vail Dyne, L. H. Carpenter, and W. W. Mautz. Energy costs of some activities of mule deer fawns. J. Wildl. Manage. This manuscript has been submitted to JWM and is in the review process. 3. Torbit, S. C., L. H. Carpenter, D. M. Swift, and A. W. Alldredge. Body composition estimation in mule deer; a comparison of methods. J. Wildl. Manage. This manuscript was intended as one publication product of Torbit's Ph.D. Thesis. No progress was made on the actual manuscript although the Ph.D. Dissertation is completed and has been accepted. 4. Torbit, S. C., L. H. Carpenter, D. ~. Swift, and A. W. Alldredge. Depletion dynamics of mule deer energy reserves. J. Wildl. Manage. Same comments as previous article. 5. Bartmann, R. M. An evaluation of winter selection by tame mule deer. J. Range Manage. This manuscript was submitted to JWM and has been accepted for publication. 6. Bartmann, R. M., and L. H. Carpenter. Influence of prior foraging experience on tame mule deer forage selections. J. Wildl._Manage. This manuscript has been accepted for publication by JWM. 7. Bartmann, R. M. Distribution and movements of mule deer in the Piceance Basin. Colo. Div. Wildl. Spec. Rep. This manuscript was published as: �23 Bartmann, R. M .• and S. F. Steinert. 1981. Distribution and movements of mule deer in the.White River Drainage, Colorado. Colo. Diy. Wildl. Spec. Rep. No. 51. 12pp. 8. Bartmann, R. M. Winter severity indices and mule deer winter mortality in the Piceance Basin, Colorado. J. Wildl. Manage. This manuscript is half-way through the 1st draft edition. 9. Bartmann , R. M. Tame mule deer forage selections and diet qualities on pinyon-juniper winter. Colo. Div. Wildl. Spec. Rep_ Data analyses have been completed for this manuscript, but the narrative has not yet begun. 10. Rutherford, W. H., and W. D. Snyder. Guidelines for wildlife habitat manipulation - a manual. Colo. Div. Wildl. Spec. Rep. A 1st draft edition of this manuscript is prepared, but a decision from Wildlife Research Chief, R. M. Hopper, on whether to proceed with publication, is pending. 11. Milchunas, D. G., and D. L. Baker. In vitro digestion with respect to wild ruminant nutritional analyses. J. Wildl. Manage. This has been accepted for publication by the Journal of Range Manage~ ment as follows: Milchunas, D. G., and D. L. Baker. In vitro digestion--sources of within and between trial variability. J. Range Manage. 12. Hobbs, N. T., D. L. Baker, and R. B. Gill. Dietary overlap of elk, deer, and bighorn sheep in Rocky Mountain National Park. Ecol. Monogr. A draft copy of this manuscript has been prepared, has passed through internal DOW review and has been submitted to selected outside reviewers prior to submission for publication. Currently, the manuscript has been reentitled and plans are to submit it to the JWM. New title is as follows: Hobbs, N. T., D. L. Baker, and R. B. Gill. Comparative nutritional ecology of montane ungulates during winter. J. Wildl. Manage. 13. Hobbs, N. T., and D. L. Baker. Energy and nitrogen based estimates of elk winter range carrying capacity. J. Wildl. Manage. This manuscript has been accepted for publication in JWM as: Hobbs, N. T., D. L. Baker, J. E. Ellis, D. M. Swift and R. A Green. 1982. Energy and nitrogen based estimates of elk winter range carrying capacity. J. Wildl. }~nage. 14. Baker, D. L., and N. T. Hobbs. Nutritional quality of elk diets in alpine and subalpine habitats, Rocky Mountain National Park, Colorado. J. Wildl. Manage. �24 This manuscript has been accepted for publication in JWM as: Baker, D. L., and N. T. Hobbs. summer diets in Colorado. 15. Composition and quality of elk J. Wildl. Manage. Baker, D. L., and N. T. Hobbs. Botanical composition and nutritional quality of elk winter diets in the upper montane zone, Colorado. J. Wildl. Manage. This manuscript has been published as: Hobbs, N. T., D. L. Baker, J. E. Ellis, and D. M. Swift. 1980. Composition and quality of elk winter diets in Colorado. J. Wildl. Manage. 45(1):156-171. 16. Big Game Investigations Federal Aid Job Progress Reports. These reports have been printed in: Colorado Division of Wildlife. 1-403. Wildl. Res. Repts. July Part 1 and 2: Additional publications which were not scheduled but which were published or accepted for publication during the 1980-81 Segment are listed below: Anderson, A. E. 1981. Morphological and physiological characteristics. pp. 27-97. In O. C. Wallmo (ed.). Mule deer and black-tailed deer of North America. Univ. Nebraska Press, Lincoln. 605pp. Carpenter, L. H. 1981. Impacts of grazing intensity and specialized grazing systems on faunal composition and productivity - a response to a paper presented by W. Evans. In Impacts of grazing and specialized grazing systems on use and value of rangelands. Natl. Acad. Sci. Workshop III. El Paso, Texas. March 16-17 (publication in process). Carpenter, L. H., and o. C. Wallmo. 1981. Habitat evaluation and management. pp 399-421. In O. C. Wallmo (ed.). Mule deer and black-tailed deer of North America. Univ. Nebraska Press, Lincoln. 605pp. Ellis, J. E., D. M. Swift, N. T. Hobbs, R. G. Woodmansee, and D. L. Baker. 1980. Estimation of nitrogen transport by large animals in seasonal ecosystems. Bull Ecol. Soc. Amer. 61(2):113. (Abstract). Hobbs, N. T., and D. C. Bowden. Confidence intervals on preference indices. J. Wildl. Manage. (accepted for publication). Hobbs, N. T., D. L. Baker, D. M. Swift, and J. E. Ellis. 1980. Energy and nitrogen based estimates of ungulate carrying capacity. Bull. Ecol. Soc. Amer. 61(2):26 (Abstract). �25 Kautz~ M. A.~ W •• W. Mautz, and L. R. Carpenter. 1980. Heart rate as a predic tor of energy expenditure. J. Wildl. Manage. 45 (3):715-721. Kufeld, R. C.~ M. L. Stevens, and D. C. Bowden. 1981. Winter variation in nutrient and fiber content and in vitro digestibility of Gambel Oak (Quercus gambelii) and big sagebrush (Artemisia tridentata) from diversified sites in Colorado. J. Range Manage. 34(2):149-151. Pojar, T. M. 1981. A management perspective of population modeling. In C. W. Fowler, and T. D. Smith (eds.). Dynamics in large mammal populations. John Wiley and Sons, Inc. New York, N.Y. Pusateri, F. M., C. P. Hibler, and T. M. Pojar. Oral administration of diazepam and promazine hydrochloride to immobilize pronghorn. J. Wildl. Dis. (accepted for publication). Reed, D. F. 1980. Road-killed deer--variability and related parameters. Central Mtns. Plains Sect. TWS, Proc. 25:8 (Abstract). Reed, D. F. 1981. Conflicts with civilization. pp 509-535. In O. C. Wallmo (ed.). Mule and black-tailed deer of North America. Univ. Nebraska Press. Lincoln, Nebr. 605pp. Reed, D. F. 1981. Mule deer behavior at a highway underpass exit. Wildl. Manage. 45(2):542-543. J. Reed, D. F., and T. N. Woodard. 1981. Effectiveness of highway lighting in reducing deer-vehicle accidents. J. Wild1. Manage. 45(3):721-726 . .' \ '; / Prepared by I ~~Z~,_,_'~~,~l_~·_~~~~~~~~--~~~~~'-R. Bruce Gill Big Game Section Chief ��27 JOB PROGRESS REPORT State of Colorado ---------------------- Proje~t No. W-126-R-4 -------------- Work Plan No. 1 Big Game Investigations Job No. 8 Job Title: Multispecies Investigations - Big Game Publication and Editing and Library Services Period Covered: Personnel: July 1, 1980 through June 30, 1981 M. W. Hershcopf, N. W. McEwen, Dr. O. B. Cope, R. B. Gill, A. E. Anderson, D. L. Baker, R. M. Bartmann, G. D. Bear, T. D. I. Beck, Dr. L. H. Carpenter, D. J. Freddy, Dr. N. T. Hobbs, R. C. Kufeld, P. H. Neil, T. M. Pojar, D. F. Reed, W. H. Rutherford, T. V. Dailey, S. Wielgopolan. ABSTRACT During Segment 4 of W-126-R, one Division Special Report was published. Twenty-three books and theses were purchased for permanent reference by DOW researchers. Thirty-nine theses were obtained on inter-library loan for research use. Three literature searches were performed by library staff for research use. Approximately 690 references requested by Big Game Research personnel were located by library staff, and approximately 15,000 pages of published material were provided for Big Game Research personnel. ��29 BIG GAME PUBLICATION EDITING AND LIBRARY SERVICES , , R. B. Gill P. N. OBJECTIVE To provide a centralized support program for Big Game Research technical editing and l~brary services so that Big Game Research scientists can allocate additional time to the conduct of actual research. SEGMENT OBJECTIVES To provide coordinated, efficient, and economic editing and library services to all Colorado Big Game Research program (Federal Aid Project W-126-R). SUMMARY OF SERVICES Publications Submitted for Editing and DOW Publication Bartma~n, R. M., and S. F. Steinert. 1981. Distribution and movements of mule deer in the White River Drainage, Colorado. Colo. Div. Wildl. Spec. Rep. No. 51. l2pp. Publications Library Purchased with W-126-R Funds and Placed ,in Research Center Amason, A. N., and L. Baniuk. 1978. POPAN-2; a data maintenance analysis system for mark-recapture data. The Charles Babbage Research Centre, St. Pierre, Manitoba, Canada. 269pp. and Church, D. ,C. 1979. Digestive ~hysiology and nutrition of ruminants. Vol. 2 - Nutrition, 2nd ed. 0 & B Bcoks, Inc., Corvallis" Oregon. 452pp. Dean, R. E. 1973. Nutritional aspects of artifically feeding captive and wild deer. Ph.D. Dissertation, Oregon State Univ., Corvallis. 96pp'. (Authorized facsimile by Univ. Microfilms, IntI.) Goldstein, M., and L' F. Goldstein. 1978. How we know; an exploration of the scientific process. Plenum Press, New York. 357pp. Hutchinson, T. C., and M. Havas (eds.). 1980. Effects of acid precipitation on terrestrial ecosystem. Proceedings of NATO Conference on effects of acid precipitation on vegetation and soils, Toronto, May 21-27, 1978. (NATO Conference Series 1: Ecology; v.4) Plenum Press, N.Y. 654pp. �30 McCullough, D. R. 1979. The George Reserve deer herd; population ecology of a K-selected species. The University of Michigan Press, Ann Arbor. 27lpp. McCullough, Y. B. 1980. Niche separation of seven North American ungulates on the National Bison Range, Montana. Ph.D. Dissertation, University of Michigan, Ann Arbor. 226pp. (Authorized facsimile by University Microfilms IntI.) Martinka, C. J., and K. L. McArthur. 1980. Bears--their biology and management. Papers of the Fourth International Conf. on Bear Research and Management, Kalispell, Montana, February 1977. The Bear Biology Association (Bear Biology Association Conference Series No.3). 375pp. Monson, G., and L. Summer (eds.). 1980. The desert bighorn; its life history, ecology, and management. The University of Arizona Press, Tucson. 370pp. Moen, A. N. parts. 1981. The biology and management of wild ruminants. CornerBrook Press, Lansing, N.Y. Four Mould, E. D. 1980. Aspects of elk (Cervus canadensis nelsoni) nutrition and associated analytical procedures. Ph.D. Dissertation, Washington State Univ., Pullman. 72pp. Pigden, W. J., C. C. Balch, and M. Graham (eds.). 1980. Standardization of analytical methodology for feeds: proceedings of a workshop held in Ottawa, 12-14 March 1979. Interntl. Devel. Research Centre, Ottawa. l28pp. Pielou, C. E. 1979. Biogeography. New York. 35lpp. Wiley-Interscience Publication, Reimers, E, et al. (eds.). 1979. Proceedings of the 2nd reindeer/ caribou symposium, 17-21 September 1979, Rros, Norway. Direktoratet for Vilt og Ferskvannsfisk, Trondheim. Part A and B. 799pp. Rogers, L. L. 1977. Social relationships, movements, and population dynamics of black bears in northwestern Minnesota. Ph.D. Dissertation, Univ. Minn. 203pp. (Authorized facsimile by Univ. Microfilms IntI.) Roughgarden, J. 1979. Theory of population genetics and evolutionary ecology: an introduction. MacMillan Publishing Co., Inc., New York. ~34pp. Ruckebusch, Y., and P. Thivend (eds.). 1980. Digestive physiology and metabolism in ruminants. Avi Publishing Co., Inc., Westport, Conn. 854pp. �31 Schemnitz, S. D. (ed.). 1980. Wildlife management techniques manual. 4th ed., revd. The Wildlife Society, Washington, D.C. 686pp. Smith, M. H., and J. Joule (eds.). 1981. Mammalian The Univ. of Georgia Press, Athens. 380pp. Spiess, E. B. 780pp. 1977. Genes iripopulations. population genetics. John Wiley and Sons, N.Y. Stelfox, J. G. 1975. Range ecology of Rocky Mountain bighorn sheep in Canadian National Parks. Ph.D. Dissertation, Univ. Montana, Missoula. 234pp. (Authorized facsimile by Univ. Microfilms IntI.) 't Mannetje, L. (ed.). 1978. Measurement of grassland vegetation and animal production. Commonwealth Agricultural Bureaux, Farnham Royal, England (Commonwealth Agricultural Bureaux, Bull. No. 52). 260pp. Thompson, R. W. 1981. Ecology of Rocky Mountain goats introduced to the Eagles Nest Wilderness, Colorado and some factors of geographic variation in the lambing season of bighorn sheep. M.S. Thesis, Univ. Wyoming, Laramie. 359pp. Publications Obtained Free or at Low Cost In addition to books purchased with W-126-R funds, about 65 free reports and short publications from state or federal agencies, and from other sources, were located, ordered and obtained for use by big game research personnel. Theses Obtained on Interlibrary Loan for Use by Researchers Acosta-Gonzalez, R. A. 1976. Chemical composition of esophageal-fistula forage samples as influenced by drying method, salivary leaching and sample preparation. M.S. Thesis, Texas A & M Univ., College Station. Anderson, D. M. 1977. Standing crop, diets, travel and weight changes under short duration and continuous grazing. Ph.D. Dissertation, Texas A & M Univ., College Station. l80pp. Barnes, V. G., Jr. 1967. Activities of black bears in Yellowstone National Park. M.S. Thesis, Colorado State Univ., Ft. Collins. 116pp. Bauer, W. A. 1977. Forage preference of tame deer in the aspen type of northern Michigan. M.S. Thesis, Michigan Tech. Univ., Houghton. 85pp. �32 Belovsky, G. E. 1977. Optimal behavior of a generalist herbivore. Ph.D. Dissertation, Harvard Univ., Cambridge, Mass. Bray, O. E. 1967. A population study of black bears in Yellowstone National Park. M.S. Thesis, Colorado State Univ., Ft. Collins. 96pp. Chappel, R. W. 1978. Bioenergetics of Rocky Mountain bighorn sheep. M.S. thesis, University of Alberta, Edmonton, Alberta, Canada. Clark, A. C. 1977. Nutrient intake of white-tailed deer in winter and determination of the feeding capacity of.deer range. Ph.D. Dissertation, Pennsylvania State University, University Park. Croyle, R. C. 1969. Nutrient requirements of young white-tailed deer for growth and antler development. M.S. Thesis, Pennsylvania State University, University Park. l03pp. Eagle, T. C. 1979. Foods of black bears in the Great Smoky Mountains National Park. M.S. Thesis, University of Tennessee, Knoxville. l04pp. Eslinger, D. H. 1976. Form, function, and biological role in the locomotory apparatus of the genus Odocoileus in Alberta. M.S. Thesis, University of Calgary, Calgary, Alberta, Canada. l37pp. Eubanks, A. L. 1976. Movements and activities of the black bear in the Great Smoky Mountains National Park. M.S. Thesis, University of Tennessee, Knoxville. 83pp. Fairbanks, R. L. 1979. An evaluation of the pellet-group survey as a deer and elk census method in western Washington. M.S. Thesis, University of Washington, Seattle. Garshelis, D. L. 1978. Movement ecology and activity behavior of black bears in the Great Smoky Mountains National Park. M,S. Thesis, University of Tennessee, Knoxville. l17pp. Gephart, Glenn. 1979. Development and validation test of a mule deer habitat rule. M.S. Thesis, Utah State University, Logan. 97pp. Grimes, J. L. 1968. Nutritive evaluation of various deer foods. M.S. Thesis, Pennsylvania State University, University Park. 97pp. Hardy, D. M. 1974. Habitat requirements of the black bear in Dare County, North Carolina. M.S. Thesis, Virginia Polytechnic Institute and State University, Blacksburg. l2lpp. Harrington, F. A., Jr. 1978. Ecological segregation of ungulates in alpine and subalpine communities. Ph.D. Dissertation, Colorado State University, Ft. Collins. l52pp. �33 Hebert, Daryll M. 1973. Altitudinal migration as a factor in the nutrition of bighorn sheep. Ph.D. Dissertation, University of British Columbia, Vancouver, British Columbia, Canada. 355pp. Jense, G. K. 1968. Food habits and energy utilization of badgers. Thesis, South Dakota State University, Brookings. 39pp. M.S. Johnson, D. G. 1978. Den ecology of black bears (Q.~.)in the Great Smoky Mountains National Park. M.S. Thesis, University of Tennessee, Knoxville. l07pp. Jorgenson, C. 1979. Bear-livestock interactions, Targhee National M.S. Thesis, University of Montana, Missoula. l62pp. Forest. Meneely, S. 1978. Chemical composition and in vitro digestibility of browse three years after a wild fire. M.S. Thesis, New Mexico State University, Las Cruces. 99pp. Moen, A. N. 1966. Factors affecting the energy exchange and movements of white-tailed deer, western Minnesota. Ph.D. Dissertation, University of Minnesota, St. Paul. Novick, H. J., III. 1979. Home range and habitat preferences of black bears (U.a.) in the San Bernardino Mountains of southern California. M.S. Thesis, California State Polytechnic University, Pomona. 58pp. Overmire, T. G. 1963. The effects of grazing upon habitat utilization the dickcissel (Spiza americana) and Bell's vireo (Vireo bellii) northcentral Oklahoma. Ph.D. Dissertation, Oklahoma State University, Stillwater. 65pp. of in Owens, R. A. 1971. The effects of several agricultural regimes upon populations of native passerine birds of an Alberta fescue grassland. M.S. Thesis, University of Calgary, Alberta, Canada. Pallesfen, J. 1979. Nutritive value of mule deer and elk diets and forages in lodgepole pine habitats of Utah. M.S. Thesis, Utah State University, Logan. Razmi, K. 1978. Feeding, behavior of Sheep with respect to food-related cues in the environment. Ph.D. Dissertation, Utah State University, Logan. l25pp. Rideout, C. B. 1974. A radio telemetry study on the ecology and behavior of the mountain goat. Ph.D. Dissertation, University of Kansas, Lawrence. l46pp. Schoenfeldt, R. C. 1975. Evaluation of winter deer habitat in southwestern Pennsylvania using tame white-tailed deer. M.S. Thesis, Pennsylvania State University, University Park. 60pp. �34 Schommer, T. J. 1978. Seasonal in vitro digestion coefficients for energy and protein of central Washington elk diets. M.S. Thesis, Washington State University, Pullman. 57pp. Shaffer, M. L. 1978. Determining minimum viable population sizes: a case study of a grizzly bear (Ursus arctos ~.). Ph.D. Dissertation, Duke University, Durham, N.C. 206pp. Shult, M. J. 1968. Incidence of deer in the Nebraska concrete-lined Ainsworth irrigation canal. M.S. Thesis, Iowa State University, Ames. 82pp. Siperek, J. M. 1979. Physical characteristics and blood analysis of black bears (Ursus americanus) in the San Bernadino Mountains of Southern California. M.S. Thesis, California State Polytechnic University, Pomona. Sivinski, R. A. 1979. A multivariate analysis of summer habitat partitioning between elk (Cervus elaphus) and mule deer. M.S. Thesis, New Mexico State University, Las Cruces. 42pp. Stager, D. W. Mule deer response to successional changes in the pinyonjuniper vegetation type after wildfire. M.S. Thesis, University of Nevada, Reno. Stelter, L. H. 1979. Inventory, food habits, and trace elements levels of selected fauna of Colorado's oil shale region. Ph.D. Dissertation, Colorado State University, Ft. Collins. 100pp. Willms, W. 1979. The effects of fall burning on grazing on Agropyron spicatum (Prusch. Schribn, and Smith) and its selection by deer and cattle. Ph.D. Dissertation, University of Alberta, Edmonton, Alberta, Canada. List of Literature Searches Performed Research Center Library for Big Game Researchers by the Diseases of pronghorn Effects of logging and roads on elk Harassment of deer and elk Reference Document Location and Delivery The Research Center Library staff also located about 690 references on request for the Big Game Research section during this segment; about 30 of these were obtained through interlibrary loans. rf'·. C\ ;~"'" Prepared i by__ '·'_'~_\_'-_· '_~_(~_. ~~~_\_' R. Bruce Gill Big Game Research \ '_'_:)'-o-"",U:",--=-S_} __ Leader �July 1981 35 JOB PROGRESS State of Colorado Project No. ------- W-126-R-4 Work Plan No. Job Title: REPORT 1 ---------- Big Game Investigations Job No. Mult~~_ecie~Inv~~tigations ----------- 9 - Energy Development and Big Game Habitats Period Covered: Personnel: July 1, 1980 through June 30, 1981 W. H. Rutherford, H. M. Swope, R. B. Gill, P. D. Olson, W. D. Clark, A. F. Whitaker ABSTRACT This research program was discontinued because management and research personnel concurred that the activities requested of the Terrestrial Wildlife Research Section were more appropriate to the Northwest Region and Ecological Services Section. W. H. Rutherford was reassigned. ��37 ENERGY DEVELOPMENT AND BIG GAME HABITATS W. H. Rutherford P. N. OBJECTIVE Prepare a detailed PROGRAM NARRATIVE and SEGMENT NARRATIVE for Segment 5 of Project W-126-R describing in detail specific studies, their objectives, procedures, schedules, costs, etc. SEGMENT OBJECTIVE 1. Prepare a study plan. RESULTS During the segment, W. H. Rutherford met several times with representatives of Division of Wildlife's Ecological Services Section, Northwest Region, and Terrestrial Wildlife Research Section to define Research's role in energy development impacts and big game mitigations. Following those several discussions it was concluded that Ecological Services and the Northwest Region could resolve the anticipated wildlife problems asso~ ciated with energy development without additional research involvement. Therefore, W. H. Rutherford was reassigned to an evaluation of moose trans locations in Colorado. '7 Prepared by: . ~1!/:II )J,;;?l·t-h-~} W. H. RuthTrford / Wildlife Researcher ��July 1981 39 JOB PROGRESS Colorado ----------------------- State of Project REPORT No. W-126-R-4 ~rk?l~ Job Title: ---------- ~. 2 Period Covered: Personnel: 1 Job No. Deer Investigations Capacity Big Game Investigations o~~~nter - Nutritional Ranges Basis for Quantifying to .~3~p~p~o_r __t__ D_e_e_r_. ~ July 1, 1980 through June 30, 1981 Dr. L. Carpenter, S. Torbit, D. Freddy, D. Swift ABSTRACT Body composition of hand-reared mule deer was studied in vivo by use of tritiated water. Three treatment groups were established and energy intake was controlled at different levels for these groups. All treatment groups received submaintenance rations to insure catabolism of energy reserves. Changes in total body water over time, indicated significant protein depletion occurred before fat reserves were exhausted. In order to validate in vivo estimates, 4 animals were euthanized and the carcass chemically analyzed. Regression analysis revealed that the chemical and in vivo estimates were highly correlated. Significant protein catabolism of malnourished deer has important implications for the survival and productivity of overwintering animals. _ ��41 NUTRITIONAL BASIS FOR QUANTIFYING CAPACITY OF WINTER RANGES TO SUPPORT DEER Dr. Len Carpenter and Steve Torbit P. N. OBJECTIVE To determine if a system can be developed to estimate the number of deer winter ranges are capable of supporting. SEGMENT OBJECTIVES 1. Test the efficacy of a computer simulation model for estimating carrying capacity of winter ranges. 2. Determine body composition of mule deer on various nutritional levels. 3. Prepare manuscript on maintenance energy requirements of adult mule deer. 4. Prepare manuscript on activity patterns of mule deer at pasture. METHODS AND MATERIALS Most of the effort this segment was spent on objective 3. A change in job assignments and duties for the principal investigator resulted in moderate progress for the 3 other objectives. However, manuscript preparation is continuing and drafts should be completed during the next segment. A dissertation by Torbit fully covering objective 2 is in draft stages and will be completed soon. Methodology for work on objective 2 was previously presented by Carpenter and Torbit (1980). Certain additions and modifications follow. Immediately after the last body composition trial, 3 deer were euthanized with T-6l (Tetracaine hydrochloride, Taylor Pharmacal Co.), the 4th deer of the group died shortly after an in vivo trial. Immediately after death all animals were weighed, wrapped in-plastic bags and frozen. Animals were thawed individually and each was shaved completely with an Oster animal clipper equipped with a surgical blade. A random sample of hair from each deer was saved for determination of water, protein, and ash content. All deer were eviscerated, the gastro-intestinal tract was cleaned completely and contents weighed. Amount of ingesta was used to determine the ingestafree weight of the animal and triplicate samples were collected and later analyzed for percent water. The entire carcass, less hair and gastrointestinal tract contents, was ground 4 times with a large commercial carcass grinder. Three random samples of approximately 1 kg were taken from the minced animal. Each sample was placed in a tared, labelled plastic bag and frozen. Duplicate subsamples from each of the larger samples were dried for 24 hours in a 60 C vacuum oven and ground through a Wiley Mill with �42 dry ice. Nitrogen content was determined by macro-Kjeldahl procedures and fat was extracted by a methanol-choloroform mixture (Spence and Wolfe 1967). Ash was determined by combustion of the sample in a muffle furnace for 6 hours at 500 C. RESULTS AND DISCUSSION Mean weight changes for the 3 groups of deer demonstrated, that desired results were obtained with the ration and feeding levels chosen (Fig. 1). All 3 groups exhibited significant losses of body fat during the trial (Fig. 2). However, even at the conclusion of the trial, fat stores were not totally depleted. Total reserves of body protein exhibited similar responses (Fig. 3). There was a significant (p < 0.05) protein loss during the first 30 days of the trial for both high and low intake groups. A similar test was not made for the medium intake group due to the death of 1 animal in this group. These results indicated that proteinaceous energy reserves of mule deer are much more labile than previously hypothesized. This has important implications for overwinter survival and productivity of deer since protein catabolism could have deleterious effects upon an animal's health manifested through decreased disease resistance and reproductive success. Results of the comparison of estimates between the tritiated water technique and chemical analysis of the minced carcass indicated that tritiated water was an excellent tracer of total body water and thus body composition. The relationship between HTO estimated body water and chemically estimated body water was nearly 1:1 (Fig. 4). Relationships between the 2 estimates for total body fat and total body protein were equally as good (Figs. 5 and 6). Body fat and body protein are considered to be the most important indicators of an animal's nutritional health. Therefore, estimates of these 2 components were of greatest interest in this study where we are attempting to understand dynamics of tissue catabolism. Finally, regression analyses also indicated a strong relationship between the two methods in estimating total body ash (Fig. 7). This study was an attempt to validate the tritiated water technique of estimating body composition. Although sample size was small, evidence suggested that the tritium technique was highly accurate as an in vivo estimate. Future work should proceed by establishing a data base for interrelating the 4 basic components (fat, water, protein, and ash) of body composition for mule deer specifically. After these data are collected, more rigorous testing of the tritium technique for estimating body composition of mule deer should proceed. �43 LITERATURE CITED C~rpenter, L. H. and S. Torbit. 1980. Nutritional basis of quantifying capacity of winger ranges to support deer. Colo. Div. Wildl. Game Res. Rep. July, Part 1:83-96. Robbins, C. T., Moen, A. N. and Reid, J. T. 1974. Body composition of white-tailed deer. J. An. Sci. 38(4):871-876. Spence, M. W., and L. S. Wolfe. 1967. Gangliosides in developing rat brain. Isolation and composition of subcellar membranes enriched in gangliosides. Can. J. Biochem. 45:671-688. Prepared by _,{_e...... __ #_._U_I2_,.;;J::___ _ Dr. L. Carpenter Wildlife Research Leader Prepared by ~~~~~~~--~~~~ S. T9r it Gradvate Research Assistant �44 • P High Intake Medium Intake Low Intake E R C E -5 N T C H A -15 N C E ,,, ,,, ,,, , ,, I N B -25 o o e, ' .. ,.• "'.. ", Y .. W T ", -, ~ -35 1 Ie 2t 3a 4111 !5e ;, 71 81 QI "" til 12' DAYS Figure 1. Mean weight change (percent) of mule deer fed 3 different 1eve1~ of food during 120-day body composition trial, day l=Jan. 2, 1980. �45 8 7 -, •x C1 High Intake Medium Intake Low Intake <, <, <, F , A <,, T , G H T . <, , , \J E I - -_ -_ -.. ..._ 6 <, , , 1i.. •• , <, , .......... .......... ~ , <, , <, , 1 <, ~ K G -_ --- - ...... ..__,_ .• 2 ""fl DAYS Figure. 2. Mean total HTO estimated body fat (Kg) of adult mule deer fed 3 levels of experimental ration during l20-day body composition trial. �46 • x o 11 <, .•. P R o <, .•. II High Intake Medium Intake Low Intake <, .•. T ---_ .••.... _ E J N -- -- ---.c W E I G 8 H T 7 K G Ii " 182m3" 41 51 61 7a 81 ;" Isa 110 1211 DAYS Figure 3. Mean total HTO estimated body protein (Kg) of adult mule deer fed 3 levels of experimental ration during l20-day body composition trial. �47 sa 28 tc. 26 C 2. B 22 0 0 y 2e 18 \I A Hi T •• E R 2 R S.L '2 .e . C H E = 99.6 ;::::0.41 P<o.oos 8 6 M E S T •• 2 e e 2 • 6 B '2 12 KG BOJY '4 16 18 \.lATER 29 22 24 26 28 32 HTO EST Figure 4. Relationship between HTO estimated body water and chemically estimated body water for 4 mule deer at the conclusion of body composition trials. �48 ~ G B 0 0 y 4 F A T C H E 3 2 M R2 = 96.1 S.E. = 0.35 P<O.025 E S T 2 3 KG BODY FAT !5 6 HTO EST Figure 5. Relationship between HTO estimated body fat and chemically estimated carcass fat for 4 mule deer at the conclusion of body composition trials. �49 '" K C B 0 8 0 y 7 P R 6 0 T E I N 4 R2 = 99.3 s. E • = o. C 19 P<OoOOS H - .. •.. M 2 E S T " 2 3 6 KG BODY PROTEIN HTO EST Figure 6. Relationship between HTO estimated body protein and chemically estimated body protein for 4 mule deer at the conclusion of body composition trials. �50 .' 2 K C B 0 0 1.5 y A S H C H E M R2 = S.L e ~ E S 99.6 = 0.03 P<o.oos T B " 1.5 KG BODY ASH 2 HTO EST Figure 7. Relationship between HTO estimated body ash and chemically estimated body ash for 4 mule deer at the conclusion of body composition trials. �July 1981 51 JOB PROGRESS State of __ ~C~o~l~o~r~a~d~o~ REPORT _ Big Game Investigations Pro jec t No. ..:.;W:.....--=1=-=2:.:::6:.....-.!::R~-_.....:4 _ Work Plan No. Job Title: Period Snowmobile Covered: Personnel: Job No. 2 Harassment of Mule Deer on Cold Winter July 1, 1980 through D. Freddy, M. Fowler, National Laboratory. 2 Ranges June 30, 1981 D. Bowden, and G. White of Los Alamos ABSTRACT A 16-mm computer-generated film, approximately 20 minutes in length, was completed in cooperation with Los Alamos National Laboratory. Ten harassment bouts were selected to show varied responses of deer to human stimuli. A manuscript summarizing heart rates of mule deer in various activities at pasture is in progress and will be submitted to a physiological journal in Fall 1981. ��53 SNOWMOBILE HARASSMENT OF MULE DEER ON COLD WINTER RANGES David J. Freddy P. N. OBJECTIVE Evaluate whether snowmobile activity on winter ranges inhabited by deer decreases the ability of deer to survive winter by modifying activities of deer so as to significantly increase energy expended. SEGMENT OBJECTIVES 1. Prepare manuscript at pasture." entitled 2. Prepare rough draft of manuscript rate of mule deer at pasture." 3. Summarize and organize data regarding harassment for conceptualization into manuscript. 4. Prepare 16-mm movie depicting reactions in cooperation with computer facilities Laboratory. METHODS Methods are presented "Heart rates for activities entitled "Seasonal of mule deer changes in heart of wild mule deer of wild mule deer to harassment at Los Alamos National AND MATERIALS in detail by Freddy (1978, 1980). RESULTS AND DISCUSSION Objectives 1 and 2 will be combined into 1 manuscript which is currently in progress and it is anticipated this manuscript will be submitted to a physiological journal during Fall 1981. Further summarization of harassment data collected in 1979 and 1980 was delayed because of a transformation error found in all estimates of distances moved by deer during harassment bouts. All distance calculations were corrected and the data set should be in manuscript form by early 1982. A 16-mm computer-generated film, about 20 minutes in length, was completed in cooperation with Los Alamos National Laboratory. Ten harassment bouts conducted in 1979 and 1980 were selected to represent varied responses of mule deer to persons walking and snowmobiles. This movie will be used to Z) illustrate the usefulness of filming computer processed telemetry data and 2) illustrate to the .public the reactions of mule deer to harassment. �54 LITERATURE CITED Freddy, D. J. 1978. Snowmobile harassment of mule deer on cold winter ranges. Colo. Div. Wildl. Game Res. Rep. July, Part 2:137-144. 1980. Snowmobile harassment of mule deer on cold winter ranges. Colo. Div. Wildl. Game Res. Rep. July, Part 1:99-111 • • �July 1981 55 JOB FINAL REPORT State of Project Colorado No. W-126-R-4 Work Plan No. Big Game Investigations Job No. 2 Job Title: Deer Investigations - Estimating Basin Deer Population Period Covered: Personnel: 3 Parameters of Piceance Dynamics July I, 1980 through June 30, 1981 R. Bartmann and J. Ellenberger ABSTRACT 2 Post-season deer classifications on randomly located l-mi. quadrats in Piceance Basin were flown in December 1981 with the Northwest Region biologist to instruct him in precedures for conducting classifications and analyzing data. There were 37 adult males, 518 adult females and 326 fawns classified for a buck:doe ratio of 7.2 + 3.0:100 and a fawn: doe ratio of 63.2 ± 5.2:100 at the 90 percent confidence level. This job is now the responsibility of the Northwest Region. The quadrat deer census, also to be flown with the Northwest Region biologist, was cancelled for lack of snow. It is planned again for next winter. Division Special Report #51, Distribution and Movements of Mule Deer in the White River Drainage, Colorado authored by R.M. Bartmann and Steven F. Steinert, has been published and the abstract is in Appendix A. Work is continuing on analysis and publication of the deer census and winter mortality portions of the study which, with the final census flight, are included under Work Plan 2, Job 9. ��57 ESTIMATING PARAMETERS OF PICEANCE BASIN DEER POPULATION DYNAMICS Richard M. Bartmann P. N. OBJECTIVE To develop and test a method for estimating density of over-winter mule deer populations in the Piceance Basin. METHODS AND MATERIALS Methods and materials have been previously described (Bartmann 1974). LITERATURE CITED Bartmann, R. M. structure. 1974. Piceance deer study-population density and Colo. Div. Wildl. Game Res. Rep. July. Part 2:363-370. Prepared bY//~~·-~ !Richard M. Bartmann Wildlife Researcher C ~ ��59 APPENDIX A �60 DISTRIBUTION AND MOVEMENTS OF MULE DEER IN THE WHITE RIVER DRAINAGE, COLORADO ABSTRACT Distribution of mule deer (Odocoileus hemionus) in the Piceance Basin in the White River drainage during the winters of 1971-72, 1972-73, and 1973-74 was defined by assessing deer track densities from the air after fresh snowfalls. Upper limits of deer distribution were near 2,440 m (8,000 ft) in December but dropped to around 2,135 m (7,000 ft) in February during two severe winters. Snow depth and its physical characteristics seemed major influences on deer distribution. Major concentration areas were in the eastern half and northwest corner of the basin. Information on deer movements was gained by marking 1,923 deer in the White River drainage during five winters, 1972-1976. This sample provided 281 recoveries and 1,053 sightings. Two deer subpopulations were identified based on use of different summer ranges, but they could not be readily distinguished on winter range. Returns from outside the White River area were common except in winter, indicating permanent egress is negligible. Considerable movement of deer on winter range was evident, and partly resulted from effects of weather and late migration. Only 15 percent of returns during fall hunting seasons originated outside the White River area. Banding data indicate current Game Management Units define no discrete subpopulations and that deer wintering in the White River drainage should be managed as one population. Recommendations concerning population boundaries and data collection systems to accomplish this are presented. �July 1981 61 J.oB:FINAL REPORT State of Colorado ---------------------- Project No. W-126-R-4 2 Work Plan No. Big Game Investigations .Job No. 4 Job Title: Deer Investigations - Forage Preferences of Piceance Basin Deer Period Covered: Personnel: July 1, 1980 through June 30, 1981 R. Bartmann, L. Carpenter, A. Alldredge, M. Stevens, J. Ritchie, C. We inland , D. Bowden ABSTRACT Two manuscripts, Evaluation of Winter Forage Choices by Tame Mule Deer and Effects of Foraging Experience on Food Selectivity of Tame Mule Deer were accepted for publication by The Journal of Wildlife Management. Winter food habits and forage quality analysis work will be further analyzed and published under work Plan 2, Job 9. ��63 FORAGE PREFERENCES OF PICEANCE BASIN DEER Richard M. Bartmann P. N. OBJECTIVE To identify major forage species in the winter-long diets of mule deer on the Piceance pinyon-juniper winter range and to estimate the nutritional characteristics of the diet. Prepared ��65 July 1981 JOB FINAL REPORT State of Project Colorado ---------------------- 2 Work Plan No. Job Title: Period Job No. Deer Investigations Covered: Personnel: Big Game Investigations W-126-R-4 No. 5NE - Experimental Deer Inventory NE Region July 1, 1980 through June 30., 1981 P. Abbot, C. Albright, A. Anderson, D. Bowden, H. Geduldig, P. Konieczny, B. Parmenter, B. Pendrey, D. Schrupp, C. Smith, J. Stiver. ABSTRACT Five estimates of deer and elk densities (1978-81) are presented based on biannual counts and removals of pellet groups from 10-m2, permanent, circular plots arranged in a stratified, proportionally allocated, random, multistage sampling scheme on about 309 mile2 (800 kro2) of deer winter range using the square mile as the sample unit. Mean (95% confidence .intervals) deer densities thus derived ranged from 18.6 (13.6-23.6) deer during the "summer" of 1980 to 31.3 (17.4-45.2) deer/mile2 during the winter of 1978-79. Mean deer densities differed significantly (P < 0.05) only between the "summer" of 1980 and the "winter" of 1980-81. Mean (95% confidence interval) elk densities were from 2.4 (1.1-3.8) elk/mile2 during the "winter" 1980-81 to 5.4 (3.0-7.9) elk/mile2 during the"winter" of 1979-80. This difference was significant (P < 0.05). Sample sizes (number of sample units) were adequate to be within 20 to 25 percent of the true mean deer densities at a = .20 or a = .10 in 3 of 5 estimates presented. Current sample sizes were grossly inadequate to reliably sample elk densities. Number and size of openings within coniferous forest and mean elevation, slope azimuth, and slope gradient are described statistically for .each.of the 30 sample units. Recommendations are made to continue the study, with NE regional personnel participation, until the spring of 1983. ��67 EXPERIMENTAL DEER INVENTORY NORTHEAST REGION Allen E. Anderson P. N. OBJECTIVE To design appropriate sampling and reliably estimate deer numbers and based on a preliminary sampling of of the four administrative regions analytical procedures necessary to buck:doe:fawn ratios1 or winter range a problem management unit within each of the state. METHODS AND MATERIALS The sampling plan is stratified, random, multistage (Fig. 1) and involves counting and removing deer and elk fecal pellet groups on 800-1200 permanent sample plots (10-m2, 107.64 ft2) sample plots on G~1U-20 deer winter range during the spring and fall each year (Anderson and Bowden 1979, Anderson 1980). Definitions and procedures used in counting pellet groups are in Appendix A. Data were recorded on prepared forms (Appendix B). I report on the deer and elk fecal pellet group deposition rates during the "summer" of 1980 and the "winter" of 1980-81 and estimates of deer and elk densities calculated from those deposition rates. In addition, I present means and variances of deer and elk deposition rates by strata, 1978-80 and sample size requirements for estimating deer and elk densities on G.M.U. 20 deer winter range. Four site factors were measured within each of 30 square mile (2.59 km2) sample units on 1:24000, 40 ft (12.2 m) contour interval USGS Topographic Quadrangles. These include numbers and size (acres) of openings in coniferous cover using computer techniques2, and elevation, slope azimuth and slope gradient from a transparent, overlay plastic grid of 80 sample points and also from both ends of each of the 4, 10-plot, 660 ft (201.2 m), randomly located pellet group transects using the technique of Lee (1963). RESULTS AND DISCUSSION Deer and elk pellet group deposition rates and mean densities calculated from those rates are listed by strata for the "summer" of 1980 and the "winter" of 1980-81 (Table 1). Note that 95% confidence intervals overlapped between "summer" and "winter" densities for both deer and elk. A summary of similar estimates of deer and elk densities for 1978-81 shows that 95% confidence limits overlapped between the two "summer" and three lCompleted and final results reported 2Work by Carol Smith and Don Schrupp, in: CDW. Anderson (1980). �68 N I t--i MILE 4 •• ------,-I ~ ~ Figure 1. BERTHOUD Thirty proportionally allocated and randomly selected square mile (2.59 km2) sample units among 11 strata sampling about 309 mile2 (800 km2) of G.M.U. 20 winter range whose upper limit is approximated by the 8,500 ft (2,591 m) contour line. During the spring and fall, deer and elk pellet groups are counted and removed from 40, 107.64 ft2 (10 m2), circular, permanent plots randomly located in 4, la-plot segments within each sample unit. �69 "winter" (Table 2). A series of "t" test comparisons of differences between mean densities, however showed that these differences were significant (P < 0.05) only for deer during the "summer" of 1980 and the "winter" of 1980-81 and elk during the "winters" of 1979-80 and 1980-81. The similarity between seasonal mean deer densities is surprising since I know of no other mule deer winter range where this has been reported. Severe weather during the winter of 1979-80 may have lowered the average elevational range of elk occupancy of deer winter range thus accounting for the significant (P < 0.05) difference between mean elk densities during that winter and the exceptionally mild winter of 1980-81. Means and variances of pellet group deposition rates, 1978-80, are listed for deer and elk in Tables 3 and 4, respectively. With the data presented in Table 1 it is possible to trace the effects of adding, during the autumn of 1979 and the spring of 1980, 2 sample units to stratum 5, 1 sample unit to stratum 6, and 1 sample unit to stratum 9. What results is a somewhat inconsistent reduction in variance relative to the mean, particularly during the "winter" period. It is obvious that strata 7 and 8 have had consistently large variances. The overall variance of mean deer density might be decreased by increasing number of sample units in those strata. Adequacy of sample size (number of sample units) is estimated for deer and elk (Table 5). Current sample sizes are grossly inadequate for estimating mean densities of elk. Mean densities of deer were estimated within 25% of the true mean at a = .20 during the winter of 1978-79 (Anderson and Bowden 1979:197) and within 20% of the true mean at a = .20 or 25% of the true mean at a = .10 during the "summer" of 1980 and the "winter" of 1980-81. Thus, the present level of sampling should provide estimates of deer density sufficiently reliable for management purposes. Current limitations of time and budget preclude, however, a level of sampling which would substantially increase the reliability of the mean deer density estimate (Table 5). Since the 30 sample units have been operational only since the "summer" of 1980, I recommend that this experiment be carried out until the spring of 1983 to further substantiate its reliability under wider fluctuations in environmental conditions and deer population levels. In a preliminary effort to provide an interpretive ecological base for assessing distribution and abundance of deer and elk on Game Management Unit 20, we presented a statistical description of the total deer and elk pellet groups counted and removed during sample plot establishment during 1978. These descriptions are presented in relation to 28 habitats and also in relation to each sample unit (Anderson and Bowden 1979:105, 127, 134-187). An additional description of the habitats within each of 30 sample units is given for numbers and size of openings in coniferous cover (Appendix C), mean elevation (Appendix D), mean slope azimuth (Appendix E), and mean slope gradient (Appendix F). For the 3 latter site factors, I also present similar information on each randomly located, 10-plot, pellet group transect. These comparisons suggest that pellet group transects are reasonably representative of the mean elevation, slope azimuth, and slope gradient of most sample units. �Table 1. Estimated pellet group deposition rates and densities of deer and elk during the "summer" 1980 and the "winter" 1980-81 on G.M.U. 20 deer winter range based on counts of pellet groups on 40 permanent, randomly selected 107.64 ft2 (10 m2) circular plots within each of 30 stratified, proportionally allocated and randomly selected square mile (2.59 km2) sample units. Summer, 1980a Species Deer No. Square Miles Strata 26 25 34 27 29 30 30 28 32 22 26 309 1 2 3 4 5 6 7 8 9 10 11 No. Sample Hiles 2 2 3 2 4 4 3 2 4 2 2 30 }{ P.G./10 Plots/day .00610 .00600 .00470 .00433 ' .00582 .01119 .01012 .01220 .02195 .00603 .01210 Winter, 1980-81b s200-5) 0 0 1.01120 1.02400 2.21840 3.03600 5.40230 0.00784 25.72820 3.14720 0.81225 x: P.G./10 Plots/day .00880 .00210 .01800 .00770 .01600 .01760 .01340 .01740 .02670 .01960 .01390 s200-5) 1.36900 0 22.50000 1.36900 25.92100 28.22400 21.60900 24.96400 0.25600 1.02400 6.40000 Deer Per Square Mile (2.59 km2 c L Mean SE 95% Conf. Interval Elk 1 2 3 4 5 6 7 8 9 10 11 26' 25 34 27 29 30 30 28 32 22 26 309 2 2 3 2 4 4 3 2 4 2 2 30 18.60 2.15 13.56, 23.64 .00203 .00200 .00525 0 .00060 0 .00068 .00100 0 0 0 29.85 4.09 20.57, 39.12 0.82369 0 3.61200 0 0.14400 0 0.13.689 0.19881 0 0 0 .00155 .00104 .00414 .00206 .00157 .00078 .00068 .00052 .00026 0 0 .47960 .21610 .72900 .00004 .98596 .02704 .13924 .05476 .02704 0 0 Elk Per Sguare Mile (2.59 km2)d Mean SE 95% Conf. Interval 2.45 .87 -.24, 5.15 2.42 .57 1.07,3.76 aCounting periods: ~fuy 16, 1980 - June 19, 1980 to Sept. 16, 1980 - Oct. 14, 1980, Mean ± SD 122.8 ± 7.0 days of pellet group deposition per 10-plot segment. bcounting periods: Sept. 16, 1980 - Oct. 14, 1980 to May 18, 1981 - June 10, 1981, Mean ± SD 241.8 ± 2.1 days of pellet group deposition per 10-plot segment. c ' Based on a mean defecation rate of 13.0 groups per deer per day as generalized from information in Neff (1968). dBased on a mean defecation rate of 12.0 groups per elk per day as generalized from information in Neff (1968). ....., 0 �71 Table 2. Estimates of deer and elk densities on about 309 square miles (800 km2) of G.M.D. 20 deer winter range, 1978-81 based on counts of pellet groups. Animals per square mile Animal Season and Year. Mean Deer Winter Summer Winter Summer Winter 1978-79 1979 1979-80 1980 1980-81 31.3 22.4 32.8 18.6* 29.8* Elk Winter Summer Winter Summer Winter 1978-79 1979 1979-80 1980 1980-81 4.1 2.6 5.4 2.4 2.4 aAsterisk bCalculated indicates means whose difference value was less than 0.0. SE (2.59 km2)a 95% Conf. Interval 5.0 4.1 5.8 ·2.2 4.1 17.4b 0.0 15.3 13.6 20.6 1.2 0.5 1.1 0.9 0.6 .30 .12 3.00 O.Ob was significant 1.07 45.2 47.6 50.2 23.6 39.1 7.0 5.0 7.9* 5.2 3.8* at P < 0.05. �72 Table 4. Means and variances of elk pellet group deposition rates on 800-1200 sample plots randomly located on G.M.U. 20 deer winter range, 1978-80. b n Strata Mean pellet groups per 10 plots per day Winter, 1 2 3 4 5 6 7 8 9 10 11 26 25 34 27 29 30 30 28 32 22 26 309 2 2 3 .0050955 .0020905 .0028236 .0027105 .0006155 .0013733 .0018583 .0055685 2 2 3 3 2 3 2 2 26 1978-79 5.19282 0.15824 0.18597 0.00530 0.07576 0.56581 0.29329 4.05090 o o .0006250 .07812 o o Summer, 1 2 3 4 5 6 7 8 9 10 11 26 25 34 27 29 30 30 28 32 2 2 2 1 1 2 2 2 22 1 2 26 309 3 .000725 .001690 .009840 1 3 4 5 6 7 8 9 10 11 26 25 34 27 29 30 30 28 32 22 26 309 aNumber .105125 .571220 .784080 o o o o o o o o o o o o o o o o 20 Winter, 2 1979 2 2 2 1 3 3 .004445 .002830 .000550 .248645 .064980 .060500 o o .005005 o 2 2 .892003 o .002705 .005320 .001650 .009670 3 1 2 1979-80 o 1.46341 3.51122 .81675 o o 23 of square miles (2.59 km2, 258.99 ha). bThe number of proportionally allocated and randomly selected square mile sample units differ because snow precluded counts on 6 sample units and 3 sample units were added, 1979-80. �73 Table 5. Number of square mile (2.59 km2) sample units required to be within specified percentages of the true mean density with confidence levels (I-a) for deer and elk on G.M.U. 20 deer winter range. (l-a) Percent .95 .90 5 10037 2510 1116 678 402 3765 942 419 236 151 1310 328 146 82 53 2626 657 292 165 106 1483 371 165 93 60 739 185 83 47 30 Deer - "winter"a Oct. 5, 1979 - June 19, 1980 Based on 23 sample units 780 200 89 50 32 530 133 59 34 22 304 76 34 19 13 Deer - "summer" May 16 - Oct. 14, 1980 Based on 30 sample units 1053 264 117 66 43 707 177 79 45 29 411 103 46 26 17 Deer - "winter"b Sept. 16, 1980 - June 10, 1981 Based on 30 sample units 7223 1806 803 452 289 3200 800 366 200 128 1290 323 144 81 52 3469 868 386 217 139 1905 477 212 120 77 426 232 103 58 38 17215 4304 1913 1076 699 9600 2400 1067 600 384 4729 1183 526 296 190 Elk - "summer" May 16 - Oct. 14, 1980 Based on 30 sample units 3016 754 336 189 121 1572 393 175 99 63 Elk - "winter"b Sept. 16, 1980 - June 10, 1981 Based on 30 sample units 10 15 20 25 5 10 15 20 25 5 10 15 20 25 5 10 15 20 25 5 10 15 20 25 5 10 15 20 25 5 10 15 20 25 5 10 15 20 25 5074 1269. 564 318 203 .80 Species and dates pellet groups were counted and removed on the first and last transect completed Deer - "summer"a April 24 - Nov. 19, 1979 Based on 20 sample units b (dvf , (d f , i 9.97) = = 12.87) Elk - "summer"a April 24 - Nov. 19, 1979 Based on 2.0 sample units Elk - "winter"a Oct. 4, 1979 - June 19, 1980 Based on 23 sample units b (d f . = i aCorrected values for those presented in Anderson adjusted degrees of freedom (d.f.). bMean-adjusted degrees of freedom (d.f.) (d.L = 3.2) 4.16) (1980:140) using variance- ��75 APPENDICES �76 APPENDIX A COUNTING DEER PELLET GROUPS Definitions and Procedures Deer Pellet Group - Five or more deer pellets of the same general size, shape, hardness and color judged to have been continuously voided at the place where observed. Pellets redistributed by water of other agents to within the plot are not recorded. Pellets strewn across the plot are counted as a group if about one-half of the total linear distance or its midpoint falls within the plot as measured with a steel pocket tape. Groups occurring on the plot periphery are counted if about one-half of their total area falls within the plot. "New" Deer Pellet Group - Pellets are typically deposited during the previous year. shiny, often soft and "Old" Deer Pellet Group - Pellets are typically not shiny, generally hard and sometimes wholly or partially concealed by litter. Search Procedures - The area of search is defined by the light metal chain, 1.784 meters in length revolving about the metal rod fitted over the angle-iron stake marking the center of each plot. The plot size is 0.001 hectare or 10 square meters or 107.64 square feet. On steep slopes the chain is held horizontal. When necessary, the plot boundary is defined by dropping a pebble from the 1.784 meter mark on the chain. Each plot is searched twice (clockwise and counterclockwise) with the two observers changing positions with the change in search direction. �77 APPENDIX B FORM FOR RECORDING PELLET GROUP COUNTS G.M.U. 20 PELLET GROUP COUNTS W-126-R WP2-J5 SAMPLE UNIT STRATUM ------ _ TRANSECT (S) _ ~-------- OBSERVERS -------------- DATE TIME BEGAN'----TRAN. Plot DEER New TOTAL--- TIME FINISHED ---ELK Old New Old TRAN. Plot 1 11 2 12 3 13 4 14 5 15 6 16 7 17 8 18 9 19 10 20 Total Total Comments, Weather, etc. HR -- MIN---- DEER New ELK Old New Old �a APPENDIX C. Legal description, actual area, and numbers and size of openings in coniferous forest characteristic of 30 randomly selected stratified "square mile" sample units from a universe of 309 square miles of G.M.U. 20 deer winter range. Sample unit Legal description Strata No. U.S.G.S. guadrangle(s) 1 1 2 2 3 3 3 4 4 5 5 5 5 6 6 6 6 7 7 7 8 8 9 9 9 9 10 10 11 11 18 22 4 11 4 8 13 5 22 6 13 21 24 4 12 17 28 8 19 29 11 28 7 14 20 22 13 22 Glenhaven Glenhaven Glenhaven Panorama Peak Panorama Peak Panorama Peak, Raymond Raymond Buckhorn Mtn.,Crystal Mtn. Buckhorn Mtn. Drake Drake Drake Glenhaven Drake Drake Glenhaven, Drake Pinewood Lake Pinewood Lake Panorama Peak Pinewood Lake Lyons Raymond Horsetooth Horsetooth Horsetooth Horsetooth Drake Drake Masonville Pinewood Lake, Carter Lake Res. 8 14 Total acres of sample Size of oEenings X (acres) b S TwpN 26 35 15 27 29 1 8 17 36 16 21 34 31 9 13 20 33 8 19 29 9 31 12 15 21 26 30 7 22 6 6 5 5 4 3 3 7 7 6 6 6 6 5 15 6 5 4 4 4 3 3 7 7 7 7 6 5 5 72 72 72 72 72 72 72 71 71 71 71 71 71 71 71 71 71 71 71 71 71 71 70 70 70 70 70 70 70 574.57 625.55 623.65 654.63 650.85 635.29 650.18 608.75 655.90 669.23 623.64 625.67 648.48 639.99 549.14 575.56 635.22 645.44 593.04 654.29 627.14 638.82 630.30 680.19 651.81 653.61 0 - - - - - - No openings - - - - - 2.37 8 4.05 1.07 7.38 3.12 3.70 13.11 0.39 4 5 16.60 2.07 48.41 18.7 22.13 34.9 62.43 3 1.47 8 3.40 3.68 0.75 10.88 8.60 27.10 4 12.35 1.58 2 64.82 86.38 3.74 125.90 2 115.95 159.37 3.26 228.64 4 3.17 1.77 1.26 4.85 2 22.36 2.01 42.70 28.77 20.90 19.52 2.51 52.69 7 91. 18 2.02 3 149.10 263.30 9 16.14 24.61 0.60 65.39 5 51.18 104.94 1.51 238.82 4 5.61 5.43 1.24 13.52 23.12 8 30.80 1.94 97.00 28.01 57.59 1.60 157.86 7 5.19 3.66 0.22 9.03 4 141.67 85.97 1.98 249.54 3 8 8.52 11. 72 1.20 36.51 4 5.52 4.81 1.35 12.37 - - - No coniferous forestC 124.69 181.20 332.99 3 3.39 1 575.73 --d 8 27.93 68.20 0.70 196.6 8 9.37 8.10 1.69 27.39 6 32.65 61.30 0.50 154.60 e 1 611.07 -- 5 4 70 519.56 6 Range W un t tb 625.65 537.35 630.0B- N SD Min Max --- 40.46 87.36 2.42 218.70 aU.S• Geological Survey (1967:1) criteria for "woodland" stipulate that the growth must be "at least 6 ft tall and dense enough to afford cover for troops." The minimum crown density requirement is from 20 to 35 percent vegetative cover; 20 percent is where the average open-space distance between the crowns is equal to the average crown diameter" and 35 percent exists where the "average open space between the crowns is equal to one-half the average crown diameter." Woodland areas of 40,000 square ft ormore (about 1 acre) are mapped and clearings or openings within woodland also have the 40,000 square ft or more requirement. bDerived by computer techniques from the 7.5 minute U.S.G.S. Topographic Quadrangles cIncludes 55.95 acres of Horsetooth Reservoir. dTwo patches totaling 63.09 acres of coniferous forest. eOne patch of 42.54 acres of coniferous forest. listed. ......, 00 �APPENDIX D. Mean elevations (ft)a of 30 sample units of 1 square mile (258.99 ha) each sampled with an 80 point grid and at each end of 4,660 ft (201.2 m) randomly located pellet group transects. Sample unit Strata no. 1-18 1-22 2-4 2-11 3- 4 3-8 3-l3 4-5 4-22 5- 6 5-13 5-21 5-24 6-4 6-12 6-17 6-28 7-8 7-19 7-29 8-11 8-28 9- 7 9-14 9-20 9-22 10-13 10-22 11-8 11-14 Total square mile - 80 sample points Mean 8049 8026 7693 7934 8443 7641 8488 7246 6694 8424 8428 7030 7113 7196 7173 8313 7956 7916 7300 6841 6859 8008 5545 6548 6548 6558 6354 6307 5588 6095 aRead to the nearest SD 268.2 285.0 238.6 198.2 203.8 147.6 233.3 308.7 l33.0 322.7 316.2 215.1 433.6 462.7 320.1 242.2 228.9 267.6 331.4 235.5 307.1 336.5 122.6 265.2 261.2 413.6 193.0 393.0 155.8 162.2 40 ft contour interval Min 7520 7400 7240 7480 8140 7270 8070 6740 6420 7920 7900 6360 6480 6410 6520 7660 7520 7300 6770 6510 6410 7020 5400 6040 6020 5800 6020 5680 5350 5810 (Lee 1963). 4 Transects Max 8600 . 8580 8240 8340 8890 8000 9020 7840 6990 9200 9180 7380 8120 8020 7720 8590 8360 8410 8030 7440 7640 8390 5930 6960 6960 7130 6780 7080 6020 6380 To convert - 8 sample points Mean SD Min Max 8138 7910 7780 7848 8378 7588 8615 7255 6604 8394 8315 6793 7146 7188 7202 8328 7878 7895 7302 6805 6929 8020 5501 6566 6152 6665 6405 6561 5632 6032 236.5 325.8 164.3 226.8 186.3 205.1 233.8 392.8 158.2 307.2 228.5 227.8 438.5 527.0 423.4 215.1 185.9 338.0 269.3 113.0 283.2 307.6 58.7 306.1 156.4 234.9 162.4 372.8 206.5 176.9 7760 7420 7560 7600 8110 7300 8340 6820 6420 7960 8000 6440 6650 6500 6520 8080 7600 7360 6980 6580 6630 7440 5420 6180 5960 6280 6120 5980 5410 5800 8440 8380 8000 8200 8620 7860 9040 7780 6840 8800 8600 7000 7720 7960 7640 8570 8160 8320 7800 6920 7440 8320 5600 6920 6400 6880 6620 7100 6000 6760 to meters divide by 3.2808. -...J \0 �Mean slope azimuth (degrees)a of 30 sample units of 1 square mile (258.99 ha) each sampled with an 80 point grid and at each end of 4,660 ft (201.2 m) randomly located pellet group transects. APPENDIX E. Sample unit strata & no. 1-18 1-22 2-4 2-11 3-4 3-8 3-l3 4-5 4-22 5-6 5-l3 5-21 5-24 6- 4 6-12 6-17 6-28 7- 8 7-19 7-29 8-11 8-28 9- 7 9-14 9-20 9-22 10-13 10-22 11 - 8 11-14 a Read to nearest 4 Transects Total square mile - 80 sample points Mean SD l32.8 193.7 155.2 142.3 100.5 182.9 104.0 141.7 142.7 112.7 114.9 208.0 121.4 198.3 158.6 205.3 127.5 154.8 l34.6 159.0 146.7 175.1 125.6 216.0 115.2 208.2 162.3 134.6 148.8 104.0 81.7 l31.4 66.2 108.4 69.1 98.5 61.7 114.2 106.4 72.4 59.6 91.5 ll8.5 114.8 89.5 114.7 105.8 124.8 88.7 93.1 69.9 102.0 85.3 77 .1 83.8 90.4 95.2 123.3 80.6 69.4 5 degrees Min Max Mean SD 0 0 5 0 0 0 15 0 0 10 20 15 0 10 0 0 0 0 0 0 0 5 15 10 5 0 5 0 0 0 280 355 340 355 345 355 340 355 355 355 260 345 355 340 355 355 355 355 350 355 340 355 335 345 355 335 350 355 285 345 108.1 196.9 191.9 159.4 129.4 205.6 161.2 172.5 58.8 108.8 90.6 153.1 98.1 175.6 133.8 ll2.5 92.5 141.9 193.1 113.8 159.4 153.1 90.6 246.2 98.1 223.1 86.2 210.0 143.1 110.6 63.0 127.5 92.0 135.3 47.6 99.2 71.6 132.9 42.5 54.8 44.8 67.6 122.1 111.1 77 .1 110.5 109.4 139.2 116.2 49.8 105.7 98.3 30.6 72.0 105.8 35.6 73.1 99.6 79.3 99.6 (Lee 1963); 360 degrees read as 0 degrees. - 8 sample points Min Max 40 20 • 30 25 80 70 85 30 10 35 30 35 5 45 20 25 10 0 35 45 20 15 40 180 5 165 10 110 60 20 210 340 320 340 215 320 300 335 155 185 165 240 330 310 230 270 305 355 350 175 320 325 150 355 340 280 215 355 245 320 00 0 �81 Table 3. Means and variances of deer pellet group deposition rates on 800-1200 sample plots randomly located on G.M.U. 20 deer winter range, 1978-80. Strata n b Mean pellet groups per 10 plots per day Winter, 1978-79 1 2 3 4 5 6 7 8 9 10 11 26 25 34 27 29 30 30 28 32 22 26 309 2 2 3 2 2 3 3 2 3 2 2 .0077305 .0119177 .0108380 .0029583 .0191740 .0228510 .0127344 .0226472 .0266496 .0195380 .0143766 6.090106 4.923084 3.317595 0.280987 73.528455 27.260672 15.511025 11.572833 26.812108 13.883112 0.447872 26 Summer, 1979 9 26 25 34 27 29 30 30 28 32 10 22 11 26 309 1 2 3 4 5 6 7 8 2 2 2 1 1 2 2 2 3 1 2 .001575 .005640 .004050 .005810 .018460 .017945 .017305 .025800 .014426 .005490 .004045 .003125 .089780 .089780 o o 8.307610 8.281850 83.558700 .263243 o ..001050 20 Winter, 1979-80 1 2 3 4 5 6 7 8 9 26 25 34 27 29· 30 30 28 32 10 22 11 26 309 2 2 2 1 3 3 2 2 3 1 2 .004445 .014190 .003285 .002290 .014767 .023337 .025755 .021960 .026093 .030390 .016010 .990215 7.663220 .244205 o 38.02190 13.86210 81.56760 60.97030 7.18476 o 2.72322 23 aNumber of square miles (2.59 km2, 258.99 ha). bThe number of proportionally allocated and randomly selected square mile sample units differ because snow precluded counts on 6 sample units and 3 sample units were added, 1979-80. �APPENDIX F. Mean slope gradient (percent)a of 30 sample units of 1 square mile (258.99 ha) each sampled with an 80 point grid and at each end of 4,660 ft (201.2 m) randomly located pellet group transects. Sample unit strata & no. 1-18 1-22 2-4 2-11 3-4 3-8 3-13 4-5 4-22 5- 6 5-13 5-21 5-24 6-4 6-12 6-17 6-28 7- 8 7-19 7-29 8-11 8-28 9- 7 9-14 9-20 9-22 10-13 10-22 11- 8 11-14 a Read to nearest 4 Transects Total sguare mile - 80 sample points Mean SD 46.0 39.1 36.7 30.9 25.4 29.8 23.3 43.6 30.6 31.0 23.8 36.8 45.7 57.2 45.9 32.6 30.7 45.6 50.5 37.9 34.4 38.3 20.4 35.4 28.9 45.1 37.0 49.8 23.3 21.8 l3.7 12.9 10.9 11.9 14.1 10.4 9.9 15.2 14.6 15.0 10.3 18.4 16.0 17.3 13.2 17.4 10.8 18.0 18.9 17.9 14.5 21.8 12.1 15.5 15.2 18.8 12.1 16.9 11.5 9.9 10 percent (Lee 1963). Min 20 10 10 10 0 10 0 10 10 10 0 10 10 10 20 0 10 10 10 0 0 0 0 0 10 20 10 20 0 10 Max Mean SD 80 70 60 60 70 50 40 90 70 80 50 80 80 80 70 70 60 80 100 70 60 80 50 80 70 100 70 90 50 40 37.5 36.2 32.5 30.0 27.5 31.2 28.8 38.8 27.5 40.0 23.8 43.8 45.0 58.8 46.3 31.2 36.2 38.7 51.2 31.2 41.3 32.5 13.8 32.5 32.5 42.5 31.2 37.5 30.0 21.2 16.7 13.0 10.4 12.0 22.5 8.4 9.9 11.3 14.9 12.0 15.1 13.0 15.1 17.3 18.5 12.5 15.1 18.1 18.8 11.3 14.6 21.9 5.2 10.4 10.4 15.8 12.5 12.8 12.0 11.3 - 8 sample points Min Max 20 20 20 20 10 20 20 20 10 30 0 30 20 30 20 10 20 10 20 20 20 10 10 20 20 20 20 20 10 10 60 60 50 50 80 40 50 50 60 60 40 70 60 80 60 50 60 70 80 50 60 80 20 50 50 70 50 50 50 40 CXl N �83 LITERATURE CITED Anderson, A. E. 1980. Experimental deer inventory, Game Res. Rep. July Part 1:1-195. northeast , and D. C. Bowden. 1979. Experimental deer inventory, region. Game Res. Rep. July Part 1:1-230. ---- Lee, R. 1963. The "topographic sampler." J. Forestry region. northeast 61(12):922-923. Neff, D. J. 1968. The pellet group count technique for big game trend, census, and distribution: a review. J. Wildl. Manage. 32(3):597-614. U.S. Geological Survey. 1967. Topographic instructions of the United States Geologic Survey. Book 3 - Mapping procedures, Part 3A Standards for the treatment of map features, Chapter 3A8 - Woodland. U.S. Dept. Interior, Washington, D.C. ii + 4p. Prepared by --'---QtM~U!IJJ.J"",~~~=-=-· ~t2c.;.~~"-c~;..o!!:A:&oo.l{ •• -<bt~!o...o/~ Allen E. Anderson Wildlife Researcher C _ ��July 1981 85 JOB FINAL REPORT State of Colorado --~~~~~---------- Project No. W-126-R-4 _Big Game Investigations Work Plan No. Job No. 2 _ 5NW Job Title: Dee£_In~estigatio~s - Experimental Deer Inventory NW Region Period Covered: Personnel: July 1, 1980 through June 30, 1981 D. J. Freddy, D. C. Bowden ABSTRACT Two manuscripts were completed during this segment: "Sainp1ingmule deer pellet group densities in juniper-pinyon woodland" and "Efficacy of permanent and temporary pellet plots in juniper-pinyon woodland." These manuscripts will be submitted to the Journal of Wildlife Management. ��87 EXPERIMENTAL DEER INVENTORY - PICEANCE BASIN - NORTHWEST REGION David J. Freddy RECOMMENDATIONS 1. A cadastral mi2 (2.59 km2), when utilized as the primary sampling unit of a 2-stage sampling system, effectively estimated densities of mule deer pellet groups in the Piceance Basin. The 2-stage sampling system provided precision estimates of ± 10%, 90 percent of the time when 100 O.OOl-ha pellet plots were used to subsample primary sampling units an2 4% of the potential primary sampling units were sampled. The mi unit should be used in future measurements of pellet group densities. 2. Analysis of subsampling systems using less than 100 pellet plots/ primary sampling unit showed that the most effective plan for personnel logistics and precision was 4 transects of 10 plots spaced at 0.16 km intervals. Transects would be paired at distances not exceeding 0.32 km and transect pairs would be selected from 5 possible pairs denoted for each sample unit. However, when subsampling was reduced below 100 pellet plots/sampling unit, to maintain the same precision levels it became necessary to increase number of sample units. 3. In a 3-year test to compare effectiveness of permanently marked and temporary unmarked O.OOl-ha plots in estimating densities of mule deer pellet groups on 10-mi2 (2.59 km2) sample units in the Piceance Basin it was found that there were no differences between plot types within years (P > 0.05) or p09led over 3 years (P > 0.05). Completion time for transects of 10 temporary plots was 16% less than time required for transects of 10 permenent plots. Because temporary plots provided equitable estimates of pellet group density for less cost they are recommended in areas where length of deer occupancy can be reasonably estimated and new and old pellet groups differentiated. Effectiveness of temporary plots in this study was likely obtained by carefully differentiating new from old pellet groups. To differentiate new and old groups, pellet group aging plots were established to create criteria upon which ages of groups could be estimated. These aging plots are highly recommended for future studies utilizing temporary plots. P. N. OBJECTIVE To design appropriate sampling and analytical procedures necessary to reliably estimate deer numbers and buck:doe:fawn ratios on winter range based on a preliminary sampling of a problem management unit within each of the four administrative regions of the state. �88 METHODS AND MATERIALS Methods have been presented in detail by Freddy (1977, 1978). RESULTS AND DISCUSSION Two manuscripts were completed during this segment: "Sampling mule deer pellet group densities in juniper-pinyon woodland" and "Efficacy of permanent and temporary pellet plots in juniper-pinyon woodland." These manuscripts will be submitted to the Journal of Wildlife Management. A third manuscript "Aerial and ground sampling systems for estimating mule deer sex and age ratios" will be completed in cooperation with project W-126-R, Work Plan 2 Job 9, Piceance Deer Study. LITERATURE CITED Freddy, D. J. 1977. Experimental deer inventory - Piceance Basin. Div. Wildl. Game Res. Rep. July, Part 2:254-273. 1978. Experimental deer inventory - Piceance Basin. Wildl. Game Res. Rep. July, Part 2:245-263. Colo. Colo. Div. �July 1981 89 JOB FINAL REPORT State of Colorado Project No. W-126-R-4 Work Plan No. 2 Big Game Investigations Job No. SSE Job Title: Deer Investigations - Experimental Deer Inventory .Period Covered: Personnel: SE Region July 1, 1980 through June 30, 1981 Jake Rodriquez, Bill Kendall, Stan Ogilvie, Dwayne Finch, Thomas Pojar ABSTRACT Through suppor t of management personnel census of mule deer (Odocoileus hemionus) using the pellet group technique was completed on the Canon City area during this segment. Previous work indicated that definition of number of pellets that constitute a group had a significant bearing on the population estimate. Therefore, during this census, only "obvious" groups were counted. The resulting average population estimate for GMU's 58 and 581 was 12,062 with 90% confidence limits of + 3,737 deer. ��91 EXPERIMENTAL DEER INVENTORY SOUTHEAST REGION Thomas M. Pojar A final Federal Aid report was submitted for this work plan and job last segment. However, through the cooperation of management personnel it was possible to execute this census again during this segment. Therefore, the final publication for the project was postponed so that this year1s data could be included. P. N. OBJECTIVE Design appropriate sampling and analytical procedures necessary to reliably estimate deer numbers and buck:doe:fawn ratios of selected management units in the southeast region of the state. SEGMENT OBJECTIVE 1. Clear and read pellet group count plots in Game Management and 581. METHODS Units 58 AND MATERIALS Sampling design and methods have been described in previous Federal Aid reports (Pojar 1977, Pojar 1978, Pojar 1979, Pojar 1980). Number of pellets per group deposited by mule deer (Odocoileus hemionus) ranged from 42 to 320 (~ = 132, SE = 12, n = 34) in Northwest Colorado' (Strong and Freddy 1979) and from 69 to 303 (~= 157, SE = 12, n = 89) in Southeast Colorado (Pojar 1980). Based on this information, groups were defined as 30 or more pellets of similar size, shape and color that were judged to have been deposited in a single defecation. Groups fitting this description are usually obvious and require minimal search time under conditions of slight vegetative ground cover encountered on the study area. This definition of a group was most practical for the Canon, City area because of the apparent high likelihood of pellets being washed onto the cleared plots from the surrounding area. Heavy summer rainstorms, steep terrain and lack of vegetative ground cove~ allow pellets to be transported quite readily from their original place of deposition. Permanent plots are read and cleared only once a year because the area is considered year-round range for mule deer and the longer time interval enhances chances for pellets from outside the plot to be transported onto the cleared plots. Management personnel executed the census during this segment with the assistance of one individual that had worked with the research project for the two previous years. �92 On each of the 64 sample transects, the first obvious intact group encountered was collected in a plastic bag. The group was then re-deposited 5.352m east of the center of the plot on which it was found. The group location was marked with a steel rebar stake, painted with blaze orange spray paint and the general area was marked with orange plastic tape. The purpose of this exercise was to monitor the dispersal rate of pellet groups throughout the area. RESULTS AND DISCUSSION Of 5,120 permanent plots on the area, 428 were destroyed from various activities such as mining, logging and road building. Therefore total number of plots read was 4,692,(_Table 1). Fifty;-.onetest groups were marked for monitoring dispersal rates. Population size estimate using only "obvious" pellet groups was 12,062 deer + 3,737 (90% confidence interval). Mean number of pellets per group-of the 51 test groups was 168.1 (SE = 8.4) • • A total of 97.9 person-days was spent to execute the census on the 1,026 square mile area and 4,569 vehicle miles were driven~ LITERATURE CITED Pojar, T. M. 1977 . Experimental deer inventory - Southeast Region. Div. Wild 1. Game Res. Rep. July, Part 2:275-286. Colo. Pojar, T. M. 1978. Experimental deer inventory - Southeast Region. Div. Wildl. Game Res. Rep. July, Part 2:265-277. Colo. Pojar, T. M. 1979. Experimental deer inventory - Southeast Region. Div. Wildl. Game Res. Rep. July, Part 1:193-207. Colo. Pojar, T. M. 1980. Experiment~l,deer inventory - Southeast Region. Div. Wildl. Game Res. Rep. July, Part 1:169-191. Colo. Strong, L. L. and D. J. Freddy. 1979. Number of pellets per mule deer defecation. J. Wildl. Manage. 43:563-564. ( "- Prepared by'• 1 ,,/ ........ ./" l. '/ / "",~<,~~~ " '-C.,.';' ~ L..- 4'-;' ~/~1 /' , �Table 1. Date of search and number of "obvious" pellet groups observed. Block No. (mi2) Section Transect 1(78) 5 5 10 8-24 8-24 63 2 7 52 67 2(74) No. of Days Between Readings No. of Plots Missing No. of "Obvious" Groups No. of Groups Adjusted to 365 Days 8-30 8-30 371 371 0 0 7 6 7 6 6-29 7-1 8-31 8-31 428 426 0 0 6 3 5 3 1 6 7-12 7-13 8-26 8-26 410 409 8 1 16 10 14 9 5 10 6-28 6-27 9-10(est) 8-1 439 400 0 1 3 4 2 4 Date Read 1979 1980 1.0 w 3(72) 4(77) 5(80) 36 1 6 7-10 7-11 8-20 8-20 406 405 0 1 0 22 0 20 64 1 6 6-13 6-13 8-16 8-16 429 429 36 58 3 0 3 0 7 5 10 7-23 7-25 7-25 7-25 367 365 0 0 0 0 0 0 54 3 8 7-26 7-27 8-6 8-6 367 375 0 1 6 7 6 7 26 5 10 8-22 8-21 9-3 9-3 377 376 0 0 34 16 33 16 67 2 7 5-25 5-28 7-23 7-24 424 422 2 1 6 5 9 8 �Table 1. Block No. (mi2) 6(78) 7(78) Date of search and number of "obvious" pellet groups observed. Section Transect Date Read 1979 1980 No. of Days Between Readings No. of Plots Missing (con't) No. of "Obvious" Groups No. of Groups Adjusted to 365 Days 6 5 10 6-25 6-26 8-28 8-28 429 428 16 0 35 49 30 42 33 5 10 8-29 8-30 8-14 8-14 349 350 3 4 25 30 26 31 70 2 7 8-15 8-16 8-21 8-21 359 360 1 0 24 35 24 36 21 4 9 8-23 8-23 9-2 9-2 375 375 0 2 3 46 3 45 51 5 10 7-18 7-16 8-11 8-11 389 391 1 14 35 20 33 19 75 1 6 6-11 6-12 8-18 8-18 433 432 5 10 12 17 10 14 18 5 10 7-19 7-24 7-30 7-29 376 370 0 1 13 8 13 8 35 4 9 8-2 8-9 8-19 8-19 382 375 0 4 29 39 28 38 65 5 10 8-8 8-7 8-31 9-1 388 390 0 10 19 18 18 17 1 6 6-21 6-22 9-10 9-10 415 414 1 0 3 5 3 4 \0 ~ 8(70) 9(80) 48 �Table 1. Block No. (mf2) 10(64) 11(81) 12(66) 13(57) Date of search and number of "obvious" pellet groups observed. Section Transect Date Read 1979 1980 No. of Days Between Readings No. of Plots Missing (con't) No. of -"Obvious" Groups No. of Groups Adjusted to 365 Days 22 19 20 17 -80(A11) 0 19 17 '--,- 396 395 63 16 5 46 5 42 8-27 8-27 418 417 7 6 50 19 44 17 0 2 31 34 27 29 410 405 78 3 8 7-25 7-30 18 5 10 -- -- -- 8-1 9-10 405 , 39 5 10 7-3 7-4 8-3 8-3 43 1 6 7-5 7-6 9-8 9-8 0 0 19 5 10 6-6 6-7 8-4 8-4 424 423 47 1 6 5-31 6-1 7-31 7-31 426 425 0 36 17 6 15 5 45 5 10 5-23 5-22 7-28 7-24 431 428 0 0 22 10 19 8 46 1 6 5-29 6-4 9-5 8-18 474 440 0 0 20 29 15 24 10 5 10 --- --- 80 80 8-25 8-25 432 436 2 11 19 22 16 18 36 1 6 --6-19 6-15 1.0 \J1 �Table 1. Block No. (mi2) 14(71) Date of search and number of "obvious" pellet groups observed. Section 49 59 Transect Date Read 1979 1980 10 --- 1 6 6-14 6-18 5 -_•... 9.... 6 8-12 (can't) No. of Days Between Readings No. of Plots Missing No. of "Obvious" Groups No. of Groups Adjusted to 365 Days -- 80 80 -- -- 408 420 58 47 9 15 8 13 '1""'- \C 0\ �July 1981 97 JOB PROGRESS State of Project REPORT Colorado ---- Work Plan No. Job Title: Deer Investigations W-126-R-4 No. 2 Winter Habitat Job No. Selection 6 and Activity Patterns of Mule Deer in Front Range Shrub land and Forest Habitats Period Covered: Personnel: July 1, 1980 through June 30, 1981 R. Kufeld, T. Fowler ABSTRACT Tests were conducted on Horsetooth Mountain, 3 miles west of Fort Collins, Colorado, to determine accuracy of Telonics telemetry equipment in locating a transmitter in rugged mountain terrain, and the ability of an observer using that equipment to correctly assess deer activity patterns. Telemetry signals were separated into classes based on signal quality. Mean differences between 2 observers, using a vehicle-mounted, precision null antenna, in . determining signal direction for Class 1, 2 and 3 signals were 0.'0, 0.2 arid 0.0 degrees, respectively. A total of III transmitter points distributed randomly throughout the study area were sampled with 4 signals (one from each of 4 spots situated within a S-foot radium of the location center) being received from each of 2 triangulation points. The compass bearing from each transmitter location consisted of the mean bearing of the 4 spot Signals. Mean biases + SD for Class 1, 2 and 3 signals were + 0.5 + 1.3, + 0.1 + 2.4, and 2.9 +-9.6 degrees respectively. Terrain, as measured by Signal-clearance (in feet) of the highest terrain feature which occurred between the receiver and transmitter, appeared to be the primary factor affecting signal quality and directionality .. Distance between receiver and transmitter was not an important factor influencing signal quality or directionality on the study area. Four tame deer, equipped with activity radiocollars, were monitored visually and electronically for 50.8 hours to assemble a file of telemetry strip charts depicting known deer activity patterns. Three tame deer were then monitored for an additional 30.8 hours to test if a second, trained observer could assess deer activity patterns from strip charts. The chart reader correctly assessed 94% of the total 1,847 minutes of deer activity. ��99 WINTER HABITAT SELECTION AND ACTIVITY PATTERNS OF MULE DEER IN FRONT RANGE SHRUBLAND AND FOREST HABITATS Roland C. Kufeld P. N. OBJECTIVE 1. To test Telonics telemetry equipment to determine its accuracy at various distances in locating a transmitter on the Horsetooth Mountain study area, and the ability of an observer using that equipment to correctly detect deer activity patterns by habitat type. 2. To determine habitat selection and activity patterns of mule deer within habitat types in the Horsetooth Mountain area during winter. SEGMENT OBJECTIVES 1. To test Telonics telemetry equipment to determine its accuracy at various distances in locating a transmitter on the Horsetooth Mountain study area, and the ability of an observer using that equipment to detect deer acti~ity patterns by habitat type. 2. To capture deer on the study area and instrument them with Telonics activity collars. 3. To monitor radio-collared deer to determine habitat selection and activity patterns. METHODS AND MATERIALS This report only describes progress on Segment Objective 1. Testing took longer than expected due to unanticipated complications. Therefore, trapping and radio-collaring of wild deer was postponed until next segment. Determination of Telemetry Directional Accuracy Sampling Design, Procedures, and Initial Testing A number of potential telemetry reciever points were selected near the perimeter of the study area. Efforts were made to select sites that offered wide visibility of the area, and were as free as possible.of nearby terrain features, which could cause an incoming signal to bounce. Subsequently testing resulted in some of these being found unsuitable. Seven sites were retained, of which 4 were located on the east side, and 3 were on the west side of Horsetooth Reservoir. �100 Transmitter points were located using a 2.5-acre grid system which encompassed the entire study area. The center point of each randomly selected 2.5-acre cell became a transmitter location. A total of 111 transmitter points were sampled with the signal from each point being received from each of 2 receiver points in order to achieve triangulation. Receiver and transmitter points were plotted on aerial photos. One observer located transmitter points in the field while the other operated the receiver. Observers communicated by hand-held 2-way radios. The radio collar was positioned about 3 feet above the ground at each of 4 spots within as-foot radius of the center of each transmitter location. The signal bearing from each spot was recorded by the receiver operator. The compass bearing from a given transmitter location consisted of the mean bearing of the 4 spot signals from that location. Testing was initiated using an 8-element yagi antenna with l3-foot boom and a telescoping mast mounted on a pickup truck. The antenna was raised to 12 feet above the vehicle when in use. The mast was equipped with a compass rose and pointer for determining antenna direction. Antenna attitude was initially measured, and the compass rose set accordingly with a hand held ccmpassr This was subsequently changed as later discussed. Signals were received using a Telonics TR-2 receiver. Relative signal strength was displayed by a Telonics digital processor. Testing with the 8-clement, yagi antenna quickly revealed serious problems with signal bounce. These problems could not be resolved using that antenna given the terrain pattern on the study area. A precision null antenna, which had been developed and successfully tested in rugged mountain terrain by John Siperek of California Department of Fish and Game was constructed. The mast system was the same as used with the 8-element yagi antenna. The precision null antenna was initially oriented and calibrated with the compass rose using a hand compass. Following Mr. Siperik's advise we began orienting the antenna electronically using radio collars placed in trees in the north and south ends of Horsetooth Mountain as beacons. This improved accuracy in measuring signal ~irection by about 2 degrees. The 8-element, yagi antenna was oriented toward the transmitter at the point of strongest signal. With the precision null antenna and using a peak combiner, a pattern is established which consists of 4 peaks and 3 nulls. The antenna is oriented toward the transmitter when receiving the center null. This null is very narrow; from less than a degree to 5 or 6 degrees wide depending on signal quality. A peak occurs 15 degrees to the right and left of the center null. A null occurs 15 degrees beyond each of those peaks. Another peak occurs 15 more degrees beyond each of those 2 nulls. To the right and left of the outer peaks are nulls which are about 90 degrees in width. A similar pattern of peaks and nulls occurs on the reversed side of the antenna, but much reduced and poorly defined, so it is not easily confused with the pattern facing the transmitter. �101 Signal Classification System A classification system for signals received with the precision null antenna was devised after much experimentation as it became apparent that directional accuracy was influenced by signal quality. This system is as follows: Signal Classes I.' Loud and clear with very little or no background noise. Patterns obvious and easy to read. Good symmetrical patterns with 4 peaks and nulls. Peaks 30° apart and nulls 30° apart. Relative difference between peaks and nulls is great. Narrow V-shaped center null. 2. Loud but with background noise. Pattern fairly obvious and fairly easy to read. Symmetrical patterns with 4 peaks and 3 nulls. Peaks 30° apart and nulls 30° apart. Relative difference between peaks and nulls is great. Center null may be narrow and V-shaped or slightly ~ __ JI shaped. It may be necessary to bracket the center null by listening to,signal pickup levels 3 or 4 degrees either side of it, but it is still relatively easy to determine the null centerpoint. 3. Weak signal with much background noise. Gain must be turned up. Pattern difficult to discern and difficult to read. Requires much time to establish the pattern. Relative difference between peaks and nulls is small. Peaks are low and valleys are high. Peaks may not be symmetrical in height and peaks and nulls may be more or less than 30° apart. Nulls are wide and it is very hard to determine the center;.. point of the null. This category includes any signal with low peaks and high valleys and those where much time is needed to establish the pattern. 4. No signal or signal too weak to establish any pattern at all. 5. Signal with 2, 4 or more nulls regardless of signal strength. NOTE: A transmitter "location" consists of the average of 4 "shots" taken as,quickly as possible while the transmitter is stationary. Regardless of signal strength any transmitter location from which the bearings of the 4 shots vary more than 6 degrees from the lowest to highest bearing angle is given the overall signal classification of class 3. Measurements of Telemetry Directional Bias on the Horsetooth Mountain Study Area All 111 transmitter locations were subsequently sampled as previously described, but in addition the precision null antenna, beacons for antenna orientation, the signal classification system, and 2 triangulation points for each transmitter location were used. The recorded bearing from each transmitter location is a mean of signals from 4 spots within a 5-foot �102 radius of the transmitter location center. The digital processor was used only to record decibel levels of peaks. The center null was located by ear by sweeping a few degrees to the right and left of the null until an equal signal pitch was heard; then the 2 bearings were bisected. Comparison of Observer Ability to Measure Telemetry Signal Directionality Since both observers took turns operating the receiver and placing the transmitter during sampling of the 111 transmitter locations, a test was conducted to determine degree of directional bias between observers. Three people were required for this test. One individual placed the radio collar transmitter at each of 20 locations situated at 100-foot intervals in a basin area on the west side of Horsetooth Reservoir. Some of the locations were line of sight from the receiving point, which was located 0.5 mile away on the east side of the reservoir. Some transmitter locations were below a 25-foot cliff in the b~sin itself, from which the signal had to go over the cliff to reach the transmitter. Procedures for this test were exactly the same as used for sampling directional bias on the study area, except that signals were received by both of the observers who conducted the overall study area directional tests. The signal from each spot within a 5-foot radius of the transmitter location center was received and recorded by both obervers before the radio collar was moved to the next of the 4 location spots, or to another of the 20 transmitter locations. The recorded bearing from each transmitter location for each observer' is the mean bearing of his signals from the 4 spots. Measurement of the Influence and Directionality of Distance and Terrain on Signal Quality The 111 randomly selected transmitter locations and those receiver locations used to sample them were plotted on a U.S.G.S. topographic map with 40 foot contours. Receiver-transmitter dist?nce (in feet), and signal clearance (in feet) of the highest terrain feature which occurred between the receiver and transmitter, were measured on the contour map. To determine signal clearance for line-of-sight signals, the vertical distance (in feet) between the sight line and the top of the highest terrain feature, which occurred between the receiver and transmitter, was recorded and assigned a plus (+). If the signal path was not line-of-sight, indicating the signal had to pass over a terrain obstacle, the distance (in feet) between top of the highest terrain feature and a straight line between the receiver and transmitter, through the terrain obstacle, was recorded and assigned a minus (-). Directional bias within signal class was then compared with receiver-transmitter distance and with signal clearance. Determination of Tame Deer Activity Establishment of a File of Telemetry Activity Patterns Strip Charts Patterns Showing Observed Deer Four radio-collared tame deer were monitored electronically and visually for a total of 50.8 hours. One deer was monitored at a time. Telemetry �103 data and written visual observations were recorded on a Rustrak strip chart recorder attached to a Telonics receiver and digital processor, to develop a file of charts which depicited known activity patterns of deer. Deer were equipped with Telonics activity collars which indicated head-up head-down activity. The strip chart recorder recorded radio signals indicating if the head was up or down, and if the animal was stationary or moving. Receiving equipment was mounted in a metal box attached to a pack frame so the deer could be constantly observed. The observer marked and recorded the activity on the strip chart each time the deer changed its activity pattern. Activity patterns were categorized as follows: roving; roving emphasis standing; roving emphasis feeding; feeding; and resting. The deer was considered to be engaging in a particular activity when more than half of its effort was devoted to that activity. For example, the deer was considered to be in a feeding pattern if feeding actively, and feeding was its primary concern. A feeding pattern included short intervals of walking from plant to plant or standing and looking around between a series of bites. Some feeding also occurred while roving, but the deer was considered to be in a roving pattern if primarily engaged in anything other than feeding actively or resting. The deer was considered to be resting only when lying down. Monitoring was done at 2 locations. One spot was adjacent to the Colorado Division of Wildlife research pens on the northwest edge of Fort Collins where the tame deer were housed. The other site was within Lory State Park on the Horsetooth Mountain study area. Monitoring sites were in the mountain mahogany vegetation type where deer had a wide choice of grass and browse. This allowed the deer maximum opportunity to feed head-up or head-down. Monitoring was done in February and March, 1981. Monitoring was done by 2 observers so that both could learn to recognize activity patterns when observing the deer and by reading strip charts. This was necessary to test observer ability in assessing deer activity patterns from strip chart reading. Determination of Observer Ability to Read Strip Charts and Accurately Assess Deer Activity Patterns Three radio-collared tame deer were monitored electronically and visually for a total of 30.8 additional hours, during 8 different days. Monitoring was done during March, 1981. Procedures were the same except that monitoring was done by only 1 observer, and code numbers, known only to him, were entered on the chart to identify periods of observed deer activity. These charts were later read by the second observer who had attained a degree of proficiency in chart reading from his involvement in the earlier period of tame deer monitoring. His assessment of deer activity was compared with actual activity recorded in coded numbers by the observer who monitored the animals. To further test the second observer's ability to assess deer activity that observer, using a second receiver, digital processor, and strip chart �104 recorder, simultaneously monitored the deer (not seen) from a vehicle 0.25 miles away. This was done for 6.5 of the total 30.8 hours of deer monitoring. This provided a second strip chart on which no marks or code numbers had been entered to indicate deer activity. The second observer made his assessment of deer activities from this chart, and compared his evaluation with the actual activities as recorded by the observer who accompanied the deer. DESCRIPTION OF AREA The study area is located approximately 3 miles west of Fort Collins, Colorado, and includes all of the east side of Horsetooth Mountain. The area is bounded on the east by Horsetooth Reservoir and on the west by the Crest of Horsetooth Mountain. The southern boundary is the surfaced county road which traverses the south side of the reservoir, and Empire Gulch is the northern boundary. The area encompasses about 9 square miles. Elevation ranges from 5,400 feet at the reservoir shoreline to 7,255 feet at the highest point on Horsetooth Mountain. Terrain is characterized by numerous rock outcrops, ridges, and canyons. Vegetation types include Ponderosa pine (Pinus ponderosa) with various canopy densities, and interspersed with Douglas fir (Pseudostuga men~iesii); mountain shrub consisting primarily of mountain mahogany (Cercocarpus montanus); wet meadow; grassland; and a riparian type with cottonwood (Populus sargentii), chokecherry, (Prunus virginiana), and wild plum (Prunus americana). RESULTS AND DISCUSSION Determination of Telemetry Directional Accuracy Accuracy Comparisons Between Observers Mean differences between observers in determining telemetry signal direction for class 1, 2 and 3 signals were 6.0, 0.2, and 0.0 degrees, respectively (Table 1). Observer differences for individual and combined signal classes were not significant (p > 0.5). There was 95% agreement by observers on classification of signals among the 20 locations. This suggests signal classes are well defined, and signals of each class can be readily recognized. �105 Table 1. Signal Class a 1 2 3 Comparison of telemetry directional accuracy between 2 observers. No. of Transmitter Locations Total No. of Signals 3 12 10 7 40 28 Degrees of Observer bias per Location Statistic Observer 1 Observer 2 Difference x + 0.10 + 0.10 0:00 s 0.78 0.58 0.15 x + 0.40 + 0.20 0.20 s 1.27 1.22 0.61 x + 6.20 + 6.20 0.00 s 6.03 5.70 0.31 a There was 95% agreement by observers on signal classification. Accuracy of Signals from Transmitter Locations Selected at Random Throughout the Study Area Differences in determination of directional accuracy occurred between signal classes (Table 2). Mean biases for class 1, 2 and 3 signals were + 0.5, + 0.1 and - 2.9 degrees, respectively. Acceptable standard deviations of 1.31 and 2.36, respectively, were recorded for class 1 and 2 signals. The large standard deviation of 9.59 for class 3 signals results in unacceptable accuracy for that class of signal. Standard deviations were used to compute 75 and 90% tolerance levels (nixon and Massey, 1957), which permit the statement that one can be 90% confident (using the same equipment and procedures, and under existing terrain conditions) that 75 or 90% of all signals of a particular class will be within X number of degrees of the actual transmitter location (Table 2). With 90% confidence the computed 75 and 90% tolerance limits for class 1 signals were + 1.7 and + 2.4 degrees respectively. For class 2 signals the 75% tolerance limit was + 3.1 degrees, and the 90% limit was + 4.4 degrees. Tolerance limits for-class 3 signals were very large; 12.6 and 18.0 degrees, respectively, for the 75 and 90% tolerance levels. Thus, during future monitoring of radio-collared deer it is recommended that class 3 signals be discarded. �106 Table 2. Directional accuracy of telemetry signals received from randomly selected transmitter locations on the Horsetooth Mountain study area. Signal Class No. of Transmitter to Receiver Location a "Shots" Degrees of bias per Transmitter Location x s Tolerance limits at b 90% Confidence level 90% 75% Tolerance Tolerance 1 77 + 0.5 1.31 + lor 2 71 + 0.1 2.36 3 65 - 2.9 9.59 + 3.10 +12.60 4 4 5 3 c + 2.40 + 4.40 +18.00 a Each of III transmitter points, was sampled from 2 receiver points totalling 222 "Shots". b See page 130, Dixon and Massey (1957). c Two more class 3 signals received, however, quality was so poor compass bearing could not be determined. Influence of Distance and Terrain on Signal Quality and Directionality Distance of the transmitter from the receiver did not appear to be an important factor influencing signal quality or directionality on the study area. There was no correlation between distance and directional bias of signals within individual classes (class 1, r = 0.02; class 2, r= -0.06; class 3, r= 0.07). Class 1 signals, which had the narrowest tolerance limits, averaged 7,132 feet between receiver and transmitter, while class 2 signals, with slightly wider tolerance limits, averaged 8,626 feet. Mean distance for class 3 signals, with very wide tolerance limits, was 9,537 feet (Table 3). However, despite the increased mean distance with decreased signal quality and directional accuracy, there was much overlap in distances among the 3 signal classes. Class 1 signals were received from as close as 1,843 feet and as far as 13,568 feet, while class 2 and 3 signals ranged between 2,078 to 14,156 feet, and 3,412 to 15,059 feet, respectively. The increased mean distance for poorer quality signals seems to be more a function of the terrain pattern on the study area. The farther the transmitter was from the receiver the higher it was on Horsetooth Mountain, and this increased the amount of rugged terrain over which the signal had to pass in order to reach the receiver. Terrain appeared to be the primary factor affecting signal quality and directionality. Seventy-one percent of all class 1 signals were line-ofsight, and the mean clearance for all signals in class 1 was + 71 feet (Table 4). Among those class 1 signals which were not line-of-sight, 22.5% were transmitted from within 100 feet of the top of the highest �107 terrain feature between receiver and transmitter, and 14% were emitted from 100 to 400 feet below the top. Among class 2 signals, 36.5% were line-of-sight, and the mean clearance was -15 feet. Among class 2 signals which were not line-of-sight, 46.6% were sent from within 100 feet of the top of the highest terrain feature, and 16.9% were sent from 100 to 500 feet below the top. Only 9% of class 3 signals were line-of-sight, and the mean clearance was -136 feet. Among those class 3 signals not lineof-sight, 46.2% were transmitted from within 100 feet of the top of the highest terrain feature, and 44.8% were emitted from 100 to 500 feet below the top. Only 4 class 4 signals were received from the randomly selected transmitter locations on the study area, but all 4 were emitted from 300 to 500 feet below the top of the highest terrain feature between receiver and transmitter. Class 1 and 2 signals were not significantly different in terrain clearance (P > 0.05), while significant differences (p < 0.05) were shown between class 1 and 3, and class 2 and 3 signals. This attempt to measure influence of terrain on signal quality and directionality considers only the single, and most obvious' terrain feature, which is the highest feature, elevationally, that occurs between receiver and transmitter. Terrain features come in many sizes, shapes, and patterns, and this attempt to show effect of terrain on signal quality and directionality by measuring only one feature is very crude. The writer knows of no way to measure influence of multiple terrain features on an invisible radio signal. However, if only one terrain feature has as much influence on signal quality and directionality as is shown in Table 4, then it can be' assumed that an even closer relationship between terrain, and signal quality and directionality, would have been shown if influence of all existing terrain features could have been measured. Determination of Tame Deer Activity Establishment of a File of Telemetry Activity Patterns Strip Charts Patterns Showing Observed Deer The strip chart was divided into 2 parts and the recorder employed 2 separate printing styluses. The stylus on the "period" side printed within an inch~wide bracket. A line down the left side of the bracket reflected head-up activity, while a line to the right side indicated head-down. Dots in the middle indicated head bobbing, or moving up to down or down to up. The stylus on the "amplitude" side recorded movement. The wider the pattern of dots at a given point on the chart the greater the movement. A narrow pattern of dots forming a line indicated no movement. Resting: When lying down the head was always up except for rare occasions when the deer lowered its head momentarily to nibble some~. thing. Period side shows a narrow line on the left side of the bracket. Few or no dots occur in the middle of the bracket. Amplitude side shows a narrow, straight line. Resting activity usually lasts more than 10 minutes. �108 Table 3. Distances (in feet) between receiver and transmitter for randomly selected transmitter locations on the Horsetooth Mountain study areaa. Signal Class 1 2 3 Mean 7132 8626 9537 Minimum 1843 2078 3412 Maximum 13568 14156 15059 Distance (Feet) a Each of III transmitter points were sampled from 2 receiver points tota:lling 213 "Shots" which resulted in class 1, 2 and useable class 3 signals. Table 4. Percentage of signals in each of 4 classes which occurred foot interval categories above (+) and below (-) line of sight. Signal Clearance in feet a in 100- Signal Class 2 1 3 4 + 401' to + 500' 0.0 0.0 3.9 0·0 301' to + 400' 5.2 5.6 0.0 0.0 + + 201' to + 300' 0.0 0.0 5.2 1.4 + 101' to + 200' 0.0 5.2 7.0 0.0 0' to + 100' 22.5 0.0 51.9 9.0 -------------------------------------------------_"_----------------------Top of Highest Terrain Feature Between Receiver and Transmitter -------------------------------------------------------------------------- - - 0' 101' 201' 301' 401' to to to to to - 100' - 200' - 300' - 400' - 500' Total + Total Total + and - Total No. of Signals Mean Clearance a . F eet.b 1n 22.5 7.0 1.4 5.6 0.0 46.6 8.5 5.6 1.4 1.4 46.2 17.9 14.9 6.0 6.0 0.0 0.0 0.0 50.0 50.0 71.4 '28.6 100.0 36.5 63.5 100.0 9.0 91.0 100.0 0·0 100.0 100.0 77 71 65 4 +71 -15 -136 -400 Signal clearance of the highest terrain feature between the receiver and transmitter in feet. Plus (+) values are above line of sight, minus (-) values are below line of sight. b Class 1 and 2 signals are not significantly different in terrain clearance (p > 0.05), Class 1 and 3 signals are significantly different in terrain clearance (p < 0.05), Class 2 and ~ignals are significantly different in terrain clearance (p < 0.05). --- �109 Roving: Stan~ing in a fixed stare produces the same chart pattern as resting except that activity rarely lasts more than 8 to 10 minutes. When walking or running, amplitude is relatively wide, and the period side shows head is up or bobbing. When walking downhill, the period side shows head is sometimes down, but movement is indicated by a wide amplitude pattern. When feeding intermittently, while in a roving pattern, the head is up, then down, and there is much head bobbing. A roving pattern is usually characterized by numerous dots in the center of the period bracket unless the deer is standing and looking. Feeding: Amplitude is usually narrow indicating the deer is moving very little except for short periods of walking. A feeding pattern is always indicated by much head-down activity. Deer may feed head-up or down, but spend a period of time at each. The period bracket in the chart is relatively clean in the middle compared with a roving pattern, indicating occupation with either head-up or head-down feeding. With tame deer, feeding patterns often occurred immediately after a period of resting. The key to differentiating between actively feeding, and roving with emphasis on feeding, is whether the deer was primarily concerned with feeding or with roving. Determination of Observer Ability to Read Telemetry Strip Charts and Accurately Assess Deer Activity Patterns Using the strip chart on which observed activities of 3 tame deer had been identified in coded numbers by an observer who monitored the deer, a second observer correctly assessed 1734 minutes or 94% of the total 1847 minutes of charted deer activity (Table 5). Within the roving category, 95% was assessed correctly as roving, and 5% was called feeding. Within the Table 5. Assessment of activity patterns of 3 radio-collared tame deer by a second observer from a strip chart on which activities had been marked and coded. Observed Activity Minutes Roving Feeding Resting all 1007 350 490 1847 Assessed Minutes Activity Accuracy of Assessment % Incorrect % Correct Roving Feeding Resting 959 48 0 95 Feeding Roving Resting 334 16 0 95 Resting Roving Feeding 441 46 3 90 all 1734 94% 5 5 9 1 6% �110 feeding category 95% was assessed correctly as feeding, and 5% was called roving. Within the resting category, 90% was correctly assessed as resting, 9% was called roving, and 1% was called feeding. The 9% error in the resting category occurred when the chart reader, in several instances, thought a deer was resting when it was actually standing in a fixed stare for an unusually long period of time. Standing fits into the roving category. Using the chart from the vehicle-mounted recorder which had no coded numbers or marks, the same observer was 95% co~rect in his assessment of roving activity, 98% correct on feeding, and 99% correct on identifying resting activity (Table 6). Table 6. Assessment of activity patterns of 3 radio-collared tame deer by a second observer from a strip chart on which activities had not been denoted.a Observed Activity Roving Feeding Resting Minutes of Activity Actual Assessed 97 167 128 102 163 127 % Correct Assessment 95 98 99 a This chart was made simultaneously and later compared with another chart where deer activities had been indicated in coded numbers by an observer. LITERATURE CITED Dixon, W. J., and F. J. Massey, Jr. 1957. Introduction to statistical analysis. McGraw-Hill, N. Y. 2nd Ed. 488pp. Prepared by '/ k:t_{ 6A-1-c{ ~r_ed . C~ <-1/ /) Roland C. Kufeld Wildlife Researcher C �July 1981 111 JOB FINAL REPORT Colorado State of Project __ No. W-126-R-4 Work Plan No. Job Title: Period - Job No. Deer Investigations Covered: Personnel: 2 Big Game Invest_i~g_a_t_i_o_n_s - Experimental _ 7 Range Fertilization July 1, 1980 through June 30, 1981 Dr. L. Carpenter, P. Boultz. L. Strong, D. Bowden, G. Kontour, A. Foster, ABSTRACT All data were gathered and analyses made of the la-year yield response of vegetation on 3 sagebrush winter range sites in Colorado to a single application of 2,4-D herbicide and nitrogen fertilizer. Work was begun on summary of the data and subsequent preparation of a manuscript to be submitted to the Journal of Range Management. ��113 EXPERIMENTAL RANGE FERTILIZATION Dr. Len H. Carpenter P. N. OBJECTIVE To measure the effects of a single application of nitrogen fertilizer and 2,4-D herbicide (applied singly and together) on the production and composition of mule deer forages on sagebrush winter range sites. METHODS Methods and procedures AND MATERIALS for this study were detailed by Carpenter 1977. RESULTS AND DISCUSSION The copious amount of yield data obtained in this lO-year study has been analyzed and mostly summarized. Each year's data was subjected to various analyses of covariance and linear regression. A change in job assignments and duties for the principal investigator has resulted in this job not being completed at this time. Final results will be presented in a manuscript submitted to the Journal of Range Management entitled "Ten-year Response of Sagebrush Rangeland to a Single Application of 2,4-D Herbicide and Nitrogen Fertilizer." This will be done in the next segment. LITERATURE CITED Carpenter, L. H. 1977. Middle Park deer study - range fertilization. Colo. Div. Wildl. Game Res. Rep. July. Part 1:43-59. Prepared by _A_..Q..",_, __ #_· (q_~---,~,__ Len H. Carpenter Big Game Research _ Leader ��July 1981 ll5 JOB PROGRESS State of Project Colorado No. Jc;b Title: ----- 3 _ Elk Investigations Period Covered: Personnel: ------ W-126-R-4 Work Plan No. ___ REPORT Big Game Investigations Job No. - Elk Population 2 and Ecological Studies July 1, 1980 - June 30, 1981 G. Bear, R. Green, other personnel are listed in Appendix B ABSTRACT Radio-collared elk wintering near Masonville migrated to the northern section of Rocky Mountain National Park for the summer, while elk wintering near Lyons migrated to the southern portion of the Park for the summer. Radio-collared elk wintering in the Estes Valley migrated to summer ranges along the Colorado River (Kawuneeche Valley), Specimen Mountain and the headwaters of Fall River, and Forest Canyon. Population estimates indicate approximately 2,000 elk winter in the Estes Valley near Estes Park. Aerial and ground surveys indicated calf:cow ratios of 41:100. Known mortality of radio-collared calves was determined to be 29.6%, with deaths attributed to predators, PI3 virus, starvation, thyroid atrophy, and fences. Mortality of ear-tagged elk was determined to be 8.1% hunter harvest, 2.0% malnutrition, 0.5% auto accident, and 3.0% other causes. Dominant habitat types used by elk during summer were krumholz (74.7%) and spruce-fir (21.9%). The main habitat type used during winter was ponderosapine shrub (49.2%), coniferous (12.4%) and wet meadow (19.7%) differences in seasonal and daily activity patterns were apparent with summer being the period of highest grazing activity and winter the period of highest resting activity. Daily cycles in activity were also exhibited with crepuscular and nocturnal feeding periods and diurnal resting patterns. ��117 ELK POPULATION AND ECOLOGICAL STUDIES George D. Bear and Ronald A. Green P. N. OBJECTIVES 1. Develop techniques to more accurately and precisely estimate elk population levels. 2. Define natality and mortality problems of selected elk populations. 3. Determine seasonal movements, daily activity patterns, and habitat preferences of elk in Rocky Mountain National Park and adjacent seasonal ranges. SEGMENT OBJECTIVES 1. Capture and mark 20 elk calves with mortality collars. 2. Trap and mark up to 200 elk on the winter range. 3. Monitor telemetry collared elk to determine mortality rates and probable causes, seasonal and daily activities, and habitat selection and preferences. 4. Conduct aerial censuses and production surveys. METHODS AND MATERIALS Methods and materials have been previously described in detail by Bear and Green (1980). A master's thesis on habitat selection of elk by Ronald Green is in preparation and will be on file at Colorado State University. RESULTS AND DISCUSSION Population Distribution Only 2 of the 5 elk marked with telemetry collars near Masonville in 1978-79 were functional during this segment. These animals exhibited similar seasonal migration patterns to those reported previously (Bear and Green 1980). Elk summered in the northeast corner of Rocky Mountain National Park in the Stormy Peaks area, returning to winter range on Green Ridge in the fall. One collared elk was harvested by a hunter during the hunting season leaving only 1 radio-collared elk remaining in the Masonville herd. Three of 4 radios placed on elk wintering near Lyons in 1979 were still active during this segment. Migration patterns of these elk were identical to those previously reported. They wintered on �118 the ridges between Longmont Reservoir and Big Elk Meadows, migrating to alpine ranges at the headwaters of the St. Vrain River in the southeast portion of Rocky Mountain National Park for the summer. These elk moved back to Big Elk Meadows in the fall. One elk lost its collar during migration to the winter range, so only 2 collars are still active in this group. Movements of 48 radio-collared elk were monitored in Estes Valley and Rocky Mountain National Park. Migration patterns were similar to those reported for these animals in previous years. They wintered in the eastern portion of Rocky Mountain National Park (Horseshoe Park, Beaver Meadows, and Morraine Park) and throughout Estes Valley to Crosier Mountain and the Crocker Ranch east of Estes Park. In late May and early June elk migrated to summer ranges along the Colorado River drainage, and to alpine ranges at the heads of Forest Canyon, Poudre RiverSpecimen Mountain and Roaring River. In the fall (September-October) these migration patterns were reversed, and elk returned to winter ranges in Horseshoe Park, Beaver Meadows, and Morraine Park. The 1980-81 \vinter was very mild with little snow accumulation as compared to the winter 1979-80. There was a distinct difference in the eastward movement of the elk. In 1979-80,52% of the radio-collared elk moved eastward from Rocky Mountain National Park to the east side of Estes Valley and in addition several others moved down to the eastern portion of Crosier Mountain. In 1980-81 only 40% of the radio-collared elk moved eastward out of the Park and many of these were only brief visits before moving back to winter ranges within the-Park. Also,· the eastward movement extended only to the very western portion of Crosier Mountain •. Snow melt on the summer ranges was earlier in 1981 and elk migrated 7-10 days earlier than in 1980. Most elk migrating to the summer ranges along the Colorado river drainage did so between May 29 to June 6, 1980. Then 83% moved to the winter range during the period October 16 to October 23, 1980. Most elk migrating to the summer range at the heads of Forest Canyon and Poudre River during the period from June 15 to June 30, 1980. Eighty-one migrated to the winter range during the period October 16 to October 23, 1980. Many of these migrations, required only 2-4 days. Four elk were radio-collared on the Crocker Ranch (east of Estes Park) in February 1980. Migration. patterns for these animals during this segment were similar to those reported last year. One animal migrated to the Colorado River, and the remaining 3 summered south of the Crocker Ranch near Twin Sisters Peak. Population Estimation Two hundred and one elk (98 cows, 70 calves, and 33 bulls) were captured and marked during the period from December 9, 1980 to January 22, 1981. �119 Overall trapping success was 54%. The area was free of snow throughout trapping with temperatures above seasonal normals. Twenty-eight were recaptures from last year, and 27 were recaptures of elk tagged this year (Appendix A). Two elk (1 adult male and 1 adult cow) died during trapping. Three population surveys were flown during February and March (Table 1). Efforts to obtain a helicopter when census conditions were optimum (solid snow cover, calm winds, clear skies) were fruitless. Also, an effort to increase sample size with additional flights was hindered, since a helicopter was not obtainable. The average population estimate was 2,648 (+ 966) elk. This may be misleading since 2 of the surveys yielded esti;ates of 2,029 and 2,154 elk based on observation of blue tags (1980-81) and 2,211 elk based on orange tags (1979-80). Whereas, the 3rd estimates based on the blue tags was 3,761 elk. The blue ear tags used this year were very difficult to detect, thus when large herds were surveyed (as occurred on the later f1ight or estimate mentioned) it was believed the marked animals were not detected as readily as on the other flights when fewer animals and smaller groups were surveyed. A series of ground or roadside surveys was attempted during February-May to obtain independent population estimates (Table 2). These surveys were limited to large open areas and roadsides; thus sample sizes were small. Average population estimate based on 1980-81 blue tags was 1,462. Table 1. : Population estimate of elk based on aerial surveys winter range during February and March, 1981. on the Total Counted Blue 2/23/81 451 44 2029 3/10/81 930 49 3761 3/19/81 383 35 Date Tags Observed Orange 24 Population Estimate 2l54(B) 2211(0) �120 Table 2. Population of elk based on ground surveys on the winter range during February-April, 1981. Date Total Counted Tags Observed Orange Blue Population Estimate Orange Blue 2/25/81 486 65 3/17/81 576 78 25 1475 3195 4/8/81 351 57 27 1226 1810 4/21/81 154 21 5 1423 3720 5/18/81 193 22 17 1694 1551 5/27 to 6/11/8la 512 64 41 1880 2041 a 1491 Random observations on upper winter range during calving season. A second series of ground surveys or observations was recorded during the first half of June before elk left the winter range. Animal observations were recorded without regard to duplication on a day-to-day basis. A total of 512 elk was classified with a popula t Lon estimate of 1880 elk based on observations of blue ear-tags and 2,041 elk-based on orange ear-tags. These population estimates were more similar to ~ach other and aerial surveys than roadside surveys mentioned earlier. The author believes there was a thorough mixing of individual animals as they congregated for the spring migration. This was based on observations of elk tagged on the east side of Estes Valley near Beaver Meadows and Horseshoe Park within Rocky Mountain National Park. This m1x1ng may have resulted in a random sample of marked elk approximated on the aerial surveys. Hopefully aerial sampling can be improved during the next segment. First a more observable ear;tag or color of ear-tag will be used. Secondly, helicopter services should be improved. Also, a population estimate of 1706 elk was derived from the number of elk recaptured during this segment (1980-81). These elk were first tagged a year ago. However, this estimate would be based on the assumption all deaths or removals of orange ear-tags are known, which is unlikely. ALthough a large percentage of the tags may have been found and reported since hunter harvest is a large percentage of known mortality and winter loss was easily detected on the winter range when elk were concentrated. �121 Mortality Twenty-nine elk calves were captured and collared in June-July, 1980. Two of the calves lost their collars within a few weeks. Six (22.2%) of the remaining 27 calves died within a month after they were collared, while the remainder survived into the winter period. Death of these calves was attributed to predator (1), PI3 virus (1), starvation (2), thyroid atrophy (1), and unknown (1). In late winter 1 calf died in a fence, and another died of malnutrition. Twelve other calves lost their collars during the winter and spring period; 8 of these pulled the collar~ off in fences. Four collars have ceased to function; while 3 are still active. Measurements of Calf:cow ratios conducted on the summer range appeared low. Ground surveys indicated 41 calves:100 cows, while aerial surveys indicated 41 and 44 ~lves:lOO cows for 2 surveys (Tables 3 and 4). Ground surveys also indicated a very low yearling:c0w ratio (18:100 cows). Data from radio-collared calves indicated a good survival rate for calves (70%) following birth, thus low calf :cow ratios may be due to prenatal mortality or to calf mortality in the first 1 to 2 days after birth. Table 3. Ground-classification of elk, July-October 1980. Month Location Cows Calves Ypsilon Chapin Creek Poudre River Specimen Mountain Total 33 25 29 49 136 12 11 13 18 54 58 39 14 III 18 20 10 48 390 160 July a Yearlings Cows Bulls 2 4 5 9 20 1 4 3 "8 Bulls 3 3 10 16 Calves :'100cows: Yearlings :'100cows: 21 August a Forest Canyon Specimen Mountain Poudre River Total 6 8 2 10 6 1 7 '6 6 13 CalvesilOO cows:43 Yearlings :'10cows:15 OctoberNovember Horseshoe Park to Beaver Meadows Calves I: 10 cows:41 a Overall average for summer = Calves:lOO cows:4l Yearlings:lOO cows:18 �122 Table 4. Aerial classification of elk on the summer range. Date Total Counted Cows Calves Bulls Spikes Adult Calf: Cow Ratio 8/22/80 575 336 148 5 86 44 9/4/80 689 356 146 34 153 41 Eighteen calves were captured and collared in 1981 (Table 5). There were 11 males, 6 females, and 1 unknown (escaped before the sex was determined). Due to lack of snow accumulation last winter, snow melt and "green-up" was earlier this year. Consequently elk moved to the summer range earlier and calving occurred on alpine ranges and willow flats. All calves were captured on the summer range, whereas, in other years approx~mately half the calves were marked while still on the winter range. Miscellaneous observations of mortality indicated losses were extremely light during the i980-8l winter. Four dead elk were reported by Park Rangers, 3 deaths were attributed to malnutrition and 1 to coyotes. All 4 were calves. In addition there were the 2 radio-collared calves men~ tioned earlier, 1 adult cow crippled by an automobile, and another adult cow was found with a broken hind leg. Both of these animals were destroyed by Park Rangers. Mortality of banded animals was light (12.5%) for the 198 elk tagged in the last segment (1979-80). Hunters harvested 8.1% (16 elk), 2.0% (4 elk) died of malnutrition, 0.5% (1 elk) died as a result of an auto accident, and 3.0% (6 elk) died of unknown causes. Only 4 adult elk wearing radio-collars were shot by hunters. This was 10.5% of adult animals, radio-collared. Light hunter harvest can be attributed to minimal migration outside Rocky Mountain National Park, as indicated earlier. Habitat Selection Summer Range Estimates of habitat selection were obtained for each of the 4 months that elk occupied their summer ranges. Estimates were summarized by month and time of day. The krumholz type was the most widely utilized habitat throughout the summer (74.7%) (Table 6). Use of the krumholz was greatest during June (94.7%) and lowest during July (60.3%) and August (65.5%). During September, use of the krumholz areas increased to nearly 80% (Table 1). Decreased use of krumholz types and increased use of sprucefir during July and August may be a reaction to warm summer temperatures. �123 Table 5. Elk calves captured and collared in Rocky Mountain National Park, 1981. Collar Number 1 3 6 7 8 9 11 14 16 15 20 67 69 17 22 70 83 84 Date Collared 6-5 6-10 6-10 6-11 6-11 6-16 6-16 6-17 6-18 6-18 6-18 6-18 6-18 6-18 6-18 6-18 6-18 6-23 Sex M F M M ND F M F F F M M M M F M M M Weight (kg) Shoulder Ht. (cm) 19.1 22.7 25.5 22.7 21.8 21.4 21.4 ND 20.9 21.4 30.5 25.0 29.5 23.6 26.4 29.1 17.7 33.6 73 71 75 74 71 72 72 95 84 83 79 76 84 83 84 83 72 83 Girth (cm) 72 72 74 72 64 64 68 82 67 74 77 67 78 65 75 74 63 83 Total Lg (cm) Est. Age (days) 107 107 114 114 111 112 108 137 111 117 124 117 128 114 131 124 96 127 2 7 7-10 7-10 5 3 5 20+ 3 3 14+ 5-7 14+ 7 14 10-14 1 20+ Use of spruce-fir was highest during daylight hours in July (45.3%) and August (68.5%) (Table 7). Alpine meadows comprised only a small portion of total habitat use on the summer range. Use of alpine areas never contributed more than 4% to total use during any 1 month (Table 6). No documented use of the willow-park type by radio-collared elk was recorded during 1980. Although there seemed to be considerable use of the willowpark areas along the Poudre River east of Specimen Mountain during June as documented by visual sightings of elk feeding in these areas during early morning hours. Although the willow-park type is apparently used on summer range, its contribution to total habitat use during summer appears small. Elk apparently used the willow-park type in June primarily as a feeding site. Even though .krumholz sites dominated habitat use, differences in habitat selectivity were apparent between different time of the day (Table 7-10). Use of krumholz areas was dominant throughout the day, spruce-fir was used more during daylight hours (38.8%) than any other time (Table 7). Use of spruce-fir declined and use of krumholz types increased during the evening hours (Table 8). Habitat use during night hours was nearly all krumholz (88.2%) with only little use of spruce-fir (7.1%) and alpine meadows (4.7%) (Table 9). During mornings hours, alpine meadows received higher use than during any other time of day even though it comprised only 13.7% of total use (Table 10). �124 Attractiveness of the krumholz type as.a major summer range habitat may be related to its large structural diversity. This type affords both resting and feeding sites. Within the krUII}holztype,numerous small streams and marshy areas occur to provide ample forage and water. Engelmann spruce, subalpine fir and willows offer cover from severe weather or shade to escape warm summer temperatures. Table 6. Selection (%) of various habitat types by elk during JuneSeptember 1980, Rocky Mountain National Park. September Habitat Type June July August Krumholz 94.7 60.3 65.5 78.6 74.7 Spruce-fir 1.8 37.8 30.7 17.3 21.9 Willow-park 0.0 0.0 0.0 0.0 0.0 Alpine 3.5 1.9 3.8 4.1 3.4 X Table 7. Selection (%) of various habitats by elk during daylight hours for period June-September 1980, Rocky Mountain National Park. Habitat Type June July. August September X Krumholz 96.1 54.7 31.5 62.5 61.2 Spruce-fir 3.9 45.3 68.5 37.5 38.8 Willow-park 0.0 0.0 0.0 0.0 0.0 Alpine 0.0 0.0 0.0 0.0 0.0 Table 8. Selection (%) of various habitats by elk during evening hours for period June~September 1980. Rock.v Mountain National park. Habitat Type June July August September X 100.0 68.0 81.3 75.0 81.1 Spruce-fir 0.0 32.0 18.7 25.0 18.9 Willow-park 0.0 0.0 0.0 0.0 0.0 Alpine 0.0 0.0 0.0 0.0 0.0 Krumholz �125 Table 9. Selection (%) of various habitat types by elk during nighttime hours for period June-September 1980, Rocky Mountain National Park. Habitat Type June July August September X Krumholz 88.9 71.4 92.5 100.0 88.2 Spruce-fir 0.0 28.6 0.0 0.0 7.1 Willow-park 0.0 0.0 0.0 0.0 0.0 11.1 0.0 7.5 0.0 4.7 Alpine Table 10. Selection (%) of various habitat types by elk during early morning hours for period June-September 1980, Rocky Mountain National Park. Habitat Type June July August September X Krumholz 92.9 52.9 88.9 75.0 77 .4 Spruce-fir 0.0 35.7 0.0 0.0 8.9 Willow park 0.0 O.Q 0.0 0.0 0.0 Alpine 7.1 11.4 11.1 25.0 13.7 Winter Range Habitat use on the winter range was dominated by ponderosa-pine shrub type (49.2%) (Table 1). West meadow and coniferous types ranked 2nd and 3rd overall with percentages of 19.7% and 12.4%, respectively. The other 4 types contributed less than 10% to total habitat use on the winter range for all 8 months. No definitive month-to-month trend in habitat use was apparent. It appeared that elk generally selected habitat types relatively independent of months (Table 11). . Considerable differences in habitat selectivity were apparent when data were summarized by time of day (Tables 12-15). Habitat use during daylight hours was almost exclusively restricted to coniferous-pine (35.3%) and ponderosa-pine shrub (52.7%) types (Table 12). This corresponded to a period of resting activity for which these 2 types were used heavily. During evening hours, when feeding activity was high, habitat use shifted primarily to ponderosa-pine shrub (48.7%),wet meadow (16.2%) and willow (11.6%) (Table 13). Use during the night was primarily in the �126 ponderosa-pine shrub and wet meadow types (Table 14). Habitat use during morning houts was similar to that in the evening with ponderosa-pine shrub (43.9%), wet meadow (23.1%), and willow (11.6%) types dominating. In summary, winter habitat use by elk in Rocky Mountain National Park was dominated by the coniferous, ponderosa-pine shrub and wet meadow types. Although monthly trends in use were not readily apparent, daily trends were quite evident with habitat use shifting with daily activity cycles. Table 11. Selection (%) of various habitat types by elk during winter on winter range, Rocky Mountain National Park. Jan. Feb. March April X 8.7 >1 20.7 22.1 10.0 12.4 34.0 55.8 93.2 50.8 34.5 40.0 49.2 16.6 10.9 9.2 0.0 3.0 0.0 15.5 7.4 4.6 1.2 7.6 9.0 2.2 6.4 1.6 0.0 4.2 21.0 23.5 25.0 6.2 4.5 15.7 27.7 26.2 19.7 Willow 3.9 8.7 9.5 7.3 0.0 2.0 9.6 8.3 5.1 Grassland 6.0 0.0 0.0 3.8 0.0 1.6 4.4 0.0 2.0 Habitat Type May Oct. Nov. Dec. Coniferous pine 21.5 3.4 13.0 Ponderosa pineshrub 39.0 46.6 Ponderosa pinegrassland 4.0 Aspen Wet Meadow Table 12. Selection (%) of various habitats by elk on winter range during daylight hours, Rocky Mountain National Park. May Oct. Nov. Dec. Jan. Feb. March April X Coniferous pine 48.6 8.7 43.2 16.7 0.0 51.5 58.3 54.8 35.3 Ponderosa pineshrub 43.1 51.3 37.3 69.0 100.0 42.9 40.8 37.2 52.7 Ponderosa pinegrassland 0.0 40.0 9.6 14.3 0.0 0.0 0.0 8.0 9.0 Aspen 8.3 0.0 9.9 0.0 0.0 5.6 >1 0.0 3.0 Wet Meadow 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Willow 0.0 0.0 0.0 0.0 0.0 0,0 0.0 0.0 0.0 Grassland 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Habitat Type �127 Table 13. Selection (%) of various habitat types by elk on winter range during evening hours, Rocky Mountain National Park. Habitat Type May Oct. Nov. Dec. Jan. Feb. Coniferous pine 0.0 >1 12.5 7.1 1.7 Ponderosa pineshrub 42.9 58.7 32.1 60.7 Ponderosa pinegrassland 23.8 16.7 12.5 2.4 1.6 21.4 Willow Grassland Aspen Wet Meadow March April X 9.0 14.4 0.0 5.6 98.3 24.1 37.5 35.0 48.7 14.3 0.0 0.0 0.0 5.6 9.1 5.3 7.1 0.0 22.2 6.3 0.0 5.6 6.5 5.3 7.1 0.0 28.6 25.9 34.8 16.2 9.5 26.8 33.9 0.0 0.0 7.1 11.3 17.6 13.3 0.0 0.0 0.0 3.6 0.0 9.0 4.7 7.0 3.0 Table 14. Selection (%) of various habitat types by elk on winter range during nighttime hours, Rocky Mountain National Park. Habitat Type May Oct. Nov. Dec. Coniferous pine 0.0 4.8 0.0 3.8 >1 Ponderosa pineshrub 28.6 44.5 37.3 50.5 Ponderosa pinegrassland 0.0 0.0 9.6 Aspen 0.0 0.0 54.0 Wet Meadow Willow Grassland March April X 0.0 0.0 0.0 1.2 86.2 71.4 25.1 30.4 46.7 2.8 0.0 0.0 0.0 0.0 1.5 9.9 10.7 2.9 3.6 3.6 0.0 3.4 45.0 43.2 9.0 10.2 25.0 25.0 61.5 37.5 0.0 5.7 0.0 15.7 0.0 0.0 0.0 8.1 5.4 17.4 0.0 0.0 7.4 0.0 0.0 0.0 0.0 4.3 Jan. Feb. �128 Table 15. Selection (%) of various habitats by elk on winter range during early morning hours, Rocky Mountain National Park. Habitat Type May Oct. Nov. Dec. Jan. Feb. March April X Coniferous pine 4.2 >1.0 1.8 12.3 2.9 29.1 7.1 0.0 7.2 Ponderosa pineshrub 39.6 31.3 23.7 46.9 91.4 42.9 40.5 35.0 43.9 Ponderosa pinegrassland 0.0 11.8 14.3 14.3 .O~O 17.9 0.0 10.4 8.6 Aspen 4.2 6.2 0.0 18.4 5.7 0.0 0.0 0.0 4.3 Wet Meadow 27.5 35.5 37.0 6.1 0.0 4.1 34.5 40.0 23.1 Willow 13.7 14.2 23.2 2.0 0.0 6.1 17.9 15.6 11.6 Grassland 10.8 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1.3 Activity Patterns Trends in activity patterns were apparent across both months and time of day (Figs. 1-5). Grazing activity was highest on the summer range (JuneSeptember 1980) and lowest on the winter range (May 1980, October-April 1980-81) (Fig. 1). Resting activity was opposite, being lowest in summer and highest in winter. These seasonal changes in activities correspond with changes in forage availability and energy and nitrogen supplies (Baker 1980). Periods of high energy and nitrogen supplies corresponded to periods of high grazing activity (Spring and Summer) while low grazing activity was observed during the winter months when range nutrient supplies were lowest. Daily changes in activity were also observed throughout the year (Figs. 2-5). Elk exhibited diurnal resting patterns and crepuscular feeding patterns. During summer and winter, elk remained active grazers during morning and evening hours (Figs. 3 and 5). Time spent grazing averaged 64.3% and 74.4% during the morning and evening, respectively. Although elk remained active during the night, the general activity pattern was one of alternating feeding and resting periods of 1-3 hours in length. Except for months of June-August, daylight hours were a period of relative inactivity (Fig. 2). Resting activity during November, December and January comprised 87%, 95.5% and 83.3% of the daylight hours, respectively. In summary, elk exhibited seasonal, monthly and daily variations in feeding, resting and moving activity patterns. Seasonal and monthly changes seem to be correlated with changes in forage energy and available supply of nitrogen. Daily changes in activity conformed to consistent circadian patterns with major feeding periods in the morning and evening and a resting period during the daylight hours. �I-' N ~ MAY HJN :JUL AUG GRAZING SEP OCT N0V RESTING DEC o JAN FEB MAR APR x MOVING Figure 1. Monthly activity patterns (% time) of elk in Rocky Mountain National Park for period May 1980 to June 1981. �I-' VJ o MAY lUL AUG GRAZING SEP OCT NaV RESTING DEC o JAN FEB MAR APR x MOVING Figure 2. Activity patterns (% time) of elk during day light hours for each month in the period May 1980 "to June 1981, Rocky Mountain National Park. �> -> •••• •••• (.) ct •••• Z I-' W I-' LIJ (.) a:: LIJ Q. SEP GRAZING OCT N0V RESTING DEC o JAN FEB MAR APR x MOVING Figure 3. Activity patterns (% time) of elk during evening hours for each month in the period May 1980 to June 1981, Rocky Mountain National Park. �•.... W N MAY J~N ~UL AUG GRAZING SEP OCT N0V RESTING DEC o JAN FEB MAR APR x MOVING Figure 4. Activity patterns (% time) of elk during nighttime hours for each month in the period May 1980 to June 1981, Rocky Mountain National Park. �•.... w w SEP MAY GRAZING OCT NOV RESTING DEC o JAN FEB MAR APR x MOVING Figure 5. Activity patterns (% time) of elk during early morning hours for each month in the period May 1980 to June 1981, Rocky Mountain National Park. �134 LITERATURE CITED Baker, D. L. 1980. Simulations of the carrying capacity of the Rocky Mountain National Park elk winter range. Colo. Div. Wildl. Game Res. Rep. July Part 2:197-219. Bear, G. D. and R. A. Green. 1980. Elk Population and Ecology Studies. Colo. Div. Wild!. Game Res. Rep. July Part 2:221-313 .. Prepared by: G.>b~ Wildlife Researcher R. Green Graduate Research Assistant �135 APPENDIX A �136 ELK TRAPPING - ESTES PARK VALLEY Summary Duration: December 9, 1980 - January 22, 1981 Trap Sites: Location 1. No. of Clover Traps Beaver Meadows Entrance Station (BME): at jct. of Bear Lake Rd. and Deer Ridge Rd., North side of road in trees 2. Morraine Park (MP): in Morraine Park Campground on Loop A, meadow near site 92 3. 4. 5. 3 5 Morraine Park - South (MP-S): at end of dirt road on south side of Morraine Park 3 Beaver Meadows (BM): approximately 100-400 yds. west of turn-around at west end in timber 5 Little Horseshoe Park (LH): on ridge 100 yds. NE of barn and in park at base of that ridge (approx. 1/4 mile NE of barn) 5 Horseshoe Park (HS): on bench 50 yds. north of road at east end 2 McGregor Ranch (McG): in mouth of Black Canyon, along creek bottom, 100 yds. west of Park Boundry 5 Crocker Ranch (CR): just west of yellow house along edge of meadow (SE base of Mt. Olympus) 6 6. 7. 8. Marking of elk: Blue eartags (maxi A1f1ex livestock plastic tags) with black numerals. Trapping Crew: Rocky Mountain National Park DOW: George Bear, Paul Neil, Gary Pollock, Bruce Gill �137 RMNP: Dave Essex George Wagner Jack Gartner Burt McLaren Lorin Casebeer Charles Logan James Wilson Bob Seibert Jim Protto Larry Van Slyke Ron Maitland Ed Menning Doug Bueler Skip Betts Ricky Nichols Dave Adams CDC: Robert McLean Crocker Ranch DOW George Bear Gary Pollock Carl Leonard Fran Marcoux Mike Babler Frank Rinella Richard Hopper RMNP Charles Logan Ron Mai tland Jack Gartner Elk Trapped-Marked:** Cows Yearling Adult Total RMNP Crocker 11 82 0 93 Total 5 5 11 87 98 26 44 70 Calves Males Females Total 67 2 1 3 Bulls Yearling Adult Total 12 19 31 2 0 2 14 19 33 191 10 201 TOTALS 24 43 ** Cattle were released in the trapping area on the McGregor Ranch thus the traps were not set. �138 Trapping Success: 54% Recaptures: First tagged 1978~79 First tagged 1979~80 Calves collared 1980 spring First tagged 1980~81 winter Blood Samples: 7 28 2 27 70 Mortality: One adult cow, one adult bull, and one trapper (Mike Babler) �139 Elk trapped and marked in the Estes Park Valley December 9, 1980 - January 22, 1981. Ear Tag Number Date Sex Age Trap Location M 6 7 8 9 10 12/9 12/9 12/9 12/9 12/9 12/9 12/9 12/9 12/9 12/9 Calf Calf Calf Calf Adult Adult Adult Adult Calf Calf BME BME BME BME BME MP BM LH LH LH 11 12 13 14 15 16 17 18 19 20 12/9 12/9 12/9 12/10 12/10 12/10 12/10 12/10 12/10 12/10 F Adult Adult Adult Yr1g Adult Calf Calf Calf Calf Calf LH LH LH BME BME BME BME MP-S MP-S MP 12/10 12/10 12/10 12/10 12/10 12/10 12/10 12/10 12/10 12/10 M Yr1g Adult Adult Adult Yr1g Adult Adult Calf Calf Yr1g MP MP MP MP MP BM BM LH LH LH 12/10 12/10 12/10 12/11 12/11 12/11 12/11 12/11 12/11 12/11 F F F F F Adult Calf Calf Calf Adult Calf Adult Calf Adult Adult LH HS HS BME BME ~ME MP MP MP HS 1 2 3 4 5 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 F M F F F M F F F M M M F F F F F F F F F F F M M F M M F F F F Remarks RC 1150 Org #23 - Rt. antler growing down Org #150 - Lf. tag pulled out Lf. antler broken at skull Org #97 4pt. bull RC #149.03 RC #200 RC 11260 ------------------------------------------------------------------------------ �140 Elk trapped and marked in the Estes Park Valley December 9, 1980 - January 22, 1981. (con't) Ear Tag Number Date 41 42 43 44 45 46 47 48 49 50 12/12 12/12 12/12 12/12 12/12 12/12 12/12 12/12 12/12 12/12 5J 52 53 54 55 56 57 58 59 60 12/12 12/12 12/12 12/12 12/12 12/12 12/12 12/12 12/16 12/16 61 62 63 64 65 66 67 68 69 70 12/16 12/16 12/16 12/16 12/16 12/16 12/16 12/17 12/17 12/17 71 72 73 74 75 76 '77 78 79 80 12/17 12/17 12/17 12/17 12/17 12/17 12/17 12/17 12/17 12/18 Sex Age Trap Location F F F F Yr1g Adult Adult Yrlg Adult Adult Calf Yr1g Yr1g Adult BM EM BM BM BM BME BME BME BME MP-S Adult Adult Calf Calf Adult Adult Calf Adult Adult Calf MP-S MP-S MP MP MP LH LH LH BME BME Calf Yr1g Yr1g Adult· Calf Calf Adult Adult Calf Adult MP BM BM BM LH LH HS BME BME. MP Adult Calf Adult Adult Adult Adult Adult Adult Yr1g Adult MP MP MP MP VM VM LH LH LH BME M F F F F F F F F M F F F M F F F F M M H M F F F F F F F F M F M F M F Remarks Org 1147 Org #86 RC 11149.13 Org 11148 Org 11161 Inj. Lf. foreleg living trap RC #320 Org #29 Org #42 Org #145 ------------------------------------------------------------------------------ �141 Elk trapped and marked in the Estes Park Valley December 9, 1980 - January 22, 1981. (con't) Ear Tag Number Date Sex Age Trap Location 81 82 83 84 85 86 87 88 89 90 12/18 12/18 12/18 12/18 12/18 12/19 12/19 12/19 12/19 12/19 F F Adult Calf Adult Calf Adult Adult Adult Adult Adult Yr1g. HP-S HP-S BH LH HS BHE BHE BME MP-S MP-S 91 92 93 94 95 96 97 98 99 100 12/19 12/19 12/19 12/19 12/19 12/19 12/19 1/6 1/6 1/6 Adult Adult Yr1g Calf Adult Adult Adult Adult Calf Calf HP HP HP MP BH BM HS BME BHE MP-S 101 102 103 104 105 106 107 108 109 110 1/6 1/6 1/6 1/6 1/6 1/6 1/6 1/6 1/6 1/7 H H Calf .calf Calf Calf Calf Adult Adult Adult Calf Calf HP-S HP-S HP-S BM BM BM LH LH LH BME Org 1190, cut Rt. hind leg Ye1 1116 (B) (T-B) Org 11159, broken Rt. antler (B) (T-B) (B) (B) 111 112 113 114 115 116 117 118 119 120 1/7 1/7 1/7 1/7 1/7 1/7 1/7 1/7 1/7 1/8 F Calf Calf Adult Adult Adult Adult Adult Adult Adult Adult HP-S :MP-S HP HP VM VM LH LH LH BME (B) (T-B) (B) (T-B) (B) (B) (B) (B) (B) (B) Org 1169 M F. F F F F F F F F M F H M 1<' F F M F H H H F F F F H F F M M F M F F Remarks (B) (T-B) Org 11110 Org 1124 Org 11143 (RC 11220) Org 11123 Org 1189 Very thin Org 1111 tag lying in trap ---------------------------------------------------------------------------~~--- �142 Elk trapped and marked in the Estes Park Valley December 9, 1980 - January 22, 1981. (can't) Ear Tag Number Date Sex Age Trap Location Calf Adult Calf Yrlg Adult Calf Adult Adult Yr1g Adult BME MP MP MP-S MP-S MP-S LH LH LH LH (B) (B) (B) (T-B) (B) (B) (B) (T-B) RC #30 (B) Org #96 (B) Org 1174 Yr1g Yr1g Adult Adult Calf Calf Adult Calf Calf Calf LH HS VME MP MP MP MP MP BM BM (B) (B) (T-B) (B) (B) (B) (B) (B) (B) (B) Calf Yr1g Adult Calf Adult Adult Adult Adult Adult Adult BM BM LH LH LH LH LH BME MP-S MP-S (T-B) (B) (T-B) (B) (B) Org 11142 (B) (B) Org #80 (T-B) (B) (B) Calf Calf Adult Adult Calf Adult AdultAdult Calf Adult HP-S MP MP MP VM VM VM LH LH LH (T-B) (T-B) (B) (B) (B) (T-B) (B) (B) (T-B) (B) Org #115 121 122 123 124 125 126 127 128 129 130 1/8 1/8 1/8 1/8 1/8 1/8 1/8 1/8 1/8 1/8 M 131 132 133 134 135 136 137 138 139 140 1/8 1/8 1/9 1/9 1/9 1/9 1/9 1/9 1/9 1/9 F F F F F 141 142 143 144 145 146 147 148 149 150 1/9 1/9 1/9 1/9 1/9 1/9 1/9 1/13 1/13 1/13 151 152 153 154 155 156 157 158 159 160 1/13 1/13 1/13 1/13 1/13 1/13 1/13 1/13 1/13 1/13 F F M F F F F M F M F M M M F M F F F F F F F F M M F F F F F F F F Remarks --------------------------------------------------------------------------- �143 Elk trapped and marked in the Estes Park Valley December 9, 1980 - January 22, 1981. (con't) Ear Tag Number Date 161 162 163 164 165 166 167 168 169 170 1/l3 1/14 1/14 1/14 1/14 .1/14 1/14 1/14 1/14 1/14 171 172 173 174 175 176 177 178 179 180 1/14 i/14 1/15 1/15 1/15 1/15 1/15 1/15 1/15 1/15 181 182 183 184 185 186 187 188 189 190 1/15 1/15 1/15 1/16 1/16 1/16 1/16 1/16 1/16 1/16 191 192 193 194 195 196 197 198 199 200 201 1/16 1/20 1/20 1/20 1/20 1/20 1/21 1/21 1/22 1/22 1/22 Sex Age Trap Location M Yr1g Adult Adult Calf Adult Adult Calf Adult Adult Adult LH BME BME MP MP MP BM LH LH LH Calf Calf Adult Calf Adult Calf Calf Calf Calf Yr1g HS HS BME MP-S BM BM BM LH LH LH Calf Calf Adult _ Yr1g Calf Yr1g Adult Adult Calf Adult HS HS HS MP MP BM BM BM LH LH Adult Calf . Adult Adult Adult Adult Calf Calf Yr1g Adult Yr1g LH CR CR CR CR CR CR CR CR CR CR F F F F F M F F F F M F M M F· M M F F F F F F F M F M M F F M F F F F M F M F M Remarks (B) (B) Org 1199 Org #98 (T-B) (B) (T) (B) (T-B) (B) Org 115 Org 11132 (T-B) (B) (T-B) (B) (T) (T-B) (B) �144 Recapture of elk marked with blue earraga (1980~81). Date 12/11 12/16 12/17 12/18 12/19 1/6 1/6 1/6 1/8 1/8 1/8 1/9 1/9 1/9 1/9 1/13 1/13 1/13 1/13 1/13 1/14 1/14 1/15 1/15 1/15 1/16 1/16 1/16 1/16 Eartag Number Location 28 33 10 12 59 LH LH HS LH MP 60 62 16 82 1 BME BME LH MP MP-S 60 26 17 34 2 MP BM BM LH LH 99 121 138 69 36 MP-S BM BM LH HS 6 97 121 15 167 MP LH MP-S MP MP 167 144 130 70 MP LH HS HS �July 1981 145 JOB PROGRESS Colorado State of Project W'.)rkPlan No. Job Title: ------ W-126-R-4 No. Elk Investigations Covered: Personnel: Big Game Investigations 3 _----_._-- Nutritional Period REPORT 3 Job No. - Evaluation of Factors Status and Population Influencing Elk Performance July 1, 1980 through June 30, 1981 D. Baker, T. Hobbs, D. Swift, J. Ellis, J. Ritchie L. Stevens ABSTRACT Four manuscripts regarding elk diet selection and nutritional ecology were prepared and submitted to the Journal of Wildlife Management. Results of these investigations advance our knowledge of elk nutrition considerably and suggest fruitful areas of research to further define interrelationships between nutritional quality of ungulate diets and digestive morphology. Detailed comparative digestion studies are planned to more clearly define nutrient utilization and intake of wild ruminants. Preliminary results indicate that elk are more efficient than mule deer in digesting fibrous components of grass hay. ��147 EVALUATION INFLUENCING ELK NUTRITIONAL OF FACTORS STATUS AND POPULATION PERFORMANCE D. L. Baker and N. T. Hobbs P. N. OBJECTIVE 1. To develop and test a system for evaluating to support elk. the" potential of habitats 2. To improve the predictive capability of this system by identifying and and experimentally quantifying nutritional requirements and physiological mechanisms of elk. SEGMENT OBJECTIVES 1. Prepare final publications describing a) elk winter and suwmer diet selection in Rocky Mountain National Park and b) the potential of this ecosystem to support elk. 2. Complete and mule carrying by Rocky 3. Refine the model of carrying capacity of elk winter habitats and test the model with emphasis on improving digestive physiology and nutrient requirement submodels. 4. Prepare detailed cedures relevent 5. Begin initial investigations tive physiology. analysis deer and capacity Mountain of dietary relations between elk, bighorn sheep, assess the effects of those relationships on the of each species in Front Range habitats typified National Park. study plans outlining specific objectives and proto the investigation of elk nutritional physiology. METHODS of elk nitrogen requirements and diges- AND MATERIALS Results pertinent to Segment Objectives 1 and 2 are described in 4 manuscripts which have either been published, accepted for publication, or submitted for publication to the Journal of Wildlife Management. An abstract of each of these manuscripts is presented in Appendix A. Segment Objective 3 was addressed indirectly in each of the 3 previous objectives. A detailed study plan (Segment Objective 4) describing a proposed deer-elk nutritional investigation has been completed and appended to the program narrative of work plan 3, job 3, job title: Evaluation of Factors Influencing Elk Nutritional Status and Population Performance. Results of initial investigations into comparative deer-elk digestive physiology and voluntary intake are discussed (Segment Objective 5). �148 Three elk and 3 mule deer were fed a native grass hay to compare the relative digestive efficiency of these animals. Baled grass hay was chopped in a hammer mill to a uniform length of 5 cm and stored in burlap bags until fed. Chemical composition of this ration is presented in Table 1. All animals were fed this ration ad libitum, 5 weeks preceding the trial. Ten days prior to the trial, all animals were separated into isolation pens where daily food intake could be measured. Following this period animals were weighed, then moved into digestion cages for a 10 day collection of feed, feces, and orts. Weight, sex, and age of each experimental animal are shown in Table 2. Samples of feed, feces and orts were subsampled for each animal and analyzed for dry matter (A.O.A.C. 1965) neutral detergent fiber (NDF) and acid detergent fiber (ADF) (Bailey and Ulyatt 1970). Differences in utilization of nutrients between deer and elk were analyzed with unpaired t-tests. Table 1. Composition of grass hay fed to deer and elk. Composition Percent Dry Matter Ash Crude Protein NDF ADF Lignin Gross Energy kcal/gm Table 2. Species Elk Elk Elk 92.0 14.0 12.0 56.0 34.0 5.3 4.3 Weight, sex, and age of experimental animals. Sex Female Male (Castrate) Male (Castrate) Number Age (yrs) Weight (kg) Pretrial Postrial 75 4 277 270 90 4 308 309 93 4 312 307 ------------------------------------------------------------------------Deer Deer Deer Male (Castrate) Male (Castrate) Female 510 4 63 63 180 189 4 4 59 56 60 56 �149 RESULTS AND DISCUSSION Results of these digestion trials indicate that elk are more efficient than mule deer in utilizing nutrients in grass hay (Table 3.) Elk digestion coefficients of dry matter, energy, NDF, and ADF were significantly higher (p < 0.05) than mule deer. These preliminary results are concordant with theoretical predictions for large and small bodied ruminants (Hoffman 1973, Janis 1976, Short et a1. 1965, Church and Hines 1978). Elk with relatively large rumens and physical barriers for slowing food passage derive a substantial part of their energy from fermentation of fiber. In contrast, deer with smaller rumens and faster turnover rates are less adapted for rumen fermentation of forages high in cellulose. Deer and elk appeared to readily adapt to conditions of confinement. Intake in digestion cages were similiar to those in isolation pens. All animals performed reasonably well given the short period of conditioning to digestion cages. Deer showed greater individual variation in digestion parameters than elk indicating the need for increased numbers of deer in future experiments. Table 3. a Comparison between deer and elk apparent digestion coefficients . Constituent Mule Deer % Dry Matter 57.0b + 2.2 47.0b + 1.4 46.0b 2.0 NDF ADF + Elk % c + 0.5 58.0 + 1.5 c 57.0 + 1.6 62.0 c a Mean + standard error b,c Means having unlike subscripts differ LITERATURE CITED A. O. A. C. 1965. Official methods of analysis. Chemist. Washington, D.C. 957pp. 10th ed. Assoc. Off. Bailey, R. W., and M. J. Ulyatt. 1970. Pasture quality and ruminant nutrition. II. Carbolydrate and lignin composition of detergent extracted residues from pasture grasses. New Zealand J. Agric. Res. 13:591-604. Church, D. C. and W. H. Hines. 1978. Rumino-reticular characteristics of elk. J. Wildl. Manage. 42:654-659. �150 Hoffman, R. R. 1973. The ruminant stomach. East African Mono. in BioI. 2. East African Literature Bureau. 350pp. Janis, C. 1976. The evolutionary strategy of equidae and the origins of rumen and cecal digestion. Evolution 20:757-774. Short, H. L., D. E. Medin, and A. E. Anderson. characteristics of mule deer. J. Mammal. Prepared by ..p.3~· .fl!!.L.""::':"'~!....L.L~.-I-g.=::!_c...\.a.A.=::.!:l-=,..,_._. Bake~ Wildlife Researcher _ 1965. Ruminoreticular 46:196-199. �151 APPENDIX A �152 Hobbs, N. T., D. L. Baker, J. E. Ellis, and D. M. Swift. sition and quality of elk winter diets in Colorado. Manage. 45(1):156-171. 1981. Compo- J. Wildl. ABSTRACT We related the botanical composition and nutritional quality of the diets of 5 tame elk (Cervus elaphus nelsoni) to forage quality, advancing season, and plant communities of upper montane winter range during 1976-78. Graminoids dominated diets (x = 61%) during 1976-77, but declined in amount from 66% of observed bites in November to 44% in February as browse consumption increased. Browse and grass (accounting for >90% of bites each month) contributed equally to diets through winter 1977-78. Browse species contained more crude protein, cell solubles, and lignin, and less cellulose and in vitro digestible dry matter (IVDDM) than grasses. Nutritional quality of browse remained constant with advancing season, whereas grasses decreased in IVDDM and crude protein, and increased in fiber content. Grasses and browse contained less crude protein, and browse less IVDDM, during. 1977-78 than during the previous year. Diet quality was influenced by changes in forage quality; during November-March 1976-77, dietary crude protein declined from 5.8 to 4.9%, and IVDDM declined from 49 to 40%. Over the same period in the following year, dietary crude protein declined from 5.4 to 4.6%; IVDDM from 42 to 37%. During 1976-77, mean overwinter dietary protein ranged from 6.3% in dry grassland to 4.6% in mesic meadow communities, whereas dietary IVDDM ranged from 47% in aspen to 35% in grasslands. During 1977-78, protein was highest (5.6%) in willow and lowest (4.6%) in mesic meadow, and IVDDM was highest (39%) in aspen and lowest (35%) in willow. Despite year-to-year variation in forage quality, elk maintained relatively stable diet quality over time and space by shifting the forage-class mix of diets. Hobbs, N. T., D. L. Baker, J. E. Ellis, D. M. Swift, and R. A. Green. 1981. Energy and nitrogen based estimates of elk winter range carrying capacity. J. Wildl. Manage. (in press). ABSTRACT Winter range carrying capacity for elk was estimated based on range supply of energy and nitrogen during 1976-78 in Rocky Mountain National Park. Elk forage supplies contained 2.45 x 109 kcal of metabolizeable energy during 1976-77 and 1.64 x 109 kcal the following year. Nitrogen supply was 11 x 103 kg and 6.3 x 103 kg during the two years. Based on energy requirements of a 200 kg elk, winter range carrying capacity was 1481 + 158 animals during year 1 and 991 ± 102 year 2. During both years nitrogen based estimates were similar to energy based predictions: 1674 + 165 elk (year 1) and 994 ± 101 (year 2). These estimates compare favorably with independent estimates of population size. Individual habitat types differed in carrying capacity. Willow, wet meadow, and wet shrub meadow could support the most elk. Aspen and mesic meadow was intermediate �153 in carrying capacity. Sagebrush, grassland, and Ponderosa-pine-shrub could carry the fewest animals. Habitat carrying capacities were poorly correlated with estimates of elk diet protein and in vitro digestible dry matter content. Sources of bias in carrying capacity estimates are discussed. Sensitivity analysis showed predictions of the range supplyanimal demand model of nutritional carrying capacity to be strongly influenced by small changes in elk metabolic fecal nitrogen excretion rates. We conclude: 1. Temporal variation in carrying capacity is important in managing harvest of elk populations and in planning future carrying capacity research; 2. Estimates of nutritional carrying capacity are viable habitat evaluation procedures particularly when used for comparative purposes. Baker, D. L., and N. T. Hobbs. summer diets in Colorado. 1982. Composition and quality J. Wildl. Manage. (in press). of elk ABSTRACT Tame, trained elk (Cervus elaphus nelsoni) were used to estimate the botanical composition and nutritional quality of elk diets on alpine and subalpine summer ranges in Rocky Mountain National Park during 1977 and 1978. Graminoids were the most frequently chosen food ~tem in all habitats sampled during both summers; shrubs and forbs contributed lesser, equal amounts. Forage class composition did not change with advancing season. Grasses contained more hemicellulose, cellulose, in vitro digestible dry matter (IVDDM) and less lignin than shrubs and forbs. Forbs contained the hIghest percentage of cell solubles and crude protein. Diet crude portein and IVDDM were similar among habitat types and between years. Elk diets averaged approximately 60% digestibility and 13% crude protein across all plant communities during year 1 and 54% digestibility and 13% crude protein during year 2. Dietary crude protein and IVDDM declined during July to September 1977 from 16 to 10% and 64 to 50%, respectively. The following year dietary crude protein decreased from 15 to 10% and IVDDM from 57 to 50%, during the same time period. Summer ranges appeared to provide energy and protein in excess of maintenance requirements, however our data suggest that, depending on weather conditions, the period of high diet quality can be brief. Hobbs, N. T., D. L. Baker and R. Bruce Gill. 1982. tional ecology of montane ungulates during winter. of Wildlife Management). Comparative nutri(Submitted to Journal ABSTRACT Comparisons of botanical and nutritional characteristics of winter diets of elk (Cervus elaphus nelsoni), mule deer (Odocoileus hemionus hemionus), and Rocky Mountain bighorn sheep (Ovis canadensis canadensis) revealed sharp divergence in food niches. Deer diets contained the most browse and lignin, were intermediate in crude protein, and contained the least �154 in vitro digestible dry matter (IVDDM). Bighorn sheep diets were dominated by forbs and were consistently highest in protein and IVDDM. Cell solubles were higher, and non~lignified cell wall lower in mule deer and bighorn sheep diets compared with elk diets. Elk diets contained the most grass, were intermediate in IVDDM, and were lowest in crude protein. Protein and cell soluble content of diets were correlated with diet selectivity. Ungulates ate less grass and more dicots when the crude protein and cell soluble content of grasses declined. Our findings are discussed in context of current theory on trophic ecology of wild ungulates. We propose that mule deer fit some aspects of this theory poorly and suggest that recent findings on their digestive physiology may explain this inconsistency. � https://cpw.cvlcollections.org/files/original/19c70d041ac4cb5d6fc783031d2f3b0c.pdf cb6ea189874d7d1693c18b75d7b9ee44 PDF Text Text 155 JOB PROGRESS State of REPORT Colorado ----------------------- Big Game Investigations Pro j ec t No. _::W:....-..=1;.;:2:...::6_-.=_R: _ Work Plan No. Job Title: Period Prescribed Covered: Personnel: Job No. 4 ----------------Burning July 1, 1980 T. Hobbs, to Imprcve through R. Spowart, 1 and Enlarge Bighorn Sheep Ranges June 30, 1981 L. Stevens, J. Ritchie, J. Gross ABSTRACT Prescribed burning exerted strong effects on structure and function of grassland and mountain shrub communities and on the composition and quality of winter diets of ungulates feeding in those communities. Mule deer and bighorn sheep consumed diets containing greater numbers of total bites; and quantities of grass, crude protein, and in vitro digestible organic matter (IVDOM) from burned areas compared with controls. The magnitude of these diffcrer.ces tended to be greater in mountain shrub than grassland and was dependent on month that diets were observed. Burning resulted in substantial decrements in the above ground standing crop of herbage drymatter, nitrogen, and IVDOM, but markedly increased the proportion of nutritious forage present in the total herbage on offer. Nitrogen fixation appeared to be suppressed by burning in mountain shrub, but was not affected in grassland. Soil mineral nitrogen increased dramatically in response to burning mountain shrub communities. ��157 PRESCRIBED BURNING TO IMPROVE AND ENLARGE BIGHORN SHEEP RANGES N. T. Hobbs and R. A. Spowart P. N. OBJECTIVES 1. Quantify the effects of burning mountain shrub and grassland communities on nutritional status of mule deer and bighorn sheep during winter. 2. Determine the change in nutritional carrying capacity of mountain shrub and grassland winter range for mule deer and bighorn sheep which is brought about by burning. 3. Examine the effect of fire on food niche relations separation of mule deer and bighorn sheep. 4. Explain changes in responses of forage resources, both quantity quality, in tenns of pro~esses in the nitrogen cycle. and ecological and SEGMENT OBJECTIVES 1. Conduct first year post-burn quality measurements. vegetation 2. Conduct first year post-burn preference measurements. bighorn cover, herbage sheep food habits yield, and and treatment ~ffiTHODSAND MATERIALS Treatment Structure Prescribed burns were conducted during October of 1979 in three 0.3 ha plots in grassland and three .9 ha plots in a mountain shrub community of deer and bighorn sheep winter range south of Wintersteen Park near Rustic, Colorado. Adjacent to each burned plot was a paired unburned control plot of equal size; replicates were paired by apparent homogeneity with burning randomly assigned to one plot in each replicate. Fire Science Personnel from Colorado State University measured pre- and post-fire fuel loadings, and estimated fire intensity during' burning. Investigations Experiment i) 1: Effects on Diet Quality Hypothesis: Nitrogen and in vitro digestible dry matter (IVDDM) are more concentrated in mule deer and bighorn sheep diets chosen from burned plant communities. �158 ii) iii) Rationale: Ungulates in the Rocky Mountain region experience nutritional deprivation during winter (Hobbs et al. 1981, Milchunas et al. 1978, Wallmo et al. 1977). If burning winter range is to substantially improve the performance of these populations, then it must elevate the nutritional status of individual animals by improving the quality of their diets, particularly the concentrations of usable dietary energy and nitrogen. Methods: Dietary nitrogen and IVDDM will be measured on samples of forage choices of 4 tame bighorn sheep and 4 tame mule deer. Diets ,,,,ill be sampled during 5, 6-day grazing trials each winter for two years. Approximate dates for trials will be 3-10 November, 15-22 December, 10-18 January, 18-25 March, 15-22 May. For 2 days preceeding each trial, experimental animals will be allowed to feed in areas to be sampled. These pre-trial periods will allow the animal to become familiar with new surroundings and available forage. No data will be collected during pre-trials. During the subsequent 6 days, pairs of animals (2 sheep or 2 deer) will be released in a randomly chosen replicate and allowed to graze for 50 minutes in each plot (treatment and control) of the replicate. Plots will be fenced with nylon netting strung between steel posts to insure control over the animals location and thus allow equal sampling of all treatment and control plots. Animals will be observed in pairs. All animals will graze once each day. Initial starting points will be the approximate center of a plot; starts in treatment or control plots will be alternated. If tractability permits, animals will be switched between plots of the replicate every 25 minutes; otherwise, they will change plots only once, after 50 minutes of feeding. As each pair of animals feeds, two observers will record (on a tape recorder) the plot number where the animals are feeding, the number of bites of each plant species eaten and a description of the plant parts selected. Simultaneously, a third observer will collect samples of forages consumed, either from the plant being grazed, or, when it is entirely eaten, from adjacent plants of the same species. This observer will pay particular attention to including in the sample the plant parts taken by the animal. Forage samples will be collected for all species contributing 2% or more of total bites observed in each plot. Fifty grams of plant material will be composited into plastic bags and frozen. Bite weights will be estimated by hand-plucking an additional 25 sample "bites" of these species, drying the samples of 100 C for 24 hr, weighing the sample to the nearest ± 0.01 g and dividing the total weight by 25. Forage samples will be analyzed for gross energy, ash, dry matter and crude protein (Kjeldahl N x 6.25) according to procedures described by A.O.A.C. (1965). In vitro dry matter digestibility will be determined by techniques described by Tilley and Terry (1963) and Pearson (1970). Rumen fluid for in vitro determinations will be collected from a holstein cow fed native grass hay. �159 Botanical composition of diets will be calculated as percentages of observed bites. Nitrogen and IVDDM content of diets will be calculated as the sum of forage analysis values times weighted diet percentages. Diet percentages will be weighted by multiplying bite frequency times bite weight and normalizing to sum to 100 (Hobbs et al. 1979). iv) Analysis: Treatments will be allocated to experimental units according to the following split-split plot design, and differences determined with analysis of variance. Source Whole Plot Split Plot Degrees of Freedom Habitat Type (H) Replications (R) RH I Treatment RT HT 1 I I I 2 2 (T) mT Split Split Plot Animal Species (A) Individual Animals (I) AA TA ~ RHT ART RAHT Error Total 19 1 3 1 1 I I I I Not Retrievable Differences in diet quality will be determined with analysis of variance according to the above design. Main effects examined will be animal species, habitat types, and treatments. In addition, deer and sheep dietary IVDDM and nitrogen content will be regressed against measures of fire intensity (rate of spread, fuel losses, flame height). Equations will be developed for each vegetation type, if these models are not different, then data will be pooled to estimate a single best-fit equation. If significant, this model will provide substantial insight into the fire prescription necessary to achieve desired effects on ungulate nutrition. Experiment 2. Simulation Modeling of Animal Condition. i) Hypothesis: Differences in diet quality between burned and unburned areas will be sufficiently great to cause biologically meaningful differences in animal condition during the winter period. ii) Rationale: Statistically significant differences in diet quality may or may not be large enough to result in biologically significant differences in animal condition. That is, although measurements may �160 be repeatable, these precise differences may be so small as to be of no consequence to the animal. The change in animal condition incurred by dietary regimes of different quality must be described to make diet quality data directly meaningful in terms of benefits to wintering ungulates. iii) Methods: A generalized ruminant energy and nitrogen balance model will be used to estimate overwinter changes in body composition of deer and bighorns grazing on burned and unburned areas. This model has seen wide application in investigations of trophic ecology of wild and domestic ungulates (Hobbs 1979, Ellis and Parton 1978). Input data for the model will be dietary nitrogen and digestible dry matter content; for examples of data necessary to adopt the model to species specific attributes (see Hobbs 1979, Table 4). iv) Analy~is: Model output which results from significant differences in input variables will be assumed to be repeatable. The model will be used to estimate the magnitude of differences in diet quality necessary to achieve various specific levels of animal condition. These estimates, compared with predictable animal condition resulting from observed diets can be used to evaluate the efficiency of burning in improving ungulate nutritional status. Experiment 3: Effects on Nutrient Distributions i) Hypothesis: The biomass distribution of nutrient concentrations will be skewed to the right on burned plots. That is, there will be a larger proportion of the biomass in burned areas which contains intermediate and high concentrations of nutrients than in unburned areas. ii) Rationale: The fraction of herbage biomass which can be used as food by ungulate herbivores has not been quantitatively described in any ecosystem. Qualitatively, it is known that within the total potential forage available there is a portion of very low quality foods (woody stems, mature seedheads) a fraction of foods of intermediate quality (mature leaves, forbs) and a small amount of highly nutritious forage (terminal buds, new leaves, fruits). The relative size of these fractions depends on a variety of phenological, structural, and successional attributes of the plant community as a whole. This relationship describing the amount of food relative to its quality is an important emergent property of ecosystems, a characteristic which determines habitat suitability for herbivores; For example, small ~odied African ungulates who require relatively small amounts of highly nutritious forage can occupy woodlands which contain an enormous amount of herbage of very low quality, but also, a small, but obtainable portion of buds and fruits with very high concentrations of nutrients (Jarman 1974). Larger bodied herbivores would starve in these areas because their total intake requirements �161 could not be met by the thinly dispersed food items present there. However, grasslands, which contain large amounts of herbage of intermediate quality and relatively small amounts of food at the high and low and of the nutrient concentration gradient, provide sufficient forage of adequate quality to these animals. How can these relationships be quantified? In the continuous case, where the number of nutrient categories characterizing forage is increased many times, this relationship can be described by a function which relates the biomass of forage to its nutrient concentration (Figure 1). This function has not been described. It has great potential to assist in predicting supportable animal density because it allows merger of constraints on nutritional status (i.e. an animal must obtain a diet exceeding 6% protein) with a range supply and animal requirement model of nutritional carrying capacity. This is possible because, by describing the distribution of nutrients, the amounts of food containing different levels of nutrients are known. Carrying capacity can then be estimated based on a much more resolute prediction of range supply. iii) Methods and Analysis: Current production of forage will be estimated at the end of the growing season using clip and weigh methods. Herbaceous vegetation in seventy-five randomly located ~ m2 plots will be clipped to ground level in each treatment and control replicate, for a total sample of 900 plots. Shrub material will be harvested from 20, 2 m2 circular plots. Dates for herbaceous biomass sampling will be August 20-30. Harvested material will be separated into the following 1. Graminoid leaves, mature. 2. Graminoid culms and infloresences. 3. Graminoid tillers and new leaves. 4. Shrub current growth, last 2 cm. 5. Shrub current growth, all but last 2 cm. 6. Shrub previous year's growth. 7. Selected 8. Other forbs. categories: forbs by species. These categories, rather than taxanomic groups, will be used because: 1) there is probably greater heterogeneity among different plant parts of the same species than among whole plants of different species (Arnold 1964, Bailey 1967); 2) the unit of dietselection for ungulate herbivores is probably more closely related to plant parts (Bell 1971, Arnold 1967) and to groups of �162 plants (Jarman 1974, Kossak 1976) than to taxonomically distinct individual plant species; 3) by pooling forages into larger categories than species, precision of biomass estimation is improved; and,4) this sampling scheme is well suited to description of the biomass distribution of forage nutrient concentrations (See Experiment 4). Nitrogen concentration, in vitro digestible dry matter, and gross energy will be estimated for each forage category described above according to procedures detailed in Experiment 1. A best fit equation will the dependence of forage bility and crude protein predict how much forage of protein on digestible Experiment 4: be developed by regression to describe biomass on forage dry matter digesticontent. In short these functions will is present (y value) with a given amount dry matter (X value). Effects on Ecological Separation i) Hypothesis: Dietary overlap between mule deer and bighorn will be greater on burned than unburned plots. sheep ii) Rationale: Bighorn sheep are thought to be best adapted to inhabiting climax plant communities, while deer are viewed as being better able to exploit early and intermediate stages of succession (Klein 1965, Geist 1971). If these generalizations are true then niche separation of bighorns and deer may be primarily dependent on divergence along successional gradients. Given no history of competition for food resources in burned communities, mechanisms of food niche separation in these areas should be less well established than in areas of traditional sympatry. As a result, dietary overlap should be greater on burned sites. If this is the case, it has important implications for use of fire for improving bighorn ranges. Although burned communities might be better habitat for bighorns in isolation, this improvement may be diluted or extinguished by competitive interactions in multispecies communities. iii) Methods: Botanical composition of mule deer and bighorn sheep diets on burned and unburned plots will be determined in Experiment 1. These data will be used to compute Horns index of similarity (Horn 1966) for comparisons of food niche overlap. iv) Analysis: procedures Indexes of similarity will be compared using analysis outlined in Garrat and Steinhorst (1976). �163 Experiment 5: Effects on Soil Water and Nitrogen Status i) Hypothesis: Water potential and total nitrogen will be greater in soil of control plots. Available nitrogen will be greater in soil of treatment plots. ii) Rationale: This experiment will provide correlative evidence to explain changes in plant production and quality, and this carrying capacity of ungulate winter range. iii) Methods: Predawn water potential will be measured for C3 and C4 grasses in all plots using a Scholander pressure bomb fitted with a dissecting scope. Ten measurements will be made on each plot. Four composite samples containing 5 individual soil cores taken at 0-5 cm depth will be collected from each plot during mid-June. Samples will be sieved through a 2 mm screen and analyzed for dry matter, organic matter, and Kjeldahl nitrogen by procedures outlined in A.O.A.C. Sample ammonium content will be determined by a calorometric ammonium reaction with salicylate nitroprusside at pH 13; nitrate + nitrate be determined by a cadmium reduction using a continuously copperized reduction column. All analyses will be conducted by Bob Woodmansee's laboratory at the Natural Resource Ecology Laboratory at Colorado State University. Differences in soil water, total nitrogen, and extractible nitrogen between treatment and control and between habitat types will be examined with analysis of variance according to the following design: Source Habitat (H) Replications (R) RH Treatment (T) RT HT RHT Error Experiment 6: Degree of Freedom 1 2 2 1 I 1 1 Total 10 Not retrievable Effects on Nitrogen Mineralization i) Hypothesis: Rates of transformation of organic nitrogen to inorganic forms will be more rapid on burned plots compared with controls. ii) Rationale: It is not known whether the increase in plant production and quality, is directly attributable to increased nitrogen availability, results from deposition of mineral nitrogen in ash,or increased mineralization of organic matter in the soil. Answering this question will enhance the capability to predict outcomes of prescribed burning. �164 CJ) CJ) <l: ~ o en HERBAGE NUTRIENT CONCENTRATION Figure 1. Hypothetical distribution of nutrients in herbage biomass. Point X represents a critical nutrient concentration for maintenance or production, the shaded area is a theoretical amount of food available to meet that cQncentration requirement. �165 iii) Methods: Buried bag techniques will be used to estimate rates of nitrogen mineralization. Twenty soil cores (0-5 cm) will be taken from each plot in early June. Half of the soil from each core will be sieved through a 2 mm screen and analyzed for organic matter, ammonium and nitrate + nitrate according to procedures outlined in Experiment 6. The remainder will be enclosed in labeled polyethelene bags and returned to the soil at the site of the original core. Locations of buried bags will be clearly marked. Six weeks later these buried samples will be retrieved, sieved and analyzed for ammonium and nitrate + nitrate. The increase in inorganic forms of N divided by the elapsed time (6 weeks) will yield indexes of rate of ammonification and nitrification. iv) Analysis: Statistical in Experiment 5. Experiment 7: analysis will follow the design outlined Effects on Nitrogen Fixation i) Hypothesis: Rates of biological fixation of nitrogen in soils of burned plots will exceed fixation rates in soils of controls. ii) Rationale: As has been shown, burning can cause large losses of nutrients, particularly nitrogen, from ecosystems. It may be that these losses are recouped through stimulatory effects of fire on nitrogen additions, primarily through increased nitrogen fixation by plant-microorganism symbiants and free-living bacteria. If additions as well as losses result from burning, the prescribed burns can be used with less danger of nutrient depeletion of burned sites. iii) Methods: Acetylene reduction (Hardy et al. 1973) will be used to estimate rates of nitrogen fixation by free-living bacteria and plant microorganism associations in soil of burned and unburned plots. Soil sampling procedures will follow methods outlined in Experiment 6 with the following modifications. To preserve soil moisture, all samples will be collected in double, sealed plastic bags with a moist paper towel between the inner and outer bag. Analysis will be performed by Don Klein in the Microbiology Department at Colorado State University. iv) Analysis: Experimental in Experiment 5. design and analysis will follow procedures �166 RESULTS AND DISCUSSION Prescribed Burning Effects on Diet Quality Effects of prescribed burning on nutritional quality of diets of bighorn sheep and mule deer were dependent on the month diets were observed, species of animal feeding, and the type of plant communities where animals fed (Table 1). Although formal statistical treatment of these data await completion of laboratory analysis of forage samples from the January and March grazing trials, preliminary examination of means and their standard errors suggests several important differences. Prescribed burning resulted in substantial increments in the IVDOM content of mule deer diets during both November and March in grassland and mountain shrub communities (Table 1). Table 1. Concentration of IVDOM (%) in diets of mule deer and bighorn sheep feeding in burned and unburned mountain shrub and grassland communities during winter 1980-81. Site Mule deer November March X SE X SE Bighorn November X SE Grassland Burn Control 48 31 4 3 57 34 4 6 53 31 Mountain Shrub Burn Control 54 33 4 1 60 31 2 3 45 40 sheeE March X SE 1 1 48 49 1 1 5 1 52 42 1 2 This treatment effect was particularly large on burned mountain shrub plots where mule deer selected diets which were almost twice as digestible as diets selected from controls. Effects of prescribed burning on IVDOM content of bighorn sheep diets were less marked. Apparently, there were no differences in digestibility of sheep diets between burned and unburned plots in the mountain shrub community during November and in grassland during March. Effects of treatment on diet IVDOM, when they were observed, tended to be smaller for bighorn sheep than for mule deer. There was no consistent difference in diet IVDOM between grassland and mountain shrub for deer or sheep during either month. Diet digestibility remained constant or increased between November and March for both deer and sheep on all plots. �167 Diets of bighorn sheep in both grassland and mountain shrub contained consistently higher concentrations of crude protein on burned plots than diets from controls (Table 2). However, this difference did not appear to be significant in March. Table 2. Concentration of crude protein (percent of organic matter) diets of mule deer and bighorn sheep feeding in burned and unburned mountain shrub and grassland communities during winter, 1980-81. Nov. Mule deer Jan. Mar. x SE x SE x SE Grassland Burn Control 11.1 1.6 0.6 15.7 0.6 2.7 22.8 7.5 7.5 0.1 0.9 Mt. shrub Burn Control 14.0 1.6 19.4 16.1 5.7 9.6 7.2 0.4 0.3 1.5 5.4 0.2 0.3 4.1 Site 10.4 Bighorn sheep Jan. Nov. x in Mar. SE x SE x 0.5 0.9 5.3 0.6 4.4 0.2 11.3 0.1 0.8 0.6 14.2 0.3 8.7 2.1 0.4 14.8 7.5 2.1 1.4 7.3 SE Mule deer diets selected from burned plots in mountain shrub contained almost 3 times more crude protein than diets from controls. Treatment effects on crude protein content of deer diets were less marked in grassland than mountain shrub during November and January, but were similar to effects in shrub communities during March. With few exceptions, mule deer selected diets higher in crude protein than diets chosen by bighorn sheep. There was no consistent difference in crude protein of ungulate between mountain shrub and grassland communities. diets Crude protein content of mule deer and bighorn sheep diets tended to be greater in January than November. This seemingly anomalous result can be exp1ained,by variation in weather. During November, 12 inches of snow covered the study area. This snowfall appeared to hinder deer and and sheep in selecting many low growing forages, particularly green grass. Unusually warm temperatures during December and January were associated with initiation of growth of tillers of Poa £Iatensis, Agropyron spp. and Bromus tectorum. Moreover, there was no snow present on the study area in January to prevent animals from finding these green, high quality foods. Although statistical analysis are incomplete, these preliminary results suggest that mule deer and bighorn sheep consume nutritionally superior diets from burned mountain shrub and grassland communities during winter one year following treatment. �168 Prescribed Burning Effects on Total Bites and Diet Botanical Composition Effects on Total Bites Total observed bites for mule deer and bighorn sheep were greater on burned plots compared with controls (P < .05, Tables 3, 4, 5). There was no habitat effect on total bites taken by mule deer (P = .42) but bighorn sheep ate more bites in grassland than in mountain shrub (P = .04). Table 3. Probability values for main effects and 2~ay interactions of winter diets of bighorn sheep a in mountain shrub and grassland habitats in Colorado. P Values % Forbs AOV category % Grass % Browse Main effectb Month Treatment Habitat .1085 .0076 .5123 .0023 .0031 .8647 .1559 .0263 .3989 .00009 .0004 .0410 2-way interactions Month x Habitat Month x Treatment Habitat x Treatment .7259 .0407 .0884 .6338 .0318 .0111 .8598 .1416 .2765 .0274 .0076 .7676 a b n = Total bites '4. Main effects averaged over all interactions (Month = Nov. vs. Jan. vs. March vs. May; Treatment = burned vs. unburned, Habitat = grassland vs. shrub). �169 Table 4. Probability values for main effects and 2-way interactions of winter diets of mule deera in mountain shrub and grassland habitats in Colorado. P values AOV Category % Grass % Browse Main effectsb Month Treatment Habitat .3907 .0016 .6068 .0004 .0020 .5854 .8 x 10 .8893 .4499 2-way Interactions Month x Habitat Month x Treatment Habitat x Treatment .9728 .0004 .2648 .88/+1 .00004 .7310 .1799 .0076 .6972 a n % Forbs Total Bites -8 1 x 10-9 .0033 .4249 .0299 .0010 .5896 = 4. b Main effects averaged over all interactions (Month = Nov. vs. Jan. vs. Mar. vs. May; Treatment = burned vs. unburned, Habitat = grassland vs. shrub) . Total bites taken by both species were dependent on the month animals fed (month effect P < .0009). More bites were consumed during January than November. This difference in eating rate may have been caused by effects of snow described previously. Both mule deer and bighorn sheep ate more bites during May. This increase in eating rate appeared to be caused by green up of vegetation on burned and unburned plots. We observed month by habitat and month by treatment interactions for total bites eaten by both species (P < .05, Tables 3, 4). During all months, more bites were taken by mule deer and bighorn sheep on burned plots than controls, but the magnitude of this difference was variable. In contrast, we found that the magnitude of treatment effect on total bites was independent of habitat (habitat x treatment interaction P > .57). Thus, the effect of burning on eating rates was similar in grassland compared to mountain shrub but was variable with month. This temporal dependence of treatment effects is attributable to monthly differences in snow cover and plant phenology. �a b Table 5. Mean total bites of mule deer and bighorn sheep winter diets in mountain shrub and grassland habitats in Colorado. Site Nov. X SE Mule deer Jan. Mar. X SE X SE May X SE Nov. X SE Bighorn shee12 Jan. Mar. SE X X SE May X SE ~ -...J 0 Grassland Burn Control 441 335 38 50 787 231 106 42 370 168 48 26 546 402 52 61 725 486 55 51 104 266 .97 41 346 76 22 26 1,112 899 84 60 Mountain Shrub Burn Control 507 226 62 19 660 183 80 23 379 127 52 19 526 321 52 70 759 278 86 30 804 207 110 33 506 236 55 56 634 487 84 93 a n = 4. b Grazlng . . = 45 mln. ~ tlme �171 Effects on Diet Botanical Composition Diet botanical composition of bighorn sheep and mule deer was dependent on treatment and month but not on habitat type (Tables 3 and 4). Diets of mule deer and bighorn sheep contained a larger proportion of grass on burned plots than diets selected on control plots (P = .008, Tables 4 and 6) and larger percentages of browse on control plots compared with burn plots (P = .002, Tables 4 and 7). Bighorn sheep consumed a higher percentage of forbs on control plots than on burned plots (P = .03, Tables, 3 and 8). There was no treatment effect on the percentages of forbs in mule deer diets (P = .89, Tables 4 and 8). There were no significant habitat effects on diet botanical composition for either species (P > .38). The proportion of grass in mule deer and bighorn sheep diets did not change during winter (month effect P > .10). However, both species consumed the greatest percentage of browse during March, and substantially decreased browse intake during May. There was a significant month effect on the percentage of forbs in mule deer diets, but the percentage of forbs in bighorn sheep diets remained constant over winter. a Table 6. Mean percentage grass of mule deer and bighorn sheep winter diets in mountain shrub and grassland habitats in Colorado. Nov. Site X SE Grassland Burn Control 52 30 Mt. Shrub Burn Control 57 21 a n 4. Mule deer Jan. Mar. Nov. Bighorn sheeE Mar. Jan. May X SE X SE X SE X SE May SE - X SE· X SE 7 7 76 25 5 6 72 19 4 4 50 42 4 4 63 61 5 8 75 44 7 7 52 44 5 5 67 50 4 6 7 6 83 10 4 4 76 11 5 4 54 38 6 7 86 57 4 7 84 36 5 8 82 28 4 6 66 49 5 7 X �172 Table 7. Meana percentage browse of mule deer and bighorn sheep winter diets in mountain shrub and grassland habitats in Colorado. Site Mule deer Nov. Jan. Mar. X SE X SE X SE May X SE Grassland Burn Control 28 41 3 7 7 57 21 69 4 4 11 9 3 9 6 3 6 1 3 9 2 3 18 26 4 4 2 1 1 1 Mt. Shrub Burn Control 27 49 7 6 9 58 3 7 15 71 3 5 18 12 6 3 2 14 1 3 2 17 1 5 3 31 1 6 5 4 2 2 a 3 2 Nov. X SE Bighorn sheeE Mar. May Jan. X SE X SE X SE n = 4. a Table 8. Mean percentage forbs of mule deer and bighorn sheep winter diets in mountain shrub and grassland habitats in Colorado. Site Nov. X SE Mule deer Mar. Jan. X SE X SE May X SE Nov. X SE Bighorn sheep Mar. Jan. May X SE X SE X SE Grassland Burn Control 19 27 5 6 16 17 3 3 7 11 2 4 39 50 4 28 32 6 8 23 45 6 6 31 29 4 5 31 48 4 5 Mt. Shrub Burn Control 16 16 3 5 8 15 2 4 9 18 2 5 28 49 5 7 10 28 3 6 14 45 4 7 14 39 3 5 29 47 4 6 a n = 4. 3 �173 Effects of month on diet botanical composition were independent of habitats where diets were observed (month x habitat interaction P > .17, Tables3 and 4). Thus, the observed monthly changes in diet composition were proportionally equal in grassland compared to mountain shrub. Magnitude of treatment effects on ungulate diet composition depended on month (month x treatment interaction P > .05, Tables 3 and 4). However, during all months, both mule deer and bighorn sheep consumed more grass and less browse on burned plots comp~red with controls. Moreover, mule deer diets selected on burned plots contained a smaller contribution of forbs than controls. It follows that ungulate species increased grass consumption at the expense of dicotyledon consumption in response to burning. However, the increase in grass and corresponding reduction in browse in bighorn sheep diets which was attributable to burning were relatively greater on shrub plots than grassland plots (habitat x treatment interacton P < .05). Thus, burning exerted stronger effects on bighorn diets. in mountain shrub than grassland. In contrast, the magnitude of treatment effect on composition of mule deer diets was independent of habitat type (habitat x treatment interaction P > .26). Prescribed Burning Effects on Ecological Separation Estimates of diet overlap are frequently used to predict the potential for interspecific competition for food. To quantify effects of prescribed burning on diet botanical composition and diet overlap, diets of both species from burned plots were compared to diets from control plots for both habitat types across all months. The most striking change in ungulate diet botanical composition was the increase in percentage of grass consumed on burned plots compared to control plots. That increase in percentage of grass was relatively greater in the diets of mule deer than in the diets of bighorn sheep. Consequently, diets of both species contained similar proportions of grass consumed from the same grass species on burned plots. On controls, sheep tended to eat more grass and less browse than deer. Thus, prescribed burning substantially increased the similarity of botanical composition of mule deer and bighorn sheep diets. Assessment of the importance of this change for competition between the 2 species awaits further analysis. Effects of Prescribed Burning on Herbage Nutrient Distributions Prescribed burning substantially reduced the total amount of herbage dry matter, nitrogen, and IVDOM present at the end of the growing season in grassland and mountain shrub communities the year following treatment (Table 9). These reductions, however, were accompanied by large increase in the density of Nand IVDOM in the standing crop on burned plots. More than 62% of the herbage drymatter on burned plots in both mountain shrub and grassland communities exceeded 40% IVDOM (F~s. 2 and 3). On control plots, only 20% of the standing crop of herbage exceeded 40% IVDOM. This effect was even greater for nitrogen density; in burned mountain shrub �174 a Table 9. Standing crops (g/m2) herbage drymatter, nitrogen and IVDOM on burned and unburned plots in mountain shrub and grassland communities during August 1980. Herbage drymatter Site IvnOM Nitrogen X SE X SE X SE Grassland Burn Control 37 197 5 25 0.4 1.8 0.04 0.4 17 38 1 11 Mt. Shrub Burn Control 19 419 3 60 0.4 3.5 0.1 0.4 8 10 2 6 aIn vitro digestible organic matter. 71% of the herbage standing crop contained nitrogen concentrations greater than 1.2% N (7.5% crude protein, Fig. 4). Similar increments in nitrogen density were observed in herbage in burned grassland (Fig. 5). Consequently, although prescribed burning reduced the total amount of herbage present in grassland and mountain shrub communities the year following treatment, the nutritious portion of the standing crop was increased substantially. Statistical analysis of these data is in progress. Effects of Prescribed Burning on Nitrogen Fixation and Soil Mineral Nitrogen Prescribed burning resulted in decreased nitrogen in the mountain shrub community rates of fixation of atmospheric (P = .06, Table 10). There was Table 10. Nitrogen fixation rates (p moles/g OM/hr) on burned and control plots in mountain shrub and grassland communities during July 1980. Site X SE Grassland Burn Control 229 311 102 37 Mountain Shrub Burn Control 164 283 26 17 �60 UNBURNED SHRU B T 50 T 40 T 30 - 1 20 o, 10 0:: o o U <.!) T I o ..•. ~ I I 10 20 30 40 50 60 Z o z « l- I-' -....J VI (/) LL. o 50 BURNED SHRUB ~ 40 30 20 10 o o 10 20 30 40 50 60 70 80 0/0 IVDOM IN PLANT TISSUE Figure 2. Distribution of IVDOM in herbage standing crops on burned shrub plots during August 1980. Vertical lines = 2 SE. and unburned mountain �70 UNBURNED GRASSLAND 60 50 40 30 a.. ~ 20 T 1 10 (.) (!) ..•.. .•. 0 Z 0 o 10 20 or 50 40 30 60 70 •.... -....J 0\ Z « ~ 50 BURNED GRASSLAND LL o 40 o~ 30 20 o T T 10 1 1 T 1 T J. - o 10 20 30 % 40 50 i - 60 70 I 80 IVDOM IN PLANT TISSUE Figure 3. Distribution of IVDOM in herbage standing crops on burned and unburned grassland plots during August 1980. 90 �50 UNBURNED SHRU B 40 T . 1 30 20 a.. o 10 T T a:: I (.) 0 0 (!) Z 0.3 0.6 0.9 .•. _T ..L 1.5 1.2 T I 1.8 I ..L I I 2.4 2.1 o z ~ '"'-l '"'-l ~ C/) u, 40 BURNED SHRU o e;!!. 30 .•. .J. 20 T 10 o J. T .. 1 o 0.3 0.6 0.9 1.2 I , 1.5 T .•. 1 1.8 2.1 2.4 2.7 3.0 , -,I 3.3 3.6 0/0 NITROGEN IN PLANT TISSUE Figure 4. Distribution of nitrogen in herbage standing crops on burned and unburned mountain shrub plots during August 1980. �50 UNBURNED GRASSLAND T 1 40 30 20 o 10 U 0 Q. 0::: T - T T .1 0 (.!) T l. ----------- 0.3 ---- 0.6 1.2 0.9 1.8 1.5 I 2.1 2.4 Z o z ~ ~ en " 00 LL 50 BURNED GRASSLAND o o~ 40 30 20 T 1 10 I o I o 0.3 0.6 % 0.9 1.2 1.5 T ~T .1 1.8 2.1 OF NITROGEN IN PLANT TISSUE Figure 5. Distribution of nitrogen in herbage grassland plots during August 1980. standing crops on burned and unburned 2.4 �179 no difference in nitrogen fixation rates between burned and control grasslands (P = .56). This differential effect of treatment appeared to be related to soil mineral N concentration. Nitrogen contained in ammonium and nitrate and nitrite was more concentrated in soils of burned shrub plots than in soils of controls (P < .001, Table 11). There was no treatment effect on soil mineral N in grassland (P > .05, Table 11). Nitrogen fixation by free-living bacteria and plant symbionts is suppressed when large quantities of N available in mineral form are present. Table 11. Mineral nitrogen content soil from burned and unburned (~g/g dm as NH4 + grassland and mountain and N0 + N02) of 3 shrub communities during June 1980. NH4+ Site Grassland Burn Control Mountain shrub Burn Control NOZ + NO) X SE X SE 6.6 5.8 1.5 0.6 6.2 5.1 0.9 0.3 13.0 4.9 2.4 0.4 7.9 5.6 0.4 0.7 �180 LITERATURE CITED A.O.A.C. 1965. Official methods of analysis. 10th ed. Association Official Agricultural Chemists, Washington, D.C. 957pp. of Arnold, G. W. 1964. Factors within plant associations affecting the behavior and performance of grazing animals. In D. J. Crisp, ed. Grazing in terrestrial and marine environmentS: Blackwell Scientific. Oxford. Bailey, J. A. 1967. Sampling deer browse for crude protein. Manage. 31:437-442. Bell, R.H.V. 1971. 255:86-93. A grazing ecosystem in the Serengeti. J. Wildl. Sci. Am. Ellis, J. E., and W. J. Parton. 1978. The impacts of strip mine relationship: A simulation study. Final report to the Western Energy and Land Use Team, Office of Biological Services, U.S. Fish and Wildl. Service. Ft. Collins, CO 265pp. Garrat, M. W., and R. K. Steinhorst. 1976. Morisita's, Horn's and related measures 96: 245-251. Testing for significance of of overlap. Am. MidI. Nat. Geist, V. 1971. Mountain sheep: A study in behavior Univ. Chicago Press, Chicago. 383pp. and evolution. Hardy, R. F., R. C. Burns, and R. D. Holstein. 1973. Application of the acetylene reduction technique for measurements of N2 fixation. Soil. BioI. Biochem. 5:47-81. Hobbs, N. T. 1979. Winter diet quality and nutritional status of elk in the upper montane zone. Colorado Ph.D. Dissertation. Colorado State Univ.,Ft. Collins. --- , D. L. Baker, J. E. Ellis, and D. M. Swift. 1979. Composition and quality of elk diets during winter and summer: A preliminary analysis. Pp. 54-62 in M. S. Boyce and L. D. Hayden-Wing, eds. North American elk:ecology, behavior, and management. Univ. Wyo., Laramie. 294pp. and winter diets in Colorado. 1981. Composition and quality of elk J. Wild1. Hanage. 45:156-171/ Horn, H. S. 1966. Measurement of niche overlap in comparative ecological studies. Am. Nat. 100:419-424. Jarman, P. J., and M. V. Jarman. 1974. The social organization of antelope in relation to their ecology. Behavior 48(3):215-267. �181 Klein, D. R. 1965. Ecology of deer range in Alaska. 35(3) :259-284. Eco1. Monogr. Kossak, S. 1976. The complex character of the food preference of Cervidae and phytocenosis structure. Acta Therio1ogica 21, 27: 359-373. Mi1chunas, D. G., M. 1. Dyer, O. C. Wa1lmo, and D. E. JohnsOn. 1978. In vivo/in vitro relationships of Colorado mule deer forages. Colo. Div. Wild1. Spec. Rep. No. 43. 44pp. Pearson, H. A. 1970. Digestibility trials: in vitro techniques. Pp. 85-92 in H. A. Paulsen, Jr., and E. H. Reid (co-chairmen) Range and wildlife habitat evaluation: a research symposium. USDA For. Servo Misc. Publ. No. 1147. 220pp. Tilley, J.M.A., and R. A. Terry. 1963. A two-stage technique for the in vitro digestion of forage crops. J. Brit. Grassl. Soc. 18:104-111. Wallmo, o. C., L. H. Carpenter, W. L. Regelin, R. B. Gill, and D. L.· Baker. 1977. Evaluation of deer habitat on a nutritional basis. J. Range Manage. 30(2):122-127 .. v-- tUL Prepared by __ ._\ _ N. T. Hobbs Wildlife Researcher and R. A. Spowar Graduate Research Assistant ��July 1981 183 JOB PROGRESS State of Project C~o~l~o~r~a~d~o No. W-126-R-4 Work Plan No. Job Title: Pronghorn Period Covered: Personnel: 5 REPORT _ Big Game Investigations Job No. Investigations 1 - Pronghorn Population Study July 1, 1980 through June _30, 1981 Gordon East, Bill Knight, Paul Neil, Barbara Hilton-Calm, Lisa Wolfe, Nanci Laurenson, Mike Miller, Lyn Stevens, Bert Widhalm, John Ellenberger, Dan Parkinson, Dr. Beth Williams, Leslie Johnston, Dr. Donald Nash. ABSTRACT There are several aspects to this study because it is in the initial year and all portions of it are being reported on under this job. Much of the work reported here is exploratory in nature, with the ultimate objective of . investigating reasons for reduced reproductive rates. Six pronghorn fawns were hand-reared to be used in nutrition and breeding experiments. Two mature does were determined to have multiple estrous cycles (4 and 3) with termination of cycling in late December. Cycles varied from 25 to 31 days. Blood samples were collected from wild-trapped animals from four areas of the state. Blood urea nitrogen (BUN) was examined as an indicator of nutritional status. There were significant (p < .05) differences in BUN's among areas and between years on one area. Trapping and transplant records were assembled to determine the populations that may have a common gene pool. Genetic variability will be estimated by electrophoresis of blood and tissue samples collected from various parts of the state. Trace mineral levels were estimated for zinc and copper from four areas of the state. There were significant (P < .001) differences in serum zinc levels between areas but it is unlikely Colorado pronghorn are ztnc d~fi~ient; -There w~re no significant (p > .05) differences in serum copper levels among areas and the values were well within the range considered to be normal. ��185 PRONGHORN POPULATION STUDY Thomas M. Pojar Since this single job involves several studies (see Segment Objectives), the format of this report is arranged so' that each study is reported independently. For reading continuity, the complete report, including methods, is included under each study heading. P. N. OBJECTIVE To deterTIine causes for low fawn: doe ratios of pronghorn Southeastern Colorado. populations in SEGMENT OBJECTIVES 1. Rear pronghorn fawns for blood parameter 2. Conduct pronghorn 3. Collect and analyze blood from wild populations nutritional status. 4. Assemble 5. Collect and analyze serum from wild pronghorn genetic variability. 6. Collect and analyze pronghorn pronghorn breeding and nutritional studies. trials. trapping and transplanting and associate it with records. populations to estimate tissue samples for trace elements. TRACE MINERALS Zinc and copper levels were measured in 10 serum samples each from four areas of the state. The blood samples were collected from live-trapped animals from the following areas: Saguache, Rocky Ford, Hugo, and Villa Grove. Since there are no established base levels published for these minerals in pronghorn serum, their samples were analyzed to provide an indication of the variability that might be encountered from different ranges and general baseline values for pronghorn in Colorado in winter. Zinc There is great variability in the concentration of Zn in various tissues in the mamalian body. Highest concentrations are found in the liver and kidney (Underwood 1977) and values that are reported in published literature for pronghorn are from liver samples (Stoszek et al. 1978, Munshower �186 and Newman 1979). Zinc is not stored in the blood, therefore short-term changes can be expected with changes in the Zn level of the diet. Evans and Hahn (1974) ~bserved a drastic decline in the serum Zn levels of rats that were fed a Zn deficient diet for thirteen days (0.62 ppm) compared with rats maintained on a stock diet (1.24 ppm). Serum Zn levels may be better indicators of the adequacy of the diet than organ Zn levels because, according to Hambidge (1974:171) " •••an inadequate dietary supply of zinc is not compensated by mobilization of zinc from liver or bone •.•". Although the mean Zn concentrations observed in this study (Table 1) are well above what would be considered deficient for other mammalian species, it must be kept in mind that these samples were collected in late autumn and winter. It is possible that during other seasons deficiencies could occur. Growth retardation is one of the first and most important manifestations of zinc deficip.ncy (Hambidge 1974). Underwood (1977) reviewed several studies in an attempt to determine what plasma levels of Zn constitute a deficiency. He concluded that no deficiency symptoms were observed in sheep with plasma levels ranging from 0.53 to 0.89 ppm. It must be noted that the values obtained in this study are from serum so they are not directly comparable to Underwood's findings for plasma. In human serum, Zn concentrations were consistently higher than plasma concentrations by an average of 16 percent (Underwood 1977), so this adjustment could be made for a more direct comparison. Stoszek et ale (1978) found no significant difference (P > .01) in the Zn level of pronghorn liver samples from three eastern Idaho populations and one Montana population. Their results are as follnws (in ppm): Montana, 100.3 (n=16), Pahsimeroi River 82.1 (n=17), Birch Creek 83.9 (n=9) and Little Lost River 91.6 (n=13). From a sample of 20 pronghorn in Southeastern Montana, Munshower and Neuman (1979) obtained a mean of 84.8 (S.D. = 28.4) ppm. It seems unlikely that Colorado pronghorn populations are adversely affected by deficiencies in zinc in view of the survey results presented in Table 1. Older literature contends that deficiency of this element in grazing animals in unlikely because of the relatively high amounts present in natural pastures (McDonald et ale 1969). However, Maynard et ale (1979) cite more recent studies that suggest borderline deficiencies may be much more common than previously thought. Although all areas in Colorado that were sampled appear to have sufficient levels of Zn there was a highly significant (P < .001) difference among areas (Table 2). Underwood (1977) cites significant regional differences in the United States in plasma Zn levels and presumes that it is a reflection of differences in dietary Zn intakes. It is interesting to note that the concentration of this element is significantly different (P < 0.01) between the Villa Grove and Sagauche areas (Table 3). These trap sites are only about 10 miles apart, both in the San Luis Valley, but the samples were taken at Saguache on 6 February 1980 and at Villa Grove 22 November 1980. This indicates that either large differences can occur in proximate geographic areas or that large temporal changes can be expected. The Rocky Ford sample was the only area that had sufficient males for a comparison between sexes. A t-test was used for the comparison which showed no significant (p > .05) difference between sexes. �Table 1. Zinc and copper levels in serum collected from live-trapped pronghorn antelope from four areas of Colorado. Saguache - TraEped 6 Feb. 1980 Rocky Ford Trapped 18 Jan. 1980 PPM Field ID No. S-15005 S-15047 S-15056 S-15065 S-15066 S-15068 S-1508l S-15094 S-15097 S-D2 Sex Female Female Female Female Female Female Female Female Female Unk Age Mature Fawn Yearling Mature Mature Mature Fawn Hature Yearling Unk PPM Zn Cu 9.3 5.8 3.6 5.8 1.2 6.6 3.6 6.0 5.8 3.2 0.3 0.6 1.2 0.6 <0.3 0.3 0.6 0.3 <0.3 0.3 x 5.0900 S.E. 0.7109 Field ID No. Sex RF-075 Male Hale Hale Female Female Male Female Hale Hale FemClle RF-On RF-122 RF-125 RF-126 RF-129 RF-l30 RF-l35 RF-l37 RF-149 Age Hature Fawn Nature Fawn Hature Hature Nature Nature Fawn Fawn Zn eu 4.5 1.2 2.4 1.2 1.8 2.1 1.2 1.2 1.5 1.5 0.9 0.9 0.6 0.9 0.3 0.3 0.9 1.2 0.9 O~ x 1.8600 S.E. 0.3208 0.4800 0.0916 Villa Grove - Trapped 22 Nov. 1980 Hugo - TraEEed 10 Jan. 1980 PPM Field ID No. H-42 H-45 H-48 H-l35 H-143 H-147 H-15l H-155 H-163 H-182 Sex Female Female Female Female Female Female Hale Female Female Female Age Mature Yearling Mature Mature Mature Fawn Mature Yearling Yearling Fawn x S.E. 0.7800 0.0916 PPM Zn Cu 12.9 12.9 16.2 5.1 6.6 2.1 8.8 12.9 8.1 1.5 0.6 1.5 1.5 0.3 0.9 <0.3 0.9 1.2 0.9 <0.3 8.7100 1.5716 Field ID No. 0.8400 0.1469 ~~---~-.-----.~-..__..-. VG-4 VG-16 VG-22 VG-24 VG-33 VG-34 VG-42 VG-43 VG-48 VG-65 Sex Hale Female Female Female Female Male Female Female Female Female Zn Age Yearling Nature Yearling Hature Hature Mature Fawn Hature Yearling Mature <0.3 1.2 1.8 <0.3 3.0 1.5 2.4 1.5 2.1 0.6 x 1.4700 S.E. 0.2844 CIl 0.3 0.6 0.9 0.3 0.6 0.3 0.6 1.5 0.2 0.3 O.5nOO 0.1248 I-' 00 ...•.• �188 Table 2. Colorado. Analysis of variance of serum zinc levels from four areas of Log transformation on the data has been ~ade. AOV Source DF -------- Total Area Error F for Bartlett's = 0.13310 -------------- SS F MS 39 35.5558 3 19.6817 6.5606 36 15.8741 0.4409 test for homogeneous variances 14.8784 1.8666 p Table 3. Comparison of log transformed data means. means that are not different at P < 0.05. Area Mean Villa Grove 0.2168 Lines connect those Rocky Ford Saguache Hugo 0.6154 1.5475 1.9682 �189 Copper Underwood (1977) cites several studies that demonstrate Cu deficiencies result in reproductive failure due to fetal death and resorption. He also cites papers that associate Cu-deficient pastures with delayed or depressed estrus in cattle. Infertility that is accompanied with small dead aborted fetuses has been elicited from experimentally Cu deficient ewes. Copper deficiency can be manifested in many ways depending on the severity and duration of the deficiency and with the sex, age, and species of animal. Some additional affects of Cu deficiency that have been observed are: Anemia and impaired iron metabolism, bone disorders, neonatal ataxia and lesions of the central neverous system, impaired keratinization [pronghorn horns are made up of Keratin (O'Gara and Matson 1975)], and reduced pigmentation of hair. Maynard et ale (1979) refer to a study from Northern Europe in the early 1920's where copper deficiencies in the forage resulted in what was called a "wasting" disease in cattle and sheep. The symptoms were characterized by diarrhea, loss of appetite, and anemia. Other cases of a wasting disease in which nervous disorders have been symptomatic have been reported in Australia, South Africa, New Zealand, Scotland and in Southeastern United States. Copper deficiency of range plants were implicated in these situations with the possibility of excess molybdenum causing a conditioned copper deficiency. The normal range of Cu concentration in the blood of healthy animals is 0.5 to 1.5 ppm (Underwood 1977). The serum samples collected from the four areas of Colorado had mean values that ranged from 0.48 ppm at Saguache to 0.84 ppm at Hugo (Table 1). There were no significant (P < .05) differences in means from the four areas. Selenium Stoszek et ale (1978) implicated selenium deficiency as a cause of poor health and reduced survival of pronghorn fawns in Idaho. They compared Se levels of 3 Idaho pronghorn populations that had low reproductive rates with the Se level of a Montana population with more normal reproduction. There were no significant (P > .01) differences in Se levels of the 3 Idaho populations but all were significantly (P < 0.01) less than the Montana population. It was decided not to measure the Se levels of Colorado pronghorn blood samples because all of the state is classified as having adequate to high Se in forages. Some of the state is even characterized as having excess Se (Maynard et ale (1979). PRONGHORN FAWN REARING Six pronghorn fawns were raised at the Colorado Division of Wildlife Foothills Facility near Fort Collins. All of the fawns, except 1 came from the Pawnee National Grasslands enclosure located north of Nunn, Colorado. The exception was obtained from the Pueblo area. Thirteen fawns were captured. The mortalities were all related to digestive upset and diarrhea. The care and feeding protocol followed very closely that described by Splichal (N.D.). �190 Since newborn ruminants. depend almost entirely on maternal colostrum for antibodies (Trindle et al. 1979), we attempted to leave the fawns with their mothers for 24 to 48 hours. A blood sample was drawn as soon as the fawns arrived at the rearing facilities. Total serum protein was estimated using a refractometer. In mule deer (Odocoileus hemionus) fawns, the amount of passive immunity obtained from ingestion of colos~ trum is indicated by the serum protein level (Trind1e et al. 1978). The serum protein levels and colostrum ingestion relationship has not been established for pronghorn antelope. Therefore, the values in Table 4, are indicative only of those encountered in fawns that have been left with dams 1 to 7 days. The two exceptions to this are fawns numbered 8 and 9. These fawns were born to a doe that was held in captivity so it was possible to get blood samples at birth and 6 days later when they were removed from the doe. The initial serum protein value for number 8 was 4.0 g/lOO m1 at birth, but before nursing, and increased to 4.6 g/lOO'ml 6 days later. For number 9 the initial value was 4.1 g/lDO ml and increased to 4.8 g/lOO ml 6 days later. The total serum protein values reported in the literature for mule deer fawns, in general, is somewhat higher than that observed in the pronghorn fawns. In four groups of hand-reared mule deer fawns Trindle et al. (1978) measured average total serum protein values of 7.1, 6.4, 6.9 and 6.6 g/lOO ml. Halford (1974) found significantly (p < .05) higher serum protein (6.5 g/lOO ml, n=19) in dam-raised fawns compared to hand-raised fawns (4.6 g/lOO ml, n=32). He attributed this difference to better overall nutrition of the dam-reared fawns. Youatt et al. (1965) took total serum protein determination on 5 white-tailed deer (0. virginanus) fawns at various postpartum intervals. The mean values for 1, 3, 7, and 21 days postpartum were 5.3, 5.0, 6.1 and 5.5 g/lOO ml respectively. BREEDING EXPERIMENT A hand-reared 2 year-old buck was vasectomized and kept with two tame sexually mature does. The buck was fitted with a leather breeding harness that holds colored wax-base marking material which marks the does when they are mounted. The marker color was changed monthly. Pronghorn courtship behavior (territory establishment and harem formation) can begin many weeks before the breeding season (Kitchen 1974), therefore the buck was kept with the does throughout the summer so no phase of the courtship process was missed. Consistent and typical courtship behavior was observed in late August and the harness was put on the buck on 28 August. Brief notes on the behavior of the 3 animals were taken during 15 minutes to one hour observation periods in the morning and afternoon. A summary of these observations is presented in Table 5. �Table 4. Statistics on fawns captured and hand-reared in 1980. Age at Body Wt. at Est. Capture Date of (lbs) Birth Blood Values at Capture Total Gamma Protein Globulin (g/lOO ml) (g/lOO ml) Fawn No.a Sex Capture (Days) 1 F 3 7.53 6/2 2 F 3 6.63 6/2 3 M 1 5.55 6/3 4.0 4 F 1 6.17 6/3 5 M 7+b 8.29 6 F 2 7 M 2 Remarks 6/20 Healthy survivor 6/9 Healthy surviror 0.0 Unk Died 6/7 4.5 0.5 Unk Died 6/6 5/30b 4.2 1.2 None Always healthy 4.00 6/10 4.8 0.5 6/14 Intermittent diarrhea but survived 5.83 6/10 5.7 2.0 6/19 Intermittent diarrhea but healthy survivor 8 M 6 6.85 6/12 9 F 6 6.15 10 M 3 11 M 12 13 4.4 0.9 no sample c - never sucked 0.4 6/18 Died 8/24 6/12 4.6 c 4.8 0.8 6/29 Chronic diarrhea, suryided but stunted 6.25 6/14 5.1 0.7 6/22 Died 6/30 3 6.00 6/15 5.3 1.3 6/29 Died 7/2 M 2 6.25 6/18 M 3 6.15 6/20 x = 6.28 SD = 1.00 a b c Date of First Morbidity no sample Died 4.5 1.0 4.72 0.50 2.78 2.04 6/23 Died 7/3 The following fawns are siblings, 1 and 2, 3 and 4, 6 and 7, 8 and 9. The exact date of birth and age at capture are unknown - therefore these are estimates. Since these fawns were born in captivity these blood parameters were estimated at birth also, before they nursed. The values for #8 was 4.0 and 0.0; and for #9 they were 4.1 and 0.3. I-' \0 I-' �192 Table 5. Notes on behavior of captive pronghorn antelope during breeding season. Notes beginning in a particular column pertain to the animal at the head of that column. Date ~ 11224 Mandy ~ 11225 Hanna 8/28 (f'Rocky Put breeding harness on Rocky 9/20 Rocky chasing and vocalizing 9/24 Marked lightly 9/25 ~~Moremarkings Following Mandy with some courtship display 9/27 Vocalizing and following Hanna w/courtship display anc territorial display 9/28 Courtship display to Mandy. Started following Hanna. Much herding of does 9/29 Herding and protective of does 10/1 Aggressive to caretaker and buck in adjacent pen 10/3 Very aggressive - does avoid him 10/4 Very aggressive, herding and courtship display 10/4 Briefly mounted twice - no marks �193 Table 5. Continued Date ~ 1/224 Mandy 10/7 *Marked - would not permit mounting ~ 1/225 Hanna cfRocky 10/10 Shows little interest in either doe, some herding 10/11 Shows little interest in does 10/12 Aggression toward caretaker and adjacent buck 10/13 Aggressive, but little interest in does 10/14 Very aggressive and vocalization, herding of does 10/15 *Light1y marked Still very aggressive 10/16 Reduced aggressiveness 10/19 Mellow and showing little interest in does 10/20 Not aggressive 10/22 Aggressive to caretaker and other adjacent buck, some herding of does 10/23 Aggressive, does 10/25 Less aggressive, herding 10/27 Not aggressive, little interest in does herding some �194 Table 5. Continued Date .'f. 11224 Mandy .'f. 11225 Hanna cl'Rocky 10/30 Increased aggressiveness 10/31 Very aggressive to caretaker and adjacent buck, vocalization 10/31 Marked 11/1 11/2 Vocalizing ar.dvery aggressive *More markings 11/3 Vocalizing and very aggressive 11/9 Light markings 11/10 *More light markings, held tail up when approached by Rocky 11/10 Very aggressive, precopulatory displays, mounted Hanna 11/12 Still aggressive 11/14 Decreased aggressiveness 11/16 Not aggressive, little interest in does 11/18 Lost one horn sheath 11/19 Lost other horn sheath 11/22 Slight aggressiveness 11/25 Increased aggressiveness --------------------------------------------------------------------------- �195 Table 5. Continued Date ~ 11224 Mandy ~ 11225 Hanna 11/26 11/28 ci"Rocky Aggressive and herding does Marked 12/1 Accompanies Mandy 12/3 *Ho1ds tail up when approached by Rocky 12/4 Very aggressive 12/8 Very aggressive 12/10 *Marked 12/12 Playful, not very aggressive 12/14 Not aggressive 12/16 Some herding of does 12/18 Very aggressive to caretaker 12/19 Herding does 12/21 Not aggressive 12/24 Increased aggressiveness 12/26 Mounted Mandy 12/27 Marked 12/27 12/28 Very aggressive *More markings-many 12/29 Herding the does 12/31 Aggressive to caretaker and adjacent buck -----------------------------------------------.-'!"'""-----.,...,-----.----.~----.---,....- �196 Table 5. Continued Date ~ /1224 Mandy ~ 11225 Hanna cl'Rocky 1/3 Decreased ness 1/4 New mark 1/5 New mark 1/6 New mark ~(light. aggressive- single marks -.not typical of estrus) 1/12 Some herding 1/16 Little 1/20 Increased aggressiveness, follows Hanna 1/23 Little 1/27 Not aggressive and little interaction 2/3 Aggressive caretakers 2/5 No aggressive behavior 2/15 Some herding intense interaction interaction toward but not 2/183/11 Little interaction no aggressive behavior 3/16 Some vocalization "snort-wheeze" 3/20 Vocalization 3/22 Vocalization 3/24 Pursued does 3/28 Some chasing of does - one of the ---------------------------------------------------------------------_----- �197 Table 5. Continued Date ~ #224 Mandy ~ 11225 Hanna 4/3 5/12 * c!'Rocky Some chasing of does but no signs of mounting Best estimate of time of estrus based on markings and behavior. Table 6. Record of estrus dates, estimated length of estrous cycles and potential parturition datesa of two pronghorn antelope does. Reproductive Female Numbers #224 #225 Hanna Mandy Events b 1st Estrus Potential date 25 Sep 4 June 15 Oct 24 June 2nd Estrus Length of estrous cycle Potential parturition date 7 Oct 12 16 June 10 Nov 26 20 July 3rd Estrus Length of estrous cycle Potential parturition date 2 Nov 26 12 July 10 Dec 30 19 Aug 4th Estrus Length of estrous cycle Potential parturition date 3 Dec 31 12 Aug 5th Estrus Length of estrous cycle Potential parturition date 28 Dec 25 6 Sep aGestation parturition period (days) is assumed to be 252 days (Hepworth and Blunt 1966). bIt is not certain that this was a true estrus in view of the fact that the second, and obvious, estrus was only 12 days later. �198 is unlikely an estrous period passed without noticeable mark-ing, although it was only possible to discern the general time frame of the estrous period. It is also likely that light markings were made during normal courtship activity which do not necessarily represent copulation. It is possible this is what happened in the first recorded estrus of number 224 since the next markings were only 12 days later (Table 6). Disregarding the first markings of number 224 the dates of first potential breeding of the two does were 7 October and 15 October for does numbered 224 and 225, respectively. Assuming a gestation period of 252 days (Hepworth and Blunt 1966), the fawning dates would have been the 16th and 24th of June. The last obse.rved estrus was 28 December which would have resulted in a parturition date of 6 September (Table 6). One of the does had 4 estrous periods and the other had 3. The apparent estrous cycles ranged in length from 25 to 31 days. Most of this variability is probably due to the detection method used. PRONGHORN BLOOD ANALYSIS Blood assays have been used to indicate the nutritional status of wild ungulates as it relates to available food sources (Seal et al. 1978, Seal and Hoskinson 1978). Researchers have found a significant relationship between diet protein levels and blood urea nitrogen '(BUN) in wild ruminants held in captivity: white-tailed deer, (Odoceoileus virginianus) (Seal et al. 1972, Kirkpatrick et al. 1975, Bahnak et al. 1979) and bison (Bison bison) (Keith et al. 1978). Unlike many blood parameters, BUN is quite stable under temporary stress (Barrett and Chalmers 1977, Trout 1976, and Franzman and Thorne 1970). Capture stress can be at least crudely assessed through analysis of intracellular enzymes such as lactic dehydrogenase (LDH) , serum glutamic oxaloacetic transaminase (SGOT) and creatine phosphokinase (CPK). These substances are released into the circulatory system when there is injury to cells (Coles 1967). Skeletal muscle, myocardium and the brain contain high concentrations of CPK and elevated values of CPK are observed after various types of physical trauma including parturition (Nieminen 1980) and after vigorous physical exercise (Savignano et al. 1969). Two very tame female pronghorn were bled via the jugular vein within one minute of being physically restrained. They were then immobilized with 4 mg of M99 and 40 mg of Rompun. They were then used in an experiment of detecting pregnancy with X-Ray and Ultra-sound. This was done on 1 May and both were determined to be carrying twin fetuses. They were immobilized for approximately one hour and just prior to administering the antidote M50-50 (8 mg) the second blood sample was taken. The Hycel values from this sample are presented in Table 7 in the "after" column. All of the intracellular enzymes showed drastic increases. Seal et al. (1972) observed spectacular increases (3-to 200-fold) in SGOT, CPK, and LDH after restraining white-tailed deer for 45 minutes either with drugs or manually. The mean values of these enzymes of wild-trapped pronghorn are all higher than those of the tame animals immediately upon capture (Table 8). �199 Table 7. Hycel blood values of tame pronghorn immediately upon restraint and after about 1 hour of immobilization with M99 and Rompun. Blood Value Glucose (mg/dl) Blood Urea N(mg/dl) Pron~horn 11224 After at ImmobiliCapture zation 68 111 Pronghorn #225 After Immobiliat Capture zation 92 150 18.0 21.2 21.8 24.0 Globulin (Gm/dl) 2.9 2.8 2.3 2.6 Total Protein (g/dl) 6.4 6.0 5.5 5.9 Chloride (mg/dl) 112 112 120 119 Cholesterol (mg/dl) 50 50 50 61 Potassium (mEq/L) 6.40 5.40. 6.10 5.95 Sodium (mEq/L) Total :gilirubin (mg/dl) 147 0.95 146 1.30 146 1.30 146 2.20 Alk. Phosphatose (U/L) 12.5 13.5 l3.0 16.5 SGOT (U/L) 100 186 125 300 SGPT (U/L) 47 56 54 92 CPK (u/L) 96 152 96 191 Inorganic Phosphorus (mg/dl) 7.5 LDH (U/L) 380 5.9 775 8.1 450 6.1 1150 Calcium (mg/dl) 9.6 9.2 10.1 10.5 Creatinine (mg/dl) 1.0 1.2 1.1 1.7 �Table 8. Hycel serum values of wild-trapped Area Date TraEEed Blood Parameter pronghorn antelope. LaJunta 1-18-80 Craig U-29-Z9 n x Glucose mg/dl 2l 139.09(59.38) BUN mg/dl 21 19.62(5.87) (SO) n x (SO) No Data 30 20.83(5.99) n 24 24 x Villa Grove 11-22-80 Saguache 2-6-80 Hugo 1-10-80 (SO) 193.42(110.21) 13.3(6.12) n 30 30 x (SO) 167.23 (l05. 58) 25.66(6.48) x n (SO) 37 n x {SO) 24 185.58(84.51) 33.36 (11. 33) 28 28.75(9.12) 211.32(95.26) 37 Hugo _ 1-28-81 , 21 2.07(.82) No Data 24 2.18(0,51) 30 2.35 (.64) 37 2.37(0.64) 25 2.84(1.08) Total Prot. g/dl 21 5.28(1. 56) N0 Data 24 6.72(0.54) 30 7.14(.69) 37 6.81(0.53) 25 7.69(1.27) Chloride mg/dl 23 95.26(13.40) No Data 24 105.33(2.88) 30 108.4(6.14) 37 113.54(4.14) 2l 109.71(4.04) Cholesterol 22 55.18(12.31) No Data 24 62.12(6.98) 30 71.00(14.54) 37 69.70(10.03) 25 77.32(27.08) K mEq./L 21 5.53 (1.01) 26 5.75(1.07) 24 5.72(0.71) 30 6.90(1.60) 37 6.55(0.91) 18 9.12(2.58) Na mEq./L 22 144.23(12.05) 28 151.16(8.02) 24 153.15(2.52) 30 150.18(4.24) 37 22 149.54(3.87) Tot. Bilirubin mg/dl 24 0.82(0.18) 35 0.98(.24) 24 0.94(0.22) 30 1.83 (0.92) V 1.28(0.41) 26 2.05(2.12) Alk. Phos. U/L 23 28.95(13.15) 35 25.57(i2.79) 24 26.48(13.44) 30 27.67(14.23) 37 31. 84 (16.91) 26 41.44(26.17) SGOT U/L 23 206(127.19) 30 175(26.69) 23 SGPT U/L 24 119.29(42.45) 35 103.08(31.23) 24 CPK U/L 24 110.8(92.09) 35 101.1(55.92) 24 116.2(34.14) 30 Inorgan phos. mg/d1 25 5.81(1.75) 35 5.56(1.99) 24 6.85(2.03) 30 LDH U/L 21 No Data 23 Calcium mg/dl 25 9.62(2.32) 34 12.67(1. 20) 24 10.92(0.80) 30 Creatinine 25 2.36(0.48) 36 3.31(0.49) 24 2.50(0.54) 30 Globulin gm/ dl mg/dl mg/dl 890(380.28) 305(195.55) 47.25(11.48) 497(222.26) 30 30 413(143.98) 92.73(47.46) 154.7(98.47) 4.37 (1.01) 158.05 (4.07) 37 339(204.93) 27 384 (290.07) 37 177 .86(68.08) 27 123.89(90.24) 36 168.0(132.13) 27 125.4 (56.27) 37 6.86 (1.24) 28 5.52(+.73) 37 1315(1070.00) 25 1607(582.05) 13.08(1. 56) 37 13.10(1.19) 27 15.25(4.68) 2.69(0.35) 37 3.24(0.59) 28 3.93(2.29) 30 1079(534.90) N 0 0 �201 Table 9. Analysis of variance table of BUN values. AOV Source DF Total 163 13936.55 5 158 Area Error SS MS F 5392.34 1078.47 19.94 8544.22 54.08 F for Bartlett's test for homogeneous variances P=0.05495 Table 10. Comparison of mean BUN values. are not different at P < 0.05. Area Year Mean a a Hugo Craig LaJunta (80) (79) 19.86 13.35 Year trapped 2.16 Lines connect those means that Saquache Hugo Villa Grove (80) (80) (81) (80) 20.83 25.53 28.27 30.69 �202 BUN values from wild trapped pronghorns were significantly (P < .05) different among some areas and between years on 1 area (Tables 9 and 10). BUN values from pronghorn of the Hugo area were significantly different (P < 0.005) from 1980 to 1981 (Table 10). The trapping operation in the Hugo area took place in 1980 following several weeks when forage availability and pronghorn mobility were restricted because of snow. Eight to 12 inches of crusted snow covered the low growing vegetation. The trapping in 1981 on the same area was after many weeks of very mild weather and virtually no snow cover. Comparison of BUN values from different areas should not be attempted without consideration of recent. foraging conditions. PRONGHORN TRAPPING AND TRANSPLANT RECORDS In order to better assess the genetic variability data it was necessary to determine if the populations being compared had a common gene pool. There has been a great deal of pronghorn transplanting in Colorado (Bear 1969). Trapping and transplanting records from all four Division Regional offices and the Denver office were examined. In addition, some of the key Division employees who were directly associated with pronghorn transplants were interviewed in person. The data thus collected will be tabulated in terms of trap site, release site, number and sex of animals moved, and relative success of the transplant. GENETIC VARIABILITY Electrophoresis analyses were run on blood and tissue samples collected from four areas of the state. Heart, liver, kidney and muscle tissues were collected from hunter-killed animals from game management units A3 (Craig) and A56 (LaJunta). Serum samples were collected from these same units but from live-trapped animals. Blood samples (including the red blood cell component) were collected from live trapped pronghorn from the Hugo and Saguache areas. A total of 396 electrophoresis runs were made on 15 different isozymps from red blood cell and tissue samples of the four areas. Six isozymes from serum were run on 10 samples from each of the four areas for a total of 240 runs on serum proteins. In total, 636 electrophoresis runs were completed (Table.). Analysis of these data are not yet complete0. (I �203 II Table •. Material, sample size, and isozyme used in electrophoresis analysis. Isozyme a Hugo Sample Size Tissue Saguache Sample Size Tissue Craig Sample Size Tissue LaJunta Sample Tissue Size PGM-l 7 RBC 10 RBC 10 Liver 10 Liver PGM-2 7 RBC 10 RBC 10 Liver 10 Liver LDH 7 RBC 10 RBC 10 Kidney 10 Kidney MDHS 7 RBC 10 RBC 10 Liver 10 Liver 10 Liver 10 Liver 10 Liver 10 Liver SOD 10 Liver 10 Liver GPD 10 Liver 10 Liver 10 Liver 10 Liver G6PD 10 Liver 10 Liver ACP 10 Liver 10 Liver SDH 10 Liver 10 Liver 10 Liver 10 Liver lCS S PG1-l SODA 7 RBC 10 RBC 7 RBC 10 RBC B l PGD 7 ESA 3 7 RBC RBC 10 10 RBC RBC ESA4 ALP 10 Serum 10 Serum 10 Serum 10 Serum ESD 10 Serum 10 Serum 10 Serum 10 Serum CAT 10 Serum 10 Serum 10 Serum. 10 Serum TF 10 Serum 10 SerUTIl 10 Serum 10 Serum LAP 10 Serum 10 Serum 10 Serum 10 Serum GP 10 Serum 10 Serum 10 Serum 10 Serum a See Harris and Hopkinson (1976) for full name of isozyme. �204 LITERATURE CITED Bahnak, B. R., J. C. Holland, L. J. Verme, and J. J. Ozoga. 1979. Seasonal and nutritional effects on serum nitrogen constituents in white-tailed deer. J. Wi1d1. Manage. 43(2):454-460. Barrett, M. W. and G. A. Chalmers. 1977. C1inicochemica1 values for adult free-ranging pronghorns. Can. J. Zool. 55:1252-1260. Bear, G. D. 1969. Antelope transplants in Colorado. Game Information Leaflet No. 70. Colo. Div. of Game, Fish and Parks. 3pp. Evans, G. W. and Carole Hahn. 1974. Albumin as a possible site for copper-zinc interaction. In: W. G. Hoekstra, J. W. Suttie, H. E. Ganther, and Walter Mertz (Eds.). Trace Element Metabolism in Animals - 2. Univ. Park Press, Baltimore. 775pp. Franzmann, A. W. and E. T. Thorne. 1970. Physiologic values in wild bighorn sheep (Ovis canadensis canadensis) at capture, after handling, and after captivity. J. American Vet. Med. Assoc. 157(5):647-650. Halford, Douglas K. 1974. A method for artificially raLsLng cmu Le deer: fawns. M. S. Thesis, Colo. State Univ., Fort Collins. 21pp. Hambidge, K. M. 1974. Zinc deficiency in children. In: W. G. Hoekstra, J. W. Suttie, H. E. Ganther, and Walter Mertz (Ed~), Trace Element Metabolism in Animals - 2. Univ. Park Press, Baltimore. 775pp. Harris, H. and D. A. Hopkinson. 1976. Handbook of enzyme e1ectropheresis in human genetics. North-Holland Publishing Co., New York. Hepworth, W. and F. Blunt. 1966. Research findings on Wyoming antelope. Wyoming Wi1d1., Special Antelope Issue. 30(6):24-29. Keith, E. 0., J. E. Ellis, R. W. Phillips, and M. M. Benjamin. 1978. Serologic and hematologic values of bison in Colorado. J. Wi1d1. Dis. 14:493-499. Kitchen, D. W. 1974. Social behavior and ecology of the pronghorn. Monograph No. 38. 96pp. Wild1. Kirkpatrick, R. L., D. E. Buck1ard, W. A. Abler, P. F. Scanlon, J. B. Whelan, and H. E. Burkhart. 1975. Energy and protein influences on blood urea nitrogen of white-tailed deer fawns. J. Wi1d1. Manage. 39(4):692-698. Maynard, L. A., J. K. Loos1i, H. F. Hintz, R. G. Warner. 1979. Animal nutrition. Seventh Edition. McGraw-Hill Book Co., New York. 602pp. McDonald, P., R. A. Edwards, J. F. D. Greenhalgh. Oliver and Boyd, Edinburgh. 407pp. 1969. Animal nutrition. �205 Munshower, Frank F. and Dennis R. Neuman. 1979. Metals in soft tissues of mule deer and antelope. Bull. Environ. Contam. Taxicol. 22:827-832. Nieminen, M. 1980. Nutritional and seasonal effects on the haematology and blood chemistry in reindeer, (Rangifer tarandus trandus L.) Compo Biochem. Physiol. 66A(3):399-4l3. O'Gara, B. W. and G. Matson. 1975. Growth and casting of horns by pronghorn and exfoliation of horns by bovids. J. Mammal. 56(4):829-846. Savignano, Thomas, Albert Hanok, and Jeremiah Kuo. 1969. Creative phosphokinase activity. A study of normal and abnormal levels. Am. J. Clin. Pathology. 51(1):76-85 Seal, U. S. and R. L. Hoskinson. 1978. Metabolic indicators of habitat condition and capture stress in pronghorns. J. WIldl. Manage. 42(4):755-763. ------- , L. J. Verme, J. J. Ozoga, and A. W. Erickson. 1972. Nutritional effects on thyroid activity and blood of white-tailed deer. J. Wildl. Manage. 36(4):1041-1052. ------~, M. E. Nelson, L. D. Mech, R. L. Hoskinson. 1978. Metabolic indicators of habitat differences in four Minnesota deer populations. J. Wildl. Manage. 42(4):746-754. Splichal, Cathy. N. D. Pronghorn antelope fawn rearing. Special Studies Report. Department of Fishery and Wildlife Biology, Colo. State Univ., Fort Collins. 15 xerox papers. Stoszek, M. J., W. B. Kessler, and H. Willmes. 1978. Trace mineral content of antelope tissues. Proceeding of the Eighth Biennial Pronghorn Antelope Workshop. May 2-4, Jasper Park Lodge, Jasper, Alberta, Canada. 156-161. Trindle, Bruce D., Lon D. Lewis and Lloyd H. Lauerman. 1978. Evaluation of stress and its effects on the immune system of hand-reared mule deer f~wns (Odocoileus hemionus). J. Wildl. Dis. 14:523-537. 1979. Techniques for evaluating humoral ------- , and cell-mediated immunity in mule deer fawns (Odocoileus hemionus). J. Wildl. Diseases. 15:25-31. Trout, L. E. 1976. Blood analysis of Idaho pronghorn. Proceedings of the Seventh Biennial Pronghorn Antelope Workshop. l22-l26pp. �206 Underwood, Eric J. 1977. Trace elements in human and animal nutrition. Academic PRess, New York. 545pp. Youatt, William G., Louis J. Verme, and Duane E. U11rey. 1965. Composition of milk and blood in nursing white-tailed does and blood composition of their fawns. J. Wi1d1. Manage. 29(1):79-84. �July 1981 207 JOB PROGRESS State of Project Colorado No. Job Title: ~ocky Covered: ~---------------- 6 Job No. Mountain Goat Ecology Personnel: Big Game Investigations W-126-R-4 Work Plan No. Period REPORT Goat Investigations 2 - Rocky Mountain Study July 1, 1980 through June 30, 1981 Dale F. Reed ABSTRACT Twenty-one mountain goats were marked with radio telemetry collars, neck bands, or ear tags. No bighorn sheep were trapped or marked. Census routes were established for locating mountain goats and bighorn sheep in the Mt. Evans area primarily along the highway. Winter conditions periodically precluded access to some of the routes. Mountain goat habitat preferences during the winter did not vary greatly from preferences during non-winter periods. During the year all marked groups of mountain goats except one either remained in the alpine or returned to the alpine after brief excursions into the subalpine. The excepted group moved to a subalpine habitat soon after having been trapped and banded and remained there throughout the winter and spring. Conspicuous periods occurred when bighorn sheep were either not readily observed or not present on the same alpine ranges as the mountain goats. ��209 ROCKY MOUNTAIN GOAT ECOLOGY STUDY Dale F. Reed P. N. OBJECTIVE To describe mountain goat and bighorn sheep habitat preferences within sympatric ranges, to determine if a system can be developed to estimate the intensity and nature of competitive interactions, and to measure dispersal rates of dispersing or translocated mountain goats. SEGMENT OBJECTIVES 1. Develop census routes to accurately and precisely goats and bighorn sheep on sympatric ranges. locate mountain 2. Estimate habitat preferences sheep populations. goat and bighorn 3. Estimate dispersal populations. of selected mountain rates of dispersing or translocated mountain goat METHODS AND MATERIALS Mountain goats were trapped and banded in three selected areas where they were known to come to saltlicks (Baumann undated) during the spring and summer· (Herbert and Cowan 1971). One to three clover traps were placed to cover as much of each lick as possible. The traps were regularly baited with salt, and periodically during the fall, with grain, apple pomace, or alfalfa when responses to salt diminished. The traps ~re set only when project personnel were in the area so that banding and tagging, collections, and measurements could be accomplished promptly. Trapped mountain goats were handled by roping and tying their horns and legs (strongly recommended in that order). A short piece of garden hose was placed over the horns for safety and a blindfold was pulled over the eyes to keep the animal as calm as possible. Selected measurements and live-animal weights (when taken): were made with a steel tape and a tripodscale (300 lb. Chat ilIon) assembly, respectively. Also, selected blood and fecal samples were collected for lymphocyte blastogenesis tests and fecal cultures (tests for Johne's disease) at Colorado State University Wild Animal Disease Center. Kids and male goats were ear tagged because of small size and potential trophy status of males. Yearling and 2-year old females were collared with individually numbered or color coded neck bands for purposes of identification. Adult females were outfitted with telemetry collars (Telonics, 1300 West University Drive, Mesa, AZ 85201) using 148 and 172 MHz. The first six telemetry collars placed on mountain goats were used for locating and eventual sight confirmation of the animals. �210 Later in the segment, three telemetry activity (tip switch) collars to be used in conjunction with a remote, portable telemetry data site were outfitted on mountain goats. Habitat utilization or preference was estimated by observing banded and tagged individuals of both species and noting their spatial and temporal relationships to sites which had been described previously as frequently used areas (Baumann undated). Movements within and between these areas were determined by direct observation or sight confirmation of marked animals often widely separated by time. Additionally, the location (± 10m to ± lOOm), date, and time of all observed mountain goats and bighorn sheep were recorded. UTM (Universal Transmercator) coordinates were measured (estimated to the nearest 10 m by extrapolating between 100 m grid lines (Fig. 1) for each of the locations for purposes of generating a computer data base and for future habitat utilization analyses. Censuses were conducted on a regular basis (2-3 times per month, weekly, or more often for selected routes or seasons) to detect changes in habitat utilization. Since work during this segment was preliminary, the area considered was all the alpine and any of the subalpine areas which mountain goats and bighorn sheep used sympatrically within a 100 km2 area (10 x 10 km square, UTM right 41-50, up 78-87) (Fig. 1) which includes the Mt. Evans shelter house on the east, Hells Hole and Bierstadt on the west, Goliath Peak on the north, and Rosalie Peak on the south. The sample unit used was the square km (1000 m2) and the next lower metric scale (100 m2). Sub-sampling at the 10 m2 scale in high density or high utilization strata was not used during this segment. Each km2 was designated by the 1000 m UTM grid lines drawn from the UTM trick marks on 1:24,000 or 1:62,500 topographic maps. For example the km2 that includes Lincoln Lake (Fig. 1) was designated 48-85. An area which includes Goliath Peak, Lincoln Lake, Rogers Peak, Mt. Warren, Chicago Creek, Mt. Spalding, Mt. Evans, Tumbling Creek, and Epaulet Mountain was covered during census routes. These routes were developed based on their accessibility and the likelihood of encountering habitats used by one or both species. RESULTS AND DISCUSSION Mountain Goats Twenty-one mountain goats were marked with radio telemetry collars, neck bands, or ear tags (Table 1). Of the 13 mountain goats trapped and banded during the summer and fall of 1980, 12 were females. The only male was one of four kids (Table 1). This was a disproportionate sex ratio compared to the eight mountain goats trapped and banded earlier in the year during June of 1981 when a 1:1 male:female sex ratio was obtained. During this earlier trapping in June, parous females were conspicuously absent and the group composition around the trapping or saltlick areas was predominantly yearlings, 2-year old animals, and adult males. It was hypothesized that the adult females began returning from their kidding areas during late June to early July when the new kids had become more mobile, and that after returning, they again exhibited their dominance (Chadwick 1977, Dane 1977) �Table 1. Date, age, sex, collar or tag, sample number, and selected measurements of mountain goats trapped in the Mt. Evans area. Sample No Girth (em) Length (em) Horn length Left Right (em) wt. (kg) No Date 1 22 Aug SO Y F Black collar 1 (Harvested 8 Sep SO) 2 22 Aug SO K F White ear tag 1 3 22 Aug SO A F White te1e 14S.13 4 lS Sep SO A F Blue te1e 148.30 114 159 5 22 Sep SO A F Yellow tele 14S.31 116 165 6 29 Sep SO K F Orange ear tag OOS 2 77 102 7 29 Sep SO A(6/ F Green 2 te1e (Ch 2) 3 116 152 S 30 Sep SO 2yr F Yellow diag (Ch 1) 4 94 l1S 9 30 Sep SO K F Yellow ear tag 79 5 69 99 2.S 3.3 10 30 Sep SO A(S) F Black collar 2 6 116 142 22.9 23.1 11 7 Oct SO K M Yellow ear tag (no No.) S 72 113 4.4 4.3 .25 12 7 Oct SO 2yr F Black collar 3 9 95 12S 13.2 12.7 41 13 7 Oct SO A(7) F Green 3 te1e (Ch 3) 10 121 147 21.3 20.3 77 14 9 Jun Sl 2yr F Red-White-Blue 11 100 143 21.0 21.5 15 10 Jun Sl Y M Yellow w/Green "-" (ear tag) 12 S9 125 14.0 14.1 16 10 Jun Sl A M Yellow w/Green 13 119 lS3 25.0 17 16 Jun Sl A(3) F Blue diag (Ch 4) 14 102 149 IS 17 Jun Sl A(3) M Yellow w/B1ue "+" (ear tag) 15 114 19 17 Jun Sl Y F Black collar 4 16 20 29 Jun Sl A(6) F Black te1e (Ch 6) 17 21 29 Jun Sl K M Yellow w/Blue "_" Age Sex Collar/tag tele (Ch 2) "+" 16.5 Horn to muzzle (em) Front leg rmsc-apu1a Hind foot (em) 16.5 27 21.6 21. 3 74 23.9 79.4 41 N f-' f-' 21 22.2 SO.O 60.0 30.0 lS.5 57.0 27.0 24.S 26.0 S8.0 34.0 21. 7 22.4 19.0 ·73.0 30.5 147 23.S 23.0 25.5 S7.0 31. 5 90 126 15.0 15.5 19.0 51.5 27.0 111 157 24.0 23.S 25.0 73.0 31.0 51 Sl 0.4 0.4 12.0 42.0 21.0 9.9 �212 s . 'II'" 6 . 4 2 o ' 2 4 6 a 1 62.500 Figure 1. 1:62,500 topographic map of the Mt. Evans study area showing the VTM 1000 m (1.0 km) grid lines. A 100 m grid scale is shown on the lower left. �213 at preferred sites; thus becoming more susceptible to being trapped at the saltlicks than the subordinate males, 2-year old animals, and yearlings. Blood and/or fecal samples were collected from most of the trapped mountain goats (Table 1). Of nine'samples either lymphocyte-blastogenesis tested or cultured, eight were negative and one positive for presence of Paratuberculosis (Johne's disease) (Williams personal comm.). Additional fecal cultures have yet to be completed. Measurements taken on most of the trapped mountain goats (Table 1) will be analyzed when greater sample sizes are obtained. Such measurements as girth may be used to estimate weight (Rideout and Horthen 1975) when the latter measurement cannot be obtained. Nine of the 21 marked mountain goats were outfitted with radio telemetry collars (Table 2). Of these, Blue tele and her cohorts were located and observed more frequently throughout the fall, winter, and spring than any other telemetered animal. Of 53 locations (Table 3), 52 were in alpine habitat and one was well into (1,040 m from and 207 m below nearest alpine) a subalpine burn where the "understory" vegetation was luxuriant. On several occasions during weekly censuses Blue tele and her group were not located even with the aid of radio telemetry. However, since she and her group were later "discovered" in the Lincoln Lake burn during the second week of May, it is attractive to conclude that she and her group were there during the previous periods when no telemetry signal was detected. If this is a reasonable assumption, then Blue tele and her group (included Yellow diag and Black collar 3; group size = 14) were in the burn approximately 15 days, 30 April through 14 May 1981. Blue tele and her cohorts used the area above Lincoln Lake the most extensively, the area southeast of Rogers Peak during mid-winter, and the subalpine burn during late winter to spring (Fig. 2). Table 2. Telemetry collar, channel, frequency, pulse, and activation for radios outfitted on mountain goats in the Mt. Evans area. Telemetry collar It.1hi te tele a Blue tele Yellow tele Yellow diagb Green 2 Green 3 Red-White-Blue Blue diag Black tele tele Channel 1 2 3 2 4 6 a Tele denotes telemetry b Diag denotes diagonal c Activity (tip switch) collars; Frequency (r-fllz) 148.1300 148.3000 148.3100 172.2375 172.2625 172.2875 172.2625 172.3125 172.3875 65 ppm Pulse per min. (ppm) 78 76 72 11 48 82 65-90c 65-90 65-90 head up, 90 ppm date Activation date 22 18 22 30 29 7 9 16 29 Aug Sep Sep Sep Sep Oct Jun Jun Jun 80 80 80 80 80 80 81 81 81 head down. �Table 3. Observed locations of Blue telemetry (Blue tele) and associated group classifications in the Mt. Evans area. Location Telemetry Collar Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele 18 22 22 23 1 1 2 4 6 10 11 12 14 15 23 31 6 11 19 20 20 25 26 3 11 22 13 20 Date General Sep Sep Sep Sep Oct Oct Oct Oct Oct Oct Oct Oct Oct Oct Oct Oct Nov Nov Nov Nov Nov Nov Nov Dec Dec Dec Jan Jan NW NE N NE SE SW SE SW NE SW SE SW SE SE NW N N NE NE NE NE N N NW NW NW NW N 80 80 80 80 80 80 80 80 80 80 80 80 80 80 80 80 80 80 80 80 80 80 80 80 80 80 81 81 Lincoln Lincoln Lincoln Lincoln Rogers Lincoln Rogers Lincoln Lincoln Rogers Rogers Lincoln Rogers Rogers Lincoln Lincoln Lincoln Lincoln Lincoln Lincoln Lincoln Lincoln Lincoln Lincoln Lincoln Lincoln Lincoln Lincoln a GrouE Classification UTM 4875 4920 4875 4910 4780 4810 4720 4845 4900 4670 4730 4820 4730 4775 4820 4845 4900 4910 4905 4900 4885 4835 4840 4850 4830 4820 4815 4871 8565 8545 8565 8560 8470 8490 8485 8495 8550 8440 8460 8515 8490 8470 8550 8550 8560 8550 8555 8560 8555 8550 8555 8570 8550 8548 8580 8560 K y~ 1 1 5 5 4 4 4 4 1 5 5 4 5 5 4 4 6 6 4 1 7 6 7 4 3 1 7 2 2 Yrf' YU 2 A~ 3 1 5 4 Ad' 1 1 1 1 1 1 1 2 2 3 1 2 5 1 7 3 2 3 3 4 4 7 6 3 2 5 3 8 2 4 4 9 4 U 2 1 1 2 2 5 AU b 4 5 5 1 4 4 4 3 5 1 1 2 4 1 3 1 3 1 1 1 ·7 2 2 2 5 3 6 10 7 4 12 14 5 5 3 6 -------------------------------------------------------------------_._--------------------------------- Total 5 4 15 14 11 11 11 14 2 13 13 11 12 14 13 13 23 24 20 8 29 24 23 15 14 11 23 8 N I-' ~ �Table 3. Continued Location Telemetry Collar Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Blue Date tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele tele 21 26 27 5 11 12 18 18 19 23 24 19 20 5 5 14 20 26 27 28 5 21 21 30 30 Jan Jan Jan Feb Feb Feb Feb Feb Feb Feb Feb Mar Mar Apr Apr May May May May May Jun Jun Jun Jun Jun General 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 NE NW N N N N N N N NW NW NW NW N N LL N NE NE NW SE NW NW N NE a Group Classification UTM Lincoln Lincoln Lincoln Rogers Lincoln Lincoln Lincoln Lincoln Lincoln Lincoln Lincoln Lincoln Lincoln Rogers Rogers Burn Lincoln Lincoln Lincoln Lincoln Warren Lincoln Lincoln Lincoln Lincoln 4905 4790 4870 4710 4848 4848 4838 4838 4827 4820 4793 4835 4835 4703 4718 4993 4860 4922 4870 4805 4580 4813 4810 4875 4900 8560 8560 8552 8705 8569 8569 8575 8552 8569 8568 8581 8554 8550 8697 8705 8406 8542 8553 8540 8532 8385 8535 8540 8565 8550 K Y~ 3 9 9 2 3 3 3 3 3 8 7 7 7 3 6 2 2 3 5 9 2 2 2 Yc!' YU 1 2 2 2 2 3 7 7 1 1 1 2 1 1 3 1 A~ 3 11 11 6 3 3 4 4 4 2 4 6 4 2 4 2 2 4 4 1 3 1 2 4 1 Ad" Au 1 7 7 1 b U Total 2 11 29 29 10 6 6 9 9 9 24 19 18 18 9 18 12 10 16 17 15 5 7 7 5 1 2 1 1 14 7 5 6 4 8 6 3 2 4 1 2 2 a First and second UTM coordinates are preceded by 4 and 43, respectively (i.e. 44800 438500). b K = kid, Y = yearling, A = adult, and U = classified. Blue tele is included in each group classification. N •... VI �216 Other telemetered and marked mountain goats were often associated with Blue tele, namely, Black collar 3, Yellow diag, Yellow ear tag plain, Yellow tele, Yellow ear tag 79, and Orange ear tag 008, in that order (Table 4). Consequently, Blue tele's movements were influenced by those of other animals as well. Hence, an analysis of their movements which basically constitutes an analysis of group dynamics - interchanges and changing group sizes per season and habitat - will be conducted in future segments. In addition to those mountain goats often associated with Blue tele and the Lincoln Lake area, White tele moved into the Chicago Creek area (Chicago Basin and north cirque) soon after being banded near Lincoln Lake and remained there until June 1981 when she came back to the trap site or saltlick. Green 2 was also banded in the Lincoln Lake area, but moved approximately 5.5km to several rock outcrops in the subalpine. She remained there with three other unmarked goats throughout the winter and had not returned to the alpine habitat as of the end of this segment. Black collar 2 and Green 3 were trapped and banded above Tumbling Creek and have remained in that area throughout the fall, winter, and spring. No movements of mountain goats have been detected between Tumbling Creek and Lincoln Lake-Chicago Creek areas. Three mountain goats were banded with activity collars (Red-White-Blue, Blue diag, and Black tele) (Table 2). Only preliminary testing of head-up and head-down (65 and 90 pulses per minute) activity was conducted by the end of the segment. Table 4. Frequency of observed associations in the Lincoln Lake group of marked mountain goats (total of each column = 100%). BT BT Marked animalsa B3 YP YT YD Y79 08 .19 .25 .32 .23 .23 .21 .19 .l3 .21 .12 .14 .21 .17 .18 .21 .25 .23 .21 .12 .07 YT .l3 YD .21 .23 B3 .32 .23 .25 YP .16 .19 .l3 .16 Y79 .10 .08 .09 .10 .08 08 .08 .08 .09 .08 .04 a BT YT YD B3 YP Y7908 - Blue tele Yellow tele Yellow diag Black collar 3 Yellow ear tag plain Yellow ear tag 79 Orange ear tag 008 .14 .1- �217 ':11·: 4. "1 o . . . .7 -l t) 8 1 62.S00 Figure 2. Estimated movements of Blue tele mountain goat based on observed locations except those shown with question marks. Question marks indicate location was based on telemetry and sightings at distances too far for collar observation. See Table 3 for data points. �218 Mountain goat reproduction as indicated by neonates obse1;"vedclosely associated with adult females appeared relatively high. Of the 12 animals collared, the status was determined on 11, of which, 7 should have been ' parturient (Table 5). However, only six apparently had kids, two (Yellow tele and Black collar 2) having had twins and another (White tele) apparently losing her kid (estimated mortality) within 27 days. Bighorn Sheep No bighorn sheep .were trapped or marked. Few bighorn sheep came into the traps baited with salt (compared to the vigorous response of mountain goats during the spring and summer), hay, or grain. Some response was detected with apple pomace baited during the fall. Five bighorn sheep banded during lungworm treatments (1 Feb 1977) were observed in the Mt. Evans area (Table 6). Only two, Yellow collar 17 and Yellow collar 19 (females, estimated age = 9) have been observed in the alpine ranges over extended periods. Their movements, based on limited data (Table 6), suggest that they move beyond or use habitat somewhat different than that of the mountain goats in the Lincoln Lake area (Figs. 3 and 4). Although most of the sightings of Yellow 1.7aRd -Yellow 19 were at the Lincoln Lake area, Yellow 17 used the Tumbling Creek area and Yellow 19 used the Goliath Peak area where no mountain goats have been observed. Table 5. Reproductive status of collared female mountain goats in the Mt. Evans study area during June 1981. No Collar 1 2 3 4 White te1e Blue tele Yellow tele Green 2 Yellow diag Black collar 2 Black collar 3 Green 3 Red-White-Blue Blue diag Black collar 4, Black tele 5 6 7 8 9 10 11 12 Date Banded Estimated Age No. Kids Aug Sep Sep Sep Sep Sep Oct Oct Jun Jun Jun Jun 4 > 4 > 4 7 3 9 3 8 -3 3 1 6 1 0 2 22 18 22 29 30 30 7 7 9 16 17 29 80 80 80 80 80 80 80 80 81 81 81 81 > 8 Unk 0 2 0 1 0 1 0 1 Kidding Site .b Chicago BaS1n Chicago Basin Unk Subalpine East Lincoln Burn Tumbling Creek Chicago Basin2 Tumbling Creek Unk Unk a Confirmed with 1 kid 2 June 81, without kid 29 June 81 and thereafter. b Moved to apparent kidding site despite absence of parturition. �Table 6. Observed locations of collared and ear-tagged female bighorn sheep and associated group classifications in the Mt. Evans area. Location a Collar or Ear Tag Yellow collar Yellow collar Yellow collar Yellow collar Yellow collar Yellow collar Yellow collar Yellow collar Yellow collar Yellow collar Yellow collar Yellow collar Yellow collar Yellow collar Yellow collar Yellow collar Yellow collar Yellow collar Yellow collar Yellow collar Yellow collar Yellow collar Yellow collar Yellow collar Yellow collar Yellow collar Red collar Ear tag 4 Ear tag 18 Ear tag 4 Ear tag 18 17 17 17 17 17 17 17 17 19 19 19 19 19 19 19 19 19 19 19 19 19 19 19 19 19 19 21 Date 11 22 30 4 5 11 12 28 15 28 29 23 10 13 20 4 5 12 23 24 5 20 12 20 28 28 30 15 15 16 16 Nov Dec Dec Feb Feb Feb Feb Jun Sep Sep Sep Oct Dec Jan Jan Feb Feb Feb Feb Feb Mar Mar May May May May Dec Sep Sep Sep Sep General 80 80 80 81 81 81 81 81 80 80 80 80 80 81 81 81 81 81 81 81 81 81 81 81 81 81 80 80 80 80 80 NW Lincoln Tumbling SE Rogers NW Lincoln SE Rogers N Lincoln SW Rogers SW Rogers NW Lincoln NE Lincoln N Lincoln NW Lincoln NW Lincoln NW Lincoln NE Lincoln. N Lincoln SW Goliath SW Goliath NW Lincoln SW Goliath N Rogers N Rogers N Lincoln N Lincoln NE Lincoln N Lincoln SE Evans NW Lincoln NW Lincoln NW Lincoln NW Lincoln GrouE C1assificatiorl 5 UTM 4775 4625 4742 4785 4725 4850 4670 4660 4780 4915 4865 4855 4815 4780 4905 4848 4832 4840 4832 4795 4722 4675 4820 4852 4780 4840 4521 4780 4780 4660 4660 8565 8153 8492 8570 8502 8568 8473 8453 8580 8560 8565 8560 8500 8569 8560 8570 8718 8770 8573 8718 8640 8585 8554 8547 8560 8571 8085 8580 8580 8460 8460 L Y~ yrf YU A~ Ac!' 1 2 2 1 3 1 2 2 1 1 2 1 4 3 6 4 6 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 3 3 2 1 2 1 1 4 2 1 2 2 1 1 1 1 1 1 1 1 1 1 4 4 4 1 3 4 3 3 4 1 1 7 2 5 2 1 5 6 1 4 4 12 12 1 3 3 AU U 7 1 1 2 15 6 1 3 3 3 2 1 1 2 1 1 17 2 15 15 Total 16 10 13 11 12 1 1 1 21 8 8 8 5 10 8 9 8 3 4 14 23 6 2 2 8 9 3 21 21 14 14 a First and second UTM coordinates are preceded by 4 and 43. respectively (i.e. 44800 438500). b K = kid~ Y - Yearling, A = Adult, and U = unclassified. tagged animals. Group classification includes collared or tv I-' 1.0 �220 lall·· 6 4 . . 2 o ;> -l Ii a 1 62.500 Figure 3. Estimated movements of Yellow collar 17 bighorn sheep based on observed locations. See Table 6 for data points. �221 ;811··.··· 6· . 4. 2 o . : • 2 -l 6 8 1 62.500 Figure 4. Estimated movements of Yellow collar 19 bighorn sheep based on observed locations. See Table 6 for data points. �222 Yellow 17 had not been sighted since 12 Feb 1981. Yellow 19 was not sighted during April 1981. Likewise, other unmarked sheep were conspicuously absent during April 1981 in the Goliath-Lincoln Lake-Rogers area. Use of radio telemetry will be necessary to obtain an adequate sample of movement data on these sheep. Trapping efforts will need to be intensified in future segments. Species Interaction Thirty-one instances of direct interaction have been noted between mountain goats and bighorn sheep in the Mt. Evans area. The intensity of these interactions primarily ranged from neutral or no overt response to moderate or intense flight reaction. Seventeen interactions were neutral, three resulted in apparent deterrence of bighorn sheep from salt lick use, one resulted in bighorn sheep walking away from mountain goats that came within 5-10 m, two resulted in slight flight reaction (walk away from approaching animal) of bjghorn sheep, three resulted in moderate to intense flight reaction (trot or run away from approaching or chasing animal) of bighorn sheep, one resulted in a bighorn sheep following several mountain goats with no apparent response by the mountain goats (considered positive interaction), and four resulted in slight to moderate flight reaction in mountain goats from the presence of bighorn sheep. In all agonistic interspecific behaviors observed, the mountain goats initiated the encounters and the bighorn sheep were the recipients. Although it could be suggested from these data that approximately 29 percent of mountain goat-bighorn sheep interactions result in bighorn sheep yielding space or other resources, such small-sample-size data should be used with caution. LITERATURE CITED Baumann, T. G. Undated. Final report for the 1978 Mount Evans bighorn sheep and mountain goat study. 38(56)pp. Typescript. Chadwick, D. H. 1977. The influence of mountain goat social relationships on population size and distribution. Proc. Int. Mountain Goat Symp. 1:74-91. Dane, B. 1977. Mountain goat social behavior: "play" behavior as affected by dominance. Symp. 1:92-106. social play structure and Proc. Int. Mountain Goat Herbert, D., and McT. Cowan. 1971. Natural saltlicks as a part of the ecology of the mountain goat. Can. J. Zool. 49:605-610. Rideout, C. B., and G. L. Worthen. estimating weight of mountain .•••.. 1975. goats. .•..j // Prepared by ( / --;,~ '._..-.~:' (,.. 'T'" Dale F. Reed Wildlife Researcher C ( Use of girth measurements for J. Wildl. Manage. 39:705~708 • �223 JOB FINAL REPORT State of Colorado Big Game Investigations Pro j ec t No. -'-'W_-.:.:cl_;_2.:....6-_:R:..;_-_4.:.__ _ 6 Work Plan No. Job No. Job Title: Rocky Mountain Composition Period Covered: Personnel: 3 ------~--------------------- Goat Investigations Comparisons - Dietary Quali~y and of Tame and Wild Rocky Mountain Goats July 1, 1980 through June 30, 1981 T. Dailey, R. Binford, and B. Gill ABSTRACT Diet quality and composition comparisons between tame and wild Rocky Mountain goats were not made. Wild goats could not be approached closely enough to accurately describe diet selection of plant parts within plant species. Therefore, only diet selection dynamics of tame goats are reported. Botanical and nutritive composition of diets of tame mountain goats (Oreamnos americanus) in alpine tundra of Colorado were studied during November 1978-July 1980. Goat diets were also compared with diets of tame bighorn sheep (Ovis canadensis canadensis) observed simultaneously in the alpine. Diets of goats during winter (November, January, March) contained 50% forbs, 38% graminoids, and 12% woody plants. The most frequently chosen individual forages were the graminoid Carex rupestris (20% of diets) and the shrub Salix (11%). During summer (July, August, September), forbs comprised 92% of the diets, graminoids 6%, and woody plants 2%. Principal forages were Trifolium, Polygonum, and Mertensia. Dicots contained more cell solubles, lignin, and crude protein, and less in vitro digestible dry matter (IVDDM) than graminoids. Animals altered diet choices in an apparent response to differences in nutrient composition among forages; in September increased consumption of Agropyron was correlated with its high concentrations of protein and IVDDM compared to other forages, and in March goats ate more forbs, and hence maintained stable dietary levels of protein and cell solubles, at the expense of reduced diet IVDDM. Diet IVDDM, diet cell solubles, and diet crude protein all decreased substantially through summer, but changed little through winter. Food niche breadth was inversely related to the quantity and quality of forage available. ��225 DIET QUALITY AND COMPOSITION COMPARISONS OF TAME AND WILD ROCKY MOUNTAIN GOATS Thomas V. Dailey P. N. OBJECTIVES 1. 2. Ha: Diet composition of tame and wild mountain Ho: No difference goats. Ha: Dietary quality of tame and wild mountain Ho: No difference in chemical constitution and apparent digestibility of diets selected by tame and wild mountain goats. in diet selection goats is dissimilar. of tame and wild Rocky Mountain goats is dissimilar. SEGMENT OBJECTIVES 1. Conduct Rocky Mountain habitat types. goat grazing trials in selected 2. Compile data, analyze results, seasonal and prepare a final report. METHODS AND MATERIALS Tame and Wild Mountain Goat Comparisons The successful completion of this experiment was dependent upon a major tentative assumption, i.e. that wild mountain goats could be habituated to an observer's presence at a distance close enough to provide an accurate bite count and plant identification. Provided this assumption could be validated, we intended to observe the forage selection of 6-7 tame and wild mountain goats during 1 summer grazing trial on Mt. Evans. The daily data collection was to consist of 1-2 observers counting from a close distance the number of bites by plant species consumed by 1 goat feeding for 1-2 hours or until the animal lost interest in feeding. On the starting date the tame animals were to be transported from Fort Collins to holding pens near Mt. Evans. One to 2 days of pre-trial grazing were to be conducted to allow familiarization of the animals to available forage. The observations of the 2 groups were to be alternated on a daily basis. This design would minimize phenological variation between observation periods for tame and wild animals. Each tame animal was to be grazed once each day. Little, if any, experimental control was exerted over �226 wild mountain goats; however, an attempt was made to observe the same goats during each day's grazing trial. Data collection was to continue until at least 3,000 bites per animal were gathered. Forage Evaluation Nutritional values of individual plants eaten by goats were to be assessed by chemical determinations of percent dry matter, nitrogen, acid detergent fiber, lignin, cell wall constituents and in vitro digestibility of species contributing at least 2 percent of total intake. Fifty-gram samples were to be handplucked from plants grazed by tame and wild mountain goats during the grazing trial; plant parts taken were to simulate as closely as possible those selected by the goats. If entire plants were taken by the animals, plants immediatay adjacent to those selected were to be collected. At least 25 individual plants were to be composited to f0rm each sample; these were to be frozen for subsequent analysis. Triplicate 10-gram samples from each composite were to be analyzed for chemical constituents. Dry matter digestibility was to be measured in vitro using procedures described by Tilley and Terry (1963) and modified by Pearson (1970). Triplicate 0.5 g subsamples of each sample were to be digested with rumen inoculum collected from a cow-maintained on a diet which was nutritionally similar to a summer time mountain goat diet. In addition 25 simulated bites were to be collected in paper bags for each species comprising 2 percent or more of the diet in order to obtain an average weight per bite. Estimates of diet composition based on these weights were to be compared to diet composition derived from bite counts. Study Area Selection The investigation was conducted in the Mt. Evans area. goats in this area were thought to be approachable. The wild mountain Also, the area was accessible to the tame mountain goat transport vehicle. Wild mountain goat foraging areas were located as the site for observing both tame and wild mountain goats. Composition and Quality of Tame Mountain Goat Diets Study Area Studies were conducted at the University of Colorado Mountain Research Station at Niwot Ridge, 24 km west of Boulder (Fig. 1). Shelters, oversnow vehicles, and transportation routes maintained at the station made it possible to work year round above timberline. �227 t. ~ • FORT COLLINS UNIVERSITY OF COLORADO~ MOUNTAIN RESEARCH./I • BOULDER STATION f\ 1\ f\ A '\ '\r-. -. COLORADO "--."'A. A. I\..... Figure 1. Geographic location of University Mountain Research Station at Niwot Ridge. _ of Colorado �228 Niwot Ridge is an interfluve, approximately 8 km long, on the eastern slope of the Continental Divide. The bedrock of the ridge is igneous and metamorphic mainly of Precambrian age (Webber and May 1977). During the late Pleistocene, glaciers occupied valleys south and north of Niwot Ridge. The ridge was not glaciated (Webber and May 1977). Soils are coarse-textured, well-drained, mostly acidic, and have thin surface-horizons rich in organic matter (Webber and May 1977). Plant matter decomposes slowly due to lack of moisture (Webber and May 1977). Permafrost on the ridge is sporadic (Ives and Fahey 1971) and too deep to affect vegetation (Webber and May 1977). Mean temperatures for July and January are 8.5 C and -13.2 C, respectively (Webber and May 1977). Mean precipitation during July is 72.2 mm and during January is 131.3 mm (Webber and May 1977). Wind clears snow from exposed ground and accumulates ~t in sheltered areas. Peak winds are stronger in January (49.9 km h-l) than in July (20.9 km h-l) (Barry 1973). The ridge is bounded on the north, south and east by a Picea engelmanniiAbies 1asiocarpa forest (Webber and May 1977). The flora of the ridge is composed of some 200 vascular species, 100 bryophytes and 50 lichens (Webber and May 1977). Vegetation on Niwot Ridge is characterized by closed meadows on level ground with an abundance of herbs which gives way to disrupted mats where solifluction is prevalent (Webber and May 1977). Fe11fie1d communities, which are characterized by Trifolium dasyphy11um and Selaginella densa, comprise approximately 40% of the ridge; 20% is covered with meadows dominated by Kobresia myosuroides (Komarkova and Webber 1978). Both fel1fields and Kobresia meadows are dry and lack winter snow cover (Webber and May 1977). The remainder· of the vegetation consists of wet meadows, snowbeds, and shrub tundra (Webber and May 1977). The major abiotic factor affecting distribution of vegetation on Niwot Ridge is availability of moisture which is influenced by snow distribution and wind (Osburn 1958, Marr 1961). Aboveground productivity of vascular plants on Niwot Ridge ranges from a few grams per square meter per year in stony fel1fields to about 400 g m-2 yr-l for shrub communities (Webber and May 1977). The estimated mean value for the entire ridge system above treeline is 172.5 g m-2 yr-l (Komarkova and Webber 1978). Average length of the growing season is 90 days (Webber and May 1977). Komarkova (1976) constructed a Braun-Blanquet vegetation unit hierarchy for Niwot Ridge and provided a vegetation map and description of its vegetation units. A description of the units in the winter and summer study sites is presented in Appendix I. A plant species list, comprised of taxa identified during mountain goat grazing trials, and by Marr (1961) is presented in Appendix II. Marr (1961) sampled flora at a location 300 m east of the summer study area. He sampled the same slopes found in the study area, and thus the plant communities (as effected by moisture availability) of the 2 sites are grossly similar. �229 Osburn (1958) investigated winter snow-free areas on Niwot Ridge, and recognized 4 vegetation stands; Kobresia myosuroides meadow, gopher gravel mulch, gravel mulch--cushion plant and Kobresia-gravel mulch. Kobresia myosuroides meadow is the climatic climax on winter snow-free areas, but is displaced by the other stands as a result of wind and the burrowing of pocket gophers (Thomomys talpoides) (Osburn 1958). Gophers reduce the thick soil "A" horizon essential for growth of Kobresia. Consequently, other plants invade these disturbed areas. Following disturbance, vegetation once again converges toward Kobresia meadows as soil organic matter is replaced (Osburn 1958). Osburn indicated that the sites selected for intensive study of each stand type, were representative of the stands in general. This assertion, coupled with my observations of gopher disturbance and plant composition in the winter study area led me to believe that Osburn's stand descriptions could be applied to this investigation. It appeared that each of the successional stages described by Osburn (1958) were present on the south slope of the winter study area. A list of plant species from Osburn (1958) is presented in Appendix III. The vegetation on the ridge has been grazed little by ungulates. Sheep and cattle grazing has not been permitted on Niwot Ridge since 1946, and a large part of the summer study area may never have been grazed by these animals (Osburn 1958). A few mule deer and elk inhabit the area. Bighorn sheep are absent now, but were plentiful until 1890 (Ives 1942). Rocky Mountain goats have never occupied the area; the nearest documented populations occur about 55 and 80 km to the north and south, respectively (Denney 1977). Why was Niwot Ridge chosen for this investigation? The access and facilities available were important considerations, especially for winter field work. Most important, the vegetative composition of Niwot Ridge is similar to that of areas occupied by wild mountain goats in Colorado. Plant taxa abundant in alpine habitat of wild goats (Agropyron, Artemisia, Carex, Festuca, Geum, Kobresia, Oreoxis, Paronychia, Potentilla, Salix, Trifolium) (Hibbs 1965:107, Adams 1981:93, Thompson 1981:94) were all common on Niwot Ridge (Appendices I, II, III). Mountain goats in Colorado generally occupy alpine terrain year round (Hibbs 1965, Adams 1981). During winter they are concentrated on windswept snowfree ridge tops, where vegetation is exposed (Hibbs 1965, Adams 1981). Separate winter and summer study areas were used in since access to the summer site was doubtful during elevation of 3,660 m the summer study area contained south facing slopes separated by a relatively level study area, 3,500 m in elevation, had areas free of winter. the alpine (Fig. 2), winter. At an steep north and area. The winter snow most of the �230 .••.. a •.•. .•••• _. ', ~""" :f Winter (foreground) evation 3,500 m) and summer (background) (3,660 m) study areas for tame mountain goat diet study, Niwot Ridge, Colorado. Photograph by J. D. Ives. �231 Diet Composition Diet composition was estimated by the bite count method; e.g., counting bites of forage of tame, trained mountain goats. Seven animals were used for grazing trials during the 2 years of sampling. Three goats who began trials as kids were observed during the first year (November 1978 - September 1979). During the second year (November 1979 - July 1980) 4 kids were added to the original experimental group so that as many as 3 yearlings and 4 kids were observed during each sampling period. During the final period (July 1980),6 animals were sampled; 3 were 25 months old and 3 were 13 months old. Diets were determined in ten 3 to 8-day sampling periods as follows: 25-27 November 1978, 17-19 January 1979, 13-16 March 1979, 1-3 August 1979, 28-31 August 1979, 25-28 September 1979, 6-8 November 1979 (yearlings), 10-11 November 1979 (kids), 9-15 January 1980 (yearlings), 17-19 January 1980 (kids), 15-18 March 1980 (yearlings), 20-22 March 1980 (kids), 1-3 July 1980 (2-year olds), 5-6 July 1980 (yearlings). Animals were not observed in May due to poor road conditions in 1979 and accumulation of snow on the study area in 1980. Between sampling periods, the animals were kept in Fort Collins, and fed alfalfa hay and grain ad libitum. One week prior to each winter sampling period, the amount of hay and grain was reduced in an attempt to increase the animals' appetite for natural forage. Similarly, during sampling periods, food intake was limited to the forage ingested in the field, and a small quantity of grain, raisins, and in winter, alfalfa hay. Whether or not these procedures affected diet preferences of mountain goats is uncertain. Previous workers found that artificial supplementation had no significant effects on torage choices of tame pronghorn antelope and mule deer (Schwartz and Nagy 1973; Regelin et al. 1976, Bartmann et al. 1982). One day prior to each sampling period, animals were transported to the study area and allowed to graze freely. Familiarity of tamed animals with available forage is considered to be an important factor in the use of tame animals to estimate diets of their wild counterparts (Bartmann et al. 1982, Bartmann and Carpenter 1982). Following this pretrial period, daily grazing trials were conducted. During each sampling period, I attempted to observe 3,000 bites per animal (discussed later). Each animal usually was observed once per day for a 1-2 hour period. If a goat ate little in its' initial grazing trial, it would sometimes be grazed a second time. For each animal, the number of days per sampling period required to obtain. sufficient bites varied from 2 to 4, depending on weather conditions and animal eating rate. Total observation time for grazing trials was 159 hours. I initiated grazing trials during the first winter and both summers at the same point in the study areas. However, starting points were alternated on a daily basis between 2 locations during the second winter to minimize grazing impact at starting points since 10 experimental animals (7 mountain goats, 3 bighorn sheep) fed in these areas. �232 The animals were normally allowed to feed and wander freely. During a 1-2 hour grazing trial a hungry animal would wander little. When not interested in feeding however, animals occasionally wandered aimlessly. In this case they would be returned to a central portion of the study area and if the animal continued to wander, the grazing trial would be terminated. I usually had only one animal in the field at a time. This prevented individuals from influencing each other's diet choices. During winter grazing trials, however, there were times when animals were restless and would not eat. Their interest in feeding was often restored by bringing a second goat along during a trial as a companion. In January, March, and Jllly 1980 only 6 mountain goats completed a grazing trial sequence. These failures were caused by insufficient time to observe 7 animals and nervous animals who ate little. As each animal grazed, I documented on a tape recorder the number of bites of each plant species consumed, and a description of plant parts selected. A bite was defined as each separate removal of plant tissue. Cues helpful in identifying a bite i~cluded jerking of the animals head and the sound of the severing vegetation. Distance from observer to animal varied, but most often was less than 2 m. Comparative Nutritional Ecology Comparative analyses were based on diets of simultaneously observed tame mountain goats and tame bighorn sheep. Three goats and 5 sheep were observed during the first 9 sampling periods previously described. The 3 goats that began trials as kids in November 1978 participated in all sampling periods. Because of differences in diets between kids and yearlings observed in the second winter (November 1979-March 1980), these kids were not included in the comparative sheep-goat analyses. Thus, from November 1978 to September 1979, the 3 goats (Table 5; Lanea, Shava, Oreo) and 3 sheep (2 ewes, 1 ram) used were all the same age. Prior to the second winter, the intact ram became untractable, and was replaced by 2 castrated 2-year-old rams. One was used in November 1979, and the other in January 1980 and March 1980. Thus, diet comparisons during each month of winter 1979-80 involved 3 yearling goats, 2 yearling ewes, and one 2-year-old ram. All animals received the same pre-trial treatment, and grazed at the same location. Forage and Diet Quality I tried to collect forage samples for all taxa contributing 2% or more of total bites ob£erved in each sampling period. Due to incomplete collections and the high diversity of goat diets, the plants collected comprised only 79% of total bites recorded during the study. Among sampling periods the plants collected comprised as little as 43% of the diets in November 1978 and as much as 90% of diets in March 1980. �233 Plant material was collected in plastic bags, put on ice, and later frozen in Fort Collins. Mean bite weights for each species were estimated by hand-plucking 25 sample "bites" of each forage, drying the samples at 100 C for 24 hours and weighing the entire sample to the nearest ± 0.0019. Forage quality samples were dried at 55 C and ground in a Wiley Mill using a 1 mm-mesh screen. Forage crude protein (Kjeldahl N x 6.25), ash, and dry matter were determined by procedures described by AOAC (1965). Cell wall constituents (CWC), acid detergent fiber (ADF), and acid detergent lignin. were determined by the procedures summarized by Colburn and Evans (1967) and Bailey and Ulyatt (1970). Analyses were performed sequentially on single samples, as recommended by Robbins et al. (1975). In vitro digestible dry matter (IVDDM) was estimated by the methods of Tilley and Terry (1963) and Pearson (1970). Two in vitro trials were conducted; 1 each for summer and winter samples. Mean bite weights were used for estimates of percent dry weight of each plant taxon in the diets. The proportion of each taxon in the diet was normalized to sum to 100%, since bite weights were not estimated for all taxon. Diet quality constituents were calculated with formulae used by Hobbs et al. (1979). For example, diet IVDDM was calculated as follows: DIG. JP i 2: (DDM. x W.. ) lp lJP l,k p where DIGj? is digestibility of diet of animal j during grazing trial p, DDM. is dlgestibility of plant taxon i during grazing trial p, Wijp = per~gnt by weight of plant taxon i in diet of animal j during grazlng trial p, and kp is number of plant taxon collected during grazing trial p. Rumen inocula were obtained from a fistulated cow maintained on alfalfa hay prior to summer trial collections and native grass hay prior to winter trial collections. The inoculum source may influence IVDDM results (Robbins et al. 1975, Palmer et al. 1976, Oldemyer et al. 1977, Bryant et al. 1980, Person et al. 1980b, Milchunas and Baker 1981). Several workers have concluded that animals used as an inoculum-source should be fed a diet mix comparable in nutritional value to that of the forage being analyzed (Pearson 1970, Grant et al. 1974, Person et al. 1980b, Trudell et al. 1980). This is important since diet nitrogen concentration affects the activity of fiber digesting bacteria (Van Gylswyk 1970, Schwartz and Gilchrist 1975). In this study, diets of inoculum donor cows contained nitrogen concentrations similar to that of the forages analyzed. The in vitro values from this experiment were not correlated with ~n vivo digestion values, and hence by themselves they can only be used in a relative comparison of forages, and extrapolation to animals on the range should be made cautiously (Tilley and Terry 1963, Milchunas 1977:7, Burns 1978). Forage and diet chemical constituents and IVDDM levels obtained in this study may be unrealistically low since workers have �234 found that plant samples selected by animals were more nutritious than those collected by hand (Weir and Torell 1959, Bredon et ale 1967, Rao et ale 1973, Ishizaki et ale 1981). Additionally, IVDDM may be underestimated due to inhibitory effects of toxic plant compounds. In the in vitro (i.e., closed) system toxins may accumulate and adversely affect rumen microorganisms (Person et ale 1980a, Turdell et ale 1980). Statistical Analysis Diet composition was calculated on the basis of percentages of total observed bites. Confidence intervals on diet percentages were calculated as ~ intervals. Use of ~ intervals is permissible because number of bites observed per animal during each sampling period was large (about 3,000); this percentages can be treated as normally distributed variates (Simpson et ale 1960:156, Hobbs et ale 1979:49). ~ercentages were not transformed; Butchner and Kemp (1974, cited by Hobbs et ale 1981) found no difference in power between analysis of variance using transformed and untransformed data, and no effect of unequal numbers of observations among percentages. Similarly, Hobbs et ale (1981) found that arc-sine transformation did not alter estimates of the variance beyond the third decimal place. Differences in diet botanical composition and diet qua.lity between age classes of animals were evaluated with simple and paired ~ tests. Unless indicated otherwise, all differences are for 2-tailed tests. Some t values were calculated with variances pooled across winter months; equality of variances (p < 0.05) was tested by procedures of Dixon and Massey (1951:90). Dietary overlap was estimated as the percent dry weight of principal plant taxa (those collected for nutrient analysis) common to sheep and goat diets (Kulcynski's overlap index, Oosting 1956:77). These principal taxa,making up 35 of 61 taxa eaten by goats, comprised 80% of total bites for goats and 82% for sheep. Differences in diets were analyzed with a factorial analysis of variance for a repeated measures design repeating over 6 (winter) and 3 (summer) months with individual animals as replicates, and animal species and months as factors of interest. Diet botanical composition in this analysis was estimated by bite counts. Differences in forage chemical constituents between winters were analyzed with paired ~ tests; comparisons of slopes and intercepts were made with small-sample ~ tests. Seasonal trends in chemical content of diets and forages were examined with simple linear and polynomial regressions. Differences differences in diet components (e.g., an increase indicated as "% points" refer to absolute from 10 to 15% is a 5% point increase). �235 RESULTS Tame and Wild Mountain Goat Comparisons Wild mountain goats were located and followed continuously through the daylight hours during a 2-week period from July 14-25, 1980. Wild goats would tolerate observers to within 2-3 m before retreating or displaying threatening behaviors. Beyond 3 m goats displayed few overt avoidance behaviors to the observers. But diet choices of wild mountain goats could not be identified to species and so bites of individual plant parts of each species could not be quantified accurately enough to compare with tame goat forage selections. The essential assumption of the tame-wild goat food comparisons was that wild goats could be approached closely enough to ohtain data comparable to tame goat forage selection data. This was not possible so the remainder of the comparative experiment was discontinued. The time which had been programmed for these activities was reallocated to a thorough analysis and reporting of tame mountain goat forage selection data from the preceding year. A final report of those activities follows. Composition and Quality of Tame Mountain Goat Diets Forage Quality Forage quality varied substantially between summer and winter. During summer the order of CWC levels among forage classes was graminoids > woody plants> forbs (Table 1). Hemicellulose (CWC-ADF) and cellulose (ADFLignin) levels were highest for graminoids and about equal for forbs and woody plants. Lignin levels were substantially higher for woody plants and similar for graminoids and forbs. Forbs and graminoids were more digestible than woody plants. The order of crude-protein concentrations was forbs > woody plants> graminoids. During winter CWC levels were lower in forbs and woody plants than in graminoids (Table 1). Graminoids were more digestible than forbs; forbs contained more IVDDM than woody plants. Lignin levels were substantially higher for woody plants, intermediate for forbs and low for graminoids. The order of crude-protein concentrations in winter was woody plants > forbs graminoids. P'rct e Ln , IVDDM, and CWC levels were greater for Salix stems than for leaves. All values during winter were similar both years (P < 0.05). .'> During plant maturation and dormancy in late summer and winter I observed reductions in digestibility and protein concentrations; concurrently, lignin and CWC increased. The chemical composition of forbs, graminoids, and woody plants showed different patterns of change with advancing season. Temporal changes in nutrient quality of woody plants were �Table l. Structural composition, crude protein content, and in vitro digestibility of forage species consumed by tame mountain goats in alpine tundra, Colorado. Season- Samples Year a (N) Taxon CWC5 X SE Percentage of dry matter ADFc Lignin Crude Erotein X SE X SE SE X IVDDMd X SE FORBS Arenaria fend1eri W-1 W-2 2 3 66 64 A. obtusiloba W-1 W-2 1 2 63 70 3.0 2.4 44 37 0.4 33 41 2.8 1.4 7.1 7.8 0.5 1.1 7.2 6.3 2.4 0.4 46 52 1.1 4.4 0.1 7.6 23 1.2 8.1 6.2 0.6 33 27 1.2 7.6 0.5 66 3.4 8.1 6.6 14 0.5 0.5 3.1 51 51 66 2.9 2.0 3.4 N e ·3 45 1.6 30 1.9 9.7 1.2 3 3 3 52 54 36 4.3 3.1 3.8 40 39 23 2.1 3.1 3.6 10 13 8.3 0.4 1.3 0.4 S 1 43 30 7.3 17 59 Cerastium arvense S 1 52 23 7.5 14 55 Erysimum niva1e S 1 22 16 3.8 27 78 Erigeron simE1ex W7-2 3 47 1.0 29 1.0 8.2 1.1 6.2 0.2 55 1.8 Geum rossii W-1 W-2 S 2 3 1 30 35 17 0.6 1.3 22 21 12 0.3 0.9 6.7 6.2 2.8 1.0 0.9 4.5 5.0 20 0.4 0.5 34 35 55 0.8 1.2 Artemisia spp. W-2 Campanu1a rot_u_nd:i._folia W-1 W-2 S Castilleja spp. f -------------------------------------------~---------------------------------------------------------------- w 0"> �Table 1. (continued) Season- Samples a Year (N) Taxon X CWCb SE ADFc X Percentage of dry matter Lignin Crude protein SE X SE X SE Heuchera parvifo1ia W-1 W-2 S 1 1 1 27 38 39 21 26 20 6.2 11· 7.7 Lewisia pygmaea S 1 26 19 11 Mertensia spp.g W-2 S 1 4 46 36 2.2 28 22 2.0 6.1 5.4 30 32 31 6.1 4.9 8.5 69 24 0.7 7.2 18 IVDDMu SE X 4.8 57 60 4.5 tv W W-l W-2 S 1 2 1 49 48 57 Oxyria dygina S 1 40 Paronychia pu1vinata W-2 2 56 Pedicu1aris parryi W-2 1 65 Phace1ia sericea S 3 42 Po1emonium viscosum W-2 S 1 1 53 44 Po1ygonum bistortoides W-l W-2 S 1 2 4 44 46 38 Oreoxis a1pina - -----_._--- 0.6 30 32 32 0.1 39 2.3 24 0.8 36 30 2.0 2.9 28 30 20 27 9.7 5.5 4.6 9.3 7.9 0.1 8.4 0.8 14 0.4 6.8 8.7 19 2.3 28 4.1 46 1.9 5.3 7.1 1.0 0.9 57 63 59 54 9.7 9.6 8.3 0.1 1.1 5.1 .6.6 8.2 15 11 39 4.4 2.7 11 24 5.3 6.6 '10 0.1 -..,J 58 6.7 53 53 0.2 4.0 46 48 53 ------------------------------------------------------------------------------------------------------------ 0.6 4.3 �(continued) Table 1. Taxon Season- Samples a (N) Year CWCb SE X 6.8 Percentage of dry matter c ADF Lignin Crude protein X SE X SE X SE Potenti11a diversifo1ia S 4 41 Primu1a angustifo1ia S 1 33 25 14 18 58 Sedum 'stenopeta1um W-2 1 28 19 10 12 64 Senecio canus W-1 W-2 1 2 43 43 6.6 22 33 33 4.6 5.0 5.8 5.1 8.8 1.8 IVDDMd X SE 1.4 16 6.0 7.6 3.7 0.4 60 61 58 4.5 0.1 N w W-2 S 1 3 44 38 W-1 W-2 S 1 3 4 T. nanum S !.~i S X W-1 W-2 S Solida~ spathu1ata Trifolium dasyphy11um Forbs 4.2 4.9 13 11 20 0.3 3.2 44 57 66 4.6 1.4 1.6 17 3.7 56 3.1 7.0 1.7 22 3.8 67 3.6 7.5 12 7.6 1.0 1.4 0.8 7.2 7.2 16 0.8 0.5 1.2 45 50 59 3.6 3.0 2.2 0.5 14 8.6 5.5 1.0 0.5 1.7 8.8 18 1.5 32 32 22 2.6 1.6 1.1 2.2 54 46 38 1.9 1.8 4 32 4 9 17 19 (X) 56 66 3.5 14 32 25 0.3 18 12 39 29 21 1.8 0.6 4.0 22 35 3.1 48 49 37 4.4 2.6 2.1 ------------------------------------------------------------------------------------------------------------ �Table 1. (continued) Taxon Season- Samples a Year (N) CWCb Percentage of drz matter ------------:::-:0 Lignin .Crude Erotein IVDDM SE X SE X SE SE X ADFc X SE X GRAMINOIDS AgroEyron scribneri W-1 W-2 S 2 3 4 72 68 68 2.0 1.3 1.5 41 37 33 1.4 1.4 1.6 2.7 5.2 4.2 0.2 0.7 1.2 7.6 8.8 14 1.1 003 4.6 59 62 65 0.5 1.6 3.5 Ca1amagrostis purpurascens W-1 W-2 3 3 73 76 1.1 1.0 Lf3 1.5 1.8 3.3 5.3 0.4 0.5 6.3 5.6 1.2 0.8 55 52 1.3 2.1 44 Carex rUEestris W-1 W-2 2 3 68 70 0.2 0.2 36 37 2.1 1.5 4.0 5.3 0.5 0.8 7.0 6.0 0.4 0.8 59 57 0.8 0.6 Carex spp. S 3 67 0.4 33 2.0 4.4 1.0 9.0 2.2 53 3.0 Festuca brachYEhz11a W-1 1 77 Kobresia myosuroides W-1 W-2 3 2 68 73 Poa spp. S 1 66 33 6.7 13 57 Trisetum ~icatum S 1 61 33 8.7 12 62 X Graminoids W-1 W-2 S 5 4 4 72 72 66 46 1.4 2.7 1.7 1.8 1.6 36 37 40 39 33 3.1 3.5 1.3 2.0 1.8 1.5 4.0 3.1 3.4 4.7 6.0 4.7 1.5 0.3 0.3 0.5 1.1 8.0 7.4 6.7 7.0 12 50 0.1 0.2 0.6 0.7 1.1 ------------------------------------------------------------------------------------------------------------ 54 54 55 56 59 0.9 3.5 1.7 2.2 2.7 N W \0 �Table 1. (continued) Taxon Season- Samples a (N) Year 6 CWC X Percentage of dry matter Lignin Crude Erotein SE X SE SE X ADFc X SE IVDDMd X SE ~.;roODY PLANTS W-l W-2 1 1 64 54 49 38 33 23 S 1 36 24 13 W-l W-2 1 2 34 44 0.8 42 31 0.6 14 17 0.4 6.4 5.2 1.2 31 38 5.6 W-l W-2 1 2 49 54 3.2 38 40 1.8 20 23 0.5 8.5 7.8 0.8 34 34 0.7 Vaccinium caesEitosium S 1 65 X Woody plants W-l W-2 S 3 3 2 49 51 50 Dr~ octoEetala Salix nivale S. spp. S. spp. h h leaves woody 45 8.7 3.3 14 43 36 34 ~-l = winter of 1978-79, W-2 = winter of 1979-80, S July 1980. bCe11-wall constituents. cAcid detergent fiber. dIn vitro digestible dry matter. ;Includes A. scoEulorum and ~. arctica. ~. occidentalis and f. Euberula. gMostly M. viridis with some M. ciliata. hMostly :[. E1anifolia with so;e ~. brachycarEa. 9.9 8.1 23 25 3.7 2.7 10 22 21 19 13 26 50 22 8.7 5.6 2.0 6.0 8.3 7.0 16 1.0 0.9 7.2 26 33 36 early and late August and September 1979 and 6.6 3.5 14 N .j:'0 �241 difficult to assess since the 4 taxa analyzed occurred irregularly in goat diets; thus discussion of woody plant changes is less thorough than in graminoids and forbs. Reductions in digestibility during summer were more rapid for graminoids than for forbs (Fig. 3). However, for those taxa in each forage class that were common in analyses for July and September, the magnitude of the decrease in digestibility for graminoids (Agr~~ scribneri) and forbs differed by only 1%. Digestibility during winter increased slightly for graminoids and decreased for forbs (Fig. 1). Salix leaves and catkins showed a 38% increase in digestibility from November to March. Increases in CWC during summer were greater in forbs than in graminoids (Fig. 4). During winter, changes in CWC levels of forbs and graminoids (taxa common in analyses of all 3 months) were small (difference of 1% in January). CWC levels changed little for Salix during winter. Crude protein levels during summer declined sharply in forbs and graminoids (Fig. 5). From July to September, protein levels decreased 64% in forbs (those taxa common in both month's analyses) and 47% in Agropyron scribneri. From early to late winter protein concentrations increased in forbs and decreased in graminoids (Fig. 5). As winter progressed, crude protein concentrations increased in Dryas octopeta1a and in Salix leaves, catkins and stems. Chemical composition (IVDDM, CWC, crude protein) of graminoids and forbs did not change significantly (p = 0.05) from the first to the second winter. Diet Composition I observed 160,254 bites of forage intake of 7 mountain goats. Seventysix plant species were eaten, including 55 forbs, 10 graminoids, 8 woody plants, and 3 cryptogams (Appendix II). The mean number of plant taxa eaten per sampling period varied from 23 in November 1978 to 37 in late August. During all sampling periods, the average proportion of bites of unidentified plants was 2.5%. Calculated over all sampling periods, forbs accounted for 69% of diets, graminoids 23%, and woody plants 8%. Forage class composition of diets varied by month and season (Fig. 6). During both winters forbs dominated goat diets, accounting for 50% of observed bites. Graminoids constituted 38% of winter diets and woody plants 12%. Despite the general importance of forbs in winter, the most frequently chosen individual forages were Carex rupestris (20% of diets) and Salix (11%). Other important forages were Campanu1a, Ca1amagrostis, Trifolium, Kobresia and Geum (Tables 2, 3). Forbs contributed 92% of the observed bites during summer. Graminoids constituted 6% of the diets and woody plants 2%. Important forages were Trifolium nanum, !. dasyphy11um, !. parryi, Po1ygonum, and Mertensia (Tables 4, 5). Agropyron scribneri accounted for most of the graminoids eaten. �242 DIGESTIBILITY 75 70 65 -~ o 60 GRAMINOIDS 55 50 45~~--~~~~----~2~2~--~17~--~14 NOV JAN MAR Figure 3. In vitro digestible dry matter (IVDDM) for graminoids and forbs eaten by mountain goats in alpine tundra, Colorado. Includes all taxa collected each month. Regression equation for graminoids is y = 84.6 - 0.52X + 0.003X2 - 0.0000053(R2 = 0.44, p = 0.025, SE of estimate = 4.8); for forbs y = 73.3 - 0.20X + 0.0004X2(R2 = 0.34, p = 0.001, SE of estimate = 9.9). Independent variable is days where 7 June = day 1. �243 CELL WALL -- 60 ~ 0 _J _J ~ _J _J 55 FORBS ".---, /' ",. 50 /' / w U 40 / /' / / / / / / / / / "<, "" " // / /' 30 5 2 30 27 22 17 JUL AUG AUG SEP 14 NOV JAN MAR Figure 4. Cell-wall content of graminoids and forbs eaten by mountain goats in alpine tundra, Colorado. Includes'all taxa collected each month. Regression equation for graminoids is y = 70.0 - 0.04X + 0.0005X2 - 0.000001X3 (R2 = 0.36, p = 0.05, SE of estimate = 3.3); for forbs y = 30.9 - 0.007X + 0.001X2 0.000004X3 (R2 = 0.40, p = 0.001, SE of estimate = 8.9). Independent variable is days where 7 June = day 1. �244 PROTEIN 26 24 \ \ \ \ \ \ - - 20 ~ 0 z 18 •••• 0 16 w r:r Q. w 0 :::> r:r (.) \ \ \FORBS \ \ \ \ \ \ \ \ \ \ \. -, -, <, <, ~ ...•.... _--.I 5 2 30 27 JUL AUG AUG SEP 22 NOV 17 JAN 14 MAR Figure 5. Pr~tein content of graminoids and forbs eaten by mountain goats in alpine tundra, Colorado. Includes'all taxa collected each month. Regression equation for graminoids is .y = 30.6 - 0.34X + 0.002X2 - 0.000003X3 (R2 = 0.78, p = 0.001, SE of estimate = 2.4); for forbs y = 34.3 - 0.33X + 0.001X2 0.000002X3 (R2 = 0.86, p = 0.001, SE of estimate = 2.8). Independent variable is days where 7 June = day 1. �tlLJ o Z N FORBS tZ .l:V! lLJ (.) a:: lLJ Q_ o ~~la~111'11III!.I_Jl~", AUG MAR NOV ~«"".'.'.'.'."'·········1···· JAN JUL 1-7 1-3 "",,,,iW#1il@Wri]@!UMbW;;;m;,;;;J AUG SEP 28-30 25-28 Figure 6. Seasonal diets by forage class of tame mountain goats during 1978-1.980 in Colorado alpine tundra. Where one month was sampled for 2 years (November, January, February), means for the month were used. �246 Table 2. Mean percentage of observed bites of principal plant taxa in diets of 3 tame kid mountain goats on alpine tundra in Colorado during winter 1978-79. Taxon a November January March All months FORBS Trifolium dasyphyllum Campanula rotundifolia Oreoxis alpina Geum rossii Arenaria obtusiloba A. fendleri Senecio canus 2.8(± 3.6)b l1.l(±lO .4)c 2.0(± 2.8) 24.2(±24.5) 1!'9(± 3.2) 4.7(± 8.8) 4.7(± 4.4) l6.4(± 9.2) 4.l(± 3.7) 7.5(± 3.4) 4.0(± 6.1) 9.2(±10.5) 6.2(± 4.4) 10.3(± 1.2) 6.3(± 5.9) 6.3 (± 3.3) trd 5.4(± 4.1) tr 3.4(± 6.8) 2.6(± 1.6) tr 6.3(± 9.8) 3.l(± 0.7) 4.4(± 9.4) 3.7(± 4.6) 3.5(± 1.5) 2.2(± 5.6) l2.2(±12.2) 22.4(±37.6) 9.5(±12.l) 14.7(±20.5) l5.2(± 2.4) l2.4(±14.4) 3.4(± 3.3) 9.5(± 6.6) l5.8(±1!.1) 8.5C± 7.3) 2.5(± 2.5) 8.0(± 2.9) 9.l(± 8.2) 4.0(± 5.9) 7.0(± 9.0) 6.5(± 3.6) 5.5(± 1.2) 2.7(± 1.9) 1.l(± 2.0) 2.7(± 2.6) 37.8(± 4.0)- 44.6(±25.9) 72.7(±20.2) 54.2(±18.4) 47.3(± 2.0) 47.7(±20.3) 19.1(± 9.1) 36.1(±12.2) 14.9(±15.5) 7.7(± 5.6) 8.2(±11.1) 9.7(± 5.5) 7,193 9,950 GRAMINOIDS Carex rupestris Calamagrostis purpurascens Kobresia myosuroides WOODY PLANTS Salix spp.e leaves, catkins Salix spp.e stems Forbs f . Ld s f GramlnOl Woody plants f Total bites 11,495 28,638 aTaxa include those contributing 2% or more to total'winter diet. b90% confidence intervals calculated across 3 animals. cMeans and 90% confidence intervals calculated across 3 animals' diet percentages, where percentages were pooled for each animal over all months. d Trace = <1%. eDiets consisted mostly of S. planifolia with some ~. brachycarpa. f Includes all taxa eaten, not just those in Table. �247 Table 3. Mean percentages of observed bites of principal plant taxa in diets of 6-7a tame kid and yearling mountain goats on alpine tundra in Colorado during winter 1979-80. b Taxon November January March All months FORBS CamEanula rotundifolia Geum rossii Trifolium dasY2hyllum Arenaria obtusiloba A. fendleri Paronychia 2ulvinata Artemisia spp.g Solidago s2athulata e l3.l(± 5.7)c 4.l(± 4.4) 10.8(± 5.5) 3.9(± 1.9) 4.2(± 2.0) 6.5(± 3.8) 8.5(± 3.8)d 5.0(± 2.6) 9.4 4.8 2.2(± 1.5) 6.2(± 2.7) 5.8(± 3.7) 4.9(± 1.6) 4.7 f tr 2.6(± 1.8) 8.8(± 7.4) 4.3(± 2.9) 3.4(± 1.3) 2.7(± 3.2) 3.8(± 3.4) 3.7(± 1.6) 4.5 3.2 2.7(± 1.7) 6.7 (± 6.3) 1.7(± 1.5) 1.8(± 1.7) 2.2(± 1.4) 2.7 (± 3.3) 2.2(± 1.4) 3.1 2.2 5.5(± 5.5) 2.4(± 3.0) 2.0 tr tr l5.2(±1l.8) l7.2(± 8.7) 36.2(±16.5) 22.3(±13.9) 22.6 7.0(± 5.2) l4.4(±10.6) 1.6(± 0.9) 8.3(± 3.9) 7.8 5.8 (± 5.9) 10.3(± 6.4) 1.2(± 0.9) 6.3(± 5.3) 5.8 tr 9.3(± 6.4) l1.2(± 8.7) 11.2 GRAMINOIDS Carex rU2estris Calamagrostis EurEurascens Kobresia myosuroides WOODY PLANTS Salix spp.h 24.4(±13.7) leaves, catkins Forbs i Graminoidsi Woody plants Total bites a i 43.0(±10.8) 52.3(±21.4) 46.8(±12.6) 47.l(±17.5) 47.5 30.5 (±21.l) 45.0(±17.5) 41.6(±17.2) 39.9 (±26.7) 39.0 26.5(±13.5) 2.7(± 1.6) l1.6(± 6.8) 13.0(± 8.5) 13.5 18,743 46,961 58,123 19,322 19,825 Number of goats observed was 7 in November and 6 each in January and M"irch. bTaxa include those contributing 2% or more to total winter diet. c90% confidence intervals calculated across animals observed. dMean and 90% confidence intervals calculated across 5 animals' (those observed in all months) diet percentages, where percentages were pooled for each animal over all months. eMean across all animals observed during trials. fTrace = <1%. gIncludes !. arctica and !. scoEulorum. ~Diets consisted mostly of ~. Elanifolia with some S. brachycarEa. ~Includes all taxa eaten, not just those in Table. �248 Table 4. Mean percentage of observed bites of principal plant taxa in· diets of 3 tame yearling mountain goats on alpine tundra in Colorado during summer 1979. Taxon August 1-3 a August 28-30 September 25-28 All months FORBS Polygonum bistortoides Trifolium dasYI~hyl1um T. :earryi d Mertensia spp. Trifolium nanum Cam:eanula rotundifolia Phacelia sericea Potentilla diversifo1ia Heuchera :earvifo1ia f Castilleja spp. 21.2(±31.5) 22.8(±13.5) l4.9(±20.4) l7.0(±16.6) 9.2(± 2.3) 16.3 (±41.8) 9.8(±13.8) 10.2(± 8.2) 13.0(± 6.3) 6.6(±22.6) 2.4(± 6.4) 3.0 (± 5.7) 4.7(± 3.9) 12.4(± 8.8(± 6.2(± 3.4(± 16.4(±13.6)c 7.5) 6.6) 2.5) 8.2) 12.3(±12.9) 11.9(± 8.0) 9.6(± 2.8) 8.6(±20.8) 9.6(±12.7) 2.9(± 3.0) 4.6(± 6.5) 8.0(±17.2) 5.7 (± 7.7) 4.6(± 7.9) 2.4(± 4.3) 2.6(± 6.0) 3.0(± 2.0) 2.6(± 2.6) e tr 1.6(± 1.2) tr 4.0(± 2.6) 7.1(± 1.9) 1.8(± 1.4) 2.6(± 0.8) 2.5 (± 2.1) GRAMINOIDS Agro:eyron scribneri Forbsg tr tr 8.5(± 6.0) 3.0(± 2.7) 98.0(± 1.6) 93.5(± 8.6) 83.9(± 9.8) 91.8(± 2.1) tr 2.8(± 5.9) l3.8(±10.5) 5.8(± 4.8) 1.l(± 2.5) 3.7 (± 4.3) 2.3(± 3.7) 2.4(± 2.7) 15,090 16,879 15,481 47,450 g Graminoids Woody plantsg Total bites aTaxa include those contributing 2% or more to total summer diet. b90% confidence intervals calculated across 3 animals. cMeans and 90% confidence intervals calculated across 3 animals' diet percentages, where percentages were pooled for each animal over all months. dDiets consist mostly of ~. viridis with some M. ciliata. eTrace 1%. f~. occidentalis or ~. puberula. gInc1udes all taxa eaten, not just those in Table. �249 Table 5. Percentage of observed bites of principal plant taxa in diets of 6 tame yearling and 2-year-old mountain goats on alpine tundra in Colorado during July 1980. Taxona Lanea Shava Animals Oreo Bucko Biff Pete X(±90%CI) 54.8 1.3 25.6 52.2 6.1 23.5 3.4 2.3 10.8 35.7 17.2 8.2 33.4 28.5 5.4 37.8 21.7 4.4 36.8(±16.4) 13.5(±15.6) 12.9(± 7.7) 2.5 1.5 4.6 0.2 10.3 0.8 0.8 11.8 3.7 3.5 3.2 6.4 4.2 (± 2.7) 3.9 (± 3.7) 0.1 6.9 7.7 0.6 1.8 2.3 3.4(± 2.2) a o a o FORBS Trifolium nanum T. dasyphyllum Mertensia spp. Polygonum bistortoides Geum rossii Potentilla diversifolia Solidago spathulata Trifolium parryi 1.5 o 22.2 0.4 6.5 0.8 3.5 0.7 3.4(± 7.4) 2.0(± 2.0) a 0.9 a 0.7 7.9 1.8 2.1(± 2.5) 99.5 98.9 94.1 86.4 88.0 87.5 92.4(± 4.9) 0.2 1.1 0.3 7.7 11.0 11.6 5.6(± 4.5) 0.3 a 5.6 5.9 1.0 0.9 2.1(± 2.2) 3,631 5,192 3,816 3,913 4,839 4,652 GRAMINOIDS Agropyron scribneri Forbsc Graminoidsc Woody plants Total bites c 26,043d aTaxa include those contributing 2% or more to mean diet. bDiets consisted largely of M. viridis with some M. ciliata. clncludes all taxa consumed, not just those in Table. dSum of total bites counted in July. �250 Diet Quality Diet quality, like forage quality, exhibited seasonal changes similar to those found for other ungulates (Schwartz et ale 1977, Hobbs et ale 1981, Baker and Hobbs 1982). Quality of diets was greater in summer than in winter (Table 6). Diet IVDDM, cell solubles, and crude protein all decreased from July to March (Figs. 7-9). Changes in these diet quality constituents were more pronounced during summer (July-September), than during winter (Figs. 7-9). a Table 6. Mean percentage structural composition, crude-protein content, and in vitro digestibility of diets of tame mountain goats on alpine tundra in Colorado during November 1978 - July 1980. Season Winter Summer (1979) July (1980) CWCb Age class Kide f Yearling g Kid Yearling Percent of dry matter Crude protein ADFc Lignin f h 2-yr-old. 1 Yearling X SE X SE 57 53 60 1.1 2.1 1.9 37 34 35 0.5 1.0 0.7 7.2 12 7.4 0.3 1.2 0.5 51 0.5 23 0.2 6.9 0.4 34 35 1.1 2.3 20 21 0.5 0.6 7.6 7.1 0.2 0.2 X SE IVDDMd SE X SE 0.3 0.3 0.1 50 42 49 0.3 0.3 1.3 16 0.2 59 0.5 27 26 0.1 0.2 67 65 0.4 0.9 X 7.8 5.9 5.7 ~eans calculated across animals' diet percentages where percentages were pooled for each animal over all months of each season. bCell-wall constituents. cAcid detergent fiber. dIn vitro digestible dry matter. e 3 kids observed in winter 1978-79. £3 yearlings observed in summer 1979 and winter 1979-80. g3 kids observed during winter 1979-80 who did not miss a sampling period. h3 2-year-olds observed. i3 yearlings observed. �251 DIET IVDDM o I y= 60.8 R2= 0.78 + 0.34X -0.0IX2+ 3.8X3_ 6.2X4 t P < 0.001 • -- • ~ o •• ~ c c > o ~ o o o o 01~--~--~--~1--~------~1------~1------~1 2 30 27 JUL AUG AUG SEP 5 22 NOV 17 JAN 14 MAR Figure 7. Digestibility of diets of tame mountain goa~s in alpine tundra, Colorado. Symbol. = kids observed during November 1978March 1979, 0 = same animals as yearlings-during August 1979March 1980, 0 = same animals in July 1980, • = kids' observed during November 1979-March 1980, and. = same animals as yearlings in July 1980. Independent variable is days where 7 June = day 1. �252 Figure 8. Cell-soluble content of diets of tame mountain goats in alpine tundra, Colorado. Symbol. = kids observed during November 1978-March 1979, 0 = same animals as yearlings during August 1979-March 1980, 0 = same animals in July 1980, • = kids observed during November 1979-March 1980, and • = same animals as yearlings in July 1980. Independent variable is days where 7 June = day 1. �253 30 DIET PROTEIN 2 3 y= 35.3 - 0.33X+O.00IX -1.2X R2= 0.98, P( 0.001 25 - ~ 0 z 20 w ~ 0 0::: CL w 0 ::::> 15 0::: (.) 10 • • 5 Or~--~I~~~~~~ 5 2 30 27 JUL AUG AUG SEP -LI ~IL_ ~l 22 NOV 17 JAN 14 MAR DATE Figure 9. Protein content of diets of tame mountain goats in alpine tundra, Colorado. Symbol. = kids observed during November 1978-March 1979, 0 = same animals as yearlings during August 1979-March 1980, 0 = same animals in July 1980, • = kids observed during November 1979-March 1980, and. = same animals as yearlings in July 1980. Independent variable is days where 7 June = day 1. �254 Temporal patterns of change for each diet quality constituent were different. Through summer, diet IVDDM decreased 19%, diet cell solubles 30%, and diet crude protein 60%. From November to March, diet IVDDM decreased 3%, diet crude protein increased 6%, and diet cell solubles increased 4%. Diet cell solubles were highest in early August; diet crude protein and IVDDM peaked in July (Figs. 7-9). Reduction in diet cell solubles in January coincided with snow accumulation in Salix shrubbery, and its' subsequent reduction in animal diets. Crude-protein levels in tame goat diets were similar to those reported for wild mountain goats. Fecal crude-protein levels of wild goats varied from 22 to 16% during summer (July, August) (Hebert and Turnbull 1977, McFetridge 1977:33); values for tame goats ranged from 27 to 16%. During winter (November-April), feces from wild goats had crude-protein concentrations ranging from 12 to 7% (Hebert and Turnbull 1977, McFetridge 1977:33); values for tame goats ranged from 9 to 5%. Comparative Nutritional Ecology Botanical characteristics of diets. I observed substantial differences in forage class composition of sheep and goat diets. In winter, goats ate more forbs (P < 0.05) and less graminoids (P < 0.01) than did sheep, but differences varied by month for forbs (month x species interaction, P < 0.005, Fig. 10) and graminoids (P < 0.02, Fig. 11). During summer, goats ate more forbs (P < 0.11) and less graminoids (p < 0.1) than did sheep. Species differences in forb consumption ranged from 14% points in August trials to 25% points in September, while graminoids consumption differed by 16% points during August and 27% points in September. Goats ate more browse than did sheep in winter (P < 0.005) and summer (P < 0.2). Again, differences between species in winter varied by month (month x species interaction, P < 0.04, Fig. 12). Similarities in sheep and goat diets were evident not only for forage classes (Fig. 13), but also for principal taxa within forage classes. Taxa ove~lap varied by month (Fig~ 13), but overall was greater in summer (X = 59%) than in winter (X = 45%). I also observed differences between sheep and goats in plant parts selected. Goats ate more leaves and flowers, and less stem material from Polygonum bistortoides, Campanula rotundifolia, and Calamagrostis purpurascens. During months when goats and sheep selected these dissimilar diets, Polygonum made up 22% of goat diets and 29% of sheep diets; Campanula comprised 10% of goat diets and 4% of sheep diets; Calamagrostis comprised 15% of goat diets and 51% of sheep diets. Nutritional characteristics of diets. I observed substantial differences between goats and sheep in all diet quality constituents except protein. Similar to diet botanical composition, differences between species were more substantial in winter than in summer. �255 70 60 - ~ 0 GOATS 50 en 00 0:: 0 u, u, 0 40 en w •.... 00 SHEEP 30 O,-~,,~--~I------~I------_I~~~~~--~I------~I------~I NOV JAN MAR 1978 1979 1979 " NOV JAN MAR 1979 1980 1980 Figure 10. Forbs in winter diets of 3 tame bighorn mountain goats in alpine tundra, Colorado. sheep and 3 tame �256 80 SHEEP - -en ~ 0 60 Q 0 z ~ « a::: (!') LL 40 0 en w •.... m GOATS 20 -~ H O~~~,--~I------~I------~I~~H--_.I------_I~----~I NOV JAN MAR NOV JAN MAR 1978 1979 1979 1979 1980 1980 Figure 11. Graminoids in winter diets of 3 tame bighorn 3 tame mountain goats in alpine tundra, Colorado. sheep and �257 -~ 0 30 LLJ en 3: 0 a:: m LL 0 20 en LLJ t- rn 10 Figure 12. Browse in winter diets of 3 tame bighorn tame mountain goats in alpine tundra, Colorado.' sheep and 3 �258 70r- OVERLAP 60150 I40 ~ o I- 301- lOr- 100- SIMILARITY 90- - 80~ o 706050~ ~~ o ~ ~,~ ~~ ~ NOV JAN MAR AUG AUG SEP NOV JAN MAR 1978 1979 1979 1979 1979 1979 1979 1980 1980 Figure 13. Estimates of congruence between sheep and goat diet choices in alpine tundra, Colorado. Top is Ku Lcynskd' s overlap index estimated as the percent dry weight of principal individual plant taxa common to sheep and goat diets (in November 1978, overlap, calculation was based on bite counts, not bite weights). Bottom is similarity in forage class composition of diets estimated as 100 - mean of sum of differences between percent bites of forbs, graminoids, and woody plants in diets·of sheep and goats, e.g., 100 - (.[% forbs in goat diets - % forbs in sheep diets] + [repeat for graminoids and woody plants] .;.3) = percent similarity .• �259 Compared to goat diets, sheep diets contained more unlignified cell wall (fiber constituents most available for microbial fermentation, estimated as CWC-lignin) in winter (P < 0.01) and summer (P < 0.1) and also, more IVDDM in winter (P < 0.001), but equal amounts in summer (P = 1.0). The size of differences between species varied by month for both constituents during winter (month x species interaction, P < 0.002, Figs. 14, 15). Goat diets contained higher levels of cell solubles in winter (p < 0.03) and'summer (P < 0.1), and also, more lignin in winter (P < 0.01) and summer (P < 0.025). Magnitude of differences during winter varied by month for cell solubles (month x species interaction P < 0.02, Fig. 16) and lignin (P < 0.004, Fig. 17). Goat diets contained greater concentrations of protein during winter (P < 0.15) and summer (P < 0.06) than did sheep diets. The size and direction of the differences between species varied by month during winter (month x species interaction, P < 0.05, Fig. 18). Plant parts selected by goats were more nutritious than those eaten by sheep. Samples of parts of Polygonum and Campanula selected by goats in August trials, had a higher mean level of cell solubles (7% points), IVDDM (4% points), and protein (3% points). Likewise, in January, the goat sample for Calamagrostis had more IVDDM (4% points) and protein (0.5% points) than did the sample collected for sheep. DISCUSSION Forage Quality The results of this study were similar to those of other alpine investigations. The decrease in forage quality that I observed from late summer to mid winter is typical (Johnston et al. 1968, Rice et al. 1971, Milchunas et al. 1978, Hobbs et al. 1981). The enhanced nutritional qualities of graminoids (IVDDM), forbs (protein), and Salix leaves (protein, cell solubles, IVDDM) in March were likely associated with translocation of nutrients to the above ground tissues. This is often accompanied by green up in vegetation (Burzlaff 1971, Hickman 1975). In this study, only Carex rupestris showed this phenological change. Cell solubles (l-CW) in forbs (e.g. Geum, Heuchera, Paronychia), and browse leaves and stems were not completely digested, as evidenced by large differences (22-54%) between their cell-soluble levels and IVDDM coefficients (Table 1). Similar results have been reported (Mi1chunas et al. 1978:8, Hobbs et al. 1981:64). These discrepancies likely result from depressed IVDDM coefficients; possible causes of this depression include: 1) less extensive rupture of cell walls (Hobbs 1979:57), 2) high ash levels (Milchunas et al. 1978:10), 3) lack of amylolytic �260 SHEEP 60 ~ ~ ~0 ~ ~ ~ ~ ~ ~ w 50 U Q W ~ z GOATS ~ ~ z ~ 40 OL-T~l----~I------~I----~~I--~~~~I------~I-------JI ~ NOV JAN MAR 1978 1979 1979 Figure 14. Unlignified cell-wall bighorn sheep and 3 tame mountain Tr NOV 1979 JAN MAR 1980 1980 levels in winter diets ~f 3 tame goats in alpine tundra, Colorado. �261 55 SHEEP O~~,~~I , ~I ~I__ ~H~~J NOV JAN MAR 1978 1979 1979 H ~I~ ~I NOV JAN MAR 1979 1980 1980 Figure 15. IVDDM levels in winter diets of 3 tame bighorn and 3 tame mountain goats in alpine tundra, Colorado. sheep �262 55 GOATS 50 - 45 ~ 0 en w _J CD ::::> _J 40 0 en _J _J w (.) 35 SHEEP 30 25 0~~--~1------~1----~1~~7~~~~1~----~1------~1 NOV JAN MAR NOV JAN MAR 1978 1979 1979 1979,. 1980 1980 Figure 16. Cell~soluble levels in winter diets·of 3 tame. bighorn sheep and 3 tame mountain goats in alpine tundra, Colorado. �263 14 12 GOATS 10 - ~ o z z 8 (.!) _J 6 SHEEP " L~<--~'------~'------~'--~'4'~~'------~'------~! ..-- NOV JAN MAR NOV JAN MAR 1978 1979 1979 1979 1980 1980 Figure 17. Lignin levels in winter diets of 3 tame bighorn and J tame mountain goats in alpine tundra, Colorado. sheep �264 9 8 - -z ~ 0 7 w ••••• 0 a::: a. w 0 ~ 6 a::: u oL I I I NOV JAN 1978 MAR 1979 1979 Figure 18. Crude-protein sheep and 3 tame mountain .,,, H I I I NOV JAN 1979 MAR 1980 1980 ! levels in winter diets of 3 tame bighorn goats in alpine tundra, Colorado. �265 bacteria in rumen fluid from grass-fed (rich in cellulolytic bacteria) inoculum donor, and 4) inhibitory effects of plant secondary metabolites (Mould 1980:63, Person et al. 1980b, Trudell et al. 1980). Knowledge of the effects of plant secondary metabolites on wild ruminants has been limited largely to volatile oils and terpenes (Nagy et al. 1964, Oh et al. 1967, Radwan and Crouch 1974). Recent studies have examined other secondary metabolites (e.g., phenolics, alkoloids) and their effects on forage preferences (Kuropat and Bryant 1980), analytical procedures for chemical analysis of forage (Mould 1980, Person et al. 1980a, Trudell et al. 1980, White and Trudell 1980), and digestive physiology (Mould 1980). The prevalence of secondary metabolites in plants (Levin 1976) and their diverse effects on animals makes a theoretical discussion of their importance in this study difficult. Generally, forbs and shrubs possess more potent chemical defenses than graminoids (Kuropat and Bryant 1980, Mould 1980, Person et al. 1980a, Trudell et al. 1980, White and Trudell 1980). Some of the genera encountered by the tame mountain goats (Hymenoxis, Epilobium, Trifolium, Senecio, Primula, Ranunculus, Salix, Dryas, Vaccinium) have been found to contain secondary metabolites (alkaloids, isoflavones, tannins, saponins, flavonoids, phenolics) (Arnold and Hill 1972, Levin 1971, Whittaker and Feeny 1971, McKey 1974, Kuropat and Bryant 1980, Mould 1980, White and Trudell 1980). Despite these reports of secondary metabolites in forage, only Mould (1980) has demonstrated their detrimental effects on wild ruminants. However, due to the generally toxic effects of secondary metabolites (Feeny 1975), their potential influence on forage quality needs to be considered in analyses of diet quality. Diet Composition Temporal variation. I observed changes in diet forage class composition among months and between seasons (Fig. 6). These differences resulted in part from variation in forage availability and forage quality (discussed later). The decrease in consumption of forbs in winter, compared to summer, could have been caused partly by intrinsic differences in floristic composition of winter and summer sites, or by differences in plant part perseverance of forbs and graminoids, or both. In summer, forbs appeared to be more abundant in the summer study area than in the winter study area. In winter, dessication and shattering of forbs contributed to their reduction in availability in the winter study area. Snow accumulation affected forage availability and influenced goat diet choices, particularly with respect to Salix. Goats ate less Salix when these shrubs were covered with crusted snow during January trials (Tables 2, 3). Adams (1981:100) reported a similar effect of snow on browse consumption by mount n goats in alpine habitat. When Salix was not covered by snow (November 1978), accumulation (8 cm) of soft snow on· forbs and graminoids appeared to be the cause of a large increase �266 in Salix consumption (9 to 19% of diets) and a decrease in consumption of forbs and graminoids. Also, when Salix was covered by crusted snow (January 1980), effects of soft snow cover on forage choices were variable. Animals pawed to expose covered forbs and grasses and did not rely extensively on plants protruding from the snow. The animals appeared to be able to sniff preferred forages through the snow, an ability apparently shared by reindeer (Nasimovich 1954, cited by Skogland 197~). Differences in forage class composition of diets (e.g., forbs, the dominant forage items) among animals increased from sununer to winter (Table 7). Using standard deviations calculated on forb consumption as a measure of food niche breadth, my findings conform to the hypothesis that niche breadth should increase as resource availability decreases (Mac Arthur and Pianka 1966, Schoener 1971, Em1en 1973, Pianka 1976). Table 7. Standard deviations of the amount of forbs in diets of 3 mountain goats in selected periods. Period August (1-3) August (28-31) September (25-28) Winter Standard deviation 0.9 5.1 5.8 1l.3a aBased on amount of forbs in diets of each animal pooled over November 1978, January 1979, and March 1979 (i.e., "all months" category, Table 8). Ricklefs (1973:215) cautioned that an increase in niche breadth usually occurs only when the pressure of interspecific competition is low. Several workers have suggested that mountain goats are separated ecologically from their competitors by living in more precipitous terrain (Flook 1964, Geist 1971:256, Adams 1981:143). With little constraint on diet choices by interspecific competitors, mountain goats could select a botanically diverse diet that optimizes the ratio of nutri~nt and energy intake to losses. Thus, during winter, when the amount of high quality food is reduced, animals cannot afford to bypass low quality forage because mean search time per high quality food item encountered is long, and frequency of encounters with high quality food items is low (Pianka 1976:118). Therefore, diverse diets of mountain goats in periods of relative food scarcity may maximize intake of nutrients and energy per unit energy expenditure. During sununer the abundance of high quality plants enables mountain goats to be more selective in diet choices; �267 high quality plants are frequently encountered with little effort. Substantiating these contentions, I found that crude-protein concentrations among forages were relatively more uniform during periods of abundance than during periods of scarcity (e.g., in early August, the month with the smallest standard deviation for forb consumption, there was a 19% decrement (21 to 17%) in protein content from the 3 most protein-rich forages to the 3 most protein-poor forages; the corresponding decrement was 40% in late August, 42% in September and 61% during winter, i.e., during winter the plants available to mountain goats displayed greater diversity in protein contents than in summer). Also, forage protein was higher in summer than in winter (Table 1). Consistent with the hypothesis, food niche breadth was highly correlated (r = 0.99) with the variability (or equally, negatively correlated with uniformity) in protein contents of forage plants (Fig. 19). A significant correlation (r = 0.97) between food niche breadth and variability in IVDDM coefficients of forages was also found. Age class variation. Inferences on effects of age on diet choices of animals were based on comparisons among: 1) one group of goats studied as kids the first winter versus the same animals as yearlings in the second winter, and 2) kids and yearlings observed concurrently in the second winter, and observed as yearlings and 2-year-olds in July. Comparisons of diet choices between kids and yearlings in the first instance could be confounded with year effects on forage quality and availability. However, differences in forage quality between winters were not significant (P < 0.05). Likewise, I observed few differences in forage availability between winter grazing trials for the 2 years and, indirect evidence suggests that differences in forage availability were small: few differences were evident between diets of kids observed the first winter compared to kids observed the second winter (Table 8); diet differences between kids (years) during March trials were due to large amounts of Trifolium eaten by kids in first winter (Table 2); 1 year later these same animals ate 58% less (24 to 10% of bites) Trifolium in March although availability of Trifolium between years appeared similar. Assuming that the 2 groups of kids had similar intrinsic responses to nutritional characteristics of the forage, I concluded that similarity in their diet choices indicates comparability in forage availability between years. Thus, differences in diets between kids and yearlings in the first instance, can be attributed primarily to effects of age. Comparisons of diet choices of yearlings and kids observed during the same winter might be affected by differences in snow cover between their grazing trials; however, these differences appeared to be small and of no great consequence for diet comparisons. Similar to other workers, I observed diet differences attributable to age (Langlands 1969, Bergerud 1972, Tucker et al. 1977, Willms et al. 1980). Generally, kids ate more graminoids but fewer forbs and woody plants than did yearlings. During July and January, the opportunities for consuming woody plants were minimal due to their scarcity in the summer �268 12 WINTERo· 10 :I: J- 0 8 « lJJ a:: III w I 6 o SEPTEMBER (.) o LATE AUGUST Z 0 0 0 4 LL 2 O~------~--------~------~ 15 30 45 60 VARIABILITY IN PROTEIN CONCENTRATIONS OF FORAGES Figure 19. centrations Correlation of foragesa between variability in protein and food niche breadthb con- aCa1cu1ated as the percent decrement in protein concentrations from the 3 most protein-rich forages to the most protein-poor forages for each sampling period. b Based on standard deviations of the quantity of forbs eaten by 3 mountain goats during selected period. �26.9 Table 8. Significance levels for differences between age classes of mountain goats observed Colorado. in diet composition in alpine tundra, Month Age class Forage class Graminoids Woody plants Forbs November Yearlings - kids; Kids - yearlings c Kids - kids ns ns ns January Yearlings - kids Kids - yearlings Kids - kids March Yearlings - kids Kids - yearlings Kids - kids July Yearlings d 0.01 e 0.02s ns 0.01 0.02 ns 0.05 0.05 O.Ose ns O.Ose ns ns ns ns ns ns 0.03 0.03e ns 0.03 ns ns ns 0.01 0.01 ns f - 2-year-olds a3 yearlings versus 3-4 kids, observed concurrently. bSame 3 animals as kids in first winter, c3 kids observed in second winter. dNot significantly in first winter versus different at P < yearlings in second winter. 3-4 different kids observed 0.05 level. eCalculated with variances pooled over November, January, and March; used only where it detected a significant difference (P < 0.05) not previously found with simple ~ test. f 3,2-year-old animals versus 3 yearlings, observed concurrently. study area, and unavailability in January due to snow. This precluded comparing consumption of woody plants between age classes at these times. In 6 of 7 comparisons, kids ate mqre graminoids than did yearlings; conversely, yearlings ate more forbs than did kids. In each of 4 comparisons, yearlings ate more woody plants than did kids (January and July disregarded). In 11 of these 18 comparisons, age differences in diets were significant (Table 8). The consequences for diet quality resulting from these differences in diet botanical composition are discussed later. �270 Variation in consumption of woody plants between kids and yearlings may result from differences in acquired tastes as well as from differential nutritional needs of kids and yearlings. Foraging experience affects diet choices of animals faced with unfamiliar plants (Arnold and Mauer 1977, Marten 1978:1473, Bartmann et al. 1982). This may be the cause of the variability in consumption of Salix (98% of woody plants eaten in November) between age classes in November (Table 8). Overall, however, kids ate a much smaller amount of Salix than did yearlings. Kids ate little browse during practice grazing trials in summer and fall; this was not unusual in light of the greater nutritional quality of forbs compared to browse plants (Baker and Hobbs 1982, see also Table 1). Thus, initial random encounters with Salix may have involved some "palatability testing" by the animals. Conversely, experienced animals (yearlings) probably sought the conspicuous Salix plants and thus increased the frequency of encounters with it, and its importance in their diets. Nutrition related effects are discussed later. Diet selection-forage quality associations. Nutrient concentration and levels of secondary metabolites in forage affect diet choices of herbivores (Klein 1970, White 1978, Hobbs 1979; Longhurst et al. 1968, Arnold and Hill 1972, Freeland and Janzen 1974, Kuropat and Bryant 1980, Schwartz et al. 1980). However, study of animal diets has been largely unable to develop consistent predictions of diet composition based on forage characteristics (Ivins 1952, Marten and Anderson 1975, Marten 1978). Mountain goats frequently chose forages with relatively high concentrations of crude protein. In March 1979, Trifolium dasyphyllum was zealously eaten (Table 2); its crude-protein concentration was about 2 times greater than the mean crude-protein level of the other principal forages (12.9% versus 6.6%). During summer, the increased consumption of Agropyron scribneri (Table 4) was associated with differences between the crude-protein concentration of this grass and principal dietary forbs (i.e., those forbs collected for chemical analysis). In late August, these forbs contained more crude protein (3.9% points) than did Agropyron; this grass was seldom eaten (Fig. 20). Conversely, in September Agropyron contained more crude protein (2.5% points) than did forbs; consumption of Agropyron increased substantially (Fig. 20). A similar, but less pronounced association was evident between diet botanical composition and forage IVDDM. In March, two rare species, Senecio canus and Sedum stenopetalum, were avidly eaten and relatively high in nutritional quality. Senecio was relatively succulent and had IVDDM levels that were substantially higher than other principal forbs (61% versus 52%). Sedum was succulent and possessed greater crude-protein (11.7% versus 6.9%), cell-soluble (72% versus 51%), and IVDDM (64% versus 48%) levels than did other principal forbs. In addition to selection of higly nutritious forages, mountain goats avoided plants uhat are likely to contain secondary metabolites. �271 8.5 AGROPYRONIN DIETS 7.5 6.5 5.5 4.5 3.5 ~ o 2.5 \PROTEIN' LEVELS: DIETARY \ FORBS MINUS AGROPYRON' 1.5 0.5 -0.5 -1.5 -2.5 \ AUG \\SEP \ \ \ Figure 20. Association of crude-protein levels in Agropyron and dietary forbs with consumption of Agropyron. �272 Hymenoxys grandiflora was often sniffed, sometimes snipped and expelled, but rarely eaten. Hymenoxys odorata contains sesquiterpene lactones, substances thought to be objectionable to herbivores (Levin 1976:136). The animals rarely ate Ranunculus adoneus; the genera is strongly distasteful to herbivores (Whittaker and Feeny 1971:758), probably due to its alkaloid content. Conifers, which contain volatile oils, a group of plant secondary metabolites known to be objectionable to wild ruminants (Carpenter 1976, Nagy and Regelin 1974, Schwartz et al. 1980:114), were rarely eaten by the goats. In contrast to these findings, mountain goats ate Epilobium angustifolium, a species containing high levels of tannins (Mould 1980). Tannins in plants commonly deter feeding by domestic ungulates (Wilkins et al. 1953, Cooper-Driver et al. 1977), but apparently are a less effective deterent against wild ungulates since Epilobium is often eaten by mule deer and elk (Kufeld et al. 1973, Mould 1980: 11). Selective feeding by ungulates for plant parts in addition to plant species has been recognized (Arnold 1960, Gwynne and Bell 1968, Jarman 1974, Hirst 1975, Sinclair 1977). I observed a general preference by mountain goats for flowers and leaves of forbs and grasses. Similar findings were reported by Casebeer et al. (1950), Chadwick (1973), and Hjeljord (1973). Consumption of woody parts of Salix was usually limited to distal portions of twigs adjacent to buds. During summer, preferences for plant parts appeared to be associated with plant phenophase. Consumption of flowers of Castilleja, Erysimum, Geum Mertensia, Phacelia, and Polygonum declined near (or after) anthesis. Consumption of leaves of Phacelia and Heuchera was higher in September than in August. Consumption of Castilleja and Erysimum was almost solely restricted to flowers. Consumption of young flowers by mountain goats provides nutritional benefits since they are rich in proximal nutrients (e.g., nitrogen) and contain low levels of secondary metabolites (Kuropat and Bryant 1980). Avoidance of some young leaves (Phacelia, Heuchera, Castilleja, Erysimum) is likely beneficial since young leaves have been found to contain relatively high levels of secondary metabolites (McKey 1974:312, Kuropat and Bryant 1980).· During winter mountain goats avidly ate Senecio (Table 2), a genera which contains high levels of alkaloids (Freeland and Janzen 1974). However, the animals ate only leaves; most al~aloids in Senecio are stored in roots during winter (Areshkina 1957, cited by McKey 1974:313). Tame versus wild mountain goats. Tame mountain goats chose diets largely similar in botanical composition to diets of wild mountain goats during summer. Forbs in general and Trifolium, Mertensia, Polygonum and Geum in particular were important contributors to summer diets of wild mountain goats in Colorado (Moser 1955, Hibbs 1966, Johnson et al. 1978, Thompson 1981). These findings are consonant with my observations. However, diets of tame and wild mountain goats during winter in alpine habitats �273 may be less similar. Tame mountain goats ate more forbs than did wild goats (Adams 1981), and seldom ate conifers, a major component of wild goat diets (Adams 1981:89). The importance of conifers in Adam's study is probably exaggerated· since fecal analysis overestimates its consumption (Anthony and Smith 1974). Another Colorado goat study indicated little use of conifers (Hibbs 1967). Also, forb consumption may be underestimated by fecal analysis as indicated in numerous studies (Free et al. 1970, Anthony and Smith 1974, Alexander 1979, Johnson 1980). Of the taxa important in diets of wild mountain goats that were available on Niwot Ridge, only conifers were rejected by the tame animals. Evidence for conifers as emergency food for mountain goats in winter is equivocal, some workers suggesting heavy use, others, little or none (Casebeer et a1. 1950, Brandborg 1955, Geist 1971, Adams 1981; Anderson 1940, Hamre 1947, Casebeer et al. 1950, Hanson 1950, Richardson 1971, Peck 1972, Hje1jord 1973, Kuck 1973, Trout 1973). Thus, the limited use of conifers by tame animals does not appear to be an important deviation from diet choices of wild mountain goats. Diet ,Quality Forage quality-diet choice interactions. A phenomenon of altering diet choices in response to reductions in food quality has been frequently observed for ungulates and other mammals (e.g., Mattson 1980, Hobbs et al. 1981, Hobbs et al. 1982). By altering diet choices, the observed animals were able to obtain diets of relatively stable quality. Stable levels of dietary protein appears to be especially important to the animals. I observed a similar phenomenon in tame mountain goats. During summer 1979 (August-Sept~mber), diet protein levels decreased substantially (Fig. 9). This decrease reflects a reduction in protein levels of forbs; they constituted 96% of diets and 100% of forages analyzed in August trials. The decrease in protein level of forbs, and hence diets, was ameliorated to a small degree by an increase in consumption of the relatively protein-rich grass, Agropyron scribneri (Fig. 20). Consequently, from early August to September, the decrease in diet crude protein w~s about 1% less than the protein decrease in forbs. During winter, shifts in diet choices in response to forage quality were more substantial. The amount of forbs eaten increased 48% (40-59%) from November trials to March trials; consumption of graminoids decreased 22% (39-30%) and woody plants 52% (21-10%). In March, crude protein concentrations in forbs were 1.4% points greater than in commonly eaten graminoids (Carex, Calamagrostis, Kobresia), and 1% point greater than in Salix leaves. However, graminoids contained more IVDDM than forbs or Salix leaves (56,49,44%, respectively), but lesser amounts of cell solubles than these dicots (30, 52, 62%, respectively). Thus, by eating more forbs and fewer graminoids and woody plants in March, mountain goats maintained stable dietary levels of cell solubles and protein (Figs. 8, 9), at the expense of reduced diet IVDDM. �274 The relationship between diet choice and forage quality, however, was not restricted solely to shifts among forage classes. Mountain goat diet choices appeared to respond often to nutrient concentrations of individual taxa, regardless of forage class. Increased consumption of specific forbs in March (Tables 2, 3; Sedum excluded) coincided with their high levels of protein (>11%) (e.g., Trifolium, Sedum), IVDDM (56-64%), and cell solubles (51-72%) (e.g., Oreoxis, Sedum, Senecio, Solidago). Increased consumption of Carex rupestris in March (Table 9) coincided with its high levels of IVDDM (58%) and intermediate protein concentrations (7%). This contrasts with the graminoid, Calamagrostis which declined in diets from November to March (Tables 2, 3). It had comparable levels of IVDDM (54%), but 45% less protein than Carex rupestris. This supports the premise that mountain goats emphasized protein in diets at the expense of IVDDM. The. importance of Carex rupestris in diets, with its' intermediate concentrations of protein, likely reflected its' prevalence in the study area. Mountain goats could ill afford to pass up a fairly nutritious and frequently encountered forage. Nutritional status. What are the consequences of the observed diet choices of mountain goats on their nutritional status? Diets of goats during summer were highly nutritious. The protein and energy (IVDDM) their diets provided likely met their growth requirements although these needs are unknown. Ammann et al. (1973) postulated that diets containing greater than 50% digestible energy (which is highly correlated with digestible dry matter (Robbins et al. 1975, Milchunas et al. 1978) were necessary to meet maintenance energy requirements of small ruminants. Diet IVDDM for tame goats was well above this figure. It varied from 66% in July to 54% in September (Fig. 7). Dietary protein levels of 13-17% were reported to be sufficient for optimal growth of white-tailed deer (Odocoileus virginianus) (French et al. 1956, HcEwen et al. 1957, Holter, et al. 1977). Crude-protein concentrations in mountain goat diets usually met or exceeded this requirement, varying from 27% in July to 11% in September (Fig. 9). During winter, shifts in diet choices no doubt had important effects on nutritional status of animals. As previously noted, tame mountain goats appear to emphasize protein in diets in late winter, at the expense of diet digestibility. Crude-protein levels in goat diets during winter (Table 6) were similar to maintenance protein levels reported for deer (6-,7%) (Dietz 1967, French et al. 1956) and bighorn sheep (5%) (Hebert 1973:294). Specifically, the diet botanical mix in March, resulted in a diet crude-protein concentration of 6.7%. This concentration may approximate maintenance protein needs of goats in late winter. Diet IVDDM of tame goats during winter (Table 6) was at, or below, the minimal maintenance digestible energy (i.e., IVDDM) estimate of 50% (Ammann et al. 1973). This apparent deficiency in IVDDM could adversely effect the energy balance of the animals; Blaxter et al. (1961) reported a 100% weight gain in domestic sheep when diet digestibility was increased from 50 to 55%. The energy deficit in March (46% diet IVDDM) tends to support the premise that energy acquisition by ungulates is secondary to protein acquisition in late winter. �275 What is the significance of the trade-off between diet digestibility and protein resulting from the observed selectivity for forbs? As noted by Rowland (1981) the complex inter-relationship of protein and energy in intermediary metabolism precludes a simple answer to the question of which nutrient is more important to the animal's welfare. However, several effects of the observed diet mix are plausible, and were reviewed by Hobbs et al. (1981). First, the higher protein content of dicots (relative to graminoids) could contribute, in vivo, to increased digestion of protein-deficient graminoids, by creating a favorable rumen environment. Second, a deficiency in dietary protein may be more detrimental to animal condition, than energy deficits, because catabolism of surplus fat, results only in weight loss, whereas catabolism of protein usually results in loss of muscle mass and decreased resistance to disease (Harper et al. 1977:570, Swick and Benevenga 1977, cited by Hobbs et al. 1981:169). Hobbs et al. (1981) concluded that dietary protein deficiencies may be relatively more costly to the wintering animal than energy deficits. It appears that mountain goat diets reflect this premise. However, I speculate that diets of mountain goats may not be as low in energy as indicated by diet IVDDM. Digestibility is an incomplete measure of the nutritive value of a forage since food intake is not considered (Short et al. 1974, Milchunas et al. 1978). If we assume that forbs are passed rapidly from mountain goat rumens (relative to graminoids), then the large amount of cell solubles in forbs could provide a substantial amount of energy. The lignin and cellulose contents of forage are important for this consideration. Milchunas et al. (1978:28), studying mule deer, concluded that lignin may confer brittleness on plant cell walls, and thus increase the rate of passage of highly lignified forages from the rumen. Conversely, ce~lulose was thought to confer flexibility and cause decreases in rate of passage. As a consequence of these structural characteristics, total metabolizable energy (ME) intake of high lignin-low cellulose forage (Vaccinium) was similar to that of low ligninhigh cellulose forage (Agropyron) (Milchunas et al. 1978:32). This was despite the markedly lower in vivo digestibility of Vaccinium compared to Agropyron. Frobs eaten by goats had higher lignin (10 versus 3.9%) and lower cellulose (22 versus 37%) levels than did graminoids in March. Thus, forb-dominated goat diets may have provided high levels of protein without sacrificing energy input. Age class variation. Differences in diet botanical composition between kids and yearlings had important effects on diet quality during winter; these effects were less pronounced in July (Table 8). During winter, more graminoids and fewer forbs and woody plants in diets of kids, compared to yearlings, resulted in consistent differences in diet IVDDM. Kids selected diets higher in IVDDM in all of the 6 winter comparisons (i.e., 2 comparisons for each winter month); the range of significance levels was 0.08 > P > 0.005. Concomitantly, yearlings generally selected diets containing more cell solubles (l-CWC) and lignin (Table 8). Differences in protein concentrations in diets of kids and yearling were inconsistent. �276 The more digestible diets of kids, relative to their larger bodied counterparts, conforms to theory on energy requirements in relation to animal body size (Kleiber 1961). Because energy requirements per unit body mass are greater for small mammals than for larger ones, small bodied ungulates are thoughtto require greater concentrations of digestible energy in their diets (Janis 1976, Schwartz and Ellis 1981, Hobbs et al. 1982). Also, young ungulates have less body reserves than can be catabolized for energy than adults (Ammann et al. 1973), and thus diet sources of energy are more important for young animals. As a consequence of these requirements, small ungulates tend to choose more digestible diets (reviewed by Hobbs et al. 1982). Because digestible energy is highly correlated with digestible dry matter (Robbins et al. 1975, Milchunas et al. 1978), kids may have been able to ameliorate their relatively greater energy losses (requirements) by consuming more digestible diets. Conversely, ungulates could choose dicot dominated diets containing high levels of cell solubles to meet their energy requirements (Short et al. 1974,. Schwartz and Ellis 1981, Hobbs et al. 1982). This appeared to be the strategy of yearling mountain goats during winter, when they ate more forbs and woody plants (95% of which was Salix leaves and catkins) than did kids (see Table 8 for significance levels). During each winter month, diets of yearlings contained more cell solubles than diets of concurrently observed kids (significance levels for months were 0.03, 0.4, 0.03). Similarly diets of yearlings contained more lignin (P < 0.002, 0.04, 0.1) than diets of concurrently observed kids. (The same differences for kids and yearlings observed in different winters were similar, but less consistent.) As discussed previously, forbs and Salix leaves were less digestible than graminoids. Diets containing large amounts of forbs or browse, or both, (i.e., cell solubles and lignin) could be good sources of energy if these forages are passed quickly through the gastrointestinal (GI) tract. As discussed previously, Milchunas et al. (1978:26) found such a situation in mule deer. For the larger bodied yearling goats, browse and forb diets could be especially beneficial since their absolute energy requirements should be greater than those of the smaller bodied kids (Kleiber 1961). Eating large amounts of graminoids might be a less efficient way of obtaining energy because of bulk limitations on digestion associated with roughage diets (Short et al. 1974, Milchunas et al. 1978, Schwartz and Ellis 1981). Conversely, animals eating highly lignified forbs and browse 'could rapidly assimilate soluble carbohydrates, quickly excrete the remaining ingesta, and thus increase intake of food and energy on a daily basis. Also, rapid passage of food leads to an increased portion of unfermented soluble carbohydrates reaching the intestine; this contributes to an increased gain of energy because methane losses are reduced (Kaufmann et al. 1980). In addition, in the case of Salix consumption (95% of woody plants eaten) yearlings might expend less energy while foraging. This is possible, because goats often ate large amounts of Salix from readily accessible clumps of the shrub, and thus used little energy in �277 searching for plants. Salix shrubbery also creates a microclimate wind, and thus animal energy expenditure is reduced. where Why did kids eat smaller amounts of forbs and Salix? Age related differences in gut morphology may have influenced the animals diet choices. Rumens of kids, relative to yearlings may be physically less able to tolerate concentrate foods. For example, Brownlee (1956) found that rumen mucosa of calves, fed only on concentrates (low fiber content), were easily torn or eroded. Kids also may be inefficient in assimilating the large amounts of nutrients produced in short time periods by highly soluble foods. Hofmann (1973:39) indicated that larger amounts (number and size) of papillae in the rumen were associated with highly soluble, rapidly passed ingesta. These allow nutrients to be absorbed quickly. Rumen papillae were found to be 2-5 mm shorter in 4-month-old white-tailed deer than in l~-year-old deer (Short 1964). If similar differences occur in mountain goat rumens, it may be more advantageous for yearlings to select more soluble, rapidly passed forages. Comparative Nutritional Ecology Estimates of congruence (i.e., similarity) between sheep and goat diet choices differed according to the calculation method used: 1) overlap using dry weight of individual taxon, and 2) analysis of variance using percent bites of each forage class (Fig. 13). Abramsky e t al. (1979) discussed the effects of food category (e.g., species or forage class) on the estimation of diet overlap. Whether overlap is overestimated or underestimated depends on the criteria of diet selection of the animal (i.e., do animals select on the basis of plant species, plant part, forage class, chemical composition, etc.). My findings on forage qualitydiet choice relationships and observations of other workers (e.g., Klein 1970, Kuropat and Bryant 1980, Schwartz and Ellis 1981, Hobbs et al. 1982) suggest that ungulate diet choices are affected by nutrient concentrations in plants. In addition, my findings indicate that sheep and goats differ in their response to forage nutrients, as evidenced by their dissimilar diets. Because nutrient levels vary among food categories (species, parts, class) (Arnold 1960, Van Dyne and Heady 1965, Cowan et al. 1970), and because these levels change seasonally and to different degrees for each of the food categories, it may be that the animals criteria of diet selection also changes seasonally and is dissimilar for goats and sheep. If this is true the accuracy of overlap estimates would also fluctuate· (under-or overestimate) temporally. For example, during August trials and in March 1980 forage class similarity indices were alike, but overlap of individual taxa was different (Fig. 13). This discrepancy may be due to the greater variety of taxa in goat diets compared to sheep (i.e., they responsed differently to forage nutrients). Overlap in March was calculated with 8 taxa that were common to goat and sheep diets; these accounted for 63% of goat diets and 83% of sheep diets. In other words, goats ate small quantities of numerous taxa, while sheep ate large quantities of a few taxa. The numerous taxa in goat diets that were not includ~d in overlap calculations, contributed to forage class similarity estimates, and hence, in �278 March, forage class similarity between sheep and goat diets could be relatively high compared to individual taxa overlap. Note that these effects may influence estimates in other months as well (e.g., September). Estimates of congruence were also inconsistent that overlap was overestimated. in January 1979; I believe Overlap indices are commonly used with little deference given to their prec1S10n. Because overlap calculations do not account for intraspecies (animal) variability, as analysis of variance (AOV) tests do, nO statistical confidence can be placed on their estimates (Abrams 1980, Lawlor 1980). Thus, inferences drawn from overlap estimates are not as reliable as those drawn from AOV tests. This is particularly important, because overlap was calculated on the basis of individual plant taxa in animal diets; intraspecies variation in selection of individual taxa was greater than variation in selection of forage classes (see Table 2, January column). To illustrate this problem, I recalculated overlap in January using the outermost diet percentages for sheep and goats (i.e., I calculated overlap for each plant taxon using the goat and sheep with the least amount of a taxon in their diets relative to their conspecifics; I repeated this for individuals with the largest amount of each taxon in their diets (Table 9). Calculated in this way low overlap (26%) and high overlap (100%) differed substantially from that of the original (64%). Table 9. Hypothetical calculation of diet overlap (Kulcynski's index) using the outermost percentages of 1 forage taxon common to sheep and goat diets. % polygonum in diets Animals Goats A B C 30 Sheep 14 4 26 5 10 High overlap pair: Goat B and Sheep C; Overlap Low Overlap Pair: Goat A and Sheep B; Overlap = = 26. 4. In conclusion, by using two methods to estimate diet differences, a more substantive evaluation of diet interactions could be made. In this regard, it was fairly clear (November 1978 excepted) that differences between sheep and goats in diet botanical composition and nutritional characteristics of diets were greater in winter than in summer. These findings agree with those of other workers who have observed that niche shifts in ungulate communities occur in response to decreases in the quality and quantity of available forage; diet overlap decreases as food becomes more limited (Longhurst et al. 1968, Sinclair 1977:80, Willms et al. 1980, Hobbs et al. 1982). �279 The similarity of diets of lambs and kids in Novmeber 1978 was striking. Similar to my previous suggestions lack of experience by animals with alpine vegetation, or characteristics of rumens of immature animals, or both, may have affected their diet choices. Regarding the former premise, it is possible that the food niches of tame animals became more established as their experience with available forage increased. Most interspecies differences in nutritional ecology of ungulates have been related to body size, and digestive and mouth morphology (e.g., Schwartz and Ellis 1981). Mouth morphology may playa minor role in sheep-goat comparisons, since their mouth parts are similar. Because energy and protein requirements of smaller mammals are greater per unit body weight than for larger ones (Bell 1970:119), small bodied ungulates are thought to require greater concentrations of protein and soluble carbohydrates in their diets, at the expense of fiber (Bell 1970, Short et al. 1974, Schwartz and Ellis 1981). However, total requirements for energy and protein are greater for larger animals then for smaller ones (Bell 1971). Hofmann (1973) related gut morphology and function to diet choices of African wild ruminants. He classified animals as follows: roughage feeders (eating mostly graminoids), mixed feeders (eating graminoids and browse). and concentrate feeders (eating browse tips, forbs and new growth of graminoids). Concentrate feeders tend to have a small, unobstructed (few pillars, large orificies between digestive organs) forestomach which allows ingesta to pass rapidly. The forestomach is characterized by rich and complete papillation which allows for rapid absorption of volatile fatty acids. These morphological features dictate that highly soluble, rapidly passed foods can best be utilized by concentrate feeders. Conversely, roughage and mixed feeders have large rumens with substantial obstructions to slow passage of food and allow more complete fermentation of a fibrous diet. Bighorn sheep and luountain goats display diet characteristics that place them along a diet quality gradient between concentrate and mixed feeders. Sheep diets, containing more graminoids than did goat diets, were more like diets of mixed feeders. At the same time, forb and browse dominated goat diets relate more to the concentrate feeding style. The nutritional benefits of these diets relative to the body sizes of sheep and goats are somewhat inconsistent. Slightly larger concentrations of protein in goat diets, relative to sheep diets, agrees with the theoretical predictions that the smaller bodied goats should be more selective for protein than the larger sheep. However, as noted previously, the energy benefits of goat diets, which contained large amounts of cell solubles and lignin, but small amounts of IVDDM, are equivocal. If we assume that goats and sheep conform to theory on body size-gut morphology-energy relationships, and that they optimized energy or nutrients, or both in selecting food, then we would expect goats to have smaller rumens with fewer physical barriers than do sheep. In this res- �280 pect, goats would appear to share some of the nutritional characteristics attributed to mule deer by Mi1chunas et al. (1978), and Hobbs et al. (1982). That is, they choose highly soluble but lignified forages, which yield sufficient amounts of energy per unit time due to their rapid passage from the rumen. Similarly, Short et a1. (1974) stressed the importance of rate of digestibility to the well-being of small ruminants. They concluded that mature forbs and browse, relative to mature graminoids, offer more digestible nutrients during the short retention times typical of small ruminants. Considering the ungulates studied by Hobbs et a1. (1982) and all goats and sheep observed in this study, theoretical predictions for relationships among body size-gut morphology-energy and protein requirements become less clear. Unlignified fiber (CWC-lignin) and cell soluble levels in diets were different than expected on the basis of body size (Table lQ. Along a body size gradient (or mixed to concentrate feeder gradient) the expected order would be elk (yearlings), sheep (mature), goats (mature), and sheep (lambs), goat (kids), or mule deer (fawns) (latter 3 have similar body sizes and are interchangeable). Note though, that fiber levels in Table l~ may be somewhat misleading for the sheep (lamb, mature)-elk comparison and the lamb-mature goat comparison. The higher concentrations of protein in sheep diets, relative to diets of elk and goats (Table l~ could increase digestion of un1ignified fiber and therefore reduce the real differences in the negative effects of fiber on retention time of ingesta. Protein concentrations in diets better paral1ed predictions based on body size of animals, with the most discrepant diet being that of mule deer fawns and bighorn lambs (alpine). Concentrations of protein in deer diets were at least 1% point lower than those of similar sized kids and lambs (Table 10). �281 Table 10. Un1ignified fiber and cell soluble levels in diets of tamed wild ungu1atesa and related hypothetical feeding types. Sample Species (Age class) size Habitat % CWC-b lignin c W_Sc W % cell solub1es W % crude Erotein W-S W Sheep (Lambs, e mature) 3 Alpine 60 51 34 43 6.3 Elk (Yearlings) 4 Montanealpinef 54 46 34 44 4.5 Goats (Kids) 7 Alpine 51 41 6.8 Sheep (Lambs) 3 Montane 49 42 7.0 Goats (Mature)g 3 Alpine 41 Mule deer (Fawns) 4 Montane 39 37 47 49 54 Feeding d types Mixed 5.9 5.8 Concentrate a Elk (montane), sheep (montane) and mule deer from Hobbs et a1. (1982); elk (alpine) from Baker and Hobbs (1982). bFiber constituents most available for microbial fermentation. Cw = mean for November, January, March trials, W-S = mean for winter and summer (July, August, September). d Concentrate feeder; eats mostly the leaves, flowers and fruits of forbs and browse, and some new growth of graminoids. Mixed feeder: eats graminoids and browse (Hofmann 1973). eDiets of lambs, yearlings and 2-year-01ds were similar. fWinter values from montane, W-S values are means of ~inter in montane and summer in alpine. gYearlings and 2-year-01ds. �282 Why was there so much variability in these body size-diet quality relationships? First, note that these ungulates are likely, in absolute terms, mixed feeders. They all show high variability in forage ~lass composition of diets within and between seasons (Todd 1972, Kufeld 1973, Kufeld et al. 1973). Thus, their digestive systems should be able to adapt to various food types. The adaptability of mixed feeders to a diverse diet is likely made possible by apparently reversible adjustments in the structural components of the stomach, especially the size and distribution of rumen papillation (Hofmann 1973). Rumen pap illation differs characteristically with food types (Kay et al. 1980). High quality diets are associated with greater quantities of papillae. Increased surface area of papillae results in greater absorption of nutrients, and hence, soluble ingesta passed rapidly through the rumen can be more efficiently utilized (Hofmann 1973). Conversely, roughage diets result in keratinized mucosa which severly limits absorptive capabilities of these structures (Hofmann 1973). Lack of information on the relative ability of these ungulates to efficiently use different types of forage, precludes a conclusive evaluation of their comparative nutritional ecology. Knowledge of the limitations of the digestive systems of mountain goats and bighorn sheep is a prerequisite to predicting the outcome of their competitive interactions. If and when the winter food supply is so reduced that their food niches approach unity, the animal that can best conserve, or obtain energy and nutrients, or both, could eliminate the other. SUMMARY 1. Diets of tame mountain goats during winter contained 50% forbs, 38% graminoids, and 12% woody plants. Despite the importance of forbs in winter, the most frequently chosen individual forages were the graminoid Carex rupestris (20% of diets) and the shrub Salix (11%). Other principal forages were (Campanula, Calamagrostis, Trifolium, Kobresia, and Geum. 2. During summer, forbs comprised 92% of the diets, graminoids 6%, and woody plants 2%. Principal forages were Trifolium nanum, I. dasyphyllum, !. parryi, Polygonum, and Mertensia. Agropyron scribneri accounted for most of the graminoids eaten. 3. Forbs and woody plants contained more cell solubles and lignin than did graminoids throughout the year. During summer, forbs' and graminoids were more digestible than woody plants. In winter, graminoids were more digestible than forbs; forbs contained more IVDDM than woody plants. The order of crude-protein concentrations among forage classes was forbs ~ woody plants> graminoids in summer, and woody plants> forbs > graminoids in winter. �283 4. Protein, IVDDM, and cell soluble levels decreased through summer for all forages. Increased consumption of Agropyron scribneri in late summer was associated with its higher concentrations of protein and IVDDM relative to other forage plants. 5. Through winter, digestibility increased for graminoids and woody plants, and decreased for forbs. Concurrently, protein concentrations increased in forbs and woody plants, and decreased in graminoids. ~pparently in response to these changes in forage quality, animals in late winter increased their consumption of forbs, perhaps maintaining stable dietary levels of cell solubles and protein, at the expense of reduced diet IVDDM. 6. Diet IVDDM, diet cell solubles, and diet crude protein all decreased from July to March. Decreases in these diet quality constituents were more pronounced during summer, than during winter. 7. The energy likely met goat diets inadequate 8. Snow affected forage availability, and hence diet choices, particularly with respect to Salix. Goats ate less Salix when these shrubs were covered with snow. When forbs and graminoids were covered with snow and Salix was not, animals ate more Salix. 9. Food niche breadth was inversely of forage available. (IVDDM, cell solubles) and protein the goat diets provided their growth requirements during summer. During winter, appeared to provide sufficient amounts of protein, but amounts of energy for maintenance. related to the quantity and quality 10. Mountain goat yearlings ate more forbs and Salix than did kids. Consequently, yearling diets contained less IVDDM, but more cell solubles and lignin. These diets may be good sources of energy and protein if forbs and browse are passed quickly through the gastrointestinal tract. 11. 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Wildl. Spec. Rep. No. 27. 2lpp. Trout, L. E. 1973. Big game rumen sample analysis. P-R Job Completion Rep. Study II, Job 1. Proj. W-14l-R-3 Idaho Fish and Game Dep. 4pp. �294 Trudell, J., et al. 1980. Comparison of some factors affecting the in vitro digestibility estimate of reindeer forages. Pages 262-273 in E. Reimers, E. Gaare, and S. Skjenneberg, eds. Proc. Second Int. Reindeer/Caribou Symp. Direktoratet For Vilt Og Ferskvannsfisk, Trondheim, Norway. Tucker, R., A. McLean, and D. E. Waldren. 1977. Relative preference by mule deer for six shrubs near Kamloops, British Columbia. Can. J. Anim. Sci. 57:375-377. Van Dyne, G. M., and H. F. Heady. 1965. Botanical composition of sheep and cattle diets on a mature annual range. Hilgardia 36:465-492. Van Gylswyk, N. O. 1970. The effect of supplementing a low-protein hay on the cellulolytic bacteria in the rumen of sheep and on the digestibility of cellulose and hemicellulose. J. Agric. Sci. 74:169-180. Webber, P. J., and D. E. May. 1977. The magnitude and distribution of below ground plant structures in the alpine tundra of Niwot Ridge, Colorado. Arct. Alp. Res. 9(2):156-174. Weir, W. C., and D. T. Torell. 1959. Selective grazing by sheep as shown by a comparison of the chemical composition of range and pasture forage obtained by hand clipping and that collected by esophageal fistulated sheep. J. Anim. Sci. 18:641-649. White, R. G., and J. Trudell. 1980. Patterns of herbivory and nutrient intake of reindeer grazing tundra vegetation. Pages 180-195 in E. Reimers, E. Gaare, and S. Skjenneberg, eds. Proc. Second Int. Reindeer/Caribou Symp. Direktoratet For Vilt Og Ferskvannsfisk, Trondheim, Norway. White, T. C. R. 1978. The importance of a relative shortage of food in animal ecology. Oecologia 33:71-86. Whittaker, R. H., and P. P. Feeny. interactions between species. 1971. Allelochemics: Science 171:757-770. chemical Wilkins, H. L., R. P. Bates, R. P. Henson, I. L. Lindahl, and R. E. Davis. 1953. Tannin and palatability in sericea lespedeza, L. cuneata. Agron. J. 45:335-336. Willms, W., A. McLean, R. Tucker, and R. Ritcey. 1980. Deer and cattle diets on summer range in British Columbia. J. Range Manage. 33:55-59. �295 Prepared by -r~vUet?-1 ,Thomas V. Dailey Graduate (\ ,\ Prepared by ~\. Research \' \ ) \)~,), R. Bruce Gill Big Game Research £1..- -Jrlw ~ Assistant \ j~ Leader ��297 APPENDICES �298 Appendix I. Braun-Blanquet vegetation description of winter and summer study areas (derived from a study by Kom~rkova 1976). Study areaa W,S Description Alliance Kobresio-caricion rupestris Diagnostic taxa: Erithichum, Haplopappus, Draba, Trifolium, Artemisia, Potentilla, w Association Eritricho-Dryadetum Diagnostic Optimum species: taxa: Paronychia, Kobresia octopetalae Dryas octopetala Draba, Potentilla Constants: Festuca, Silene, Lloydia, Arenaria, Carex, Phlox, Hymenoxys, Erithichum, Draba S Association Trifolietum Diagnostic species: Optimum taxa: Hymenoxys dasyphyllum Trifolium T. dasyphyllum, dasyphyllum Helictotrichon, Constants: Geum, Silene, Sedum, Castilleja, Mertensia, Oreoxis, Poa, Carex, Hymenoxys, Eritrichum, Paronychia S Association Silene, Arenaria, Constants: Geum, Festuca, Luzula, Haplopappus W,S Association Selaginello-Kobresietum Dominant Optimum Arenaria, Sileno-Paronychietum Dominant taxa: Paronychia species: taxa: Constants: Oreoxis, Poa, Kobresia Lloydia, Trifolium, Polygonum, myosuroidis myosuroides Draba, Arenaria Geum, Polygonum, Arenaria, Poa, Carex, Eritrichum Lloydia, �299 Appendix I. (continued) Study areaa S Description Association Caricetum elynoidis Dominant species: Optimum taxa: Carex elynoides Carex, Agropyron, Oreoxis, Potentilla, Poa Constants: Trisetum, Arenaria, Mertensia, Oreoxis, Poa, Agropyron W,S Alliance Deschampsio-Trifolion parryi Diagnostic taxa: Deschampsia, Erigeron, Trifolium, Agrostis, Gentianella, Stellaria W,S Association Acomastylidetum (Geum) rossi1 Dominant species: Constants: W,S Geum rossii Festuca brachyphylla Association Toninio-Sibbaldietum Dominant species: Sibaldia procumbens Constants: Sibaldia, Ranunculus, Geum, Festuca, Luzula, Polygonum, Artemisia, Arenaria, Lewisia, Agrostis, Antennaria S Association Caricetum pyrenaicae Dominant species: Optimum taxa: Constants: S C. pyrenaica, Erigeron Juncus, Antennaria Association Juncetum drummondii Dominant species: Optimum taxa: Constants: W Carex pyrenaica Juncus drummondii J. drummondii, Carex Sibbaldia, Antennaria, Epilobium Alliance Salicion planifolio-villosae Diagnostic species: winter, S summer. Salix villosa, S. planifolia �300 Appendix II. Taxa found in or near Niwot Ridge study areas and forages eaten by tame mountain goats in alpine tundra, Colorado. Scientific name a Conunon name Study areab. Goat foodc W W * * Sourced FORBS Achillea lanulosa Allium sp. Androsace septentrionalis Anemone canadensis Angelica grayi Antennaria sp. Arenaria fenderi A. obtusiloba A. sp. Artemisia arcticae A. scopulorum Besseya alpina Cal~ha leptosepala Campanula rotundifolia C. uniflora Castilleja occidentalis C. puberula Cerastium arvense Chionophila jamesii Cirsium scopulorum Crepis runcinata Cryptogranuna acrostichoides Draba crassifolia D. fladnizensis Epilobium angustifolium Erigeron peregrinus E. pinnatisectus E. simplex E. sp. Eritrichum aretoidese Erysimum nivale Gentianodes algidae Geum rossii --Haplopappus pygmaeus Heuchera parvifoliae Hymenoxys acaulis H. grandiflora Lewisia 'p~a Yarrow Wild onion Rock jasmine Meadow an emone Angelica Pussy toes Fendlers sandwort Alpine sandwort Sandwort Arctic sage Alpine sage Alpine besseya Marsh marigold Conunon harebell Alpine harebell Western yellow paintbrush Alpine paintbrush Mouse-ear chickweed Snowlover Thistle Hawksbeard Rockbrake Thick leaved whitlow grass White arctic draba Fireweed Subalpine daisy Daisy One-headed daisy Daisy Alpine forget-me-not Wallflower Arctic gentian Alpine avens Haplopappus Conunon alum-root Hymenoxys Old man of the mountain Pigmy bitterroot S S S W S ,W S ,W S W S,W S S S,W S S,W S ,W S,W S S S D D M M * * * * *,'e * * * * * * * * D D D D M D D D D D M D D D D D D S D S S S S S M S,W S,W S ,W S ,W S S,W S S,W S,W S,W S M * * * * * * * * * * * * D D D D D D D D D D D D D D �301 Appendix II. Scientific (continued) namea Lloydia serotina Mertensia ciliata M. viridis Oreoxis alpina Oxyria digyna Paronychia pulvinata Pedicularis· groenlandica P. parryi Penstemon whippleanus Phlox caespitosa Phacelia sericea Polemonium viscosum Polygonum bistortoides R_. viviparum Potentilla diversifolia P. nivea P. rubricaulis Primula angustifolia Pseudocymopteris montanus Ranuculus adoneus Saxifraga clebilis S. rhomboidea Sedum integrifolium S. stenopetalum f Selaginella densa Senecio canus S. carthamoides Sibbaldia procumbens Silene acaulis S. scouleri Solidago spathulatae Stellaria longipes Swertia perrennis Taraxacum ceratophorum Thlaspi alpestre Trifolium dasyphyllum T. ---. nanum T. parryi f Woodsia oregana Zygadenus elegans Common name Alp lilly Tall mertensia Green mertensia Alpine parsley Alpine sorrel Nail wort Elephantella Lousewort Beard-tongue Phlox Purple fringe Sky pilot American bistort Viviparous bistort Blueleaf cinquefoil Snow cinquefoil Cinquefoil Alpine primrose Yellow mountain par s Ley Snow buttercup Pygmy saxifrage Saxifrage King's crown Yellow stonecrop Rock selaginella Golden ragwort Ragwort Sibbaldia Moss campion Scouler's catchfly Golden rod Long-stalked stitchwort Star gentian Alpine dandelion Wild candytuft Whiproot clover Dwarf clover Parry clover Oregon woodsia Death camas Stud?: area Goat foodc S S * * * * * * S,W S ,W S S,W S W S S,W S S ,W S ,W S S ,W S S S Sourced * * * * **. * * s,w S,W S S,W S S,W S,W W S S ,W S ,W S S,W * * * * * * * * * * S S S S M D D D D D D D D M M D D D M D D D D D D D D M D M S S S S S ,W D D D D D D M M .* * * * * ----------------------------------------------------------------------- D D D D D M �302 Appendix II. Scientific (continued) namea Common name Studt; area Goat foodc S ,W * * * * Sourced GRAMINOIDS Agropyron scribneri Calamagrostis purpurascens Carex rupestris f. sPP. Danthonia intermedia Deschampsia caespitosa Festuca brachyphylla Juncus drummondii Kobresia myosuroidese K. spp. Luzula spicata Poa spp. Trisetum spicatum Scribner's wheatgrass Purple reed-grass Sedge Sedge Timber danthonia Tufted hairgrass Fescue Rush Kobresia Kobresia Spike wood-rush Bluegrass Spike trisetum S ,W S,W S,W W W S,W S,W S,W S,W S,W S,W S,W * * * * * )'t D D D D D D D D D D D D D WOODY PLANTS Abies lasiocarpa Dryas octopetala f Parmelia conspersa Picea engelmannii Potentilla fruticosa Salix arctica S. brachycarpa S. nivalis S. planifolia Vaccinium caespitosum w Subalpine fir Mountain dryad Lichen Engelmann spruce Shrubby cinquefoil Willow Willow Snow willow Planeleaf willow Dwarf bilberry W W W S,W S W S S,W S ~omenclature from Harrington (1964). b Study area; W = winter, S = summer. CEaten by tame mountain goats. dCollector = of plants; D eNomenclature fCryptogam. from Weber Dailey, M (1976). = Marr (1961). * * * * * * * D D D D D M D D D D �303 Appendix III. Plant taxa common in winter study area on Niwot Ridge. Dominance estimates are means calculated across dominance values for each of 4 stand types studied by Osburn '.1958). Dominance Taxon Kobresia myosuroides 3.6 Arenaria obtusiloba 2.6 Geum rossii 2.2 Lloydia 2.1 serotina Phlox spp. 1.9 Poa spp. 1.8 Trifolium dasyphyllum 1.6 Silene acaulis 1.6 Eritrichum 1.6 aretoides Oreoxis alpina 1.2 Festuca brachyphylla 1.2 Sedum stenopetalum 1.2 Polygonum spp. 1.2 Hymenoxys spp. 1.2 Arenaria fendleri 1.1 Campanula rotundifolia 1.0 Cerastium arvense 1.0 0.9 Carex spp. Agropyron scribneri Calamagrostis Paronychia purpurascens. pulvinata 0.8 0.6 ��July 1981 305 JOB PROGRESS State of Project REPORT Colorado ~]o. W-126-R-4 ----- 7 ~.J'ork Plan No. Job Title: Job No. 1 Black Bear Investigations Period Covered: Personnel: Big Game Investigations July 1, 1980 through June 30, 1981 T. D. I. Beck, R. B. Gill, R. M. Hopper, L. H. Carpenter, D. L. Baker, M. L. Stevens, R. Danvir, M. Haroldson, G. H. Bock, D. Millers D. Sizemore, D. Coven. ABSTRACT Thirty-seven black bears were captured during the 1980 season. Thirty-one of these were initial captures and 6 were recaptures. Analysis of age distribution data from captured bears indicated a population subject to heavy exploitation. Sixty percent of the capture sample was comprised of subadults. Other areas where bears have been hunted only lightly or not at all report subadult components ranging from 20-25% of the population. Our data are disturbing because our bear study area has been closed to hunting for 3 years. Denning activity began the 3rd week in October. Bears remained denned until March 13, 1981 when the first radio-collared bear left his den for the remainder of the spring-summer period. Most of the radiocollared bears did not leave dens until after April 13, 1981. Females .generally entered dens earlier and abandoned dens later than males. Nineteen den sites were located during winter-spring 1980-81. Physical characteristics of den sites were described. ��307 BLACK BEAR INVESTIGATIONS Thomas D. I. Beck P. N. OBJECTIVES 1. Develop techniques to accurately and precisely estimate black bear populationS. 2. Determine habitat preferences of selected black bear populations. 3. Describe black bear population dynamics sufficiently to allow analysis of various harvest and habitat manipulations. SEGMENT OBJECTIVES 1. Develop techniques to accurately and precisely estimate black bear populations. 2. Determine habitat preferences of selected black bear populations. 3. Describe black bear population dynamics sufficiently to allow analysis of various harvest and habitat manipulations. METHODS AND MATERIALS Study Area Delineation Delineation of the study area will be based upon movements of radiocollared female black bears. A study area boundary will be established' to include nearly all the seasonal movements of the collared females. It is assumed that a substantial amount of the male black bear ranges will also be within the study area but preliminary data indicate very large ranges for males. Capture and Marking Capture efforts lasted from 27 May to 8 October 1980. All captures were made using Aldrich spring-activated snares and basic techniques described by Flowers (1977). All snare sets were out of sight from roads and maintained trails and were checked daily. Snared bears were immobilized with a combination of ketamine hydrochloride and xylazine hydrochloride in a mixture of 180 mg ketamine and 90 mg xylazine per cubic centimeter. Drug was administered with use of a 6-foot jab pole. Bears were tattooed, ear-tagged with calf-size Ritchey rubber ear tags, and some were instrumented with a Telonics radio transmitter collar. Numerous physical measurements were taken and a premolar removed for subsequent aging by cementum annuli counts. �308 Habitat Use Data on habitat use was collected primarily from ground tracking radiocollared bears supplemented by aerial tracking. Den sites were located in November and December 1980 by both ground and aerial tracking. Dens located from the air were subsequently searched for in February and March 1981. All located dens were visited in February-April 1981 and numerous physical measurements taken of the dens and general descriptions of the den site were made. Data on relative use of community types was of insufficient quantity to warrant analysis during this segment. RESULTS AND DISCUSSION Study Area Selection The area closed to bear hunting for purposes of this study proved to be most fortuitous in regards to final study area selection. Initially it was hoped the hunting closure would extend beyond the study area. However, to include all of the females captured within the study area required enlarging the study area. There are still 3 females who spend substantial time out of the study area. The study area includes all of Game Management Unit 53 south of the Smith Fork, Sink Creek and West Elk Mountains, all of GMU 63 east of Colorado Highway 92 and Crystal Creek, and GMU 54 west of Soap Creek. Thus the principle drainages are Soap, Curecanti, Mesa, Crystal, Dyer, Muddy, Clear Fork, Virginia, and South Smith Fork Creeks (Figs. 1 and 2). The area encompasses approximately 190 square miles. Capture and Marking Thirty-seven black bear captures were made during the 1980 season of which 31 were initial captures, 3 were recaptures of bears marked in 1979, and 3 were recaptures of bears marked in 1980. A summary of all bear captures in 1979, 1980, and June 1981 is provided in Tables 1 and 2. Estimated ages are from cementum annuli counts. Estimates of age by cementum annuli counts has been quite variable in bears over age class 6. However, the younger age classes have been consistent when tested in 2 labs using multiple teeth. Lab estimates agree well with field estimates on the younger bears and when varied the lab estimate is consistently 1 year higher. Extreme wariness (trap-shy) by once-handled bears is evident from the number of recaptures. This must be considered an un~sual phenomenon in light of other bear studies. Analysis of age distribution of trap sample is disturbing. One of the reasons for selecting the area for study was the belief that there was no illegal killing of bears by livestock owners and the licensed harvest was historically low. The age distribution presented in Table 3 shatters any such belief. Data from literature indicates the percent of subadult bears generally ranges around 20-25% in unhunted and lightly hunted populations and 40-48% in heavily hunted, over exploited populations (Beecham 1980, �July 1 through August 28 w o 1.0 Figure 2. Colorado Black Bear hunting seasons and location of study area (closed to hunting). �310 Figure 1. Black Mesa, black bear study area (outlined with stippled line). �311 Table 1. Estimated age and weight of captured male black bears, Black Mesa study area, Colorado. LD. No. Capture Date Estimated Age Weight (kg) M-l 5-29-79 7 141 H-l M-2 M-2 H-3 6-29-79 6-13-79 6-27-79 7-25-79 7 4 4 132 68 66 121 M-4 M-5 M-5 M-5 M-6 7-30-79 8-2-79 6-2-80 5-27-81 9-11-79 6 M-7 8-11-79 8-27-79 9-20-79 9-12-80 5-28-80 6-3-80 5 H-8 H-9 M-9 H-IO M-Dead M-11 H-12 M-12 M-13 6-3-80 6-4-80 10-7-80 6-8-80 H-14 H-14 6-16-80 9-18-80 H-15 M-16 6-18:"'80 6-19-80 6-25-80 7-1-80 7-3-80 7-12-80 6-24-81 8-13-80 8-20-80 9-14-80 9-25-80 9-25-80 10-6-80 6-10-81 6-22-81 M-17 M-18 M-19 H-20 H-20 H-21 M-22 M-23 M-24 M-25 M-26 M-28 H-29 12 2 3 4 3 107 41 76 86 71 5 9 84 52 48 61 127 159 4 5 5 6 52 91 118 130 3 3 48 3 68 139 2 1 2 9 71 3 3 3 3 4 55 2 2 46 34 82 80 32 80 4 3 1 4 Remarks Killed 7-12-79, 9 miles south of study area Illegally killed November 1980 in study area Killed 7-14-80 approx. 90 miles SW of study area. Cub of the year Illegally killed while in snare Killed 6-16-81, 0.2 N of area Illegally killed 10-13-80 in study area 57 59 64 66 1 11 3 61 miles Cub of the year Cub of the year �312 Table 2. Estimated age and weight of captured female black bears, Black· Mesa study area, Colorado. Capture Date Estimated Age Weight (kg) F-1 F-1 F-2 F-3 F-4 F-4 F-5 F-6 F-7 F-7 F-8 6-13-79 6-8-81 6-17-79 7-1-79 7-18-79 8-16-79 7-19-79 7-20-79 7-27-79 7-9-80 8-1-79 6 8 1 1 2 2 2 4 6 7 5 70 82 F-9 F-9 F-10 F-11 F-12 F-13 F-14 F-15 F-16 F-16 F-17 F-18 F-18 F-19 F-20 F-2l F-22 F-23 8-1-79 6-10-81 9-10-79 9-14-79 9-17-79 5-25-80 6-7-80 6-8-80 6-19-80 9-5-80 6-21-80 6-24-80 6-20-81 6-25-80 6-27-80 6-30-80 7-1-80 7-9-80 7 9 3 12 3 4 4 10 4 4 6 1 2 3 5 2 9 1 68 61 59 107 50 57 57 98 45 54 76 15 27 43 61 30 91 13 F-24 F-25 F-26 9-3-80 10-4-80 6-19-81 5 2 2 52 36 31 LD. No. 11 13 27 33 27 48 75 64 43 Remarks Cub of the year Cub of the year Killed 10-22-79, 5 feet N of study area Cub of the year Cub of the year; cub of F-7 Lindzey and Meslow 1980). Our capture sample is comprised of 60% subadults. There is no reason to believe we are achieving a skewed capture frequency favoring the younger animals. Subadult females are generally believed most difficult to capture because of smaller ranges and this group accounts for 57% of the females caught (cubs excluded). The high subadult percentage, combined with the absence of any old bears (oldest bear caught was 12), leads one to believe this population has been subjected to severe over-exploitation during the past decade. The �313 consequences of these data are relevant to our research program and mana-, gement. From a research perspective, we have only a small number of reproductive-age bears thus obtaining good samples of reproductive rates will take longer than originally planned. From a management view, one must wonder what level of illegal killing is occurring throughout the state, especially in areas where sheepmen are known to be killing a lot of bears. Table 3. Age distribution of black bears caught in Black Mesa study area, 1979 and 1980 (all ages of 1979 caught bears corrected to 1980). Age Class Cubs 1 Subadult 2 3 4 5 6-10 11-15 Males 1 3 9 4 3 6 1 Females 2 4 4 5 2 7 1 Of the 52 bears caught in 1979 and 1980, a m1n1mum of 7 have been killed; 4 legally and 3 illegally. I have an unsubstantiated report of another collared bear being illegally killed in 1980. The 3 illegal kills were 1 male shot while in a snare, 1 male shot and abandoned in the closed area, and 1 male shot and taken out of closed area. Also, 2 bears were reported killed in 1979 in the closed area by a Colorado Division of Animal Industries trapper but it is not known if these were marked bears. Even with the large closure to bear hunting the loss of marked bears is a major problem to our research effort. This loss is es.pecially important because the study population is so young and probably at lower density than the habitat can support as a result of illegal killing throughout the 1960's and 1970's. Habitat Use Collared black bears began entering dens during the 3rd week of October with the last to den doing so during a major snowstorm December 8-10 (Table 4). Emergence began in mid-April and continued into mtd-May. Only one collared bear left a den prior to 13 April (Table 5). A 3-year old male left his den on 13 March and stayed out. His den was located on a steep south facing slope in the Black Canyon and subject to direct warming from the sun. March 1980 was unusually mild with temperatures reaching 50-550 F during many afternoons. A 4-year old male changed dens sometime in January or early February. His den was also in a steep, south facing rock slope with direct exposure to the sun. Both dens were shallow natural cavities in the rocks and could have easily warmed up to a temperature of 500 F or better. �314 Eight dens used by male black bears have been located and described. Six. dens were in natural rock cavities, 1 was dug in soil under a clump of Prunus virginiana, and one was an open bed under the low limbs of a Picea engelmanni. Two of the rock dens showed evidence of use by.bears in previous years (Table 6). Table 4. Number of collared bears entering dens by I-week periods, 1980. 3rd wk Oct. 4th wk Oct. 1st wk Nov. 2nd wk Nov. 3rd wk Nov. 4th wk Nov. 1st wk Dec. 2nd wk Dec. Males o o 2 o 3 2 o 2 Females 3 4 2 a 1 o 2 o Table 5. Number of collared bears leaving dens by I-week periods, 1981. 2nd wk March 1st wk April 2nd wk April 3rd wk April 4th wk April 1st wk May 2nd wk May Males 1 0 2 1 2 0 0 Females 0 0 0 3 4 5 0 Eleven dens used by female black bears have been located and described (Table 7). Six dens were in natural rock cavities, 3 were dug under the base of conifers, and 2 were dug under the bas~ of serviceberry shrubs (Amelanchier sp). The 3 females that had cubs in 1980 were in excavated dens; 2 under shrubs, one under a spruce. Each of the females with young had litters of 3. Two females with yearlings used rock cavity dens; one had 2 yearlings, 1 had a single yearling. �315 Table 6. Site characteristics of dens used by male black bears. Den type Rock cavitya Rock cavity Excavated in soil Rock cavity Rock cavity Rock cavity Rock cavitya Above ground bed Overstory Vegetation Elevation (feet) Aspect (degrees) Slope (degrees) 8470 9300. 8380 9400 9550 8450 8650 10300 360 297 20 112 224 352 60 185 22 51 15 35 45 57 80 Gambel oak Douglas fir Gambel oak Douglas fir Douglas fir Douglas fir Douglas fir Engelmann spruce 3 a Evidence of use by bears in previous years Table 7. Site characteristics of dens used by female black bears. Den type Excavated-tree base Rockcavitya Rock cavitya Rock cavitya Rock cavity Excavated-tree base Excavated-under shrubs Excavated-under shrubs Rock cavitya Rock cavity Excavated-tree base Overstory Vegetation Aspect (degrees) Slope (degrees) 10400 9550 10750 10750 250 210 180 232 36 65 45 46 9350 10050 8650 260 160 -112 48 49 27 8250 330 35 9650 8950 11225 130 112 240 70 50 53 Elevation (feet) Engelmann spruce Douglas fir open rock cliff Engelmann sprucesubalpine fir Douglas fir Douglas fir Gambel oak Gambel oak Gambel oak Rcok slide Engelmann spruce a Evidence of use by bears in previous years �316 LITERATURE CITED Beecham, J. J. 1980. Some population characteristics of two black bear populations in Idaho. Proc. IntI. COnf. Bear Res. and Manage. 4:201-204. Flowers, R. 1977. The art and technique of snaring bears. Forest Protection Assoc., 37pp. Washington Lindzey, F. G. and E. C. Mes1ow. 1980. Harvest and population characteristics of black bears in Oregon (1971-74). Proc. IntI. Conf. Bear Res. and Manage. 4:213-220. Prepared by JLt>-4fl.J£!d. Thomas D. I. Beck Wildlife Researcher �317 July 1981 JOB PROGRESS State of Project C_o_l_o_r_a_d_o No. W-126-R-4 Work Plan No. Job Title: Mountain Period Covered: Personnel: 8 REPORT _ Big Game Investigations Job No. Lion Investigations 1 - Mountain Lion Population Dynamics July 1, 1980 through june ?O, 1981 Allen E. Anderson, Chuck Anderson, Craig Albright, Jeff Brent, Ingrid Hamann, Donald Masden, Jim alterman, Geoff Tischbein ABSTRACT About 20 percent of the first draft of the comprehensive review of literature on the puma has been written and essentially all tables typed. A study plan was written and accepted for Program Narrative. Game Management Unit 62 on the east slope of the Uncompahgre Plateau was selected as a study area and one immature, female puma was captured, radiocollared, ear-tatooed and released on April 16, 1981 to test equipment, logistics, and sampling methodology. Body and dental measurements, hematology, pulse and respiratory rates, and rectal temperatures are reported for this puma. Six aerial and ground relocations using radiotelemetry indicated that the puma moved about 11 miles (29 km) south and gained about 2,600 ft (793 m) in elevation over a two-month period. ��319 MOUNTAIN LION POPULATION DYNAMICS Allen E. Anderson P. N. OBJECTIVES 1. Estimate density and population populations. 2. Develop an improved method to reliably estimate mountain densities which is also economically feasible to apply. 3. Assess mortality rates of young mountain maternal-filial bond separation. 4. Assess dispersal 5. Estimate the impact of hunting and removal on mountain population dynamics. 6. Measure the inter-relationships between mountain and large ungulate prey populations. and subsequent size of selected mountain lion lion lions after the period of fate of sub-adult mountain lion. lion lion populations SEGMENT OBJECTIVES 1. Complete and submit a 'review of literature for publication. 2. Complete a draft study plan outlining procedures by July 31, 1980. 3. Examine several potential study areas and from these select 2 or more for evaluation by professional lion hunters as their potential for studying mountain lion population dynamics. 4. Select final study area and become thoroughly topography, vegetation composition, etc. 5. Field test equipment, drugs, and capture techniques by capturing, radio-collaring, and radio-tracking one or more mountain lions. METHODS on mountain lion ecology specific objectives and familiar' with access, AND MATERIALS A comprehensive review of the published and unpublished the puma was undertaken as the first step. literature on �320 RESULTS AND DISCUSSION All tables of the comprehensive puma literature review were essentially completed and typed and the first drafts of chapters on evolution genetics and taxonomy were written. Three detailed, documented distribution maps for the puma in North America were completed by Ingrid Hamann. A draft for peer review is targeted for completion during the 1981-82 segment. A revised Program Narrative was submitted and accepted for the 1981-82 segment. Detailed study plans each specific investigation will be pre-:pared and appended to the Program Narrative before the investigations are initiated. Three candidate study areas (Colorado Fuel and Iron Corp. lands southwest of Trinidad, the Southern Ute Indian Reservation, and Game Management Unit 62 on the east slope of the Uncompahgre Plateau) were examined and the latter area selected during March 1981. The principal reasons for selection of GMU 62 were that C.F. and I did not want the study on their lands, the Southern Ute Indian Reservation was not enthused either, and G.M.U. 62 was mostly public land, reasonably accessible, apparently supported moderate numbers of puma in the opinion of professional hunters Chuck Anderson and Jeff Brent, and had a good deer census system which estimated 10.4 deer/km2 (Kufeld et al. 1980). A total of 16 days during January, February, and March 1981 were spent in the field with either Anderson or Brent and C.D.O.W. personnel familiar with the Uncompahgre in an attempt to assess the relative status of the puma population and the overall suitability of the Uncompahgre Plateau for long-term research on puma. Game Management Unit 62 includes the entire east slope of the Uncompahgre Plateau to approximately the west banks of the Uncompahgre and Gunnison rivers; an area of about 3,263 km2• About 1305 km2 are BLM lands, 1305 km2 U.S. Forest Service 65 km2 are other public lands, and 588 km2 are privately owned. The average elevational range approximates 1,734 m to 2,896 m. The Plateau is dissected by many canyons whose permanent or intermittent streams flow easterly in dendritic drainage patterns. Some canyons are over 300 m deep and most are steep-sided; characterized by extensive cliffs, rims, and exposed sandstone. The vegetation types range from sparse desert shrub at the lowest elevations, changing to pinyon-juniper woodland, ponderosa pine-Douglas fir forest and spruce-fir forest with increasing elevation. Gambels oak and aspen are important components of the upper elevation plant communities. Program Narrative Objective 5 to; "Estimate the impact of hunting and removal on puma population dynamics." requires the closure of G.M.U. 62 to sport hunting for 10 years. Even though the unit has been hunted only three seasons with about 16 puma recorded killed by the sport hunting, the Wildlife Commission rejected our request for closure at a public meeting on March 19, 1981. Additional efforts will be made during 1982 for enactment of this regulation. �3n Mainly as a learning exercise for me about 168 km2 were hunted with professional hunter Chuck Anderson and a pack of up to 8 dogs April 9-16 and April 22-29, 1981 (Table 1). Sixteen days (about 167 hr) of hunting resulted in the capture of one immature female on April 16, 1981. In spite of an inadvertent overdose of immobilizing drug (Table 2), resulting in an 8 or 37 minutes recovery time; the puma survived and one of its first conscious efforts after walking away from me was an unsuccessful attempt to force the radio collar from its neck. During immobilization pulse rates at 1 hr 29 minutes and 7 hr 52 minutes from starting were 84 and 68 beats/min, respectively. Respiration was an almost constant 20 exhalations per minute, and 3 rectal temperatures; 1 hr 35 minutes, 2 hr 35 minutes, and 3 hr 32 minutes from darting were 38.3 e, 37.3 e, and 37.1 e, respectively. Body and dentition measurements of puma number 1 are listed in Table 2. Her hematology is compared with that from two other immature but confined puma in Table 3. All measurements are within the range of values for apparently healthy puma reported in the literature. Tests for haemobartenella and dirofilaria in puma number 1 were negative. .~. Table 1. Legal descriptions of areas hunted for puma at least twice April 9-16, 1981 and April 22-29, 1981, Uncompahgre Plateau. Total Sections Sections Hunted T~ Range 15S 15S 96w 97W 31,30 36 2 1 49N 49N 49N 12W 13W 14w 7,8,18 3,4,5,6,7,10,11,12,13. 2,11,12,13,14 3 9 5 50N 50N 12W 13W 2 19 50N 14W 5,6 2,5,6,7,10,11,14,15,20,21,22,27,28,29,30,31, 34,35,36 2,11,12,13,14 51N 51N 12W 13W 10,11,15,16,20,21,29,30 12,13,14,23,26,27,32,33,34,35,36 8 11 Total 5 65 �322 Table 2. captured, Location Body and dentition measurements of an immature female puma tatooed (#1) and radiocollared at 0923 on April 16, 1981. of capture Drug and dosage: Radiocollar site: NW~, S10, T50N, R13W (Cottonwood Basin, U.S.G.S. Quadrangle) about 20 m N of Dry Fork Escalante Creek, 6060 ft (1847 m) elevation. 27.1 mg/kg (12.3 mg/lb) Serial No. 8389, Transmitter Measurements: of 1:2 mixture frequency of Rompun/Ketamine1 149.9550 MHz (1005-1050) Body wt (kg) Total body length (cm) Tail length (cm) Head-body length (cm) Chest girth (cm) Neck circumference (cm) Height at shoulder (cm) Head length Zygomatic breadth Ear length (cm) Hindfoot length (cm) Hind paw length (cm) Heel pads; max diameters (cm) Left front:anterior-posterior transverse Left rear:anterior-posterior transverse Teeth (cm) Maxillary toothrow (canine-premolar) Upper second premolar:crown length :crown width Lower first molar crown length Upper canine anterior-posterior diameter Gumline Recession (mm) Upper canine Lower canine 2nd upper premolar 1st lower premolar 33.12 172.5 74.0 98.5 55.0 31.0 69.0 3 __ 3 9.0 26.5 8.0 3.7 5.5 3.5 4.8 3.60 1.50 0.72 1. 63 1.29 3 3 1.5 1.5 lAdministered with powder-propelled, 3 cc capacity, barbed dart from a Cap Chur rifle at a distance of about 25 ft (7.6 m). The dart hit the area of the biceps femoris muscle in the right rear leg. Furacin was applied to dart wound. 2weighed to the nearest 3Wood swelling impossible. pound on a spring scale and converted in our fabricated wood-metal to kg. caliper made measurement �323 Table 3. Hematology immature puma. of one free-ranging (puma #1) and two confined Date completed Sex Packed cell vol (%) Hemoglobin (g/100 ml) Erythrocytes (106/mm3) Mean corpuscular vol (~3) M.C.H.C. (%)1 Total protein (g/100 ml) Fibrinogen (mg/l00 ml) Leukocytes (Total count) Differential Segmented neutrophils Lymphocytes Eosinophils IMean corpuscular 4-20-81 F 48 16.2 10.8 44 34 8.0 300 12,100 No. hemoglobin 8833 3025 242 34 23 12 3-16-81 M 3-30-81 F 38 13.2 7.9 30 35 6.8 100 14,800 % 73 25 2 No. 9916 4588 296 14.8 10.3 % 67 31 2 concentration. 2Free ranging 12-21 months of age; radiocollared taken April 16, 1981. and blood samples 3Confined, captured near Delta, CO, about 5 months of age. 4Confined, captured near Nucla, CO, about 3 months of age. Relocations of puma number 1 by aerial and ground telemetry are summarized in Table 4. One week after capture the puma was radiolocated and also seen from the air in the same canyon (Dry Fork Escalante Creek) and about one-half mile (805 m) west of the capture site. Three relocations during June 1981 were within about 6.5 km of each other. In about two months, puma number one moved about 29 airline kilometers south and gained about 793 m in elevation. From the air, I guess that relocations have been within one km2. As yet, we lack sufficient experience to estimate relocation accuracy on the ground. Triangulation techniques have not been possible since the puma apparently has been in almost constant motion when radiotracked on the ground. The major objective of ground tracking is to find prey killed by puma. But poor tracking conditions in the lush vegetation characteristic of the upper elevations now being used by the radiocollared puma make this a fairly remote possibility without an intensive triangulation effort. �Table 4. Movements of female puma no. 1 as detected by aerial and ground telemetry at approximate weekly intervals. Legal DescriEtion Date ~ S Twp 4-16-81 NW 10 50N 13W 4-23-81 SE 9 50N 13W 5-19-81 SW 36 50N 13W Monitor Creek 5-28-81 SW 6 48N 13W Monitor Mesa 6-08-81 NW 32 49N 13W Monitor Creek " 6-15-81 NW 4 48N 13W Criswell Creek 6-22-81 NE 5 48N 13W 7N Mesa Drainage or Locale R Dry Fork Escalante Creek " " " Cottonwood Basin " ft m Distance from Erevious location ft m 6060 1847 (capture site) Elevation U.S.C.S. Quadrangle " " 6200 1890 3168 966 " " 7320 2231 23886 7282 8920 2720 42436 12938 " 8680 2646 13035 3974 " " 8720 2658 33660 10262 " " 8800 2682 9240 2817 Moore Mesa (..oJ N .j:- �325 LITERATURE CITED Kufeld, R. C., J. H. Olterman, and D. C. Bowden. 1980. A helicopter quadrat census for mule deer on Uncompahgre Plateau, Colorado. J. Wildl. Manage. 44(3):632-639. c.., ~) Prepared by ~~.~~. ._~ ~ ,7~. .__~__. Allen E. Anderson Wildlife Researcher C ~ ~A ) .~ __.__,__ ~__ . _ � https://cpw.cvlcollections.org/files/original/6b1d5ef931c3f93ad74c26f678fc3ea9.pdf af67dd4e2340620c6ea4a7f35fb0333d PDF Text Text 1 JOB PROGRESS State of Project October 1981 REPORT Colorado ----~~~~~-No. 1 Work Plan No. Job Title: Waterfowl Period Covered: Personnel :. Migratory W-88-R-26 Bird Investigations Job No. Production 21 April 1 Surveys 1980 to 20 June 1980 Those cooperating in the 1980 survey included: M. Nail and staff, Monte Vista National Wildlife Refuge; J. Creasy and staff, Brown's Park National Wildlife Refuge; S. Petersburg, Dinosaur National Monument; F. N. Folks, Utah Division of Wildlife Resources; M. Bauman, J. Corey, M. DePra, H. Donoho, J. Ellenberger, J. Frothingham, J. Gray, D. Kenvin, J. Lorentzson, D. Masden; K. MOser, T. Rauch, W. Russell, G. Saville, S. Steinert, M. Szymczak, Colorado Division of Wildlife. ABSTRACT Water conditions for duck production were improved over 1979 in all areas of the state. Conditions for Canada goose (Branta canadensis) production were variable. Northcentral Colorado had excellent conditions, with the rest of the state having normal to poor conditions. The total number of duck breeding pairs was estimated at 55,326, 6% below the long term average. The mallard (Arias platyrhynchos) was the major breeding species. Five new transects were evaluated for inclusion in the air-ground count to determine visibility ratios in the San Luis Valley. The total estimated p.ost-nesting season Canada goose·populatibn in northwest ·Colorado was down 22% "from 1979, but near the long-term average. An air-ground Canada goose breeding pair survey was conducted on the Yampa and Little Snake rivers to evaluate a proposed change in survey techniques. Aerial counts on the Colorado, Gunnison and White rivers continue to show an increasing trend. Canada goose breeding pairs in the San Luis Valley increased 42% over 1979. �2 RECOMMENDATIONS San Luis Valley Duck Breeding Pair Survey: a. Extend transect air-ground H, 3 miles to the west, making transect H 10 linear miles. The additional area will cover the south 1/8 of a mile of sections 16, 17 and. 18 and the north 1/8 of a mile of sections 19, 20 and 21 in Township 38N, Range BE. b. Continue to count new air-ground transect BWT on an experimental basis. The use of the data from BWT will be at the discretion of the principle compiler of the entire survey and will depend upon whether· the distribution of ducks in respect to BWT transect is representative of duck distribution in the entire sample area. Transect BWT is 2 linear miles covering the north 1/4 mile of sections 15 and 16 in Township 3BN, Range 9E. c. Continue to conduct the San Luis Valley duck breeding the 2nd full week of May. d. Explore the possibility of redefining the waterfowl habitat with possible stratification of sample areas according to quality of habitat or duck distribution. Northwest a. Colorado pair survey Goose Surveys: .Conduct an aerial survey of Canada geese during the last week in April on the Yampa and Little Snake rivers. Record all geese observed as either lone birds, lone.birds with nest, pairs, pairs with nests or groups. Compile data collected according to the following river segments: Little Snake River - (1) Baggs, Wyoming to Powder Wash Bridge, (2) Powder Wash Bridge to Highway 31B Bridge, (3) Highway 31B Bridge to confluence with the Yampa River; Yampa River - (1) Highway 131 Bridge to Steamboat Springs, (2) Steamboat Springs to Hayden, (3) Hayden to Craig, (4) Craig to Juniper Canyon, (5) Juniper Canyon to Sunbeam, (6) Sunbeam to Cross Mountain Canyon, (7) Cross Mountain Canyon to Yampa Canyon. Every 5th year an airboat survey will be conducted f~om Craig to Yampa Canyon on the Yampa River and from the Powder Wash Bridge to the confluence with the Yampa River on the Little Snake River. �3 WATERFOWL PRODUCTION SURVEYS Michael R. Szymczak Steven F. Steinert P.N. OBJECTIVES 1. To estimate the number of duck breeding pairs, by species, major waterf~wl nesting areas in Colorado. on selected 2. To estimate the number of goose breeding pairs, and in some cases, obtain production data on selected goose nesting areas in Colorado. 3. Compile data and submit repor t s to appropriate state personnel and federal agencies for use in establishing hunting season recommendations. SEGMENT OBJECTIVES 1. To estimate the number of duck breeding pairs, by species, in the San Luis," Cache la Poudre, South Platte and Yampa Valleys, and in North Park and Brown's Park, using procedures presented in the PROGRAM NARRATIVE. 2. To estimate the number of goose breeding pairs in the San Luis Valley and obtain goose breeding pair and production data on the Yampa, Little Snake and Green rivers in northwest Colorado and in northcentral and west central Colorado using procedures presented in the PROGRAM NARRATIVE. 3. To evaluate aerial counts as a means to monitoring the status of the Canada goose nesting population on the Yampa and Little Snake rivers. 4. To reevalu~te the positioning in the San Luis Valley. 5. Compile data and submit appropriate METHODS of air-ground duck breeding pair transects reports. AND MATERIALS Present duck breeding pairs and production surveys consist of a breeding pair inventory of only major production areas. The 1980 duck breeding pair surveys were conducted during the period of May 13 to June 20. Surveys in North Park and the.Cache la Poudre and South Platte valleys were conducted exclusively from the air. Ground counts were made in the Yampa Valley and Brown's Park. �4 In the San Luis Valley aerial counts were adjusted for visibility by air-ground comparison ratios. Five new air-ground transects located in high duck density areas in the San Luis Valley were counted in an experimental attempt to increase the sample size of observed ducks on air-ground transects. A second flight was also conducted on selected transects on June 4th to see what changes there would be in number and composition of breeding pairs. Pair estimates for the Monte Vista National Wildlife Refuge in the San Luis Valley were obtained from nesting transects. All survey methods and sample areas for ducks remained the same as in previous years. Surveys of Canada goose production were conducted within the period of April 21 to June 20. On April 28 an aerial Canada goose breeding pair survey was conducted on the Yampa and Little Snake rivers. Geese observed were classified as singles, breeding pairs, non-breeding pairs and groups. The results of the April survey were compared with the results of the May ground survey to determine if aerial survey data will be sufficient to monitor the breeding population. Production estimates for Moffat County presented in this report including the Yampa and Little Snake rivers were made from as complete a count as possible of hatched or active nests and brood sizes. Population estimates for northcentral Colorado were obtained from counts of goslings and adults conducted from the ground during the period in which the birds were flightless. Estimates of the Colorado, Gunnison and White river populations were obtained from direct counts from a fixed-wing aircraft. In the San Luis Valley, a survey of expected productive Canada goose breeding pairs was conducted by flying transects in defined sample blocks located in production areas throughout the Valley. All flying was accomplished with a Cessna 185 aircraft. Two observers were used when sampling by transect while one observer was used when sampling sections or flying river courses. RESULTS AND DISCUSSION Water conditions for duck production are improved over 1979 in all areas of the state. Surface water in the San Luis Valley showed only minor improvement. Extensive spring rains in the eastern portion of the state created excellent conditions not only in the irrigated portion of the Cache la Poudre and South Platte valleys, but also in the adjacent prairie areas. Water conditions in North Park are excellent, however the nesting season will be delayed as a result of a later than usual spring in that area. Atypically early high water resulted in Canada goose nest flooding along. the rivers in the western portion of the state. Some nest flooding also occurred along the Rio Grande River in the San Luis Valley. Reservoirs were at capacity in northcentral Colorado and major spring storms did not occur, thus producing excellent conditions for nesting geese. �5 The total estimated number of duck breeding pairs increased to 55,326, 7% above the 1979 level but 6% below the long-term average (Table 1). The Cache la Poudre Valley population nearly doubled. The San Luis Valley population also increased but remained below the long-term average. All other areas declined from 1979 levels. The estimated mallard breeding population increased slightly over 1979 record low levels while redhead estimates remained at an atypically high level (Table 2). Table 1. Summary of Colorado's selected areas, 1980. duck breeding Total estimated San Luis Valley North Park2 South Platte Valley Cache la Poudre Valley Yampa Valley Brown's Park Total breeding 1980 1979 21,571 13,644 8,318 9,418 1,486 889 55,326 17,140 16,822 9,794 5,018 1,897 953 51,624 Area pair population :eairs Long-term average 1 27,228 16,467 7,416 4,094 2,653 1,119 58,977 estimates Percent in 1979 change From long-term average +25.9 -18.9 -15.1 +87.7 -21. 7 - 6.7 + 7.2 - 20.8 - 17.1 + 12.2 +130.0 - 44.0 - 20.6 6.2 From 1San Luis Valley and North Park averages are based on results of 1964 through 1979 and 1968 through 1979 surveys, respectively, because of changes in survey methods utilized prior to those dates. Figures for other areas are 24 year averages. 2Aerial counts corrected the San Luis Valley. by species Table 2. Species population. of Colorado's composition from visibility ratios obtained 1980 duck breeding Number of breeding :eairs pair Percent species composition 1954-79 1954-79 SEecies Mallard Blue-winged and cinnamon teal Gadwall Pintail Shoveler Green-winged teal Redhead American wigeon Other divers Total 1979 15,980 15,456 7,485 8,805 2,904 6,251 3,501 9,120 146 1,134 55,326 6,349 4,838 3,297 2,891 5,719 9,707 1,204 2,163 51,624 1980 1979 27,075 28.9 29.9 49.8 5,931 5,641 3,592 3,518 3,284 2,548 1,134 1,685 54,4081 13.5 15.9 5.2 11.3 6.3 16.5 0.3 2.0 12.3 9.4 6.4 5.6 11.1 18.8 2.3 4.2 10.9 10.4 6.6 6.5 6.0 4.7 2.1 3.1 average average. composition computed from data from all areas for the 24 year regardless of changes in survey method. lSpecies period 1980 in �6 The estimated post-nesting season Canada goose population located along river systems in northwest Colorado is down 22% from 1979 but near the 1967-79 average (Table 3). The decline resulted primarily from a reduction in gosling production on all surveyed sections (Table 4). Number of geese observed by ground counts are recorded by river section in Table 5. Comparison of the results of two independent surveys (aerial and ground) are presented in Table 6. The only area of agreement that might be expected between these two surveys is in the estimated number of breeding pairs. Normally the estimated breeding pairs obtained during the aerial count will exceed the estimate obtained from post hatch ground surveys because (1) all nests are not found since only islands are searched and (2) high water some time between nest establishment and the ground survey may destroy nests. Beginning in 1981, ground surveys will be conducted only every 3rd year on the Green River and every 5th year on the Yampa and Little Snake rivers. Aerial counts will be conducted each year on the Yampa and Little Snake rivers. Table 3. Total estimated trend areas, 1980. number of Canada geese observed, Moffat County Percent Area Yampa River Green River Brown's Park Dinosaur National Monument (Colorado-Utah)l Little Snake River Total lNot surveyed Table 4. Estimated areas, 1980. until change From 1967-79 average From 1979 1980 1979 1967-79 average 707 854 598 -17.2 +18.2 434 559 379 -22.4 +14.5 257 276 389 338 386 291 -33.9 -18.3 -33.4 - 5.2 1,674 2,140 1,654 -21.8 + 1.2 1970. numbers of Canada goose goslings, Moffat County trend Percent Area Yampa River Green River Brown's Park Dinosaur National Monument (Colorado-Utah)l Little Snake River Total lNot surveyed until 1970. 1967-79 average From 1979 change From 1967-79 average 1980 1979 98 199 174 -50.8 -43.7 170 225 154 -24.4 +10.4 139 60 172 104 160 94 -23.7 -42.3 -13.1 -36.2 467 700 582 -33.3 -19.8 �7 Table 5. Number of Canada geese observed areas in Moffat County, Colorado, 1980. Nonnesting adults Nesting pairs Area Yampa River Craig-Juniper Springs Juniper Springs-Cross Mtn. Lily Park Subtotal Green River Brown's Park Dinosaur National Monument (Colorado, Utah) Subtotal Little Snake River Total and estimated Total adults production on trend Estimated no. goslings1 Total geese 11 15 12 38 165 225 143 533 187 255 167 609 27 47 24 98 214 302 191 707 60 144 264 170 434 35 95 48 192 118 382 139 309 257 691 16 184 216 60 276 149 909 1,207 467 1,674 nests lCalculated using average brood size and number of successful number of broods observed, whichever is greater. or Table 6. Comparison of the number of geese observed on the April 28, 1980 aerial count with estimates obtained from the May 20-22, 1980 nest search and ground count, according to breeding status designation. Breeding pairs Area Yampa River Craig-Juniper Springs 15 Juniper SpringsCross Mountain 31 Lily Park 6 Total 52pr Little Snake Total Air count NonBreeding pairs Groups Total Breeding pairs 11 Ground NonBreeding pairs count Groups Total 4 157 187 16 3 65 30 22 ~pr 22 38 63 144 94 303 15 12 38pr 10 0 14pr 205 143 505 255 167 609 21 30 40 142 16 l3 158 216 73pr 98pr 103 445 54pr 27pr 663 825 �8 The aerial breeding pair survey of Canada geese in west central Colorado was conducted on April 21 and 22, 1980, about the same time as the 1978 survey was conducted. The 1977 and 1979 surveys were conducted in early May. Comparing 1980 results with 1978 indicates the population is continuing to increase (Table 7). Counts by river section are presented in Table 8. Table 7. on aerial Comparison of the number of Canada goose singles and pairs observed surveys in west central Colorado, spring, 1977-1980. Number 1980 Area White River Roaring Fork River Colorado River Glenwood Springs-5th Bridge Bridge-Utah Line Gunnison River Hotchkiss-Delta Delta-Grand Junction Total Table 8. West central Singles 1979 1978 observed 1977 1980 Pairs 1979 1978 1977 30 6 46 3 31 2 15 2 39 1 50 7 26 0 32 6 47 6 41 4 30 4 13 2 74 11 77 12 35 4 33 7 2 3 94 0 0 94 0 4 71 0 0 32 5 8 138 1 5 153 0 5 70 0 11 89 St. Colorado Canada goose breeding Area White River Meeker-Rio Blanco L. Rio Blanco L.-Rangely Rangely-Utah State Line Subtotal Colorado River Newcastle-Silt Silt-Rifle Rifle-Grand Valley Grand Valley-Debeque Debeque-Palisade Palisade-Grand Ave. Bridge Grand Ave. Bridge-Fruita Fruita-Horsethief Canyon Horsethief Canyon-Utah State Line Subtotal Roaring Fork River EI Jebel-Carbondale Gunnison River Hotchkiss-Delta Delta-Mesa County Line Mesa County Line-Whitewater Whitewater-Grand Junction Subtotal Total Singles pair survey, Pairs 1980. Groups 7 5 o 19 4 30 31 3 39 35 15 50 4 22 21 4 6 16 15 28 22 16 3 6 41 3 o o o 3 53 8 85 18 108 6 1 10 2 3 5 10 6 o o o 4 2 1 2 5 94 1 o 7 1 13 138 o o o 16 184 �9 The estimated number of productive breeding pairs of Canada geese in the San Luis Valley increased 42% from the 1979 level (Table 9). Increases in the number of nesting pair are substantial along the Rio Grande River between Del Norte and Monte Vista and on the Alamosa National Wildlife Refuge. Estimates of non-nesting pairs also increased substantially. Table 9. Results of San Luis Valley Year Canada goose breeding Nesting Projected pairs pair survey. total number Non-nesting 1975 Helicopter 59 59 1976 Helicopter Fixed-wing 92 77 63 69 1977 Helicopter Fixed-wing 100 100 101 91 1978 No counts 1979 Fixed-wing 99 96 1980 Fixed-wing 141 226 pairs Results of the 1979 Canada goose production survey in northcentral Colorado are presented in Table 10. The number of adult geef:e observed on trend areas is 14% above the 1979 level and 0% above the 11-y~ar avera~e (Table 11). Gosling production reached a record high level in 1980 as all trend areas except Loveland recorded increases over 1979 (Table 12). Table 10. 1980. Production area 'Nellington Results of northcentral Colorado No. broods Water area Terry Lake Launer Pond Douglas Lake Dry Creek Reservoir North Poudre No. 2 North Poudre No. 5 North Poudre No. 6 Bureau of Standards Divide No.8 Elder Reservoir Annex No.8 Van Sant Pond Cobb Lake Dale Pond Watson Lake Curtis Lake Beghtol Lake Subtotal goose census, 12 2 Pond 1 2 0 0 0 1 4 1 0 0 5 0 2 Total no. goslings 53 47 11 47 7 0 0 0 6 18 3 0 76 18 43 0 11 340 June 12 and 13, Total no. adults and yearlings 60 24 23 18 4 4 16 25 68 30 32 5 246 10 147 57 4 773 Total birds 113 71 34 65 11 4 16 25 74 48 35 5 322 28 190 57 15 1,113 �10 Table 10. Results 1980. (continued) Production area Fort Collins Loveland Boulder of nbrthcentral Colorado Water area Peterson Ponds Dixon Reservoir Miller Ponds College Lake Dean Acres Andrijeski Marsh Claymore Sterling Gravel Pits Lindenmeier Grey Lakes Novaks Flatiron Anderson's Parkwood Kitchel Timnath Romily Fossil Creek Schuelke Lake Wolaver Pond Subtotal goose census, No. broods 0 3 7 2 0 1 4 4 5 2 0 6 6 8 20 2 5 3 3 Flatiron Reservoir Boedecker Reservoir Marina Reservoir Flatiron Gravel Pits Kauffman Gravel Pits Big Thompson River McNeil Reservoir Welch Reservoir Reservoir No. 12 Subtotal 0 Ish Lake Crystal Lake Terry Lake Faivre Pond Rest Home Pond Valmont Reservoir. Boulder Valley Farm Pond King Pond Eddy Pond Angus Ranch Pond Subtotal 3 0 9 1 5 3 0 8 6 3 3 0 1 1 June 12 and l3, Total no. goslings Total no. adults and yearlings Total birds 0 10 29· 34 0 4 13 108 11 21 8 0 27 28 34 95 7 14 14 10 467 17 18 32 244 28 2 41 198 42 36 4 8 45 64 19 97 25 27 13 10 970 17 28 61 278 28 6 54 306 53 57 12 8 72 92 53 192 32 41 27 20 1,437 0 28 2 15 11 0 33 25 12 126 11 50 4 39 16 12 163 50 63 408 11 78 6 54 27 12 196 75 75 534 15 0 50 24 31 83 9 0 5 3 220 6 23 73 41 119 l30 6 2 5 2 407 21 23 123 65 150 213 15 2 10 5 627 �11 Table 10. Results 1980. (continued) Production area of northcentral Total no. goslings No. broods Water area 14 3 0 6 8 8 11 3 0 0 8 1 2 0 7 5 5 0 2 Ketring Centennial Columbine C.C. Chatfield Golf Course Bowles Lake King's Pond Tule Lakes Grant Ponds Marston Reservoir Pinehurst C.C. Clarefield Reservoir Ward Reservoir Kendrick Lake Federal Center Pond Sloan's Lake Stanley Lake Denver City Park Colorado Blvd. @ Quincy Blackmer Reservoir Subtotal Denver goose census, Colorado Grand Total Table 11. production Number of adult Canada geese observed trend areas, 1980. June 12 and 13, Total no. adults and yearlings Total birds ---- 50 15 0 27 38 30 44 11 0 0 28 2 9 0 26 22 18 0 6 326 71 23 31 54 127 113 66 98 15 1 36 2 24 2 277 45 57 14 5 1,061 121 38 31 81 165 143 l10 109 15 1 64 4 33 2 303 67 75 14 l1 1,387 1,479 3,619 5,098 in northcentral Colorado 1969-79 Percent From No. of adults change From 1980 1979 average 1979 Wellington Fort Collins Loveland Boulder Denver 773 970 408 407 1,061 711 663 374 286 1,147 724 705 232 547 1,220 + 8.7 +46.4 + 9.1 +42.3 - 7.5 + 6.8 +37.6 +75.9 -25.6 -l3.0 Total 3,619 3,181 3,428 +13.8 + 5.6 Area 1.969-79 �12 Table 12. production Number of Canada goose goslings produced trend areas, 1980. in northcentral No. of goslings Area Wellington Fort Collins Loveland Boulder Denver Total EVALUATION Air-Ground OF EXPERIMENTAL Colorado 1969-79 Percent From 1980 1979 average 1979 340 467 126 220 326 290 378 158 181 310 252 293 105 208 281 +17.2 +23.5 -20.3 +21.5 + 5.2 +35.0 +59.4 +20.0 + 5.8 +16.0 1,479 1,317 1,l39 +12.3 +29.9 CHANGES change From 1969-79 IN THE SAN LUIS VALLEY DUCK SURVEY Transects Permanent changes in the methods of irrigation along with almost continuous drought in the San Luis Valley have resulted in degradation of habitat for waterfowl in recent years. The methods of estimating the San Luis Valley duck breeding pair population have remained essentially the same since 1963. Recently, confidence in the estimates has been declining. One problem has been the reduction in the number of ducks on air-ground comparison transects. Ducks observed on these transects are used to calculate visibility ratios for the various species. Therefore, a reduction in numbers of ducks means a reduction in the sample size used to calc~late the ratios increasing the possibility of error in the ratio. In 1979, a total of only 279 indicated breeding pairs were observed from the ground on these transects. From 1972 through 1978, the total indicated breeding pairs were 876, 635, 900, 646, 591, 385 and 435, respectively. In 1980, 4 additional air-ground transects were established and one of the original transects was extended for 2 miles, creating an additional 16 linear miles of sample area 1/4 mile wide. Portions of two transects were located on the Monte Vista National Wildlife Refuge and the two others were located through currently good wetland habitat in areas adjacent to the Rio Grande River. A total of 414 indicated breeding pairs was observed from the ground on the 5 new areas or 24.4 pairs/linear mile. Only 3.5 pairs/ linear mile were observed on the original transects that are counted each year. The new transects, if utilized, would have essentially doubled the sample from the 428 pairs observed on the original transects to 842 pairs. In 1980, only the data collected on the original transects were used in calculating the visibility ratio. �13 An evaluation of the 5 new segments follows: Experimental Transects 1 and 2 -- These transects are 5 and 4 miles in length with 3 and 2 miles, respectively, being on the Monte Vista Refuge. Most of the birds on both transects were on the refuge segments. Even though the transects were selected to extend through areas of the refuge with minimum water development, duck densities were quite high. Therefore, these transects are not considered useful in establishing air-ground ratios, since the habitat, and resulting duck densities, are not representative of what occurs across the remainder of the valley. Experimental Extended H -- This segment is located on the west end of airground H. The eastern 1 mile contains very little waterfowl habitat and the west 2 miles, when surveyed in 1980, contained numerous ponds, ditches and flooded meadows. All of the ducks observed were on the western section. The habitat is considered representative of waterfowl habitat in the valley. Duck densities will vary depending on the status of irrigation during the census. However, it is appropriate to utilize counts on these habitat types to establish visibility ratios. It is recommended that this three mile segment be retained (see recommendations for appropriate description). Experimental Management Area -- This transect is approximately 3 linear miles along the southern boundary of the Rio Grande Management Area. The transect is considered typical of river bottom habitat in the valley. Unfortunately, the course of the Rio Grande River made it impossible to count all portions of the transect and the section should not be used for future counts. Experimental Blue-winged Teal Club -- This 2 mile transect includes pond and flooded meadow habitat adjac~nt to the river bottom. In 1980, 123 pairs were recorded along this transect from the ground. If these ducks would have been used in calculation of the visibility ratio, 22% of the pairs used to establish the ratio would have been from this 2 mile segment. The current distribution of ducks throughout the valley is not known. It has been assumed that degradation of habitat throughout the valley has resulted in concentrating birds in the remaining wetland areas such as river bottom habitat. If 22% of the total birds observed on all transects in the San Luis Valley occur in river bottom areas such as represented by this transect, then calculating visibility ratios using data from this transect is valid. We recommend that counts on this transect be continued experimentally. Until more information is gathered concerning the distribution of ducks in the valley, the decision as whether to use the data from this transect should be made on a year to year basis. SURVEY TIMING Historically, the San Luis Valley duck breeding pair survey has been conducted during the second week in May. Recently some people have suggested that, because in changes in irrigation practices, a later survey would provide a better measure of the population. In 1980, an aerial count of breeding pairs on selected transects was conducted on June 4 and the results compared with those obtained on May 12 (Table 13). More birds �14 were observed on June 4 and there was considerable variation in the species composition between the 2 periods. More redheads (Aythya americana) were observed than any other species and most of those were found in flocks. Shoveler flocks were prevalent on May 12. There was essentially no variation in the mallard count between the two periods. Since the mallard is the major species and visibility ratios are more stable than for other species, we recommend that the survey continue to be conducted during the second week in May. Table 13. Comparison of breeding pairs observed varying dates in the San Luis Valley, 1980. Total breeding May 12 Species Mallard Pintail Cinnamon Teal Shoveler Gadwall Blue-winged Teal Redheads Green-winged Teal i''''Z''/;J'' " r , :_-,;7 by -> ~ tr . ./ (_ ':-/ 5r7:-?V'<-e ;7 Michael R. Szymczak Wildlife Researcher » ;:;:::::r C ~~@!~ Steven F. Steinerf Wildlife Technician III pairs observed June 4 1 16 6 75 o 3 120 14 248 320 ./ ." <#~~ '1t0~ y ...-; surveys on 85 88 8 10 67 39 8 28 Total Pairs Prepared on aerial ,'/.' / �15 October 1981 JOB FINAL REPORT State of Colorado ---------------------W-88-R-26 Proj~ct: No. 2 Work Plan No. Job Title: Migratory Job No. Bird Investigations 6 Studies of Canada Goose Populations in Colorado Transplant Areas Period Covered: Personnel: April 1, 1970 to March 31, 1981 J. Corey, Dale Flenthrope, Dean Flenthrope, R. Hopper, W. Rutherford, S. Steinert, and M. Szymczak, Colorado Division of Wildlife. ABSTR..<\CT Of the 3,121 large Canada geese banded in northcentral Colorado, posthunting season 1968 through 1975, 822 were reported recovered through 1978. Fifty-seven percent of those recoveries occurred in Colorado; all but 4 percent in the traditional Hi-Line range. Saskatchewan accounted for 17 percent of the recoveries, followed by Alberta and Montana with 8 percent each, Wyoming 3 percent, and Nebraska and New Mexico 1 percent each. One hundred thirty-nine of the 769 small Canadas banded posthunting season 1971 through 1975 in northcentral Colorado were recovered through 1978. Colorado accounted for 41 percent of the recoveries which, geographically, were equally divided between northcentral Colorado and the Shortgrass Prairie Population range in southeast Colorado. Alberta recorded 31 percent of the recoveries and Saskatchewan 15 percent. The small Canadas banded post-season in northcentral Colorado seem to be oriented more toward the western portion of the Shortgrass Prairie Population range.than those banded in sou(heast Colorado. The survival rate of large Canadas banded post-hunting season in northcentral Colorado in 1970 through 1975 was estimated to be 73.23 + 2.22 percent, with an estimated mean recovery rate of 7.92 + 0.44 perc_ent and a mean life span of 3.21 + 0.31 years. A total of 180 Canada geese, 39 adults and 141 goslings were banded on brood rearing areas in the San Luis Valley. Trapping since 1974 has resulted in a total of 150 adults and 575 goslings being banded. The following is a summarized list of findings along with the appropriate publication in which the results can be found for objectives covered under this study since 1970. �16 Hi-Line Population Harvest hunting pressure and season recommendations -- 1970-1973: Szymczak, M. R. 1975. Canada goose restoration along the foothills of Colorado. Colo. Div. Wildl. Publ. No. 31. 63p. Fall and winter food as indicated by analysis of fecal droppings: Szymczak, M. R., and R. C. Staffon. 1975. Studies of Canada goose populations in Colorado transplant areas. Colo. Div. Wildl. Fed. Aid Proj. W-88-R, Work Plan 2, Job 6 Progress Rep., Oct.: 53-Ill. Staffon, R. C. 1976. Local movements and food habits of Canada geese wintering in Colorado. M.S. Thesis. Colo. State Univ., Fort Collins. 127p. Breeding range of Hi-Line Canada geese: Szymczak, M. R. 1980. Studies of Canada goose population in Colorado transplant areas. Colo. Div. Wildl. Fed. Aid Proj. W-88-R, Work Plan 2, Job 6 Progress Rep., Oct.: 15-34. Distribution of harvest and estimate of survival of resident geese: Staffon, R. C. 1976. Local movements and food habits of Canada geese wintering in Colorado. M.S. Thesis. Colo. State Univ., Fort Collins. 127p. Szymczak, M. R., R. C. Staffon, and J. F. Corey. 1981. Distribution and harvest of Canada geese nesting along the foothills of Colorado. Colo. Div. Wildl., Spec. Rep. No. 49. In press. San Luis Valley Population Harvest, hunting pressure and season recommendations 1970-1975: Szymczak, M. R. 1976a. Studies of Canada goose populations in Colorado transplant areas. Colo. Div. Wildl. Fed. Aid Proj. W-88-R, Work Plan 2, Job 6 Progress Rep., Oct.: 41-63. Migration routes and harvest patterns of wintering geese: Szymczak, M. R. 1976b. Investigations of Canada geese in the San Luis Valley. Colo. Div. Wildl. Fed. Aid Proj. W-88-R, Work Plan 2, Job 8 Final Rep., Oct.:67-86. Westcentral Harvest, Population hunting pressure and season recommendations 1971-1975: Szymczak, M. R. 1976a. Studies of Canada goose populations in Colorado transplant areas. Colo. Div. Wildl. Fed. Aid Proj. W-88-R, Work Plan 2, Job 6 Progress Rep., Oct.:41-63. �17 STUDIES OF CANADA GOOSE POPULATIONS IN COLORADO TRANSPLANT AREAS Michael R. Szymczak These investigations, which began in 1970, were initiated in order to provide information needed to establish sound hunting regulations on an annual basis as well as ascertain various ecological facts which are pertinent to long-term management for goose populations which had been established through restoration programs in Colorado. The result has been a number of unrelated studies being conducted under a single job. Many of these studies have resulted in recommendations for population management which have been adopted by biologists and administrators responsible for management of the various populations. Other studies have resulted in information that will be used in the future. All of the results will be referred to in this report, but not presented in detail, since most have been previously published in some form. Publication of this final report does not mean that we have completed all studies that are needed on the populations established through transplants. Future studies will be conducted under separate jobs that will deal more specifically with particular objectives. OBJECTIVES There were two overall objectives during the term of this job. which was in effect from April 1970 through March 1976, was: The first, To investigate the status of resident and migrant Canada goose flocks and their interrelationships in areas in which populations have been established through transplant programs in Colorado. The second objective, in effect from April more specific and had three parts: 1976 through March 1981, was 1. To document the breeding range of Canada geese wintering in northcentral Colorado. 2. To document the distribution of harvest survival rate of the resident foothills population. and estimate the Canada goose 3. To document the distribution of harvest nesting in the San Luis Valley. of Canada geese METHODS AND MATERIALS Because of the diverse nature of this study, the procedures to obtain specific objectives varied considerably. Most of these procedures have been well documented in reports and publications reSUlting from the study and therefore will not be repeated here. However, the final results of �18 two objectives, (1) harvest patterns of wintering Hi-Line Canada geese and (2) estimates of mortality of that population, have not appeared in previous reports. Harvest patterns were documented by recording the distribution of harvest by degree block of Canada geese banded post-season in northcentral Colorado from 1968 through 1975 and recovered through the 1978-79 hunting season. The estimates of mortality were obtained by recording the distribution of recoveries over time for large Canada geese banded post-season in northcentral Colorado from January 1970 through January 1975. Because of small sample sizes and the inability to positively identify immature females at the time of banding, all ages and sexes were pooled for this analysis. The resulting recovery matrix was analyzed using computer program ESTIMATE and the best model selected as discussed in Brownie et al. (1978). In this report, mortality estimates will be referred to in the reciprocal, as estimates of survival. All recovery information was compiled from annual summary reports to the bander supplied by the U.S. Fish and Wildlife Service Bird Banding Laboratory. From 1971 through 1980, adult and young Canada geese were trapped annually in mid to late June on brood rearing areas in the San Luis Valley in order to obtain information on the distribution of harvest produced on the various production areas. In 1980, birds were trapped during the period of June 17 to June 20. Banding schedules were submitted to the Bird Banding Laboratory. RESULTS AND DISCUSSION Studies under this job were conducted on three populations: (1) the Hi-Line Population in northcentral Colorado, (2) the San Luis Valley Population in southcentral Colorado, and (3) the Westcentral Population located along the Colorado River drainage in the westcentral portion of the state. The results will be discussed on the basis of the specific populations. Hi-Line Harvest, Hunting Pressure Population and Season Recommendations Harvest and hunting pressure was estimated annually from 1970 through the 1973 hunting seasons under this job utilizing a survey of hunters who obtained a special permit indicating their intention to hunt geese in the northcentral Colorado area. The survey was first established in 1964 and was terminated in 1973. The results of all 10 years of this survey appear in Szymczak (1975). Input into annual season recommendation began under this project in 1970 and terminated in respect to these project documents in 1975. Input included an analysis of the harvest survey, winter inventory data and reports of nesting and production surveys. Some aspects of regulation changes are referred to in Szymczak (1975). �19 Migration Routes and/or Harvest Patterns From 1968 through 1975, 3,890 Canada geese were banded post-season in northcentral Colorado. Eighty percent of the birds were classified as large Canada geese, probably members of the Hi-Line Population, while 20 percent, all which were banded during the 1971-1975 period, were small Canada geese, assumed to be members of the Shortgrass Prairie nesting population. Over 26 percent, or 822, of the large Canadas were reported ,harvested through the 1978-79 hunting season and the geographic distribution of harvest is presented in Figure 1. Colorado was the major state of harvest accounting for 57 percent of the recoveries. About 96 percent of the Colorado recoveries occurred in the traditional Hi-Line Population range in the northcentral part of the state. Saskatchewan accounted for 17 percent of the recoveries with 82 percent reported taken in the traditional Hi-Line area in the 4 degree blocks in the extreme southwest portion of the Province. Alberta and Montana each accounted for about 8 percent of the recoveries. About 63 percent of the Montana recoveries occurred in the historical Hi-Line range in the northcentral portion of the state, while 33 percent were reported taken in the Yellowstone-Big Horn river area in southeastern Montana. Wyoming accounted for 3 percent of the recoveries with 71 percent reported taken in the Hi-Line wintering range in extreme southeast portion of the state. Two percent of the recoveries were reported taken in Nebraska and only 1 percent in New Mexico. One hundred, thirty-nine of the 769 small Canada geese banded from 1971 through 1975 were reported recovered through the 1978-79 hunting season. The geographic distribution of the recoveries is presented in Figure 2. The distribution generally followed the range of the Shortgrass Prairie Population as reported by Grieb (1970). However, in Colorado, the numbers of recoveries were equally divided between the Shortgrass Prairie Population range in southeast Colorado and the Hi-Line Population range in northcentral Colorado. Colorado accounted for 41 percent of the recoveries, followed by Alberta with 31 percent and Saskatchewan with 15 percent. Texas, a major wintering area for a portion of the Shortgrass Prairie geese, accounted for only 2 percent of the recoveries from Colorado bandings. For small Canada geese banded post-season in southeast Colorado since 1971, Colorado has accounted for 38 percent of the recoveries, but only 14 percent of the Colorado recoveries occurred in the northcentral portion of the state (Szymczak 1979). In Canada, recoveries of southeast Colorado banded birds were more equally distributed between Alberta (21%) and Saskatchewan (20%) than those from northcentral Colorado bandings. The small birds in northcentral Colorado seem to be more westerly oriented than their counterparts wintering in the southeast portion of the state even though considerable interchange between the two wintering areas does occur from year to year as indicated by band recovery information (Szymczak 1979). �Figure 1. Distribution hunting season, resulting $ .ts. ! $ 1\ I '1- \ / / ot ~22 recoveries of large Canada fro~ 3,121 birds banded post$eason /"'1---1 ... --L._ / (f E '0' ~ 11--I--LL /) / r § geese reported taken through the 197e-79 in north-central Colorado 1968-1975. ~ ~ 8, ~ 100' t.. 'j.. I I ~. ..•.•...._/ ,, / /... _ i, �N 4- o t/) (I)~ ~ ...•••. ~-....;_ --ot, '.,.. C "'0 (I) 10 1- ._ ..0 g; -&" o 1u·_ <o r-, 0"\ (1)..0 10"\ CV'\ . ~'. ':' .•... ,' �22 Survival and Recovery Rates Subjecting the banding and recovery matrix (Table 1) of geese classified as large Canadas at time of banding to program ESTIMATE (Brownie, et al. 1978) resulted in the selection of estimates indicated under Model 2, which assumes constant survival but time-specific recovery rates. The constant survival estimate is 73.23 + 2.22 percent. The mean recovery rate is 7.92 + 0.44 percent while the mean life span was 3.21 + 0.31 years. Th~ "half-life" or an estimate of when 50 percent of the p~pulation will be dead is 2.21 years (MLS x 0.69, Brownie et a l . 1978). A major change in hunting regulations occurred in northcentral Colorado between the 1972 and 1973 hunting seasons when the seasonal bag limit of 6 years was eliminated. A test of recovery rates before and after the regulation change indicated no difference in rates between the two periods (Z = -.0919, p > 0.46). . Table 1. Banding and recovery matrix and estimates of recovery rates, survival rate and mean life span of large Canada geese banded in northcentral Colorado, post-season 1970-1975 and recovered through the 1979-80 hunting season. .YEAR 1970 1971 1972 1973 1974 1975 NUMBER BANDED 185 745 579 170 . 672 187 -- - RECOVERY - 8 0 0 0 0 0 16 86 0 0 0 0 -- 6 49 60 0 0 0 - - 5 40 36 9 0 0 - 0 26 13 9 52 0 - RECOVERY I 1. 2 3 4 5 6 ESTIMATE STANDARD 4.180 11.634 9.353 8.481 6.925 6.946 - 1 12 16 12 28 13 7 19 .11 10 28 12 1 7 11 8 17 4 1 7 7 3 22 4 2 1 4 0 17 4 - RATE ERROR F{I) (PCT) 95% CONFIDENCE 1.464 1.084 .865 .944 .718 .821 ARITHMETIC MEAN RECOVERY RATE 7.92 STANDARD ERROR OF MEAN RECOVERY RATE .44 95% CONFIDENCE INTERVAL FOR MEAN RECOVERY RATE 1.311 9.509 7.658 6.632 5.519 5.338 INTERVAL - 7.048 - 13.759 - 11. 048 - 10.331 - 8.332 - 8.555 7.05 - 8.79 CONSTANT SURVIVAL RATE (PCT) - 73.23 STANDARD ERROR OF THE CONSTANT SURVIVAL RATE 2.22 95% CONFIDENCE INTERVAL.FOR THE CONSTANT SURVIVAL RATE MEAN LIFE SPAN AS AN ADULT 3.21 STANDARD ERROR OF THE MEAN LiFE SPAN 95% CONFIDENCE INTERVAL OF LIFE SPAN MATRIX .31 2.68 - 3.94 68.88 - 77.59 �23 The survival of adult large geese of the northcentral Colorado resident population, banded during the post-nesting season period from 1974 through 1978, was estimated to be 68.88 + 3.1% (Szymczak et al., In Press 1981); numerically lower but not signifIcantly different (Z = 1.14, p > 12) than the birds banded post-season which included migrants as well as residents. In contrast, the mean recovery rate of the resident adult birds was si~nificantly lower (5.55 ± 0.43%, Z = -3.94, p < 0.01) than the winter banded birds. The survival rate of northcentral Colorado banded birds was lower than the estimate for small Canada geese banded in southeast Colorado during the same years (77.96 + 2.33%), however, statistically, the hypothesis of no difference could not be rejected (Z = 1.47, P > 0.07). The mean recovery rate (5.05 ± 0.29%) of southeast Colorado banded birds was significantly lower (Z = 5.71, p < 0.01) than those banded in northcentral Colorado. Breeding Range From 1970 through 1979, an attempt was made to define the breeding range of the Hi-Line Canada Goose Population wintering in northcentral Colorado through analysis of banding and recovery information. Birds banded outside of Colorado that were reported recovered during the hunting season or recaptured during post-season banding operations in northcentral Colorado were recorded according to area of banding and age at time of banding. The results, as presented in depth in Szymczak (1980) were hampered by the lack of banding in· breeding areas, particularly Saskatchewan. Yet, some good information was obtained. Fall and Winter Food as Indicated by Analysis of Fecal Droppings The complete findings of this portion of the study which was conducted in the fall and winter of 1974 can be found in Szymczak and Staffon (1975) and Staffon (1976). A formal publication is now in preparation. Distribution of Harvest and Estimate of Survival of Resident Geese The primary results of the five-year banding program as well as the observation of collared geese as they relate to this objective are published in Szymczak et a~. (1981). Additional information concerning the distribution of resident birds is presented in Staffon (1976). San Luis Valley Population Harvest, Hunting Pressure and Season Recommendations Harvest and hunting pressure we re estimated annually from 1970 through the 1975 hunting seasons under this job utilizing a survey of hunters who obtained a special permit indicating their intentions to hunt geese in the San Luis Valley. The results of the survey for those five years are presented in Szymczak (1976a)~ The survey has been continued, as recommended, by Colorado southwest regional personnel. �24 Input into annual season recommendations began under this project in 1970 and terminated after the 1975 hunting season. The recommendations were based on harvest, hunting success, winter counts and estimates of breeding population. Some aspects of the regulation changes are referred to in Szymczak (1976). Migration Routes and Harvest Patterns All information concerning migration ing geese. were presented in Szymczak Distribution of Harvest of Nesting of Wintering Geese routes and harvest (1976b). patterns of winter- Geese A total of 180 Canada geese were banded in the San Luis Valley on 7 different areas (Table 2). The total number banded since 1974 was 725 birds (Table 3). The recoveries from these bandings were scheduled to be analyzed this year. However, the analysis will be delayed until at least 1 year's recoveries from the 1980 bandings are available. The results will be published under W-88-R, Work Plan 2, Job 10, Distributional Characteristics of Some Populations of Canada Geese Inhabiting Colorado. Table 2. Location and number, by age and sex, of Canada geese banded on production areas in the San Luis Valley 1980. Adults Male Female Location Monte Vista National Refuge Rio Grande Management National Blanca Wildlife Russell Lakes Baca Grande Total Total Wildlife Area Kempf Farm Alamosa Goslings Male Female 3 4 21 13 41 0 0 5 3 8 5 7 8 12 32 Wildlife Refuge 3 3 11 10 27 Management Area 2 3 6 5 16 4 5 26 16 51 0 0 3 2 5 17 22 80 61 180 �25 Table 3. Numbers of Canada geese banded on production in the San Luis Valley, 1974-1980. and molting areas Year Adults Goslings Total 1974 1975 1976 1977 1978 1979 1980 4 14 55 4 18 16 39 6 5 149 33 121 120 141 10 19 204 37 139 136 180 150 575 725 Totals Westcentral Harvest, Hunting Pressure Colorado and Season Recommendations Harvest and hunting pressure was estimated annually from 1971 through the 1975 hunting season under this job utilizing a survey of hunters who obtained a special permit indicating their intentions to hunt geese in westcentral Colorado. The results of the five years of survey are presented in Szymczak (1976a). The survey has been continued, as recommended, by Colorado southwest and northwest regional personnel. Input into annual season recommendations began in 1971 and terminated under the project in 1975. The recommendations were based on harvest, hunting success, winter counts and estimates of breeding populations through the entire Rocky Mountain Population range of which westcentral Colorado is a portion. These recommendations have continued, in part, but not as an objective of this study. LITERATURE CITED Brownie, Cavell, D. R. Anderson, K. P. Burnham, and D. S. Robson. Statistical inference from band recovery data - a handbook. Interior, Fish Wildl. Servo Resource Publ. No. 131. 212p. Grieb, J. R. 1970. The shortgrass Monogr. No. 22:1-49. prairie 1978. U.S. Dep. Canada goose population. Staffon, R. C. 1976. Local movements and food habits of Canada geese wintering in Colorado. M.S. Thesis. Colo. State Univ., Fort Collins. 127p. Wildl. �26 Szymczak, M. R. 1975. Canada goose restoration Colorado. Colo. Div. Wildl. Publ. No. 31. along the foothills 63p. of 1976a. Studies of Canada goose populations in Colorado transplant areas. Colo. Div. Wildl. Fed. Aid Proj. W-88-R, Work Plan 2, Job 6 Progress Rep., Oct.:41-63. 1976b. Investigations of Canada geese in the San Luis Valley. Colo. Div. Wildl. Fed. Aid Proj. W-88-R, Work Plan 2, Job 8 Final Rep., Oct. :67-86. 1979. Monitor banding of the shortgrass pra1r1e Canada goose population in southeastern Colorado. Colo. Div. Wildl. Fed. Aid Proj. W-88-R, Work Plan 2, Job 9 Progress Rep., Oct.:27-39. 1980. Studies of Canada goose populations in Colorado transplant areas. Colo. Div. Wildl. Fed. Aid Proj. W-88-R, Work Plan 2, Job 6 Progress Rep., Oct.:15-34. _______ , and R. Staffon. 1975. Colorado transplant areas. Work Plan 2, Job 6 Progress Studies of Canada goose populations in Colo. Div. Wildl. Fed. Aid Proj. W-88-R, Rep., Oct.:53-111. ------- , R. C. Staffon, and J. F. Corey. 1981. of Canada geese nesting along the foothills Wildl. Spec. Rep. No. 49. In Press. Prepared by _,,/ m,--,-..I&-U=, =''\..4.-=-,,,,=,,,,,,,,!p~1-:::7?"''''''::'_~~,-,,::""~f-/h....-.""'7=~:::.r....::_:::l=i) __ Michael R. Si~~ Wildlife Researcher C Distribution of Colorado. and harvest Colo. Div. �October 1981 27 JOB PROGRESS REPORT State of Colorado Project No. w-88-R-26 2 Work Plan No. Job Title: _______Mi_·~gratory Bird Inyesti~atio~~ Job No. 9 Monitor Banding of the Shortgrass Prairie Canada Goose Population in Southeastern Colorado Period Covered: Personnel: 19 January 1981 - 31 March 1981 Jennifer Slater, B. McCloskey, J. Corey, S. Steinert, M. Potter, J. Young, and M. Szymczak, Colorado Division of Wildlife. ABSTRACT Post-season trapping in southeastern Colorado resulted in 416 geese being banded in January 1981. In 1978-79, according to the distribution of recoveries,harvest in Canada north of 530 latitude was at its lowest level since 1971-72 while harvest in Texas was at a record high 15 percent. In 1979-80, an atypically high 57 percent of the recoveries occurred in Canada while a record low 19 percent was reported taken in Colorado. The mean survival rate during the 1967-79 period was 73.78 ± 1.21 percent compared to 74.44 ± 1.61 during the 1958-65 period. The mean recovery rate was 4.80 ± 0.19 percent during the 1967-79 period compared to 8.78 ± 0.37 during the 1958-65 period. _ ��29 MONITOR BANDING OF THE SHORTGRASS PRAIRIE CANADA GOOSE POPULATION IN SOUTHEASTERN COLORADO Michael R. Szymczak P. N. OBJECTIVE To continually document, through monitor banding and analysis of recovery data the annual and long term status of the southeastern Colorado (Arkansas Valley) segment of the shortgrass prairie Canada goose population. SEGMENT OBJECTIVES 1. Band a nu.namum of 1,000 Canada geese in southeastern the post-season period. 2. Prepare and submit banding schedules, reports, and progress reports. 3. Analyze band recovery data for geese banded in southeast Colorado to determine (1) distribution of harvest, (2) recovery rate, and (3) survival rate. METHODS band recovery Colorado during and return AND MATERIALS All birds banded in southeast Colorado in 1981 were captured with baited cannon nets, and the age and sex. determined through cloacal and tail feather examination. All banding schedules, including recapture information, were submitted to the U.S. Fish and Wildlife Service's Bird Banding Laboratory, Patuxent, Maryland. Information on the distribution of harvest and the scheme of recoveries by year of harvest was obtained from computer printouts that are provided periodically by the Bird Banding Laboratory. The banding and recovery matrices were subjected to programs BROWNIE and ESTIMATE in order to obtain estimates of recovery and survival rates (Brownie et al. 1978). RESULTS AND DISCUSSION Trapping and Banding Trapping efforts at Turk's Pond in southeast Colorado resulted in 416 geese being banded in late January 1981. The number of birds banded by age and sex are as follows: adult males 103, adult females 134, immature males 78, immature females 86, adult unknown 14, and imma ture unknown 1. �30 Distribution of Harvest The distribution of harvest during the 1978-79 season and 1979-80 season were quite different (Table 1). On a percent basis, harvest in areas north of 530 latitude in 1978-79 was at its lowest level since 1971-72 while harvest in Texas accounted for a record high 15 percent of the recoveries. Harvest in the north-central portion of Colorado reached a record high 10.3 percent during the 1978-79 season. In 1979-80, an atypically high 58 percent of the recoveries occurred in Canada while a record low 19 percent was reported taken in Colorado. The total annual number of recoveries continues to reflect poor trapping success in southeast Colorado in recent years. Recovery and Survival Rates Banding and recovery summaries from bandings beginning in 1967 were analyzed to obtain recovery and survival rate estimates. Since two age groups, adults (ASY) and juveniles (SY), were identified during post-season banding, the data were first tested to see whether juveniles and adults have similar recovery and survival rates. The tests involved model HO' which assumes recovery and survival rates to be independent of age, against HI, which permits those parameters to differ between juveniles and adults (Brownie et al. 1978, p. 56-113). Both models allow recovery and survival rates to vary from year to year. The results of the contingency chi-square tests of data sets provided no evidence that recovery and survival rates for juveniles and adults were different (Table 2). Additional analysis, using a Z-test statistic also indicated there is no evidence that survival rates of the two age groups are different, but some indication of a difference in juvenile and adult recovery rates. Data for 1958-65 bandings which was analyzed by Szymczak (1979) also indicated ages should be pooled (Table 2). Adult and juvenile bandings and recoveries for the 1967-79 period were then pooled (Table 3) and tested for model fitness using models developed for bandings of adult birds only (Brownie et al. 1978, p. 13-55). Analysis and testing of 1967-79 bandings indicate Modell, which assumes time-specific survival and recovery rates is the appropriate model for the data set. The same model was appropriate for 1958-65 data (Szymczak 1979). Survival and recovery estimates under Modell are presented in Table 4. Average survival during the two banding periods was nearly equal (74.44 ± 1.61 - 1958-65; 73.78 ± 1.21 - 1967-79, Table 4). Recovery rates were considerably different (8.78 ± 0.37 - 1958-65; 4.80 ± 0.19 1967-79) with only nhe recovery rate in 1979 approaching the pre-1965 level. A test of the difference between the two recovery rate data sets was highly significant (Z = 8.33, p < .0001). �Table Percentages of total band recoveries, Arkansas Valley post-season bandings, by area and year of recovery, all bandings. 1. Area 195155 Five ~ear averages 19561966196160 70 65 Recovery year 19741975197675 76 77 197175 197172 197273 197374 0.5 2.4 0.5 3.3 2.8 4.2 1.4 8.5 0.4 5.3 1.3 7.1 17.6 16.9 16.0 14.7 1.3 19.5 16.8 2.2 1.8 197778 197879 197980 Total no. recoveries % of total recoveries Far North Above 530 N.W. Territories Alberta Sa s ka t chewan Total --- ---- 9.8 --- -- -- --- 7.1 10.2 9.1 1.0 5.1 0.8 7.0 28.6 18.3 0.1 35.3 10.5 28.9 11.5 19.0 21.8 0.8 15.6 18.3 0.6 11.9 21.8 0.5 0.8 0.1 0.7 0.1 5.4 0.4 0.3 0.5 0.3 5.8 0.7 0.3 0.5 0.3 6.3 1.4 0.1 1.3 0.5 4.9 1.9 1.0 1.4 5.2 --- -- 44.6 1.9 46.5 1.0 0.5 0.5 24.6 5.6 30.3 -- 0.9 7.9 8.8 0.9 8.3 0.9 10.1 1.2 4.3 0.6 6.1 1.1 12.2 1.1 14.4 2.6 1.3 3.9 6.0 2.4 8.3 15.6 22.0 0.9 17.8 19.6 0.6 12.2 28.9 15.4 20.5 20.2 29.8 356 8.6 1,008 713 10 24.5 17.3 0.2 28 6 26 16 239 0.7 0.2 0.6 0.4 5.8 1.289 20 34 39 31.3 0.5 0.8 1.0 4.4 1.4 0.6 6.6 Provinces Below 530 Alberta Saskatchewan B.C., Manitoba, Ontario Central_Elv\~ay Montana North Dakota Wyoming South Dakota Nebraska Colorado Southeast and Other :Inrth Central Total Kansas Oklahoma New Nexico Texas Panhandle Waggoner Ranch Gulf Coast Total -- 0.6 6.5 -- -- 0.7 10.6 -- -- 25.5 0.4 0.9 1.5 30.9 0.3 0.3 0.7 28.4 0.5 0.4 0.4 31.4 0.2 0.4 0.4 34.3 5.0 39.3 0.9 1.4 0.6 2.3 2.0 0.8 5.1 2.8 2.2 0.2 5.2 4.6 1.5 0.5 6.6 5.5 1.2 0.8 7.5 5.6 1.0 0.5 7.1 2.4 1.0 3.4 8.5 0.7 2.8 . 12.0 Pacific~ 2.4 2.2 0.8 0.7 0.9 1.4 1.4 HississiEpi Fl~way 0.1 0.3 0.1 Mexico 0.1 Total number of recoveries 748 aLess than 0.1 percent. 1.3 0.6 1.4 0.7 2.7 0.4 2.2 0.9 0.9 5.3 37.8 5.8 43.6 1.8 0.9 27.4 7.1 34.5 5 •. 1 1.3 7.9 0.9 6.4 0.9 5.5 1.2 . 2.2 6.7 8.8 7.3 6.7 2.2 0.4 1.8 0.9 1.8 26.6 6.1, 33.0 0.9 3.7 2.8 8.6 27.6 7.4 34.4 1.8 0.6 1.8 2.2 7.8 17.8 10.0 27.8 2.2 2.2 0.9 0.9 3.6 6.0 26.9 10.3 37.2 2.6 1.3 14.3 4.8 19.1 10.3 2.6 2.6 15.4 8.3 1.2 9.5 183 58 24 270 2.3 1.2 62 1.5 5 0.1 Ta 2.4 0.6 J 677 770 712 800 211 142 225 113 109 163 90 78 84 4,122 w •....• �Table 2. Comparison of juvenile and adult recovery and survival rate estimates for Canada geese banded post-season in eastern Colorado. Banded 1967-79 Difference Parameter or Test Subadult Adult Mean recovery rate 6.16 ± 0.60 4.97 ± 0.26 Mean survival rate 82.50 ± 7.94 75.88 ± 2.87 Test of HO vs HI X2 = 27.54, df = 25, P = 0.33 Goodness of fit test of HO X2 =118.40, df =117, P = 0.45 Z Value P -1.19 -1.69 0.09 -6.62 -0.13 0.90 W N Banded 1958-65 Mean recovery rate 8.51 ± 0.69 8.43 ± 0.40 - .08 -0.10 0.92 Mean survival rate 73.50 ± 4.63 74.08 ± 2.07 .38 0.12 0.90 Test of HO vs HI X2 = 20.72, df = 16, P = 0.19 Goodness of fit test of HO X2 =154.44, df =130, P = 0.07 �Table 3. Year Number of Canada geese banded in southeast Colorado and resulting recoveries 1967-79. Number banded 1967 1968 1969 1970 Number and year recovered 1975 1971 1972 1973 1974 _---_---- ---- 1976 1977 1978 1979 1967 674 47 36 15 21 17 14 11 6 5 1 3 0 0 1968 922 0 51 26 37 29 16 13 4 8 6 3 4 1 1969 1,089 0 0 31 50 53 18 33 15 10 8 8 4 8 1970; 599 0 0 0 39 30 10 19 4 2 9 5 4 1 1971 836 0 0 0 0 42 26 34 20 13 14 2 5 5 1972 1,032 0 0 0 0 0 42 51 24 18 25 14 8 5 1973 564 0 0 0 0 0 0 35 16 19 11 8 9 2 1974 337 0 0 0 0 0 0 0 14 11 13 5 4 1 1975 428 0 0 0 0 0 0 0 0 22 10 7 10 4 1976 920 0 0 0 0 0 0 0 0 0 38 19 15 12 1977 353 0 0 0 0 0 0 0 0 0 0 12 6 9 1978 236 0 0 0 0 0 0 0 0 0 0 0 10 7 1979 353 0 0 0 0 0 0 0 0 0 0 0 0 31 w w �34 Table 4. Recovery and survival rates of Canada geese banded in southeast Colorado 1958-65 and 1967-79. Year Recovery rate Estimate (±SE) 95% C. I. Survival rate Estimate (±SE) 95% C. I. 1958 10.33 ± 1.33 7.72 - 12.93 87.54 ± 9.67 68.59 - 106.48 1959 9.13 ± 1.04 7.10 - 11.15 47.91 ± 5.34 37.44 - 1960 10.27 ± 1.14 8.03 - 12.50 83.34 ± 8.53 66.62 - 100.07 1961 8.10 ± 0.86 6.41 - 9.79 71.47 ± 7.49 56.79 - 86.16 1962 6.05 ± 0.74 4.61 - 7.49 80.89 ± 7.88 65.45 - 96.33 1963 9.13 ± 0.78 7.61 - 10.65 64.31 ± 5.92 52.71 - 75.91 1964 8.45 ± 0.79 6.90 - 10.01 85.63 ± 7.76 70.43 - 100.84 1965 5.53 ± 0.52 4.51 - 58.39 6.54 Arithmetic Mean 8.78 ± 0.37 74.44 ± 1.61 95% C.I. 8.06 - 9.50 71.29 - 77.60 1967 6.97 ± 0.98 5.05 - 8.90 88.77 ± 8.98 71.17 - 106.37 1968 5.71 ± 0.64 4.47 - 6.96 71.88 ± 6.57 59.00 - 1969 3.29 ± 0.40 2.50 - 4.08 89.82 ± 9.03 72.11 - 107.53 1970 5.71 ± 0.61 4.51 - 6.90 76.61 ± 8.46 60.02 - 93.20 1971 6.07 ± 0.58 4.94 - 7.19 72.49 ± 7.34 58.12 - 86.87 1972 4.08 ± 0.42 3.26 - 4.91 78.52 ± 9.00 60.89 - 96.15 1973 6.52 ± 0.70 5.15 - 7.89 77.33 ± 12.75 52.34 - 102.32 1974 3.81 ± 0.62 2.63 - 4.99 82.88 ± 15.58 52.34 - 113.42 1975 3.99 ± 0.60 2.81 - 5.17 90.92 ± 15.42 60.69 - 121.14 1976 3.96 ± 0.49 3.01 - 4.92 65.33 ± 14.24 37.42 - 93.23 1977 3.24 ± 0.66 1.96 - 4.53 58.04 ± 17.64 23.46 - 92.63 1978 4.25 ± 1.04 2.21 - 6.28 32.71 ± 10.07 12.98 - 52.44 1979 8.78 ± 1.51 5.83 - 11.73 Arithmetic Mean 4.80 ± 0.19 73.78 ± 1.21 95% C.I. 4.42 - 5.18 71.41 - 76.14 84.76 �35 LITERATURE CITED Brownie, C., D. R. Anderson, K. P. Burnham, and D. S. Robson. 1978. Statistical inference from band recovery data - A handbook. U.s. Fish and Wildl. Serv., Resour. Publ. 131. 212pp. Szymczak, M. R. 1979. Monitor banding of the shortgrass prairie Canada goose population in southeastern Colorado. Colo. Div. o~ Wildl., Game Res. Rept., Fed. Aid Proj. W-88-R. Oct. pp . 27-39. ��October 37 1981 INTERIM JOB FINAL REPORT State of Project Colorado ----------------------- 3 Work Plan No. Job Title: Covered: Personnel: Job No. Population Central Period Migratory W-88-R-26 No. Characteristics Bird Investigations 7 of Mallards Wintering in West Colorado April 1, 1980 to March 31, 1981 G. Bock, J. Corey, D. Coven, J. Ellenberger, J. Frothingham, J. Gray, J. Gumber, D. Schaefer, S. Steinert, and R. Hopper. ABSTRACT The eighth consecutive year of the study of wintering mallards in west central Colorado was completed in Segment 26. The 1981 January inventory produced an estimated 14,695 mallards in the study area, which was considerably lower than the 8-year average. Sex ratio counts of 3,000 birds yielded 99.1 males:l00 females, the lowest ratio since the first year of counts in 1976. Fifteen hundred mallards were banded in 1981, bringing the total to 12,840 for the eight years. An analysis of the band recoveries from the first six years of banding (1974-197"9) was conducted during the segment. This analysis was based on 724 recoveries from 10,141 bandings. Over 60 percent of the total recoveries occurred in western Colorado. Adult females showed the strongest affinity for this area. This wintering population is strongly Pacific Flyway oriented, with nearly 80 percent of the total recoveries in this flyway. In addition to western Colorado, Utah and Idaho were important recovery locations in the Pacific Flyway. Only about 11 percent of the total recoveries occurred in the Central Flyway, while less them one percent occurred in the Mississippi Flyway. Canada (mostly Alberta) accounted for only 9.2 percent of the total recoveries, less than was the case for east slope wintering mallards. Mean recovery rates for males and females were estimated at 4.40 and 2.58 perce~t, respectively. Mean survival rate estimates were about 68 percent for males and 60 percent for females. These estimates were somewhat lower than those obtained for the east slope population. West slope estimates showed much wider confidence intervals, reflecting the need for additional years of recovery data. A final report will be in the form of a publication under Work Plan 6, Job 1 in Segment 27. �38 RECOMMENDATIONS The above discussion points to the need for additional years of recovery data for the west slope population. One more year of recoveries should help in reducing the size of standard errors and in narrowing confidence intervals. Banding was continued in 1981 and probably will be continued beyond 1981 by Region personnel as a monitor program. This means that by October of 1981 an additional year of recoveries, or a total of seven, will be available for analysis. At that time, an updated analysis covering all seven years should be conducted by this project in an effort to add precision to parameter estimates. It is also recommended that: 1. The Migratory Bird Project (W-88-R) terminate its responsibility for trapping and banding wintering mallards in west central Colorado following the January-February 1981 banding period. 2. The Northwest and Southwest regions take over this trapping and banding responsibility by initiating a monitor banding project beginning in January 1982 as part of their overall waterfowl management program. 3. Each region select a crew leader and make him available to assist research personnel in 1981 so that final instructions can be given for an orderly transfer of responsibilities. A detailed proposal covering recommendations for this monitor banding project was submitted to the two regions in the spring of 1980. �39 POPULATION CHARACTERISTICS OF MALLARDS WINTERING IN WEST CENTRAL COLORADO Richard M. Hopper This report presents results of the eighth consecutive year of study of the mallard population wintering in west central Colorado. Included is a summary of all data collected since initiation of this investigation during the 1973-74 segment year. In addition, an analysis of the first six years (1974-1979) of band recovery data is included. P. N. OBJECTIVES 1. To estimate population parameters of mallards wintering in west central Colorado by age and sex category; specifically (1) population size, (2) recovery rates, (3) survival rates, (4) mean life span, (5) sex ratio, and (6) geographic distribution of the harvest. 2. To develop a management plan for the population wintering in west central Colorado. of mallards SEGMENT OBJECTIVES 1. Conduct an aerial count of wintering numbers of mallards during the first or second week of January as part of the annual midwinter survey conducted through the United States. 2. Conduct a minimum of two ground counts (sex ratio) of 500 mallards each in each of the following two concentration areas: (1) Grand Junction-Highline Lake area, and (2) Montrose-Delta area. 3. Trap and band 1,500 mallards during the postseason period, including 750 in the Grand Junction-Highline Lake area, and 750 in the Montrose-Delta area. Distribute the. sample equally among the four age and sex classes. 4. Conduct an analysis of band recovery data for the first six recovery years, including the estimation of the following major population parameters by age and sex class: (1) distribution of the harvest, (2) recovery rates, and (3) survival rates. 5. Prepare and submit banding schedules and band recovery and return reports. Also prepare a progress or final report; the latter being dependent upon the existence of sufficient recovery information to yield accurate estimates of survival rates, etc. 6. Publish findings in appropriate technical is sufficient to justify publication. journals if information �40 METHODS AND MATERIALS Field methods remained the same as presented in Segment 22 (Hopper 1977). The winter aerial survey was conducted on January 6, 1981. Personnel of the Northwest Region again made the count, utilizing a Cessna 185 aircraft and a crew consisting of a pilot and two observers. Sex ratio counts were made during the postseason banding period on January 20-22, 1981 at Highline Lake and on January 22-24, 1981 at Sweitzer Lake. Salt Plains type traps, baited with corn, were again used to capture mallards for banding after the close of the hunting season. Banding dates included the period January 21-30, 1981. All birds were aged as usual by the wing-aging technique (Carney 1964), and recapture data were kept for each banding location. Banding schedules and recovery reports were prepared and submitted to the Bird Banding Laboratory. The banding analysis utilized only band recoveries that resulted from the first six years of banding, 1974-1979, or recovery years 1974-75 through 1979-80. Also, bands recovered anywhere were included (but only from those birds shot and/or found dead during the hunting season of September 1 to January 31). Recovery distributions and recovery year information were tabulated by hand from recovery cards received periodically from the Bird Banding Laboratory. Recovery and survival rate estimates were obtained from computer programs using two new analysis methods developed by: (1) Brownie and Robson (1976), and (2) Seber (1970) and Robson and Youngs (1971).' These methods provided for the calculation of confidence limits to apply to these rates. RESULTS AND DISCUSSION Aerial Census Results of the 1981 January inventory of wintering mallards in west central Colorado, along with counts from the previous seven years, are presented in Table 1. The estimate of 10,403 in 1981 was down about 4,300 birds from 1980 (14,695), and well below the 8-year average (1974-1981) of 14,000. The area covered in this count was presented in more detail in an earlier report for this job (Hopper 1977). Sex Ratio Counts Sex ratio counts taken in 1981 are shown in Table 2 by location and date. Overall ratios found during the previous five years (1976-1980) are also included for comparison. Three thousand mallards were classified in 1981. The overall ratio in 1981 was 99.1 males:100 females, with northern and southern areas producing different ratios, 104.9 and 93.5 males per 100 females, respectively. In previous years, counts have produced overall sex ratios ranging from 99.2 (1976) to 114.5 (1978) males per 100 females. �Table 1. January inventory of mallards wintering in west central Colorado, 1974-198l. Location 1974 1975 1976 1977 1978 1979 1980 1981 Highline Lake 2,400 4,700 10,200 6,900 6,100 4,200 3,050 4,100 Colorado River 3,llO 1,830 1,545 3,390 3,046 2,856 3,599 2,687 Gunnison River 700 1,345 9,100 2,215 3,756 2,850 4,002 1,164 Uncompahgre River 0 485 270 225 208 1,785 179 172 Sweitzer Lake 0 0 1,900 1,800 450 0 3,400 2,000 .j;:- •.... Burlingame Road 2,975 3,400 llO 1,750 300 1,000 465 280 Total 9,185 ll,760 23,125 16,280 13,860 12,691 14,695 10,403 8-year Mean (1974-1981) = 14,000 �42 Table 2. Sex ratio counts of mallards in west central Colorado during January 1981, with overall ratios from 1976-1980 for comparison. Location Highline Lake Subtotal Sweitzer No. Males No. Females Total Males: 100 Females 01-20-81 01-21-81 01-22-81 235 267 266 265 233 234 500 500 500 88.7 114.6 113.7 768 732 1,500 104.9 252 231 242 248 269 258 500 500 500 101. 6 85.9 93.8 725 775 1,500 93.5 1,493 1,507 3,000 99.1 1,741 1,375 2,816 1,761 1,316 1,755 1,279 2,460 1,599 1,273 3,496 2,654 5,276 3,360 2,589 99.2 107.5 114.5 110.1 103.4 (North) 01-22-81 01-23-81 01-24-81 Lake Subtotal Date (South) TOTAL 1976 1977 1978 1979 1980 Trapping and Banding Trapping efforts in 1981 represented the eighth consecutive year of the postseason banding program in west central Colorado, and resulted in the banding of 1,500 mallards (Table 3). Success was higher in the Grand Junction area (900) than in the Delta area (600) because of a shortage of birds on major trapping sites in the latter area. This year's bandings brought the total banded sample to 12,840 for the eight years (1974-1981). Table 3. Number and age and sex composition of mallards banded postseason in west central Colorado, 1981, with totals for the period 1974-1981. AM Year and Location 1981 Grand Junction Delta Area Total 1974-1981 Grand Junction Delta Area Total ~~e Number banded and Sex AF SF Total Area 241 154 395 246 159 405 224 145 369 189 142 331 900 600 1,500 Area 1,994 1,474 3,468 1,881 1,770 3,651 1,284 1,015 2,299 1,885 1,537 3,422 7,044 5,796 12,840 �43 Analysis of Band Recovery Data Table 4 shows the first six years of banding and recovery data utilized as the basis for this band recovery analyses. A total of 10,141 bandings yielded 724 recoveries during the 1974-75 through 1979-80 hunting seasons. There was considerable variation in the number of recoveries by age and sex class, even though the number banded was similar except for adult females. Table 4. Number of mallards banded in west central Colorado, 1974-1979, and number recovered from these bandings during the 1974-75 through 1979-80 hunting seasons. Age and Sex Adult male Subadult male Adult female Subadult female Total Geographic Distribution No. Banded No. Recovered 2,769 2,946 1,727 2,699 10,141 221 286 83 134 724 of Band Recoveries The locations of the 724 band recoveries are presented in Table 5 according to the percent of ~otal recoveries by age and sex class. Western Colorado, or the general vicinity of banding represented the major recovery location for all four age and sex classes, accounting for from 58.2 to 71.1 percent of the total recoveries. Adult females appeared more likely to be recovered in this area than males or subadult females. The population of mallards wintering in west central Colorado is definitely Pacific Flyway-oriented, as indicated by the occurrence of nearly 80 percent of the total recoveries in this Flyway. Utah and Idaho were the only other Pacific Flyway states of importance as recovery locations. The relationship of these birds to the Central Flyway was small, with only about 11 percent of the total recoveries occurring here (Table 5). Adult females, especially, seemed to avoid the Central Flyway. The Mississippi Flyway appeared to be insignificant as a recovery area for this population, as only three (0.4%) of the 724 recover~es came from this flyway. There was not a strong relationship between Western Slope wintering mallards and Canadian production areas, with the possible exception of subadult females. Only 9.2 percent of the total recoveries occurred in Canada, mostly in the province of Alberta, but about 13 percent of all recoveries of subadult females were taken here. �44 Table 5. Recovery distribution of mallards banded postseason in west central Colorado, 1974-1979, and recovered through the 1979-80 hunting season (all years). Recovery location AM Percent of total recoveries Age and sex AF SF SM Total Canada Alberta British Columbia Manitoba Saskatchewan Subtotal 6.3 0.4 0 3.2 10.0 4.9 0.3 0.3 1.7 7.3 4.8 0 0 3.6 8.4 9.0 0.7 0 3.0 12.7 6.1 0.4 0.1 2.6 9.2 Pacific Flyway Arizona Colorado Idaho Montana New Mexico Oregon Utah Washington Wyoming Subtotal 0 58.8 6.8 1.4 0.9 0 6.3 0 2.3 76.5 0.7 60.8 5.6 2.4 0.7 0.3 6.6 0 1.7 79.1 0 71.1 3.6 1.2 1.2 2.4 6.0 1.2 1.2 88.0 0 58.2 4.5 0 0.7 0 11.9 0 1.5 76.8 0.3 60.9 5.5 1.5 0.8 0.4 7.4 0.1 1.8 78.9 Central Flyway Colorado Kansas Montana Nebraska New Mexico North Dakota South Dakota Texas Wyoming Subtotal 6.8 0 2.7 0.4 0 0.9 0 0.4 0.9 12.2 6.6 0.7 0.3 2.1 0.7 0 0.3 0.7 1.7 13.3 0 0 1.2 0 1.2 0 0 0 0 2.4 3.7 0.7 1.5 0 0.7 0 0.7 0 1.5 9.0 5.4 0.4 1.4 1.0 0.6 0.3 0.3 0.4 1.2 10.9 0 0.4 0.3 0 0 1.2 Cf:3 1:2 0 0 0 0.1 0.3 0.4 0 0 1.5 0.6 100.0 100.0 100.0 100.0 100.0 221 286 83 l34 724 Mississippi Flyway Iowa Louisiana Subtotal 0.9 Unknown Total Number ----0.4 of recoveries �45 Some interesting comparisons are shown in Table 6 between recovery distributions of west slope and east slope (Central Flyway) wintering populations of mallards in Colorado. The east slope data represent unpublished information covering the period 1964-1976. While western slope wintering mallards are not entirely independent of Canadian production areas (9.2% of total recoveries), this relationships does not appear to be as strong as that between eastern slope wintering mallards and Canadian production areas (17.3% of total recoveries). Females banded as subadults in both portions of Colorado seemed to show the closest association to Canada of the four age and sex classes. Alberta was the major Canadian province of recovery for both groups, and the recovery distribution within Alberta appeared to be similar for the two groups. Western slope mallards were more likely than eastern slope mallards to be recovered in their respective general areas of banding, with the exception of the adult males. Only about 50 percent of the recoveries from eastern slope subadult males occurred in eastern Colorado (Central Flyway portion), while about 61 percent of those from western slope subadult males occurred in western Colorado (Pacific Flyway portion). Figures for adult females were about 54 and 71 percent, respectively, for the east and west slopes, and 36 and 58 percent for subadult females. Similar percentages of total adult male recoveries from east and west slope bandings, 59 and 64, respectively, were taken in the portion of the state where banded. The same pattern as above existed between the two areas in regard to flyway of recovery. With the exception of adult males, mallards banded in 'western Colorado were taken to a greater extent in the Pacific Flyway than were eastern Colorado banded mallards taken in the Central Flyway. These differences no doubt reflect to some degree the larger proportion of eastern slope recoveries occurring in Canada, thereby leaving a smaller proportion available to be recovered in the state and flyway of banding. Recovery and Survival Rate Estimates The initial step in the analysis involved the question of whether or not subadult and adult mallards have similar recovery and survival rates. The answer to this question determines if the two age groups should be pooled and the data analyzed using models that assume parameters are age-independent. First, this involved a test between two hypotheses or models (HO vs. HI' Brownie and Robson 1976). Both models allow recovery and surv1val rates to vary from year to year, but HO assumes these parameters to be independent of age (i.e., subadults and adults), whereas HI permits them to differ between the two. The results of contingency chi-square tests of Ho vs. Hl are shown in Table 7 for both males and females. These results prov1de no evidence that subadults and adults of either sex were significantly different in regard to recovery or survival rates. The "Goodness of Fit Test of HO" also supported this conclusion. Hopper et al. (1978) came to this same conclusion with regard to studies of the eastern Colorado wintering mallard population. �Table 6. Comparison of recovery distributions between wintering populations of mallards on the west slope (1974-1979) and east slope (1964-1976) of Colorado. General Recovery Location AM SM WS Total .ES WS ES Canada 10.0 15.0 7.3 16.7 8.4 19.8 12.7 24.3 9.2 17.3 Pacific Flyway 76.5 2.5 79.1 5.8 88.0 3.1 76.8 6.6 78.9 4.2 Colorado 58.8 0.2 60.8 0.6 71.1 0.5 58.2 0.4 60.9 0.4 Central Flyway 12.2 81.3 13.3 74.5 2.4 75.1 9.0 63.5 10.9 76.1 Colorado 6.8 63.6 6.6 50.5 0 54.5 3.7 35.6 5.4 54.5 Mississippi Flyway 0.4 1.2 0.3 2.9 1.2 2.0 0 5.3 0.4 2.4 Atlantic Flyway 0 0 0 0 0 0 0 0.2 0 T Alaska 0 0 0 0 0 0 0 0.1 0 T Mexico 0 0 0 0.1 0 0 0 0 0 T Unknown 0.9 0 0 0 0 0 1.5 0 0.6 100.0 100.0 100.0 100.0 100.0 100.0 100.0 100.0 100.0 100.0 221 2,707 286 2,294 83 651 134 828 724 6,480 Total Number of recoveries WS Percent of Total Recoveries AF SF WS ES ES WS ES 0 ~ 0'\ �Table 7. Comparison of subadult, and adult recovery and survival rate estimates (%) for mallards banded postseason in west central Colorado, 1974-1979. Parameter or test Males Difference ..z. value Subadult Adult Mean recovery rate (± SE) 4.36 ± 0.39 4.01 ± 0.33 - 0.35 - 0.68 0.25 Mean survival rate (± SE) 81.50 ± 8.54 57.08 ± 3.73 -24.42 - 2.50 0.01 Test of HO vs. HI a P x2 = 16.37, df = 11, P = 0.13 X2 = 40.34, df = 27, P = 0.05 Goodness of Fit Test of HO - --Parameter or test Females Difference Subadult Adult Mean recovery rate (± SE) 2.38 ± 0.31 2.77 ± 0.37 0.39 0.81 0.21 Mean survival rate (± SE) 67.51 ±13.38 54.73 ± 8.35 -12.78 - 0.81 0.21 Test of HO vs. HI a Goodness of Fit Test of HO X2 X2 Z value = 16.19, df = 11, P - 0.13 = 20.50, df = 23, P 0.61 aTest of the null hypothesis that there is no difference in survival and/or recovery rates between subadults and adults. P ~ .....• �48 There is one set of data in Table 7 that fails to support the above conclusion; that being that the mean survival rate estimates for the two age classes of males appear to be quite different (86.57 and 58.80%). A hypothesis similar to that above was tested for these survival rates by means of the ~ test. This test showed evidence of an age-specific survival rate for males. However, this could be a reflection of small banded samples over a short period of years. Hopefully, additional years of data will resolve this apparent conflict in the data. Despite this complication, it was decided to pool the age classes by sex and proceed with the analysis based on the assumption of ageindependent parameters. The pooled recovery data were subjected to a computer program that presented a detailed analysis of four models, each based on several assumptions (Brownie et ale 1978). Under each model, parameters were estimated using the theory of maximum likelihood, and the assumptions were tested statistically. The computer program processes the data through a sequence progressing from a general model, based on relatively few assumptions, to a model which makes very simple, but quite restrictive assumptions. Results of the above analysis showed that both the male and female data sets were best described by the assumptions under Model 3. Model 3 is the simplest possible age-independent model of band recoveries (Brownie et ale 1978). It is based on the assumption that recovery rates (and therefore harvest rates and 'band reporting rates) and survival rates are constant from year to year and independent of the age of the bird or its capture history. This model is based on only two parameters, the constant recovery rate and the constant survival rate (Table 8). A third parameter estimate, mean life span, is common to all models. Table 8. Population parameter central Colorado, 1974-1979. Population estimates of mallards Male Mean recovery rate (%) Estimate Standard error 95% confidence interval 4.40 0.24 3.93-4.88 Mean survival rate (%) Estimate Standard error 95% confidence interval 68.25 2.71 62.94-73.55 Mean life span (years) Estimate Standard error 95% confidence interval 2.62 0.27 2.16-3.25 banded in west Female 2.58 0.22 2.16-3.01 60.10 3.86 52.52-67.67 1.96 0.25 1.55-2. 56 �49 Mean recovery rates for west slope mallards were estimated at 4.40 and 2.58 percent, respectively, for males and females (Table 8). Mean survival rate estimates were about 68 percent for males and 60 percent for females. Lower recovery and survival rates for females than for males is typical for most duck populations. Females are simply subjected to higher mortality than males, and this situation is reflected in the lower mean recovery rate and shorter mean life span for females. The estimates of population parameters presented above for west slope mallards differed somewhat in most respects from those obtained for the east slope population (Tables 8-9). Survival rates were lower for west slope birds and the mean recovery rate for west slope females (2.58%) appeared to be higher than that for east slope females (1.95%). Standard errors and confidence intervals for the parameter estimates are important in that they provide an indication of the precision of the estimates. Parameter estimates for the west slope appeared to be much less precise than those for the east slope. Again, sample sizes probably reflected this difference, since the east slope study involved 13 years of banding and over 6,400 recoveries, compared to only six years of banding and 724 recoveries for the west slope study. Thus, one should be cautious in comparing estimates of population parameters between the two populations. Table 9. Population parameter eastern Colorado, 1964-1976. Population estimates parameter for mallards Male banded in Female Mean recovery rate (%) Estimate Standard error 95% confidence interval 4.32 0.14 4.05-4.58 1.95 0.07 1.81-2.08 Mean survival rate (%) Estimate Standard error 95% confidence interval 72.00 1.35 69.35-74.65 65.12 1.05 63.07-67.18 Mean life span (years) Estimate Standard error 95% confidence interval 3.04 0.17 2.73-3.42 2.33 0.09 2.17-2.51 LITERATURE CITED Brownie, C., and D. S. Robson. 1976. Models allowing for age dependent survival rates for band-return data. Biometrics 32(2):305-323. ________ , D. R. Anderson, K. P. Burnham, and D. S. Robson. 1978. Statistical inference from band recovery data -- a handbook. U.S. Fish and Wildl. Servo Resour. Publ. No. 131. 212pp. �50 Carney, S. M. 1964. Preliminary keys to waterfowl age and sex identification by means of wing plumage. u.s. Fish and Wildl. Servo Spec. Sci. Rept. - Wildl. No. 82. 47pp. Hopper, R. M. 1977. Population characteristics of mallards wintering in west central Colorado. Colo. Div. of Wildl. Fed. Aid Game Res. Rept., Oct. pp. 33-39. , H. D. Funk, and D. R. Anderson. 1978. Age specificity mallards banded postseason in eastern Colorado. J. Wildl. Manage. 42(2):263-270. ---- in Robson, D. S., and W. D. Youngs. 1971. Statistical analysis of reported tag-recaptures in the harvest from an exploited population. Biometrics Unit, Cornell Univ., Itahca, N.Y. BU-369-M. 15pp. Seber, G. A. F. 1970. Estimating time-specific survival and reporting rates for adult birds from band returns. Biometrika 57(2):313-318. �51 October JOB PROGRESS State of 1q81 REPORT Colorado ------~~~~-------W-88-R-26 Project No. 3 Work Plan No. Job Title: Monitor ---- Period Covered: Personnel: Migratory 8 Job No. Banding of Eastern Bird Investigations Colorado Mallard Populations )0 April 1980 - 31 March 1981 M. Babler, G. Brown, L. Budde, J. Corey, C. Crawford, D. Crawford, L. Crooks, K. Dillinger, M. Etl, D. Gardner, D. Homan, J. Jackson, T. Lynch, F. Marcoux, R. Oehlkers, J. Pogorelz, F. Rinella, H. Spear, S. Steinert, M. Szymczak, E. Wagner, Colorado Division of Wildlife. ABSTRACT A total of 4,089 mallards was banded postseason in seven locations of eastern Colorado in Segment 26. The updated analysis of post banding data was essentially finalized in Segment 25 and some was updated in Segment 26. Preparation of the final report was started, with completion scheduled for next segment in the form of a publication. ��53 MONITOR BANDING OF EASTERN COLORADO WINTERING MALLARD POPULATIONS Gerald Lorentzson This report summarizes Segment 26 results of the program involving monitor banding of mallard populations in eastern Colorado during the post season period. Hopper (1977) described the purpose of the original study, general accomplishments, reasons for transferring field responsibilities to the regional management system, and the role of the Research Section in the program. Hopper (1978) presented results of the first phase of this investigation, which related to the first P.N. Objective listed below. These results concluded that the transfer of monitor banding responsibilities from research to management was not complete, and that the first objective of this study was thereby met. The 1978 report also outlined the responsibilities of the Research Section with regard to this banding program. Thus, the Segment 26 report presented here addresses the second P.N. Objective. P. N. OBJECTIVES 1. To establish monitor banding of wintering mallard populations eastern Colorado as an annual management function. in 2. To continually document, through monitor banding and analysis of recovery data, the annual and long term status of eastern Colorado wintering mallard to provide a basis for annual hunting season recommendations. SEGMENT OBJECTIVES 1. Band a m~n~mum of 4,000 mallards during the post season period, including a minimum of 500 birds in each of the following general areas of the South Platte Valley and Arkansas Valley: (a) DenverGreeley, (b) Fort Collins-Loveland-Windsor, (c) Greeley-Fort Morgan, (d) Fort Morgan-Sterling, (e) Sterling-Julesburg, (f) Bonny Reservoir, (g) Manzanola-Lamar, and (h) Two Buttes Reservoir area. Divide the banded sample in each area equally among the four age and sex classes. 2. Conduct an updated analysis of band recovery data, including the following major determinations for important population segments of mallards wintering in eastern Colorado: (a) distribution of the harvest, (b) recovery rates, and (c) survival rates. 3. Prepare and submit banding schedules, reports, and progress report. band recovery and return �54 METHODS AND MATERIALS Procedures and equipment remained essentially the same as in the previous year (Hopper 1977), with the exception that participation in the actual banding effort by research personnel (Federal Aid Project W-88-R) was limited to Bonny Reservoir. The Research Section has retained responsibility for banding at Bonny Reservoir because of closely related studies anticipated for the future. RESULTS AND DISCUSSION Banding Nearly 4,100 mallards were banded postseason in January of 1981, or in Segment 26 (Table 1). This number was again in excess of the 4,000 quota originally set for eastern Colorado. An eighth location, the Arkansas Valley did not contribute a sample during this segment. The four age and sex classes were all well represented in the banded sample. All banding schedules and recovery reports were prepared and submitted to the Bird Banding Laboratory. Table 1. Mallards banded postseason by age and sex in seven eastern Colorado banding areas, January 1981. Banding Number of ducks banded Age and sex SM AF SF Unk Total Area AM Bonny Reservoir Sterling-Julesburg Fort Mogran-Sterling Greeley-Fort Morgan Denver-Greeley Fort Collins-LovelandWindsor 'l'woButtes Area 252 126 129 126 132 272 125 129 129 137 325 125 157 111 113 233 124 84 134 117 128 270 131 27 137 154 104 56 2 500 509 950 1,122 852 2 4,089 Total 1,163 Total 1,082 500 499 500 499 �55 Updated Analysis of Band Recovery Data The previous segment report indicated the progress made through Segment 24 in regard to the updated analysis of banding data from eastern Colorado wintering mallard populatlons (Hopper 1979). This same report discussed the steps followed in the analysis, as well as the quantity of data used. Most of the analysis was finalized during Segment 25, and considerable progress was made in writing portions of the final manuscript. Some additional progress was made in Segment 26. However, preparation of a final report by way of a publication will be completed during the next segment under Work Plan 6, Job 1 and submitted for publication in an appropriate technical series. LITERATURE CITED Hopper, R. M. 1977. Monitor banding of eastern Colorado wintering Mallard populations. Colo. Div. of Wildl. Fed. Aid Game Res. Rept., Oct. pp. 41-47. 1978. Monitor banding of eastern Colorado wintering mallard populations. Colo. Div. of Wildl. Fed. Aid Game Res. Rept., Oct. pp. 41-45. 1979. Monitor banding of eastern Colorado wintering mallard populations. Colo. Div. of Wildl. Fed. Aid Game Res. Rept., Oct. pp. 49-56. , Prepared " :J~~~)A~~~~~_V'~:~~~~~~==~ _ by ,~__ Gerald Lorentzson Senior Wildlife Biologist ��October 57 1981 INTERIM JOB FINAL REPORT State of Project Colorado ----~~~~~-------No. W-88_-_R__26 3 Work Plan No. Migration Job Title: Migratory _ Job No. and Mortality in the Inter-mountain Period Covered: Personnel: Bird Investigations 9 Characteristics of Duck Populations Valleys of Colorado 1 April 1980 to 31 11arch 1981 M. Nail and staff, Monte Vista and Alamosa National Wildlife Refuges; J. Corey, Dale Flenthrope, Dean Flenthrope, K. Karrow, M. Jancowsky, S. Steinert and M. Szymczak, Colorado Division of Hildlife. ABSTR..A.CT Totals of 3,205, 2,688, and 3,736 ducks were banded in North Park, South Park and the San Luis Valley, respectively, in 1980. Since the coordinated banding program was initiated in 1971, over 103,000 ducks have been banded. In 1980, the gadwall (18%) was the most numerous species in the harvest during the early season in North Park followed by the mallard (16%). In South Park, the mallard made up 47 percent of the harvest followed by the green-winged teal (25%) and the bluewinged and/or cinnamon teal (15%). The mallard comprised 43 percent of the harvest in the San Luis Valley followed by the gadwall (23%) and. the green winged teal (19%). In all years of collection the composition of the harvest during the early season for the major species was as follows: North Park - gadwall (21%), wigeon (16%), blue-winged and/or cinnamon teal (13%), mallard (10%); South Park - mallard (36%), greenwinged teal (17%), blue-winged and/or·cinnamon teal (14%)-; wigeon (12%), pintail (10%); San Luis Valley - mallard (47%), gadwall (16%), green-winged teal (12%). Population information for the major species in the three banding areas is summarized and comments and conclusions concerning early seasons are presented. RECOMMENDATIONS 1. Discontinue preseason duck banding in North Park with the exception of an increased effort to band adult gadwall on molting areas. Gadwall banding will be conducted under a different job. 2. Discontinue preseason duck banding in South Park. �58 3. Continue preseason duck banding in the San Luis Valley under another job, mainly as a means to monitor waterfowl population in respect to the continued decline of quantity and quality of Valley waterfowl habitat. 4. Continue waterfowl hunting seasons in North Park, South Park and the San Luis Valley which begin the Saturday nearest to October 1. 5. If experimental seasons should be proposed in North Park, South Park or the San Luis Valley which begin earlier than the Saturday nearest October 1, the size, species, age and sex composition of the harvest should be monitored. Consider the possibility of banding of key species to monitor survival. 6. If seasons should be proposed in North Park or South Park in September or October without concurrent seasons in the area adjacent to the foothills, some controls on the number of hunters participating in those seasons should be considered. �59 MIGRATION AND MORTALITY CHARACTERISTICS OF DUCK POPULATIONS IN THE INTER-MOUNTAIN VALLEYS OF COLORADO Michael R. Szymczak This job was initiated in 1976 to cover pre-season duck banding in the inter-mountain valleys of Colorado, analysis of recovery data resulting from those bandings and evaluation of those data in reference to establishing special high mountain duck seasons. The actual banding began at earlier dates; 1963 in the San Luis Valley, 1968 in South Park and 1971 in North Park. Some pre-season duck banding began even earlier in the San Luis Valley and in North Park, but the coordinated effort in all three areas began in 1971. Analysis of recovery data began in 1977 and has continued sporadically since that time. Collection of data, including banding and recovery data and species composition of the harvest, continued through 1980. The formal publication has not been completed. However, an evaluation of the status of the banding program, in reference to the objectives, was undertaken. The result was substantial changes in the program and therefore the job will be terminated and the aspects that will be continued will be written as separate jobs. P. N. OBJECTIVES 1. To investigate migration, mortality, recovery distribution and relationships among populations of selected species of ducks present in North Park, South Park and the San Luis Valley during the midJuly through mid-September period. 2. To document the species composition of ducks harvested during early October seasons, should seasons occur in North Park, South Park and the San Luis Valley. 3. To examine the feasibility of establishing early special duck seasons in September in the high mountain part areas or to examine the feasibility of special early October seasons in South and North Park similar to what has been recommended for the San Luis Valley. 4. To establish procedures to monitor the effect of early special duck seasons on duck populations in the high mountain park areas if granted. SEGMENT OBJECTIVES 1. Trap and band ducks in North Park, South Park, and the San Luis Valley during the mid-July through mid-September period as designated in Program Narrative Outline. Complete, submit and file appropriate banding schedules and recovery cards. �60 2. Collect and analyze data concerning the species composition of the harvest during early October seasons in North Park, South Park and the San Luis Valley utilizing wing collection barrels and results of the U. S. Fish and Wildlife Service's Parts Collection Survey. 3. Analyze band recovery data through the 1978 recovery year for mallards, pintail and green-winged teal banded during the preseason period .in North Park, South Park and the San Luis Valley to determine the feasibility of requesting early seasons in the high mountain area. 4. Prepare 5. If a special high mountain duck season is requested and granted, design a program for population monitoring which may include banding, harvest surveys, wing collection or other activities. progress report. METHODS AND MATERIALS Ducks were captured from August 9 - September 17 in South Park, August 6 - September 22 in the San Luis Valley and July 24 - September 18 in North Park. Primarily, "salt plains" type bait traps were used (Szymczak and Corey 1976). In North Park, night lighting from airboats was used to capture most gadwall and wigeon. The age and sex of all birds captured and banded were determined. Wings from ducks bagged during the October 4, 1980 through October 19, 1980 period were collected in North and South Parks through the use of voluntary collection barrels (Hoffman and Braun 1975). Barrels were placed at Walden Reservoir (3 barrels) and Lake John Annex (1 barrel) and at Cowdrey (1 barrel) in North Park, and at Antero Reservoir (3 barrels) in South Park. Some additional wings were collected at Hebron Ponds in North Park. All wings collected were classified by species, age, sex and location, and in some cases, periods of harvest. The species composition of the harvest in the San Luis Valley was obtained at the Central Flyway, u.S. Fish and Wildlife Service's Parts Collection "Wing Bee." Computerized analysis of banding and recovery tapes of birds marked through 1975 in North Park, South Park, the San Luis Valley and the high country areas west of the San Luis Valley was continued using programs outlined by Szymczak (1978) as well as programs ESTIMATE and BROWNIE for estimating survival and recovery rates as described by Brownie et al. (1978). In addition, most results for mallard, pintail and green-winged teal were updated through the 1978-79 recovery year utilizing periodic printouts of recovery information received from the Bird Banding Laboratory. �61 RESULTS AND DISCUSSION Banding Totals of 3,205, 2,688 and 3,736 ducks were banded in North Park, South Park and the San Luis Valley, respectively, in 1980. The species composition of the birds banded in each area are presented in Tables 1, 2 and 3. Quotas of 300 mallards of each sex of adult and immature birds were met for adult and immature males in all three areas, for immature females in the San Luis Valley, but not for adult females in any area. The North Park gadwall quota of 300 adults of each sex was met. A total of 9,629 birds was banded (Table 4). Since coordinated banding programs were initiated in the three areas, more than 103,000 ducks have been banded, with mallard and pintail being the most numerous. A summary of the number of birds banded of the target species is presented by year and area in Table 5. Species Composition of the Harvest A total of 429 wings was collected during the 1980 early duck hunting season in North Park (Table 6). Seventy-one percent of the wings were collected at Walden Reservoir. Eighty-one percent of the. wings were obtained during the opening weekend (Table 7). More gadwall were harvested than any other species, but the mallard was nearly as numerous. The percent of mallards in the harvest has been increasing since 1977 after being fairly stable during the first 3 years of collection (Table 8). .~ In South Park, the mallard made up 47 percent of the harvest followed by the green-winged teal with 25 percent and the blue-winged and/or cinnamon teal at 15 percent (Table 8). A record number 116 wings were collected during the 1980 season. In the San Luis Valley, the percent of mallards in the harvest declined from the atypically high 1979 level (Table 8). The mallard still remained as the major species of harvest followed by the gadwall and the green-winged teal. . , In all years of collection since 1975 combined, the mallard was the dominant species of harvest in South Park and the San Luis Valley (Table 8). In addition, the green-winged teal, blue-winged/cinnamon teal, pintail and wigeon each made up more than 10 percent of the harvest in South Park while the gadwall and green-winged teal were the species in that same category in the San Luis Valley. Harvest in North Park was more equally distributed among species with the gadwall the most numerous followed by the wigeon, blue-winged/cinnamon teal and the mallards. Diving ducks were more prominent in the harvest in North Park (20%) than in South Park (4%) or the San Luis Valley (1%). �62 Table 1. Number of ducks banded, by species in North Park during the pre-season period, 1980. Species Pintail Mallard Gadwall Green-winged teal Wigeon Blue-winged and/or cinnamon teal Redhead Totals Age and sex 1M IF LM LF Total 409 194 306 16 6 206 319 2 19 3 216 262 0 23 3 4 19 33 0 6 2 11 34 0 2 1,246 1,107 695 88 42 3 9 1 1 8 0 5 0 0 0 0 0 17 10 1,095 933 557 509 62 49 3,205 AM AF 409 302 320 30 22 Table 2. Number of ducks banded, by species, pre-season period, 1980. Species Pintail Mallard Green-winged teal Blue-winged and/or cinnamon teal Redhead Wigeon Gadwall Canvasback Totals in South Park during the Age and sex 1M IF AM AF 524 300 162 209 191 60 186 299 112 71 4 3 0 2 35 7 0 0 0 1,066 502 1M LF Total 110 209 56 0 5 0 0 8 0 1,029 1,012 390 70 2 1 0 0 58 0 2 2 0 0 0 0 0 0 0 0 0 0 0 234 13 6 2 2 670 437 5 8 2,688 �63 Table 3. Number of ducks banded, by species, during the pre-season period, 1980.~/ Species Mallard Pintail Green-winged teal Blue-winged and/or cinnamon teal Redhead Gadwall Totals in the San Luis Valley Age and sex IF 1M LM LF 747 213 51 18 2 22 3 a a a a 189 25 8 128 18 3 2 22 4 0 12 a a 1,030 1,160 48 37 AM AF 501 247 122 267 177 53 516 216 76 55 3 35 a a a 928 532 UU 1 a a/ - Includes 1,677 ducks banded by Monte Vista and Alamosa Wildlife Refuge personnel. Table 4. Number of ducks banded, by species, in North/Park, and the San Luis Valley during the pre-season period.~ Species Mallard Pintail Green-winged teal Gadwall Blue-winged and/or cinnamon teal Wigeon Redhead Canvasback Totals Age and sex 1M IF AM AF 1,103 1,180 314 320 652 795 129 306 1,134 608 207 10 129 25 16 2 71 6 8 3,089 Total 2,072 858 302 0 409 80 15 1 3,736 National South Park, LM LF UU 1,218 539 130 5 42 6 37 41 5 0 34 a a a 4,191 3,133 780 712 267 4 27 191 5 18 a a a 2 6 22 0 0 2 12 0 a a a a 660 48 103 2 1,967 2,257 2,106 115 94 1 9,629 a/ - Includes 1,677 ducks banded by.Monte Wildlif e Refuge personnel. a Vista and Alamosa 1 Total National �Table 5. Number of ducks of the major species banded in North Park, South Park and the San Luis Valley during the pre-season banding period, 1971 through 1980. -----NaLl.a r d Blue-winged and cinnamon teal Green-winged SLV NP NP Pintail teal NP SP SLV 1971 1,102 1,078 1,223 888 244 1,249 33 93 131 232 163 421 1972 1,114 867 1,408 1,101 521 1,333 76 83 303 245 138 673 123 1973 1,149 745 1,789 1,660 565 1,502 41 124 455 65 563 119 1974 1,304 993 1,684 1,779 580 1,725 81 333 599 367 753 1975 1,133 656 2,291 1,482 962 1,592 16 557 396 372 1976 1,146 651 1,647 855 1,082 1,231 25 323 625 1977 1,162 897 1,533 1,643 1,000 1,075 27 235 1978 1,085 1,049 2,295 1,046 1,090 965 15 1979 1,367 908 2,190 1,321 1,581 1,294 1980 1,107 1,012 2,072 1,246 1,029 11,669 8,856 18,132 13,021 8,654 Total NP SP SP SLY SP SLY A. Gadwall Year . NP SP Wigeon SLY NP SP 20 13 3 94 27 21 110 12 805 15 18 1,273 438 484 0 277 1,546 1,364 847 350 178 2,017 1,006 301 878 187 844 5 240 610 85 858 17 234 409 12,824 336 2,523 4,756 Redhead Total SLY NP SP 16 24 o 53 2,293 1,582 3,113 10 72 6 41 2,758 1,616 3,862 6 3 31 32 339 2,979 2,036 4,317 23 0 0 21 5 91 3,590 2,665 4,922 34 55 4 5 5 7 32 3,547 3,459 4,788 3 38 118 0 10 18 19 62 3,286 3,624 4,977 721 0 45 107 0 7 39 39 93 3,877 4,188 4,109 608 451 3 133 69 6 24 22 22 57 2,875 3,315 4,960 425 257 802 107 83 8 6 12 267 3,669 3,168 4,733 88 390 302 695 2 15 42 6 0 10 13 80 3,205 2,686 3,736 2,096 8,112 5,993 4,160 12 614 549 27 83 248 155 1,115 32,079 28,339 43,517 0 SLY NP SP SLY ~ ~ �Table 6. Species composition of the harvest in North Park during the October 4 - October early duck hunting seasons according to wings voluntarily placed in collection barrels. Walden Reservoir Species Area Lake John Annex 19, 1980 Hebron Ponds Cowdrey Total Gadwall 49(16.1) 6 (8.8) 19(40.4) 2(22.2) 76(17.7) Mallard 41(13.4) 14(20.6) 8(17.0) 7(77.8) 70(16.3) Blue winged or Cinnamon teal 35 (11. 5) l3(19.1) 4 (8.5) (0.0) 52(12.1) Lesser 48(15.7) 2 (2.9) 1 (2.1) (0.0) 51(11.9) 24 (7.9) 8(11.8) 8(17.0) (0.0) 40 (9.3) 29 (9.5) 8(11.8) 2 (4.3) (0.0) 39 (9.1) teal 30 (9.8) 4 (5.9) 3 (6.4) (0.0) 37 (8.6) shoveler 22 (7.2) 2 (2.9) 2 (4.3) (0.0) 26 (6.1) 10 (3.--3) 8(11.8) (0.0) 18 (4.2) 6 (2.0) 3 (4.4) (0.0) 9 (2.1) Bufflehead 5 (1. 6) (0.0) 5 (1. 2) Ruddy. 4 (1.3) (0.0) (0.0) 4 (0.9) C. Merganser 2 (0.7) o o o o o o o (0.0) 0_ (0.0) o o o o o o o o o o o (0.0) 2 (0.5) scaup American Wigeon Pintail Green-winged Northern Redhead Ring-necked duck 30.5 68 (0.0) 47 (0.0) (0.0) (0.0) (0.0) 9 429 0"> Vl �Table 7. Species composition of the duck harvest by time period at Walden Reservoir, Lake John Annex, Hebron Ponds and the Cowdrey area in North Park during the October 4 - October 19, 1980 duck season. Species Oct. 4-5 No. Percent Oct. 6-12 No. Percent Oct. 12-19 No. Percent Entire season No. Percent Gadwall 58 16.8 16 24.2 2 11.8 76 17.7 Mallard 57 16.5 11 16.7 2 11.8 70 16.3 Blue winged or Cinnamon teal 46 13.3 4 6.1 2 11.8 52 12.1 Lesser scaup 40 11.6 11 16.7 0 0.0 51 11.9 American wigeon 29 8.4 5 7.6 6 35.3 40 9.3 Pintail 32 9.2 5 7.6 2 11.8 39 9.1 Green-winged teal 34 9.8 3 4.5 0 0.0 37 8.6 Northern shoveler 21 6.1 4 6.1 1 5.9 26 6.1 Redhead 17 4.9 1 1.5 0 0.0 18 4.2 Ring-necked duck 7 2.0 2 3.0 0 0.0 9 2.1 Bufflehead 1 0.3 3 4.5 1 5.9 5 1.2 Ruddy 3 0.9 1 1.5 0 0.0 4 0.9 C. Merganser 1 0.3 0 0.0 1 5.9 2 0.5 Total 346 66 17 429 0\ 0\ �Table 8. Percent species composition of the harvest in North Park and South Park during ealy October duck hunting seasons 1976-1980according to independent wing barrel surveys and in the San Luis Valley based on U.S. Fish and Wildlife Service's Parts Collection Survey. Species NP Hallard 6.9 Gadwall 20.6 Green-winged Amer ican teal w i geon Blue \~inged/ Cinnamon teal 7.7 28.9 1975 Spi SLY 47.1 1978 SP SLY 1980 SP SLY All years SP SLY 1976 SP SLY NP 6.1 8.7 44.9 6.0 55.9 43.7 11.4 43.5 40.0 12.7 28.6 67.3 16.4 46.6 43.1 10.3 36.3 47.2 20.6 28.1 4.4 1977 SP SLY NP 2.2 6.7 25.4 7.2 25.0 19.1 2.9 21.4 28.3 4.9 4.5 14.9 NP 8.8 12.7 20.0 NP 6.5 24.4 17.0 1979 SP SLY NP 9.5 15.4 17.8 2.6 23.1 20.9 4.9 12.7 5.2 12.0 7.1 3.8 8.7 25.0 18.5 4.4 7.0 9.0 12.0 11.1 12.3 14.3 7.7 9.4 1.4 10.4 7.6 4.4 7.3 NP 5.5 16.2 6.9 17.3 11.5 5.1 0.0 0.0 15.8 11.5 8.9 18.5 19.0 1.9 11.5 14.7 3.1 12.6 14.2 6.1 0.0 5.4 17.9 0.0 9.2 6.9 7.7 5.8 10.2 8.2 8.5 3.6 1.9 6.1 0.9 4.6 8.9 4.4 15.4 5.9 8.8 25.0 13.5 8.6 2.9 Pintail 5.1 11.8 4.8 3.3 7.9 5.6 8.8 16.9 N0rthern shoveler 3.4 4.4 2.7 2.2 2.2 11.2 0.0 4.2 18.4 Redhead 4.3 1.5 11.2 5.4 0.0 10.1 2.9 0.0 6.9 0.0 0.0 9.6 0.0 1.9 4.2 0.0 0.0 7.4 1.5 0.5 Ruddy 1.6 0.0 1.6 0.0 0.0 2.6 1.5 0.0 1.3 0.0 0.0 0.4 0.0 0.0 0.9 0.0 0.0 1.3 0.2 0.0 Ring-necked duck 0.2 0.0 1.1 0.0 1.1 0.7 0.0 1.4 1.8 0.0 0.0 2.1 0.0 0.0 2.1 2.6 0.0 1.4 0.7 0.5 5.0 15.2 1.1 11.1 1.5 Lesser scaup 5.1 0.0 7.0 0.0 0.0 9.7 0.0 0.0 10.4 0.0 0.0 5.6 0.0 0.0 12.0 0.0 0.0 8.2 0.0 0.0 Common merganser 0.0 0.0 0.0 1.1 0.0 0.4 1.5 0.0 0.0 2.2 0.0 0.0 0.0 0.0 0.5 0.0 0.0 1.1 0.9 0.0 Hooded merganser 0.0 1.5 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 0.3 Bufflehead 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Q.6 0.0 0.0 1.2 0.0 0.0 0.3 0.0 0.0 Canvasback 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.9 0.0 0.0 0.0 0.0 Unknown 0.8 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 0.0 0.0 Total no. wings examined 494 68 374 1 \Jings not collected in South Park in 1975. 92 89 268 68 71 597 92 45 481 84 52 426 116 65 2,640 452 390 0\ -....I �68 Early Special Duck Seasons The feasibility of establishing early duck seasons in the high mountain park areas of Colorado was examined in light of population information collected and analyzed to date. A review of some of the population information with an emphasis on the mallard follows: Mallard - North Park 1. At least 65 percent of all direct recoveries of each age and sex class occurs in Colorado. Colorado takes nearly 60 percent of the indirect adult male recoveries, 47 percent of the indirect immature male recoveries, and about 70 percent of either age female recoveries. 2. A slight trend indicates increasing Mexico and reductions in Colorado. 3. The area adjacent to the foothills, west of Greeley and north of Denver, the San Luis Valley and southeast Colorado are the major recovery areas in the state for males. Females are taken primarily in the foothills area, Colorado River area and North Park. 4. A portion of the North Park mallard of North Park prior to October 1. 5. North Park mallards are in Colorado throughout the fall and winter periods. However, 70 percent of the direct and 60 percent of the indirect recoveries are reported taken in Colorado prior to November 21. 6. Species composition data has indicated mallards generally make up less than 15% of the harvest during.the early season in North Park. However, there is an increasing trend in mallard harvest. 7. Adult males banded in North Park in August are not recovered there during the early season. However, 25 percent of the mallard harvest during the early season is composed of adult males. Apparently the adult male population is changing during August-September and therefore those adult males banded in August are not available for harvest in October in the banding area. Mallard recoveries in Texas and New banded population migrates out - South Park 1. At least 70 percent of all direct recoveries of each age and sex group are taken in Colorado. Of the indirect recoveries, Colorado takes 56 percent of these for adult males, 46 percent of the immature males, 70 percent of the adult females and 58 percent of the immature males. 2. New Mexico and Texas are important males. recovery areas, particularly for �69 3. About 20 percent of the indirect recoveries of both sexes of immatures are recovered in states north of Colorado. 4. The San Luis Valley, South Park, southeast Colorado and the foothills area are the primary recovery areas in the state for both males and females. 5. The chronological distribution of recoveries indicate a portion of the South Park banded mallard population is in Colorado throughout the hunting season. 6. Some birds move out of South Park into the San Luis Valley prior to the hunting season. Apparently birds move into the foothills area at a later date than their counterparts from North Park. 7. Species composition data since 1977 indicate mallards to 50% of the harvest during the early season. make up 25% Mallard - San Luis Valley 1. The San Luis Valley is the major recovery in the San Luis Valley. area for mallards banded 2. New Mexico is now a more important recovery area for San Luis Valley birds than it was during the 1963-70 period. 3. San Luis Valley throughout the the direct and taken prior to 4. There was no evidence indicating a change in survival between the 1963-70 period and the 1971-78 period for both ages of males and adult females. Immature female survival increased. Recovery rates declined significantly between the two periods for all ages and sexes. banded mallards are taken in the San Luis Valley hunting season, but 84 percent and 79 perc.ent of indirect recoveries, respectively, are reported December 1. Pintail - North Park 1. Since 1975, pintail have comprised during the early season. less than 6 percent of the harvest 2. Most of the pintail harvest during the early season has been composed of adult females and young. 3. The major harvest areas are the Gulf Coast of Texas and Louisiana, California and Mexico. �70 Pintail - South Park 1. Since 1976, pintail have comprised during the early season. 2. As in North Park, most of the pintail harvest during the early season was composed of adult females and young. 3. The major harvest areas are the Gulf Coast of Texas and Louisiana, California and Mexico. Pintail about 10 percent of the harvest - San Luis Valley 1. Since 1975, pintail have comprised during the early season. 2. The San Luis Valley is a much more important recovery area for pintail banded in the San Luis Valley than are the other two park areas for birds banded in those respective areas. The Valley is a particularly important recovery area for females. 3. California, of note. Green-winged Texas, Louisiana about 8 percent of the harvest and Mexico are other recovery teal - North Park 1. Since 1975, green-winged teal have comprised the harvest during the early season. 2. Although recovery 3. Texas and California Green-winged areas about 7 percent of total recoveries are few, North Park is only a minor area for birds banded in that area. are important recovery areas. teal - South Park 1. Since 1976, about 16 percent of the harvest during the early season in South Park has been composed of green-winged teal. 2. This area of banding was of major of adult females. 3. California, Texas, Louisiana of importance. importance only for the recovery and Mexico were other recovery areas �71 Green-winged teal - San Luis Valley 1. Since 1975, 12 percent of the harvest during the early season in the San Luis Valley was composed of green-winged teal. 2. This area of banding is a major harvest area for adult females and immatures recovered directly but of minor importance on an indirect basis. 3. California, areas. Texas and Louisiana were other significant recovery Gadwall - North Park 1. Since 1975, the gadwall has been the most numerous species of harvest during the early seasons, making up about 21 percent of the harvest. 2. This area of banding is a substantial area of recovery, for immatures (which were banded as locals). particularly Gadwall - South Park 1. Gadwall are not an important hunting season period. species during the banding or early Gadwall - San Luis Valley 1. Since 1975, 18 percent of the duck harvest during the early season has been composed of gadwall, resulting in that species being the second most numerous species of harvest. 2. This area of banding is the major area of recovery for gadwall. New Mexico and Texas are other recovery areas of note. Blue-winged 1. and Cinnamon Teal - North Park Blue-winged and/or cinnamon teal made up 13 percent of the harvest during the early seasons 1975 through 1980. It was the most . numerous species in the bag in 1979. Blue-winged and Cinnamon Teal - South Park 1. The percent of harvest during the early season composed of bluewinged and/or cinnamon teal averaged 14 percent since 1976 but varied considerably from year to year. 2. South Park was not a major recovery banded in South Park. area for blue-winged/cinnamon �72 Blue-winged 1. and Cinnamon Teal - San Luis Valley Blue-winged/cinnamon teal made up about 6 percent of the harvest during the early season from 1976 through 1980. General Comments Concerning Early High Mountain Seasons All Areas 1. There is no evidence to indicate that recent early hunting seasons (~ October) have been detrimental to any waterfowl population in any of the three high mountain banding areas. 2. Changes in point values could be used during early seasons to orient the harvest toward or away from selected species and sexes, if needed. 3. If early seasons were established in high mountain areas without accompanying seasons in the area directly east of the foothills, some methods of controlling the number of hunters would be needed in North Park, probably needed in South Park but not in the San Luis Valley. The degree of control is based on available harvest habitat as well as distance from population centers. 4. A season beginning earlier than about October 1 in the 3 park areas, cannot be specifically evaluated without it actually occurring. Therefore, following comments in respect to such a season, although based on some supporting data, are speculative. North Park 1. A hunting effects: season earlier than October 1 could have the following a. Increase the harvest of mallards with the greatest effect on adult hens and young. Additional harvest of adult hens would most likely be detrimental to the local breeding population whereas harvest of males probably would not. b. Increase the harvest of pintail and blue-winged/cinnamon teal which would not be detrimental to those populations. c. Increase the harvest of late nesting adult hens and flightless broods, on production areas, primarily gadwall and lesser scaup, which could be detrimental to the population. d. Increase mortality, through harvest or harassment, molting adult gadwall hens. of late �73 2. Since the mallard population is harvested primarily in Colorado, any additional early season harvest in North Park would reduce harvest opportunity in other recovery areas within the State. 3. Using wing collections as an index, the level of harvest during the early season has varied somewhat since 1975, but there is no indication of an increasing or decreasing trend. South Park 1. A hunting effects: season earlier than October 1 could have the following a. Increase the harvest of mallards. The increase might be oriented toward adult hens and young. Although South Park cannot be considered a major production area, the effect could be detrimental to the local breeding population. Additional harvest of males would probably not be detrimental to the population. b. Increase the harvest of pintail and blue-winged/cinnamon which would not be detrimental to those populatipns. c. Increase the harvest of green-winged teal with a probable disproportionate increase· in adult females. teal 2. Since the mallard population is harvested primarily in Colorado, additional early season harvest in South Park would reduce harvest opportunities for mallards in other recovery areas in the state. 3. Using wing collections has been stable. as an index, harvest during the early season San Luis Valley 1. A hunting effects: season earlier than October 1 could have the following a. A disproportionate increase in the harvest of adult hens of the prominent breeding species in the Valley. Harvest during a season earlier than October 1 would take place mainly on production areas and therefore would result in harvest of adult hens associated with those production areas. This harvest could be detrimental to the local breeding populations. Additional harvest of males would not be detrimental to the population. b. Increase the harvest of blue-winged/cinnamon not be detrimental to the population. teal which would �74 2. In the case of survival rates a reduction in indicates that mallards, there has been no significant change in between the 1963-70 and 1971-78 period in spite of recovery rates during the latter period. This additional harvest may not affect survival. 3. Since most of the harvest of the San Luis Valley breeding mallard population occurs in the Valley any alterations in the population resulting from changes in survival, recovery rates or harvest chronology will have an effect mainly on harvest opportunity in the Valley. Conclusions Concerning Early Seasons 1. On the basis of available information, hunting seasons beginning on the Saturday nearest October 1 in North Park, South Park and the San Luis Valley are not detrimental to populations of any of the affected species given these current harvest levels. 2. Increased harvest on the various populations during the early October period would not be detrimental to the populations affected as long as the sex and age composition of the harvest remained the same. 3. Seasons beginning earlier than the Saturday nearest October 1 would probably change the species as well as the sex and age composition of the harvest. Some changes would be detrimental to various populations and therefore the effects of an early season should be monitored. Control over the species and sex composition could possibly be obtained through bag limit changes. 4. Current hunting season frameworks allow for sufficient early utilization of the waterfowl resource in the three areas. Therefore, special high mountain seasons will not be requested at this time. LITERATURE CITED Brownie, Cavell, D. R. Anderson, K.P. Burnham, and D. S. Robson. 1978. Statistical inference from band recovery data -- a handbook. U.S. Dept. Interior, Fish Wildl. Servo Resource Publ. No. 131. 212pp. Hoffman, R. W., and C. E. Braun. 1975. A volunteer wing collection station. Colo. Div. Wildl. Game Inform. Leafl. No. 101. 3pp. �75 Szymczak, M. R. 1978. Migration and mortality characteristics of duck populations in the Inter-mountain valleys of Colorado. Colo. Div. of Wildl., Game Res. Rep., Fed. Aid Proj. W-88-R. October. p. 47-59. , and J. F. Corey. 1976. Construction and use of the Salt Plains duck .t rap in Colorado. Colo. Div. \\Tildl.Div. Rep. No.6. 13pp. ---- Prepared by: ��October 1981 77 Job Final Report Colorado State of Project Covered: Bird Investigations 7 Job No. Effects Job Title: Personnel: ----- 4 Work Plan No. Period Migratory W-88-R-26 No. of Land Use Changes 1 January 1978 through on Mourning 15 October Doves 1980 T. E. Olson, R. A. Ryder, P. D. Curtis, L. Sweanor, D. A. Rein, V. E. Youngman, Colorado State University; C. E. Braun, R. D. Funk, Colorado Division of Wildlife. ABSTRACT The objectives of this study have been fully achieved. Data were collected during 3 field seasons (April through October 1978, 1979, and 1980). Manuscripts covering study objectives have either been prepared, or are currently in preparation. Those submitted, published, or in progress are listed below. Olson, T. E. 1980. Mourning Colorado. M.S. Thesis. doves and land use changes in eastern Colorado State Univ. Fort Collins. 119pp. 1980. Agricultural land use changes and mourning dove production in northeastern Colorado. Proc. 3rd, Joint Mtg. of the Cooper and Wilson Ornithological Societies, 20 March, Corpus Christi, Texas. Abstract. 1980. Patterns of agricultural land use changes and effects on avian species in northeastern Colorado. Proc. 89th Stated Mtg. of the American Ornithologists' Union, 12 August, Fort Collins, Colorado. Abstract. 1981. Doves and agriculture. Colorado and C. E. Braun. 1979. Land use changes J. Colo.-Wyo. Acad. Sci. 11:95. Abstract. Outdoors 31:8-11. and mournin$ doves. �78 ________, and Effects of changes in agricultural land use on mourning dove nesting and production in eastern Colorado. J. Wildl. Manage. (in prep.). , and ---mourning Relation between cooing activity and nesting of doves in eastern Colorado. J. Wildl. Manage. (in prep.). ----- , and doves in Colorado. Crop gland activity of September harvested mourning Wilson Bull. (in prep.). ------- , and Characteristics of mourning dove nest sites on the eastern Colorado plains. Southwes. Nat. (in prepv), _____ , and R. A. Ryder. 1980. Avian nesting studies on agricultural land in northeastern Colorado. J. Colo.-Wyo. Acad. Sci. 12:44. Abstract. , and ---avian species Prepared by Agricultural land use and effects on nesting of in eastern Colorado. Prairie Nat. (in prep.). 71-- t)£;A< ,if /lp;c.= Thomas E. Olson Graduate Research Assistant Approved by fZ~(;:'/~£~~~ Nu')~ Clait E. Braun . Wildlife Research Leader �79 October 1981 JOB FINAL REPORT State of Project Colorado No. Migratory W-88-R-25 Work Plan No. Job No. 4 Evaluation Job Title: Bird Investigations of Daily Counts of Band-Tailed 8 Pigeons as a Census Method Period Covered: Personnel: 1 January 1980 to 31 March 1981 P. D. Curtis, R. A. Ryder, J. Ellis, T. E. Olson, Colorado State University; C. E. Braun, H. D. Funk, Colorado Division of Wildlife; C. J. Curtis. ABSTRACT Methods of censusing band-tailed pigeons (Columba fasciata) at artificial bait sites were examined with emphasis on daily counts. During 1979-80, 1,342 pigeons were trapped near Evergreen and Niwot, Colorado, primarily with cannon nets. Of this total, 983 birds were newly banded with 862 color-marked with alpha-numerically coded wing tags made from SAFLAG. Investigations of bandtails at study sites were made on alternate days from April through August. With a sample size of 12 days, a 20% change (95% CI) in the mean number of band-tailed pigeons counted per day could be detected at Evergreen while 14 days would be required for counts during 20 May - 10 June. Counts during 8 days would be required to detect a 20% change in the mean number of pigeon coos heard from sunrise to 1600 MDT. Jolly-Seber mark-recapture analyses indicated that numbers of pigeons at Evergreen and Niwot remained stable during 1970-80. Due to year-to-year variability in estimates of population size, these estimates should only be used as indices. of pigeon abundance. Time budget data from Evergreen in 1980 indicated there were no differences (I-way ANOVA, f > 0.05) in the distribution of time spent among 6 behavioral categories between tagged and nontagged bandtails. Most pigeons spent more than 70% of their time at the feeding site in resting or maintenance activity. ��81 INTRODUCTION The band-tailed pigeon is a major migratory western North America with annual·harvests birds (Jeffrey Mexico. 1977). game species in approaching Two populations are recognized The Coastal population (C. !_. north (C. f. fasciata) Columbia. occurs in the forested of monilis) occurs primarily west of the Sierra and Cascade mountains in California, Washington, into central British 500,000 The Interior Oregon, population mountains of Utah, Colorado, New Mexico and Arizona (Braun et ale 1975). Despite the abundance and wide distribution of the species, little information is available on population size and annual changes in breeding technique densities. Pigeon cooing, in reference for the Coastal population, to a census was described (1968), Keppie et al , (1970) and Keppie (973). Jeffrey (1980) reported by Sisson McCaughran and that the Poisson probability mass function best fit pigeon call count frequency data. centration have also been attempted (Jeffrey sites and mineral springs 1977). (Fitzhugh Braun been unsuccessful 1974, Braun et al. due to apparent artificial bait sites. Previously, and banding popu- low calling 1975). (1976) and F. J. Ward (unpubl. tailed pigeons can be attracted for trapping in winter con- Call counts within the range of the Interior lation have previously rates Censuses data) found that band- to and held for long periods at these sites have been established (Braun 1976, Kautz 1977) and several �82 subpopulations of pigeons in Colorado have been studied extensively since 1969 with large samples of birds being banded annually. This study was designed to evaluate the r-eliabrlrty of counts of band-tailed pigeons at artificial bait sites as a census technique. Changes in numbers of color-marked pigeons by daily timed intervals were ascertained from April through for most migrant bandtails Ziegler 1971). 1. September, the breeding (Glover 1953, March and Sadleir 1970, Specific hypotheses examined were: Changes in population size can be reliably detected counts of barrd+tafled pigeons made at intervals out the breeding 2. from through- season. The ratio of pigeons marked in each time interval are present season in the next time interval that does not change from spring to fall. 3. Sex ratios of marked and unmarked pigeons are constant for each daily interval 4. from spring to fall. Timing of pigeon arrival to and departure from feeding sites is constant by age and sex from spring 5. Survival of color-marked differ (patagial tags) until fall. pigeons does not (P > 0.05) from pigeons marked with only leg bands. �83 DESCRIPTION OF STUDY AREAS Data were collected at 3 sites near Evergreen Niwot, Colorado (Fig. 1). they have fairly discrete (Braun 1972). These 2 areas were selected subpopulations Also, continuous owned by Forest trapping, data were available for banding Neff (1947). and observation range and encinal pigeons spp.) The dominant overstory tree pine (Pinus ponderosa). menziesii), center for the Evergreen of band-tailed species area outlined by the breeding montane at Forest Heights was Smaller amounts of Douglas-fir (Picea engelmannii), aspen Few berry-producing are found in this area (Braun (Populus trees and 1973) and pigeons rely heavily by local residents. Climate of the Evergreen annual precipitation on land woodlands. Engelmann spruce were also present. upon grain supplied pigeons described lodgepole pine (Pinus contort a) and quaking tremuloides) Colorado. as Rocky Mountain (petran) (Quercus (Pseudotsuga County, This site was chosen as the main Brown and Lowe (l974~,b) of band-tailed ponderosa 105°18'W) at 2,146 m elevation Jefferson Heights School. and is within the breeding shrubs pigeons and 1970 (Niwot). Heights site (39°38'N, was 0.6 km south of Evergreen, forest because Heights The Forest habitat of band-tailed banding both sites since 1969 (Evergreen) Forest and 1 site near area is dry and cool with average of 40 em and an average mean temperature of �84 • NIWOT JEFFERSON COLORADO Fig. 1. 1979-80. Band-tailed pigeon study areas, northcentral Color ado , �85 6.3 C (U.S. Dep , Commerce 1973). Over two-thirds of this moisture falls as rain between April and September. Upper Bear Creek This site (39°38'N. Creek-Jefferson (Braun 105024'W) was 0.8 km west of the Clear County boundary 1976). on the Upper Bear Creek Road Pigeons fed on grain provided and geese at the Jon-Hill Estate. Forest This site was 6.4 km west of Heights at about 2.100 m elevation. characteristics for domestic ducks were similar to the Forest The habitat Heights and climate site. Conifer The R. Wood residence (39°32'N. west of Conifer near the intersection Mountain Road at an elevation southwest of the Forest of Griffen Drive and Black of 2.743 m approximately Heights site. species was lodgepole pine. 105°23'W) was 4.8 km north- The dominant overstory The area was burned ago and thick pine regeneration shrubs or trees. ground surface slightly cooler than the other has prevented Little understory vegetation 2 Evergreen range by other was present and the The climate is sites. described about 20 years reinvasion was matted with pine needles. within the pigeon breeding 8.4 km but Conifer is by Neff (1947). Niwot The Knaus farm (40006'N, 105°10'W) site was 1.6 km northwest of Niwot. Boulder County. Colorado where pigeons grain storage 1976). area (Braun and was 58 km north fed in a livestock This area was at 1,551 m elevation of the Evergreen study areas. Bandtails �86 follow Left Hand Creek east from the foothills to the feed storage area. The dominant overs tory tree species was cottonwood -(Populus spp.) growing along Left Hand Creek. primarily cropland with corn, barley, alfalfa being the major cultivated tion of 32 cm (U.S. from April through wheat, sugarbeets than the Evergreen and study areas of 9.2 C and mean annual precipita- Dep , Commerce 1973). September. oats, area was crops. This site is warmer and drier with a mean yearly temperature The surrounding Most precipitation occurs This site is outside typical band- tailed pigeon breeding range. No nesting cottonwoods bordering Left Hand Creek. is known to occur in the �87 METHODSAND MATERIALS Baiting Students of the Forest Heights School baited the site daily with 1. 4 (mid-March until 14 May) or 2.8 kg (after corn per day in 1979. were provided During 1980. 2.8 kg of whole corn per day from early April through amount of corn provided maintained throughout was increased 25 May. On 26 May the to 5.6 kg per day, a level the summer. In 1979 and 1980, the groundskeeper (Upper Bear Creek) 14 May) of whole provided domestic ducks and geese. milo, wheat and proso millet. of the Jon-Hill Estate 1. 4 kg of mixed grain daily for The mixture consisted Bandtails of cracked corn. used this food when domestic fowl were not feeding at the site. During 1979 and 1980, the R. Wood family (Conifer) approximately 5 feeders. 4.2 kg of cracked seeds daily in Some pigeons also used small amounts of a peanut butter-cornmeal Bandtails mix. at Niwot fed at a grain storage unlimited food supply. until 7 June, ground a rolled corn-barley the rolled corn-barley sharply corn and sunflower supplied after area and had an In 1979, from the time of arrival whole ear corn was available. mix was stored at the site. mix, as numbers the change in feed. corn was available until 5 June. observed During After (mid-April) After 7 June Pigeons preferred daily increased 1980, ground 5 June separate whole ear piles of �88 cracked corn and barley disliked the cracked nesting areas, were stored at the site. corn or found alternate for smaller numbers Bandtails apparently feeding sites closer to of pigeons used the site in 1980 than in 1979. Trapping Band-tailed pigeons were trapped at Forest Heights and Niwot with cannon nets in both 1979 and 1980. Small numbers were also captured Heights in 1979 with a drop net. at Conifer and Forest Trapping was not attempted used were those described Information location, by Braun of pigeons· at Upper Bear Creek. (1976). collected for each bird handled included: age, time, U. S. Fish and Wildlife Service band number, and color if newly banded, a subadult), weight, primary and crop gland activity. Bandtails 1976, White and Braun activity by palpation Fitzhugh 1974). a Hanson dietetic as Crop gland of the crop lining Weights were recorded molt if were classified 1978). sex, tag number molt (secondary to age and sex (Braun was determined Methods to the nearest (Zeigler 1971, 2 grams using platform scale (Drewien et al , 1966). Tagging Unbanded pigeons captured Service size 5) and color-marked. II SAFLAGII material Pawtucket, R.1. were banded (U .S. Fish and Wildlife Markers were fabricated (The Safety Flag Co. of America, P.O. 02862). from Box 1005, The tag style was a modification of that used by Hewitt and Austin-Smith (1966). Two wing markers placed on each bird and held in place by cutting were a 6-7 mm slit in �89 the tags. pulling the tab on the tag strap and secUring the tab with a single staple Forest Heights were coded with a letter Niwot with 2 letters. through (Fig. 2). this opening, Tags used at and a number, those at while tags at Conifer were coded with 2 numbers. Data Collection Counts of pigeons were made twice weekly at Forest Heights and Niwot in 1979 and 1980, and once weekly at Upper Bear Creek in 1979. Counts were made daily at Conifer in 1979 and 1980 by the R. Wood family. The most accurate pigeons present present were made during fed during each session. estimates of the number of feeding sessions since most birds At this time. a count estimate of the total number feeding was made and tag color and code of marked individuals were recorded. Sex and age ratio estimates were made from counts of groups of feeding band-tailed pigeons. In both 1979 and 1980, the number of different cooing was recorded throughout examination of the relationship the day. pigeons heard These data allowed between pigeon cooing and numbers observed. In both years, the number of different feeding each day was estimated by adjusting birds observed pigeons observed the total number of feeding based on tagged bird reobservations the formula m y X r n=1 P -n (n ) using �90 4 jE-2.8CM SCM I 7MM 12CM I ~~ -5CM-~1 LEFT RIGHT PULLTAB------~~---+~~~ FOLD OVER WING NEXT TO BODY THROUGH SLIT AND SECURE WITH STAPLE PLACE STRAP BETWEEN TERTIALS AND BODY 2. Tag design used method of attachment.. Fig. to color+ma r k band-tailed pi gcons and �91 where X estimated total number of different =: birds observed feeding per day, Y = total number observed n = number of times a tagged feeding (1, 2, ••. feeding per day, individual n), m = maximum r = number of tagged individuals t = total number of tagged individuals P = -n was observed number of times a tagged seen pigeon was observed, n times, seen per day, then -n r It, proportion -n - (including This formula provided proportion of tagged repeats) birds same number of times. birds observed feeding n times. reliable estimates if it was assumed that the reobserved not differ from the proportion in the population of daily sum of tags observed a given number of times did of nontagged The greater the percentage (and correspondingly each day), In 1980, band-tailed pigeons reobserved of tagged the more different the more accurate pigeon behavioral the estimate. data were collected sample of focal (a particular individual during sampling period) at the Forest site. Each pigeon selected with all behaviors observed observed Heights for 10 minutes and the time of change between behavior's a digital stopwatch) being recorded. used between observation by the bandtails individuals was continuously periods. were color-coded. birds tagged from a stratified an entire the Two minute test periods Five trees most frequently Since it was difficult (using were used to randomize �92 selection of individual pigeons, selection of trees was randomized using a random. numbers table. The most obvious pigeon of the correct code. age and sex was selected from the tree with the proper Nontagged male pigeons were observed and nontagged females from 1100 to 1400. birds became visible, from 0630 to 0930 MDT Any time that a tagged it was included in the next vation period regardless of sex. lO-minute obser- Immatures were included in the sample whenever they were observed in a randomly selected tree. Five color-coded flags were placed near the bait to randomize selection of feeding pigeons. The closest bird of the proper age and sex to the colored flag was selected. Population Modeling Banding data from 1970-80 at Forest Heights and Niwot were analyzed with the Jolly-Seber stochastic immigration (Jolly 1965, Seber 1973:219). model including The Fortran death and program used was written by Kautz (1977) as modified from the original published by Davies (1971:431-438). were made using recapture Annual population estimates data from trap samples. Immature band- tails were not included in these analyses because of differences behavior patterns 1977)• and the likelihood of lower survival rates This population model also gave estimates of survival probability (l-[mortality + emigration». in (Kautz �93 R'.:i;.S'JLTS Trapping During 1979, Heights, 1,076 band-tailed Of the 1,076 pigeons handled. of birds banded from 1970 through (9.2%) captures 298 (27.4%) 1979. Sixty-four tagged in 1979 were recaptured site where banded. Forest at Forest Niwot, and Conifer (Table 1) of which 778 were newly banded. birds pigeons were trapped occurred were recaptures of the 698 at least once at the Fifty-two of the 64 (81. 3%) tagged pigeon re- at Niwot, while only 12 of 64 (18.7%) occurred 266 bandtails Heights Heights. During 1980, were trapped at Forest and Niwot (Table 1) and 205 pigeons were newly banded. 266 pigeons, through 61 (22.9%) 1980. recaptured (54.5%) at were recaptures Eleven of the 164 birds of birds (6.7%) banded tagged at least once at the site where banded. tagged pigeon recaptures remaining 5 (45.5%) occurred occurred at Forest Of the from 1970 in 1980 were Six of the 11 at Niwot, while the Heights. Of the 778 pigeons newly banded in 1979 (Table 2), were males and 421 (54.1%) were females. were too young to be classified 778 birds as to sex. 312 (40.1%) Forty-five (5.8%) immatures Sixty-six (8.5%) of the newly banded were immatures. In 1980, 85 (41. 5%) of the 205 newly banded bandtails males and 118 (57.6%) were females (Table 3). were Two (1. 0%) immatures �Table 1. Band-tailed Date pigeon Fa N N F N F N F N C F 1980 19 May 20 May 24 May 08 Jun 05 Sep Subtotals Totals success Total pigeons captured Site 1979 14 May 15 May 22 May 04 Jun 05 Jun 25 Jun 26 Jun 27 Jul 28 Jul c 27 Aug 28 Aug Subtotals trapping F N N F N at Niwot, Forest Heights, RecaEtures Tagged N - 9; N - 0 25 106 63 97 147 121 190 82 198 40 7 1,076 9 28 18 29 34 27 66 13 69 5 0 298 36.0 '26.4 28.6 29.9 23.1 22.3 34.7 15.8 34.8 12.5 0.0 27.7 23 58 12 146 27 266 1,342 7 19 2 30 3 61 359 30.4 32.8 16.7 20.5 11.1 22.9 26.8 1b 2 6 3 21 5 24 % 1.6 2.1 4.1 2.5 11.1 6.1 12.1 0 64 0.0 5.9 5 8.6 5 1 11 75 3.4 3.7 4.1 5.6 and Conifer, Total birds tagged 16 78 45 68 100 94 101 68 93 35 1979-80. Color Total controls banded RED YELLOW ORANGE BLACK GREEN WHITE BLUE YELLOW RED CHROME 13 23 1 36 7 80 698 16 39 10 99 PINK YELLOW YELLOW BLACK 17 24 41 121 164 862 ----~~ aF = Forest bInc1udes Heights, only pigeons cAll pigeons captured N = Niwot, C = Conifer. tagged except at the site where retrapped. those on 27-28 August (drop trap) were caught in cannon nets. 1.0 +:- �Table 2. Age and sex of band-tailed Site pigeons AHY Date Forest Heights 14 May 04 Jun 25 Jun 27 Jul 28 Aug captured a New Recaps New Recaps New Recaps New Recaps New Recaps Subtotals 6 2 12 5 38 17 10 6 1 Males SY at Niwot, HY 3 1 19 8 1 2 Forest AHY 9 7 46 23 22 8 14 5 Heights, Females SY 32 32 11 10 5 33 9 42 30 33 39 244 1 20 4 14 2 52 5 1 3 1 347 88 HY Unknown sex HY 13 1 1 2 17 332 3 134 44 34 17 24 13 60 24 45 32 38 22 309 11 23 6 78 2 18 2 18 4 4 2 3 1 6 10 11 449 126 10 45 6 Totals 1 7 12 2 21 1 1979. 16 9 68 29 94 27 69 13 7 1 4 97 and Conifer, Niwot 15 May 22 May 05 Jun 26 Jun 28 Jul New Recaps New Recaps New Recaps New Recaps New Recaps Subtotals 27 Aug Conifer New Recaps Totals aNew = newly banded, recaps = previously 5 6 banded. 1 1 6 14 1 17 78 28 45 18 113 34 124 66 129 69 704 35 5 1,076 1.0 V1 �Table 3. Age and Site Forest sex of band-tailed Date Heights 19 May 08 Jun AHY a New Recaps New Recaps Subtotals Niwot 20 May newly Males SY 4 2 35 14 55 14 18 2 at Niwot HY 1 and Females AHY SY 10 5 47 16 78 15 Forest Heights, HY 1980. Unknown sex HY Totals 16 7 116 30 169 1 20 21 1 5 7 3 5 2 Subtotals 1 1 33 3 5 2 44 5 5 2 Totals 88 18 5 122 26 5 2 266 05 Sep = New Recaps New Recaps New Recaps captured 39 19 10 2 24 3 97 24 May aNew pigeons banded, recaps 19 10 4 2 9 4 = previously banded. 2 1.0 0\ �97 were too young to be classified as to sex. Twelve (5.9%) of the 205 newly banded pigeons were nmnatures. In 1979, percent captured) decreased recaptures (total recaptures with each successive Heights while the opposite occurred tagged recaptures increased trapping attempt at Forest at Niwot (Table 1). (total tagged recaptures with each successive -;-total pigeons trapping Percent -;- total pigeons captured) attempt at both Forest Heights and Niwot. Percent recaptures decreased attempt at both sites in 1980. with each successive Percent tagged recaptures trapping were lower in 1980 trap samples at Niwot and Forest Heights than the levels observed at the end of the 1979 trapping season, even though more pigeons were tagged. Daily Site Attendance During 1979 and 1980 at both Niwot and Forest 95%confidence limits were wide for the average number of birds present variation per hour. fewer bandtails numbers during the count estimates of There was much day-to-day in the number of pigeons counted during Generally, Heights, were observed any hour period. in the morning with peak midday or afternoon intervals. At Niwot, during May 1979, daily peaks in the number of pigeons present occurred between 1000 and 1300 MDT (Fig. 3). During the - same period in 1980, far fewer (95%CL, P < 0.05) bandtails were observed numbers and no discernible of pigeons were observed 1979 (Fig. peaks were recorded. during corresponding Greater intervals in June 4) than in May 1979, although this difference was not �98 o rt') CD o o o o rt') rt') rt') ~ CO m o o o o rt') o o o j'(') j'(') N o j'(') V TIME Fig. 3. Band-tailed pigeon daily attendance patterns, Niwot, May 1979-80. Horizontal center line mean, vertical bar = 95%confidence interval. = �99 01979 f?&f1980 II: :::) o :I: ......40 ••• z '" UJ IIJ II: 0UJ Z o IoU C) 0LL. o a: w m ~ :;:) z o w ~ ::E I- en IoU UJ C) .r <t 0:: UJ ~ o ,." co o o ,." ••••• o o rt) CX) o o ,." en o o o ,." o ,." o ,." N o o ,." v ,." ,." TIME Fig. 4. Band-tailed pigeon daily attendance patterns, June 1979-80. Horizontal center line mean, vertical 95%confidence interval. = Niwot, bar = �100 significant occurred (95% CI, P > 0.05). The hourly peak during June 1979 between 1400 and 1500. during June The number of pigeons counted 1980 at Niwot was lower (95% CI, P < 0.05) number counted. in June 1979. No discernible than the peaks were recorded and the number counted in June was not greater (95% CI, P > 0.05) than in May 1980. At Forest Heights in May 19.79, daily peaks in the number of bandtails present occurred During May 1980, lower during between 1200 and 1300 MDT (Fig. pigeon numbers were greater in the morning and midday although these differences were not significant (95% CI~ P > 0.05). Greater counted during corresponding in May 1979, numbers of band-tailed intervals but this difference was not significant in June during 1980 as compared to June 1979. differences (95% CI, P > 0.05) 6) than (95% CI, P > 0.05). between 1300 and 1400, while the peak in 1980 was between 1400 and 1500. lower (95% CI, P > 0.05) pigeons were in June 1979 (Fig. The hourly peak during June 1979 occurred were slightly 5). Pigeon numbers midday and afternoon In general, no 'significant were found between years or months at Forest Heights. Confidence limits for the 1980 data were narrower because of the increased 20 in 1979). Daily observations sites in 1980 during established cooing. number of observation the intensive to investigate than for 1979 days (30 in 1980 vs. were made simultaneously study the relationship period at both (25 May - _7Jun ), between pigeon counts and �101 01979 ml980 0: ~ o :r: .•.•. ~ Z l&J (I) l&.I 0:: Q. (I) Z o l&J C) Q. lL o 0:: lAJ ~ 80 ~ z o lAJ ~ :t 60 i= (I) lAJ W C) 40 <{ 0:: W > <{ 20 o If') <D o o If') Q) o o o 0\ o rn o rn o rn o o N '¢ rn rn TIME Fig. 5. Band-tailed pigeon daily attendance patterns, Forest Heights. May 1979-80. Horizontal center line = mean. vertical bar 95%confidence interval. = �102 210 01979 19 Ffal980 a: :l 0 ::z:: '" •••• Z LLI C/) LLI a: Q. C/) Z 0 LLI C) Q. u, 0 110 a:: lIJ CD ~ :l 90 Z C LLI •••• ~ :e 70 i= C/) LLI LLI 50 C) ct a:: LLI > ~ 30 10 o f(') CD o o f(') o f(') o f(') o f(') ••••• (J) m o o o o o f(') o f(') o f(') N v TIME Fig. 6. Band-tailed pigeon daily attendance patterns, Forest Heights, June 1979-80. Horizontal center line = mean, vertical bar = 95%confidence interval. �103 Evening observations during July and August, sites until sunset, were made at both Niwot and Forest 1979. most birds Although bandtails had departed Heights remained at both the site by 1600. Most feeding sessions after this time numbered less than 20 pigeons and few tagged" birds earlier arrived at the site which had not been observed ".i·nevalue oi e vezun g coun cs 'rlas .uw, ci.uu cne aame day. none was made in 1980. Seasonal Site Attendance At Niwot in 1979, more bandtails were observed during 3rd week of June than at any other time throughout (Fig. 7). Nearly 1,400 different birds 2nd peak in numbers was observed more than 1,000 different birds were observed count of the summer. occurred during 400 birds sharply birds the last week of May, the lowest During 1980, the mean number counted per day decreased per day were observed (95%CI, P < 0.05) The mean number of pigeons observed per day was 743 in 1979 but only 55 in 1980. birds Less than 200 the 1st and 2nd weeks of August when only about from 1979 (Table 4). attendance A Another large decrease in site attendance per day were observed. of different per day. the 3rd week of July when pigeons were seen daily. per day during the summer were observed during the More than only during the 3rd week in July. 150 pigeons Pigeon was low most of the summer and less than 80 different were counted most observation days. �104 o .l&J 1400 1979 > Q: l&J fI) CD o fI) 1000 800 z fil<!) 0. ••• Z l&J Q: l&J I&. I&. o 360 I&. 320 Q: 280 m 240 :> 200 o l&J 2! z 1 1980 I~ ,,,,,, ,, ,,, ,,, , , f , ,, I I I I I o IIJ ti I I I 2! , ~ I I t; au •••• , ,'.... 7 15 22 MAY Fig. 7. Estimated numbers daily at Niwot, 1979-80. 7 I .•.. , .•.. ~J. ..., 15 22 JUN 7 of different band-tailed 15 22 JUL \ \ \, 'I ~' ""- \\ 7 \ -.4 '' , 15 22 AUG pigeons observed �105 pigeons observed Site Year Days (N) Mean number of different pigeons observed I daya ' 95%CI Niwot 1979 23 743 624-862 1980 26 55 27-83 1979 21 154 107-201 1980 33 116 74-158 1979 72 81 70-92 1980 76 66 54-78 1979 8 65 31-99 Band-tailed Table 4. 1979-80. Forest Heights Conifer Upper Bear Creek aDaily totals At Forest during different birds occurred during summer. using formula on pg. the 3rd week of June were counted per day. during (Table 4). Approximately increase Less than occurred 350 in numbers 20 pigeons More birds were observed 15 June fewer bandtails summer. A small increase weeks of July but during during per day the 2nd week of In 1980, the mean number per day decreased but after pigeons also the 3rd week of May, the lowest count of the Other marked decreases observed 8). Another the 3rd week in July. July and the 1st week of August. pigeons (Fig. sites, 9. Heights in 1979, the peak in pigeon numbers occurred were counted calculated per day at all study slightly during (95% CI, P > 0.05) late May and early June, were counted was observed during the rest of the the 2nd and 3rd the 1st 2 weeks of August, per day were counted. of less than 10 �1919-1980----- o W I > ,I,I" lLQ: OW Q:C/) m wO m " T', I ,, :sCI) I zO I ;:)z OW LLJ~ •.... t-Q. <tt- ~z W LLJW o 0'\ \ \ \ tCl)Q: u, u, o \ \ \ \ 80 \ \ \ , '" Fig. 8. 1979-80. Estimated numbers of different , .... ...•.. ...J band-tailed pigeons observed dally at Forest. Heights, �107 At Conifer in 1979, large day-to-day number of different observed birds counted (Fig. variation occurred 9). More bandtails during late summer and the peak count occurred the last week of August. at this time. were during Over 240 pigeons per day were counted No unusually low counts were recorded. the mean number of birds observed (95%CI, P > 0.05) (Table 4). per day decreased Peak counts occurred 3rd week of May (more than 240 pigeons per day) Generally, in the pigeon numbers were highest lower during July and August. During 1980, slishtly during the (Fig. 10). during May and June and The same pattern was also observed at Forest Heights in 1980. Site attendance during 1979. No large differences June and July, (Fig. 11). at Upper Bear Creek was checked periodically in numbers were observed and 50-130 different Numbers decreased bandtarls were counted daily during August and less than 20 pigeons per day were observed. Since fewer birds at this site in 1979 than at the other 2 Evergreen counts were discontinued keeper indicated in 1979. after 1979. that fewer bandtails This reduction areas, Discussions with the groundswere present during 1980 than late sites. (Seber 1973) mark and recapture used to estimate. population size and survival Capture study Population Estimates The Jolly-Seber band-tailed were observed in numbers in 1980, especially during summer, was similar at all study Jolly-Seber during model was probabilities pigeons for the Niwot and Evergreen of adult subpopulations. data (Table 5) from 1970-80 were obtained from the original �300 0 l&J > 240 u..a:: owrn a:: Wen O en :Ern 180 :::>z zO w o(!) w- ti Q. A , 120~ :E~ -w tia:: wwu, u, - n 1\1_ fan IV 1J daily at 60 0 Fig. 9. Conifer, Estimated 1979. numbers of different band-tailed pigeons observed •.... 0 OJ �0 w > 240 lLQ:: OW Q::en wOm m :e en 180 :::>z z@ O(!) w- tt 0.. 120, ~I_z ~ 11,11 111\/\ 1\ 1\ ,_. 0 ~ \0 I-W Cf)Q:: wwu, u, -0 60 Fig. 10. Estimated numbers daily at Conifer, 1980. of different band-tailed pigeons observed �130 0 W 110 > LLO:: OW en m Wo men 0:: 90 ~z z 0 70 o(!) WW ~Q. ~ ~ 50 1 -w •..• 0:: f3wu, 0 .-.- \ 0 LL 10 7 15 22 MAY Fig. Bear 11. Estimated Creek, 1979. numbers 7 15 22 JUN of different 7 15 22 JUL band-tailed pigeons 7 10 22 AUG observed daily at Upper �111 banding records trapping data from the Knaus farm site were included. Evergreen estimates, data were pooled for 14 trap Number Year Captured Released Ever- 1970 green 1971 1972 1973 1974 1975 1976 1977 1978 1979 1980 321 297 244 147 149 171 157 153· 165 580 84 301 294 243 145 149 168 138 153 165 579 83 309 223 357 158 318 49 104 143 156 254 156 303 218 355 158 314 49 104 143 156 252 156 Recaptures 70 73 36 25 43 4 .11 29 7 3 5 3 3 10 5 2 1 2 2 0 2 2 0 2 4 3 0 0 0 14 8 5 3 3 5 0 29 30 4 3 0 0 1 0 0 0 Probability errors. the estimates of survival 71 52 8 13 2 0 0 0 0 0 38 35 4 1 1 1 0 1 23 3 2 5 1 2 3 increased 12 13 12 4 5 0 4 11 10 2 0 15 10 6 0 6 2 2 6 10 8 13 1 0 9 5 1 0 5 4 9 26 4 6 20 2 37 3 13 14 8 4 from 1970 to 1980 at have large standard was variable and by year of last caEture· 74 75 76 77 78· 79 72 Numbers of pigeons apparently Niwot; however, sub- pigeons trapped Table 5. Capture data for adult band-tailed banded at Niwot and Evergreen, 1970-80. 1971 1972 1973 1974 1975 1976 1977 1978 1979 1980 locations 1972). (Braun Niwot 1970 For the Heights and Conifer) within the Evergreen (including Forest population banding only For the Niwot estimates, and field forms. errors (Table 6). (58.3-111.5%) with large standard Although biologically impossible, survival estimates and �112 standard errors greater than unity were reported as computed to show their imprecision. Table 6. Jolly-Seber estimates of population parameters for adult band-tailed pigeons captured and banded at Niwot and Evergreen. 1970-80. Year Niwot Evergreen Marked Proportion Total 1970 1971 1972 1973 1974 1975 1976 1977 1978 1979 1980 0.121 0.279 0.300 0.195 0.211 0.242 0.170 0.170 0.191 0~274 1970 1971 1972 1973 1974 1975 1976 1977 1978 1979 1980 0.130 0.230 0.317 0.233 0.490 0.144 0.196 0.083 0.189 0.154 0.0 185.0 266.8 257.7 271.1 387.3 314.0 300.6 362.1 556.4 0.0 139.5 217.9 301. 2 291.4 252.7 177,8 186.3 339.2 1,308. 0 Total number (N) Probabili ty of survival (PHI) a 1,526 957 861 1,393 1.840 1,2.97 1,769 2,134 2~907 1,072 949 952 1,252 516 1,233 952 4.070 6.922 0.615 0.602 0.583 0.756 0.990 0.605 0.726 0.847 1.115 Standard N 0.090 0.090 0.102 0.144 0.200 0.129 0.148 0.162 0.391 300 154 164 306 409 277 411 484 992 0.460 0.663 0.614 0.712 0.476 0.640 0.698 1.126 2.713 error PHI 0.065 0.089 0.099 0.148 0.152 0.225 0.174 0.294 1.484 215 133 175 245 173 387 241 1.416 3,726 aSurvival probabilities and standard errors > 1.0 are reported computed to show their imprecision. It would be more proper biologically to report these as 1. o. No consistent trend in numbers was apparent 1970 to 1980 except for an increase during crease was probably were variable were large. an artifact 1978 and 1979. of sampling. (46.0-271.3%) and standard at Evergreen from This in- Survival probabilities errors as of these estimates �113 Since trapping Heights, was conducted 5 times at Niwot and 4 at Forest it was possible to obtain Jolly-Seber ture data of adult bandtails estimates from recap- newly banded in 1979. (Table 7) were obtained primarily from wing-tagged banded controls were also included. leased at Niwot (391 vs. (14.3. vs. data pigeons, but More marked adults were re- 180) and the recapture rate was higher 5.0%) than at Forest Heights. Table 7. Capture data for adult band-tailed banded at Niwot and Forest Heights, 1979. pigeons trapped Number Captured. Released Date Niwot Capture and Recaptures by time of last capture May June 15 22 05 26 15 May 79 78 22 May 46 46 1 05 Jun 119 119 3 3 26 Jun 148 148 9 3 12 28 JuI 133 133 2 3 6 14 14 May 04 Jun 25 Jun Forest Heights 14 May 16 16 04 Jun 70 70 2 25 Jun 95 94 1 2 27 JuI 58 57 0 2 Jolly-Seber estimates of population parameters banded in 1979 were of little value. Standard for pigeons errors were large 2 �114 (Table 8) and estimates obtained mended that this technique over a short were meaningless. It is not r'ecom- be used on small data sets collected period of time. Table 8. Jolly-Seber estimates of population parameters for adult band-tailed pigeons captured and banded at Niwot and Forest Heights, 1979. Date Niwot Forest Heights Total Probability number . of survival (PHI) a (N) Marked Proportion Total 15 May 0.0 Standard N 0.930 error PHI 0.353 22 May 0.022 72.6 3,338 1.007 3,513 0.379 05 Jun 0.050 118.4 2,348 0.606 1,145 0.216 26 Jun 0.162 140.3 865 28 Jul 0.188 14 May 312 0.0 1.219 04 Jun 0.029 19.5 683 25 Jun 0.032 97.0 3,072 27 Jul 0.069 1.109 1.180 803 1.089 3,409 aSurvival probabilities and standard errors > 1.0 are reported as computed to show their imprecision. It would be more proper biologically to report these as 1.o. Daily Cooing Patterns Band-tailed pigeon cooing in May 1979 at Niwot, peaked between .1200 and 1300 MDT (Fig. 12). In 1980, the peak in the average number of coos heard per hour occurred with a lower level of cooing intensity P > 0.05). between between 1000 and 1100, 1100 and 1500 (95% CI. The peak in pigeon cooing at Niwot during June 1979 �llS 0:: 50 01979 :;:) 0 :x: ..••.. ~1980 c 40 0:: « l&.I ::I: (/) 0 0 30 0 u, 0 a:: w m 20 :E ::l Z l&.I <!) 10 <t 0: w > <t 0 ,.,., 0 ,.,., 0 ,.,., to ~ 0 CD 0 0 0 rt') 0 rt') m rt') 0 0 0 0 ,.,., 0 ,.,., ,.,., N 0 ,.,., '¢' TIME Fig. 12. Band-tailed pigeon daily cooing patterns, Niwot, May 1979-80. Horizontal center line mean, vertical bar 95% confidence interval. = = �116 occurred between P < 0.05) Greater 1200 and 1300 (Fig. peaks were recorded. numbers of coos were heard between 1000 and 1500 in June May 1979 (95% CI, P > 0.05). Peaks in pigeon cooing occurred May 1979 at Forest Heights in 1980~ the peak occurred coos were heard during similar between years 1300 and 1400 (Fig. in both years. In 1979, occurred 14). between the greatest 15), 1300 and 1400. Although more in 1980 ~ the pattern At Forest was Heights during number of coos was heard between and patterns were similar (95% CI, P > 0.05) numbers of coos were heard between 1000 1979 vs , May 1979 (95% CI~ P > 0 •.05). no differences (95% CI, P > 0.05) between Niwot and Forest Band-tailed in May and June 1200 and 1300 during During the same time period all time periods Greater and 1500 in June (Fig. between (95% CI, P > 0.05). 1979 and 1980, June. In 1980~ fewer (95% CI, coos were heard and no discernible 1979 vs. June 13). in cooing patterns Heights during pigeons cooed less either Mayor (95% CI, P < 0.05) 1980 than those at Forest at Niwot Heights. Seasonal Cooing Patterns The peak in band-tailed Heights in 1979 occurred 17) • (95% Cooing intensity cr, P < 0.05) (Table 9). pigeon cooing at Niwot, and Forest during decreased the 2nd week of June in August P < 0.05) 1979 at Niwot; however, number !: > (95% cr, both years. 0.05) June or July coos were recorded summer 1980 vs. approximately of coos was heard at Forest Fewer coos (95% CL, P < 0.05) 16, at both sites and fewer pigeons were heard than in either Fewer (95% cr, (Figs. during the same Heights in were recorded only �117 60 01979 0::: ::) 0 z ~1980 50 .•.•.••.. 0 0::: .:( IJJ :I: 40 (/) 0 0 0 LL 0 30 0::: IJJ m ~ ::) z 20 IJJ C) .:( 0::: IJJ > 10 .:( o If') U) o o ,... rt'l o o If') o If') ex> 0') o o o rt'l o o If') o If') N o If') v TIME Fig. 13. Band-tailed pigeon daily cooing patterns, Niwot, June 1979- 80. Horizontal center line = mean, vertical bar = 95% confidence interval. �118 0:: 50 01979 :::l 0 ::I: .•.... c 40 0:: ~1980 « w ::I: en 0 0 30 (.) u, 0 0:: lIJ CD 20 :IE :::l Z w C) « 10 0:: LIJ > « 0 0 ro U) 0 0 ro •••• 0 0 ro eo 0 0 ro m 0 ro 0 0 ro 0 0 N ,.,., ro j"f') 0 f1j) V 0 TIME Fig. 14. Band-tailed pigeon daily cooing patterns. Forest May 1979- 80. Horizon tal center line = mean, vertical bar confidence interval. = Heights, 95% �119 o I'f") U) o o ro ,.... o o ro Q) o o I'f") m o o o ro o I'f") o ro N TIME Fig. 15. Band-tailed pigeon daily cooing patterns. Forest June 1979-80. Horizontal center line = mean. vertical bar confidence interval. Heights, 95% = �200 ~ 0 •.•... 0 0: c:t 160J I \ lLJ J: ~ ...J 19791980----- 120 c:t 0 u, 0 0: lLJ OJ 80 1 v I <, to-' N 0 :E :::> z ...J 40 ~ 0 •••• Fig. 16. Band-tailed pigeon seasonal cooing patterns, Niwot 1979-80. �I, " 240 . 19791980----- II I II ?i 0 ••... 0 a: II +, , 'V' ,: , '.' 200 I" II" . ct ," I I , , I, W :x: en , I, ,, I ct , it/ " o u, 0 II I I I I I I a: W en :E ::J " I I I I I • ..J 40 , I , ~ .\ •... \ \ I I I I ~ ~ I .•. A. '\ " ,( ,, " , '.•..•.....••.•. Fig. 17. •.... N •.... , , , ,, ,~ I Z ~ " I' " ," ..J ..J ~ .It , I Band-tailed pigeon seasonal cooing patterns, _ Forest Heights ]979-80. �122 during July (95% Cl, Forest 1980 vs. P > 0.05) Heights Heights. of coos were heard (N = 1,747) far rmre(95% Cl, Heights 1979 at Forest P < 0.05) (N = 2,433) Table 9. Band-tailed Heights, 1979-80. Niwot summer 1979 at and Niwot (N = 1.732). pigeons were heard During 1980, cooing at Forest than at Niwot (N = 197). pigeon coos heard Days Site during Similar numbers per day at Niwot and Forest Mean number coos/day Date (N) May Jun Jul Aug 4 4 5 5 76 136 64 8 0-156 40-232 24-104 0-24 May-Aug 18 67 38-97 6 6 7 6 15 4 7 1 4-27 0-9 0-15 0-4 25 7 3-11 May Jun Jul Aug 3 4 4 4 74 167 100 9 19-131 102-232 48-151 0-30 May-Aug 15 88 52-125 95% Cl 1979 1980 May Jun Jul Aug May-Aug Forest Heights 1979 1980 -..-May Jun Jul Aug 9 9 8 5 127 121 1l.5 1.8 77-177 59-183 0-23 0-4 May-Aug 31 75 46-105 �123 Relationship Between Cooing and Numbers There was a strong correlation between the average coos heard per hour and the· average number of birds = hour in May 1979 at both Forest Heights (1:_2 (~2 = 0.650) (Fig. was greater 18). The y-intercept p < 0.05) for Niwot than Forest were present June. (!_ During Heights indicating was found for both sites 1980. counts of pigeons observed were made simultaneously 25 May - 7 June. the relationship at Niwot and Forest This provided increased average number of bandtails (~2 = 0.363) and Niwot (~2 period was weaker 1979 (Fig. greater (!_ more birds The slope of observed = (Fig. 19). and calls heard Heights during The relationship per hour and the per hour at Forest 0.143) during the intensive (Ffsher+s ~-transformation, Heights. During sample sizes to verify between calls and numbers. between the average number of coos heard per (! test. p > 0.05) for both sites. test. a weaker correlation present 0.873) and Niwot at that site at low cooing intensities. the lines was. similar number of Heights study ~ < 0.05) than in 20). For Forest the y-intercept test. P < 0.05) in 1980 (45.7) than in 1979 (24.0) in- dicating that more pigeons were attending the site. of the line was The opposite was true for Niwot (5.48 in 1980, 97.7 in 1979) indicating fewer (t test. P < 0.05) birds the lines were also less The relationship (!_ test. attended the site. that The slopes of P < 0.05) for both sites in 1980. between the total number of coos heard hour and the maximum number of pigeons observed during same hour was examined between 0600-0800 MDT (Fig. the 21) and per �130 N / N 0: N :J 0 J o::I: N W ...... m t- 90 :sZ :JW Zen W wO::: C!)Q. eten O:::z W 0 50 ,_. N ~~ Q. LL 30 ~ o E ~ 1\ N 10~ ~ ~ r2=0.650 NIWOT Y=49.9+4.482S, EVERGREEN Y" = 10.3+ 5.23~, r2=0.873 1 5 10 15 20 26 AVERAGE NUMBER OF COOS HEARD/HOUR Fig. May 18. Relationship 1979. between band-tailed pigeon calling and numbers, �5 10 15 20 26 30 AVERAGE NUMBER OF COOS HEARD/HOUR Fig. June 19. Relationship 1979. between band-tailed pigeon calling and numbers, 35 �126 0: ::l 60 0 l: ••..•. IZ W (f) W 0: 1\ Q. Y = 45.7 + 0.78 X ,r_2= (f) z 0.363 0 w (!) E Q. u, 0 0: w m 30 5 15 25 35 45 55 20 ~ :::> z w 15 (!) c:( 0: W ~ 10 2 y= 5.48+ 1.28X, r = 0.143 1\ 5 N -- - N 2 4 6 8 10 AVERAGE NUMBER OF COOS HEARD/HOUR Fig. 20. Relationship between average number of coos heard per hour and average number of pigeons present per hour at Forest Heights (E) and Niwot (N), 25 May - 7 June 1980. �127 0: 120 :l 0 :I: ••••• l(I) bJ 0: 0.. en Z • l&J 0.. (!) lL 0 0: • • •• 80 0 • • Z 100 bJ • 60 l&J CD ::E ::::l 40 A Y=41.8+ 1.97~,!2= 0.216 Z :E ::::l :E x « :E 20 • •• • 10 20 30 TOTAL NUMBER OF COOS HEARD/ HOUR Fig. 21. Relationship between total number of coos heard per hour and maximum number of pigeons present per hour for 0600-0800 MDT, Forest Heights, 25 May - 7' June 1980. �128 1l00-1400 (Fig. 22). during the intensive Height s , The rintercepts equations indicating were similar test. the strength ~-transformation. the morning period indicating present P > 0.05) at Niwot (r2 = 0.854) (t test. = observed during the intensive number of birds observed Statistical was P < 0.05) increase for in number between the total number of coos heard sample size (N 2 The r2 values between morning and there was a greater per day and the total number of birds was weaker (r were present for an equal increase in cooing intensity. A strong relationship With a larger for both of the relationship The slope of the line was greater of birds P > 0.05) during both time periods. were also similar indicating midday. (!_ that similar numbers of bandtails at low cooing intensities equal (Fisher's study period at Forest = 8 vs. N = per day was found study period (Fig. 6) and a greater per day at Forest Heights. 23). (P < 0.05) the relationship 0.189). Properties of Count Data Changes in population size could not be reliably detected direct counts of band-tailed 1979. from pigeons at Niwot or Forest Heights in Due to small sample sizes. variances observed in daily counts of pigeons (Table 10) were large and the 95% confidence intervals for both May and June were wide. Counts of pigeons cooing were less variable at both sites during May. To increase instantaneous sample sizes, the assumption of sampling. counts were made simultaneously at Niwot and Forest Heights during 1980. yet satisfy Means. variances, a 2-week intensive standard errors, study period in and 95% confidence �129 a:: ::J 0 120 ::J: '" lZ W U) 100 W 0: • • • • • • Q. en Z 0 LLJ ~ •• 80 • Q. u, 0 a= lLJ CD ~ ::> z • •• 60 40 ~ ::::> :E • • • x ~ ~ 20 1\ y= 43.5+0.99X, - - -r2=O.247 . • 20 40 60 80 TOTAL NUMBER OF COOS HEARD/ HOUR Fig. 22. Relationship between total number of coos heard per hour and maximum number of pigeons present per hour for 1100-1400 MDT, Forest Heights 25 May - 7 June 1980. �130 350 A 300 ?i a E X,.r.2=O.l89 Y =88.2+0.67 250 ..•.•. 0 l&J > a: lLJ en 200 CD 0 en z 150 0 l&J (!) a. lL 100 50 0 100 150 200 250 a: l&J CD :i! 60, z 50-1 ::> ..J ~ 40 •••• 30 0 20 -y"= 10 10 5.28+1.75X, 20 2=0.854 --r 30 40 50 TOTAL NUMBER OF COOS HEARD I DAY Fig. 23. Relationship between total number of coos heard per day and total number of pigeons observed per day at Forest Heights (E) and Niwot (N). 25 May - 7 June 1980. �131 Table 10. Statistics for the total number of calls heard and estimated number of different band-tailed pigeons observed daily at Niwot and Forest Heights, May and June 1979. Forest Heights Calls! Pigeons counted! day (N) day Calls! day (~) Niwot Pigeons countedl . day May -a x 74 117 76 456 S2 506 14,068 2,530 56,694 SE 10 53 25 cr, 95% 31-119 N, Days 0-264 119 0-156 77-835 5 5 4 4 167 241 136 881 S2 1,647 7,516 3,636 92,385 SE 18 39 30 124 June x cr. 95% 85-249 N, Days ax confidence = 40-232 133-349 5 4 5 mean, S2 interval:- = variance, = SE limits for the total number of calls heard both sites of pigeons was less were calculated (22.8) during June calling per day was less (12.4 vs. standard 136) than in 1979. 1979. error, and pigeons for this time period. per day (Table 6 Cl = observed at The mean number 11) at Niwot during (95% cr. p < 0.05) than the mean number day (881). counted heard counted 562-1200 of birds The mean number (95% cr. P < O.OS) during 1980 per of pigeons 1980 �132 Table 11. Statistics for the total number of calls heard mated number of different band-tailed pigeons observed Niwot and Forest Heights, 25 May - 7 June 1980. Forest Calls I day (N) Heights Pigeons counted I day Calls} day (N) and estidaily at Niwot Pigeons counted I day -a x 170.1 202.0 12.4 22.8 S2 1,439.0 3,900.7 99.8 341.1 SE 13.4 18.0 4.5 6.5 cr, 95% l38. 4-201. 8 162.4":241. 6 8 N. Days similar number between change Heights, study period of pigeons years Tagged Tagging = standard 1979 were compared. of bandtails per day was pigeons at Niwot and Forest throughout Pigeons present Heights (95% cr, P > 0.05) per day. in the mean number of Band-tailed The mean a 20% with a at the 95%confidence with a sample size of 8 days, was continued Cl Consequently, counted would be detected of tagged error, of calls heard per day was also similar A 20%change Proportions were monitored 8 (202.0 in 1980, 241 in 1979). in 1980. Proportions 5 (170.1 in 1980, 167 in 1979) when the and June in the mean number calling per day, SE the mean number counted samp le size of 12 days, interval variance, (95% cr, P > 0.05) intensive. 7.4-38.2 12 a2 x = mean, S confidence interval-:- At Forest 0.0-24.9 of birds heard could also be detected. Observed during during feeding sessions 1979 and 1980. the summer at both sites (Table 1). �133 At Niwot in 1979, tagged 5 June, then stabilized again increased were detected tions at Forest Heights, 21 May through served 18 July, other 24). of birds present between years at Niwot. Tagged proporfrom after which they declined during (Fig. the re- (95% cr. P > 0.05) were No differences Lower tagged 25). Tagged percentages were higher at Niwot than Forest (95%CL, P < 0.05) were ob- proportions Heights from 5 June through differences Tagged proportions (95% cr, P > 0.05) in at feeding sessions at Forest (Fig. No differences found within or between years Heights. until 3 August when they in both 1979 and 1980, increased mainder of the summer. observed from 21 May to in 1980 due to fewer numbers feeding sessions. proportions increased with no changes (95%Cf , P > 0.05) were more variable during proportions 3 July (P > 0.05) were found between sites 1979. No during 1979 or 1980. Tagged percentages by color were also examined throughout summer 1979 at Niwot and Forest tagged proportions changed. tagged Lower percentages RED during 2-week intervals greater of bandtails differences marked during May at Forest (Student's !_ test, marked in. May were present whether specific time intervals P < 0.05) of pigeons Heights were recorded !_ test, June was observed (Student's !_ test, (Student's from 19 July through percentage WHITE during Heights to ascertain 1 September during (Table 12). ~ < 0.025) of birds during 4-18 July. ~ > 0.05) were found. 3 A tagged No o_ther Fewer pigeons during, late summer, therefore the ratio �134 0.24 t--I95CfoCI 0.20 MEAN • -1979 0.38 0.27 0.30 ••• • 0.45 ••• • ••• • •• •• --1980 z o 0.16 ._ a:: o a. o a:: a. o 0.12 LLJ C) C) <t I- 0.08 / / / / 0.04 Z ...J ::J ..., NV Fig. 24. observed 1(1) IO_ ::J ..., 1m V Average daily tagged proportions of band-tailed during 2-week intervals at Niwot, 1979-80. I f'()CJ) pigeons �l35 0.16 ~ 9s%Cl • MEAN z 0 0.12 t- -1979 --1980 a:: 0 0.. 0 a:: 0.08 Q.. 0 W <!) <!) oct 0.04 t- 0.00 , >-2 ~.., <:l NV Fig. 25. observed I 2 .., 2....1 .., .., :l :l:l en 0 Nrt) . I I()_ ....I :l ..., ICD V_ ~<!) <!) :l:l ..,< « :l 10') ~N rt)_ Average daily tagged proportions of band-tailed pigeons during 2-week intervals at Forest Heights, 1979-80. �Table 12. Tagged band-tailed pigeons 21 May-~- 54 Jun 19 Jun Forest observed daily at Forest 20 Jun3 Jul Heights and Niwot, Tagged EroEortion (%) 419 JulIS Jul 2 Aug 1979. 317 Aug 18 Aug1 Sep Heights Total birds observed 617 RED 0.8 BLACK 646 O.Sa 2.0 WHITE 611 414 0.8 2.8 60 0.5 0.5 0.0* 0.0* 0.0* 3.9 3.9 2.6 3.3 1.7 1.4 2.7** 1.4 0.4 3.3 2.1 - 1.2 1.7 6.1 4.9 6.7 YELLOW All colors 246 500 5.8 7.1 Niwot Total birds observed 1,794 2,813 2,442 2,327 2,481 1,493 641 YELLOW 4.1 4.2 4.0 '3.4 2.4*** 2.1*** 2.2*** ORANGE 2.3 2.9 2.5 2.4 1. 5*** 1.1 *** O.S*** 6.2 - 4.4 4.6 3.4* 4.1 3.9 4.5 4.8 3.5** 4.5 4.7 5.8 S.4 7.2 16.6 20.4 18.8 GREEN BLUE RED I All colors aStudent's t tests * Significant - at 6.4 13.3 for differences ex = 0.05. ** Significant at ex = 0.025. *** Significant at ex = 0.01. 15.4 15.2 by color were based on the underlined value. ,_. w 0\ �137 of birds marked in each time interval next time interval changed Lower proportions that were present from spring (Student's!. to fall. test, P < 0.01) of band-tailed pigeons tagged YELLOWand ORANGE at Niwot during were recorded during 1 September. Lower percentages birds tagged during 3 2-week intervals were not present 2 August. time interval June were observed at the site after mid-July, changed from spring P < 0.05) of f test, Many bandtails pigeons marked in each time interval therefore, fewer birds 8 June at either attended daily were based on reobservation birds, reobservation in 1979. was (Fig. 26). at Forest different (X 2 = bandtails proportions at Niwot and Forest distribution and far late summer. served Reobservation were were color- Heights. Since the estimates of the number of different examined. in the next Similar analyses Niwot or Forest both sites during patterns the ratio of that were present to fall. only marked in May not performed on the 1980 data because no bandtails marked after May 1979 from 19 July through (Studerrt's GREEN and BLUE during 19 July through in the ob- of tagged Heights were in 1979 of pigeons tagged 55.2, P < 0.005) at these At Niwot. more pigeons were seen after sites marking than Heights. No differences distribution (X2 = O. 01. ~ >. O. 90) in the reobservation in 1980 of bandtails Niwot and Forest Heights (Fig. 15+ were pooled so the expected tagged in 1980 were found between 27). Data from classes frequencies This pooling may have masked differences 1-5 through in all cells exceeded in some reobservation 5. clas ses , �138 NIWOT ~ ~ = 417 (217) FOREST HEIGHTS o 1-5 6-10 11-15 0 ~= 246 15+ NUMBER OF DAYS REOBSERVED Fig. 26. Reobservations of band-tailed 1979 at Niwot and Forest Heights. pigeons in 1979, tagged in �l39 o 60 NIWOT~ I!J <.!) <.!) FOREST HEIGHTS N=49 0~ = 115 ~ _. 40 ~ ~ IL o •... 20 z I!J o Q: I!J 0.. o ~~~~~~L-'_~_'_'~~~ o 1-5 6~10 __~_'~ __ 11-15 15+ NUMBER OF DAYS REOBSERVED Fig. 27. Reobservations 1980 at Niwot and Forest of band-tailed Heights. pigeons in 1980, tagged in �140 2 (X = 3.68, Slight significance between the reobservation observed classes distribution in 1980 at Niwot and Forest 6-10 through Differences observed 0.05 < ~ < 0.10) was found of birds Heights marked in 1979 but (Fig. 28). Data from 15+ were pooled to raise the expected frequencies. were found in pooled cells as Niwot had fewer pigeons re- than Forest Heights. 1979 were reobserved Band-tailed Generally, in 1980 at Forest more bandtails marked in Heights than at Niwot. Pigeon Sex and Age Ratios Braun et ale (1975) documented changes in sex ratios of pigeons according trapped adults to time of day. Males comprised about 68%of the adults prior to 1000 MDT, females comprised about 65%of the trapped captured transition after between 1600. periods. and subadults) 1000 arid 1600, and equal proportions I further During trapped trapped after between Pigeons trapped bandtails captured during males (adults before periods before 1001-1400, and 25%of the Females comprised approxi0830, 80%of the pigeons after 1530. were more than 70%females. Heights, males comprised 75%of the 0830, 30%of the birds 1001-1400, while no birds were trapped after Females comprised about 25%of the bandtails and 70%of the birds before the morning (0830-1000) and afternoon 1980 at Forest caught Heights, 1001-1400, and 75%of those caught (1401-1530) transition During between 1530 (Table 13). mately 20%of the birds trapped 1979 at Forest the day to separate comprised about 80%of the pigeons trapped 0830, 20%of the birds bandtails subdivided were caught between trapped between 1530 (Table 14). trapped 1001-1400. before Forty-five 0830, percent �141 . 10 NIWOT Q 8 FOREST ~ HEIGHTS ~ 1&1 e e s s e ...J DN=28 ~ (Tagged )=417 ~ = 30 ~ (Togged) = 246 6 •••• o 4 I- Z I&J U Q: IIJ 0.. 1-5 NUMBER OF DAYS REOBSERVED Fig. 28. Reobservations in 1980 of bandtails Niwot and Forest Heights. tagged in 1979 at �Table 13. Sex and age of band-tailed < 0810·· pigeons from trap 0830-1000 N % samples by time of day and site, Time of traEEing 1001-1400, N % 1401-1530 % 1979. > 1530 ' % N - % Forest Heights Males Adults Subadults 61 19 62.9 19.6 5 1 22.7 4.6 13 10 11.3 8.7 12 2 24.0 4.0 6 25.0 Subtotals 80 82.5 6 27.3 23 20.0 14 28.0 6 25.0 6 11 6.2 11. 3 15 1 68.1 4.6 70 22 60.9 19.1 26 10 "52.0 20.0 18 75.0 Subtotals 17 17.5 16 72.7 92 80.0 36 72.0 18 75.0 Totals 97 100.0 22 100.0 115 100.0 50 100.0 24 100.0 106 25 63.9 15.0 29 8 27.6 7.6 104 17 27.7 4.6 5 2 13.5 5.4 131 78.9 37 35.2 121 32.3 7 18.9 19 16 11.4 9.7 47 21 44.8 20.0 217 37 57.9 9.8 26 4 70.3 10,8 Subtotals 35 21.1 68 64.8 254 67.7 30 81.1 Totals 166 100.0 105 100.0 375 100.0 37 100;0 Females Adults Subadults Niwot Males Adults Subadults Subtotals Females Adults Subadults - - N - N ~ N �143 of the pigeons caught trapped during during the afternoon and 80% the morning transition transition Table 14. Sex and age of band-tailed time of day and site. 1989. were females. pigeons from trap samples by Time of traEEin g < 0830 N Forest 90 0830-1000 N % N 1001-1400 % 1401-1530 9- N 0 Heights Males Adults Subadults Subtotals 10 2 62.5 12.5 26 6- 44.8 10.4 17 4 24.6 .5.8 2 3 7.4 11.1 12 75.0 32 55.2 21 30.4 5 18.5 1 3 6.2 18.8 19 7 32.8 12.0 41 7 59.4 10.2 16 6 59.3 4 25.0 26 44.8 48 69.6 22 81.5 16 100.0 58 100.0 69 100.0 27 100.0 8 88.9 23 2 37.7 3.3 2 1 13.3 6.7 8 88.9 25 41.0 3 20.0 1 11.1 34 2 55.7 3.3 9 3 60.0 20.0 Females Adults Subadults Subtotals Totals 22.2 Niwot Males Adults Subadults Subtotals' Females Adults Subadults Subtotals Totals 1 11.1 36 59.0 12 80.0 9 100.0 61 100.0 15 100.0 About 80% of the bandtails Niwot were males (Table 13). between captured Thirty before percent 1001 and 1400, and 20% trapped after 0830 during of the birds 1979 at trapped 1530 were males. �144 During the morning transition were females. No birds tion.. about 65% of the pigeons caught were trapped during the afternoon transi- In 1980 almost 90% of the pigeons caught before 0830 and 40% trapped trapped between after 1001 and 1400 were males. 1530 or during No birds the morning transition. percent -of the pigeons captured were Eighty during the afternoon transition were females. Sex ratios of unmarked pigeons were constant interval 1979 from spring (Tables 15, served for each daily to fall at both Niwot and Forest Heights during 16). Before 0830. about 80% of the bandtails ob- were males. while from 1001 to 1400 and 1401 to 1530 about 70% of the birds seen during variable, observed were females. the morning transition but the summer average Sex ratios of pigeons and after 1530 were most for both periods at both sites was about 50% male - 50% female. Sex ratios of unmarked bandtails each daily interval Heights (Tables 17. from spring 18). in 1980 were constant to fall at both Niwot and Forest Before 0830; nearly 80%· of the birds seen were males, while from 1001 to 1400 and 1401 70% of the plgeoIis observed were females. served after 15~Oj oUt sex ratios were again quite variable. for to 1530 about No prgeone Wei'e ob= dUrlhg the th6rfiiiig trart§ition The ratio during the morning transition at Forest Heights for the summer was 50%male - 50%female, but at Niwot over 60% of the pigeons observed were females. �Table 15. Band-tailed pigeons observed dai~y by sex and time of day, 1979. Niwot Time of observation Month Niwot May Sex Males Jul Aug Females 29 417 88 Males 323 83.2 Females 65 115 43 16.8 72.8 909 225 1,134 80.2 Males Males Females May-Aug 54 65.1 34.9 82.6 17.4 Females Jun < OS-30 N % Males Females Totals 27.2 19.8 100.0 -0830-TOO-0' 1001-1400 N - % N 39 26 83 60.0 40.0 46.4 53.6 96 42 42 116 50.0 50.0 79 280 59.5 40.5 53.5 243 523 223 569 224 631 294 814 269 702 > 1530 1401-1530 % N - % 29.1 70.9 26.2 73.8 64 138 16 53 31.7 68.3 23.2 76.8 26.5 73.5 27.7 65 26.7 178 40 73.3 172 162 30.1 71 69.9 28.6 98 287 71.4 100.0 337 624 72.3 27.3 46.5 1,020 2,716 72.7 93 185 462 100.0 3,736 100.0 647 N % 44 36.4 63.6 51.5 77 48.5 42.0 58.0 46.0 54.0 100.0 •.... .l:- V! �Table 16. Band-tailed pigeons observed daily by sex and time of day, Forest Heights Month Forest Heights May Sex < 0830 N % Males Females 51 20 71.8 28.2 Jun Males 86.5 Jul Females Males 90 14 Aug Females Males Females May-Aug Males Females Totals 116 23 56 20 13.5 83.5 0830-1000 N % 57 36 155 70 61.3 38.7 68.9 31.1 16.5 73.7 29 50.0 77 19.7 29 241 135 50.0 313 26.3 80.3 390 100.0 376 64.1 35.9 100.0 Time of observation 1001-1400 1401-1530 N N % % 81 180 113 225 31.0 69.0 33.4 66.6 28.0 89 229 72.0 41 87 32.0 68.0 324 721 1,045 31.0 69.0 100.0 10 24 51 144 27 79 1 5 89 252 341 1979. > 1530 N % 53 77.9 22.1 29.4 70.6 26.2 73.8 25.5 74.5 16.7 83.3 26.1 73.9 100.0 15 26 53 79 68 147 32.9 67.1 53.7 '46.3 100.0 ,_. .p.. 0\ �147 Table 17. Band-tailed pigeons observed day, Niwot 1980. < 0830 Month Sex % N daily :by sex and time of Time of observation 0830-1000 1001-1400 N N % % 1401-.1530 0 N 1) May Males Females 31 15 67.4 32.6 8 20 28.6 71.4 48 136 26.1 73.9 32 58 35.6 64.4 Jun Males Females 2 100.0 9 10 47.4 52.6 37 38 49.3 50~7 27 42 39.1 60.9 Jul Males Females 30 5 85.7 14.3 7 10 41,.2 ·58.8 156 489 24.2 75.8 87 242 26.4 73.6 Aug Males Females 26 14 65.0 35.0 13 24 35.1 64.9 54 129 29.5 70.5 36 91 28.3 71.7 Males Females 89 34 72.4 27.6 37 64 36.6 63.4 295 792 27.1 72.9 182 .433 . 29.6 70.4 Totals 123 100.0 101 100.0 1,087 100.0 615 100.0 MayAug Table 18. Band-tailed pigeons observed day. Forest Heights 1980. Month Sex < 0830 N % Time of observation 0830-1000 1001-1400 N N % % 8 21 May Males Females 48 11 81.4 18.6 Jun Males Females 21 4 84.0 16.0 Ju1 Males Females 109 15 87.9 12.1 22 12 Aug Males Females 20 2 90.9 .9.1 Males Females 198 32 Totals 230 MayAug daily by sex and time of 27.6 72.4 17 45 27.4 72.6 16 27 37.2 62.8 64.7 35.3 50 139 8 5 61.5 38.5 86.1 13.9 38 38 100.0 76 1401-1530 N % 7 8 46.•7 53.3 26.5 73.5 18 58 23.7 76.3 6 9 40.0 60.0 14 20 41.2 58.8 50.0 50.0 89 220 28.8 71.2 39 86 31.2 68.8 100.0 309 100.0 125 100.0 �Table 19. Sex and age of tagged band-tailed pigeons observed daily at Niwot, 1979. Time of observation -N- < 0830 0830-1000 1001-1400 % N 100 34.1 45 21.8 39 13.3 15 Adults 91 31.1 Subadults 41 May-Aug 1401':'1530 % % N 268 23.0 132 7.3 83 7.1 108 52.4 625 14.0 32 15.5 3 l.0 1 19 6.5 293 100.0 > 1530 N % 20.6 20 22.5 48 7.5 10 11.2 53.6 328 51.2 44 49.4 141 12.1 93 14.5 9 10.1 0.5 7 0.6 6 0.9 5 2.5 42 3.6 34 5.3 6 6.8 206 100.0 1,166 100.0 641 100.0 89 100.0 "6 N Males Adults Sub adults Females Immatures Males Females Unknown Totals ~ 00 �Table 20. Sex and age of tagged < 0830 May-Aug band-tailed pigeons 0830-1000 N % observed daily at Forest Time of observation 1001-1400 N % 'N 1401-1530 % 1979. > 1530 N % 11. 5 1 16.7 1 3.9 2 33.3 58.1 16 61. 5 1 16.6 12.9 5 19.2 1 3.9 1 16.7 1 16.7 6 100.0 N % 11 44.0 4 18.2 14 22.6 3 2 8.0 1 4.5 4 6.4 Adults 8 32.0 15 68.2 36 Sub adults 4 16.0 2 9.1 8 - Heights, Males, Adults Sub adults Females t-' Immatures Males Females Unknown Totals , 25 100.0 22 100.0 62 100.0 26 100.0 ~ ",. �150 Sex and age ratios of tagged during feeding sessions 0830. served During were females. bandtails From 0830 through were females, while after observed seen during 50%male - 50%female 1530 MDT, over 65% After 1530 about 60%were the sex ratio of tagged before 1530 only 6 tagged 0830 were males (Table 21). 100 birds} after pigeons were Heights in 1980, 74%of the the morning transition tails were observed 50%male - 50%female 1530 over 70%of the pigeons ob- At Forest of the pigeons observed Forest Heights, seen before 0830 approached seen <IV 20%females}. {IV 1979 at 1980 at Niwot, too few tagged pigeons were ob- 1979 at Forest (Table 20). birds During (N = 86) to compute proportions. During served was nearly From 0830 through of the pigeons observed females. pigeons observed were also documented. Niwot, the sex ratio observed (Table 19) before band-tailed Only 20% were females and about 40%· from 1001 to 1530 were males. 1530. No band- However, only small samples of tagged pigeons were observed each year at Heights. Table 21. Sex and age of tagged band-tailed daily at Forest Heights, 1980. < 0830 May-Aug Males Adults Subadults Females Adults Sub adults Totals N % pigeons observed Time of observation 0830-1000 1001-1400 N N % % 19 4 61.3 12.9 7 1 70.0 10.0 14 4 31. 8 9.1 8 25.8 2 20.0 20 6 45.5 13.6 44 100.0 31 100.0 10 100.0 1401-1530 N % 7 36.8 9 3 47.4 15.8 19 100.0 �151 Ratios of adults period. and immatures were estimated Immatures were easily distinguished pale gray appearance 1978). The first from adults and lack of a neck crescent immatures were observed 18 June and at Forest Heights by their (White and Braun in 1979 at Niwot on on 19 June. immatures were seen at Forest for each feeding During 1980, the first Heights on 4 June and at Niwot on 30 June. Average during daily proportions 2-week intervals and 1980 at Niwot (Fig. served of immature bandtails , in feeding were similar (95% CI, P > 0.05) for 1979 29). sessions Since the number of immatures ob- was quite variable were small, the 95%confidence Generally, immature proportions to approximately 11%during The proportions increased the first were different (95%CI, P < 1980 at Forest Heights (Fig. o. 05) 30). (95% CI, P < 0.05) proportion during the first 2 weeks of August. June 1979 and fewer (95% CI, 1980 than in 1979, a Higher proportions similar numbers increased July and early August. tion s in June were zero but increased July slightly sessions the last 2 weeks of July in 1980 (95% CI, In 1979, immature proportions during in feeding of immatures was observed 2 weeks of July in 1980. early June to '\. 7% during sessions from zero in early June Although higher though were wide. durin g July between ~ > 0.05) immatures were seen during P > 0.05). and sample sizes limits for both years of immatures observed were also seen during observed to nearly from zero in In 1980, propor- 30%in July. of immatures were present in feeding 1979 and 1980, fewer (95% CI, P < 0.05) AI- �152 0.21 1.00 t--I •• •• 95CYo cr A MEAN •• •• -1979 --1980 0.12 z Q •••• 0: 0 e, 0.10 0 0: e, 0.08 L&J a: :J •••• c:t ~ .,_ 0.06 1 .iii! 0.04 0.02 ::::> ~ len V_ Fig. 29. observed ••• •• ••• Z...J ...J ...J(!) C) ~ ~ 0 ~ ~« « 0.00 Z , •• I I ::l::l Nit) ::::> 1(1) V_ ::l ::l=> ~N II'- It)_ Average daily proportions of immature band+tailed during 2-week intervals at Niwot. 1979.... 80. pigeons �153 --.-• 0.83 0.50 ·0.40 0.30 0.20 t-t 95.,. e cr • MEAN r ., -1979 --1980 ,. ,: , I I , z I 0 e, •• ""* 'OT7 . 0.14 0 0.17 I·~ 0.16 •••• II: ••• I , 0.12 I 0 a: e, I I LIJ I I II: ~ l- I c( ~ I ::E I I I I I I I 0.04 I I 0.02 0.00 Z ;::) -, len v_ Fig. 30. observed • Z..J ..., ..., ;::);::) 0 Nrt'> •• •• • • ..J ..., ;::) leo v_ •• • 1(.') (.') -1;::) ;::) c:( ~<t I ~N •••••• rtl_ Average daily proportions of immature band-tailed pigeons during 2-week intervals at Forest Heights, 1979-80. �154 total birds were present. in immature proportions arrival This caused the apparent observed between years. to feeding sites was not constant at either site in either ·24 different total, by age from spring pigeons observed site was also ascertained. pigeons tagged both Conifer Timing of pigeon to fall year. The age and sex of band-tailed other than the banding difference During 1979, at Niwot were observed (13) or Forest Heights at a location at either (15) (Table 22). or Of this 61%were males, 32%were females, and 7%were immatures. Seventy-nine at either bandtails tagged at Forest Heights were observed or both Conifer (71) or Niwot (11). In this sample, 45%were males, 45%were females, and 10%were immatures. high interchange low interchange of distance During either between Forest Heights and Conifer. between Forest between sites (Kautz 1977). or both Conifer (3) and Forest were reobserved Thirty-five Heights at Conifer (32) or Niwot (3). (3) (Table 23), 2 at Forest 63% between Conifer . = 0.55, Heights Of this total, The interchange Heights was similar (X between years. at Niwot were seen. at pigeons tagged were females and 37%were males. and Forest and the Heights and Niwot was a function 1980, only 5 pigeons tagged and all were females. The 0.40 < P < 0.50) �Table 1979. 22. Age and sex of band tails N - tagged % of total tagged in 1979 observed at a location Age and sex other than the banding (%) AHY HY AHY 5Y 35.7 17.9 7.1 21. 4 10.8 in Sex unknown Females Males SY site HY --m Banded at Niwot Total tagged 417 a reobserved at = Forest Heights Conifer 15 3.6 13 3.1 7.1 .- 32.2% Female 60. 7% ~ale V1 V1 Banded at Forest Heights Total tagged ~ 246 reobserved at Conifer 71 28.9 Niwot 11 4.5 aSome tagged pigeons were seen at both 26.8 17.1 45.1% Male sites. 1.2 31. 7 12.2 45. 1% Female 1.2 9.8 �156 Table 23. Age and sex of bandtails tagged in 1980 observed location other than the banding site in 1980. N % of total tagged at a Age and sex (%) Males Females AHY SY AHY SY Banded at Niwot Total tagged =a49 Reobserved at Forest 3 6.1 3 6.1 32 27.8 3 2.6 Heights Conifer 80.0 20.0 100%Female Banded at Forest Heights Total tagged = 115 a Reobserved at Conifer Niwot a Some tagged 28.6 8.6 40.0 37.2% Male 22.8 62.8% Female pigeons were seen at both sites. Crop Gland Activity Crop gland activity palpation of the crop lining were classified lated of bandtails as either (Zeigler active trapped was determined 1971, Fitzhugh (thick, spongy (moderate crop lining development), 1974). or inactive "cr-op milk", 1971) used to feed squabs. During may be used as an indicator 1979, an increase higher percentage than adult females. (Table 24). material glandular activity. in the number of crops with glandular development was found at both Niwot and Forest summer progressed Thus, of breeding stimu- (no crop The crop glands slough off cheesey development Crops crop lining), lining development). (Zeigler by Heights as the During mid-summer, a slightly of adult males had active or stimulated Except for the trap attempts crops at Forest �24. Table Crop Siteb Date gland activitya of band-tailed AHY S A N pigeons N N % in 1979 trap samples. I % N - SY S A % N - % N - I % N % 56.2 1 5 4 6 11 20 6 4 100.0 100.0 100.0 100.0 57.9 83.3 85.7 25.0 14.3 6.7 23.1 25.0 18.2 37.9 1 11 6 6 14 10 12 15 17 100.0 100.0 100.0 85.7 93.3 76.9 75.0 68~2 58.6 Males 14 15 22 4 5 25 26 27 28 May May May Jun Jun Jun Jun Jul Jul F N N F N F N F N 6 34 17 15 41 60 76 25 87 1 6 2 2 12 8 15 18 6 24 14.6 3.8 11.1 16.9 '3.0 22.9 36.6 34.6 33.3 33.8 6 32 12 11 27 20 32 10 35 100.0 97.0 100.0 100.0 77.1 48.8 61.5 55.6 49.3 1 5.3 1 3 14.3 18~8 7 4 9 36.8 16.7 Females 14 15 22 4 5 25 26 27 28 May May May Jun Jun Jun Jun Jul Jul aA bF F N N F N F N F N = Active, = Forest 10 45 29. 56 77, 36 74 41 89 S = 1.7 1 10 16.6 Stimulated, Heights, 3 1 1 11 7 18 6 25 N = I = Niwot. 8.8 4.3 2.0 17.7 30.4 31.0 31.6 41.7 Inactive. 9 31 22 48 51 16 39 13 25 100.0 91.2 95.7 98.0 82.3 69.6 67.2 68.4 41.7 3 1 13.6 3.5 1 1 3 4 4 11 I-' VI --..J �158 Heights on 25 June and Niwot on 28 July, sub adult males had fewer active crops than sub adult females. In 1980, fewer pigeons were trapped gland activity trapped (Table 25). Despite small sample sizes, in May had inactive to the results and examined for crop crops, obtained in 1979. 2 similar (X = most pigeons 1. OS, 0.30 < P < 0.40) However, during early June 1980, 45%of the adult males and 31%of the adult females had active or stimulated crops while 43%of the sub adult males and 24%of the sub- adult females showed signs of crop gland activity. June 1979, no pigeons trapped During early had active or stimulated appeared that the nesting aeason started intensity in 1980 than in 1979. earlier crops. It and with greater Time Budgets Preliminary artificial data concerning bait sites were recorded sample stratification, Forest resting band-tailed in 1979. pigeon behavior Due to problems with the 1979 data may be biased. Heights usually spent at Pigeons at more than 60%of the time at the site· (Table 26) and about 3%of the total time feeding. comprised most of the maintenance category and disturbance Preening comprised most of the other category. Band-tailed pigeon time budgets examine behavioral tagged differences males, nontagged and immatures. were collected in 1980 to between 5 categories males, tagged A stratified of birds females, nontagged sample of focal individuals -females , was used. Pigeons in all 4 adult categories spent over 70%of their time in resting (Table 27) with no differences and maintenance activity �25. Crop Table gland activitya of band-tailed pigeons in 1980 trap samples. AHY Siteb Date N % N SY 5 A N - % 5 A I N - % 6 26 4 27 1 100.0 96.3 100.0 55.1 100.0 15 29 6 43 100.0 100.0 100.0 69.4 71.4 N - % N - I % N % Males 19 20 24 8 5 F N N F N May May May Jun Sep 7 29 4 63 2 14.3 7 1 3.7 15 30.6 1 6 1 42.9 100.0 2 100.0 100.0 8 57.1 1 100.0 2 100.0 76.2 33.3 Females 19 May 20 May 24 May 16 29 8 83 10 F N N F N 8 Jun 5 Sep 2 17 2 3.2 aA = Active, bF = Forest S = Stimulated, Heights, N = I = Inactive. Niwot. 27.4 28.6 5 5 2 23.8 66.7 16 1 - VI \0 �Table 26. Band-tailed 0600-0859 N=139 % Summer totals No observation pigeon time budgets, a Forest Heights 0900-1159 N=194 % 1979. 1200-1459 N-144 % 1500-1759 N-73 % - Daily average % (N=550) 27 19.4 22 11. 3 14 9.8 13 17.8 13.8 Feeding 5 3.6 5 2.6 5 3.5 2 2.7. 3.1 Resting 81 58.3 109 56.2 99 68.6 47 64.4 61.1 Maintenance 20 14.4 44 22.7 19 13.2 10 13.7 16.9 6 4.3 14 7.2 7 4.9 1 1.4 5.1 Other aThe no observation category includes were present and none was visible. times when no birds were present and times when few birds I-' ~ 0 �161 (P > 0.05) between groups approximately 60%of their (I-way ANOVA). time resting Immatures or preening. (P > 0.05) from any other category. spent not different No differences 0.05) (po> were found between any of the 5 groups in no observation. ship. and disturbance Table 27. Band-tailed activities (I-way ANOVA). pigeon time budgets. Forest Heights Percent of time Tagged Nontagged Tagged Norrtag ged male male female female (N 12)b (N 41) (N = 61) (N 9) = Activitya court- = = 1980. lIinnatures (N 16) = 14.5 18.9 13.2 9.1 23.0 Feeding 5.0 8.5 4.2 4.0 10.4c Resting 46.9 49.8 52.6 59.1 47.4 Maintenance 33.2 22.1 21. 4 8.5d 26.1 15.0 No observation Courtship Disturbance 0.4 Other 0.6 0.1 0.1 0.1 Sample size f 120 410 90 O.b 0.3 1.0 3. ge 610 160 aDifferences °between age and sex classes for each behavior category were tested with a I-way ANOVA at a 0.05. All significant F-tests were further analyzed with Tukey's Q-test. = bNumber of 10-minute observation CSignificant from all but tagged periods. female at a = O. OS, Tukey's Q- test. dData biased by 1 tagged female which formed a pair bond with a tagged male and spent a great deal of time allop reenin g , eSignificant Q-test. from only nontagged male at a fTotal number of minutes observed. = 0.05, Tukey's �162 The amount of time spent feeding did not differ (Tukey's .Q-test, .P > 0.05) between any of the 4 adult categories. Immatures spent more (P < 0.05) time feeding than all adult groups except tag ged females. All observations birds picking in the other category up grit along the roadside immatures begging for food. activities Immatures spent than nontagged Mean durations bandtails for the behavioral length bouts During by Q-test, males or females vs. for each of the 5 groups between examined. activity. No observation resting, in the and mainte- (Table, 28) • 1979 and 1989, band-tailed were noted at both. Niwot and Forest pigeon courtship Heights. interactions Data for both sites in 1979 were biased because it was not possible served courtship An effort served displays Forest Heights sample in 1980 was large enough for analysis. At Forest (Table 29). groups bouts were There was little difference for feeding, of groups. I-way ANOVA) were found between of time of each observation nance activities (Tukey's to examine differences activities longer than any other disturbed males. (P > O. OS, No differences -- more (P < 0.05) time on other of activity were calculated or birds No differences P > 0.05) were found between nontagged immatures. were in 2 groups displays. to record all ob- was made to record all ob- and their outcome at both sites in 1980. Heights in 1980, 59 males were observed Thirty-four (57. 6%) males directed females. while 12 (20.4%) cooed when there Only the displaying coos towards was no other bird �163 Table 28. Band-tailed Heights 1980. ragged male Activity pigeon mean duration of activity bouts, a Mean duration Nontagged Tagged . Nontagged male female female Forest Immatures No observation 5.9 5.2 5.9 4.3 5.3 Feeding 0.9 2.7 1.9 1.7 4.0 Resting 2.0 1.6 1.3 1.7 1.8 Maintenance 1.3 1.2 1.2 1.0 1.3 0.1 1.2b 0.3 0~3 0.1 0~3 0.3 3.1 1.4 Courtship Disturbance 0.1 Other 0.1 Sample size C 39 130 20 48 152 a1n minutes. spent bData biased by a tagged female which formed a pair bond and a large amount of time allopreening • . cTotal number of observations, Table 29. all categories. Band-tailed- pigeon courtship interactions, Forest Heights 1980. Category Total number of birds N observed Males Displayed to female, cooed Displayed to female, no cooing Cooed, no other birds nearby 34 13 12 57.6 Females Pecked at male Few to another limb or tree Slapped male with wing Accepted males' courtship advances 15 15 6 1 40.5 40.5 16.3 % 22.0 20.4 2.7 �164 within 10 m of their perch. displaying to females, usually coo was heard. displaying (22.0%) males were observed within 50 m of my vehicle, This sample indicated males either It was possible displayed that more than even though I was less than to record the reactions or wing-slaps). Fifteen other or tree apparently of 37 of 47 female Twenty-one towards the males (pecking females (40.5%) flew to another to avoid the male's courtship displays. (2. 7%) female formed a pair bond with a displaying a bout of allopreening. female-directed formation at artificial intensity This suggested male courtship that the 50 m away. to by the 59 males (Table 29). (56.8%) of the females showed aggression yet no 20%of the did not coo, or cooed so softly, coo could not be heard bandtails Thirteen bait sites despite resulted Only 1 male followed by that less than behaviors limb 3%of the in pair bond the high level of cooing by males. Band Recoveries and Returns Of the 778 pigeons newly banded in 1979, 14 were recaptured during 1980 (Table 30). were retrapped tagged tured in 1980. Overall, there season, controls percent samples than of those pigeons tagged Niwot or Evergreen, and reported was a greater (Table 31). 1 each in Colorado, 22 February during at Niwot Only 3 (1.1%) pigeons Heights and none o,f the banded From 1 May 1979 through at either (1. 4%) tagged along with 5 (6.9%) controls. at Forest (6.3) in trap Six bandtails was recap- of the controls (1. 2) . 1981, 5 bandtails banded 1979 or 1980 were recovered Three were shot during the 1979 hunting New Mexico, and Arizona. The other �165 2 were found dead (1 at a raptor of controls banded during Table 30. samples. nest) in 1979. No recoveries 1979 or 1980 have been reported. Control and tagged bird returns from 1979 in 1980 trap 1979 No. tagged No. controls banded Tagged returns Niwot 417 72 6 Forest Heights 281 8 3 698 80 9 Site Totals 1980 Control returns Percent Control Tagged returns returns Sa 1.4 6.9 1.1 S- 1.3 6~3 a Two of S (40.0%) Niwot controls recaptured were at Forest Heights on 19 May and 8 June. One Niwot control was recaptured twice but counted as a single return. Too few bands have been reported about the survival hypothesized of color-marked that survival vs. for meaningful conclusions leg-banded of color-marked pigeons. bandtails (P < O.OS) than those marked only with leg bands. _ It is will be less �Table 31. Recoveries of band-tailed pigeons banded at Niwot and Evergreen, and reported from 1 May 1979 through 22 February 1981. Year banded Site a banded 795-21821 79 N 795-21923 79 N 795-21930 79 E 8 Sep 79 Shot 795-22302 79 N 12 Oct 79 Shot 795-22408 79 N ? Sep 79 Shot Band number Date recovered How obtained 1 Jun 79 Found dead 24 Sep 79 Found dead Where found Tolland, Colorado Jamestown, Taos, Colorado New Mexico Flagstaff, Arizona Colorado and recovered Age and sex at banding Status AHY-Male 639-YAR AHY-Female 639-0BO AHY-Female 639-BA3 AHY-Male 639-BDV SY-Male 639-RZN b ,_. aN = b639 Niwot, E = = Evergreen. Experimental bird, wing-tagged. Colorado City, Colorado 0"0"- �167 DISCUSSION Trapping The reduction in percent numbers of trapping due to decreased or restriction recaptures in 1979 with increasing attempts at Forest Heights may have been site attendance. switching to natural of home range after nest establishment. pigeons recaptured during the single trapping were tagged at Forest Heights. food sources. Two of 5 effort at Conifer During both 1979 and 1980. '" 28% of the pigeons tagged at Forest Heights were reobserved once at Conifer (Tables 22. 23). Interchange at least between Forest Heights and Conifer probably varied between 25 and 40%. Reports of tagged pigeons also indicated greater reduction at Forest Heights than at Niwot in 1979. Far fewer reports tagged birds were received from observers though more pigeons were color-marked) area. However. there were apparently sites available near Niwot. since most reports in site attendance of in the Niwot area (al- than in the Evergreen fewer alternate Switching to natural feeding foods was unlikely of pigeons away from the study sites occurred at other baited areas (i. e. birdfeeders). No data were available concerning changes in size of home range after nest establishment. During 1980, decreased site attendance at Niwot than at Forest Heights. was apparently During 1979, a rolled corn-barley mixture was available at Niwot from early June throughout summer. greater the In 1980, cracked or ground corn and barley were stored �168 in separate piles in the pit from early June through Bandtails apparently preferred September. rolled corn to cracked or ground corn since fewer pigeons were. observed at Niwot in 1980. No native foods were known to be available near Niwot that may have caused food switching. pigeons were trapped Percent Due to decreased site attendance. fewer at Niwot than at Forest Heights in 1980. recaptures decreased attempt at both sites in 1980. with each successive The percent trapping of tagged recaptures was also lower in 1980 trap samples at both sites than the levels observed at the end of the 1979 trapping pigeons were tagged. probably greater apparently decreased season even though more Although decreased site attendance was at Niwot in 1980. similar levels of attendance occurred at Forest Heights between years. site attendance. mortality of tagged pigeons may have caused the lower percent Along with {Table 31} of tagged recaptures in 1980 trap samples. Daily Site Attendance Daily peaks in the numbers of pigeons pr-eserit usually occurred between 1000 and 1500 MDT at all study sites. A slight increase in pigeon numbers was noted before 0800 due to the presence presumably transition mated male pigeons (Peeters 1962). (change in sex ratios), noticeably between 0800 and 0900. through During the morning pigeon numbers decreased Numbers usually increased late morning and early afternoon as small groups females) arrived at the site. of Site attendance decreased (mostly between �169 1400 and 1500 as females departed. Most pigeons left in one large group and less than 10%of the afternoon peak remained at the site after 1600. Passmore (1977) found most pigeons that visited mineral springs in Oregon each day initially arrived few afternoon observations were made. before This was not true at artificial bait sites in Colorado as several bandtails observed each day between 1200 and 15-00. numbers was more likely to occur after Differences (95%CI, P < 0.05) at Niwot between 1979 and 1980. apparent 1200, therefore were initially The daily peak in 1200 than before. in site attendance were recorded These changes were caused by site avoidance and possibly mortality of tagged pigeons. The value of evening counts at bait sites was limited. sessions attracted tagged birds less than 20 pigeons after arrived Most feeding 1600 MDT and few at the site which had not been observed earlier the same day. Seasonal Site Attendance Peaks in pigeon numbers occurred sizeable day-to-day during the first variation. during early June, Numbers decreased 2 weeks of August. may have been caused by ripening at all sites This reduction of natural with in attendance foods which attracted pigeons from the bait sites for short periods of time. Kautz (1977) noticed evidence of both grain and native foods in band-tailed pigeon fecal material on his study have been caused by the start areas. The decrease of southward migration. may also Fewer �170 pigeons marked in May sites during (!_ test, P < 0.05) were observed late July and early August At mineral sites in Oregon, {Table 12}. Passmore (1977) observed than 10 pigeons per day prior to mid-June. these sites occurred attendance during recorded of grainfields different late August. Seasonal patterns of sites were in early August. Peak use in Colorado may 'occur from mid-summer to fall, from that observed at my study Fewer pigeons were observed At Forest at from those at Peaks in numbers at study in early June with decreases fewer Peaks in numbers at bait sites in Colorado were different mineral sites in Oregon. at the Heights, more bandtails sites. in 1980 than in 1979 at Niwot. were observed summer with a decrease in attendance August. may have been caused by site avoidance This decrease following trapping At Conifer, of 146 birds numbers increased numbers occurred Large day to day fluctuations throughout during pigeons apparently the summer. was observed used alternative Bandtails During at Forest food sources the site may have been 1980, peaks in later in the Heights, during and mid-summer. Estimates Kautz (1977) discussed resulting and were noted but early summer with decreases The same pattern Jolly-Seber July. on 8 June. to the amount of grain provided. summer. late June, 1979 was the first year pigeons visited in large numbers. adjusting during early in the the problems encountered bias from using the Jolly-Seber pigeon recapture data from Colorado. technique and the with band-tailed He concluded that: �171 1) Because tagged proportions decreased with distance from the trap site and variation occurred· in feeding flock location and size from year to year, tion of the Jolly-Seber technique that is difficult to satisfy with bandtails is that all birds, or unmarked, the assump- whether marked have the same probability of being captured (Seber 1973). 2)· Because recapture tagged proportions distance from the banding site, from trapping Jolly-Seber decrease with using banding information at 2 or more sites will result in biased. estimates and the amount and type of bias will be highly variable depending upon temporal variation in the use of the same sites. To use the Jolly-Seber estimation technique yearly variation in. population parameters as an index to of band-tailed pigeons, Kautz (1977) recommended that: 1) Trapping should be done at the same site (or sites less than 5 km apart) to prevent probability due to distance . of retraps a decrease in the .,:", 2) Flight patterns of pigeons using the trap site should be monitored each year to estimate the size of the range from which the population is being drawn. 3) Due to behavioral differences, immature populations should be estimated separately. 4) Trapping should be done thr-oughout the day to get �172 5) Trapping should be done only over a time period of a few days each year to prevent trap avoidance and to satisfy stantaneous At Evergreen, population's the birds from learning the assumption of in- sampling (Seber 1973). 14 different trap locations all within the sub- range were used between 1969 and 1980. Several of these sites were more than 5 km apart and there may have been a decrease in the probability trap sites. of recapture due to distances In most years traI?ping at both Evergreen and Niwot was done over a period of a few months so the bandtails learned trap avoidance. sampling was violated. Thus. between may have the assumption of instantaneous Flight patterns were not monitored at either site so an estimate of the range from which the population was drawn was not available. Due to large standard and survival. Jolly-Seber errors in estimates of total numbers estimates of population parameters only be used as indices of change in Colorado. The large increase in numbers at Niwot between 1978 and 1979 and at Evergreen 1977 and 1979 (Table 6), was caused by unrealistic bilities. When survival probability should survival estimates exceeded unity, between probapopulation estimates· were inflated. Survival probability estimates exceeded unity when the data set had 1 or more of the following characteristics: 1) The total marked and released at time i+1 greatly exceeded the number marked and released at time i , �173 2) The total number marked in the population prior to time !_ was 3) relatively small. The number of marked animals in the trap sample at time Kautz (pers. !_ was unusually large. commun.) believes the 3rd characteristic most important factor smce it may reflect capture for different unequal probability groups of tagged bandtails. earlier years were tagged primarily at different from more recent years at the same site, solved by capturing and releasing animals at each trapping of If pigeons from sites, and those pigeons from more recent years would have a higher probability (assuming both groups are alive). to be the of being recaptured All 3 problems could have been a given number of marked time, and trapping the same site each year. Few conclusions can be drawn from the trapping Jolly-Seber analysis for 1979 (Table 8). because pigeons captured cluded in the analysis most trap samples). at Forest Heights. data and Estimates were biased and marked prior to 1979 were not in- (even though they accounted for 20-30%of Site avoidance probably caused bias, especially This analysis with a small data set was of limited value and it is recommended that this technique be used only with large data sets from long-term studies. Relationship Between Cooing and Numbers Daily and seasonal pigeon cooing patterns site attendance, and August, were related especially during May and June. cooing intensity decreased to By late July and no relationship between �174 cooing and numbers was evident. The average number of coos heard per. hour was a good predictor pigeons present for the same period during and Forest Heights (Figs. period in 1980, patterns relationship predictor During the intensive were similar but the strength of pigeon numbers observer 1979 at both Niwot z-transformation, study of the P < 0.05), number of coos heard per hour was not a good collection during (Fig. 20). J. Ellis assisted with data 1980 at Forest Heights and despite training, bias may have accounted for some of the variability between years. ship differed 1980. 18, 19). was weaker (Fisher's and the average of the average number of At Niwot, it was not surprising that the relation- between years because few pigeons were present Due to conflicting results between years, in it cannot be con- cluded that the average number of coos heard per hour can reliably predict the average number of pigeons present per hour. Further study is required. Cooing intensity time of day. and pigeon numbers were also examined by During the morning (0600-0800), primarily mated males and unmated females were present During midday (1100-l400), intensity. present smaller percentage a greater for an equal increase This may have reflected coo less frequently 21). The slope of the regression for the morning period indicating in number of birds 1962) (Fig. the group was comprised mostly of mated females and unmated males. was greater (Peeters the hypothesis increase in level of cooing that mated males than unmated males (Sisson 1968) or that a of females was present during line the morning �175 period. Probably a combination .of these factors as peaks in calling intensity (coos/hour) was responsible usually occurred during midday. The relationship between the total number of coos heard day and the total number of birds during the intensive strong relationship = 0.854) (Fig. (::.2 Statistical Properties During detected size, Data were inconclusive used to predict was found at Forest as to- whether counts of band-tailed were made simultaneously during in 1980 at Niwot and Forest pigeons at Niwot or To increase at both sites, samplin g , counts a 2-week intensive Heights the sample study period (Table 11). the variability in the total number of coos heard per day was less than the variability the total number of pigeons counted per day. At Forest in 1980, a 20%change in the mean number of bandtails per day. calls size could not be reliably the assumption of instantaneous For both years at Niwot of Count Data Heights due to small sample sizes. yet satisfy with a sample size of 12 days, 95%confidence interval. in Heights counted would be detected at the A sample siZe::! of only 8 days would be ";.:; v required to detect heard cooing. A pigeon numbers. 1979, changes in population from direct Heights. was recorded 23) but a weak relationship (-!_2 0.189). can be reliably per day was examined period at Niwot and Forest between the 2 variables = Heights Forest study observed per a 20%change in the mean number of pigeons �176 It was not possible to establish whether or not a change in the number of coos heard could reliably predict of pigeons present. Thus, a change in the number counts of pigeons present heard should be made simultaneously at selected sites. It appeared once a sufficient changes In pigeon numbers at feeding data base is available. that changes in population size can be reliably of band-tailed was neither artificial bait that both direct counts and coo counts may be used to reliably' predict sites, and coos pigeons made at intervals accepted or rejected Tagged Proportions Thus, the hypothesis derected from counts through the breeding because of insufficient data. Observed Tagged proportions at Niwot and Forest of pigeons at feeding sessions were monitored Heights during 1979 and 1980. No differences (95%CI, P > O. OS) were found within sites between years.; 1979, higher (95%CI', P < 0.05) tagged proportions at Niwot during season 5 June through 3 July. During were observed No differences (95%CI, P > 0.05) were found between sites in 1980, although tagged proportions observed tagged proportions were usually higher at Niwot. were expected Higher at Niwot since more pigeons were tagged there in 1979 than at Forest Heights. Fewer (!_ test, p < 0.05) bandtails Niwot and Forest Heights were present 19 July through observed 1 September. differences. marked during May at both at feeding sessions Three hypotheses from may explain the First, some pigeons marked in May possibly arrived a few weeks earlier than the main group, nesting sooner, southward and started completed migration during the end �177 of July or early August. Second. effects of site avoidance may not have become apparent until late July and August. marked in May were involved in more trapping marked later in the summer. Pigeons attempts The cumulative effects than those of several trap attempts may have caused pigeons marked in May to seek alternative food sources by late summer. may have been time-dependent, summer. Bandtails of mortality until late marked in May were exposed to predation factors marked later in the summer. for longer periods nest near Jamestown, It is not known how many other tagged have been victims of avian predation. Colorado pigeons The hypothesis ratio of pigeons marked in each time interval the next time interval than The band of a tagged pigeon (Orange BO) was found at a raptor (Table 31). effects not becoming apparent and other population reducing birds Finally. may that the that are present does not change from spring in to fall was rejected. Tagged pigeons released 2 (X = at Niwot in 1979 were more likely 55.2, P < 0.005) to be reobserved 26). after marked at Forest Heights. (Fig. marked at Forest Heights were not observed tagging Over 40%of the bandtails after tagging only 18%were not seen at Niwot. Apparently occurred 1979 than at Niwot. at Forest no differences 2 (X Heights during = no difference greater 0.01, P > 0.90) in reobservability between Niwot and Forest Heights than those (Fig. 27). site avoidance During 1980, were recorded Apparently in avoidance between sites in 1980. while there was �178 2 (X Slight significance between sites for birds Generally, at Forest = 3.68, 0.05 < P < 0.10) was found marked in 1979 but observed more bandtails marked in 1979 were reobserved Heights than at Niwot. apparently greater in 1980 (Fig. 28). in 1980 Site avoidance or mortality was between years at Niwot. Sex and Age Ratios Braun et al. (1975) documented changes in sex ratios tailed pigeons in trap samples according to time of day. the day more finely to separate of the day. The results of both studies more likely to be trapped likely to be caught day more finely, during portions periods from other times indicated that males were midday or after:noon. percentages By dividing the of males and females were found of the day in this study. 75%of the pigeons observed observed I subdivided in the morning and females were more during higher transition of band- Passmore (1977) found before 0900 PDT were males and 70% between 0900 and 1200 were females at mineral sites in Oregon. No differences or Forest in sex ratios in trap samples were found at Niwot Heights within (X2 = 0.12, 2 (X = 1. 78, 0.10 < ~ < 0.20) years. between years, afternoon results 0.70 < P < 0.80) or between Although sample sizes differed were comparable . (1401-1530) transition periods Morning (0830-1000) and were most variable (Tables 13, 14). The hypothesis constant that sex ratios of unmarked bandtails for each daily interval both Niwot and Forest from spring were to fall was accepted Heights in 1979 and 1980. Transition at periods �179 were most variable from trap and results samples. obtained Males were more likely to be observed the morning and females during Sex ratios compared well with those of tagged the afternoon. pigeons observed during did not compare well with those of unmarked trap samples at Niwot or Forest During many time periods, Heights during tagged birds feeding sessions birds or those from 1979 or 1980. appeared site in groups of more random composition. to arrive by sex and age were available. of tagged hypothesis birds was altered from spring The hypothesis by color-marking. that timing of pigeon arrival unmarked bandtails = that the breeding Thus, 2.00, was accepted. August for each and departure by sex from spring until fall for There were no differences 0.10 < P < 0.20) in sex of birds May through the to fall was rejected. from feeding sites was constant 2 it is possible proportions that sex ratios of marked pigeons are constant daily interval (X Thus, at the However, it should be noted that only small sample sizes and unequal tagged status during observed daily from at Forest Heights or Niwot. Immature proportions in feeding sessions were quite variable and sample sizes were small at both Niwot and Forest Heights during 95%confidence 1979 and 1980 (Figs. limits were wide. 29, 30). At Niwot, immature proportions from early June through August tween years. Heights, proportion At Forest with no apparent a greater of immatures was observed July 1980 vs. Consequently, 1979. Therefore, during increased steadily differences (95% CIt !: < be- 0.05) feeding sessions at both sites in both years, in timing �180 of pigeon arrival spring until fall. at feeding sites was not constant Proportions by age from of immatures increased as the summer progressed. The age and sex of band-tailed pigeons observed other than banding sites were determined. between Forest Heights and Conifer. no consistent trends at locations Except for interchanges sample sizes were small and in age or sex of pigeons seen· at other sites were found between sites or between years. Interchange rates between sites were examined. 24%of the adult band-tailed pigeons recaptured 1970 and 1975 were caught '\, 8.4 km. 1979 and 28%for 1980. '\, 58 km, rate between these sites was 29%for between the Evergreen The estimates of interchange sites and Niwot was between these areas for from 3.1 to 6.1% (Tables 22,23), slightly than the 2%calculated by Kautz. During 1979, 3%of the pigeons banded at Niwot were observed at Forest Heights or Conifer. 6.1%. 9. Kautz 1977). These values are similar to the 24%calculated The distance 1979 and 1980 ranged higher 51-60 km away (Fig. between the Forest Heights and Conifer sites was The interchange by Kautz. in Colorado between from 1-10 km from the original banding site and about 2%were trapped The distance Approximately The opposite trend Heights to Niwot. occurred to 2.6%. to for movements from Forest During 1979, 4.5% of the pigeons banded at Forest Heights were observed decreased During 1980 this value increased at Niwot, while in 1980 this value These interchange rates also indicate that site �181 avoidance increased during 1980 at Niwot from that measured in 1979 although sample sizes were small. Crop Gland Activity A greater proportion or stimulated crops during period in 1979. of pigeons at Forest early June The late spring in 1979 as snow was recorded Heights had active 1980 as compared to the same weather was milder in 1980 than at Forest Heights on 30 May 1979. Although heavy snow was common in early May 1980. no snow was recorded at the site in late May. have started The nesting at least 2 weeks earlier in 1980. supported by the first immature being observed on 4 June 1980 compared to 19 June season appeared to This hypothesis was at Forest Heights 1979. Due to few pigeons feeding at Niwot in 1980. no trapping conducted during June or July. Therefore. data were available for comparison between months. 18 June However. the first was no crop gland activity 1979 and 1980 for these immature bandtail was observed on 1979 and 30 June 1980. Time Budgets More than 70%of the time for all categories immatures was spent in resting No differences or maintenance activity besides (Table 27). (I-way ANOVA. ~ > 0.05) among all 5 groups of pigeons were found for either time was allotted to resting necessarily of birds of these activities. and maintenance. had small sample sizes. biologically significant Since so much the other Several differences were not statistically significant. 5 classes that may be For example. �182 nontagged males spent nearly twice as much time feeding the other 3 adult categories significant (Tukey's ('" 4.4%), yet this difference Q-test, it was noteworthy among all 5 groups of bandtails. 96%of the time was spent in resting, for all classes. no differences These data. support (l-way This hypothesis Between 85 and maintenance and no observation the hypothesis that there birds since the and the percentage reobserved. Sample sizes were also small for the mean duration these data were a subset no differences among the 5 categories of the time budgets. this difference of bandtails. hypothesis activity, Generally, (l-way males ANOYA, P > 0.05) data were quite similar within compared among groups of pigeons. and nontagged Since greater feeding bouts 4 times longer than those for tagged that there were no differences between tagged Within a behavioral Even though was not significant due to small sample sizes. a behavioral data since (I-way ANOYA, P > 0.05) were found of immatures (4.0) lasted (0.94), of time seen per day was calculated based on the total number of pigeons observed activity, were between tagged and nontagged was an implicit assumption, estimated number of different of tagged birds how similar time ANOYA, P > 0.05) in the distribution spent among 6 behavioral categories pigeons. was not P > 0.05). Despite small sample sizes, was partitioned ('" 8.5%) as band-tailed The (P > 0.05) in behavior pigeons was supported. than 20%of the male bandtails observed dis- playing to females were not heard cooing (Table 29), any census technique relying on coo counts would underestimate the absolute �183 number of sexually active males. This assumption would hold unless females paired only with cooing males. breeding If this was true, segment of the male population attending sites would be biased towards noncooin g birds. data to either support the non- artificial bait I collected no or reject this hypothesis. Less than 3%of the observed females formed bonds with dis- playing males at Forest Heights in 1980 (Table 29). must assume that greater Therefore, than 90%of the females arriving at bait sites for the first time in early summer were already paired. formation must have occurred migration, during migration, before attending intensity one Pair either in Mexico before northward or at the nest territory a bait site. (Peeters 1962) Also, given the high rate of cooing at bait sites during 1979 and 1980, and the low acceptance rate of females, there must be unmated males in the population. Band Recoveries and Returns The differential return rate for banded controls tagged pigeons (1.3%) may reflect differences Bands from 5 tagged bandtails 1981 (Table 31). and Arizona), (6.3%) and in survival have been recovered (Table 30). as of 22 February Three were shot (1 each in New Mexico, Colorado, the band of 1 bird was found at a raptor 1 bird was found dead in Colorado. nest, and No controls banded in 1979 were recovered. Tagged birds appeared to have been more sus- ceptible to raptor predation controls. and hunting pressure but too few bands have been reported conclusions . than banded for definitive �184 Criticism of the Data Several factors that influenced the results during the study. Results of future ingful if these variables to attract bandtails studies would be more mean- could be controlled. that an unlimited food supply. were recognized It became apparent e. g.• at Niwot, was not required to an artificial bait site. In fact, too much food could reduce the accuracy of the estimate of the number of pigeons feeding at a given time. 600 individuals during a feeding session at Niwot than 125 during a session at Forest Heights. baiting was important. It was more difficult to count Also. the type of grain used for Bandtails appeared to prefer rolled or whole corn over cracked or ground corn. The influence of alternate study site was important. feeders) increase help of interested alternative feeding areas within 30 km of the Other artificial bait sites the probability observers of site avoidance. reporting (e. g. bird Without the color-marked pigeons, food sources would have been difficult to locate. the Wood family made counts of bandtails I was able to estimate t he .interchange frequenting Since their feeders. between Forest Heights and Conifer (nearly 30%). Many pigeons regularly observed at Conifer were not seen again at Forest Heights after initial trapping and tagging. Other bait sites in the immediate area also influence the amount of grain that must be supplied at the study site. appear to congregate Pigeon flocks at the closest food source that has ample grain to supply their needs. At Forest Heights, 5-6 kg of whole �185 corn per day was ample to feed a daily flock of over 200 pigeons. I would recommend this baiting level for establishing sites. future Also, I would try to locate the site in a relatively area so that the number of alternative bait unpopulated food sources would be reduced. Variability in the clumping of food sources must be considered when selecting an area for establishment The distribution of both natural Thus, bait site. and artificial foraging areas" the amount of mast or grain available. habits. of a permanent may affect flock feeding it would be best to locate permanent areas where the number of alternative and bait sites in food sources would be minimal. It may be argued that the numbers of pigeons at an artificial bait site indicate the amount of grain provided lation size of a given area. Evergreen study the site daily. Increasing site increased This relationship several alternate and not the popu- the amount of grain at the the size of the pigeon flock attending was confounded by the fact that food sources were available nearby. It would be important to reduce or eliminate these sources because a change in the amount of grain provided probably at one of the other sites would affect counts at the study site. If a given amount of grain is baited daily, and the flock is given time to find and adjust to this level of baiting. data from this study indicated in the mean number of different reliably predicted pigeons counted per day may be with a sample size of 12 days during 10 June at artificial bait sites. a 20%change 20 May - �186 It may also be argued that the land area from which pigeons are drawn to the bait site changes each year and that a difference in counts at bait sites indicates a change in land area, not population size. Approximately 95%of the band-tailed pigeon recaptures in Colorado occurred less than 60 km from the original banding site (Fig. 9, Kautz 1977). Thus it would be possible to delineate a 60 km radius circle around the bait site and be reasonably certain that the pigeons observed were drawn from this area. that a radio-telemetry I also feel nesting study would be valuable as the mean distance and 95%CI from nest site to bait site could be determined for a group of pigeons. This would provide a more accurate estimate of the area around the bait site from which bandtails were drawn, and should be attempted at permanent bait sites in conjunction with daily counts. The observability of pigeons IS when creating an artificial bait site. an important factor to consider For example. at Niwot, band- tails perched in cottonwood trees bordering Left Hand Creek.· Due to dense foliage. it was difficult to count flights of pigeons arriving at or leaving the site. .Accurate estimates of the number present could be made only during feeding sessions when most pigeons flew to the ground. At Forest Heights, ponderosa pine stand, or leaving the site. the birds perched in an open and it was easy to count groups arriving Estimates of the number of pigeons in the trees and the number observed during a feeding session usually were close. at �187 Coo counts may have been biased at Niwot because many pigeons at the site were able to perch more than 100 m from my observation point. The loudness of the coo and the distance it will carry varies between pigeons. been heard because they perched vehicle. Cooing pigeons may not have farther than 50 m from my At Forest Heights the opposite was true. Most birds perched within 50 m of my vehicle and I believe few pigeons that actually cooed while displaying were not heard. Finally. it is not desirable to trap pigeons at the artificial bait sites used for counts. This probably increased at Niwot and Forest Heights during 1979 and 1980. site avoidance �188 RECOMMENDATIONS Data from this study indicate that counts at artificial bait sites have potential as a census technique for the Interior of band-tailed pigeons. be established in Colorado. population Consequently permanent bait sites should Observations provide the long-term data required at these sites could to draw definitive conclusions about the value of either direct counts of pigeons or call counts at artificial bait sites. All factors discussed under Criticism of the Data should be evaluated when choosing and establishing a per- manent site. The Jolly-Seber mark-recapture analysis (Jolly 1965, Seber 1973) provides estimates of changes in population size at artificial 1:::·"1.1t sites, Due to variation, these estimates should only be used as indices of changes in population size. In future analyses, less variable estimates of population parameters may be obtained with the POPAN computer package (Arnason and Banruk A radio-telemetry nesting study, if initiated, 1980). would provide valuable information concerning the size of the area from which nesting bandtails are drawn to artificial bait sites. should be conducted at a permanent bait site, to ascertain in nesting The study along with counts, if changes in counts can be correlated with changes area size between years. Counts of pigeons and calls at permanent bait sites should be made during 20 May - 10 June. Data should be collected during �189 at least 14 days during this period to provide a large enough sample for statistical instantaneous analysis and yet satisfy the assumption of sampling. By this time, bandtail flocks should have completed northward migration and nesting should be well underway. The scarcity of natural foods (Braun 1973) causes pigeons to rely heavily on waste grain or grain provided at artificial food sources (i.e. bird feeders) during this period each year. Braun (1972) has shown the distribution subpopulations in Colorado. of band-tailed Permanent bait sites should be established only in flock areas which contribute the fall harvest. establishing to approximately 60 man-days annually for counts. of whole corn per day at each site, be required substantially If 4 of the 14 known flocks were selected for permanent bait sites, would be required pigeon each year. baiting of each area. At a baiting level of 5 kg about 4,000 kg of corn would Cooperation would be required for daily If a Division of Wildlife employee did not live or work near the bait site, a volunteer could be selected to pro- vide the grain for the pigeon flock each day. �190 LITERATURE CITED Amason, A. N., recapture and L. Baniuk. analysis 1980. A computer system for mark- of open populations. J. Wildie Manage. 44: Movements and hunting mortality of Colorado 325-332. Braun. C. E. 1972. band-tailed Conf. pigeons. Trans. North Am. Wildl. and Nat. Resour. 37:326-334. 1973. Colorado. Distribution Proc. and habitats West. Assoc. of band-tailed pigeons in State Game and Fish Comm. 53: 336-344. 1976. Methods for locating, tailed pigeons in Colorado. trapping and banding band- Colo. Div. Wildie Spec. Rep. 39. 20pp. ---- , D. E. Brown. J. C. Pederson, Results of the Four Corners investigation. U.S. and T. P. Zapatka. cooperative band-tailed 1975. pigeon Fish and Wildie Servo Resour. Publ. 126. 20pp. Brown, D. E., and C. H. Lowe. compatible classification the Southwest, , and potential for natural with particular Acad. Sci. 9 (Supp!. ---- 1974a. 2). 1974b. vegetation. 3). 56pp. computer- and potential vegetation reference to Arizona. in J. Ariz. llpp. The Arizona system for natural Illustrated summary through for the North American Southwest. (Supp!. A digitized J. Ariz. and the fifth digit Acad. Sci. 9 �191 Davies. R. G. 1971. Computer programming in quantitative Academic Press. London. England and New York. Dr'ewien, R. C •• R. J. Vernimen. 1966. Spring S. W. Harris. weights of band-tailed biology. N.Y. 492pp. and C. F •. Yocum. pigeons. J. Wildl. Manage. 30: 190-192. Fitzhugh. E. L. 1974. gland activity in Arizona. Glover. F. A. Chronology and breeding Ph.D. 1953. Thesis. A nesting (Columba f. fasciata) of calling. egg laying. among wild band-tailed Univ. Arizona. study pigeons Tucson. 74pp. of the band-tailed in northwestern crop California. pigeon Calif. Fish and Game 39: 397-407. Hewitt. O. H•• and P. J. Austin-Smith. for field-marking Jeffrey. fasciata). 1977. Band-tailed shore and upland game birds Assoc. Fish and Wildl. Agencies. Jolly, G. M. 30: 625-627. pigeon Pages 210-245 in G. C. Sanderson, of migratory Int. A simple wing tag J.o Wildl. Manage. birds. R. G•• Chairman. 1966. 1965. Explicit estimates (Columba ed .• Management in North America. Washington. D.C. from capture-recapture with both death and immigration - stochastic model. data Biometrika 52:225-247. Kautz, J. E. 1977. Effects of band-tailed mates of population parameters. Columbia, Vancouver. Keppie, D. M. 1973. pigeon behavior M.S. Thesis, on esti- Univ. British 70pp. Morning commencement of calling of band- tailed pigeons in Oregon. Murrelet 54:28-30. �192 Keppie, D. M., H. M. Wight. and W. S. Overton. band-tailed pigeon census, Wildl. and Nat. Resour. March, G. L., a management need. Conf. A proposed Trans. North Am. 35: 157-171- and R. M. F. S. Sadleir. tailed pigeon 1970. (Columba fasciata) 1970. in British Studies on the bandColumbia. I. Seasonal changes in gonadal development and crop gland activity. Can. J. Zo01. 48:1353-1357. McCaughran. D. A.• and R. Jeffrey. index of relative abundance 1980. of band-tailed Estimation of the audio pigeons. J. Wildl. Manage. 44: 204-209. Neff, J. A. 1947. tailed pigeon. Passmore, M. F. pigeons. Peeters, Habits, North Am. Fauna 58: 1977. M.S. Thesis. H. J. 1962. G. A. F. related Sisson, reference Univ., 1973. 1968. Bay area. U.S. Department Hafner, Climatography New York, N.Y. technique. and heating 1973. and 506pp. of band-tailed pigeons in M.S. Thesis, Oregon State N. C . Monthly normals of temperature, and cooling degree of the United States Asheville. pigeon 57pp. and Atmospheric Admin. Environ. Cen ter, of the band-tailed 56pp. Condor 64: 445-470. Calling behavior of Commerce. precipitation, Corvallis. The estimation of animal abundance to a census Corvallis. 76pp. Oregon State Univ., Nuptial behavior parameters. L. H. of the band- Utilization of mineral sites by band-tailed in the San Francisco Seber, food and economic status 9pp. days 1941-70. 81 (Colorado). Natl. Oceanic Data Serv .• Natl. Climatic �193 White. J. A•• and C. E. Braun. of juvenile band-tailed Zeigler , D. L. 1971. losses of squabs Thesis. Prepared by 1978. pigeons. Age and sex determination J. Wildl. Manage. 42: 564-569. Crop-milk cycles in band-tailed due to hunting pigeons in September. Oregon State Univ •• Corvallis. by 48pp. {!!) tki-i,!£4 ~ k~' Paul D. Curtis / Graduate 'Research Assistant Approved pigeons and _~_:::-?-:-'~_'---..:&='_Z' _';U_' ~_~_/'i._,_&_~~-'-::_.#;-=:I',--7_,,';:::-_ Clait E. Braun Wildlife Research Leader M.S. ��October 195 JOB PROGRESS State of Project REPORT Colorado -------No. W-88-R-26 Job Title: Bird Investigations 1 Job No. Migratory Covered: Personnel: Migratory 6 Work Plan No. Period 1981 Bird Publications April 1, 1980 through March 31, 1981 C. E. Braun, P. D. Curtis, W. P. Gorenzel, J. E. Kautz, T. E. Olson, R. A. Ryder and M. R. Szymczak ABSTRACT Publications planned are as follows: for and accomplished under this job for Segment Curtis, P. D. 1981. Evaluation of daily counts of band-tailed pigeons as a census method. M.S. Thesis. Colorado State Univ., Fort Collins. 113pp. , and C. E. Braun. 1980. Evaluation band-tailed pigeons as a census method. Sci. 12(1) :36-37. ---- Gorenzel, w. P. 28-32. 1980. The American Coot. of daily counts of J. Colo.-Wyo. Acad. Colo. Outdoors 29(2): __ ~ __ , R. A. Ryder, and C. E. Braun. 1981. American coot response to habitat change on a 'Colorado marsh. Southwest Nat. 26· 59-65. Kautz, J. E., and C. E. Braun. 1981. band-tailed pigeons in Colorado. Olson, T. E. 1980. Mourning Colorado. H.S. Thesis. 119pp. Survival and recovery rates of J. Wildl. Manage. 45:214-218. doves and land use changes in eastern Colorado State Univ. Fort Collins. 1980~ Patterns of agricultural land use changes and effects on avian species in northeastern Colorado. Annu. Mtg. Am. Ornithol. Union 89:Abstract. 26 �196 Olson, T. E. 8-11. 1981. Doves and agriculture. Colorado Outdoors 31(2): , and R. A. Ryder. 1980. Avian nesting studies on agricultural ----=-land in northeastern Colorado. J. Colo.-Wyo. Acad. Sci. 12(1):44. Szymczak, M. R. 1981. Distribution and harvest of Canada geese nesting along the foothills of Colorado. Colo. Div. Wildl. Spec. Rep. No. 49. In press. � https://cpw.cvlcollections.org/files/original/9971a0884de5c51f6057cd23868c112b.pdf 973d9ad02775da700496c55ccb33d7f5 PDF Text Text 1 JOB PROGRESS REPORT State of COLORADO Project No . SE- 3-4 ~~~~~~~~~~- Work Plan No. Job Title 1 Endangered Wildlife Investigations Job No. 1 Identification of Habitat Requirements and Limiting Fac tors for Colorado Sauawf ish and Humpback Chubs Period Covered: Personne l: J anuary 1, 1981 to June 30, 1982 C. Haynes, R. Muth, G. Skiba, L. Wycoff ABSTRACT Field collections were made i n two study areas in the Upper Colorado River drainage during January 1, 1981 through June 30, 1982. Field and laboratory methods are presented. Identifications, counts and measurements of 1981 samples are completed and Yampa River data has been stored on the FWS MANAGE computer pro gram (Colorado State University) . Pr eliminary l ab processing of 1982 samples will be initiated in f all , 1982 . In 1981, Yampa River sampl es were dominated by red shiner s (7563 individuals) and , overall, introduced species were mor e numerous both in number of species (11 of 18 taxa) and i ndividuals (9660 ver sus 9398 natives). In the Colorado River, 17 species were coll ected of which 11 were introduced . Red shiner s and fathead minnows were the predominant speci es wi th 24,353 and 16,019 individuals, r espectively . In 198 1, 23 Colorado squawfish were collected in the Yampa study area during July and August . All wer e collected in the lowermost 20 km of the Yampa Canyon, Dinosaur National Monument. Only one squawfish YOY was co l lected in the mainstem Colorado River . This individual was collected at a site in Mesa County wes t of Grand J unc tion in J uly. Analysis of known-age hat cher y-rear ed humpback chubs is presented and analysis of various genetic crosses of humpbacks, bonytails, and roundtails in addi tion to known-age bony t ails and roundtails will begin in fall, 1982. Those analyses will b e used to evaluate unknown field-collected chubs fo r the possible pr esence of humpbacks in the s tudy ar eas . This Job Report represents a preliminary anal ys i s and i s subj ect to change. For this r eason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permis sion of the Director. ��3 IDENTIFICATION OF HABITAT REQUIREMENTS AND LIMITING FACTORS FOR COLORADO SQUAWFISH AND HUMPBACK CHUBS Charles M. Haynes and Robert T. Muth PROGRAM NARRATIVE OBJECTIVE The overall objective of this investigation is to determine the physical/ biotic factors which limit the reproduction and distribution of Colorado squawf ish (Ptychochielus lucius) and humpback chubs (Gila cypha) in Colorado. Similarly, this project is designed to develop field/laboratory methods for evaluating squawf ish and humpback chub habitat and reproductive success which may be employed by management personnel for future habitat enhancement and/or reintroduction programs in accordance with overall recovery efforts. Information derived from this study will be made available to management and research personnel in the Division of Wildlife as well as other cooperating agencies through annual reports, presentations, and technical manuscripts. Additionally, aspects of this project relating to the humpback chub will be incorporated into a doctoral dissertation in 1984-85 by the junior author. SEGMENT OBJECTIVES 1. To identify and describe quantitatively those habitat parameters which limit the distribution and abundance of Colorado squawf ish and humpback chubs in the Upper Colorado River, with particular reference to larval and juvenile stages. 2. To develop methods for evaluating larval/juvenile squawfish and humpback chub habitat and reproductive success which may be employed by management personnel for future habitat enhancement and/or reintroduction programs. 3. To develop methods for the identification of larval and juvenile humpback chubs and differentiation of humpback chubs from immature stages of the roundtail chub and possible Gila spp. hybrids and/or intergrades. 4. To develop baseline criteria for both water quantity and quality as a tool for predicting and mitigating potential impacts from future river development. 5. Compile data and submit reports to appropriate state and federal offices. Note - These objectives are in accordance with tasks 111 and 1111 of the approved Colorado squawfish recovery plan (1978) and tasks 11, 111, 41, 42, and 43 of the approved humpback chub recovery plan (1979). �4 INTRODUCTION As a consequence of documented declines in numbers and ranges of Colorado squawfish and humpback chubs, these species have been listed as endangered by both the federal government and the State of Colorado. Accordingly, recovery plans have been approved for both species. Both plans recognize a number of factors.which appear to be responsible for the endangered status of these species, including habitat alterations (i.e. river modifications, diversions, withdrawals, etc.), competition from nonnative species, and, in the case of humpback chubs, hybridization with congeners. Since 1977, the Colorado Division of Wildlife has been investigating the status of these species via systematic sampling in the Colorado, Yampa, White, and Gunnison Rivers. Recently, studies have been conducted in cooperation with the U.S. Fish and Wildlife Service (Colorado River Fisheries Project). As a result of these efforts, the distribution, relative abundance, and status of these species within the State of Colorado have been documented and described (Wick et al., 1981; Miller et al., 1982a; Miller et al., 1982b). In Colorado, squawfish are apparently restricted to the lower and middle reaches of the Colorado, Yampa, Gunnison, and White Rivers and to a section of the Green River near its confluence with the Yampa in Dinosaur National Monument. Reproduction has been documented in a reach of the Colorado River west of Grand Junction (Mesa County) and in a reach of the Yampa River in Dinosaur National Monument (Moffat County). Humpback chubs appear to have a more restricted distribution, reaching their greatest abundance in the Black Rocks area of the Colorado River. Six humpbacks collected in 1981 in the Green-Yampa system in Dinosaur National Monument and one collected in Cross Mountain Canyon in 1980 suggests the presence of a small population in this area. Although evidently ripe humpback adults have been observed, particularly at Black Rocks, evidence of successful reproduction, via the collection of larval forms, has not been documented since adaquate features for the differentiation of humpback chubs from the more cotm11on roundtail (G. robusta) have not been developed. Additionally, considerable genetic variation appears to exist in the genus Gila in the Upper Colorado River as a consequence of sympatry between the humpback, roundtail, and the apparently extinct bonytail (Q. elegans). Although the "pure" adult forms may be readily identified, "intergrades" and/or "hybrids" are frequently encountered which cannot be readily assigned to a specific taxon. The problem is compounded relative to the identification of young, since adequate differentiation between the "pure" forms has not been accomplished, nor has intergradation or hybridization been addressed. The analysis of larval/juvenile squawfish and humpback chubs, therefore, is an essential aspect of the overall research project. This progress report includes a description of overall field/laboratory methods and a preliminary sutm11ary of data collected during the period January 1, 1981June 30, 1982. �5 STUDY AREA Field activities were conducted in the Colorado (Mesa County) and Yampa Rivers (Moffat County) (Figure 1). The Colorado River study area includes an approximately 32 km (20 mi) river reach from the vicinity of Loma to the Colorado-Utah state line. The Yampa River study area includes an approximately 95 km (59 mi) reach from the upper end of Cross Mountain Canyon to the confluence with the Green River. Approximately 75 km (46.5 mi) of the study area are within Dinosaur National Monument. METHODS General. Field sampling was conducted during periods which reflected pre-runoff, peak-runoff, and post-runoff conditions in the Colorado and Yampa study areas. Collections were made during 4-5 day field trips utilizing a power boat and inflatable rafts in the Colorado while rafts and canoes were used in the Yampa. Cross Mountain Canyon was surveyed entirely on foot since the extremely turbulent nature of the canyon made float trips unsafe. Fishes were collected with 3.0 x 1.2 m and 1.0 x 1.2 m seines (1.6 mm square mesh) and dip nets (0.79 mm square mesh). Samples were collected according to the "qualitative representative sample" approach (Hocutt et al., 1974) i.e., all available habitats that could be seined were sampled to obtain a representative collection of species at a given locality and which would provide relative abundance information, recognizing the bias of the collection technique employed. Current velocity was measured with Marsh-McBirney or Pygmy-Gurley flow meters, or float techniques. Water temperature was measured with a Taylor thermometer. In order to correlate fish distribution and relative abundance with the wide range of measurable variables, samples were collected according to a detailed habitat stratification approach which was designed to reflect the geomorphic, hydrological, and ecological variables of the drainages (Appendix A). This approach incorporated methods outlined by Herrington and Dunham (1967) and Platts (1974), but was modified to reflect conditions of larger streams. In order to interface data with both those collected by DOW personnel in the NW Region and the U.S.F.W.S., habitat designations were coded to be computer compatible with the FWS MANAGE program and data are currently being entered at the Colorado State University Computer Center, Fort Collins. Standardized field/laboratory data forms are employed. Within each study area, three types of sample sites were designated. "Intensive" sections were randomly selected prior to each field trip utilizing a table of random numbers. This process required the initial random selection of one location within the study area, corresponding with a designated river mile. This site, plus each river mile every 5 miles (8 km) above and below the random site were designated as intensive sampling sites. At each intensive site, all recognizable habitats were sampled for a distance of 0.25 (0.4 km) miles above and below the mid-point. Typically, depending upon the initial random site selection, 9-10 intensive sites were designated for the Yampa River study area and 5-6 sites for the Colorado River study area. "Intervening" sections were localities which �6 1089 1()90 107° 41° I I'8 :r <r ~ ~ ::> I .:J 10 '~by I I Canyon 3go 39° N I I I I I I J 0 0 109° Figure 1. ml 20 40 km 30 80 1oe• Upper Colorado River drainage study areas (hatched). 101• �7 were excluded during the random site selection but which were sampled due to the wide variety of habitat types available. "Special" sections (e.·g., Black Rocks) were relatively short river reaches which were exhaustively sampled due to the enormous variety of habitat features available at these localities. Catch-per-unit-effort was defined as number per species per unit area (m 2). Area was determined utilizing sein dimensions and length and width of the individual sein haul. Substrate composition was determined for each sample (Appendix A). Fish specimens were identified and assigned to 10 mm total length (TL) size increments. Larval forms which could not be identified in the field were preserved in 10% formalin and returned to the laboratory. Beginning in May (1982), ichthyoplankton nets were employed in the Black Rocks area (Colorado River) to evaluate larval drift. Nets were approximately 4 m in length with a rectangular opening of 1.5 square meters. Mesh size was 560 µ. Analyses of these samples will be initiated during fall-winter, 1982. Estimates of spawning periods were made using length-frequency information (i.e. number x 10 mm size group) in addition to indications of adult ripeness (i.e. color, tuberculation, gravidness). In addition, squawfish spawning date estimates were determined utilizing calculated ages based upon data derived from Hannnan's (1981) hatchery study. Larval growth rate (rn) was determined for time of hatching up to 45.0 mm TL from the slopes of linear regression analysis for three size increments (i.e. 7.0 - ±15.0 mm TL,> 15.0 - ±30 mm TL, and> 30.0 - ± 45.0 mm TL). Growth rate varied from 0.35 (r = 0.93), 0.36 (r = 0.99) to 0.40 mm/day post-hatching (r = 0.99), respectively. The ages derived from the following equations permitted a projection of both the earliest and latest spawning dates for both a given collection period and the year as a whole. A. + HTmin = larval B. + HTmax = Where: age (days) larval age (days) Lmin = total length (mm) of smallest individual in a collection = total length (mm) HSmax = maximum hatching Lmax of largest individual in a collection size (7 .5 mm TL) (Hamman, 1981) HSmin = minimum hatching size (6.0 HTmax = maximum HTmin = minimum hatching time (3.75 days) (Hamman, 1981) m = growth Colorado Sguawfish. mm TL) (Hannnan, 1981) hatching time (6.0 days) (Hamman, 1981) rate (0.35, 0.36, 0.40 mm/day) Larval Colorado squawfish were identified utilizing �8 known-age larval series obtained from Willow Beach National Fish Hatchery and published descriptions (Snyder, 1981). Humpback chub. At this time, it is not possible to reliably separate larval/juvenile stages of Gila either in the field or laboratory and this problem is addressed in objective three. This project is designed to evaluate three critical research areas relative to the humpback chub: 1. To evaluate the validity of standard morphomeristic techniques for the differentiation of known (hatchery-reared) YOY/juvenile roundtails, humpbacks, and bonytails. 2. To employ the results of (1) above for the identification of field collected inunature chubs. 3. To determine and describe any variation in physical habitat requirements and other biotic associations for sympatric Gila spp. In order to meet the first research need (above), known-age larval series of humpbacks and bonytails have been obtained from Willow Beach National Fish Hatchery (Arizona) and Dexter National Fish Hatchery (New Mexico). Additionally, developmental series of six hybrid crosses have been provided and will be evaluated i.e. humpback x humpback - bonytail bonytail x roundtail - bonytail roundtail x bonytail roundtail x roundtail - bonytail humpback x bonytail bonytail x humpback - bonytail The original humpback chub brood stock was captured at Black Rocks. Bonytail brood stock was from Lake Mojave, California. Unfortunately, roundtail brood stock was captured at Black Rocks and was therefore sympatric with humpback material. Therefore, we are attempting to produce a developmental series from the Juniper Springs area of the Yampa River and will also utilize a series from the White River (Rio Blanco County). Since no historical humpback records exist for these latter localities, we assume a high degree of genetic purity. Standard morphomeristic techniques are described in Appendix B. Data for each known developmental series will be statistically evaluated to determine if key identification characters can be meaningfully determined. Multivariate statistical testing will involve both principal components and discriminant function analysis, following procedures employed by Daly and Balling (1978), Davis and Baker (1974) and Smith et al. (1979). A differentiation of unknown field-collected Gila larvae into humpback, roundtail, "intergrade" and/or "hybrid" will follow the laboratory phase above and will be a function of the success of the laboratory research and the degree of morphomeristic divergence of these closely-related species. Similarly, the recognition of variation in habitat requirements will also relate to laboratory findings. �9 RESULTS AND DISCUSSION General I. Sampling Bias/Selectivity Several sources of sampling bias are inherent in the design of this study and should be acknowledged. A. B. C. II. Gear Selectivity. Built-in bias against the capture of adults of larger species is inherent in the use of relatively short (3.0 and 1.0 m) seins of small mesh (0.79 and 1.6 uun). However, gear selection is consistent with the objective of sampling young-of-the-year (YOY) fishes. Habitat Selectivity. Design and size of sampling gear is biased against the sampling of deeper swifter sites. Such sites could be more efficiently sampled with electrofishing and/or passive netting techniques; however, these methods would not capture YOY forms and most juveniles. Seining Efficiency. Efficiency varies according to both width and depth of the individual sein haul and substrate. This study is not designed to account for sein efficiency variability. Yampa River Study Area. A total of 422 samples were collected during March - August, 1981 (Table 1). Laboratory processing and computer storage of these data has been completed, as well as a preliminary data analysis. A total of 103 samples have been collected during April - June, 1982; however, these samples will be processed during fall-winter, 1982. A total of 18 species have been identified from 1981 samples, of which seven were native (Table 2). Unidentified YOY chubs (Gila spp.) await taxonomic clarification and are listed here as a single taxon. Several specimens of innnature minnows and suckers could not be positively identified due to their early stage of development and are listed as "unidentified Cyprinidae" and "unidentified Catastomidae", respectively. For the Yampa River study area as a whole, 19,058 fishes were identified in 1981. Of these, the introduced red shiner (Notropis lutrensis) was the predominant species (7563 individuals) and native unidentified chubs (Gila spp.) were the second most numerous (5823). The number of individuals of native species was slightly smaller (9398) than introduced species (9660). Based on a length-frequency analysis for YOY Yampa fishes (Appendix C), 1981 spawning periods for the eight most abundant species were determined (Figure 2). Species-habitat associations have not been completed and these will hopefully be conducted utilizing the MANAGE program. III. Colorado River Study Area A total of 354 samples were collected during April - July, 1981 (Table 3}. Sample number per habitat type has not been compiled, nor has computer �Table 1. Number of Collections By Habitat Classification, Yampa River, 1981. for Definitions of Habitat Types. Main Channel BA ED PO RI RU 4 57 3 4 2 SH 171· IP PU 2 3 EM 111 Refer to Appendix A RA RF co TOTALS 1 1 21 380 ...... 0 Side Channel Totals ) 8 1 3 2 3 15 8 - 12 58 6 6 5 186 10 3 ) 1 - - 1 42 112 1 1 22 422 . ... ) �) ) Table 2. ) Individuals per species, Yampa River study area, 1981 (N SPECIES = native) DATES July April June Catastomidae Catostomus commersoni (white sucker) C. discobolus (bluehead sucker) N C. latipinnis (flannelmouth sucker) N Unid. Catastomidae 0 1 8 0 43 37 0 1 163 46 0 3 320 88 1 9 527 179 1 Centrarchidae Lepomis cyanellus (green sunfish) 0 0 4 0 4 Cottidae Cottus bairdi (mottled sculpin) N 1 2 1 0 4 13 6 0 0 20 8 0 0 281 70 1 22 0 541 234 0 0 46 10 1176 42 0 0 4310 6391 312 956 13 1106 336 1 32 Cyprinodontidae Fundulus zebrinus (plains killif ish) 0 0 0 1 1 Ictaluridae _!_. Eunctatus (channel catfish) 0 0 0 11 11 Salmonidae Prosopium williamsoni (mountain whitefish) N 1 0 0 0 1 59 1236 3883 13881 19059 Cyprinidae Cyprinus carpio (carp) Gila spp. N Notropis lutrensis (red shiner) N. stramineus (sand shiner) Pimephales promelas (fathead minnow) Ptychochielus lucius (Colorado squawfish) N Rhinichthys osculus (speckled dace) N Richardsonius balteatus (redside shiner) Semotilus atromaculatus (creek chub) Unid. Cyprinidae Totals 1 5 2 1219 1101 72 August Total 2 5823 7563 23 620 1 32 ....... ....... �SPECIES Gila sp. Rhinichthys osculus Notropis lutrensis Notropis stramineus Pimephales promelas Catostomus discobolus Catostomus latipinnis Lepomis cyanellus E 1 M 1 L MARCH Figure 2. ) E 1 M 1 APRIL L -··---- --·-·-·-·-· ·-·-···-·---· ----·----· -·-·-·-·-·-·· -·-·-· ·----·-----·-·-· -·-·-·· -·-·-·· -· ----·---· -·-·-··-·-·-·-·-· -·-···-·-·· -· ------·-·-·-·-·-·· ··-·-···-·--·-------·-·-·· -·-·-·-·-·-·-·-·.- - --· ----·-·-·4 -·-·-· -·-·- ·-·-·-·---·--·-·-····-·-····-·-·E 1 M 1 MAY L E 1 M 1 JUNE L E 1 M 1 JULY L EI MI L AUGUST 1 E 1 SEPT. Spawning periods of selected species, Yampa River study area, 1981. Dashed line represents observed spawning season based upon length-frequency analysis. Solid line represents principal spawning period, while dotted/dashed line represents reported spawning period extremes. ) M ) L ..... N �13 ~ storage been completed. During April - June, 1982, 114 sein samples and approximately 20 one-hour drift net samples were collected. These samples will be processed and placed in computer storage during fall-winter, 1982-83. A total of 17 species have been identified from the Colorado River study area in 1981, of which only six were native to the drainage. As in the case for the Yampa data, YOY chubs were not distinguishable as either roundtailsorhumpbacks and were all grouped as Gila spp. for the purposes of this progress report. For the Colorado River study area as a whole, 48,901 fishes were processed in 1981. Of these, red shiners were the most numerous (24,353 individuals) followed by another exotic, fathead minnows (Pimephales promelas), with 16,019 individuals. Gila spp. was the most abundant native group with 3,488 individuals. Spawning periods for the eight most abundant species in this study area during 1981 have been determined (Figure 3) based upon length-frequency analyses (Appendix C). Comparisons of spawningrelated data for Colorado - Yampa River species by other researchers are presented in Table 4. Species-habitat associations will be conducted utilizing the MANAGE program. On 2 May 1981, 11 redside shiners (Richardsonius balteatus) were collected 48 km west of Grand Junction at km 230.4 - 243.4 (mi 143.1 - 151.2). This species, while relatively connnon in the Yampa River study area, had not been previously documented in the mainstem Colorado River in either the upper or lower basin. This observation was published in The Southwestern Naturalist (Haynes, et al., 1982). ~Colorado Sguawf ish In 1981, 23 larval squawfish were collected in the Yampa River study area, while only one was collected in the Colorado River study area. Yampa squawfish collected on 7/22-26/1981 were captured by NW Regional personnel and submitted to Nongarne Research staff for length-frequency analysis. Squawfish collected on 8/11-18/1981 were captured by Nongame Research personnel, as was the single Colorado River specimen. Collection data for both 1981 Yampa and Colorado River squawfish larvae are presented in Table 5. Estimated 1981 spawning periods in the Yampa River based upon maximum and minimum size per collection and estimated growth rates ranged from 25 June to 3 August. The earliest Yampa data correlates closely with both dates and locations when ripe radiotagged adults were observed over apparent spawning gravels in the same area by FWS personnel (Tyus et al., 1981) i.e., 26 June - 10 July, km 19.6 - 0.2.Juvenile (age-class I) squawfish were not collected. In July 1981, seven of the Yampa squawfish larvae were collected in main channel embayrnents, two were collected in side channel back.waters, while the remaining individual was collected in a side channel isolated pool. Eight were captured over a predominantly silt substrate whereas the remaining two were associated with a substrate that was predominantly sand. For all individuals, velocity could not be detected and was recorded as 0.0 m/sec. Depth varied from 0.1 - 0.3 meters. In August, all 13 YOY squawfish were associated with main channel embayments with �Table 3. Individuals per species, Colorado River study area, 1981. April SPECIES Catastomidae Catostomus conunersoni (white sucker) c. discobolus (bluehead sucker) N C. latipinnis (flannelmouth sucker) N Centrarchidae Lepomis cyanellus (green sunfish) Micropterus salmoides (largemouth bass) Cottidae Cottus bairdi (mottled sculpin) N Cyprinidae Cyprinus carpio (carp) Gila spp. N Notropis lutrensis (red shiner) N. stramineus (sand shiner) Pimephales promelas (fathead minnow) Ptychochielus lucius (Colorado squawf ish) N Rhinichthys osculus (speckled dace) N Richardsonius balteatus (redside shiner) • Cyprinodontidae Fundulus zebrinus (plains killifish) Ictaluridae I. melas (black bullhead) Y. punctatus (channel catfish) Total ) ) (N = native) DATES June May Total July 1 138 16 4 99 44 0 54 18 1 132 53 6 423 131 27 4 5 60 0 44 0 9 136 13 0 0 0 2 2 3 410 6115 938 2293 0 245 0 1 1230 6462 1174 2647 0 142 10 1818 11553 1293 10694 1 363 0 14 3488 24353 3409 16019 1 752 11 0 30 223 4 385 0 2 0 12 0 0 10 22 0 1 0 3 0 0 4 113 4 117 10214 11811 776 26100 48901 ...... J:-. 11 ) �) ) : J.tj . I ,i :~ :.l~: ) f,,. ---·-·-·-··-·-·- ·-·-·-· -·-·- SPECIES ··- Gila sp. - 4 -·--- Rhinichthys osculus Notropis lutrensis Notropis stramineus Pinephales promelas Catostomus discobolus Catostomus latipinnis Lepomis cyanellus EI MI L MARCH Figure 3. - E 1 M 1 APRIL L -·-·-·-··-·-·--··-·-·------·-·-·4 -·-·-·· -·-·-··-·----·-·-·· -·-·--·-·-·· -·-·-·· --·-·-·· --·-·-·· --·----·-·-· -·-·-·· -·-·-·· -·-·-·· -----·----·-•-•1 -·-·-·I -·-·-·-·-·-·· --1-----·-·-·-·· -·-·-·· -·-·-·- --·-·-· -·-·-·I -·-·-·E I M I MAY L E 1 M 1 JUNE L E 1 M 1 JULY L E 1 M 1 L AUGUST E 1 M 1 SEPT. Spawning periods of selected species, Colorado River study area, 1981 . Dashed line represents observed spawning season based upon length-frequency analysis. Solid line represents principal spawning period, while dotted/dashed line represents reported spawning period extremes. L �Table 4. Reproductive and early life history information on selected species, Colorado and Yampa Rivers. Reported Spawning Times Reported Spawning Temperatures References Size at hatching (mm SL/TL) Incubation Time and Temperature Catostomus discobolus 10-11/ 10-11 7 days -18-21°c (cultured) 7-8 days 15.5-17.8°c 13-23°c est. May-August Yampa River, CO Yampa River, co June 15-18°C Yampa River, CO Yampa River, co Carlson et al. (1979);. Holden (1973); Snyder (1981) Catostomus latiEinnis 10-11/10-11 6-7 days 15.5-17.8 0 c May-July 12°c Yampa River, co 6-12°c Yampa River, co 13-23°c est. Yampa River, co Carlson et al. (1979); Holden (1973); McAda (1977) 18°c Behnke and Benson (1980); Colorado River Fishes Recovery Team (1979; Holden (1973); Snyder (1981); Suttkus and Clemmer (1977) Species Gila C:2]~ha Gila robusta ) 6-7/7 ?/7 June-July Grand Canyon 4.8-6.7 days 19-20°C (cultured) 4.2-6 days 21-22°c (cultured) May-June 20-21°c Green River, UT (cultured) June-July 18°c Yampa River, CO Yampa River, co Green River, UT Green River, UT 15-24oc est. May-August Yampa River, CO Yampa River, co 6-8 days ) ..... °' Carlson et al. (1979); Holden (1973): Vanicek and Kramer (1969) ) �) ) ) Table 4. Cont. Size at hatching (nrrn SL/TL) Incubation Time and Tempterature Lepomis cyanellus /3.5-3.7 2-5 days May-July Wyoming Late Spring to Mid-Sunrrner 23°c Wyoming 20-28°c Balon (1975); Baxter and Simon (1970); Childers (1967); Eddy and Urderhill (1974); Meyer (1970) Notropis lutrensis 3-4/4 5 days 23.3°C May-October Kansas June-September Oklahoma June-July Wyoming 15.5-29°C Kansas 26. 7-31. 7 Oklahoma Baxter and Simons (1970); Tuber (1969); Simon (1946); Snyder (1981) Species 4 days 28.9 Reported Spawning Times Reported Spawning Temperatures References Notropis stramineus 3-4/3-4 6 days June-September 16-26°c est. Yampa River, co 21-27°c Kansas Carlson et al. (1979); Snyder 1981; Summerfelt and Minckley (1969) Pimephales promelas 3-4/3-4 5-6 days 25°c May-August 13-23° est. Yampa River, CO 13-18°c Colorado Andrews and Flickinger (1973); Carlson et al. (1979); Dobre et al. (1956); Snyder (1981); Snyder et al. (1977) P. lucius 6-7/6-7 4-5 days 20-21°c (cultured) Hanrrnon (1981); Snyder (1981); �Table 4. Cont. Size at hatching (mm SL/TL) Rhinichthys osculus 5-6/5-6 Incubation Time and Temperature 5 days 16°c Reported Spawning Times May-July 13-23°c est. Yampa River, co 12-1a0 c Baxter and Simons (1970); Carlson et al. (1979); Fuiman and Loos (1977); Minckley (1973); Snyder (1981) June-September 16-26°c est. Yampa River, CO 21-37°c Kansas Carlson et al. (1979); Summerfelt and Minckley (1969); Snyder (1981) 6 days 18-19°c (cultured) Richardsonius balteatus 5/3.9-5.S ) 3-7 days 21-23°c (cultured) ) Reported Spawning Temperatures ..... co ) �19 Table 5. River Sunnnary of Larval Colorado Squawfish collections, Yampa and Colorado River study areas, 1981. Date Locality Number (nnn) (km) Yampa 7/24 19.6 2 11.0 7/25 17.4 14.2 9.6 8.8 1 1 2 9.0 11.0 11.0-12 .o 12.0 5.2 0.2 1 2 7/26 ~ Colorado Total Length 1 10 11.0 12.0-13.0 9.0-13.0 Estimated Age (Days) Estimated Spawning Period (Days) 8-26 7 /1-7 /18, 1981 8/14 28.8 14.6 4 2 10.5-13.0 11.0-14 .o 8/15 8.5 8.4 6 12.0-16.5 22.0 10.5-22.0 12-52 6/25-8/3, 1981 18.0 34-40 6/18-6/24, 1981 7/27 243.6 1 TI 1 �20 no measurable velocity. Six individuals were associated with a predominantly silt substrate, while the remaining seven were found over sand. Depths ranged from 0.2 - 1.3 meters. A long-term analysis (1979-1983) of the possible relationships of YOY distribution and abundance with physical habitat variables (i.e. seasonal flow and temperature patterns) is being conducted and this analysis will be included in the final report. Gila spp. To date, reliable criteria for differentiation of YOY humpbacks, roundtails, and bonytails do not exist. Collections of larval and juvenile chubs have been made in both study areas and, although the specimens are probably predominantly roundtails, stored specimens are classified as Gila spp. It is anticipated that morphomeristic analysis of known-age hatchery larvae of bonytails and various genetic crosses will be completed during winterspring of 1983. Multivariate statistical analyses of "knowns" will be employed to determine those features, if any, which may be used to distinguish the species, afterwhich unknown field-collected specimens will be compared. Tables D-1 and D-2 sunnnarize morphomeristic analysis of hatchery-reared humpbacks (Little Colorado River and Black Rocks stock, respectively. Spinal deformities (lordosis) in YOY chubs were observed during 1981 August Yampa River collections. ·In 1981, 101 of 127 samples (79.5%) contained deformed chubs. A total of 4032 specimens were examined, of which 667 (16.5%) were deformed. A subsequent inspection of August 1980 Yampa samples revealed deformed chubs in 68 of 106 samples (64.1%). A total of 3497 YOY chubs was examined of which 360 (10.3%) were deformed. For 1981, YOY chubs ranged from 14.5 - 48.0 mm TL and were probably 17-84 days old. Deformed specimens were 23.0 - 41.0 mm TL suggesting that spinal curvature appeared at 34-70 days. A number of yearlings (> 48 nun TL) were collected, but lordosis was not observed in this group. Examination of cleared and stained whole specimens indicates a gradual spinal curvature beginning near the 10th trunk vertebra through the 11th caudal. Maximum ventral depression was at the 3rd and 4th caudal. Vertebral rupture, separation, or compression were not evident. Microscopic examination of saggital series has not revealed any readily noticable gross differences between deformed and normal fish. Examinations for two parasites known to be associated with fish spinal deformities (Myxosoma cerebralis and Ichthyosporidium hoferi)werenegative. At this time, neither cause-effect nor impact upon Yampa River chubs can be determined; however, it is likely that such deformities, unless reversible, reduce the fitness of individuals relative to feeding, predator escape, etc. A number of factors have been shown to cause lordosis, including water quality, genetics, radiation, and electric current. Based upon spinal deformities observed in wild populations of salmonids, pike, and herring--none of which had frequencies as high as Yampa River chubs--Bengston (1979) proposed using the frequency of deformities as a monitor of marine pollution. An evaluation of frequency of deformities in 1982 collections will be made as samples are processed. ~ �21 Razorback Sucker No razorback sucker (Xyrauchen texanus) larvae or juveniles were collected in 1981, nor were any reported by Wick et al. (1981), Miller et al. (1982a) or Miller et al. (1982b) for the years 1977-81. The collection of ripe adults by the above authors in several areas of the Colorado and Yampa Rivers suggests that, while spawning may occur, it is probably quite · limited and that hatching and/or larval survivorship is very low if it occurs at all. �22 LITERATURE CITED Bengtson, B.-E. 1979. Biological variables, especially skeletal deformities in fish, for monitoring marine pollution. Phil. Trans. R. Soc. Lond. B. 286, 457-464. Daly, H. v. and S. S. Balling. 1978. Identification of africanized honeybees in the western hemisphere by discriminant analysis. J. Kans. Ent. Soc. 51(4): 857-869. Davis, B. L. and R. J. Baker. 1974. Morphometrics, evolution, and cytotaxonomy of mainland bats of the genus Macrotus (Chiroptera: Phyllostomatidae). Syst. Zool. 23(1): 26-39. Ha11llllan, R. L. 1981. Spawning and culture of Colorado squawfish in raceways. Prog. Fish-Cult. 43(4): 173-177. Haynes, C. M., R. T. Muth and L. C. Wycoff. 1982. Range extension for the redside shiner, Richardsoni.us bal teatus (Richardson) , in the upper Colorado River drainage. The SW Nat. 27(2): 223. Herrington, R. B. and D. K. Dunham. 1967. A technique for sampling general fish habitat characteristics of streams. US For. Ser., Intermountain Forest and Range Exp. Sta., Res. Pap. INT-41. 12pp. Hocutt, C. H., R. L. Kaesler, M.·T. Masnik, and J. Cairns, Jr. 1974. Biological assessment of water quality in a large river system: an evaluation of a method for fishes. Arch. Hydrobiol'. 74(4): 448-462. Miller, W. H., Archer, D., Tyus, H. M. and R. M. McNatt. 1982(a). Yampa River Fishes Study Final Report, Colorado River Fisheries Project, FWS. 78pp. Miller, W. H., J. J. Valentine, D. L. Archer, H. M. Tyus, R. A. Valdez, and L. Kaeding. 1982(b). Colorado River Fishery Project. Final Report. Colorado River Fishery Project, FWS, 42pp. Platts, W. S. 1974. Geomorphic and aquatic conditions influencing salmonids and stream classification with application to ecosystem classification. Surface Environ. and Mining Program, U.S. For. Ser. 199pp. Smith, G. R., R. R. Miller, and W. D. Sable. 1979. Species relationships among fishes of the genus Gila in the upper Colorado River drainage. In: Linn, R. M., ed., Proc:-Tst Conf. Sci. Res. Nat. Parks 1: 613-623. U.S. Dept. Interior NPS Trans. & Proc. Ser., No. 5. Snyder, D. E. 1981. Contributions to a guide to the cypriniform fish larvae of the upper Colorado River system in Colorado. BLM Biol. Sci. Ser., 3. Denver. 81pp. �23 Tyus, H. M., E. J. Wick and D. L. Skates. 1981. A spawning migration of Colorado squawf ish (Ptychochielus lucius) in the Yampa and Green Rivers, Colorado and Utah, 1981. 13th Ann. Symp., Desert Fishes Council, Death Valley Nat. Mon., CA., 1981. Wick, E. J., T. A. Lytle and C. M. Haynes. 1981. Colorado squawfish and humpback chub population and habitat monitoring, 1979-1980. Prog. Rep., Endangered Wildlife Invest. SE-3-3. Colo. Div. Wildlife. 156pp. Prepared by: M. Haynes Aquatic Nongame Researcher �24 APPENDIX A: PHYSICAL HABITAT STRATIFICATION PROCEDURES The following habitat stratification approach is designed to reflect the geomorphic/hydrologic variability of the Upper Colorado River drainage. These habitat evaluation methods are designed to quantify physical variables in such a manner as to provide high resolution relative to fish habitat electivity. The habitat evaluation and data recording procedures are used by both the NW Region survey personnel who concentrate largely on adult fishes collected by electrofishing and research personnel who concentrate on larval forms collected largely by seining. With minor modifications, the system will interface with procedures used by FWS (Colorado River Fisheries Project) and is computer compatible with the MANAGE database system. Figure A-1 depicts most habitat descriptors in a generalized river reach. For each sample collected at an intensive, intervening, or special site, a "primary habitat" designation is assigned. Primary habitats are designed to reflect largely riverine geomorphic variation. Within each primary habitat, a variety of "specific habitat" types exist, which reflect the variability of discharge and flow. For each primary-specific habitat pair, substrate, cover, current velocity, water temperature, depth, and area sampled are determined. Strata Definitions: I. Primary habitat MC (Main Channel). That ~ection of a river which carries the greatest part of the flow during all seasons. CC (Chute Channel). High gradient secondary channel with high velocity and large substrate size in the upper section typically followed by a deep pool. Lower section is usually characterized by decreased velocity and small substrate size. SC (Side Channel). A secondary channel which may carry appreciable flow and provide either low velocity or near stagnant habitat, particularly in the lower section. Gradient and velocity are low and similar to main channel. Side channels are usually depositional with substrates of small particle size. Emmergent and/or submerged macrophytes frequently abundant. TS (Tributary Stream). ID (Irrigation Ditch). Man-made diversion from river, used chiefly for irrigation. IK (Lake). A natural lake or manmade impoundment on a permanently flowing stream. GP (Gravel Pit). Excavation for gravel mining. or permanently connected to river. An inflowing permanent or ephemeral stream. May be periodically �25 II. RP (Reservoir, Primary). stream. Reservoir on 4th order or larger permanent RS (Reservoir, Secondary). permanent stream. RT (Reservoir, Tertiary). 0I (Offstream Impoundment). channel. Reservoir on 3rd order or smaller Reservoir on ephemeral tributary stream. Excavation isolated from the river Specific Habitat BA (Backwater). A body of water off the main channel with no measurable velocity, often created by a drop in water level which partially isolates a former secondary channel or by high water levels which flood low-lying areas. ED (Eddy). Whirlpools or turbulent backcurrents created by obstructions or islands in the channel or by the juncture of two channels below an island. P0 (Pool). A portion of stream that is deep and quiet relative to the main current. RI (Riffle). A shallow rapids in open river where the water surf ace is broken into waves by obstructions or irregular substrate wholly or partially submerged. RU (Run). A stretch of relatively deep fast-flowing water with the surface essentially nonturbulent. SH (Shoreline). The shallow, low to negligible velocity waters next to shore. RA (Rapid). A relatively deep and fast flowing area characterized by standing waves and whitewater caused by channel constriction or obstructions. RF (Rubble Flat). Quiet water areas of large substrate particle size, with some silt deposition. Typically associated with island heads and chute channels, but may be located elsewhere. EM (Embayment). Shoreline concavities or depressions often created by shoreline obstruction or irregular substrate and associated with low velocity shoreline backcurrents. Length greater than width at mouth. C0 (Concavity). length. Similar to EM except width at mouth greater than· �26 III. IV. IP (Isolated Pool). blocked. A backwater pool with access to main channel PU (Puddle). Isolated on-shore bodies of water created by falling water levels, not cutoff secondary channels. Substrate SI (Silt). Fine gritty material that is suspended easily. SA (Sand). Less than 1/10 inches in diameter. (<3 nnn). CL (Clay). Compacted fine particles in bed form that are not suspended easily. OR (Organic Debris). GR (Gravel). 3/8 to 3 inches in diameter (3 nnn to 76 mm). RU (Rubble). 3 to 12 inches in diameter (76 mm to 305 mm). B0 (Boulders). Individual segments of rock larger than 12 inches in diameter {>305 mm). BE (Bedrock). Muck derived from dead decaying matter. Large solid masses of rock without individual form. Cover V (Vegetation). Living emergent vegetation as cover. B (Brush). Dead plant material, such as Russian thistle and tamarisk or other brush which may provide cover. R (Rock). Cover as gravel, rubble and/or boulders (correlated with substrate). 0 (Overhang). Overhanging cliffs and/or ledges which may shade the water at certain times. T (Turbulence). Surface disturbance or turbulence which may provide cover. S (Shade). Cover provided by riparian vegetation which may shade the water at certain times. D (Debris). Cover provided by accumulations of fine aquatic and/or terrestrial organic matter. N (None). No observable structure, irregularities, etc. �27 Figure A- 1 . Gener alized river reach in the Upper Colorado River Basin depicting physical hab i t a t descriptors. Refer to t ext, Appendix A, fo r str a t a def initions and codes . �28 APPENDIX B: METHODS FOR THE LABORATORY ANALYSIS OF GILA CYPHA, Q. ROBUSTA, AND G. ELEGANS Morphomeristic data to be compiled on known developmental series of humpback chubs, roundtail chubs, and bonytails, and genetic crosses of these species, include the following: I. Lengths A. B. C. D. E. F. G. H. II. Anterior margin of snout to: (1) anterior margin of eye (AE) (2) posterior margin of eye (PE) (3) origin of pectorals (OPl) (4) origin of pelvics (OP2) (5) posterior margin of yolk (PY) (6) origin of preanal fold (OPAF) (7) origin of dorsal f infold (ODF) (8) origin of dorsal fin (OD) (9) insertion of dorsal fin (ID) (10) posterior margin of vent (PV) (11) origin of anal fin (OA) (12) insertion of anal fin (IA) (13) posterior margin of hypural plates(PHP) (14) anterior margin of fork of caudal fin (AFC) (15) posterior margin of fork of caudal fin (PC) Yolk (Y) Pectoral fins (Pl) Pelvic fins (P2) Dorsal fin (D) Anal fin (A) Longest ray of anal fin (LAR) Longest ray of dorsal fin (LDR) Widths A. B. C. D. E. F. III. Innnediately behind posterior margin of eye (BPE) At origin of pectoral fins (OPl) At origin of dorsal fin (OD) Innnediately behind posterior margin of vent (BPV) At anterior margin of most posterior myomere (AMPM) Maximum width of yolk (Y) Myomere counts to: A. B. C. D. E. F. Posterior margin of yolk (PY) Origin of preanal finfold (DPAF) Origin of pelvic fins (OP2) Origin of dorsal fin (OD) Posterior margin of vent (PV) Total myomere number (Total) �29 IV. Fin ray counts (principal fin rays only) A. B. C. D. E. V. Oblique measures A. B. VI. Most posterior edge of opercle or origin of anal fin (POP to OA) Origin of anal fin to posterior margin of hypural plates (OA to PHP) Depths A. B. C. D. E. F. G. H. VII. ,,.,.... Caudal (C) Dorsal (D) Anal (A) Pectorals (Pl) Pelvics (P2) Immediately behind posterior margin of eye (BPE) Origin of pectoral fins (OPl) Origin of dorsal fin (OD) Immediately behind posterior margin of vent (BPV) Anterior margin of posteriormost myomere (AMPM) Maximum width of yolk (Max. Y) Middle of eye (ME) Least depth of caudal peduncle Calculated characters A. B. c. D. E. F. G. H. I. J. Eye diameter = posterior margin of eye - anterior margin of eye (PE-AE) Head length = anterior margin of snout to origin of pectorals (OP2-0P1) Pectoral fin length + anterior margin of snout to origin of pectoral fins [Pl length + (AS to OP2 - AS to OPl)] Pelvic fin length + anterior margin of snout to origin of pelvic fins anterior margin of snout to origin of pectoral fins [P2 length + (AS to OP2 - AS to OPl)] Origin of anal fin to posterior margin of hypural plates + posteriormost edge of opercle to origin of anal fin (OA-PHP + POP-OA) Anterior margin of snout to anterior margin of fork of caudal fin + anterior margin of snout to posterior margin of fork of caudal fin (AS to AFC + AS to PC) Anal fin basal length = anterior margin of snout to origin of anal fin (IA-OA) Snout length = anterior margin of snout to anterior margin of eye (AS-AE) Dorsal fin basal length = anterior margin of snout to insertion or dorsal fin - anterior margin of snout to origin of dorsal fin (ID-OD) Sum of fin lengths (including caudal) Total number of morphomeristic characters to be used for analysis of larval/juvenile humpbacks, roundtails, bonytails, and genetic crosses= 57. The number of characters vary according to developmental phase i.e. protolarvae - 30, mesolarvae - 54, metalarvae - 50, and juvenile - 47. �-I WIDTHS~ DEPTHS jE---- LENGTHS FROM SNOUT AS AE -4DEPTHS ~WIDTHS!<- Anterior Margin of Snout to: Anterior Margin of Eye (Snout Length) PE OPl Origin of Pectoral Fin (Head Length) ODF Origin of Dorsal Finfold OPAF Origin of Preanal Finfold - OD Origin of Dorsal Fin (Predorsa 1 Length) OP2 Origin of Pelvic Fin (Prepel vie Length) lr,;,ertion of D"rsal Fin (ID minus OD 1s Bare Length) PY (J.J' 0 Posterior Margin of Yolk Posterior Margin of Vent (Snout- to-Vent Length) Origin o• Anal Fin (Preanal length) IA Insertion of Anal Fin (IA minus OA is Base Length) Posterior Margin of Hypural Plates or Notochord (Stantl•rd Length) Anterior Margin of Fork of Caudal Fin (Fork Length) AMPH Posterior Margin of Caudal Fin (Total Length) ) ) ) �31 ·APPENDIX C: LENGTH FREQUENCY DISTRIBUTIONS Yampa (Figures C-1 through C-4) and Colorado River (Figures C-5 through C-9) study areas, 1981 �32 Tol•I 1 Le~ h 4122 S•mplin9 D•IH 0116-19 1122 ·26 Total 8/11-18 Lr~ 1h 10 4 11-20 88 83 21·30 31·40 108 31·40 41·50 511 41·50 51·60 19 51-60 11·20 38 21•30 61•70 8/11-18 ~ 61·70 4 11·80 11·80 81·90 81·90 91-100 91-100 101·110 101-110 111·120 111·120 121·130 121·130 131-140 131·140 141·150 1'1-150 Bluehead Sucker Carp 151· 151· Tol1I 4122 Sampling Delea 6116·19 7/22-28 Tol1I 8111·18 L~~~1h 1·10 1·10 11·20 11·20 21·30 21-30 31•40 31·40 41·50 4 51·80 61-70 61-70 11·80 71·80 81-110 81·90 91-100 91-100 101-110 101·110 111·120 111-120 121·130 121-130 131-140 131-140 141·150 4122 S1mphng Dates 8116·19 7122·26 12 41-50 51-60 141-150 Channel Catfish Ui1· Sampling D•t•• 6116-19 1/22 ·26 1·10 1·10 Lr~h 4122 Colorado Squawfish 151· Figure C-1. Length-frequency distributions for bluehead sucker, carp, channel catfish, and Colorado squawfish; Yampa River study area, 1981. �33 Tot•I Lr~r •122 S•mpltng D•tH 6/16·19 7122 ·26 8111·18 Tol•I L•n9th (INT'l C'22 S•mphng 6116·19 0~10 7'22. 26 1·10 12 1·10 16 11·20 14 •4 8111 18 1• •8 11·20 21·30 76 109 21·30 126 31·•0 101 105 31·•0 10 •1-so 40 10 •1-so 97 19 51-60 30 61-70 26 81-70 71-80 14 71-80 81-90 5 81-90 51-60 4 11 91-100 91-100 101-110 101·110 111-120 111-120 121·130 121·130 131-140 131·140 35 141-150 141-150 Sand Shiner Redside Shiner ISi- 151- ~ Tol•I Length Imm 4122 Sampling D•IH 7122-26 8/18-19 1-10 Tol•I 8111-18 L~~ft:~ 4 715 306 11-20 21-30 25 286 485 21-30 278 8 272 31-40 41-SO 177 6 38 41-50 Sl-60 39 15 51-60 61-70 4 61-70 71-80 14 71·80 8 81-90 81-90 91-100 91-100 101-110 101-110 111-120 111-120 121-130 121-130 131-140 131·140 141-150 141-150 Speckled Dace IS1- Sampling D•IH 7122 - 26 6118·19 1-10 153 11-20 31-40 4,22 White Sucker ISi· Figure C-2. Length-frequency distributions for creek chub, fathead minnow, flannelmouth sucker, and Gila spp.' Yampa River study area, 1981. 8'11·18 �34 Total Length Imm! 4122 1-10 Sampling Oalea 8118·19 7/22 ·28 lot al 8/11·18 4 Lr~i"° 4122 Sampling Datu 7122 ·28 6116·19 8/11·18 1·10 11·20 11-20 21·30 21·30 31-40 31·40 41·50 41-50 51-60 51·60 61·70 61-70 71-80 71·80 81-90 81·90 91-100 91-100 101-110 101·110 111-120 111·120 121-130 121·130 131·140 131-140 141-150 141-150 Green Sunfish Mottled Sculpin 151· 151· ~ 4122 Sampling Datea 8118-19 7/22-28 Total 8111-18 L1~1" 4122 Sampling D•tn 7122. 26 6116·19 8111·18 1-10 1-10 884 370 11·20 11-20 341 4389 21-30 21-30 25 31-40 31·40 27 41-50 387 30 145 41-50 29 761 51-80 51-60 13 300 61·70 61·70 71-80 71-80 81-90 81-90 91-100 91-100 101-110 101-110 111-120 111-120 121-130 121-130 131·140 131·140 141-150 141·150 Plains Killifish 151· 68 Red Shiner 151· Figure C-3. Length-frequency distributions for green sunfish, mottled sculpin, plains killifish, and red shiner; Yampa River study area, 1981. �35 Toto I L1!R!" 4122 Sampling DatH 6/16-1g 7/22 •26 Total 8111-18 lr::.~,h 4122 Sampling Oatea 11116-19 7122. 26 8111·18 1·10 1·10 11-20 11·20 22 227 21·30 21·30 15 554 31-40 31-40 41-50 41·50 51-60 51·60 61-70 61-70 71-80 71·80 81·90 81-90 91-100 91·100 101-110 101·110 111-120 111 ·120 121-130 121-130 131-140 131·140 141·150 127 13 4 26 ,. 141-150 Creek Chub Fathead Minnow 151· 151· ~ Tol•I l•::.R!h 4122 Sampling D•toa 6/16·19 7122 -26 Sampling Dates 21-30 28 7/22·26 Bt11·18 22 11·20 631 102 15 21-30 493 1302 23 31-40 41 2'64 38 41-50 8 112 23 51-60 4 138 4 14 22 19 9 18 20 31-40 41-50 6116-19 1·10 1-10 11-20 4•22 8/11-18 4 51-60 375 61•70 61-70 71-80 71-80 81-90 81-90 15 91-100 91-100 6 101-110 101-110 111-120 111-120 121-130 121-130 131-140 131-140 141-150 141-150 Flannelmouth Sucker 151- Gila spp. 151- Figure C-4. Length-frequency distributions for redside shiner, sand shiner, speckled dace, and white sucker; Yampa River study area, 1981. �S•mpli119 Datu 4121-512 51211-28 11/2-4 8119-27 1/8-10 1121-31 -------------------------·~ Total Length mm1 S•mphng D•tH 4/27. 5/2 51211. 28 8/2. 4 6/19 - 27 1 II 10 1·10 1-10 11- 1121-3 31-40 4 21-3 20 4 31-4 88 41-50 41-50 81-90 91-100 101-110 111-120 121 ·13 131-14 141-15 Channel Catfish Carp 151• 1'27·31 151- Figure C-5. Length-frequency distributions for black bullhead, bluehead sucker, carp, and channel catfish; Colorado River study area, 1981. �37 C/27-512 51211-28 Total S•.,.pling Dai.. 1112-• 81111•17 7127-3t 1-10 L•.::J.'" C/27· 512 5128- 28 Samph"ll O•tu 812-• 8119. 27 1-10 20 21-3 338 332 31-C 208 ••2 '1-50 10• 102 25 51-8 01·7 81-7 71-8 71- 118·10 7117·31 318 1'5 002 28 300 11178 1'1 ~o 100 12 116 lU 12 ·~ U8 18 37 81-110 111-100 101-11 131-1' 1'1- 15 Fathead Minnow Colorado Squawfish 151- 151- Total L•,::R,'" C/27- 5/1 5/28- 28 Sampling OatH 812 - ' 81111- 27 7/8-10 7127•31 C/27• 5/2 5/28- 28 Sampling DalH 1112-• 111111-27 ' 8 11 II 51·60 81-7 71· 8 71-8 81-90 81·90 111-100 91-100 101-11 101-110 111-12 111-12 121·13 121-13 131- 1C 131 ·1' 151- 1150 200 Flannelmouth Sucker 311 17 13 '1·50 141·15 7127-31 93 21-3 31·• 118-10 1-10 1-10 11· Tol•I Length mml ' " -15 111 181 33 308 ' 133 " 15 7 18 8 8 Gila spp. '51- Figure C-6. Length-frequency distributions for Colorado squawfish, fathead minnow, flannelmouth sucker, and Gila spp.; Colorado River study area, 1981. �38 1olal •·.::i"' I 4127 5.·2 :01H 28 Ill Samphn9 Uatn 8/2 4 81111 21 sa .. phn9 o., •• ,. 118 10 Cl/2. 1:21 31 c CIJtg. 27 7!8 10 7!27·31 1 10 011 II 2 ICI :n-3 20 11·2 31. c 41·50 4 4 81-7 11-8 81·90 81·110 111-100 111-100 101·1'0 11'. 120 121·13 131·14 141 - 15 141·15 Largemouth Bass Green &.tnfish 151- 101- Total L•:J,lh 4m-512 5128-18 Sampling DatH 812-4 81111-27 7/8·10 ------------------------r-11 Total L•.::R,th 4/27-512 5128-28 Sampling Da1u 812·4 81111·27 718·10 1-10 1·10 51·80 81-7 71-8 81-90 91-100 101. 110 111-12 121-13 121 ·13 131 - ,.. 131 ·14 141·15 141 -15 Mottled Sculpin 151- Plains Killifish 151- Figure C-7. Length-frequency distributions for green sunfish, largemouth bass, mottled sculpin, and plains killifish; Colorado River study area, 1981. 112i �39 Tatel L•.::~1h .,27. 512 5/28·28 S•mpllng D•IH 0/2·4 01111··21 45 1·10 51• 11&· 10 322 53 13511 1254 48 685 155 53 130 430 81 141 133 37 20 75 3 20 7127·31 3115 Tol•I L•,::!_lh 5128·20 S•mphng D•IH 812· 4 01111·27 7/11 ·IO 7'27·31 1·10 8855 '1·2 114 21 3 178 487 31 4 85 1105 •1 404 51·8 58 81 · 7 22 11·8 3 4/27· 512 01-10 141-15 141. 15 Redside Shiner Red Shiner 151· 151 • Tol•I Len91h mm) 4127•5/2 5120•28 S•mpllt1g D•tea 812-4 8119-21 718 ·10 7127·31 211 1·10 22 Total L•n9lh mm) 4127 • 512 5128. 28 S•mph119 Dlln 612·4 8119·27 118. 10 7127·31 1 ·10 1132 11•2 115 58 98 21·3 4 73 11-2 8 21-3 82 34 31-40 117 154 20 9 31·40 22 3 12 41-50 18 235 8 72 41•50 27 28 10 51-0 10 51 811 51·80 01 ·7 3 0 9 12 20 01·7 11-8 71· 8 81 ·110 81·110 111-100 01·100 101·11 101·110 111-120 111·120 121·13 131·14 Sand Shiner 141·15 Speckled Dace 151. Figure C-8. Length-frequency distributions for red shiner, redside shiner, sand shiner, and speckled dace; Colorado River study area, 1981. �40 Tol•I ......, Length 4127- 512 5129- 28 S•111pllng D•t•a 812- 4 8119-27 7'8-•0 7127-31 1-10 81•90 91·100 101-110 111-120 White Sucker 151 • Figure C-9. Length-frequency distribution for white sucker; Colorado River study area, 1981. �41 APPENDIX D: MORPHOMERISTICS, GILA CYPHA Means and ranges of selected morphomeristic values, expressed as a percent of the standard length, for hatchery-reared humpback chub larvae and juveniles. Known-age series of Little Colorado River, Arizona (Table D-1) and Colorado River (Black Rocks), Colorado (Table D-2) progeny were provided by Willow Beach National Fish Hatchery. Superscripts accompanying mean values represent the number of specimens from which the value was derived, if less than N in the column heading. �42 Size mm SL (AS to PHP) mm TL Lengths. Anterior Margin of Snout t3±1 AE 8±3 PE 6+1 (PE-AE) OPl 18±1 OP2 (OP2-0P1) OD ID (ID-OD) PV 67±3 OA IA (IA-OA) (PHP-OA) AFC PC 106±1 F in Lengt h s: g±3 Pl P2 DAI: Fin Lengths (including caudal) Fin Rav Lengths: 27.4-43;7 ,\ [I): 2-4 5-10 3-7 15-20 -3.± 1 10±2 7±1 21±2 2-5 8-16 5-12 18-26 63-76 _68±2 64-71 109 17±1 103-109 110±3 4-14 l 13±1 106-113 104-115 11-16 -· -· c- D - APl P2 - 4-6 12-14 7_g 24-28 47-52 20-24 51-55 65-68 12-15 64-69 61-6_9_ 70-79 9-12 32-35 112-115 132-152 6±1 13±1 7±0 26±1 ~8± 1 22±2 51± 1 66±1 14± 1 64±1 62±1 77±1 12±1 35±1 113±1 128±2 5-7 11-14 6-8 23-28 45-48 20-25 49-53 64-67 17±2 23±2 18+2 13-19 9-18 20-25 15-20 18±1 15±1 24±1 20±1 17-21 13-17 20-26 18-24 100±11 82-118 105±4 96-115 16±2 19+2 12-19 15-22 17±1 21±1 16-20 17-22 12-16 62-67 63'."'f)_7_ 75-80 10-15 33-37 110-115 124-131 13±1 12±1 14±1 11±1 8±1 5±1 12-16 11-14 13-18 9-14 7-13 2-7 17±1 15±0 23+1 21±3 15+2 8±0 16-18 15-16 20-25 16-27 12-18 7-9 16±1 14±1 24±1 26±2 18±1 8±0 14-19 12-16 22-26 22-30 16-19 7-8 14±1 11±1 6±1 5±1 _2±0 12-17 8-1-5___ . ..2.9±1 20±2 16+3 5-10 12±1 4-7 5+1 2-4 19-21 14-22 13-21 12-15 4-6 18±1 21±1 20±1 14±1 - 4±1 -- 16-21 19-23 18-24 13-15 3-6 ·- - - 5±0 13±0 8±0 27+1 49±1 21±1 53±1 66±1 13±1 66+2 6_6±2 77±2 11±1 34+1 113+1 130±1 .. 13±3 LAR LDR Bodv Depths at: BPE 13±2 11-17 ME 12±2 9-16 OP! 12-29 19±5 OD 12-23 19±5 [i+l BPV 6-13 AMPM 3±1 1-3 Bodv Widths at: BPE 12±2 9-16 OPl 9±1 8-13 8+3 OD 5-14 BPV 4-13 6±2 AMPM 1±0 1-2 Calculated Ratios: Pl Length f (AS to OP2-AS to OPl) P2 Length f (AS to OP2-AS to OPl) AS to AFC f AS to PC Oblique Measure: OA to PHP f POP to OA Ml'Vomeres (V erte b rae i n J uveni les ) to: OP2 OD PV 12q±1 28-30 PV to MPM 18±1 16-18 Total 46±1 45-47 F'in Ravs (P r i nc i .p 1e Fi n Ravs 0 n.v 1 ): - 10 17 ±1 29±1 _17±1 46±1 - 8-10 27-31 16-12 45-48 ~±Q 4-6 3±1 2-4 4+0 5+1 3-5 5-7 3±0 3±0 2-4 3-4 5+1 4-6 3±0 3-4 16+1 lg±o 28±0 17+0 45±1 15-17 18-20 28-29 16-17 44-46 1710±1 19 10 ±1 28 10 ±0 19 1 u±o 47 10 ±1 16-18 18-19 28-29 19-20 47-48 19±0 9±0 10±0 161;±1 gs±o 19-19 9-9 9-10 14-16 9-9 - 19±0 9±0 10±0 16±1 9±0 19-19 . 9-10 9-10 14-16 9-9 �43 Table D-2. Gila cypha (Black Rocks strain) nun SL (AS to PHP) nun TL Len2ths. Anterior Margin of Snout 3±1 AE 9±1 PE (PE-AE) 6±1 18+2 OPl OP2 (OP2-0P1) OD - Range 21.2-43.0 Size - ID - (ID-OD) PV OA IA (IA-OA) (PHP-OA) AFC PC Fin Len tbs: 65±2 1nc;±1 AS t»: 2-6 8-11 4-7 13-20 _4±1 _1_0± 1 7±1 21±2 45 2 ±0 24"±1 50 .. ±1 62'*±2 12 .. ±1 66±2 61-68 .66'+±2 74'+±2 9'*±1 34'+±2 110.l.l:!:l 1n?-tflJ 1na±4 . 2-5 8-12 5-8 18-27 45-46 23-25 50-53 60-65 11-12 62-70 66-F.P. 73-74 8-10 J2-J5 -102-112 1nt.-11Q 5±1 12± 1 7±0 26+1 47±1 21± 1 51±2 64±2 13±1 6F.±1 6'l±l 7F.± 1 11±1 1c;±1 111+1 1?t..±1 5-6 .. 11-14 6-8 -25-28 45-48 19-23 48-53 62-67 12-1 'l i:;.c;_,;8 F.4-1.P. 7!.-77 Q-11 ':t?-17 111-ll'l 11 A-128 - 6±0 12:!; 1 6± l 25± 1 411± 1 21± 1 4q±2 63± 1 14± l F.4±1 64±1 7'l±l 11±1 1F.±1 111± 1 12,;±2 5-7 10-14 5-8 21-28 42-48 18-25 44-55 61-67 11-16 62-67 61-6h 72-7P. 8-13 14-17 111-114 121-110 ~P~l~-------------------------1-o.;;..;..o,...._____....._...__--1.l~.±~l,._____~l-O_-~l6..__..f..-"-=-..._---.........t=.11..L....--~u....,......;.....____~;;;.4 ____________________________________________.,.....6. ±1 --~P2....;.._- 5-7 4 --~D-----------------------------+------------------i-l~S~4=±~3--_...1~2-~1~8'"--'-"..l.l::."'---~i.z=..jo..L...---~wi.=~----~=..a...1..-A- . 11 ±1 10-12 Bodv De ths at: BPE ME OPl OD BPV AMPM Bod Widths at: BPE OPl OD BPV AMPM Calculated Ratios: Pl Length + (AS to OP2-AS to OPl) P2 Length + (AS to OP2-AS to OPl) AS to AFC + AS to PC Oblique Measure: --~OA;.;;._;t~o_..;;;..P~HP;;;__+___;;;,P~O~P_t~o;,..._O_A_-__________________._J~4~±~1:,..___~6~-~8~---...&..::5=±~1----...:!.4-~6='-----~~3~±~1---------=2~-5::;._____ Mvomeres ( Vertebrae in J uveni 1 es ) to: 16+0 OP2 16 2 +0 11 10 ±1 16-18 15-17 16-16 19'+±1 OD 19+1 19 10 ±1 18-20 17-20 17-19 PV 29+1 29±1 29 10 ±0 28-29 29±1 28-30 28-30 28-30 18 1 u±o 17+1 17+1 17±1 PV to MPM 17-18 17-19 16-19 16-19 47+1 47+1 Total 47 10 ±1 46-48 46±1 45-48 45-48 45-48 1 ): Fi n Ravs (P rinciple Fin Ravs 0 n.y 19±0 19±0 19-19 c19-19 9.l ·9-10 9±0 D9±0 9-10 A10±1 10±1 8-10 9-10 16+0 Pl 15-17 P2 9-9 9 2 ±0 9±0 9-9 - ��45 JOB PROGRESS State of COLORADO Proj ec t No. _.;:_S.:;:E_--=.3_-_4 Work Plan No. Job Title: Period Personnel: II -~~--------------_ Nesting Covered: _ Performance March REPORT Endangered Wildlife Investigations Job No. 1 of Peregrine ~--------------- Falcons in Colorado 1, 1981- June 30, 1982 E. Bauer, D. Berger, M. Berman, E. Bowden, G. Craig, B. Grebence, R. Meese, S. Munsell, J. Rucks, and H. Stolzenburg, Colorado Division of Wildlife; J. Enderson, the Colorado College; J. Hogan and S. Petersburg, National Park Service. ABSTRACT In the 1981 and 1982 breeding seasons, nest site occupancy stabilized and augmentation efforts resulted in fledging rates of 2.50 and 2.86 young per active pair, respectively. Eggshell thickness levels improved from 16 percent thin in 1980 to approximately 10 percent in 1982. Pesticide residues did not decrease significantly, but extremely high levels were not encountered either. Photographs of adults at breeding sites permitted identification of individuals and yielded a survivorship of .76 based upon territory occupancy. This Job Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the Director. ��47 NESTING PERFORMANCE OF PEREGRINE FALCONS IN COLORADO Gerald R. Craig P. N. OBJECTIVE The objectives of this study is to annually monitor the breeding numbers and reproduction of Colorado peregrine falcons in an effort to document further population declines as well as eventually record the population's response to various recovery efforts which are implemented. Additionally, health of the population may be monitored indirectly by analyzing pesticide residue levels in the falcons' eggs and principal prey they feed upon. Information obtained from these investigations will be made available through annual reports to the Rocky Mountain/Southwest Peregrine Falcon Recovery Team as well as cooperating agencies to aid in evaluation of various recovery efforts. SEGMENT OBJECTIVE 1. Annually monitor the number of breeding reproduction in Colorado. 2. Annually monitor peregrines. 3. Monitor recruitment breeding population 4. Compile data and submit reports to appropriate state and federal personnel and the Rocky Mountain/Southwest Peregrine Falcon Recovery Team for use in evaluating recovery efforts. Note organochlorine pairs of peregrines pesticide of reintroduced of Colorado. and levels in wild breeding peregrines into the wild These objectives correspond to tasks 1113., 1114.,212.,221., 3211., 3212., and 333. of the approved American Peregrine Falcon Recovery Plan (Rocky Mountain/Southwest Populations). METHODS AND MATERIALS Methods and Materials for this study have been described (Craig and Enderson 1981). previously RESULTS AND DISCUSSION Territory Occupancy and Productivity In 1981, breeding territories were occupied by 7 adult pairs while 2 other territories were frequented by lone adults. A mixed pair consisting �48 of an adult female and yearling male occupied 1 hack site while another hack site continued to be utilized by a lone adult male release there in 1978. Six of the adult pairs successfully bred and all were manipulated to augment reproduction under Job 2 (Reintroduction and Augmentation of Peregrine Falcon Production). The pairs produced 28 eggs, 21 young were returned to them for rearing and 15 young successfully fledged (2.50 young per active/successful pair). In 1982, breeding territories were occupied by 8 adult pairs while an adult pair occupied another hack site. Three territories were occupied by lone adults (one adult male, one yearling male and one adult female). Seven of the adult pairs laid eggs and were manipulated to augment reproduction under Job 2. A total of 26 eggs were removed, 26 young were returned and 20 young were successfully fledged by 6 pairs (3.33 young per successful pair and 2.86 young per active pair). The site that was unsuccessful failed after 4 young were returned to the pair. When the site was revisited 9 days later no sign could be found of young or adults. It is possible golden eagle predation was the cause of failure although no peregrine remains were found in an active golden eagle nest in the vicinity. Although rate of occupancy remained low (Table 1) some evidence of success at release (hack) sites was evidenced by the presence of a lone adult male at 1 site and a mixed pair (adult female and yearling male) at another site in 1981. In 1982, an adult pair frequented 1 hack site but did not breed and an adult male which was released at another site in 1980 attracted a wild adult female from an adjacent site and successfully fledged 4 young. For purposes of the program, falcons which return to breed at release sites will be regarded as wild breeding. Eggshell Thickness Although the 1980 eggshell samples did not vary significantly from the 1973-79 thickness values which averaged 16% thinner than pre DDT era eggs, the 1981 samples were significantly thicker (11.7% thin) and the 1982 samples improved to 10% thin. Several investigators (Newton 1979, Ratcliffe 1980) believe that 10% thinning is the point above which a population can sustain itself. Several more years of monitoring will be necessary to establish the true nature of the shell thickness trend. Organochlorine Residues in Egg Contents DDE residue levels from 9 wild eggs collected in 1980 averaged (arithmatically) 13.6 ppm fresh, wet weight (range: 10.7 - 19.7 ppm) while 8 eggs in 1981 averaged 13.8 ppm (range: 9.27 - 20.4 ppm). The 1980 and 1981 averages were below the 1973-79 arithmatic means which ranged from 17.5 to 26.3 ppm. It is also interesting to note that for the first time since samples were collected in 1973, extremely high levels (greater than 21 ppm) were not present in 1980 or 1981 eggs. While the trend in DDE residues appears to be downward, the small sample size does not �49 Table 1. Occupancy and productivity of Colorado peregrine 1972-1982. eyries, Year 1972 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 Eyries visited 15 23 24 26 27 31 32 33 34 34 34 Occupied 11 12 9 8 8 12 11 12 13 11 11 Adult pairs 8 11 7 6 5 11 7 6 8 7 8 Immature 0 0 0 0 2 0 2 2 3 1 0 3 1 2 1 1 1 2 3 2 3 3 1 5 2 4 6 5 4 5 6 6 No. yng. fledged No. yng. augmented 2/0 11/0 5/0 6/4 11/5 16/11 12/8 16/16 15/21 20/26 Young Fledged/ adul t pair.Y 0.18 1/67 0.83 1/20 1.00 2/29 2.00 2.00 2.14 2.50 Young Fledged/ successful pair 2.00 2.20 2.50 1.50 1.83 3.20 3.00 3.20 2.50 3.33 eyries pairs!! Lone adults Successful pairs Percent of sites occupied1! 73% 52% 38% 31% 30% 39% 34% 36% 38% 32% 32% Percent of sites w/adult pairs 53% 49% 29% 27% 26% 35% 22% 18% 24% 21% 24% 1/ At least one member 1/ 1.25 young fledged per pair is considered 1/ of the pair was in juvenile plumage. normal reproduction. 80%-90% of the eyrie sites should normally be occupied year. in any particular �50 permit statistical verification. It is hoped that results from the 6 eggs obtained in the 1982 season but not yet analyzed will verify the trend. Photographic Identification of Adults Between 1980 and 1982, 30 useable photographs were obtained of individual adult peregrines at 9 breeding territories. In 17 cases, photographs permitted comparison to determine if the same individuals returned on subsequent years. Among the 17 cases, 13 represented adults which had been present the preceeding year. An annual adult survivorship rate of .76 can then be calculated if it is assumed that all adults return to the same breeding territory so long as they live. This survivorship is remarkably similar to the rate obtained from band recoveries. Cessation of territory abandonment in 1982, shell thickness improvement, absence of extremely high pesticide residues in egg contents, and reoccupancy of 2 historic sites by released adult pairs and presence of released lone adults at 2 other localities are all factors which are indicative of improvement in Coloradots peregrine population. Time will tell if this trend will hold. LITERATURE CITED Craig, G. R. and J. H. Enderson. 1981. Nesting performance of peregrine falcons in Colorado. Colo. Div. Wildl. Wildl. Res. Rep. January (1): 13-23. Newton, I. 1979. S.D. 399p. Population Ratcliffe, D. 1980. S.D. 416p. Prepared ecology'of The peregrine by Wildlife Researcher C raptors. Buteo Books, Vermillion, falcon. Buteo Books, Vermillion, �51 JOB PROGRESS COLORADO State of Project No. Job Title: II Job No. Reintroduction Covered: Personnel: Endangered SE-3-4 Work Plan No. Period REPORT March and Augmentation Wildlife Investigations 2 of Peregrine Falcon Production 1, 1981 - June 30, 1982 J. Enderson, the Colorado College; E. Bauer, D. Berger, M. Berman, E. Bowden, G. Craig, B. Grebence, D. Langlois, R. Meese, J. Rucks, H. Stolzenburg, and T. Washington. Colorado Division of Wildlife; J. Ferguson, and D. }lcVean, Bureau of Land Management; W. Burnham, W. Heinrich, D. Konkel, C. Sandfort, and S. Sherrod, the Peregrine Fund, Inc.; E. Freienmuth, the North American Peregrine Foundation; J. Hogan and S. Petersburg, National Park Service. ABSTRACT All wild breeding peregrines encountered in Colorado during the 1981 and 1982 seasons were manipulated to increase fledging success to 2.50 and 2.86 young per active pair, respectively. Thirteen captive hatched young were released at 4 hack sites and all achieved independence (3.25 young per site). Release of captive held and conditioned adult falcons was not successful either season and will be discontinued. This Job Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the Director. ��53 REINTRODUCTION AND AUGMENTATION OF PEREGRINE FALCON PRODUCTION Gerald R. Craig P. N. OBJECTIVE The objective of this program is to sustain the wild breeding peregrine falcon population in Colorado through augmentation of poor natural production and reestablishment of breeding pairs at vacant sites by release of captive produced falcons. SEGMENT OBJECTIVES 1. Augment poor wild production by placement of captive hatched wild young and captive produced young into occupied wild nests. 2. Release captive hatched wild young and captive produced young to the wild through "hacking" at potential and abandoned wild nests. 3. Develop and implement release of adult falcons at potential or abandoned nest sites and at wild nests occupied by lone adults. 4. Monitor results of the efforts, compile data and submit reports to appropriate state and federal agencies and the Rocky Mountain/ Southwest Peregrine Falcon Recovery Team. Note These objectives correspond to jobs 222., 3133., 321., 322., 331., 332., and 3211. in the approved American Peregrine Falcon Recovery Plan (Rocky Mountain/Southwest Population). METHODS AND MATERIALS la. Breeding pairs of peregrines will be kept under surveillance to determine initiation of egg laying. Shortly after completion of the clutch, the eyrie site will be visited and all the eggs removed and replaced with dummy eggs which the adults will be permitted to incubate. Since the wild eggs usually are thin-shelled they will be artificially incubated to avoid their being accidentally crushed by the adult. After several weeks, the site will be revisited and young peregrines will be exchanged for the artificial eggs. Up to four young may be placed at each site to assure that the maximum number of young are fledged. lb. On occasion, wild breeding peregrines will deposit their eggs in inferior eyrie sites. If the site cannot be improved to assure that young will successfully fledge, it will be necessary to relocate the pair. Relocation will be undertaken no later than 10 days after completion of the full clutch of eggs. Prior to that time, the eyrie will be �54 visited and all the eggs removed and artificially incubated. Approximately two weeks after removal of the eggs, the pair will relocate to another (hopefully superior) nest ledge and will recycle to lay a second clutch of eggs. The second clutch will be replaced with dummy eggs and the site will be treated as outlined in lao Every effort will be made to return the young hatched from the first and second clutches to wild eyries with comparably aged young. In some cases, it may be necessary to include odd aged wild young with broods to be released through other techniques such as hacking (see 2.). 2. Captive produced young may be released at unoccupied or potential sites without the benefit of protection or care from adults through the technique of "hacking." Young falcons of 3 to 4 weeks of age will be placed on a suitable ledge at a potential reintroduction cliff site. They will then be cared for and fed by human attendants until they are flying and capable of feeding themselves. In this manner, the young falcons will return to the site at which they were reared and hopefully breed. This approach requires constant attendance and observation in order to protect the vulnerable young and insure they have sufficient food while they are in the hack box. Because of this, the first technique will receive priority attention. If there are no additional adult breeding pairs as required by approach 1. then young will be placed into the wild using this technique. 3a. Young falcons that are too late to be hacked in the regular program and expected to survive on their own will be hacked for two weeks and retrapped. All birds will be kept together as pairs (male and female). The members of a pair will be flown together using falconry techniques. The birds will learn to hunt at this time and will become accustomed to being in the air with another falcon. A Research Associate employed by the Peregrine Fund and stationed at Ft. Collins will provide the nucleus for the conditioning program as well as the release program. He will fly three to four pairs of falcons together and prepare them for future release. If it becomes necessary to return more falcons from hack sites, a few select falconers will be asked to cooperate by flying a pair of falcons that can be recalled at any time. All falcons that are flown will be equipped with telemetry. 3b. Release of mates for lone wild adults: Wildlife Technicians will check wild eyries in early March for signs of occupancy by lone adults. The Research Associate will then construct a barred cage to fit the physical situation (approx. 1.5 m cube). An adult from one of the conditioned pairs and of the opposite sex will be placed in the cage and fed through a chute. Food will also be deposited via a chute to a shelf perch outside the cage in order to appear as a food transfer to the wild bird. A loud speaker mounted on the cage will broadcast courtship vocalizations. After 1 to 2 weeks of this treatment or after behavior indicates that the proper time has arrived, the caged bird will be released and followed with �55 telemetry. If necessary, food will continue to be provided until the pair is functioning together. Continual observation will be carried out by an attendant provided by the Peregrine Fund. 3c. Establishment of adult pairs. A small loft 2.4 m cube) will be constructed below or on top of an unoccupied eyrie. An extended window in the loft will allow maximum visual exposure to the landmarks of the area. A pair of conditioned, adult falcons will be flo,vu at the site and served with prey. At night they will be housed in the loft. After the pair of birds has become familiar with the area and begins exhibiting courtship behavior, the researcher will gradually allow them more and more freedom until they are finally left to themselves. This progress will be followed by telemetry. RESULTS AND DISCUSSION Augmentation Efforts All breeding peregrine falcons encourtered in the wild during the 1981 and 82 seasons were manipulated to increase productivity. The recycle effort was nearly eliminated since increased captive production at the Peregrine Fund's Fort Collins facility reduced the need to supplement production with wild eggs. A single site was recycled in 1981 because the pair nested 10 to 14 days early and had to be brought into synchrony with the captive population in order for young to be available for them. This pair produced 4 eggs in the first clutch and replaced them with a second 4 egg clutch when recycled. In 1981, 26 eggs were removed from 6 breeding pairs and were subsequently replaced with 23 captive hatched nestlings (3.83 young per pair). The 6 pairs successfully reared and fledged 15 young (2.50 young per pair). Despite losses of 1 member of the pair at 2 sites, the remaining adults successfully reared young. A single male reared 3 of 4 young after the female apparently was killed by a great horned owl, and a lone female reared 1 young out of a brood of 4 after the male disappeared of unknown causes. Remains of young found below the eyrie indicate they may have starved. Twenty six young (3.71 young per pair) were fostered into 7 eyries sites in 1982, with 20 young (2.86 young per pair) successfully fledged. An entire brood and both adults disappeared within 9 days after the young were placed in the eyrie. Although no evidence was present, it is speculated that golden eagle predation removed one or both adults and the young starved. At a second site, another young disappeared during the rearing period and golden eagle predation accounted for the death of another young at a third site. Fledging rates of 2.50 and 2.86 young per pair were significantly above the average of 1.25 generally accepted as necessary for a self-sustaining population. �56 Hacking Efforts In the 1981 season, 4 hack sites were operated and all 13 young (3.25 young per site) which were placed in the hack boxes successfully reached independence. A mixed pair, consisting of an adult female and a yearling male from a previous release, frequented one release site but did not interfere with the young. The lone adult male which was released at another site in 1978 returned for the third consecutive year and although a yearling female was observed with him on one occasion, no further interaction was noted. Adult Release In 1981, two territories occupied by lone adult females were selected for release of conditioned adult males to establish breeding pairs. The release attempt was discontinued at one site when a wild adult male appeared on the scene before the conditioned falcon could be released. The second release could not be initiated because the falcon intended for release was injured when it received a severe electrical shock from a power pole. Finally, release of a conditioned pair was not undertaken in 1981 since a compatible captive pair had not been established by release time. Adult release efforts in 1982 did not fare any better. Steve Sherrod, the Research Associate for the Peregrine Fund, was responsible for the project and provided the following observations and recommendations: "After two seasons of attempting to release adult peregrines at Colorado nest sites, several considerations have become apparent. "It is difficult to determine whether or not a lone adult will obtain a mate and how long one should wait before an attempt is made to supply a mate. "It is necessary to have qualified observers continually watching the nest site from dawn til dark for a period of about a week prior to any type of attempted release activities. In this way positive identification of the falcons can be made and their daily routine can be determined. "During the previous season, attempts to release a mate for a reportedly unmated wild female were thwarted when the wild bird's mate was observed copulating with her, but only after all the paraphernalia for the release had been set into position. During this season, the lone male at Site #1 was only one year old, and observers at Site #2 speculated that their bird may have been only one year one. It also appeared that the male at Site 1f3 was only a year old, and the female there was definitely only a one year old. Most peregrines breed at three years of age. Of the few birds that do breed at two, females appear more likely to breed than males. It is futile to �57 attempt releasing a two year old bird as a mate for a one year old wild bird. If the wild bird, however, were an adult which would court the captive bird, then a pair might be expected to develop especially if the two year old captive bird were a female. Similarly it is highly unlikely that two, two year old birds could be released and expected to breed. This was especially true of the male which was available this year. He vocalized during his initial flights with females, but afterwards showed no interest and soared up out of sight. A two year old male released for an adult female in the eastern U.S. this year, stayed around the nest site but did not court the female apparently because he was too young. The problem of age, however, can eventually be overcome by time, at least for the captive birds. "The task of maintaining the falcons in captivity throughout the year is very demanding. This is true especially because of the daily conditioning which is necessary if the birds are expected to survive. It is next to physically impossible for a single handler to fly more than three or four falcons on .a daily basis. Several times that many peregrines are required in order to obtain enough particular individuals which will function well in such a release program. Efforts to increase the number of peregrines which can be conditioned for release by loaning pairs to handlers has met with little success. Loaned birds in most cases have been lost, electrocuted, or shot. Of the few which have been returned several were imprinted on humans and thus of no value to the program. "In summary, there are many problems involved in this type of release program, and the program does not appear to be cost effective. For these reasons, it is suggested that the program be discontinued." Based upon these recommendations, it is likely that this particular method of release will be discontinued. Prepared by Gerald R. Craig Wildlife Researcher C ��59 JOB PROGRESS State of COLORADO Project No. SE-3-4 Work Plan No. Job Title: II Peregrine Period Covered: Personnel: REPORT March Endangered Wildlife Job No. 3 Investigations Falcon Captive Maintenance 1, 1981 - June 30, 1982 W. Burnham, D. Konkel, C. Sandfort, J. Hoolihan, D. Bay, M. Rider, The Peregrine Fund, Inc., G. Craig, Colorado Division of Wildlife. ABSTRACT In 1981-82, 35 adult anatum peregrine falcons were maintained at the Peregrine Fund, Inc's facilities at Fort Collins, Colorado. Pairs were separated and provided with new mates in an attempt to correct certain behavioral problems and promote copulation. No losses or injury were incurred during the contract period. All falcons were maintained in robust condition and good health. The security of the facility was not violated and no unusual events occurred. This Job Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the Director. ��61 PEREGRINE William FALCON CAPTIVE MAINTENANCE Burnham and Gerald R. Craig P. N. OBJECTIVE The objective of this program is to maintain a colony of adult peregrine falcons (Falco peregrinus ana tum) which is genetically representative of the wild population, free of disease and capable of reproductive activity. SEGMENT OBJECTIVES 1. Annually contract with The Peregrine Fund, Inc. to maintain adult anatum peregrine falcon in captivity. 2. Prepare an annual report of maintenance contract. METHODS efforts associated 35 with the AND MATERIALS Methods and materials for this study have been described (Burnham and Craig 1981). previously RESULTS AND DISCUSSION In 1981 and 1982, 35 adult anatum peregrine falcons were maintained at the Fort Collins, Colorado facility of the Peregrine Fund, Inc. All falcons were fed a daily diet of Coturnix quail and 5 to 6 week old cockerels. No adverse health conditions or disease problems were encountered and no losses or injury were incurred during the period. Seven pairs were separated and given new mates in order to improve their compatibility or in order to permit older, experienced individuals to teach young, inexperienced mates. When limited aggression occurred on several occasions, certain recombinations were made and the aggression subsided. The security of the facility was not compromised nor did unusual events occur during this period. LITERATURE CITED Burnham, W. and G. R. Craig. 1981. Peregrine Colo. Div. Wildl. Wildl. Res. Rep. January Prepared by ~C;~~~~~~~~~~ _ Wildlife Craig Researcher falcon captive maintenance. (1): 34-37. ��63 JOB PROGRESS REPORT State of Project No. COLORADO SE-3-4 ~~--~--------------- Endangered Wildlife Investigations Work Plan No. II Endangered Birds Job Title Development of a Preservation Program for Insular Job No. 6 Populations of Prairie Grouse Period Covered: Personnel: 1 March 1981 - 30 November 1981 L. Cordova, T. Olson, G. C. Miller, P. Svoboda, R. Calderon, W. D. Graul. ABSTRACT Research begun under this Federal Aid Project was continued under Project ~v-145-R as of 1 December 1981. All work conducted under this segment has been reported under Project 145-R (p. 65). This Job Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the Director. ��65 JOB PROGRESS State of COLORADO ~~~~---------------- Project No. ~FW~-~1~4~5_-~R~ Work Plan No. I Job Title Development _ Period Covered: Prairie Grouse Investigations 1 Job No. of a Preservation Populations Personnel: REPORT 1 December Program of Prairie for Insular Grouse 1981 - 30 June 1982 R. Calderon, W. D. Graul, S. Lustig, B. Van Santo G. C. Miller, D. Opperman, ABSTRACT This project is a continuation of work begun under Federal Aid Project SE-3. The objectives and design of the study remained the s~me. Insular populations of lesser prairie-chickens (Tympanuchus pallidicinctus) and greater prairie-chickens (T. cupido pinnatus) were investigated and evaluated for their suitability of study populations. The greater prairie-chicken population south of u.S. Route 34 and north of the Arikaree River was selected for further study. This Job Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the Director. ��67 DEVELOPMENT OF A PRESERVATION PROGRAM FOR INSULAR POPULATIONS OF PRAIRIE GROUSE G. C. Miller P. N. OBJECTIVES 1. By 1986, ascertain for 1 species of prairie grouse in Colorado the combination(s) of habitat island size, condition, inter-island distance, and number (by sex and age class) of grouse needed to establish and maintain populations with a persistence probability of P > 0.5 and extinction time of T > 100 years. SEG}lliNTOBJECTIVES 1. By July, 1982 locate, select and describe occupied (by pr aa r i.e grouse) and unoccupied prairie habitat blocks ("islands") most suitable for study of the biogeography and other aspects of prairie grouse ecology. 2. By July, 1982, ascertain the nature of occupancy of selected prairie islands by grouse and, where applicable, an index of abundance. 3. Begin to ascertain areas. 4. Publish research population characteristics of grouse in selected results. METHODS AND MATERIALS Potential study areas were surveyed for presence of grouse and cover mapped. Areas of 10 or more sections that did not contain leks and in which winter flocks were not known to occur were classified as unoccupied by grouse for the purposes of this segment. Leks were located between March and June by driving predetermined routes (generally 10 to 16 km in length), beginning no earlier than 1 h before sunrise and ending no later than 1 h after sunrise. Stops of 3 minutes duration were made along the route to listen for sounds made by displaying grouse and triangulate their locations. Stops were about 0.8 km (0.5 mi) apart; each route was traversed twice in a morning, listening at alternate stops on the first trip and the remaining stops on the return trip. Precise lek locations were ascertained later in the day, using the triangulations to locate the general vicinity and then searching for evidence of the lek (droppings, tracks, feathers, and grouse). Areas were cover mapped in several stages. First, areas were classified by general vegetation type (cropland vs. rangeland); rangelands were further classified with respect to dominant vegetation appearance--shortgrass, midgrass, tallgrass, and/or shrubs. Vegetation on randomly selected plots �68 was measured with a modification of Graul's (1978) technique. Vegetation sampling on the South Platte Wildlife Management Area continued in the fashion described previously (Miller 1981). The potential of remote sensing for detecting vegetation differences in rangeland types was evaluated through analysis of LANDSAT-generated data and vegetation data collected during the previous segment (Miller 1981). Between December, 1981 and May, 1982 various techniques of trapping and radio-fitting greater prairie-chickens were attempted and evaluated. Capture techniques included night-lighting and the use of walk-in funnel traps and cannon nets. The initial portion of this segment was conducted under Federal Aid Project SE-3-4, Work Plan II, Job 6. This was replaced by Federal Aid Project FW-145-R, Work Plan I, Job 1 on 1 December 1981. The objectives and design of the study remained the same. STUDY AREAS General study area descriptions have been provided previously (Miller 1981). Intensive work on lesser prairie-chickens (Tympanuchus pallidicinctus) was conducted on 3 apparent population centers, referred to as Southeast Springfield, East Campo, and Southwest Campo. The Southeast Springfield area was centered approximately 15 km south and 11 km east of Springfield, Baca County, Colorado. The East Campo area was centered approximately 7 km south and 16 km east of Campo, Baca County. The Southwest Campo area was centered 8 km south and 5 km west of Campo. RESULTS Lesser Prairie-Chicken The occupied range of lesser prairie-chickens did not change markedly from that reported previously (Miller 1981). The Southeast Springfield area contained 2 leks, and was separated from the nearest lek (in the East Campo area) by 21 km. The distance between the Southeast Springfield and Southwest Campo areas was 27 km. The East Campo area (15 leks) and Southwest Campo (3 leks) areas were separated by 10 km. The Southwest Campo population may have extended into Oklahoma. Areas between the population centers were classified as unoccupied. Lek locations were ascertained primarily by personnel of the Southeast Region, Colorado Division of Wildlife (R. Mellott, J. Slater, and C. Wagner). In the Southeast Springfield area, tallgrasses were dominant in a block of about 5.5 contiguous sections (1400 ha); sandsage dominated in 3 (800 ha). In the East Campo area, the largest contiguous block of tallgrassdominant vegetation type was approximately 3.5 sections (900 ha) with other blocks of approximately 300 and 100 ha. Blocks of about 10 sections �69 (2600 ha) and 7 sections (1800 ha) were dominated by sandsage. Two tallgrass-dominant blocks of about 2 sections (500 ha) each occurred in the Southwest Campo area, and sandsage dominated a 10 section (2600 ha) block. Preliminary analysis of vegetation data indicated that grasses were taller and more dense in areas within 2 km of leks than in areas >2 to 5 km of leks. USDA Forest Service-Comanche National Grassland records revealed lower grazing intensities in pastures within about 2 km of leks (means of 6.2 to 7.2 acres/ADM) than in those >2 to 5 km from leks (means of 5.1 to 5.8 ac/ADM). Greater Prairie-Chicken Maps of the distribution of greater prairie-chickens (T. cupido pinnatus) reported previously (Miller 1980, 1981) were revised following the 1981 surveys (Fig. 1). For the most part, lek locations north of U.S. Route 34 were ascertained by R. Kahn and B. Van Sant, of the Northeast Region, Colorado Division of Wildlife; those south of U. S. Route 34 by G. C. Miller and P. Svoboda. Further revisions of distribution maps will result from 1982 surveys conducted by R. Kahn and B. Van Sant (Van Sant, unpubl.). At least 4 population centers appeared to exist in the area north of the Arikaree River and south of U.S. Route 34, based upon 1981 and 1982 survey work. Blocks of occupied habitat associated with these population centers ranged in area between approximately 5 and 25 km2• The southernmost and easternmost boundaries of the occupied blocks were well delineated by changes in soils and native vegetation. In the area labelled Arikaree in Fig. 1, at least 2 leks active in 1981 were inactive in 1982. Three leks were found in 1982, however, that were not present in 1981. Attempts to trap greater prairie-chickens began in December, 1981. Bait stations were established at 7 sites where feeding flocks had been detected. Husked ear corn, small grains, and moistened alfalfa were used to attract concentrations of chickens. When walk-in funnel traps or cannon nets were set to capture the birds, however, feeding at the stations ceased. Nightlighting of birds at known winter roosts was unsuccessful, due to extremely dense vegetation. Camouflaged cannon nets were set at leks beginning in March. This was the only method by which we caught prairie-chickens. Twenty-three prairie-chickens (12 males, 11 females) were captured. Solarpowered radio transmitters were affixed to 10 birds (3 males, 7 females) with back-pack harnesses. The radio-fitted birds represented 3 leks in 2 population centers. Data from the radioed birds have not been analyzed. At the end of the reporting period, 2 male and 1 female prairie-chicken had been killed. The status of 1 male and 1 female were unknown (no signal), In at least 6 instances females reached the incubation stage; clutches hatched in 2. Two eggs did not hatch from one clutch, although the eggs were fertile. No evidence of infertile eggs was found. �70 At this time, the use of LANDSAT - generated data appears capable of differentiating various types of rangeland, both suitable and unsuitable for sustaining greater prairie-chicken populations. We attempted to differentiate 6 combinations of vegetation and range management practices and were successful in differentiating 5 (Miller and Schrupp 1981). DISCUSSION The portion of greater prairie-chicken range lying north of the Arikaree River and south of U.S. Route 34 was selected as the primary study area for development of a preservation program for insular grouse populations. At least 4 population centers exist, separated from each other by "barriers" (areas unsuitable for sustaining greater prairie-chickens) of varying sizes. In the next segment, collection of habitat and population data will continue, using radioed birds. Additionally, further refinements of the LANDSAT analysis technique for quantifying habitat conditions will be attempted. LITERATURE CITED Graul, W. D. 1978. A technique for evaluating greater pra~r~e chicken habitat in Colorado. Colo. Div. Wildlife unpubl. rep. 3pp. Miller, G. C. 1980. Development of a preservation program for three species of prairie grouse. Job Prog. Rep., Proj. SE-3-3. Pp.76-94 In 1980 Wildlife Research Report Part One, Colo. Div. Wildl., Denver. Miller, G. C. 1981. Development of a preservation program for three species of prairie grouse. Job Prog. Rep., Proj. SE-3-3. Pp. 38-52 In 1981 Wildlife Research Report Part One, Colo. Div. Wildl., Denver. Miller, G. C., and D. L. Schrupp. 1981. Characteristics of greater pra~r~e chicken range in Colorado. Proc. Prairie Grouse Technical Council Conf. 14. (abstract only). Prepared by C~C.1:n·~().<= Gary Miller Wildlife Researcher �71 .- I I I 6 ----:::I _. ~l'i.J..llt---+~-_";:'H~OI-yo.Lke{~~:...__0V••...,e,.,...jnI-rna....J LEGEND - n -. I ~ __F~_H__I__L~L • Lek -<>- Chickens heard 1--;8~! (::J746 I_P~~S ~~'~ ~ ....... Study Area Boundary - --1--:--'- --1'1.-'-.-- ---- ., G~ ~I. I I 1 o~~ ~ 0) ,'- ~I 0 ;:;1 ~I I -<>- -<>- l 00 ·r·\~. t... ~;:'''0 r ~" 0 -<>-.:: N ; -<>- ~;"'.. Po \ .• - _" 0'[ 5' i , I Clarkville r---i--·O-=c~')J=~-"--_.J ~I- ; i ~o ~,,~.s -..., •.•...... -<>- -<>- 51 W ray " , '. • 1 ':-1 . L I :. • IV". •• ~W une!a • I " I 1 0 -I ~I <01 -r---.-·---;H~Y~dle,o. __ (4241) (iv orJ1,\ _ ~ 1 I Yuma I I 49 Fig. 1. I 4s1 M A N f\<· Greater prairie-chicken distribution in Colorado, 1981. I 42. \'1 ��73 JOB PROGRESS State of COLORADO Project No. W-136-R Nongame 1 Job No. Work Plan No. Job Title REPORT Population Surveys of Selected Investigations 1 Bird and Mammal Species in Colorado Period Covered: Personnel: 1 January 1981 through 30 June 1982 W. Graul, D. Vos, G. C. Miller, P. D. Harrison, R. Calderon--Colorado Division of Wildlife. M. Kandel, ABSTRACT This report is presented in two parts: (1) the great blue heron/doublecrested cormorant statewide study, and (2) the great blue heron-human intrusion study. In the statewide study, habitat data were collected from 108 tree stands representing 38 nesting colony sites in northeastern and northwestern Colorado. Forty-three colony sites were surveyed, 11 of which were inactive during the reporting period. In the heron-human intrusion study the responses of nesting herons at four sites to various types of human activity were recorded. Data were then analyzed relative to differences in responses as a function of the type of intrusion, time of day, and time of nesting cycle. This Job Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the Director. ��75 POPULATION SURVEYS OF SELECTED BIRD AND MAMMAL G. C. Miller SPECIES IN COLORADO and Diana Vos P. N. OBJECTIVES 1. For all great blue heron and double-crested cormorant nesting colonies in Colorado, both active and inactive, document the following: (1) colony size and past population trends, and (2) distances to various forms of human development and activity. 2. For all active great blue heron and double-crested cormorant nesting colonies in Colorado document the anticipated future human development and activity patterns near the sites, and the number and condition of the trees being used as nest sites. 3. Document the reactions of nesting great blue herons and doublecrested cormorants to various types and quantities of human activities at nesting colonies. 4. Determine the availability of future alternate nest trees for the active great blue heron and double-crested cormorant colonies in Colorado. SEGMENT OBJECTIVES 1. For one-half of the great blue heron and double-crested cormorant nesting colonies in Colorado, both active and inactive, document colony size and past population trends, and the distances to various forms of human development and activity. 2. For all active great blue heron and double-crested cormorant nesting colonies in Colorado, document the anticipated future human development and activity patterns near the sites, and the number and condition of the trees being used as nest sites. 3. Document the reactions of the nesting great blue herons and doublecrested cormorants to various types and quantities of human activities at 2 nesting colonies. GREAT BLUE HERON/DOUBLE-CRESTED CORMORANT STATEWIDE STUDY INTRODUCTION The Colorado Division of Wildlife's species-ecosystem approach to nongame wildlife management keys management activities to those species exhibiting the most stringent habitat requirements. In Colorado's lowland riparian ecosystems, two such species are great blue herons (Ardea herodias) and double-crested cormorants (Phalacrocorax auritus). Both are colonially nesting species which, in Colorado, nest �76 almost exclusively in cottonwood (Populus spp.) trees. Colorado's entire breeding populations of these species are confined to relatively few (probably <50) colony sites (Graul 1980, 1981). Any preservation of, or management for, the wildlife of the riparian ecosystem, then, must address the needs of these 2 species as a mlnlmum. Previous work, under Federal Aid Project FW-22-R, gave some indication of possible determinants of colony establishment and persistence (Graul 1980). Identification of these determinants will result in a basis for the management of these species, and provide the foundation for the management of the riparian ecosystem, the most restricted ecosystem in the state (Graul and Svoboda unpubl.). The statewide study of great blue heron and double-crested cormorant colonies was designed to test various hypotheses regarding relationships between the status of colonies and characteristics of their nesting resources. Relationships between colony status and physical and biological characteristics of the colony sites may exist. Human activities may influence colony site selection and colony longevity. At least some of the factors being investigated are those that the Colorado Division of Wildlife or other agencies may have a reasonable chance to influence. The study is also designed to yield information on population trends and predictions of habitat availability. METHODS AND MATERIALS Active and abandoned great blue heron and double-crested cormorant colony sites were located from previous work (Graul 1980, 1981) and from reports by Division of Wildlife personnel, other wildlife professionals, and interested citizens. Colony status (active vs. abandoned) and numbers (by species) of individuals, pairs, occupied nests, and nest platforms were recorded during the nesting season, just prior to or at the onset of nest tree foliation (generally between mid-April and late May). Physical and biological characteristics of colony sites were measured during non-nesting periods. Colony sites, stands of trees containing colony sites, and stands of trees within 500 m of colony sites ("alternate stands") were measured and mapped. Vegetation sampling plot centers were established in a systematic (with random starts) fashion. Sampling plots for trees ~76 rom diameter at 1.4 m height were of 5.64 m radius and non-overlapping. These ~lots totalled 500 - 600 m2 for each stand >500 m2 in area; stands ~500 m in area were 100% sampled. Tree species, crown condition (% of crown dead), and tree diameter at 1.4 m height were recorded for all sampled trees >76 rnm in diameter. An increment core was taken from every 10th tree sampled (all sampling from innermost to outermost tree). Embossed, numbered aluminum tags were nailed to all nest trees, whether or not they occurred in sample plots. Species, crown condition, diameter, and number of nest platforms were recorded for all nest trees, and every 10th tree was cored. �77 Trees <76 mm in diameter were sampled from 2.82 m radius plots, using the same plot origins as the larger plots. Stems of these trees were classified by species and age class (0-2 years, 3-5 years, and 5+ years). RESULTS Status of Colonies In northeastern Colorado, platform and active nest counts were made at 19 colonies that had been active during the reporting period (Table 1). One new colony was established on the Arikaree River, and another was found in 1982 southwest of Wray in Yuma County, but has not been surveyed (R. Kahn pers. commun.). Five colonies reported as active by Graul (1981) were not used in at least one of the nesting seasons. Nesting great blue herons were not found at the Horseshoe, Sterling, Walden, Franklin, or Barbour Ponds sites. None of these sites had been known to contain more than 10 nests. Double-crested cormorants nested at 6 of the surveyed colony sites (Barr Lake, Chatfield, Empire, Milton, Jumbo. and Riverside). Cormorants may have nested at Horse Creek, but confirmation was not possible. Active cormorant nests outnumbered those of great blue herons at Barr Lake, Milton, and Riverside. In northwestern Colorado 28 reported colony sites were visited. Twenty-five were confirmed to have been used by great blue herons for nesting (Table 2), Confirmation of nesting was not possible for sites at Dotsero (Eagle Co.). Carbondale (Garfield Co.), and Little Snake (Moffat Co.). Five sites. reported as active by Graul (1981) were not known to contain nests in 1982. All except Clough Island had not been known to contain more than 10 nests. Five colonies not previously reported (Graul 1980, 1981) were located. One site (Craig), reported as inactive (abandoned) by Graul (1981), contained a nest in 1981 but not in 1982. Three inactive sites, previously unreported, were found during the survey. In 1982 occupied great blue heron nests at the surveyed colonies numbered 160; platforms numbered 235. Colony Site Characteristics Nesting resource measurements were taken from 52 stands at 18 colony sites in northeastern Colorado in 1981. Access was denied by the owner of the Barbour Ponds site. Measurements were not taken at 10 abandoned or status unknown sites that could not be located precisely. High water and earlier-than-expected nesting activity prevented some measurements of sites at Barr Lake, Chatfield, Empire, Milton, Panama Lake, and Riverside Reservoir. Stands containing active and abandoned colonies, and alternate stands, appeared to differ from each other in crown condition and diameter distribution. Regeneration of trees (primarily Populus deltoides) was found in 19% of the plots in stands with active colonies, 17% of those with abandoned sites, and in 26% of the alternate stand plots. �78 Table 1. Nest counts reported for great blue heron colonies in northeastern Colorado, 1981-82. Colony Location ConunonName R. Sec. Great Blue Heron Nest Countsl/ Year 1981 1982 County T. Adams IS IN 66W 64W 27-28 31 Barr Lake Horse Creek Res. (148) 37 29 IN 2N 69W 69W 16 35 Boulder Creek Panama Lake (232) (3) 1 95 n.d. Jefferson 6S 69W 12 Chatfield Res. 75 n.d. Larimer 4N 6N 6N 7N 9N 69W 68W 68W 68W 68W 9 16 31 24 18 Lone Tree Res. Fossil Creek Res. Horseshoe Res. Timnath Res. Wellington #3 (52) (122) 115 (4) 1 (8) 8 (56) 51 47 94 0 13 50 llN 48W l3 Jumbo Res. (142) 34 n.d. Washington 5N 54W 14 Prewitt Res. (111) 15 n.d. Weld 3N 6N 5N 3N 4N 2N 6lW 67W 67W 65H 62W 68W 1 1 33 10 12 2 Empire Res. Franklin Lk. Johnstown Milton Res. Riverside Res. Barbour Ponds 4S 46W 6 Boulder Logan Yuma Arikaree R. n.d. (7) (157) (31) 31 n.d. (2) 2 (1) 1 n.d.]} n.d. 47 0 60 n.d. (7) 7 (1) 0 (2) 2 }j Numbers in parentheses are platform counts--all others are active nests. Jj n.d. = no data collected. �79 Table 2. Nest counts reported for great blue heron colonies in northwestern Colorado, 1981-82. Great Blue Heron Nest Countsl! Year 1981 1982 Gypsum Cabin Cr. (5) /2 n.d.l County T. Colony Location R. Sec. Common Name Eagle 5S 2S 85W 85W 6S 7S 6S 6S 941-1 96W 92W 93W 29 13 10 11-12 Clough Island Grand Valley Silt Snyder Is. Grand IN 79W 18 Mesa 8S 9S 9S 9S 97W 97W Garfield Moffat Rio Blanco Routt (5) 4 5 (10) (13) (29) (10) (10) (9) (20) (17) 0 5 12 17 Krennnling (18) 12 (14) 12 97W 33 17 5 18 Debeque DeBeque DeBeque DeBeque n.d. n.d. n.d. n.d. 3 (11) 6 3 (5) 0 7N ION 7N 6N 6N 12N 6N 89W 103W 95W 91W 90W 89W 89W 3 25 30 11 6 13 4 Elkhead Res. Brown's Park Maybell Craig Moffat Airport Slater Ralph White Lk. (5) 4 (6) 6 (3) 1 (1) 1 5 (1) n.d. (4) (6) (2) (1) (19) (1) (4) IN IN IN 93W 94W 96W 33 30 5 Rattlesnake Mesa Powell Park Rio Blanco Lk. (28) (10) 6N 3N 6N 8N 85W 85W 88W 85W 9 34 8 29 Steamboat Yampa Hayden Elk R. (52) 43 (31) (6) (2) 97VJ 6 5 (7) 3 3 0 5 2 0 11 0 2 1 (9) 0 (6) 4 (55) (28) (6) (1) 47 23 0 0 1/ Numbers in parentheses are platform counts--all others are active nests. 2/ n.d. = no data collected. �80 Nesting resource measurements were taken from 56 stands at 20 colony sites in northwestern Colorado. Measurements were not taken at 8 sites because of time limitations and/or our inability to locate abandoned sites precisely. Stands containing active and abandoned colony sites, and alternate stands appeared to differ from each other in crown condition, diameter classes, and area, but not in stem densities. Regeneration of nest tree species (primarily Populus spp.) was found in 24.4% of the plots in stands containing active colonies and in 47.8% of those in stands with abandoned or status unknown sites. Regeneration data for alternate stands was collected but not summarized during the reporting period. The influences of fixed-point human activities upon colonies are unclear. Distances from active colonies to nearest habitation ranged from 90 m to 2,500 m; for abandoned colonies the range was 400 to 1,260 m. Distances to roads (not under construction) showed no clear pattern. Human activities of ephemeral nature were addressed in a different portion of the study (D. Vos unpubl.). One such event was detected during this study, however, at the Prewitt Reservoir site. Large-scale nest abandonment occurred, apparently as a result of a troop of boy scouts camping directly beneath the colony during the great blue heron incubation period (D. Homan pers. commun.). DISCUSSION Over 50% of the presently known great blue heron and double-crested cormorant colonies have been surveyed. The hypotheses developed in the initial stages of the study still appear reasonable, although perhaps not as discreet as originally proposed. For instance, tolerances of colonies to fixed-point human disturbances may be more than a function of distance and type of disturbance--factors such as colony size and the screening effects of topography or vegetation may be involved. No conclusions can be reached about trends in colony or population numbers. Several censuses over a period of time appear necessary to detect such trends. Fluctuations in numbers and colony abandonments can be precipitated by unpredicted and, I suspect, often undetected events (e.g., Prewitt Reservoir site in 1981). No statistical analysis of colony site characteristics was presented in this progress report. High variances due to small sample sizes of abandoned sites would be encountered, making such analyses invalid. Sample sizes should become adequate as the rest of the sites are surveyed and additional sites become inactive. Future work will be concentrated in the southeastern and southwestern portions of the state. Measurements missed from the colonies surveyed during this segment will be taken during the next segment. Additionally, all known sites in the state will be visited for status determination and census between late April and early June, 1983. �81 LITERATURE CITED Graul, W. D., J. Torres, and R. Denney. 1976. A species-ecosystem approach for nongame programs. Wildl. Soc. Bull. 4(2): 79-80. Graul, W. D. 1980. Population surveys of selected bird and mammal species in Colorado. Job Prog. Rep., Proj. FW-22-R. Pp. 98-123 In 1980 Wildlife Research Report Part One, Colo. Div. Hildl., Denver. Graul, W. D. 1981. Population surveys of selected bird and mammal species in Colorado. Job Prog. Rep., Proj. FW-22-R. Pp.83-129 In 1981 Wildlife Research Report Part One, Colo. Div. Wildl., Denver. Prepared by ~:~ Wildlife SJJ1.\.\u: Researcher �82 GREAT BLUE HERON-HUMAN INTRUSION STUDY INTRODUCTION A study of the reactions of nesting great blue herons (Ardea herodias) to various types of human intrusions was initiated in 1980 at the Chatfield Reservoir, Boulder Creek, Lonetree Reservoir and Fossil Creek Reservoir heronries by Elizabeth McGrath. This work was continued by Diana Vos in 1981 and 1982 at Fossil Creek Reservoir and Lonetree Reservoir as a master's project in the Department of Fishery and Wildlife Biology, Colorado State University, Fort Collins. The results presented herein summarize the findings of all work at the latter 2 heronries from 1980-1982. STUDY SITES A complete description of all 4 study sites was presented by Graul (1981). METHODS 1980 Field Season The methods for the 1980 field work were described by Graul (1981). 1981 and 1982 Field Seasons The 1981 and 1982 field season work was conducted from late February through July when most herons had departed from the heronries during both years. Between 15 March and 21 July 1981, and 6 March and 20 July 1982, Fossil Creek Reservoir and Lonetree Reservoir were each visited twice a week on a rotating schedule to include an equal number of visits on weekdays and weekends. On each day of observation the heronries were watched for periods of 6 hours, either 0600-1200 or 1200-1800. The observation periods were divided into monthly segments which corresponded with major stages of the breeding cycle as follows: March - arrival, courtship and nest construction; April - egg-laying and incubation; May - hatching and brooding of young; June - development of nestlings; and July-fledging of young. During the study, a human intrusion was considered to be any human activity within 100 m of the heronry. A distance of 100 m was selected because preliminary study showed that herons rarely reacted to human activity beyond this distance. Any human activity outside 100 m, however, which obviously did cause herons to react was recorded. Reactions of herons to human intrusions were grouped into the same response categories (minimal, local and general) as described by Graul (1981). All observations at the 2 heronries were made using a Redfield �83 15x-45x spotting scope from a distance of 250 m or more to prevent disturbance of the herons by the observer. To examine specific tolerances of great blue herons to various types of human activity, the distance, to the nearest 10 m, at which herons were first disturbed by a particular human activity was recorded. Measurements were made with a Lietz rangefinder. Herons were considered to be disturbed by human activity when 2 or more adult herons (or fledglings when present) flew from their nests while the activity was occurring. If fewer than 2 herons were present on nests during a human intrusion (common later in the breeding season when nestlings were old enough to be left unattended) the distance at which the one heron present first flew from its nest was measured. At Fossil Creek Reservoir, the heronry is spread out, following the contour of the shoreline which is U-shaped. Therefore, while monitoring human disturbances at this site, if herons proximal to a human intrusion were not disturbed, but a few herons further away (>100 m) did fly from their nests, it was presumed that the herons which flew away did so for some other reason than being disturbed by the human activity. To examine variation in heron activity during the day, the number of flights made by herons during each hour were counted in 1981. Herons of 25 active nests at both Fossil Creek Reservoir and Lonetree Reservoir were monitored each day of observation. During each hour of observation a 15 minute time segment was randomly chosen. During this 15 minute time segment all flights, including short flights to the ground beneath a heronry and long flights away from a heronry, were counted. The values obtained were multiplied by 4 to estimate an hourly rate of heron normal activity flights. Experimental Intrusion Study One major objective was to determine if there are differences in heron response relative to different types of human activity. Three other objectives related to this were to determine: i) whether heron response changes as the breeding season progresses, ii) if heron response varies at different times during the day, and iii) if heron response to the same type of human activity differs between heronries. Since the number of naturally occurring human intrusions in 1980 and 1981 were too low and random in time for a statistically valid analysis, a series of controlled experimental intrusions were conducted in 1982. The experiment was designed to require a minimum number of additional intrusions, well below the number known to have occurred in previous years. Further, the experimental intrusions were well spaced in time and short in duration to minimize disturbance to the herons. The experimental intrusions consisted of 4 types of human activity known to occur at Fossil Creek Reservoir and Lonetree Reservoir. The activities were: i) a person approaching a heronry on foot, ii) a person riding a motorcycle past a heronry, iii) a tractor being operated near a heronry, and iv) a motorized boat passing along the shore near a heronry. �84 The "person approaching the heronry on foot intrusion" consisted of Vos walking slowly towards a heronry. She wore similar clothing each time to maintain consistency in the experiment. The type of motorcycle used in the experiment was a Yamaha 175. Vos approached the heronry on the motorcycle along an existing road at the Fossil Creek heronry at a speed of approximately 16 km per hour. For the boat intrusion, a Mirror Craft, 3.7 m (12. ft.) V-bottom boat with a Johnson 9~ HP outboard motor was used. The heronry was passed at a speed of 12-16 km per hour at closer and closer distances until the herons reacted. In the tractor intrusion an Allis-Chalmers (model CA) tractor was driven towards a heronry at a speed of less than 8 km per hour. The experimental intrusions were conducted on a monthly basis to see how heron response varies throughout the breeding season. Each intrusion each month was repeated twice to form replicates to be analyzed later by an Analysis of Variance (Steel and Torrie 1980). The distance at which herons were first disturbed (defined earlier) by each experimental intrusion was measured. All 4 types of experimental human intrusions were conducted at Fossil Creek Reservoir. At Lonetree Reservoir only the "person approaching" and the boat types of intrusions could be done. The motorcycle, motorized boat and tractor experimental intrusions were ~ll conducted in the morning between 0700 and 0900. To see how heron response varies throughout the period of a day, however, the "person approaching a heronry" intrusion was conducted during 3 different time segments: i) morning - between 0700 and 0900, ii) midday - between 1100 and 1300 and iii) afternoon - between 1600 and 1800. Each "person approaching" intrusion at a specific time segment was conducted on different days to reduce the chance of herons becoming habituated to the intrusions. Population Status The number of active nests at each site was counted to estimate population levels. Reproductive success was measured by counting the number of young within nests just prior to when they began climbing onto branches near their nests. Nestlings at this stage were between 45 and 60 days of age. RESULTS AND DISCUSSION Overview of Great Blue Heron Activity Great blue herons began to arrive at Fossil Creek and Lonetree Reservoirs in late February and early March in both 1981 and 1982. In 1981, 20 great blue herons were first seen at Fossil Creek Reservoir on 28 February and at Lonetree Reservoir, 12 herons were first seen on 3 March. In 1982 herons returned to the breeding sites slightly earlier; the first �85 herons were seen on 20 February February at Lonetree Reservoir. at Fossil Creek Reservoir and on 21 During March of both years great blue herons continued to arrive at the breeding sites and pairs began forming. Nest construction and copulation ensued and clutches were initiated. During April herons were involved in incubating eggs. When clutches were being laid both adults were often seen at the nest, but during incubation only one heron was generally at the nest throughout most of the day. In May eggs began to hatch. On 4 Nay 1980 nestlings were first observed at Chatfield Reservoir and by the middle of the month young had appeared at the other 3 heronries (Graul 1981). In 1981 nestlings were first seen at Fossil Creek on 5 May and at Lonetree on 3 May. In 1982 nestlings were first observed slightly earlier; 27 April at Fossil Creek and 5 May at Lonetree. Nestlings were brooded for approximately 10 days after hatching. As they grew older, nestlings were attended by at least one parent throughout the day. By June nestlings parents foraged. were generally left unattended at the nest while both Adult herons only returned to nests to feed the young. In July nestlings fledged as th~'reach about 60 days of age. Adult herons and fledglings lingered along the shore near the heronries and in nearby trees. By mid-July most of the nests had been abandoned. Chronology of Human Intrusions Fossil Creek Reservoir: During March 1981 there was a total of 0.19 human intrusions observed per hour of observation (R) at Fossil Creek Reservoir (Table 1). Human intrusions consisted primarily of people such as nature watchers and photographers approaching the heronry as well as boats, both motorized and unmotorized, passing the heronr)·. In March 1982 (Table 2) the rate of human intrusions per hour of observation was lower = 0.037) than in 1981. One unusual intrusion in 1982 involved a hot-air balloon approaching and briefly hovering directly above the heronry. (R In April 1981 the rate of human intrusions (R) at Fossil Creek Reservoir increased from March (Table 1). During April 1982 the rate of intrusion also increased but not to as high a level as observed in 1981 (Table 2). Intrusions during April of both years consisted of people approaching the heronry on foot as well as people on horseback, trucks passing the heronry and 2 motorcycle intrusions. Also in 1982, aerial intrusions by helicopters flying less than 100 m above the heronry were observed. During May, the rate of human intrusions per hour of observation at Fossil Creek Reservoir increased in 1980 from previous months (R) (Graul 1981) �Table 1. The number and types of human intrusions per observer hour at Fossil Creek Reservoir in 1981, by month. TYPE OF HUMAN INTRUSION March --- April May LAND people on foot motor vehicle motorcycle horseback June July 0.0741 (4) WATER boats2 0.6667 (36) 1.3333 (48) co 0\ Total number of human intrusions/ month 40 48 Total hours of observation/month 36.00 48.00 48.00 54.00 36.00 Total number of human intrusions/ hour /month CR) 0.1944 0.2708 0.1666 0.7407 1.3333 1 Sample size in parentheses. 2 Includes motorized boats, unmotorized boats, sailboats and canoes. �Table 2. The number and types of human intrusions per observer hour at Fossil Creek Reservoir in 1982, by month. TYPE OF HUMAN INTRUSION LAND people on foot motor vehicle motorcycle horseback tractor March --- April May June 0.0555 (3) 0.0185 0) 0.0370 (2) 0.0370 (2) 0.0370 (2) July 0.0740 (4) 0.0185 (1)1 0.0185 (1) WATER boats2 0.1296 (7) 0.0833 (3) co '-...I AIR helicopter hot-air balloon 0.0370 (2) 0.0185 (I) -- - - - --- - - - 0.0370 (2) ------- ------ ----- - ------ Total number of human intrusions/ month 2 8 8 10 3 Total hours of observation/month 54.00 54.00 54.00 54.00 36.00 Total number of human intrusions/ hour/month (11) 0.0370 0.1481 0.1481 0.1852 0.0833 1 2 Sample size in parentheses. Includes motorized boats, unmotorized boats, sailboats and canoes. �88 In May 1981 the rate of intrusions decreased and in 1982 the May rate (R) remained the same as in April. The decrease observed in 1981 may have been associated with weather conditions. Precipitation in May 1981 totaled 4.21 inches while average precipitation for May is 2.90 inches. There were 16 days in May 1981 with measurable precipitation (0.01 of an inch or more), 8 days of which were wet enough to make the ground muddy (Mountain States Weather Services, Fort Collins). This could have influenced the number of people participating in outdoor activities. In June of all 3 years (1980-1982), the rates of human intrusion (ft) increased at Fossil Creek Reservoir from the rates in May. The highest June intrusion rate = 0.7600) took place in 1980 (Graul 1981). Human intrusions in 1980 consisted mainly of people approaching the heronry on foot (62.5%) (Graul 1981), while in 1981 and 1982 most of the intrusions in June (86.0%) were boats passing the heronry. People on foot constituted only 12% of the intrusions in June of 1981 and 1982. Many of the people approaching the heronry on foot in June did so to gather asparagus (Asparagus offincialis L.). (R During July at Fossil Creek Reservoir nestlings fledged and activity within the heronry decreased. The heronry no longer provided a subject for photographers and nature watchers and all human intrusions in July 1980-1982 were boats passing the heronry. When combining all results obtained at Fossil Creek Reservoir from 1980 to 1982 (Table 3), the overall rate of human intrusions per hour of observation increases by month reaching high levels in June and July. During all 3 breeding seasons boats constituted the major source (66.1%) of human intrusions. People approaching the heronry on foot (20.9%) was also a common type of human intrusion. Lonetree Reservoir: Rates of human intrusions appear to be relatively low in March. No human intrusions were observed at the Chatfield Reservoir and Boulder Creek heronries during March 1980 (Graul 1981). No human intrusions were observed at Lonetree Reservoir during March 1981 either. In }farch 1982 only 5 human intrusions = 0.1402) were observed (Table 4). These low levels of human intrusions at heronries in March may be a critical factor influencing whether or not herons will settle into a particular breeding site. Once established in a area, herons may be less likely to abandon the heronry even with increasing rates of human intrusion in subsequent months. (R (R During April 1981 only one human intrusion = 0.0200), a sailboat passing the heronry at a distance of 80 m, was observed at Lonetree Reservoir. Many other boats were seen on the reservoir in 1981, especially during weekends, but few came within 300 m of the heronry. In April 1982 the rate of human intrusion (R) reached the highest level �Table 3. The combined number and types of human intrusions per observer hour at Fossil Creek Reservoir for the 1980, 1981 and 1982 breeding seasons. TYPE OF HUMAN INTRUSION LAND people on foot motor vehicle motorcycle horseback tractor March --- April 0.0333 (3)1 0.0588 0.0196 0.0196 0.0098 0.0185 (1) 0.0185 (1) (6) (2) (2) (1) May June 0.0611 (7) 0.1570 (21) 0.0374 (5) 0.0075 (1) 0.0349 (4) July 0.0075 (1) WATER boats2 0.0833 (3) AIR helicopter hot-air balloon 0.0185 (1) 0.0784 (8) 0.0349 (4) 0.3734 (50) 0.5977 (52) ~ 0.0196 (2) 0.0175 (2) Total number of human intrusions/ month 9 21 17 78 52 Total hours of observation/month 90.00 102.00 114.50 133.75 87.00 Total number of human intrusions/ hour /month (:R) 0.1000 0.2059 0.1485 0.5832 0.5977 1 2 CJ:) Sample size in parentheses. Includes motorized boats, unmotorized boats, sailboats and canoes. �Table 4. The number and types of human intrusions per observer hour at Lonetree Reservoir in 1982, by month. TYPE OF HUMAN INTRUSION LAND people on foot motor vehicle horseback tractor WATER boats2 March --- April May 0.0417 (2)1 0.0417 (2) 0.0625 (3) 0.0185 (1) 0.2083 (10) July 0.0208 (1) 0.0208 (1) 0.0208 (1) 0.0208 (1) June 0.1481 (8) 0.0926 (5) 0.0833 (3) 1.0 o AIR airplane 0.0208 (1) Total number of human intrusion/ month 5 15 9 7 3 Total hours of observation/month 48.00 48.00 54.00 48.00 36.00 Total number of human intrusions/ hour /month CR) 0.1042 0.3125 0.1667 0.1458 0.0833 1 2 Sample size in parentheses Includes motorized boats, unmotorized boats, sailboats and canoes. �91 of that year (Table 4). Sixty-six percent of the April 1982 intrusions were boats passing the heronry. Also, one airplane was seen flying 60 m above the heronry. During May 1981, the rate of human intrusion (R) reached its highest level = 0.1000) at Lonetree Reservoir. In 1982 the May intrusion rate decreased from the rate in April (Table 4) but was similar to the May rates observed in 1980 (Graul 1981) and 1981. The most intrusions during all 3 years took place over Memorial Day weekend. (R In June and July of all 3 years the rates of human intrusion (R) decreased from the rates observed in previous months. In 1980 no human intrusions were observed at Lonetree Reservoir in June or July (Graul 1981). No human intrusions were observed in June or July 1981 either. In 1982 there were several human intrusions in June and July, of which 80.0% were boats passing the heronry (Table 4). The overall rate of human intrusion at Lonetree Reservoir was high in 1980 (60 intrusions, 58.3% land related) and the heronry was considered to be highly disturbed (Graul 1981). In 1981, however, only 7 human intrusions were observed and in 1982 there were 39 intrusions. In 1981 human traffic from the public campsite across the reservoir may have been better controlled. Namely, Fred Eldred, caretaker of the Mead fishing and hunting club in 1981, which leases Welch Reservoir to the south of the heronry, made a special effort to keep people away from the heronry that year. In 1982 several additional "No Trespassing" signs were posted in the area. The combined results obtained between 1980 and 1982 at Lonetree Reservoir (Table 5) shows that the rate of human intrusion at this heronry peaked in May. During all 3 years of the study, the most common type of intrusion was boats (52.8%) passing the heronry. People approaching the heronry on foot constituted 34.9% of the intrusions. Overall Intrusion Patterns: Changes in the rates of human intrusion during each month for the 4 heronries studied between 1980 and 1982 are summarized in Figure 1. As can be seen, peak and low rates of human activity did not coincide at different heronries. The low rates of intrusion at Chatfield Reservoir in May and June, however, may not be representative of the normal situation since the reservoir was closed to all boating between 8 May and 28 June, 1980 because of unusually high water levels (Graul 1981). The daily activity pattern of great blue herons when compared with human recreational patterns during 1981 and 1982 is interesting (Fig. 2). Herons were most active in the early morning and late afternoon at Fossil Creek and Lonetree Reservoirs while most human intrusions (89.6%) occurred between 0900 and 1500. �Table 5. The combined number and types of human intrusions per observer hour at Lonetree Reservoir for the 1980, 1981 and 1982 breeding seasons. TYPE OF HUMAN INTRUSION LAND people on foot motor vehicle motorcycle horseback tractor WATER boats2 March --- April May 0.0417 (2)1 0.0417 (2) 0.0306 (3) 0.0102 (1) 0.2081 0.0143 0.0072 0.0287 0.1122 (11) 0.2009 (28) 0.0208 (1) (29) (2) (1) (4) June July 0.0158 (2) 0.0125 (1) 0.0079 (1) 0.0079 (1) 0.0948 (12). 0.0500 (4) \.0 N AIR airplane 0.0102 (1) Total number of human intrusions/ month 5 16 64 16 5 Total hours of observation/month 66.00 98.00 139.33 126.50 80.00 Total number of 0.0758 0.1633 0.4593 0.1265 human intrusions/ hour/month (Ii) 1 Sample size in parentheses 2 Includes motorized boats, unmotorized boats, sailboats and canoes. 0.0625 �93 0.6 ________ Fossil Creek Reservoir Lonetree -.-. 0.5 -- l( Reservoir Chatfield (1980-1982) Reservoir Boulder Creek -X"" (1980-1982) (1980) • / (1980) A .-/ / \ II \. I ~ .~ 0.3 / \ ..•..... / I Ul ;::l l-< / +J ~ H / . 1 '3 I \ / Ul / b 0.2J i .f\/ \ / • \ · /./) -J 0.1...... iI I . J/ / -/- • / · /i U ;../"'1- . ./'" ------------r------------~------------_.--------------r_------ March April May Figure 1. The number of human intrusions observed 4 heronries studied from 1980-1982 by month. June per hour July (ft) at each of the �94 A I , / I , 25 \ \ Human Intrusions \ 20 Heron Activity '"c0 •... Q) ;;-, .0 \ Q) 15 "0 <:l E <J; .u Cc .•..• .-; ~ 10 \ \ \ \. __ / 18.0 ,/ -: /' / •... Q) ,/ .0 E ;:J Z Q) 5 14.0 0 12.0 II c c"' 0 .,.; 10.0 "' •... ;:J .u c •..... c e'"o ;:J ;r 8.0 6.0 ..... 0 .u c 4.0 Q) u •... Q) ""' 0( til •... Q) :> < 16.0 '" '"' "-'c 2.0 6-7 7-8 8-9 9-10 10-11 11-12 12-13 13-14 14-15 15-16 16-17 17-18 Time (hour intervals) Figure 2. Percent of human intrusions per hour at the Fossil Creek and Lonetree Reservoirs combined for 1981 and 1982, and the average number of normal activity flights made by' herons l per hour throughout the 1981 season for comparison. Herons from 25 active nests at both heronries monitored throughout the 1981 season. �95 The number of human intrusions on cooler days (raining and/or temperatures below SooF, n = 7) throughout the 1981 breeding season at Fossil Creek Reservoir was significantly lower (t = 1.82, p < 0.05) than the number of intrusions on warm, sunny days (n = 20). The number of intrusions on windy days (maximum wind velocity >24 km per hour, n = 6) was not significantly different (t = 1.27, p > 0.05) from the number on calm days (n = 20). The number of human intrusions on weekends at Fossil Creek and Lonetree Reservoirs during 1981 and 1982 combined was significantly higher (T = 5.18, p < 0.001) than the number on weekdays. Heron Response to Human Intrusions At the 4 heronries studied between 1980 and 1982, most human intrusions (66.9%) caused only a minimal response while local responses were elicited during 26.9% of the human intrusions (Table 6). Only 6.2% of the intrusions resulted in a general response. When examining each type of human intrusion independently, boats passing the heronry caused mainly (92.1%) minimal responses (Table 7). Only 7.9% resulted in a local response. It is interesting to note that all the local responses caused by boat intrusions involved slow moving boats and canoes which maneuvered directly under the heron trees. No boat intrusions resulted in a general response during any of the 3 years the heronries were observed. Land related intrusions at the 4 heronries resulted most often in local responses (61.4%), while minimal responses were elicited during 21.9% of the land intrusions and 16.7% caused a general response (Table 7). General responses were often elicited when people approached the heronry on foot, especially early in the breeding season. One general response was caused at Lonetree Reservoir on 2 May 1981 when 4 people approached the heronry and one of the people climbed a nest tree. Motorcycle intrusions often caused general responses as well. Air related intrusions resulted most often in minimal responses ~6.7%), although one intrusion, a hot-air balloon hovering above the Fossil Creek Reservoir heronry in March 1982, caused a general response. Overall it appears that great blue herons are most disturbed by land related intrusions and least by boat activity. Grubb (1978) found that great blue herons may become habituated to loud noises, such as that of an outboard motor. There were too few air related intrusions to determine if herons are generally bothered by them. The herons at Fossil Creek Reservoir may have habituated to aerial activity, however, because there is an airport located 3.5 km south of the reservoir and at least 10 airplanes were seen flying overhead each day of observation throughout the 1982 breeding season. �96 Table 6. Percent of types of human intrusions per type of heron response for all 4 heronries studied during 1980, 1981 and 19821. TYPE OF HERON RESPONSE2 Minimal Local General 17.34% (56) 1.24% (4) 0.31% (1) 2.67% (7) 0.62% (2) 2.67% 0.93% 2.48% 0.31% TYPE OF HUMAN INTRUSION LAND People on foot motor vehicle motorcycle horseback tractor WATER boats4 5.88% 0.62% 0.93% 0.31% (19)3 (2) (3) (1) 57.89% (187) 4.95% (16) AIR airplane helicopter hot-air-ballon 0.31% (1) 0.93% (3) 0.31% (1) Total n=323 66.88% 26.93% (7) (3) (8) (1) 6.19% 1 Fossil Creek Reservoir; 1980-1982. Lonetree Reservoir; 1980-1982. Chatfield Reservoir; 1980. Boulder Creek; 1980. 2 Defined in Graul (1981). 3 Sample size in parentheses. 4 Includes motorized boats, unmotorized boats, sailboats and canoes. �97 Table 7. Percent of heron response per response category for each type of human intrusion at the 4 heronries studied during 1980, 1981 and 19821. TYPE OF HERON RESPONSE2 Local General 8.54% (7) 33.33% (3) 66.67% (8) 11.11% (2) 21.93% 68.29% (56) 44.45% (4) 8.33% (1) 77.78% (7) 100.00% (2) 61.40% 92.12% 7.88% 100.00% (1) 75.00% (3) 25.00% (1) 66.66% 16.67% Minimal TYPE OF HUMAN INTRUSION LAND people on foot motor vehicle motorcycle horseback tractor Combined n=114 WATER boats4 n=203 AIR airplane helicopter hot-air balloon Combined n=6 23.17% 22.22% 25.00% 11.11% (19)3 (2) (3) (1) 16.67% 100.00% (1) 16.67% 1 Fossil Creek Reservoir; 1980-1982. Lonetree Reservoir; 1980-1982. Chatfield Reservoir; 1980. 2 Defined in Graul (1981). 3 Sample size in parentheses. 4 Includes motorized boats, unmotorized boats, sailboats and canoes. �98 Changes in Heron Response: Great blue herons are quite sensitive to human intrusions early in the breeding season and will flush from their nests with the slightest disturbance, returning only when the disturbing cause is no longer present (Cottrille and Cottrille 1958). In late February of 1981 and 1982 all herons present at the Fossil Creek Reservoir and Lonetree Reservoir heronries flew from their nests as I approached at a distance of more than 200 m. Attachment to the nest site, however, appeared to strengthen after eggs had been laid and young had hatched. During all 3 breeding seasons, an increasing percentage of minimal responses were elicited with human intrusions as the season progressed (Table 8). Figure 3 illustrates the monthly changes in average distances at which land related intrusions first caused herons to be disturbed during 1981 and 1982. It can be seen that herons flew less readily from their nests in response to human activity as the breeding season progressed. The average distances at which herons first responded to land related intrusions at Lonetree Reservoir each month was slightly less than the distances at Fossil Creek Reservoir (Fig. 3). A difference in heronry structure may be a factor accounting for this. The original heronry at Lonetree Reservoir is located within a dense grove of trees and intruders are not as readily visible to the herons as at Fossil Creek Reservoir. Wintering bald eagles (Haliaeetus leucocephalus) were found to be more tolerant when human intruders were partially obscured from their line of sight by buffers of vegetation (Stalmaster and Newman 1978). Experimental Intrusion Segment The average distances at which great blue herons were disturbed by the 4 types of experimental intrusions conducted at Fossil Creek Reservoir are presented in Figure 4. No clear pattern of differences in heron response to different types of human intrusions are evident. Herons reacted to the tractor and the person approaching the heronry types of intrusions at similar distances each month, and the boat intrusions resulted in heron response at lesser distances than other types of intrusions in all months except July. It is interesting to note in Figure 4, however, the definite change in heron response which occurred overall as the breeding season progressed. As can be seen, heron nest attachment increased from March through May when eggs began to hatch and then decreased in subsequent months. The experimental boat intrusions caused herons to fly from their nests during each month of the experiment while uncontrolled boat intrusions rarely did. Most uncontrolled boat intrusions at Fossil Creek Reservoir consisted of motorized boats passing the heronry at speeds of 45-55 km per hour while during the experiment boat intrusions the heronry was �Table 8. Heron response to human intrusions at all 4 heronries studied during 1980, 1981 and 19821, by month. -_ March April May June July Minimal 28.57% (4)3 64.71% (33) 51.65% (47) 66.67% (62} 95.95% (71) Local 42.86% (6) 21.57% (11) 40.66% (37) 31.18% (29) 4.05% (3) General 28.57% (4) 13.72% (7) 7.69% (7) 2.15% (2) n=323 TYPE OF HERON RESPONSE2 1 Fossil Creek Reservoir; 1980-1982. Lonetree Reservoir; 1980-1982. Chatfield Reservoir; 1980. Boulder Creek; 1980. 2 Defined in Graul (1981). 3 Sample size in parentheses. I.D I.D �100 90 n=4 n=4 Fossil Creek Reservoir 80 Lonetree D Reservoir n=10 70 60 ".......-_ S .. n=4- '-' 50 n=4 n=10 n=4 40 30 20 10 March April Hay June Figure 3. Average distance at which naturally occurring land related intrusions1 first caused herons to be disturbed at Fossil Creek and Lonetree Reservoirs during 1981 and 1982 by month2. 1 Includes people on foot, horseback, motorcycle, motor vehicle intrusions. 2 -July excluded because no land intrusions were observed. and tractor �March April May June Figure 4. Average distance at which the 4 types of experimental intrusions first caused herons to be di~turbed at Fossil Creek Reservoir by month. I Average of values from morning. midday and afternoon intrusions presented in Tahle 9. 2 Water level· too low to conduct boat f ntus i on .. 3 Fledglings only reacting, no adults present. �102 passed at a speed of only 12-16 km per hour. Furthermore, during most uncontrolled boat intrusions the boat passed only once while in the experimental intrusion the heronry was passed repeatedly at closer and closer distances until a response was elicited. During most experimental boat intrusions a response was only elicited at distances nearer to the shore than most boats could usually come safely. A comparison of the results obtained for the experimental boat intrusions at Fossil Creek and Lonetree Reservoirs (Fig. 5) showed little difference in heron response between the 2 sites to the same type of intrusion, even though boat activity is rare near the heronry at Lonetree Reservoir (most recent site). The average distances at which great blue herons responded to the "person approaching heronry" type of intrusion at different time periods during the day are presented in Table 9. There does not appear to be any consistent pattern to the way herons responded to the same type of intrusion at different times during the day. Results obtained for the "person approaching the heronry" intrusion at Fossil Creek and Lonetree Reservoirs are presented in Figure 6 for comparison. Herons at Fossil Creek Reservoir flew from their nests as a person approached later than herons at Lonetree Reservoir during all months except July. Results from July should be examined separately, however, since during July only fledglings were present within the heronry to respond to the intrusions. In March through June only adult herons could react to intrusions. Effect of Human Activity on Nesting Success At Fossil Creek Reservoir, 115 nests were occupied at one point during the 1981 breeding season. One hundred and seven pairs were successful at fledging at least one young. In 1982, 94 nests were occupied at Fossil Creek Reservoir and 92 pairs successfully produced young. At Lonetree Reservoir 27 nests were occupied at the more recent site in 1981 and 25 pairs fledged young. In 1982, 22 pairs occupied nests at the more recent site, and all produced young. Active nests at the original site at Lonetree Reservoir could not be counted without disturbing the herons but a nest count made after the herons had left the heronry each year revealed 20-25 nests in 1981 and 35-40 nests in 1982. A summary of the number of active nests at the Fossil Creek and Lonetree Reservoir heronries is presented in Table 10. An average of 2.85 young per successful nest and 2.65 per nesting attempt was produced at Fossil Creek Reservoir in 1981. In 1982 production was 2.83 young per successful nest and 2.76 per nesting attempt at Fossil Creek Reservoir. At Lonetree Reservoir (more recent site only) production was 2.86 and 2.82 young per successful nest, and 2.74 and 2.82 young per nesting attempt for 1981 and 1982 respectively. The number of young produced at both heronries during both years appears adequate when compared to the value of 1.91 young per breeding pair (nesting attempt) calculated by Henny (1972) to be necessary to maintain a stable population. �180 165 HERONRY --- 150 Fossil Creek Reservoir 135 Lonetree Reservoirl 120 D II ~ 105 r:l 0 •..l n=1 IJl ~ ~ 90 H .•.•a t-' o 75 J aJ g 11 w n=2 60 IJl orl p 45 30 15 I I ~ March Figure 5. Comparison between average distances at which the motorboat type intrusion disturbed herons at both Fossil Creek and Lonetree Reservoirs by month. I Experimental boat intrusions conducted at mos t recent heronry site at Lone t ree Reservoir only. 2 Water level too low to conduct boat i.ntrusion. 3 Fledglings only reacting, no adults present. �Table 9. Average distances1 (m) per month at which herons were first disturbed by the "person approaching" type of experimental intrusion, for 3 daily time segments. TIME SEGMENTS2 March A MD M 140 125 130 40 60 55 25 75 175 160 80 80 95 50 M AEril MD A M May MD June MD A M July3 MD A A M 40 35 65 45 55 70 70 60 40 45 60 60 55 50 60 50 HERONRY LOCATION Fossil Creek Reservoir Lonetree Reservoir (original site)4 I-' 0 Lonetree Reservoir (newer site)4 1 165 130 175 90 75 80 50 50 60 55 60 Each value based on two trials except in July (1 trial) M = morning (0700-0900), MD = midday (1100-1300), A = afternoon (1600-1800). 3 Fledglings only reacting, no adults present. 4 See description of study sites in Graul (1981). 2 60 70 60 50 ~ �n=6 HERONRY Fossil Creek Reservoir Lonetree Reservoir (original site) 2 ~ ~ ...•0., Lonetree Reservoir (newer site) 2 e •... I:l o II ISJ H .•.• 0 I-' QJ o 7S o V1 ...ij., ...• p 60 45 Figure 6. Comparison of the average distances 1 at which herons rUst reacted to the "person walking in" type of intrusion at the Fossil Creek and Lonetree Reservoir heronries by month. I 2 3 Values are the average of dlstanr.es obtained for the 3 daily time negments, presented See description of study sites In Graul (1981). Fledglings only reacting, no adults present. in Table 9. �106 Table 10. Number of active nests counted at the Fossil Creek and Lonetree Reservoir heronries between 1979 and 1982. NUMBER OF ACTIVE NESTS HERONRY LOCATION 19791 19802 1981 1982 Fossil Creek Reservoir 61-65 80-85 107-115 94 Lonetree Reservoir (original site)4 inactive3 25 (reoccupied) 20-25 35-40 Lonetree Reservoir 4 (newer site) 41-45 22 27 22 1 From Graul (1979). 2 From Graul (1981). 3 Abandoned presumably because of a shooting incidence in mid-1970's; relocated to newer site 0.5 km to the east (pers. comm. C. Cummings). 4 See description of study sites in Graul (1981). �107 LITERATURE CITED Cottrille, W. P., and B. D. Cottrille. 1959. Great blue heron: behavior at the nest. U. Mich. Zool. Misc. Publ. 102: 1-15. Graul, W. D. 1979. Population surveys of selected bird and mammal species in Colorado. Colo. Div. Wildl. Job Progress Rep., ProJ. FW-22-R. 1981. Population surveys of selected bird and mammal species in Colorado. Colo. Div. Wildl. Job Progress Rep ,, Proj. FW-22-R. Grubb, M. M. 1978. Effects of increased noise levels on nesting herons and egrets. Proc. Colonial Waterbird Group, 1978: 49-54. Henny, C. J. 1972. An analysis of the population dynamics of selected avian species, with special reference to changes during the modern pesticide era. U.S.D.I. Bureau of Sport Fisheries and Wildlife, Wildl. Res. Rep. No.1. Stalmaster, M. V., and J. R. Newman. 1978. Behavioral responses of wintering bald eagles to human activity. J. Wildl. Manage. 42: 506-513. Steel, R. G. D., and J. H. Torrie. 1980. Principles and procedures of statistics. A biometrical approach. 2nd ed. McGraw-Hill Inc. New York. Report segment prepared by: Diana Vos ��\ 109 JOB FINAL REPORT COLORADO State of Work Plan No. Job Title Investigations 2 Job No. 1 Statewide Period Covered: Personnel: Nongame W-136-R Project No. 1 January Bat Distribution Study 1981 through 31 December W. Graul, J. Freeman, L. Wunder--Colorado 1981 Division of Wildlife. ABSTRACT This study was designed to document the statewide distributions and habitat associations of the bats of Colorado. During the field seasons of 1978-81 more than 900 individuals of 14 species of bats were captured. The study documented: (1) range extensions of Myotis yumanensis, ~. californicus, Pipistrellus hesperus, Antrozous pallidus, Lasiurus cinereus, and Lasionycteris noctivagans, (2) that Tadarida brasiliensis is a summer resident of the state, and (3) habitat associations for 14 bat species. Additionally, the study provided over 325 standard museum specimens for future study and provided scientific and educational material to the Denver Museum of Natural History. This Job Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR' QUOTED without permission of the Director. ��111 STATEWIDE BAT DISTRIBUTION STUDY Jerry Freeman P. N. OBJECTIVES 1. Determine statewide bats of Colorado. distributions and habitat associations for the INTRODUCTION This project was designed to document distributional patterns and habitat associations of bat species in Colorado. The purpose of this report is to present the results of four summer field seasons of work regarding 14 of Colorado's 16 bat species. An appendix lists exact localities where each bat species was taken. It also indicates the number of individuals observed per locality as well as the number of individuals collected per locality and prepared as scientific specimens. All specimens besides 40 Tadarida brasiliensis are housed at the University of Colorado Museum, Boulder, Colorado. The 40 T. brasiliensis are in the collection of the Denver Museum of Natural History, Denver, Colorado. METHODS AND MATERIALS After reviewing the scientific literature and contacting professional mammalogists, areas were selected to be trapped for bats. Most of these areas had never been investigated or had not been investigated in the recent past. Some areas which had been studied previously were used to validate some important recorded distributions. At each area, bats were trapped by placing three to five mist nests over isolated ponds, metal tanks or along streams. Trapping was done throughout the night for one to three nights in each area. For each individual captured, species, sex, reproductive condition, time of capture, weight, and length of forearm were recorded. During four field seasons more than 175 nights (approximately 700 net-nights) of netting were completed. In addition to trapping by mist netting during the night, old buildings, cellars, mines, bridges, rock crevices and caves were searched during the day for bats. RESULTS AND DISCUSSION During the summer months (May through September) of 1978, 1979, 1980 and 1981, more than 900 individuals of 14 species (Table 1) were taken; more than 325 have been prepared as standard scientific specimens. Two species (Lasiurus borealis and Tadarida macrotis) of rare occurrence in Colorado (Armstrong, 1972) were not captured. The spotted bat, Euderma maculatum, known to occur in Colorado, also was not captured. Two other species of probable occurrence (Myotis velifer and Plecotus phyllotus) were not found. �112 Table 1. Number of individuals Species caught No. Caught Myotis lucifugus 6 Myotis vumanensis 22 ~ thysanodes californicus No. Caught Species Pipistrellus 242 Myotis per species. 22 hesperus Eptesicus °fuscus 68 Lasionvcteris 56 noctivagans 5 Lasiurus cinereus 61 90 Plecotus townsendii 10 65 Antrozous pallidus 52 49 Tadarida 165 brasiliensis 912 TOTAL Table 2. Distribution of Coloradan mammals in 14 community-t,~es. ~ (0') ~ r-- -e ~ _..... ro ....• ...• -e ~ ro ....• fIl fIl ro ... 00 -e or< S ;:l .c Species ..0 ;:l '" Myotis lucifugus Myotis yumanensis Myotis evotis Myotis thysanodes Myotis volans Myotis californicus Myotis leibii X Lasionycteris noctivagans Pipistrellus hesperus Eptesicus fuscus Lasiurus borealis Lasiurus cinereus Plecotus townsendii Antrozous pallidus _..... N fIl -e 0 0 :;. ~ -o ~ ro or< ....• '"... '"'CJ .;:;- " ~ ... _..... so ~ s: fIl ;:l ... ..0 CJ ~ or< ..... ..0 ;:l _..... II") .r: fIl ;:l 0 ... U or< ..0 CJ :>. co '" co X X fIl or< "- ~ -a ~ " c;: fIl ·M c ro ..... -e ..... x X X X eo ... OJ fIl '"'... 0 0 " ... OJ 4-< .r< c 0 fIl OJ Q) '0 or< '" '"'" CJ ~ -e N c ro -e 0 0 _..... cr:;. ~ .., ~ ... 0Q) ·rl ro fIl CJ 0 ~ or< fIl .....P- Q) '" ..0 ;:l_"'" fIlO ..... 4-< 0 ... ~ 0 CJ -o ro '"'c "~ro " '"''''' "- p., ;;: ;;: X X X >, eo >, '" CJ ~ 0 X X X X X X X X X X X X X X X 0 4-< 0 c;: 0 QJ '" .!< c ro ..0 '" <II CJ ... +J fIl OJ ~ or< Q) X X X X X Q) Ul X X ~ ..... ....• ~ .., ... ....• X X X X X ..... "d Q) X X X ~ c;: ~ eo ~ .....0- '" ..0 ;:l '" X X X X X ~ ...• ~ (0') "" ~ •..... .." 0 0 -c ro ....• .c eo .r< ::c < x X X X " " ~ OJ 0Ul X X X X .~ ;t-:r 0 •....• ..,,~ "'CJro....• CJ OJ •••• "'" c, ....• or< •..•• .... CJ <4-< �113 Table 2 provides a synopsis of the habitat association(s) of Coloradan bat species and will be referred to continually in the species accounts given below. Table 2 and figures 2-15 are taken and modified from Armstrong (1972) and used here with his permission. For a detailed discussion of community-types in Table 2 see Armstrong (1972). Figure 1 indicates areas trapped. In figures 2-15, closed circles indicate localities from which specimens were examined by Armstrong (1972), open circles are additional records as noted in Armstrong (1972) and solid triangles indicate localities where each species was observed during this study. The ranges mapped are from Armstrong. Myotis lucifugus. The little brown bat though known to occur widely in the mountains of western Colorado was captured at only two locations (Fig. 2). The location in southwest Colorado was an abandoned church building in a ponderosa pine forest which was being used by a maternity colony of approximately 200 bats. Dr. Preston Somers of Fort Lewis College in Durango noted that this location had been used in several previous summers as a maternal colony roosting site. Two~. lucifugus were captured in 1978 at Elk Springs in pinon-juniper habitat. As intensive field work at this site has been done for three field seasons (1979, 1980, 1981) without capturing further individuals and much field work has been done at lower elevations in western Colorado, it would seem that ~. lucifugus occurs at higher elevations, i.e. transitional life zone and above. Myotis yumanensis. The Yuma myotis, previously known from only the most northwestern and southwestern counties in western Colorado and three southeastern counties, was taken not only in southeastern Colorado but also along the western border of the state (Fig. 3). The Yuma myotis should be regarded as occurring along the entire Western Slope at lower elevations. ~. yumanensis was captured in semidesert scrub and pygmy conifer woodland habitats (Table 2). M. yumanensis was always caught in or near canyons or high rocky bluffs where running streams were nearby, except for six individuals which were taken from the interior of John Martin Dam in Bent County. The Yuma myotis is known to use unoccupied man-made structures within its range (Armstrong, 1972). Myotis evotis. Armstrong (1972) noted that the long-eared myotis is a bat of middle elevations generally occurring above 6,000 ft. in ponderosa pine and pygmy conifer woodlands. The results of this study (Table 2 and Fig. 4) confirm his observation. Fourteen of the 16 localities from which this species was collected were be tween 6,000 ft. and 7,500 ft. and two were above 5,200 ft. All sites were in the above habitat types. Myotis thysanodes. The fringed myotis was captured only in three locations: Moffat County (two individuals), Rio Blanco County (two individuals), and Mesa County (one individual) (Fig. 5). All five individuals were taken in pygmy conifer woodland habitat (Table 2). It seems, as suspected by Armstrong (1972), "that M. thysanodes is not especially common at the edge of its range." Myotis volans. This species, the long-legged myotis, occurs widely in wooded habitats in western Colorado (Table 2, Fig. 6). It was collected from elevations ranging from 5,000 ft. in Yellowjacket Canyon, Montezuma County, to almost 10,000 ft. in Fish Canyon, Hindsdale County. �114 Myotis californicus. The California myotis reaches the northeastern limits of its range in Colorado (Armstrong, 1972). All 49 individuals taken in this study were in semidesert scrub and pygmy conifer woodland habitats (Table 2) in northern and southern Moffat County and in Rio Blanco County (Fig. 7). These data indicate that~. californicus should be considered to occur at lower elevations along the entire Western Slope. Myotis leibii. The small-footed myotis ranges throughout the state of Colorado at lower elevations (Fig. 8). Individuals were captured in saxicoline brush and pygmy conifer woodland habitat where rocky outcrops occurred (Table 2). Lasionycteris noctivagans. The silver-haired bat is a migratory species which occurs in Colorado from late March to early October. Data from this study indicate that both sexes are in the state until late May when females go north. Females return in late August and both sexes migrate south by mid-October. L. noctivagans was captured in pygmy conifer woodland, ponderosa pine woodland, montane forest, and aspen woodland habitats (Table 2). Results of this study indicate that the silver-haired bat uses habitats at lower elevations (especially pygmy conifer woodland habitat) more than was previously known (Fig. 9). I suspect these habitats are particularly important to migrating females, perhaps due to a reduced food supply at higher elevations in late spring and early fall. Pipistrellus hesperus. The western pipistrelle, Colorado's smallest bat, occurs in saxicoline brush and pygmy conifer woodland habitats (Table 2). This species, previously known to occur along the western border as far north as the Colorado River Valley, was captured northward along the White River in Rio Blanco County (Fig. 10). I suspect it may extend further northward and be found in Brown's Park in northwestern Moffat County. Eptesicus fuscus. This species is perhaps Colorado's most common bat. It is Colorado's most widespread Chiropteran species as it occupies man's buildings (Fig. 11). It was taken during this study from the following habitats: plains riparian woodland, saxicoline brush, pygmy conifer woodland, ponderosa pine woodland, and aspen woodland (Table 2). Preliminary analysis of data indicates that the big-brown bat may show elevational movements during the year, occupying lower elevations during the colder months and moving upward as temperatures warm. Lasiurus cinereus. Like Lasionycteris noctivagans, the hoary bat is a migratory species with the females going north for the summer while the males remain in Colorado. This species occurred in the following habitats: pygmy conifer woodland, ponderosa pine woodland and aspen woodland (Table 2). Results from this study indicate that L. cinereus is more common in habitats of lower elevations than previously known (Fig. 12). Data from the Elk Springs site where netting has occurred on a monthly basis for three field seasons indicate that the intensity of use of pygmy conifer woodland by this species may vary from year to year. Like female silverhaired bats, female hoary bats may depend on the food resources of habitats at lower elevations during migratory flights. �115 Plecotus townsendii. Townsendts big-eared bat was captured only ten times at six locations. These locations were in pygmy conifer woodland and ponderosa pine woodland habitats (Fig. 13) (Table 2). Antrozous pallidus. The pallid bat occupies semidesert scrub and pygmy conifer woodlands (Table 2). All individuals observed from this study come from the above two habitats. ~. pallidus was collected in southern Moffat County and in Rio Blanco County (Fig. 14); this indicates that it can be considered to occupy lower elevations along the entire Western Slope. Tadarida brasiliensis. A colony of 9,000 Brazilian free-tailed bats in the San Luis Valley, Saguache County was reported previously (Meacham, 1974). I checked the colony in July, 1978 (Fig. 15). It had an estimated population of 50,000. Only adult males were taken in 1978. During 1979, juveniles and lactating females were also captured from the colony, then estimated at 50,000 - 75,000 bats. In 1980, the size of the colony was estimated to be 100,000 - 150,000. Juveniles, lactating females, and females with embryos were taken in addition to males. In 1981, Laurie Wunder of the Colorado Division of Wildlife made a population estimate using photographic techniques. She estimated a population size of 86,000 and although she found juveniles in the colony no lactating or pregnant females were captured. This is the northernmost breeding colony of this species east of the continental divide. These data indicate that ~. brasiliensis should be considered a resident species of the state. Populations of this species have been declining dramatically elsewhere. Data show that pesticide poisoning has contributed to this decline (Geluso et al., 1976). Due to the importance of any breeding colony of this species, I sent fecal material and bodies to Patuxent Wildlife Research Center, U.S. Fish and Wildlife Service, for pesticide analysis. Results of that analysis indicate that the colony is free of serious organochlorine contamination. Data from the analysis appear in appendix II. SPECIAL COMMENTS As part of this study, the following publication has been completed: Freeman, J. and S. Bissell, 1979. Bats. Colorado Outdoors, 28(2): 1-5. Portions of the data were presented to the Colorado Chapter of the Wildlife Society and to the Colorado-Wyoming Academy of Science in 1981. Currently we are analysing periods of bats in various food habits, reproductive patterns, habitats throughout the state. and activity We have cooperated with the Denver Museum of Natural History by providing scientific and educational material for the development of a permanent exhibit on bats at the museum. CONCLUSION This study has demonstrated: (1) range increases M. californicus, 'Pipistrellus hesperus, Antrozous of Myotis yumanensis, pallidus, Lasiurus cinereus �··116 and Lasionycteris noctivagans, (2) provided over 325 standard museum specimens for future study, (3) indicated that Tadarida brasiliensis is a resident species of the state and analyzed the pesticide load of the species, (4) validated habitat associations, and (5) provided scientific and educational material to the Denver Museum of Natural History. LITERATURE CITED Armstrong, D. M. 1972. Distribution of mammals Mus. Nat. Hist., Univ. Kans. No.3, 415pp. in Colorado. Monogr. Geluso, K. N., J. S. Altenback, and D. E. Wilson. 1976. Bat mortality: pesticide poisoning and migratory stress. Science, 194: 184-186. Meacham, J. W. 1974. A Colorado Research News, 15: 8-9. Prepared by: Jerry Freeman Wildlife Tech. I-B colony of Tadarida brasiliensis. Bat �" .AK'iI()" ".,,,,, 0 DH, h~~ tJ'-U • w.3, 'V"lIN~rq" JR.,." .711.." •• ° ,..... ,..... --.J ~/(,"O ,!,}", 0"''''' ;,..".A S••••• .~I'.I Figure 1. Locations (.) of areas trapped for bats. �107 r----------I--':'1---\-------T D' .•. I I r .J (I I. I-''I} • ,1 J r r ----' I (. ",.-_" , ./ I I 38 -i- -I-" ~ '> '""I) l. I ' j-----,-'-""\ \.1< I '; --------7---; I .:: I 1 I I fA. , .• ' I / I '---'--- I o 1 r"".... 'l., 1 I __ 50 \ \ --...1 • 0 •..,/ •.••••• 1 :. V,,"'" '"" •••• ~ I' -L - ••. 0 .J 1 ' l.,"'''''' J 1 I I 1 1 1 r-. ., co 1 ! I I -1'""--"""" I I I 1 L _..l I I I I ....• ) 1 - 105 Distribution of Myotis lucifugus in Colorado. For explanation of symbols, see text. ,_. ,_. I • I..J , J 1 '------T 1 _ I _ ~------l ----- 100 Miles 107 Figure 2. .,...._ -<In..l '----I 1 I ,,...,___ ,.oJ , ..• 1__ /'"I 40 I 1 I , .: i I .J ......_,.,""4..----..... I'I I I I I L------t---l-I! 0' I I' L1 - ~ , 1 I: I----L--.L , 1 , i : "'-------r J 1 ---- _-, __ .1. 1 -(----...1""1---1'. I, 1 I, II! 1 .1 ", .. . ..J I' ~ \ ---:_-~-.I{...' 1 ••. " I ~--------~ -I.., 'I I - -i I I1 , (~~ : ----+---,---, I I -,-- I ( \.._r'-t- ' 1 I : ••. 1 _J •••..•r------ _5-...r .•••;r 1 rJ ...• --"? r{\._.... I 1 .)-'---1 '-(/--- ....J 1------ .. I 1---- ..... 1 -----..J----1 I, • I1 ,(DJ I I _ 1 J..- r-; I ',}-.... L _.J " ••• \'"'\. r........'1,I L, 1 1 .1-------- L__.L ..J' ,---- ,..---- ./ \.,. [----1I I1 --------,--- ---- I \ 't- •...••••.'"'-_. .•.)" I----~---- __ L__,: 40 103 105 103 38 �105 107 103 --1--'.:---,-------1---1 ,JI I I }1 "rr-r+7'...,I...,I...,It---L __ , ,I •• .A. II---...r---- J----' 1 •• r r--'J.L J I. 38 ( 1-- -- c, I A 1.-----\ r J- t •••• L., I~, ..• 1--------7'---1 1 ,J I I I "I , ~ ~~_~_ o :' A--. I I 50 '--------7 I 1 .I -L _ " _ 1 1 I 1 , I 1 1 1 1 I .-- I _j' II ~ J l I I 40 I I J i l f-' f-' \0 I I -------i I I ,I ,h-----------, I I "'I, _ i,...(, I I I I 1 /L-' I I I ,..I I I" I ' I ---t---'"'j--J I , L _ 100 Miles 107 Figure 3. I - '-----\., \•...\_/" I,,, , L ..J-...._~ 1 I 1 I---L __.L I' , I. ..1. I : ~ 2.J.,. r I.... )..J .J... , J r, 1 I L.. J I 'I.". I'" I---------~ ...." )-----1 I _, __ I --.... I -1------t---I-- I "I \ ---------' I -\,-.. ': ...1 ...J __ l, I . ( L <:I I I I ,_ ,_,-"1 J-..... I L .." I 1 1 I ----I ---- 1 rJ r----"1 }' {-,r--~ ----..,,/..... .) I \' (~ I I,' 1 \..... ·J'-t,_" I I \ ," .•...• ,.J ---- \ r---' I I I I •...••...\ r"""'-", "1 r1 \._..... I •••• "" __ ) \, •• '" r--- ---, , , 1 I IA 1------ , {\ ( I r 40 I 'I ---------,--I 1 105 Distribution of Myotis yumanensis in Colorado. For explanation of symbols, see text. 103 38 �103 105 107 --1-----,--I ,Ir-----. , 1 I • •.••. I. ' I ----1.J----1 It I I I I I 40 I _______ -l , -+ I ----: I I L_~ , ,II , 1 38 I 1 I 1__ L- - - J I I 1 1 I I 1 L ,/ J N 1 J ~ 1 I I 1 ' 1 <, 1 1 I .....•...I •.....• o I 1 -1 I~ ___ -... 40 I i Ir------l ! I, I I ______ I I I I I ,I 1 I ' I I I 1 ~----L-T-J----~ I I I I t1 1 ______1 o 50 100 Miles 107 Figure 4. 105 Distribution of Myotis evotis in Colorado. For explanation of symbols, see text. 103 I I I ---...J • 38 �107 103 105 r---------,-,----,-------,---1 D r ,J I •• ' , ---.... -- _ _,-----.J ' I •••, (I r1""'\. --.. t-- ~ '" - " \... -"~ 1-----1 L....) J , .r< r .,..... , )• r -'-.... ...J --------7'---. '" ./ ( I I I " I I,- .,' __l.. I o :' I I 1 I I---L __ J.. I I ~ \ - -' I I I"0~ -1 -.... ..1_••••. II II I I I ,'L - -,' I , _L i I s f"\ I I r '1 , I h----------~ ---r-- . ..I I L' I I , I j...,""'? ,/,,)J -L - ' I, I I, t. I,..."" i, ,'. ,_. N ,_. I 'I I , _J 'I I \. 40 1 r ------l' L------.l.--, l-I, s,J I, , ! -1------,---:- , , , _J I '1 I , '. _ ____.!-----J 100 Miles 107 Figure 5. I II II ...I I \""'~l/,J1..----.... I I 1 I ' : " »<: '----'---7 ,I' ...L----L-T--'-----.I 1-, ~)O I "------ I ,I.. 1I I l _--, __ ..1. L\ I 1---------1.. 1 -----, I ,.,1---1 \ I I , I , -.J----1 '----"-, , .) \..-,.,. .... -::"-""-.. ----- 38 ""'", I J- I I" l I \ __ ,.-J?" I .A.. ••• _J" '----"'"1 (I 1 I 'I, I r-- ---rL--.L---... J..--,-"1 I , )0..... I L , ..._ \ r " .•.. ~ L, ------- r.J ,-----" I I, rJ' I 1 "l I •• \ \') I I 'r----. I ·1I l .•._ , .A , I, I (I I , .A ------------'---, 40 "" 1 ---------,--. I ,1 105 Distribution of Myotis thysanodes in Colorado. For explanation of symbols, see text. 103 I 38 �107 105 103 ~~I-------T-----~=--j ~~I 1 r---..... I' J II : , 40 _J ----1.J----1 I I ' II I_..J I I I, _J II I -1-- I I --L----t:I I L_ _J I I I , I ...")...... J 1 ) / I I I J L I .I ____ ... I' 1 I ~ t-' N N , I I ..1. 38 L- - - J i ' Ir------l ! I I : 40 ' _1_ - - - - -+ - I I , I I I I _______ I, I I ~ I I . , I I I I I I I I I 1 ' I ~----L-T--J----~ 1 I I •.••1 I t I I I 1 I ____________-----J 1 I o 50 100 Miles 107 Figure 6. 105 Distribution of Myotis volans in Colorado. For explanation of symbols, see text. 103 38 �107 105 103 ------,--I r----------,---c:.:-'1----y-------:-- D .•. .•. I I, ;1 r I 40 •• I 1 I --., J '_1r r-- ---.... ,--- I 1- - - - I 1---r-"1 ,•.•. ,\ . r " )-,~ '- ..." <c (/--- - '" r1"'"'\.-.... (\I 1 "'_r'-t- -~.... •... ;:" ( 1 ILl I ii, ...1_•.• _-1 I <- I "\ J 1 I -- •..') J ",. ~---~ L ') 1 ')...i< (J...,- 38 ,} ----,_,-'"\ I '.. ------7---"1 I I/ 1 \ ~\,. ..... ""'1.----.... t L----r---7 -. "IN 1 -L_ , - ---------'---\ o 50 / I 1 ~~""'''I I1 I •....• 1 N I W I r--- I I 1 I I , ,....,-- r' ,1 _ 1 I I I I 1 I L __ _l I ..., I,.,.) 1,/ I !I t,I I •. I1 I. I _ ~.1 (\..-____ _: ---,---- 100 Miles 107 Figure 7. I ,~I I I I I---L-_.L I I 1 ", I, ./I r ,I ,'\., '\_----\ I I-------l II L_.J: I1 1 I 1 ...1.------ ,------ 1 I I , ~ _______ s,J ,1 J I. 1 ..1. I I, ,I 1 1 40 , 1 -I- - ' 1 I _J I I 1I I •.•• 1 1 -------1 I I I 1 ----, 1 (I ,. L L, J -----',,,~ f ~ I 1 11I I 1 I --- I , J _,---- 'r 1 1 '----""1 (, 1 l, ...J r------ 1 J I --J---- LI J '). J \ 'J.. ••• -."......... _j I I \ I •• 1 __, I-:------------L.. ' -.A I (I 105 Distribution of Myotis californicus in Colorado. For explanation of symbols, see text. 103 ..J 38 �105 107 103 40 40 'I-' N .j:-- 38 38 o 50 100 Miles 107 Figure 8. 105 Distribution of Myotis leibii in Colorado. For explanation of symbols, see text. 103 �' I. \ . r----------! D .•. r \ II 1 I .•._______ , -----'---,.A. -;.a I I 40 ~~)---\------:-------:-----~--- ,.J ~ • J,-- -,I .•. 1 , I16___ -' _J /( r-r- • I I • '( 1 ) I ...,.._--,._J/" ....J.I ( ""--..\._r'\..~'- ( I I -, ____ .•. '-. , '\"1 _________ I r-----, • -'_"", ,_ ......1 rJ-.. ')..J< ~ ,.... ...,oJ tA I~, 1----L ,,,,. -- I ,... o I I II I) 50 ~____, __ 1 I , -lI I I I I I• L_..J I , I I 1 r------l I I t-' N lJ1 I r--------l ..l I J I ' I ,....,----------- I I I r.ll \'••.. /",,'"''-1 ~ •.. ---•••••• ;, --7 /~-, 1 I/ '-'I. ~ I I 1 I 1 I' I ' I I I \.. \ I J I I I '--4 I +---1---I I -- 40 I I } • J t> I ________________I 1 _j' I I1 i I1 I 105 Distribution of Lasionycteris noctivagans For explanation of symbols, see text. 38 I I I I I 1 L 100 Miles 107 Figure 9. I. ----- -,,""" II ----I ~ I.. 1..,-----L"I" I I 1 1 I '\.. I I I ...J_ '\.._\ I. _____,___________ I' --- ". I '-_ I I --- I I c; )') 1---:;&_- I .:r--":, I,' -., L 1 I 1-- .••. '" • ./ _:._ , 1 "L ., ~J 1 (\ I I I. r--.----'----....J ,-"1 I I l,__, ("I I '( -----..J----1 I : I I I r 1..1.. -,-- ~ •.•.••• 1, ~ _J-..... 1./ ~ I• 38 '\ '-(" '..... "f" (f---J 1 I -, I_____..-"'\_ I L I1 I • - r---' I : I I I ---"1 I I 1"""""- I I J 1I 1.___ 1 \ \.1 ~' J \, I '" -~"""'_-~ 1 I 103 105 107 103 in Colorado. I J' I I I _J I �---- I ------- Dr 40 ---,--- I -';-, I I ) I ~--~--"rJ \----.., .•••. I' - _- I I I I __ ----, ~ I, r- -, r '-.< -A , , ( I r f 1----4I /1 "f-- '1.--\......1' -.... I I ll,...., --, I' I •••;:.... ( " / '1---' 1"- - ----\_,-"' -, ----7'---"1 I I I ---.....-I-.J.L I" .II (J lJ/ , , '\\ --' __ /, r' I I I 107 I,I _ ' l, I I L_ I r I I I __ I II""'} I . J-,-----, I ----- I I ,, ..J T - f.\ \.l:..J I, ~ !I ,I I (J\ I , _~I N _I J t> t-' I , I .f I, _ -~-L-----t---.--J" : 40 I _ _._ _-' _ _ 100 Miles 50 10. I" I J ""------ 1_ ----L _ 'II J L 'I I I L_..J I I I _ I I I o I II I -' I .1 I I I •..• I \\ 1-----,I ,,,I ')\.r, rJ-,')., ~c,-i.---- __ ~- I __.1L...•. ~ ... I I L----r-Ir --L 38 Figure ..J -, J' __ ,:::-_s< I I ----~I - __ I iI ----1I, I , L I , I ~ r I, r---L, ----14 ------- I -----;~ " II : , , -----,I ---I I Ir- ----- I(, - ~( - iI r-'" )--- -.., , J L__ l._ __, J. •.•.. [ L r r '~"L, ..J' ,---- \. I -------~--"I ' .•• -------I I ---- \ ( Ir ,'----- 103 105 107 105 Distribution of Pipistrellus hesperus in Colorado. For explanation of symbols, see text. 103 38 �-_ --1 ----- D.r--.• -------~--, , .•• I' - 11 ..... J --- J ,.. "-, \... " ~, r (/--- ' "J < ~( - • - ,--- 1 1 ',,_,, __' ' +__ I 1 I 1 J"-L.. 1 .., _ I , 1I ' I, \ L_____ 1 " I:, -,.1 --~-,< - I ,;1 J, 1 . v. l _~ 4 -----_') .\_ ' ').,,< r -, 'I., ,~- --,-It--., 1 " ••• -? I1------.--'..... ' ------7---; -", " I -'_J 1, '.... -'(1 I---~-1 1 -J'I .L-----L_ L' 1- J - -'- o - J -- \. I L..__ '. • - 1 ~ ,I 40 I II 1 I 1 r------ 1 ,I , 1 l ~__ j _~), 0 _ 1 _~ , , I, 1 r'1-,----, : __ "'1I II ~__ .: __ __ --- Distribution of Eptesicus fuscus in Colorado. For explanation of symbols, see text. i ...,1 Ii , I ",. _ , 105 I ,L _ T- ...1 100 Miles - 1 __, 50 107 Figure 11. I I, I- , "r ,_ I.... .' ·~--~-~ -~---, U ~..._v: J --- -)--, l.,.•• 1---- ,'... __ -- I i 1 ( 11 1 __ ...I 1 , 1 , -I , I 1 1r__ 1" '.J----1 1 ~____1L_.J '(, ,--1 --,--- .__ 1 1 -'__, __ _- 11I ~--- I _(\--)'~.'! ,(\ ~ ..'I 1 ~ I,---- A, , r+: 1---,--...l ,,' /1J \.--\.•••• ,,1-<,-..." "'....... ,.-_ 1 0rJ (0' ,I. ('.I ' I '----- ., 1 __ L r- • JI 1 .J - \ I ---j .', I,-:1.- - .•• ,---- - :1.--- - -- 38 (' '. I1------ 40 ( ) r ' ---1.1, -----'. ----11 11 1 ------- I. ~--,-\ ... -';'" : _..A.' • ••• • ••••• 103 105 107 103 __ _ -' 38 I--' N " �107 r------- DI I' ---- A )11 1 I" I\,. r1 , -------~--,---{ e, -------11 -l , JI \.--~-1 A ------ 40 -';" ---,- 1 103 105 ',- 11 1 el 11 1,--- , 1 1 (, ' I, ,-----.1-I,1 -----, • r' I L11 ----11 "----1,I 1 1 ~ -----'---,- .-- .I "'"'1 , I, 1- I, 11 1 ' , 1---,--,b --------, --' 1L rJerL--L-,--" _h,' I'I -,(~ ------1---, ---&---~ ' , \.J. _ ''(I 1 1I 1I .- I ...1 ( , A -~ ------"f--' I' -~ , 38 , / I' ' ' r1 ,,-- ,.--" ------ \.. ••••1"'" I I ' -- J_, '---- ---',~ I -,& I o , ••'\ _ 50 ~ ---- - - I,I r r... I,,.,,.)J t> I I. , .__ ----- I _'I , , I I __ L : ..1 , I ...• 1, I 1 _,&, _ I, I _ , 100 Miles 105 Distribution of Lasiurus cinereus in Colorado. For explanation of symbols, see text. ,..... tv 00 II , __ I , I '------, I I , I ' 1 , , I' .....•... J / , .••.•1~__ _ r I I ------l __ ...1. I I \ ~"" ,"" 1r- , •... \.." l _ ••-__ 1 I I " I- -L J..-----L_ -- I I \ 107 Figure 12. 1 '------\, \ I I' -,' • ,---I ,--- cw ;- [.) ' ".( r J "'''',.,'~---I-J-' ,_ , r-----,~-V I' -----7'---"1 -', I' I I I I '---l I r/ I L_.l I ( '" -I-''I" --- , 1 ,J.. I' -~------~-I-,& '-, ,&' ---' I ~ 1 1 .i.; I' \... '\, 1 I1 1IIl, {\ 1-- '::, (I I --,I !... 'I 'J ~1 J, j --?--7~ I'l ~ ' /--- 40 103 , I -' _ 38 �103 105 107 ~i -------l-----~=--J, I1 '"lJ r , , , 40 r=r=':' I I I I I I I -----.J----1 1 r-- ----_!---I-....I I ______ __ -...J , 1 1 -1- - I I I I I , -,-- I --- 50 Figure 13. I - -I I I 1 I I •.....• N ..0 I I I I 1---'-- j-------I I L J I I I --l--- 100 Miles 107 I J - r-" , o I I j 38 I 40 • Ir-------l ! , < <4: I I I I I L_.J I I ~).).:'J.J. I I I 'r.""""'->.,.V-----I I I I I I - I 105 Distribution of Plecotus townsendii in Colorado. For explanation of symbols, see text. 103 I I 38 �103 105 107 r---------,-,---,-------T-I "'I .•• ~ 40 I, I .& I (I ,J ,. . .& .•• ------ I _J r-I I___ ...r----.J ----' , I I ~ "', (;- "'" __ ..J1 (I f-. \"_J" I '< _-, __ _L I.J J L_ 1 ,I 1- I.) "'():.J ----, , -'-""\,. •... Y -----7'---'1 r / I I 1 I J... r'........ 1-, I I I I ~~ 14. : I '-----\. ,I I I II I I I I I -L" -.~ 1 I -:". .•. I I I II I I I I I r- ------ll 1 ,_.. I I I I j- I I h-----T----~ r..l 1 ~ I ..! _ ) I __ -- 105 Distribution of Antrozous pallidus in Colorado. For explanation of symbols, see text. w o I I __ jI 40 I : _." ,,'.I I I I "'"1,../ """'-I•• __ l I" I 1_ _L---J 100 Miles 107 Figure I I __l__. 50 1 --------L I •.. I I---L-_J... I I l , o ~ 1 ..J. I 1 I Lj ~----I-... --I L I I I I I + 1 I _-' _t I ! _____ ~_.r< 38 1 I I I I I -I., .•... 7 "'" (I ~J .1 1 \ •••- _..,..__ I r-J/~i' ( --.. I I ----1..J----1 ----..... J---r-""1 )0..... 1 LL, r......--J ,\ •.•.• I I I r1'" \. I } I I I 1 I I ~__- .... I 1 J '----""1 (I I •• ,--- 1 I II I rL--l.-I--' rJ I I~--- \ \') \. , -------- I I I 1"--.,,_-J \ 'r I I 1 L.. __ .•• ''& I --- 103 38 �107, 105 103 r----------,--,---,--------, -------1-I"', D ,. ; I------------'---, . I' 'I ----' rJ I 40 I I" I" 1 \.1'•••••••"'_- 1 ••••, I I 1-- I --i , ~ I--------7 ~ I, I .. f .I I ••• 1 ---1 1 I I , ...., " . / o l--~ J' ') '\'(,., ,•.1 I -'-", I ... -----, I "'", , I I J t...., ' _ •.---...... l -L' - }- J II h-----------, I I .•, ~,~ - 1 ,I L. -T- _I , -., t'I 1,,- I..... 1, , , , L- I I ~ , I ')...... --- t-' W t-' I , ,,"" \ I , J , 1 I I j __ I-------l I I '-'-~ ,,/,}.1 I 1-_J I '- ,,-v 4 -------·7 I I 1 I I I, , I, I r------t---,-- 'I I' ''', ,I, 50 .._ 'I 40 , : I I I I ---------t----,---- I "'\ •••.• I I----L--..L. II' I I •..•• I I .I 1, _J---1' I ---- _J _______, ..J_""_..j ~\~, 1-, 1 •••• -- __ I, ! ~."------ I '. __J-----..J _ 100 Miles L____.. 107 Figure 15. '-........J ....(I:J~" \ , ...J l --.,--.1----1 -loa.. I, l. L L, I ',~ , \~. r " J......, '-~ r ( (~\.. \'_J',t- 1 ), 'r------ I---r-, .••• \'. 1---------1 1-----1I I , -5,...._,..,. I II ...! I 38 - "I L, \ r---.....• rJ' (f..---..J /1 \._.... I J I '----"'1 (, ,~, !------~---~, ... J \" I ,--.,' rL--L---... I I '). I 1 I I 1---- I \ I I ,'"-- I I ~---~ I I 105 Distribution of Tadarida brasiliensis in Colorado. For explanation of symbols, see text. 103 , 38 �132 Appendix I. Locality data of bat species. The first number in the parentheses refers to the number of individuals examined per site; the second, the number of individuals collected and prepared as scientific specimens per site. Myotis lucifugus 1. Bell, 13 mi. N. of Durango, La Plata County (R9W,T37N), CO. 2. Elk Springs,S mi. SE of Elk Springs, Moffat County (R9SW,T5N), co. (4-4) (2-2) Myotis yumanensis 1. Florida River, 12 mi. S. of Durango, La Plata County (R9W,T33N), CO. (1-1) 2. La Sal Creek, 4 mi. W. of Bedrock, Montrose County (R19W,T47N), CO. (3-3) 3. Douglas Pass, 2S mi. N. of Loma, Garfield County (R102W,T5S), CO. (1-1) 4. Slickrock, 1 mi. SW of Slickrock, San Miguel County (R1SW,T44N), CO. (5-1) 5. Cow Canyon, 16 mi. SW of Kim, Las Animas County (R56W,T34S), CO. (4-0) 6. Douglas Creek, IS mi. S. of Rangely, Rio Blanco County (R101W,T2S)CO.(1-1) 7. Monument Gulch, 16 mi. NE of Rangely, Rio Blanco County (R99W,T2N) (1-1) S. John Martin Dam, 3 mi. S. of Hasty, Bent County (R47W,T35S), CO. (6-6) Myotis evotis 1. Billy Creek Refuge, 10 mi. N. of Ridgeway, Ouray County (RSW,T47N)CO.(1-1) 2. Leroux Creek, S mi. NW of Hotchkiss, Delta County (R93W,T13S), CO. (2-1) 3. West Creek, 4 mi. NE of Gateway, Mesa County (R103W,T15S), CO. (1-0) 4 .. Douglas Pass, 2S mi. N. of Loma, Garfield County (R102W,T5S), CO. (9-3) 5. Alta Vista Ranch, 2 mi. SE of'Greystone, Moffat County (R100W,T7N)CO.(3-1) 6. Elk Springs,S mi. SE of Elk Springs, Moffat County (R9SW,T5N), CO. (200-42) 7. Meeker, 6 mi. SW of Meeker, Rio Blanco County (R95W,TlN), CO. (4-4) S. Sand Mountain, 5 mi. NW of Milner, Routt County (RS6W,T7N) , CO. (1-1) 9. Blacktail Mountain,S mi. E. of Oak Creek, Routt County (RS4W,T4N)CO.(1-1) 10. Pot Creek, 7 mi. SW Gates of Ladore Ranger Station, Moffat County (R103W,TSN), CO. (S-O) 11. Coon Hollow,S mi. W. of DeBeque, Mesa County (R97W,TSS), CO. (1-0) 12. Holland Draw, 6 mi. SW Greystone, Moffat County (R101W,T7N), CO. (4-4) 13. Douglas Creek, IS mi. S. of Rangely, Rio Blanco County (R101W,T2S)CO.(1-0) 14. Monument Gulch, 16 mi. NE of Rangely, Rio Blanco County (R99W,T2N)CO.(1-0) 15. Lily Park,S mi. N. of Elk Springs, Moffat County (R99W,T5N), CO. (3-0) 16. Graveyard Gulch, 8 mi. NE of Sagauche, Sagauche County (R9E,T45N),CO.(1-0) Myotis thysanodes 1. Meeker, 6 mi. SH of Meeker, Rio Blanco County (R95W,TlN), CO. 2. Elk Springs,S mi. SE of Elk Springs, Moffat County (R98W,T5N), CO. 3. West Creek, 4 mi. NE of Gateway, Mesa County (R103W,T15S), CO. (2-2) (2-1) (1-1) Hyotis volans 1. Yellowjacket Canyon, 1 mi. NE of Ismay Trading Post, Montezuma County (R20W,T36N), CO. (1-1) 2. Cahone Canyon, 4 mi. SW of Cahone, Montezuma County (R1SW,T39N), CO. (1-1) 3. La Sal Creek, 4 mi. W. of Bedrock, Montrose County (R19W,T47N), CO. (2~2) 4. Cottonwood Trail, 12 mi. NE of Nucla, Montrose County (R14W,T48N),CO.(1-1) 5. Fish Canyon, 11 mi. SE of Powderhorn, Hindsdale County (R2W,T45N),CO.(1-1) �133 6. Gold Basin, 6 mi. S. of Gunnison, Gunnison County (R1W,T49N), CO. (1-1) 7. Leroux Creek, 8 mi. NW of Hotchkiss, Delta County (R93W,T13S), CO. (3-3) 8. Grand Mesa, 6 mi. E. of Lands End Observatory, Mesa County (R96W,TllS), CO. (1-1) 9. Douglas Pass, 28 mi. N. of Loma, Garfield County (R102W,T5S), CO. (6-1) 10. Alta Vista Ranch, 2 mi. SE of Greystone, Moffat County (R100W,T7N)CO.(2-1) 11. Blacktail Mountain,S mi. E. of Oak Creek, Routt County (R84W,T4N)CO.(1-1) 12. Escalante Creek, 14 mi. SW of Delta, Delta County (R13W,T51N), CO. (3-0) 13. Pot Creek, 7 mi. SW of Gates of Ladore Ranger Station, Moffat County (R103W,T8N), CO. (3-0) 14. Holland Draw, 6 mi. SW of Greystone, Moffat County (R101W,T7N), CO. (2-2) 15. Douglas Creek, 18 mi. S. of Rangely, Rio Blanco County (R101W,T2S)CO.(3-0) 16. Monument Gulch, 16 mi. NE of Rangely, Rio Blanco County (R99W,T2N)CO.(4-4) 17. Graveyard Gulch, 8 mi. NE of Sagauche County (R9E,T45N), CO. (4-1) 18. Elk Springs,S mi. SE of Elk Springs, Moffat County (R98W,T5N), CO. (47-8) 19. Lily Park,S mi. N. of Elk Springs, Moffat County (R99W,T5N), CO. (1-1) 20. Sand Mountain,S mi. NW of Milner, Routt County (R86W,T7N), CO. (3-2) Myotis californicus 1. Holland Draw, 6 mi. SW of Greystone, Moffat County (R101W,T7N), CO. (3-3) 2. Monument Gulch, 16 mi. NE of Rangely, Rio Blanco County (R99W,T2N)CO.(1-1) 3. Lily Park,S mi. N. of Elk Springs, Moffat County (R99W,T5N), CO. (4-4) 4. Elk Springs,S mi. SE of Elk Springs, Moffat County (R98W,T5N), CO. (41-3) Myotis leibii 1. Elk Springs,S mi. SE of Elk Springs, Moffat County (R98W,T5N), CO. (55-15) 2. Slickrock, 1 mi. SW of Slickrock, San Miguel County (R18W,T44N), CO. (1-0) 3. Holland Draw, 6 mi. SW of Greystone, Moffat County (R101W,T7N), CO. (3-3) 4. Monument Gulch, 16 mi. NE of Rangely, Rio Blanco County (R99W,T2N)CO.(2-1) 5. Lily Park,S mi. N. of Elk Springs, Moffat County (R99W,T5N), CO. (4-4) Lasionycteris noctivagans 1. Florida River, 12 mi. S. of Durango, La Plata County (R9W,T33N), CO. (2-2) 2. Yellowjacket Canyon, 1 mi. NE of Ismay Trading Post, Montezuma County (R20W,T36N), CO. (1-1) 3. McElmo Canyon, 11 mi. W. of Cortez, Montezuma County (R18W,T36N),CO. (2-2) 4. Cahone Canyon, 4 mi. SW of Cahone, Montezuma County (R18W,T39N), CO. (1-1) 5. La Sal Creek, 4 mi. W. of Bedrock, Montrose County (R19W,T47N), CO. (2-2) 6. Cottonwood Trail, 12 mi. NE of Nucla, Montrose County (R14W,T48N),CO.(5-2) 7. Billy Creek Refuge, 10 mi. N. of Ridgeway, Ouray County (R8W,T47N)CO.(5-1) 8. Fish Canyon, 11 mi. SE of Powderhorn, Hindsdale County (R2W,T45N),CO.(2-2) (1-1) 9. Gold Basin, 6 mi. S. of Gunnison, Gunnison County (R1W,T49N), CO. 10. Rulison, 1 mi. E. of Rulison, Garfield County (R94W,T6S), CO. (3-3) 11. West Creek, 4 mi. NE of Gateway, Mesa County (R103W,T15S), CO. (1-0) 12. Douglas Pass, 28 mi. N. of Loma, Garfield County (R102W,T5S), CO. (12-1) 13. Blacktail Mountain,S mi. E. of Oak Creek, Routt County (R84W,T4N)CO.(1-1) 14. Mountain Ute Indian Reservation, 9 mi. SW of Towaoc, Montezuma (3-0) County (R19W,T34N), CO. (2-0) 15. Escalante Creek, 14 mi. SW of Delta, Delta County (R13W,T51N), CO. 16. Holland Draw, 6 mi. SW of Greystone, Moffat County (R101W,T7N), CO. 0-1) 17. Elk Springs,S mi. SE of Elk Springs, Moffat County (R98W,T5N), CO. 02-2) �134 Pipistrellas hesperus 1. Yellowjacket Canyon, 1 mi. NE of Ismay Trading Post, Montezuma County (R20W,T36N), CO. (4-4) 2. McElmo Canyon, 11 mi. W. of Cortez, Montezuma County (R18W,T36N),CO. (1-1) 3. La Sal Creek, 4 mi. W. of Bedrock, Montrose County (RI9W,T47N), CO. (2-1) 4. Escalante Creek, 14 mi. SW of Delta, Delta County (R13W,T51N), CO. (4-1) 5. Douglas Creek, 18 mi. S. of Rangely, Rio Blanco County (R101W,T2S)CO.(2-2) 6. West Salt Creek, 19 mi. NW of Mack, Garfield County (RI04W,T7S), CO. (8-8) 7. Monument Gulch, 16 mi. NE of Rangely, Rio Blanco County (R99W,T2N)CO.(1-0) Eptesicus fuscus 1. Florida River, 12 mi. S. of Durango, La Plata County (R9W,T33N), CO. (1-1) 2. McElmo Canyon, 11 mi. W. of Cortez, Montezuma County (R18W,T36N), CO.(l-l) 3. Cahone Canyon, 4 mi. SW of Cahone, Montezuma County (R18W,T39N), CO. (1-1) 4. Cottonwood Trail, 12 mi. NE of Nucla, Montrose County (RI9W,T47N),CO.(3-2) 5. Billy Creek Refuge, 10 mi. N. of Ridgeway, Ouray County (R8W,T47N)CO.(2-0) 6. Escalante Creek, 14 mi. SW of Delta, Delta County (R13W,T51N), CO. (4-0) 7. Rulison, 1 mi. E. of Rulison, Garfield County (R94W,T6S), CO. (1-1) 8. West Creek, 4 mi. NE of Gateway, Mesa County (R103W,T15S), CO. (2-2) 9. Douglas Pass, 28 mi. N. of Lorna,Garfield County (RI02W,T5S), CO. (1-0) 10. Alta Vista Ranch, 2 mi. SE of Greystone, Moffat County (RI00W,T7N)CO.(2-1) 11. Sand Mountain,S mi. NW of Milner, Routt County (R86W,T7N), CO. (2-1) 12. Coon Hollow,S mi. W. of DeBeque, Mesa County (R97W,T8S), CO. (3-0) 13. Holland Draw, 6 mi. SW of Greystone, Moffat County (RI01W,T7N), CO. (11-4) 14. Douglas Creek, 18 mi. S. of Rangely, Rio Blanco County (R101W,T2S)CO.(1-0) 15. Beecher Island, Grange Hall, Yuma County (R43W,T2S), CO. (12-4) 16. Elk Springs,S mi. SE of Elk Springs, Moffat County (R98W,T5N), CO. (9-2) 17. Lily Park,S mi. N. of Elk Springs, Moffat County (R99W,T5N), CO. (2-2) Lasiurus cinereus 1. McElmo Canyon, 11 mi. W. of Cortez, Montezuma County (R18W,T36N), CO.(l-l) 2. Cahone Canyon, 4 mi. SW of Cahone, Montezuma County (R18W,T39N), CO. (3-2) 3. La Sal Creek, 4 mi. W. of Bedrock, Montrose County (RI9W,T47N), CO. (2-2) 4. Cottonwood Trail, 12 mi. NE of Nucla, Montrose County (R14W,T48N),CO.(7-2) 5. Mesa de Maya, 12 mi. SW of Kim, Las Animas County (R54W,T34S), CO. (1-1) 6. Billy Creek Refuge, 10 mi. N. of Ridgeway, Ouray County (R8W,T47N)CO.(5-1) 7. Gold Basin, 6 mi. S. of Gunnison, Gunnison County (R1W,T49N), CO. (1-0) 8. Rulison, 1 mi. E. of Rulison, Garfield County (R94W,T6S), CO. (3-3) 9. Silt, 3 mi. S. of Silt, Garfield County (R92W,T6S), CO. (1-1) 10. Douglas Pass, 28 mi; N. of Lorna,Garfield County (R102W,T5S), CO. (5-1) 11. Alta Vista Ranch, 2 mi. SE of Greystone, Moffat County (R100W,T7N)CO.(1-0) 12. Escalante Creek, 14 mi. SW of Delta, Delta County (R13W,T51N), CO. (1-1) 13. Cow Canyon, 16 mi. SW of Kim, Las Animas County (R56W,T34S), CO. (1-0) 14. Elk Springs,S mi. SE of Elk Springs, Moffat County (R98W,T5N), CO. (30-7) Plecotus townsendii 1. McDermott Arroyo, 18 mi. S. of Hesperus, La Plata County (RllW,T12N) CO. (1-1) 2. Yellowjacket Canyon, 1 mi. NE of Ismay Trading Post, Montezuma County (R20W,T36N), CO. (1-1) 3. Silt, 3 mi. S. of Silt, Garfield County (R92W,T65), CO. (2-2) �135 4. Elk Springs,S mi. SE of Elk Springs, Moffat County (R98W,TsN), CO. (2-1) 5. Holland Draw, 6 mi. SW of Greystone, Moffat County (RI0IW,T7N), CO. (1-1) 6. Monument Gulch, 16 mi. NE of Rangely, Rio Blanco County (R99W,T2N)CO.(1-1) 7. West Salt Creek, 19 mi. NW of Mack, Garfield County (RI04W,T7S), CO. (2-2) Antrozous pallidus 1. Picture Canyon, 10 mi. SW of Campo, Baca County (R47W,T35S), CO. (20-0) 2. Alta Vista Ranch, 2 mi. SE of Greystone, Moffat County (RI00W,T7N)CO.(1-1) 3. Elk Springs,S mi. SE of Elk Springs, Moffat County (R98W,TsN), CO. (18-6) 4. Cobert Canyon, 16 mi. SW of Kim, Las Animas County (Rs6W,T34S), CO. (4-4) 5. Cow Canyon, 16 mi. SW of Kim, Las Animas County (R56W,T34S), CO. (2-0) 6. Monument Gulch, 16 mi. NE of Rangely, Rio Blanco County (R99W,T2N)CO.(s-3) 7. West Salt Creek, 19 mi. NW of Hack, Garfield County (R104W,T7S), CO. (2-1) Tadarida brasiliensis 1. Orient Mine, 7 mi. E. Junction of Highways #15 & #17, Sagauche County (RI0E,T46N), CO. (165-60) �136 Appendix II. Results of pesticide analysis of bat guano from the Orient Mine free-tailed bat colony. PATUXENT WILDLIFE RESEARCH CE~!ER - ANALYTICAL REPORT - PR-2l22 Submitter: Don Clark, Project Leader, Population Ecology Project, Patuxent Wildlife Research Center, Laurel, Ha ry Land , Specimen Data: Guano samples from a colony of free-tailed bats at Orient Mine, Saguache County, Colorado collected on 19 August 1980 by Jerry Freeman. Results: ppm dry weight; organochlorine compounds. reportable residues = 0.1 ppm. Sample No. 81-B-1 81-B-2 8l-B-3 Compound p,p'-:-DDE Submitter's No. Lower limit of Whole Dry Wgt.,g 1.065 8.588 7.034 1 2 3 81-B-1 81-B-2 81-B-3 0.48 0.33 0.31 p,p'-DDD p,p'-DDT Dieldrin Heptachlor epoxide Oxychlordane cis-Chlordane trans-Nonach1or cis-Nonachlor Endrin Est. Toxaphene RCB Mirex Est. PCB - = none detected 0~ w. L. Reichel, Project Leader Env. Residue Chemistry �137 JOB PROGRESS State of Project COLORADO No. W-136-R Work Plan No. Job No. Free-tailed Covered: Personnel: Nongame 1 Job Title: Period REPORT 1 January 1982 through Investigations 3 Bat Study 30 June 1982 P. Svoboda, W. Graul, S. Bissell--Colo. J. Choate--Fort Hays State Univ. Div. Wildlife; ABSTRACT Results from April through June of the 1982 field season are reported. Mexican free-tailed bats (Tadarida brasiliensis) arrived at their roost in Colorado by 12 June. Population size was approximately 30,000 by the end of June. Feeding flights after 14 June are described. Bats from the colony were trapped with mist nets; 63 adult males and 1 adult female were captured. This Job Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the Director. ��139 FREE-TAILED BAT STUDY Peggy L. Svoboda P. N. OBJECTIVES 1. Determine seasonal 2. Ascertain colony status relative 3. Determine feeding ecology patterns: where do they feed, what do they eat, when do they feed, total amount of insects consumed by the colony. 4. Ascertain the relationship of the San Luis Valley to other colonies in the United States. 5. Determine DDT. whether and daily chronology the Colorado patterns. to size and reproduction. colony relative colony is experiencing problems with SEGMENT OBJECTIVES 1. Determine seasonal and daily chronology 2. Ascertain colony status relative 3. Ascertain the relationship in the United States. 4. Determine at Orient Mine. to size and reproduction. of the San Luis colony to other colonies the diet of the bats at the Orient Mine. INTRODUCTION A colony of Brazilian free-tailed bats (Tadarida brasiliensis mexicana) lives in Colorado during the summer. Armstrong (1972) was not aware of this colony when he reported only occasional records of T. brasiliensis and did not consider the species a regular resident of the state. Meacham (1974) reported that, in August of 1967, he found about 9,000 or more ~. brasiliensis occupying a mine which had been abandoned since 1932. This mine is located at the northeast rim of the San Luis Valley in the Sangre de Cristo Mountains. The size of this population was estimated to be 75,000 to 100,000 individuals in 1981. The colony seems to be predominantly male, but a few pregnant females and juveniles also are present (J. Freeman and L. Wunder, in prep.). Unique characteristics of this colony are: 1) the colony only recently has been established (Meacham, 1974); 2) it is unusually large for a primarily male, summer roost as defined by Cockrum (1969); 3) it contains the northernmost breeding records for ~. brasiliensis east of the Continental Divide; 4) feeding areas during the summer might include intensively �·140 farmed sections of the valley where many pesticides are being used; 5) the colony is located at the northern margin of the range of the species and in between the two easternmost populations of T. b. mexicana as outlined by Cockrum (1969) (fig. 1); 6) it is in a-mo;tane valley with widely fluctuating temperature and weather patterns (Ramaley, 1942). Prior to 1982, this colony had not been studied intensively. The location of the colony on the margin of the range of the species provides an opportunity to compare characteristics of a geographically marginal colony with the better known colonies of the Southwest. The purpose of this paper is to report preliminary results of the 1982 field season spent studying free-tailed bats in the San Luis Valley. METHODS AND MATERIALS A thorough literature search for information about T. brasiliensis and the San Luis Valley was completed by April 1982. This information was used to develop a research design and to outline work to be done during the 1982 field season. The mine was checked for the presence of T. brasiliensis beginning in March 1982. A bat monitor was used in April to help detect an out-flight. After 17 May, mist nets, the bat monitor, and entry into the roost were used to determine the presence of ~. brasiliensis. A night vision scope was used several times to detect emerging bats at the main pit of the mine. During the day, mines in the San Luis Valley by Crestone, La Garita, and Bonanza were searched for sign of T. brasiliensis. Once the colony began regular out-flights, 18 ft. and 30 ft. mist nets were set at several sites along the edge of the mine to capture bats exiting and entering the roost. Mist nets were set before the out-flight began and usually were attended throughout the night. When a bat was captured, its species, sex, and age were determined. I weighed all bats with a 50 g pensola scale, measured the forearm with a metric ruler and placed a white plastic band on the right forearm. Reproductive condition, occurrence of ectoparasites, and any abnormalities were noted. Additional data collected for each night of netting included the following: ambient air temperature, relative humidity, wind speed and direction, and time of sunset and sunrise. Size of the population was determined using a modification of a photographic technique described by Humphrey (1971). A Pentax 135mm camera with a flash attachment was mounted on a tripod. The camera was placed such that the main flight of bats went between the camera and a cliff of the pit. Ropes marked the edges of the photographic frame on the cliff. The camera was loaded with Tri-X 400ASA black and white print film. An electronic, digital stopwatch was used to time the bats across the frame after each picture had been taken. After dark, a headlamp and flashlight were necessary to observe and time the bats. Pictures were taken every 2 minutes during the main out-flight. Gaps in the flight after dark were detected with the help of a bat monitor. �141 -, I I A \ \ \ I \ \ I t t ,, ORieNT I -------- MINt * , I a ~m tie 100 ~ Fig. 1. Map showing approximate limits of behaviorally (and probably genetically) separate groups of Tadarida brasiliensis in the southwestern United States (after Cockrum, 1969). �142 The area where most of the guano accumulated beneath the roost was covered with a piece of black plastic, 12 m by 24 m. It was checked about every three weeks to measure the amount of guano which had accumulated. Some bats were collected for electrophoretic analysis. Heart, liver, and kidney tissues from the bats were frozen in liquid nitrogen. Tissues from Colorado will be compared with tissues from populations to the southwest and southeast to test genetic affinities among the populations. Guano samples for food analysis were obtained from individual bats by confining them to a covered, styrofoam cup until they defecated. The bats were then released. Other bats were collected for stomach analysis. Both guano samples and stomachs were preserved in 10% formalin. A large sample of guano was taken from the entrance to the roost and stored in a glass jar. Feeding sites were located by setting nets over water or between clumps of trees at various distances from the mine. These nets were attended at least until midnight and sometimes for the entire night. Hater sources included irrigation ponds, streams, and beaver ponds. T. brasiliensis caught in the valley were processed similarly to those captured at the mine except the plastic band was placed on the left rather than the right forearm. Bat activity at various sites was recorded with a bat monitor and tape recorder. RESULTS AND DISCUSSION I visited the mine once in March and once in April 1982. Although there was the distinctive smell of !. brasiliensis coming from the mine during northerly gusts of wind, no bats were seen existing from the roost in April. I checked for an out-flight at the mine 7 times between 17 May and 9 June 1982. No T. brasiliensis were seen or caught in nets set along the edge of the main pit at the mine during this time. On 12 and 13 June, bats were observed on the roost in large numbers. On 14 June bats were observed exiting from an opening above the main pit, and on 15 June, the bats were observed emerging from the lower part of the main pit. I began capturing I. brasiliensis in mist nets on 16 June at the main pit. The colony was 98% male, with only 1 female being caught in a sample of 64. This female weighed about 4 grams heavier than the average I. brasiliensis and was caught earlier in the evening than a male of the same weight. From superficial examination, this female appeared to be pregnant. No juveniles were captured. Three other species were captured at the mine: Lasiurus cinereus, Eptesicus fuscus, and P1ecotus townsendii (Table 1). Bats generally flew from the pit in a west or southwesterly direction. Concentrated flights flew mostly to the southwest while bats in the later, more dispersed flights tended to fly in a variety of directions, mostly �143 to the west. Most bats generally flew down the ravine leading southwest from the pit to the valley floor and around a prominent rock ridge standing to the northwest of the pit. Individuals of the main flight remained even with the height of the pit (about 2877 m) for at least eight kilometers into the valley. As a result, the column of bats flew at an altitude of about 439 m above the valley floor. Later, more dispersed fliers tended to drop in altitude before scattering in southerly, westerly, and northwesterly directions. The time of emergence relative to sunset was difficult to quantify. Many times the sunset was obscured by dark clouds. The pit was in shadow from the rocky ridge to the northwest about half an hour before the sun set on the far western horizon. Flights always began after sunset. Earliest emergence time of main flight was 20:28 on 16 June and the latest was 20:58 on 30 June. Six of ten flights observed from 15 to 30 June began between 20:50 and 20:57; three began between 20:44 and 20:47. Individuals appeared in the upper pit three to eight minutes before the main flight began. Flights lasted from 36 to 73 minutes. Bats began returning to the pit between 21:25 and 22:10 and continued to return throughout the night in scattered groups. Bats were generally back into the roost by 05:45. The sun rose over the valley about 06:00 but did not reach the pit until about 08:00. Weather seemed to exert an influence on the emergence behavior of the colony. The bats would emerge if it was raining lightly. However, on one night of severe storms they failed to emerge in the characteristic pattern. Some individuals could have emerged after the storms ended. The last frost of the spring was the first night I observed a definite, early out-flight. Qualitatively, it seemed the area covered by roosting bats had doubled between 13 and 28 June. An estimation of the:population from direct observation of the roost was not possible, because I could not accurately count the number of bats per unit area or estimate the total area of the roost. At the closest observation point, I was still about 30 m from the roost. Part of the colony appeared to be obscured by large, overhanging rocks. The photographic estimation of the population size for June was as follows: 9,270 (19 June); 15,542 (24 June); and 28,265 (30 June). Several characteristics of the out-flight need to be considered when analyzing the photographic estimation of the population: bats flew out holes other than the exit in the main pit; bats did not always fly over the arch in front of the camera when they did exit from the main pit; the flash and flashlight may have affected the flow of bats; it was difficult to accurately time bats across the frame after dark; some bats were flying in as others were flying out; not all bats necessarily leave the roost nightly to feed; weather affected the concentration and length of the main flight. For these reasons, the population estimates are probably conservative. �144 Table l.--Total capture of bats at the Orient Mine: 17 May - 30 June 1982. Mean Weight (g) (range) Mean Forearm(mm) (range) Species n Tadarida brasiliensis 63 Male Adult 11. 2 (9 - 15) 43.5 (42 - 43) 1 Female Adult 15 44 Eptesicus fuscus 1 Male Adult 14 51 Lasiurus cine reus 1 Male Adult 27 Plecotus townsendii 1 Ma Le Adult 9 Sex Age �145 I set nets at five different locations in the valley but failed to capture any 1'.. brasiliensis in the nets during June. I did catch four other species of bats at two of these locations: Lasiurus cinereus, Plecotus townsendii, Motis evotis, and M. lucifugus. In June, I collected from T. brasiliensis two stomach, six guano, and eight tissue samples at the mine. These samples await future analysis. LITERATURE CITED Armstrong, D. M. 1972. Distribution of mammals Mus. Nat. Rist., Univ. Kansas, 3: 1-415. in Colorado. Monogr. Cockrum, E. L. 1969. Migration in the guano bat, Tadarida brasiliensis. Misc. Publ. Mus. Nat. Rist., Univ. Kansas, 51: 303-336. Rumphrey, S. T. 1971. Photographic estimation of population size of the Mexican free-tailed bat, Tadarida brasiliensis. Amer. Midland Nat., 86: 220-223. Meacham, J. W. Res. News, 1974. A Colorado 15: 8-9. colony of Tadarida brasiliensis. Ramaley, F. 1942. Vegetation of the San Luis Valley in southern Colorado. Univ. Colorado Studies, Sere D, 1: 231-277. Prepared by: pE@ Svoboda Wildlife Tech. 1-B Bat ��147 JOB PROGRESS State of COLORADO W-124-R Project No. Work Plan·No: Raptor Investigations II Job Title Job No. Osprey Nesting Period Covered: Personnel: REPORT January 1 ------------------------ Studies 1, 1980 through June 30, 1982 Michael Berman, Elizabeth Bowden, Gerald Claassen, Gerald Craig, James Enote, Marta McWhorter, Gail Rosendale, Wayne Russell, Brian Simmons, Herb Stolzenberg, and John Wagner, Colorado Division of Wildlife. ABSTRACT Colorado osprey productivity averaged 1.00 and 0.30 young per active nest for 1981 and 1982 respectively, with 5 of 9 active sites being successful in 1981 while 2 of 10 pairs were successful in 1982. Shell fragments of 24 eggs were obtained and measured for thickness and 12 intact nonviable eggs were collected for pesticide residue analysis. Over the two breeding seasons, 9 active nests were kept under observation to ascertain influence of human activity upon nest success. This Job Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the Director. ��149 OSPREY NESTING INVESTIGATIONS Gerald R. Craig P.N. OBJECTIVES The objectives of this study are: (1) locate previously unknown nesting ospreys and monitor productivity of all sites, (2) determine cause of reproductive failure and develop methods to restore normal reproduction, (3) determine nesting habitat requirements and implement protective measures to assure continued occupancy, and (4) compile data and prepare annual and final reports. SEGMENT OBJECTIVES lao Locate and map previously Colorado. unknown osprey nesting lb. All known nest sites will be visited reproductive success. 2a. Observe nesting pairs from concealment to determine any behavioral abnormalities, weather conditions, predation, or human disturbance which might impact reproduction. Sites in remote localities will be compared with those adjacent to public activities to determine responses to human visitation. 2b. All unhatched eggs and shell fragments encountered during nest visitations described in lb. will be collected and submitted for pesticide analysis. Shell fragments will be measured and compared with pre DDT era eggs for possible thinning. 3. Habitat features such as key hunting areas, topography, vegetative type, climate, and geology will be recorded for each nest site in an attempt to establish habitat requirements. 4. Analyze annually sites throughout to establish data and prepare a report of the findings. METHODS AND MATERIALS Previously recorded osprey nest sites were visited in late April and early May to determine occupancy and onset of incubation. Deep snow cover prevented visitation to most of Jackson County nests until late May. Upon initiation of incubation, nests were climbed and the number of eggs counted. Sites were revisited at the estimated time of hatch and all addled or broken eggs were collected for shell thickness measurement and pesticide analysis. If a site appeared to have failed any time during incubation, the nest was visited to determine cause of failure and collect eggshell fragments. Efforts also were made to visit �150 all active nests in the aftermath of severe winds to assess possible damage. Nestling osprey were banded at an appropriate age with U.S. Fish and Wildlife Service bands and the nests were kept under intermittent observation to determine fledging success. Whole, unhatched eggs were submitted to Raltech Scientific Services, Inc. for chlorinated hydrocarbon insecticide analysis in accordance with laboratory procedures standardized by the U.S. Fish and Wildlife Service Research Center at Patuxent, Maryland. Eggshell thickness measurements were obtained optically using a microscope equipped with a stage micrometer. Where possible, three measurements were taken around the waist of the egg and the results were averaged. In 1981 and 1982, 4 active nests were observed from blinds to monitor osprey responses to human activity in the vicinity of nests. Two remote nests were chosen to represent low disturbance sites and 2 nests situated adjacent to recreational use areas characterized high disturbance sites. Blinds were concealed to permit observation of the nests and surroundings without disturbance to the ospreys. Observation blocks of 4 and 6 hours were arranged to monitor each site in mornings and afternoons as well as weekends and weekdays. In addition to recording duration and intensity of osprey responses to disturbance, observers also monitored adult attentiveness, duration of egg exposure to the elements, frequency prey was brought to the nest, and responses to climatic conditions. RESULTS AND DISCUSSION Table 1 summarizes osprey nest occupancy and reproduction for Colorado while Table 2 provides information on a site by site basis. Productivity for 1981 approached that experienced by normally reproducing populations with 9 sites producing 9 young for an average of 1.00 young per active nest. Although the number of active nests increased by 1 in 1982, production declined with only 2 sites fledging a total of 3 young (0.30 young per active nest). All of the nesting failures in 1981 and 1982 occurred during incubation with 6 sites exhibiting incubation prolonged beyond normal hatch while the other 8 nests failed prior to hatch. One nest each failed in 1981 and 1982 when high winds destroyed the nests and contents. In 1981, 13 fractured or addled eggs were collected for shell thickness measurement. Contents of 7 intact eggs were submitted to Raltech Scientific Services, Inc. for chlorinated hydrocarbon insecticide analysis. In 1981, an additional 11 egg samples were collected and measured for shell thickness. Five intact eggs were retained for pesticide analysis. A total of 550 hours of blind observation was obtained from early May through July of 1981. Nests LA-2 and JA-4 were situated in remote locations adjacent to beaver ponds and were selected to represent low disturbance sites. Higher disturbance sites were represented by GR-4 �Table 1. Colorado 1973-1982 osprey reproduction, TOTAL 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 Total Nests 5 8 16 16 17 17 15 16 16 17 143 Occupied 4 6 6 8 13 6 8 13 11· 12 87 4 5 6 8 13 6 8 10 9 10 79 0 3 0 2 2 3 2 1 5 2 20 Young Hatched 0 7 a 3 3 4 2 2 10 4 35 Young Fledged a 7 a 3 3 4 2 1 9 3 32 0.00 1.17 0.00 0.38 0.23 0.67 0,25 0,08 0.82 0.25 0.37 0.00 ·1.40 0.00 0 .. 38 0.23 0.67 0.25 0.10 1.00 0.30 0.41 0.00 2.33 0.00 1.50 1.50 1.33 1.00 1.00 1.80 1.50 1.60 Active Nests Nests Successful Young Nests per Occupied Young Per Active Young Site Site per Successful Pair ,_. ,_. \.J1 �152 Table 2. Site No JA-1 JA-2 JA-3 JA-4 JA-5 JA-6 JA-7 LA-1 LA-2 GR-1 GR-2 GR-3 GR-4 GR-5 GR-6 GR-7 GR-8 GR-9 GR-10 GR-ll GR-12 GR-13 GR-14 GR-15 GR-16 GR-17 GR-18 GR-19 GR-20 GR-21 GR-22 GR-23 Colorado osprey reproduction by site, 1973-1982. 1973 1974 1975 1976 1977 1978 ?/3/3 ?/?/O ?/?/O ?/?/O ?/?/2 ?/?/2 ?/?/O ?/?/O ?/?/1+ AB UO AB uo ?/?/1+ TD 2+/0/0 ?/?/O ?/?/O TD AB OC uo uo OC ?/?/O ?/?/O ?/?/2 uo uo ?/?/1 ?/O/O OC ?/O/O ART ART 1/0/0 ?/O/O OC ?/?/O ?/O/O ?/O/O UO ART uo 1/0/0 ?/O/O 2/0/0 2/0/0 ART uo ?/O/O UO ?/?/2 OC ?/O/O ?/?/O 1+/0/0 OC ?/O/O TD ?/O/O UO 1979 uo uo uo OC ?/?/1 3/0/0 3/0/0 ?/O/O TD AB 3/0/0 UO AB AB 3/0/0 UO ?/?/2 3/0/0 3/1/1 oc ?/?/O TD 1980 UO AB 0/0/0 ?/2/1 AB uo ?/?/O TD ?/O/O 2+/0/0 2+/0/0 TD OC AB AB 3/0/0 uo UO 3/0/0 AB uo AB AB 2+/0/0 ?/O/O 0/0/0 2+/0/0 1981 uo 1982 uo UO OC TD 3/0/0 AB AB AB AB 2/0/0 TD 5/0/0 AB 4/0/0 AB AB 3/0/0 4/0/0 AB 3/1/1 AB 3/0/0 TD ?/2/1 TD 3/2/1 AB 2/0/0 2/0/0 3/0/0 AB TD TD AB 2/1/1 TEl 3/2/2 AB 3/2/2 AB AB uo 3/0/0 AB 3/3/3 AB uo uo UO 0/0/0 UO Status Codes OC = occupied breeding territory UO = unoccupied breeding territory AB = abandoned ART = artificial nest constructed TD = tree or nest blown down 3/2/1 Production Codes 3 eggs, 2 young hatched, 1 young fledged TD TD UO OC UO ?/O/O 2/0/0 �153 and GR-I0 since they were subjected to high recreational use of Shadow Mountain and Granby reservoirs. Osprey responses were recorded for intrusion by hikers, fishermen, boaters, horseback rides, trail bikes, automobiles, and aircraft. Due to personnel and travel restrictions in 1982, GR-2 and GR-12 in Grand County were substituted for LA-2 and JA-4 in North Park. Due to early failure of GR-2, GR-10 was substituted. Since the critical disturbance occurred during incubation, observation was terminated after hatch rather than continue through mid-July and August. Unfortunately, GR-2, GR-4, GR-8 and GR-12 all failed during incubation and observation was terminated prematurely. Subsequently, only 287 hours of observation were accrued in 1982. Finally, an experiment was conducted to examine the potential of egg chilling or overheating during prolonged disturbance. Extra large hen eggs were marked similarly to osprey eggs and a thermister was inserted into the core of the egg. Each egg was heated to 36.6-37.7oC, then placed in an artificial osprey nest. Egg, nest cup, and atmospheric (shaded) temperatures were monitored under conditions simulating coolsunny, cool-overcast, warm-sunny, and warm-overcast days. Temperatures were recorded at 5 minute intervals for 90 minutes, or until the temperatures stabilized. The results will aid in delineating critical temperatures and the duration osprey can remain off the eggs before egg viability is compromised. Prepared by C.R Gerald R. Craig Wildlife Researche C � https://cpw.cvlcollections.org/files/original/e046560ac8155fb6bc7c5484e900505b.pdf 2d1dcea64ab6c424f60762f5c7a19d9a PDF Text Text JOB FINAL REPORT State of Colorado ----~~~~~------------- Project No. Game Bird Survey W-37-R-35 Work Plan No. Job No. 22 Evaluation of Nesting Cover Preferences Job Title: of Pheasants in Relation to Wheat Farming Methods Period Covered: Personne 1: 1 April 1981 through 30 June 1982 David C. Bowden, Clait E. Braun, Martin Staab, and Warren D. Snyder, Colorado Division of Wildlife. ABSTRACT Ring-necked pheasant (Phasianus colchicus) hens were fitted with solar-powered radio transmitters and monitored during the nesting seasons of 1979-81 to investigate nest habitat preferences in a northeastern Colorado region dominated by dryland wheat farming. Survival status was known for 113 of 131 radio-marked hens. Of these, 74 were monitored through mid-summer and 53 (71.6%) successfully nested. Green wheat, its residual stubble, or summer fallow farmed in a biennial rotation occupied approximately 85% of the total area and provided a greater proportion of the nesting cover available in spring. The proportion of land classed as nesting cover declined rapidly to 48-56% with spring tillage of wheat stubble. Weather, primarily precipitation, through its influence on onset of wheat growth and the resulting quality of green wheat and stubble was the major factor determining selection of nest habitats. Because of the biennial wheat farming rotation, the amount of precipitation received in any 1 year partially influenced nest placement for 2 subsequent years. Drought conditions in 1978 delayed wheat growth in 1979 and most hens (p > 0.05) began nesting in wheat stubble. The 1978 drought resulted in poor stubble conditions in 1980. With better green wheat in 1980 and delayed onset of nesting, nearly all nests (p < 0.05) were placed in wheat. Wheat made excellent early growth in 1981 and stubble remaining from the 1980 harvest was of good quality. Approximately one-third of the hens nested in stubble and most remaining hens nested in green wheat in 1981. Nests in stubble, if not predated, were nearly all destroyed by stubble tillage. Timing of stubble tillage was important to pheasant production in years when hens nested in stubble. Stubble tillage in late April or early May, before incubation commenced, promoted early renesting in green wheat where most nests successfully terminated before wheat harvest. Late Mayor early June stubble tillage destroyed nearly all clutches in residual stubble. Hens moving to green wheat in late Mayor early June after nest loss to tillage or predation were placed in jeopardy of subsequent nest loss to wheat harvest. Little nest loss because of wheat harvest occurred in 1979 because harvest was delayed by cool, wet weather. Pheasant production was curtailed by nearly one-fourth in 1981 when a majority of stubble tillage was late due to wet fields. Disking or plowing of stubble in April, when possible, is �2 recommended to increase pheasant production by forcing hens into wheat earlier increasing chances for nest completion ahead of harvest. An undercutter, containing mulch treader or harrow attachments to flatten residual stubble, should be used in early spring to replace disks or plows where soil erosion is a problem. A 2nd option is to replace early spring tillage with herbicide weed control so that the stubble can remain standing until after the primary nesting season. Mixed native perennial grasses, either grazed or ungrazed, were less attractive for nesting than annual weeds and weedy residual. Weeds were used for nest placement at a greater than expected rate (p < 0.05) but weedy sites comprised only a small proportion of the area-available. Hens with broods moved from wheat to annual weeds, usually in roadsides, and then often shifted to irrigated -------G-G-~R-aRd-G{:_he_r___F_ew__c_r_ep5,___e-~ew-e-r-op__weed-i-n-~e-r-s_pers-i-Ofl-. -P-r-ed-at-i-on-of~-------hens was greatest in late winter and early spring prior to spring dispersal from wintering sites. Predation was greater than expected (p < 0.05) on or near the Sand Draw Property, with its extensive tree plantings, than elsewhere. Nest predation also occurred at higher than expected rates (p < 0.05) on or near the Sand Draw Property. Older,; radio-marked hens were heavier and lived longer than subadults. Pheasant pr'oduction in the eastern Colorado Tablelands was closely related to timing of wheat culture. Nesting chronology was closely related to wheat growth. Findings of the study were reviewed with respect to hen harvest, development of predictive models, and census. �3 EVALUATION OF NESTING COVER PREFERENCES OF PHEASANTS IN RELATION TO WHEAT FARMING METHODS Warren D. Snyder Pheasants, as products of farmland, are subject to the impacts of weather and farming throughout their range. Wheat farming on a biennial rotation with summer fallow dominates agriculture in eastern Colorado where precipitation is below optimum for both pheasants and farming. Pheasants are severely Impacted at times by summer faTlow operations, yet summer fallow provides the vigorous wheat growth pheasants need and allows wheat stubble to stand over-winter providing essential fall-winter survival and feeding cover. Thus, pheasants are dependent on wheat for their existence throughout much of eastern Colorado and densities tend to reflect or indicate the quality and productivity of the farmland. This report comprehensively summarizes findings from a 3-year (1979-81) study to provide insight as to the climatic/vegetation/farming interactions and resultant direct and indirect impacts of weather and farming on pheasants. P.N. OBJECTIVES 1. Document-the relative importance of wheat stubble, green wheat, and other vegetative cover for (a) pheasant nest site selection, and (b) successful production of young in the wheatlands of northeast Colorado. Other variables pertinent to understanding the basic ecology of the nesting pheasant in the Tablelands include determination of primary limiting factors to reproduction and brood survival. 2. Upon documentation that pheasants use wheat stubble extensively for initial spring nesting and that adequate sample sizes can be obtained in the chemical fallow treated fields and their controls, a 2nd objective will be to determine if minimum tillage fallow with herbicides (Greb 1977) will increase pheasant nesting success when compared to conventional summer fallow methods. METHODS AND MATERIALS 1. Literature review, interviews, and coordination meetings. Literature on farming and chemical fallow techniques, and telemetry procedures and equipment were reviewed. Meetings and contacts were conducted to select the study site, develop a contractual agreement with the property lessee, and to select monitoring equipment and develop monitoring procedures. �4 2. Selection of study area. Adequate numbers of pheasants, location, and access for monitoring, proper cover types, and opportunity for application of treatments were primary criteria used in selecting a study area. 3. Plot design and application of treatments. A 2,327-ha (9-section) study area was chosen to be cover-type mapped and used in monitoring hen pheasant use of different habitats in relation to ava ilab iIity. ______ Ap_p_r-oX-i-flla-tel-y_8-O-h<1-(_20-0----<3-e_r:e_s_4f--whea_t-I-an<l-On-t-Ae-D_i_v-i-s-i-0n 's---~------Sand Draw Property and on the lessee's adjacent private property were selected for use in evaluation of minimum tillage fallow with herbicides. Only one-half of this acreage (40 ha) was used each year because of the biennial wheat/summer fallow farming rotation. Chemical treatment was applied to approximately 20 ha and 20 ha served as a control to be farmed by conventional tillage summer fallow methods each year. The lessee or his contracted applicator used ground spraying equipment to treat wheat stubble in late summer after wheat harvest with atrazine preemergent herbicide, applied at 1.125 kg/ha (lIb/acre). A contact herbicide was used in the mixture. Glyphosate "Roundup" applied at the rate of 2.5 liters/ha, or 2,4-D amine applied at the rate of 1.125 kg/ha could be used as weed conditions necessitated. The lessee was not allowed to till the treated stubble prior to late June of the subsequent year. The control, or untreated stubble, was farmed using a conventional summer fallow approach wherein an offset disc or mold-board plow was used in InItial stubble tillage in May, timed to coincide with conventional summer fallow activity in the surrounding region. Repetitious tillage was continued as needed to wheat planting in September. 4. Analysis a. Hen lighting trapped trapped and Evaluation. Trapping and monitoring procedures and equipment. pheasants were trapped in late February or March using night techniques described by Hoffman (1975). Additiona-l hens were in April as needed to replace known mortalities among initially hens. Captured hens were equipped with solar-powered backpack radio transmitters which weighed approximately 15-18 grams. Flat nylon elastic material was used in harnesses. Harnesses were similar to those described by Brander (1968), but were modified in 1980 and 1981 by passing nylon elastic loops directly from the transmitter under each wing .. �5 Monitoring procedures followed those described by Kuck (1968) and others. Three 9.1-m masts were erected on high points on and near the Sand Draw Property and were positioned slightly over 1 km apart. Telescoping masts, which extended approximately 5.5 m above ground, were attached to a frame on vehicles to monitor pheasants outside the range of the stationary antennas. Both stationary and portable masts were equipped with 2.4-m yagi high-gain antennas. Hand-held 3-element antennas were used in field monitoring when on foot. Two 24-channel receivers used in 1979 were replaced with receivers having a 150.000-151.999 Mhz range in 1980 and 1981. .~ Flushing counts were initially used to determine brood size. However, these were discontinued because it was difficult to obtain accurate counts, chick -----,ll'lClL.:taliLy__JJ]a¥-hav_e_ __j_n.c-~eased-,--an.d_s-h_i_f_t-s-of_Aen_S---t-O-new-l-oca-t-i-on-s-may-ha-v resulted. Ages of egg embryos were classified using a key developed by Sandfort (1957). Backdating of clutches was based on a laying rate of 1.3 days/egg (Buss et al. 1951) and an incubation period of 23 days. b. Environmental monitoring. The Robel et al. (1970) method as modified by L. Kirsh (unpubl. rep., U.S. Fish and Wildl. Serv., Jamestown, N.D., 1977) was used to obtain height-density indices of wheat stubble, green wheat, and other vegetation at specified times each spring. Wheat was measured at 10-day intervals in late April and May. Readings were made diagonal to the direction of drill rows in wheat and stubble covers. Height only measurements were also obtained for evaluation purposes. Stubble tillage progress was surveyed at weekly intervals each spring on an established route from west of Holyoke, northeast to the study area and back toward Holyoke. Between 80 and 100 fields were included in the sample. Progress of stubble tillage was also obtained for all fields within the study area. Dates of wheat harvest initiation and progression within the study area were recorded each year. Precipitation was recorded each year on the study area and supplemented during winter months, when necessary, with information from U.S. Dep. of Commerce weather stations at Holyoke and Julesburg. Grasshopper densities were obtained in selected cover types by use of ocular 0.09':'m2(1-ft2) samples obtained at 1 pace intervals whi le walking in midsummer. The technique is that used by Colorado State University Extension personnel (E. Anderson, pers. commun.). Nine samples were combined to project an index of density/O.8-m2 (l-yd2) sample in each cover: , ,type. DESCRIPTION OF AREA The study was conducted in the tablelands of southeast Sedgwick County in extreme northeast Colorado. Major study activities were within a 2,327-ha (9-section) study area, but extended into neighboring locations with spring dispersal of some hen pheasants. The center of the study area was the 84-ha (210-acre) Sand Draw Property where Division of Wildl ife personnel had conducted extensive tree and shrub planting in 1949. This area has extensive rows of wild plum (Prunus americana) and Siberian elm (Ulmus siberica). �6 Soils are of the Rago-Richland-Kuma associations characterized as deep, nearly level, and of loam texture (Brubacher and Moore 1969). Sand Draw, which has not flowed runoff water for several years, is the northern most tributary of the Republican River drainage in Colorado. The elevation approximates 1,143 m (3,750 ft) and average annual precipitation is about 45.7 cm (18 in) most of which is received during the 143 day average annual growing season. Dryland winter wheat and summer fallow, in biennial rotation, dominate but several deep irrigation wells, installed in recent years, provide water to row crops under center-pivot systems. Only 3 occupied farm residences are within the study area attesting to low human populations and large field -------an4-Fafm-s-i-z.es-I"-F-ev-al-a-n-t-thr-oughol1t-eas-t-e-rn-€-o-1-o-ra-do,-.---------------Ring-necked pheasants are the major hunted game species in the region. Mourning doves (Zenaida macroura), desert cottontails (Sylvilagus audobonii), and numerous nongame species used the Sand Draw and surrounding areas during the year. RESULTS AND DISCUSSION Land Use, Dispersal, and Seasonal Hen Use of Cover Types Land Use. -- Rapid development of deep-well irrigation from the Ogallala aquifer has converted extensive acreages along the eastern border of Colorado into row crops and alfalfa during the past 15 years. Corn dominates on irrigated acres with lesser amounts of sugar beets, pinto beans, alfalfa, and grain sorghums. Restricted water availabil ity, depletion of the aquifer, rising pumping costs, and low prices received for crops have slowed expansion of irrigation with a slight current trend toward irrigated winter wheat and grain sorghums, which require less water, and a shut down of some marginal wells. Winter wheat remains the dominant cash crop in eastern Colorado and will probably retain that status for the foreseeable future. Percentages of crop types within the 2,327-ha (9-section) telemetry study area were compared with information obtained for a 2,566.7-km2 (991-section) area type mapped in 1963-68 (Snyder 1970) (Table 1). The 1963-68 data were obtained before major irrigation development had begun. In 1979-81, wheat still dominated the landscape in the overall area although up to one-third or more of the land was irrigated in some tableland localities. �7 Table 1. Percentages of cover types within the 23.3-km2 (9-section) telemetry study area in 1980 and a 2,S66.7-km2 (991-section) northeastern Colorado Tableland area mapped in 1963-68. Cover type Telemetry study area Wheat and summer fallow Irrigated row crops Millets and sorghums Short and midgrass pasture 86.S 6.7 1.2 3.0 1-ot-a--I---crop-type '91--;-4 Roadsides Odd areas Occupied farmyards Tree and shrub plantings 0.3 1.4 0.2 0.7 1963-68 study areaa 76-83 1 - 1.S 4.4- 4.S 8 -13. S 96-9 0.6 O.S-l.S 0.4 0.2 aOata from Snyder (1970). Over 96% of the total area was farmland or pasture during both study~periods (Table 1). Odd areas, comprised of weedy draws, unfarmed low areas, old building sites, etc., were the najor nesting covers available other than farmland. Most roadsides were farmed to the shoulder offering little nesting cover. Extensive tree and shrub plantings within the Sand Draw Property biased comparison of woody plantings between the 2 study areas (Table 1). Private lands surrounding Sand Draw, like those elsewhere in the region, were nearly treeless except for farmstead windbreaks. The telemetry study area closely resembled land use elsewhere in extreme northeast Colorado. Edge types within the 2,327-ha study area were classified in spring 1979. The average section contained approximately 12.2 km of edge although the extensive strips of woody plantings and wheat on the Sand Draw Property skewed the amount of edge upward compared to quantities of edge on private lands. Approximately 12% of the edge was classed as moderate to high value to pheasants, 49% was fair to moderate, 32% was poor to fair, and 7% was zero to poor. Both quality and quantity of edge types changed rapidly with season progression, tillage of residual covers, and with growth of wheat, row crops, and wild annuals. Spring Dispersal of Hens. -- Approximately 2S:-SO% of the hens radio marked in late winter on or near the Sand Draw Property made spring dispersal movements of at least 1.2 km (3/4 mi) or longer during the 3 years of study. Most movements in 1979 and 1981 occurred in early to mid-April. In 1980, the majority were in late April (Table 2). Dispersal, like nesting activity (Table 3) was delayed in 1980 by a major late March snowstorm. �8 Table 2. Timing of spring dispersal of radio-marked hen pheasants northeastern Colorado 1979-81.a Years Time of movement 1979 Prior to 10 Apr 10-16 Apr 17-23 Apr 24-30 Apr 2 --------T-0-ca-l-s 3 5c-----------' Percent of marked sample 27.8 1980 1981 2 1 1 9 4 3 2 8 7 3 9 1-3 9 27 50.0 29.0 Totals aAt least 1.2 km (3/4 mi) or greater distances. Table 3. Cover type of first knowna April and May pheasant nesting attempts, northeastern Colorado, 1979-81. Year Cover type 1979 Wheat stubble Green wheat 1980 Green wheat Undisturbed cover 1981 Wheat stubble Green wheat Undisturbed cover 15-30 Apr 1-15 May 16-31 May 3 7 2 8 10 1 7 3 3 4 3 2 2 1 7. 1 aFirst confirmed nest/hen. Contact was lost with some radio-marked hens because of spring dispersal even though extensive searches were made for hens each year. Emigrations of 3.2-4.8 km (2-3 mi) were common and the longest known movement approximated 10 km. These distances greatly exceeded those reported by Carter (1971) for hens in eastern South Dakota. Gates and Hale (1974) found that adult hens dispersed to specific, previously used breeding areas whereas movements of subadults were less predictable. They noted that spring movements were usually shorter than those in fall. Movements of hens were frequent through the spring and summer and were usually associated with nest predation. Hens frequently shifted 1.6-3.2 km for 1 or 2 days, but most hens returned to renest within 100-400 m of the previous nest site. Occasionally hens did not return to the area of the original nest. �9 Prelaying Use of Cover Types. -- Wheat stubble received primary use by hens in early to mid-April during dispersal, harem formation, and prelaying. Woody covers ranked 2nd in use with lesser use of residual weed and grass types. Patterns of cover use were similar each year. Location of radio-marked hens was primarily conducted between 0800 and 1700 hours, a period when most feeding activity had ceased. Movement into and use of green wheat varied from year to year in relation to its growth. No significant use of wheat occurred until early May 1979 when it was only 16.3 cm (6.4 in) tall by 1 May (Fig. 1). Hens began using green wheat about 24-28 April 1980 because of earlier growth and poor stubble quality. Still earlier use (20-24 April) occurred in 1981 under excel-------I-errt---§-Few-t:-h-eend-i-t-i-ens-(-F-i-g-. -l-)-.-Howe-ve-a,-some-hen-s-rema+n-e1:i-i-n-whe-a-T--------stubble and other covers through early spring each year. Nest Placement in Relation to Cover Availability. -- Nesting hens were not randomly distributed within the study area and some hens dispersed beyond the area boundaries prior to nesting. The tract size a hen would use when selecting a nest site was unknown so a 145.4-ha (360 acre) tract was arbitrarily selected to use in examination of cover types surrounding each nest site. This tract size provided 0.6 km (0.37 mi) or more of land in any direction from the nest site. Vegetation types were classified surrounding 99 nests during the 3 years andj divided into early spring (55 nests primarily initiated prior to stubble tillage); late spring (39 nests initiated prior to wheat harvest); and post-harvest (5 nests). Approximately 87% of the classified area contained some form of nesting cover in early spring 1979 compared to 65% in 1980 (some stubble was tilled prior to initiation of some nests), and 90% in 1981. The percent of land classed as nesting cover ranged from 48 to 56% after completion of most stubble tillage during the 3-year interval. Wheat stubble wasl classified as nesting cover if it had been undercut no more than once with a sweep plow, which allowed a majority of the stubble tQ remain standing. Stubble that was sweep-tilled more than once, or tilled once by other implements was classified as summer fallow (primarily bare ground) and was not considered suitable for pheasant nesting. However in 1981, 2 hens whose nests in stubble were destroyed by tillage, renested in disced fallow near their previous nest sites. These nests were deleted from subsequent analyses. Occasional nesting in disced fields has been reported by farmers in the past. Hens, in early spring 1979, placed most nests in wheat stubble (10 of 13). However, the proportion was not greater than expected (p > 0.05) (Table 4). In contrast, hens selected green wheat at a greater than expected rate (13 of 14) (p < 0.05) and avoided poor quality stubble in spring 1980. Green wheat was used at a higher than expected rate for nesting in early spring 1981 but the rate was not significant based on the small sample (~> 0.05). Combined early spring 1980-81 data showed wheat received greater nesting use than would be expected on a proportional availability basis (!: < 0.05) (Table 4). �10 100 -------------------------------I---------------------------~--\~e~~~'---------------------------- ,,' 25.9 em 1 MAY 1980 ~-," 80 w o Z ,w a: :::> o o / -,9) •• .....• .• 22 APRIL 1981 40 •••• o • •• .... •• 8·21 •• ••• . 22·5 APR •• •• • • •• •• •• . • •• • • . • 20 ...•.. •• •• • / a: w a, Fig. 1. " 32.6 em •... -,," " ~~ 60 o z w o / / -' ••• ~ 15.4 em WHEAT HEIGHT 1 MAY 1979 6·18 MAY 19·2 16·30 JUN Occurrence of radio-marked ring-necked pheasant hens in green wheat in relation to growth and time among years, northeastern Colorado, 1979-81. �11 Table 4. Pheasant nest placement in relation to cover availabil ity during nesting periods within the Sand Draw study area, northeastern Colorado, 1979-81. Nests Year and period Cover type 1979 Wheat stubble Green wheat a Undisturbed Early spring ________________ 1980 1981 .Suh.to.ta. _,_] Early spring Early spring 197981 197981 Early spr ing Early sp ring Late spring to summer Early spring to summer 10 3 0 6.3 5.7 1.0 o Subtotal 14 Wheat stubble Green wheat Undisturbed 14 1 8 6 8 27 7 Subtotal 44 Combined wheat and stubble Grass 48 Subtotal 49 Wheat & stubble Grass Weeds & roadside Grass Weeds and roadside Subtotal 3.9 8.8 1.4 13.4 11.5 3. 1 28 Wheat stubble Green wheat Undisturbed 1 34 2 1 P 2.19 1.30 0.98 ---'lI-.Jq_®--2-df-->--O-._O_;;, _ -I-.J- 13 Subtotal 197981 Exp. Wheat stubble Green wheat Undisturbed Subtota I 198081 Obs. 18.8 20.5 4.7 45 4 32.5 2.9 0.9 3.87 2.03 0.10 6.00 @ 2 df < 0.05 2.20 0.56 2.71 5.47 @ 2 df > 0.05 6.16 2.03 1.11 9.30 @ 2 df < 0.05 0.20 2.25 2.45 @ 1 df > 0.05 > 0.05 < 0.05 0.02 0.26 0.02 0.30 @ 3 df 37 3 8.0 3.09 7 10 2.0 12.06 aUndisturbed covers included perennial annual weeds, and roadside. 15.15 @ 1 df grazed and ungrazed grass, �12 Green and ripe wheat composed an average of 88% of the available nestinq cover for the 3 years in late spring. Nest placement was approximately in proportion to availability of different cover types with 34 of 37 nests placed in wheat (P > 0.05) (Table 4). The small sample of hens (5) nesting after wheat harvest exclusively used wheat stubble. ______ Several small tracts of mixed mid- and shortgrass were within the study area and the majority were ungrazed. These areas represented less than 8% of the total nesting area but received no nesting use by radio-marked hens during the first 2 years of;the study. Weedy cover, primarily on the Sand Draw Property, represented less than 2% and roadsides represented less than 0.25% of' the available nesting cover. Proportional nesting in grass, annual weed, and --I-Ua.ds_j...de-cov.er--S-wa-S-compa-r-e-cl-ga-s_e-cl--0n-pee-I-ed-3~y-eaF-da-t-il-. -Samp-l-e-s-i-ze·s-we·Fp-----small (Table 4) but results indicated pooled annual weeds and roadsides received greater than expected use (p < 0.05), whereas less than expected use of grass cover was evident. Some caution should be used when interpreting P values associated with X2 due to small sample sizes. Hens also tended to nest in wild annuals-where present within and along the edges of wheat fields. Quantities of roadsides and nest samples were too small to reveal reliable nest density comparisons. However, information presented by Snyder (1974) indicated hens nested in both untreated and revegetated roadsides at much higher rates than would be expected in adjoining wheatfields. Hen - Brood Cover Use. -- Green wheat, the cover type in which most nests hatched, was the major cover used by hens with broods for the first 1-2 weeks after hatching (Fig. 2). However, a marked shift to edges of wheat fields and to weed and weed-grass dominated roadsides and other unused sites was noted. Weedy cover became the major cover type used by 3-week and older broods (Fig. 2). Increased use of cornfields, sorghum, and shrubs and trees was also evident for hens with older broods especially by mid- to late August. Hens with broods were often forced from wheat fields and their edges by wheat harvest. Hens with broods on or near the Sand Draw property frequently moved to the forb-shrub-sorghum-alfalfa covers found there. Hens 0.8 km or more away from the property usually moved to roadsides and then toward irrigated row crops or weed dominated sites. A tendency to use green vegetation and to remain along the edge of large irrigated fields was noted. Free drinking water was often associated with these sites but an attraction specifically to water could not be ascertained. Grasshopper densities were consistently highest in green weed and grass dominated sites (Table 5). This food source, essential to chicks, combined with the shade and protection p rovided by sunflowers (Hel ianthus spp.), kochia (Kochia scoparia), and other forbs was a major factor in their high use by young broods. Warner (1979) considered good brood foraging habitat to be potentially critical to chick survival in Illinois with decreased grass and forb sites and increased row crops. �13 w o z w a: ::::> o o o I- Z W o a: w a, BROOD AGE F occurrence 0 f ring-nee k ed pheasant hens ·lg. 2. Percent d 1979-81. by cover types, northeastern Colora 0, WI ..th Young broods �14 Table 5. Grasshopper densities among vegetation types and years on and near the Sand Draw Property, northeastern Colorado, 1979-81. Vegetation type Sunflowers, Dock (annual forbs) . Roadside (Cheatgrass brome & sunflowers) Perennial mixed grasses Alfalfa (0Id-79, new-81) Uncut ripe wheat W-i=lea-t-s-t-l;Ib-b-l Irrigated corn Irrigated pinto beans Mill et Average density/O.8m2 19S0 1979 35.5 39.3 14.5 14.4 3.0 C.3 Trace 14.5-31.0 4.2-11.8 11.5-14.2 (1 yd2) 1981 19.0-20.0 12.4 41.4 1.0 1--;-1 Trace 0.4 0.5 Hen and brood movements to irrigated row crops often subjected them to aerially applied insecticides and fungicides in mid- to late summer. Treatments varied among fields and years. Considerable spraying of corn occurred in 1980 compared to other years. Fungicides were used on sugar beets and pinto beans. This pattern of hen and brood movement to pesticide treated irrigated fields occurs throughout eastern Colorado and undoubtedly elsewhere. Direct mortality of radio-marked hens was not detected and the fate of broods could not be determined. Their tendency to remain along field edges potentially reduced impacts of spraying and possibly permitted them to shift to other covers during spraying operations. Fall - Winter Cover Use. -- Several hens, inaccessible for capture in large cornfields, retained their transmitters until after the October corn harvest in 1980. These hens continued to reside in corn throughout the interval, often feeding in adjacent covers, but moving to the corn when pursued. Transmitters were left on surviving hens through the fall - winter interval of 1981-82 and removed in late February. Cover type occurrence of these hens was compiled by bimonthly intervals (Fig. 3). These data documented the high use of cornfields in early fall. In most years, corn was picked and the majority of the corn fields were disked and plowed by late October or early November. However, wet corn and low corn prices delayed picking in 1981 and much of the crop was still standing at the start of pheasant season on 7 November 1981. Many fields were picked in mid- to late November, but some corn protected the majority of monitored hens into early December 1981. These fields offered a sanctuary, inaccessible to most hunters, and rooster harvest was reduced. �15 100 r GRASS ---------------------_-_-_-_-_-_-_-_-_-_- --------~~-~------------------------------------------ ------------------ 80 w o z w 60 a: :::J o o o I- Z W o 40 a: w 0.. CORN s SORGHUM 20 o SEP-OCT Fi g. 3. NOV-DEC JAN-FEB Percent occurrence of radio-marked ring-necked pheasant hens among cover types through fall and early winter, northeastern Colorado, 1981-82. �16 Some hens remained in wheat stubble/weed/shrub associations throughout the fall, but most used corn when available (Fig. 3). Wheat stubble was above average in height-density rating (Fig. 4) furnishing vast areas of protective escape cover through late fall and winter of 1981-82. Only light 2.5-5.1 cm (1-2 in) snows occurred through the interval. Hens made no major shifts due to hunter harassment and made no pronounced movements to the Sand Draw property through the fall and winter of 1981-82. Lack of significant snow was considered a factor in this stability. Excellent wheat stubble and corn cover conditions were also important factors. Weather and Wheat Growth Influence on Nesting The biennial wheat - summer fallow farming cycle must be understood when examining precipitation-wheat farming-nest placement relationships. Winter wheat is harvested in July. Most farmers in northeastern Colorado usually do not disturb the stubble until the following spring when summer fallowing is initiated prior to major weed growth. Summer fallowing usually involves spring tillage of wheat stubble followed by repetitious shallow cultivation, as needed, to prevent or remove weeds and prevent wind erosion. Seed is planted in September allowing new plants to establish root systems and enough growth to partially protect the bare soil over-winter. Wheat grows vigorously as the soi I warms from Apri 1 through early June, maturing rapidly for July harvest. Precipitation can be accumulated in the soil any time during the 2-year harvest-to-harvest cycle, but most is received and accumulated during spring and summer of the summer fallow year and in spring of the year of harvest. Precipitation received in the Great Plains Region fluctuates dramatically around average annual rates and thus has major impacts on dryland wheat and other vegetation. Precipitation received from 1977 through 1981 on or near the Sand Draw property was compared with long term monthly and annual means (Table 6). Precipitation was above average in 1977, severely deficient in 1978, near average in 1979 and 1980, and above average in 1981. Weather impacts on wheat growth were tested by regression analyses combining data from the mid 1960's (Snyder 1970) with 1979-81 data from precipitation recorded during a 16 month (Jan-Dec + Jan-Apr) summer fallow-wheat growth cycle! plotted against wheat height on 10 May. The regression coefficient (r = 0.70, N = 11) indicated there was a general dependence of wheat growth on precipitation. An improved relationship (r = 0.88) was attained when average April temperature departure from the long-term mean was combined with precipitation (Fig. 5). When May precipitation was added to the preceding 16 month total, the relationship with wheat growth improved (r = 0.81), but addition of April temperature departures from the long-term mean changed it only slightly (r = 0.83). More precise weather - wheat growth relationships might be-obtained by deleting light showers and snows, which seldom add soil moisture, by increasing emphasis on moisture received near the end of the interval, and by more precise weighting of spring temperatures. However, the above data provide strong evidence that wheat growth was primarily dependent on precipitation accumulated in the soil during the summer fallow year and spring of growth. Spring temperature had a lesser impact. �17 4 E "0 3 x w o 1981 WHEAT STUBBLE z >I- 1980 1982 NEW ALFALFA & CLOVER , --- NEW ALFALFA / / / 2 / / 00 Z 1979 W o / WHEAT STUBBLE .;. /,/" ===== WHEAT STUBBLE / W I ,/ ,/ '/" » ,> " ,/ " ----- ~--~---7 ,/ 1979------, _ __ 1980WHEATsTUBBLE'- ",,""" ~f>..\..~f>.. ""0 f>.."-' ,/ -I 1981 _~ -- "" ,,0"-' •.• 9'00 - ,,"'" "-- '" .."," - 1981 --- APR Fig. 4. Comparative 10 20 MAY height-density r-==-=- MIXED GRASS MIXED GRASS • 30 --- 1980 O~~------~~-------L-- L_ 20 - -- ,/ / , I (!J ,/ / _L__L_ 30 2 JUN indices (dotted lines indicate 95% CL) for wheat stubble, native perennial grasses, and old and new seeded alfalfa in spring among years in the Sand Draw study area, northeastern 1979-81. Colorado, �18 75 '. • 1981 60 z o E o I- I CJ .1966c .1963 45 • 1966e W I t:( W I ~ z w 30 w a: CJ 1. = 0.883 X = .1965e ... -7.85 + 1.04~ .1979 15 20 16 MONTH Fig. 5. The relationship bining precipitation 25 30 PRECIPITATION + APRIL TEMPERATURE DEPARTURE between wheat height on 10 May and an index com- (in.) over a 16 month (Jan-Apr) of mean April temperatures 35 interval plus departure from the long-term average, northeastern Colorado. Data include individual samples from 1963-64, 1979-81 and 2 samples for 1965-66 and 1968. �19 Table 6. Precipitation 1977-81.a in the Sand Draw vicinity, northeastern Precieitation 1975 1979 (cm) 1980 Month 1977 Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Totals cm in 0.99 0.25 7.52 10.52 11.05 ~1 6.27 9.53 3.00 0.10 0.38 0.36 1.14 1.09 0.18 1.88 7.06 3.48 4.14 3.18 0.28 1.27 1.09 1.91 55.28 26.70 45.50 40.64 55.88 21.76 10.51 17.91 16.00 22.00 1.63 0.23 5.44 3.68 6.20 9_~68 5.89 4.50 1.65 2.54 4.06 ir 2.29 1.85 8. 13 4.57 7.09 5--38 6.91 3.89 0.28 Tr 0.25 Tr 1981 Tr Tr 10.26 7.37 14.88 ~~-6B 5.31 10.01 0.76 1.91 1.65 0.05 Colorado, Long-term mean cm in 0.79 0.74 2.18 3.89 8.36 9-.-7~ 7. 16 4.78 4.17 2.87 1. 17 0.91 b 0.31 0.29 0.86 1.53 3.29 3-.g-3 2.82 1.88 1.64 1. 13 0.46 0.36 46.75 18.40 aprecipitation for 1977, 1978, and subsequent winter months was obta ined from U.S. Department Commerce records for Julesburg and Holyoke, Colorado. bLong-term mean precipitation for Holyoke, Colorado. Similarly, 16 month precipitation + April temperature departures partially influenced subsequent post-wheat harvest stubble height (~= 0.55, ~ > 0.05) because stubble height was in part a function of spring wheat growth (r = 0.60, N = 12, P < 0.05). Weather and wheat growth impacts on stubble height were-weak for several reasons. First, 10 May wheat height may not always be indicative of final wheat height. More importantly, late Juneearly July hailstorms- that lodge ripening heads are not uncommon in northeastern Colorado. Light to moderate hailstorms, where harvestable wheat is still standing, results in combine operators cutting lower to gather lodged heads. Moderate hail losses in 1980 lowered subsequent stubble height in contrast to that expected based on weather data (Fig. 6). No hail of significance was received during the 3 other years in the study area and a more direct weather - stubble height-density relationship was evident. Accumulated precipitation through the summer-fallow cycle and temperatures during the spring period of wheat growth influenced the quality of wheat, the qual ity of wheat stubble, and the placement of early spring nests by hen pheasants (Table 7). Precipitation accumulated over a 16 month (Jan 78 - Apr 79) interval was below the expected average for that period and consequently yielded sparse, short wheat in spring 1979 (Table 7). This, in turn, impacted the quality of stubble available to nesting hens in spring 1980 (Fig. 7). Thus moisture received in 1978 influenced nest placement in both 1979 and 1980 in addition to its impact on 1978 nesting. �20 2.5 .1981 >< w o z >I- Ci5 1.5 Z W o ._!. .1978 I CJ W I 1.0 W ~ • 1980 ( HAIL) [Q [Q ::> I- en t;: 0.5 W I ~ .1979 o ~ 15 16 MONTH ~ ~ 20 ~ 25 PRECIPITATION 30 + ~ 35 APRIL TEMPERATURE DEPARTURE Fig. 6. The relationship and an index combining between the wheat stubble height-density precipitation (in.) over a 16 month (Jan-Apr) interval plus departure of mean April temperatures average, northeastern Colorado, index 1979-81~ from the long-term �21 7 GREEN WHEAT 6 5 E "'C X W 0 Z >Ien 4 Z W 0 ~ I o 3 W I WHEAT STUBBLE ~ W I 3: 2 1979 ·1981 1980 APR Fig. 7. MAY JUN Wheat growth from 1979 through 1981 (vertical lines indicate 95% CL) compared with wheat growth during the 1960's and with subsequent wheat stubble height-density, northeastern Colorado, 1979-81. �22 Table 7. Comparison of precipitation and temperature impact on wheat growth, stubble quality, and placement of early spring pheasant nests, northeastern Colorado, 1978-81. Year 1978 1979 1980 198-1 a Deeart from Apr 16 month ppt. (cm) temp{F) + 5.23 -16.66 + 8.00 Y ... -9I, + 1.3 + 1.1 - 1.4 ,f-~-.6. Height-density Wheat index (dm) Stubble Nest placement Unknown~ v o 1.3 -1.1~.3 B0-Eh 11.1~..s 1982 Stubble Wheat 2.2 Unknown aprecipitation received through the previous summer fallow year plus the first 4 months of the listed year in relation to the average expected amount for that 16 month interval. April average temperature departure from long-term expected average for the month. bArrows denote the relationship between the following year's stubble quality. year's wheat growth and April temperature departure from the mean had less impact on wheat growth than did precipitation. Increased precipitation in 1979 and spring 1980 (Table 6) stimulated better wheat growth in 1980 resulting in better stubble going into spring 1981. Most radio-marked hens selected wheat stubble for nesting in late April and early May 1979 when better quality stubble and poor quality wheat were present (Tables 3, 7; Fig. 7). The opposite occurred in 1980 when green wheat made better growth and stubble was short and sparse. However, some bias in. the 1980 data occurred because a late March snowstorm deposited approximately 0.5 m of snow which remained on the ground into early April. This apparently stressed hens delaying onset of nesting (Table 3), during which time wheat made considerable growth. In addition, some stubble near the Sand Draw property was plowedprior to initiation of some nests .. Extremely early wheat growth and average stubble qual ity conditions were present in spring 1981 (Fig. 7). More hens selected green wheat than selected stubble for early nest placement but both covers were used (Table 3). Monitoring of hen activities and chance location of several abandoned nests in wheat provided evidence that some hens abandoned initial nests in early May 1981 during a period of wet conditions accompanied by relative cold temperatures. �23 Several nests were placed in residual weed and grass cover on and near the Sand Draw property in 1981, in contrast to almost no use of these covers during the 2 previous years. Lack of over-winter compacting snows increased the qual ity of residual weed and grass cover in 1981 but other factors including increased hen density and reduced harassment by predators could have influenced nest placement. The information presented (Table 7) indicates that hens selected for increased cover quality whe~ nesting with some exceptions. One hen moved from tall green wheat, where she had apparently lost 1 or more nests, to a short, sparse, grazed pasture where she nested successfully in an exposed situation. Another hen resided in dense, tall wheat while laying a clutch --------+.-i n-a.f-l-a.d.:i-a-ceA-t-s-umme.r-=-f-a-l--l-Gwe.d---£-i-e-1-cl-t-ha-t-ha.d-gee-n-G-i-s-k-e-d--Gn~!-.------------ Height-density indices (H-DI) of wheat, wheat stubble, ungrazed stands of mid- and shortgrasses, and several small areas of old and new seeded alfalfa varied with progression of spring and among years (Figs. 4, 7). The most striking feature was the rapid spring growth of green wheat (Fig. 7) beginning in mid- to late April. Wheat growth in 1979 was markedly below that of the 2 subsequent years and the 6-year mean obtained in the mid-1960's. In contrast, wheat growth in 1981 was much advanced. The data (Fig. 7) indicates, when compared with that in Table 3, that wheat attained enough growth to be used by nesting pheasants sometime in late April or early May in most years. H-D indices above 2.5-3.0 dm probably were of little additional value to nesting pheasants. The H-DI of wheat stubble did not change significantly prior to the time it was tilled in April or May. However, volunteer wheat could increasingly alter the cover value of stubble from mid-May into June if tillage was delayed as it was in 1981. Measurements of H-D were obtained in March and April except for the 1982 measurement obtained for comparative purposes in winter 1981. Stands of mixed grasses present on the Sand Draw property ranked below wheat in height-density value all 3 years and below stubble in 1979 and 1981 (Figs. 4, 7). Grass cover consistently failed to attain tall growth and the grasses frequently lodged under drifting snow, further reducing its nesting value. Small patches of alfalfa on the Sand Draw property were partially renovated by controlled burning in 1979 and old and new plantings were measured in 1980 and 1981 (Fig. 4). The new stands provided excellent spring nesting cover in 1981 although the H-DI lagged behind those for green wheat in early spring. Comparison of the H-DI among covers may not reflect their proportional value as nesting cover. Alfalfa or grass at a lower H-DI than wheat might still be preferred for nesting (Hartman and Sheffer 1971). The biennial summer fallow-wheat rotation, characteristic of the Sand Draw property and the Central High Plains; gives way to annual wheat or annual row crop farming in higher rainfall areas of the Great Plains at about 98-1000 longitude. Pheasant nesting conditions in the Western High Plains �24 Region are unique when compared to conditions further east. First, winter wheat and its residual stubble often dominate to the near exclusion of other nesting covers in major areas of eastern Colorado, southwest Nebraska, and western Kansas. Second, this High Plains region is semi-arid and below optimum for farming and for growth of perennial nesting covers in contrast to more eastern sites. Therefore, summer fallowing and biennial crop rotation of wheat permits it to make more vigorous and attractive growth for nesting than untreated native perennials. This better cover is frequently carried over to the stubble and is often available for initiation of early nesting the following spring. Numerous studies conducted in areas which are annually cropped have indi--------\c-<a:utue~d.J__tuba:Lwheat was-oot a preferred nas.t.Lnqcover (Tralltman 1960, Hantman and Sheffer 1971, Baxter and Wolfe 1973, and others). Nests placed in wheat were sometimes considered renests after hens were forced from other covers by haying or predation (Baxter and Wolfe 1973). However, a combination of relatively high nest success and extensive areas make wheat fields important pheasant production sites in nearly all locations where wheat is grown. Although winter wheat may not furnish high quality nesting cover in eastern Colorado, the available data indicates it dominates the scene to the near exclusion of other nesting covers. Wheat fields contained 63% of known nests established by radio-marked hens during this study and contained 75% of the successful nests (Tables 8, 9). Wheat stubble may be more attractive in early spring in some years but virtually no production comes from stubble because of tillage. Perennial vegetation, unless periodically renovated or reseeded, is less attractive than wheat for nesting and acreages are relatively small. Annual weeds and their residual apparently offer better nesting habitat but acreages are extremely limited and cannot be expected to increase. Thus, High Plains pheasants are clearly a product of wheat fields. Table 8. Colorado, Radio-marked hen pheasant nesting by cover typea, northeastern 1979-81. Cover type Green wheat Residual stubble New stubble Undisturbed cover Summer fa llow Totals 1981 1980 1979 N % N % N % 17 10 58.6 34.5 25 83.3 2 6.9 3 2 10.0 6.7 24 8 1 10 2 53.3 17.8 2.2 22.2 4.4 29 a'ncludes only verified nests. and either destroyed or abandoned. 30 45 All years N % 66 18 4 14 2 63.5 17.3 3.8 13.5 1.9 104 Other nests could have been initiated ~ �8 8 7 E "'C ~ Cl Z ~ en 7 MEAN DATE OF NEST INITIATION 6 ~ ," 5 t-!I C) [ij I 1 ,, 01 // 4 z w Cl ,r--~---y· I •...&, .,,~ , \9'00,'" 3 . I I -." ~/ 2 ~~ ~~ ",/ \919 ~~ ••• "til' ••• •••• ••• •• I I • I ,,..I .•. •• •• ~ 30 10 APR 20 30 MAY Fig. 8. The relationship pheasant nest Co lorado, initiation 1979-81. • • • •• .. •• • • •• •• •• •• IZ 5 ••• •• • •• •• •• •• 0). I\. • ....•• 0). 4 • •• •• •• •• •• •• • •• • • • •• • • •• of wheat and primary 25 MAY7 JUN 3 growth period 5 JUL to mean date of ring-necke of hatch, northeastern 6·19 JUL I ~ o I. Z 2 o 22 JUN· m G) • 8-21 JUN Z C/) -i C/) • •••••••••••••• o 20 , 1 . •• /' ,I •• ----.6 I 20 JUL2 AUG N V1 �26 70 .1981 60 ~ ~ ..0 50 Z 0 .1963 .1966 E o ._ 40 .1980 .1968 I C!:) W I .1964 30 ~ .1967 W I 3: .1965 20 .1979 1 21-30 11-20 JUN 1-10 JUL PRIMARY PERIOD OF HATCH Fig. 9. Relationship between wheat height on 10 May and major period of ring-necked pheasant nest hatching, 1963-68 and 1979-81, northeastern Colorado. �27 Table 9. Occurrence of successful radio-marked hen pheasant nests by cover type, northeastern Colorado, 1979-81. Cover type Wheat Wheat stubble (residual) Undisturbed vegetationa New stubble (late summer) Totals 14 1981 1980 1979 N % 87.5 N 14 % N % 87.5 11 55.0 5.0 35.0 5.0 1 2 12.5 7 2 16 16 12.5 1 20 All years N % 39 1 9 3 75.0 1.9 17.3 5.8 52 aUndisturbed vegetation included annual and perennial forbs and grasses and small acreages of alfalfa. Pheasant nesting chronology is tied to wheat growth further illustrating the species dependence on wheat. Delayed wheat growth yields later nesting and earl ier wheat growth results in earlier nesting as was shown for the 3 years of study (Fig. 8). A scatter diagram (Fig. 9), incorporating 9 years of information from northeastern Colorado, comparing wheat height on 10 May with the major hatching period further illustrates this relationship. Baxter and Wolfe (1973) found that nesting chronology was predominately influenced by precipitation (65%) and to a lesser extent by temperature (35%) in their southeastern Nebraska study areas. Wheat growth (Fig. 7), likewise is primarily governed by precipitation while temperature changes have a lesser impact. Therefore, the 2 data sets yield similar results. Openness of wheat fields allowed nesting hens and hens with broods to move about with relative ease. No clear tendency to nest near field edges was noted as has frequently been reported (Hartman and Sheffer 1971, and others). Average distances for nest placement from edges of wheat and stubble fields approximated 114-120 m. There was a tendency for hen pheasants to select turnrows, containing greater wheat density. In 2 instances, where adjacent fields were planted with 25.4 and 36.6 cm (10-14 in) row spacing, 3 monitored hens nested in the fields with the narrower row spacing each time. Other factors could have been involved in placement of these nests as hens frequently nested in fields containing the wider row spacing. �28 Mechanical and Predation Impacts on Pheasant Nesting Stubble Tillage. -- Wheat stubble, an essential component of eastern Colorado pheasant habitat, was extensively used for feeding, loafing, roosting, and escape from late summer through winter to early spring. Tall, dense stubble increases winter survival by reducing pheasant losses to predation and bl izzards, but tall, dense stubble also attracts increased nesting use the following spring. The current practice of tilling stubble during the primary nesting season is detrimental to pheasant production. There was a trend toward later stubble tillage in 1978-81 than in the 1960's (Fig. 10). This trend may be an artifact of precipitation because wet field conditions in early May 1980 and through most of May 1981 halted all stubble tillage. However, high fuel costs, the trend toward fewer summer fallow replications, and the fact that recently developed irrigation takes priority over summer fallowing in spring, support the data available (Fig. 10). The data for 1979 through 1981 (Fig. 11) clearly show delayed stubble tillage. Known nest destruction by tillage and predation in wheat stubble and days into incubation in spring 1979 and 1981 varied (Fig. 12). Several other nests in stubble were suspected both years, but could not be confirmed before they were assumed destroyed. Nests were not found in stubble in 1980 although 1 or 2 were suspected. The primary concern with late May - early June stubble tillageiis that nest destruction and nest predation carried beyond their immediate impacts; often causing total reproductive failure among some hens for the remainder of the summer. Hens forced from stubble in late Mayor early Junel when incubating, must physiologically prepare their reproductive systems for 1-2 weeks to resume egg laying and renesting (Table 10) (Seubert 1952, Dumke and Pils 1979). Green wheat is the primary nesting cover available after stubble tillage, consequently most hens move there to renest. Subsequently, they often are in late stages of incubation of their renest efforts when the nest is jeopardized by July wheat harvest. Thus, hens losing incubated nests in stubble face a high risk of also losing a 2nd incubated nest in wheat and the majority will not renest after a late nest loss. Table 10. The relationship between incubation progression prior to nest destruction and time required for renesting, northeastern Colorado, 1979-81. Days incubated 0-2 3-9 10+ Sample size 8 2 11 Average days to renest 3.9 7.0 11.0 �100 80 0 w _J _J i= I- Z 60 W o a: a. w 40 -Jrr/ / L / I N 1..0 20 17 10 24 APR 15 MAY Fig. 10. Progression Colorado, 1963-68, 1970-76, and 1978-81. 22 15 JUN of spring plowing of wheat stubble, northeastern �30 0 W .....I .....I rrz w 0 a:: 60 40 W a.. MAY APR Fig. 11. 1979-81 Progress (vertical of wheat bars stubble indicate JUN tillage 95% CL). in northeastern Colorado, �31 100 0 W _J _J DAYS INTO 60 INCUBATION lI- Z W o a: 40 W Q.. 20 1981 MAY APR Fig. 12. Loss of ring-necked northeastern Colorado, JUN pheasant nests to tillage in stubble fields, 1979 and 1981. �32 Wheat Harvest. -- The impact of wheat harvest on uncompleted nests in wheat fields varied (Table 11). The influence of harvest was sl ight in 1979 but major in 1981. Timing of harvest, 1 ike timing of stubble tillage or nest predation can be a critical factor. For example, wheat harvest was delayed by wet, cool weather in 1979 (Table 12) which permitted several hens to hatch clutches only a few days prior to harvest. If harvest had begun around 5-6 July, as it did in subsequent years (Table 12), up to one-third of the nests could have been lost. In contrast, if rains had delayed wheat harvest by 5 days in 1981, approximately one-half of the 10 nest failures might have hatched prior to harvest. Farmers cannot be asked to delay harvest of a crop that takes 2 years to produce when a hailstorm might destroy it at anytime. Therefore, timing of stubble tillage is the only negotiable variable and it often is partially governed by the weather. Table 11. Impact of wheat harvest on radio-marked success, northeastern Colorado, 1979-81. Variable hen pheasant nesting 1979 Number of nests cutover during harvest 1980 1981 2 Number of hens returning to incubate Percent of hens being monitored whose nests were terminated by harvest Number of hens subsequently successfully Above number as a percent of available following wheat harvest Number of hens not successfully harvest 5.6 renesting renesters 100 40 10 o 3 9 21 95 14.2 25.7 renesting after Projected reduction of nesting success due to wheat harvest disruptionc 1 hen nesting 28.6 2 Eggs lost to wheat harvest not subsequently replaced by renesting alncluded 23.8 o in a turn row that was not directly cut-over. bHen resumed incubation until abandoned on 8 September. CHypothetical - some nests if not disrupted, would have probably been lost to predation. �33 Table 12. Timing of wheat harvest in the vicinity of the Sand Draw property, northeastern Colorado, 1979-81. 1979 Date of July harvest initiation Major period of harvest (Ju1) Termination of minor fields (Ju1) Comparison with average 12 15-22 26-27 Late (1 week) 1980 1981 5 8-15 22 Normal 6 7-14 19-20 Normal Most nests lost during wheat harvest were either crushed by combine wheels or were covered by straw forcing abandonment after the hen was flushed. One hen returned to resume incubation in each of the 3 years (Table 11), however, in 1980, the hen nested in a weedy turnrow where the combine missed the nest. A hen, returning to her nest in 1981 failed to hatch her clutch (the embryos were potentially stressed during harvest) and she continued incubation until 8 September -- a 2 month interval. Two other nests, not significantly covered by straw, were in early incubation stages when abandoned during the 1981 harvest. Linder et al. (1960) reported most hens returned to complete incubated clutches after wheat harvest in their south central Nebraska study area. However, they searched only the portion of stubble not covered by straw during 2 years of their 5-year study and straw spreaders may have not been used on combines at that time and locat ion. Four of 16 radio-marked hens (25%) that lost nests during wheat harvest during the 3 years subsequently renested successfully. Extremely hot, dry weather prevailed in July 1980 when 2 of 5 hens successfully renested, but weather conditions were more moderate in July 1981 when only 1 of 9 hens successfully renested. However, cover type data (Table 3), body weight comparisons, and observations indicated egg laying began much earlier in 1981, possibly prompting earlier cessation of nesting effort. All known post-harvest renest efforts were successful among hens represented in Table 11. Post-harvest sweep tillage of stubble to control weeds is a common practice among eastern Colorado farmers, especially after mid summer rains. Some loss of late nests undoubtedly occurs, but since the practice usually impacts less than one-third of the stubble and nest densities are low at this time, the effect on pheasant production is low. Nest Predation.--Predators destroyed fewer nests (6) than were destroyed by cultivation (10) among documented nests in stubble fields during 1979 and 1981 when nests were found in stubble. Presumably all nests lost to predators would have subsequently been lost to tillage since only 1 of 18 nests placed in stubble hatched successfully. Thus, tillage destruction was the dominant factor in stubble field nest losses. Nest predation in stubble may have benefitted some hens by forcing them to renest in wheat at earlier dates. �34 Most confirmed nest losses to predation, machinery, or abandonment had progressed into incubation before the loss occurred. Numerous nest predations and several losses to tillage were suspected, primarily in April and May when hens were laying and nest sites could not be located by monitoring of the hen. Therefore, nest predation statistics are considered incomplete. Documented nest predation in green wheat approximated only about one-half of that verified in wheat stubble or undisturbed covers (Table 13). Linder et al. (1960) reported similar findings in their south central Nebraska study area. Greater predation in stubble and undisturbed covers probably occurred because a majority of nest placement there was earlier in the season when mammalian nest predators were feeding young. Rodent populations in stubble were also higher than in green wheat, potentially attracting increased predator activity. Avian nest predators, primarily blackbilled magpies (Pica~) and American crows (Crovus brachyrhynchos) can also find nests in open stubble more readily than after green wheat and other green vegetation provided concealment (Chesness et al. 1968, Jones and Hungerford 1972). Although documented nest predation in wheat was not high, it did force some late renesting in wheat, thus subjecting these renest efforts to potential nest loss by wheat harvest. The same could be said for nest predations in undisturbed sites with subsequent late renestina in wheat fields. Table 13. Relationship of predation of nests of radio-marked to cover type location, northeastern Colorado, 1979-81. Cover type Green wheat a Wheat stubble Wheat stubble (post-harvest) Undisturbed covers Total nests 66 18 4 14 hen pheasants Total nests lost 10 5 0 5 Percent 15. 1 27.8 0 35.7 alncludes standing residual stubble, sweep+t-ll led stubble, and 2 nests in disced stubble. Most or all predations in stubble would have subsequently been lost to stubble tillage. Pheasant nests initiated in May (most were placed in wheat) were most sutcessful in contrast to those initiated in April or June (Table 14)~ Predation and tillage had equal impacts on April initiated nests resulting in about 50% success. July wheat harvest severely impacted June initiated nests so that nesting success was low (Table 14). Nest predation rates were fairly uniform among months. �35 Table 14. Predator destroyed, machinery destroyed, and successful pheasant nests in relation to month initiated, northeastern Colorado, 1979-81. Destroyed Mechanical N % Predator N % 3 10 6 2 25.0 20.4 22.2 20.0 Month Apr May Jun Ju 1 25.0 14.3 51.9 20.0 3 7 14 2 Successful N % 6 32 7 6 Totals 12 49 27 10 50.0 65.3 25.9 60.0 Nest predation on or near (within 0.6 km) the Sand Draw property occurred at a higher than expected rate (33%) when compared to losses at more distant locations (14%) (Table 15). Nest predation was higher (p < 0.05) near the Sand Draw property in 1979 and the combined 3-year interval based on total nests found. Significantly increased predation on or near the Sand Draw property was not detected for individual years based on days of monitored hen occurrence, but the combined 3-year data were significant (p < 0.05) (Table 15). Table 15. Relationship of pheasant nest predation and nesting to the Sand Draw Property, northeastern Colorado, 1979-81.a Proximal locations Year Total nests 1979 1980 1981 8 14 20 42 Totals Distal locations Hen daysb of occur. Total nests 4 4 6 688 938 3,373 22 16 26 2 4 3 1,379 1,880 5,059 14 4,999 64 9 8,318 Predated nests % predator destroyed Predated nests Hen days of occur. 14. 1 33.3 Chi -square values Predation/nests 1979 1980 1981 6.08,': 0.06 2.48 .::. 3-year 8.62"~ 3 d.f . 3-yea r combined 5.581~ d.f. Predation/hen days Total nests/hen days 3.00 0.95 2.67 Tr Tr Tr 5.41 1 d.f. 0.19 1 d.f. aNests and predator destroyed nests were classified as on or near the Sand Draw property « 0.6 km) or away from the property (> 0.6 km). bHen days represent the total days of monitored hen presence on or near (proximal) or away (distal) from the Sand Draw property. �36 Both avian and mammal ian nest predators affected nest success on and near the Sand Draw. Several hens in the northeast part of the Sand Draw property in 1980 and to the west in 1981 were unable to advance nests to the incubation stage because of excessive nest predation. However, few of their nest establ ishment efforts were documented, if they occurred. An analysis comparing total nests on or near the Sand Draw property with those at more distant locations, based on hen days of occurrence at the 2 locations, revealed no evidence that less than the expected number of nests were placed in the Sand Draw vicinity (~> 0.05) (Table 15). Nest Success.--The proportion of pheasant hens that nest successfully is the primary factor responsible for the size of the fall population (Linder et al. 1960, Snyder 1970). Clutch size and hatching success (Table 16) have far less impact on the annual increment. Percent nesting success, based on hens surviving at mid-summer completion of major nesting effort, varied (Table 17). Highest nesting success occurred in 1979 when 94.4% of the surviving hens were successful, with progressively lower success in 1980 (76.2%) and 1981 (57.1%). The primary variable responsible for this decline was the impact of wheat harvest on nesting (Table 11). That, in turn, resulted because late stubble tillage and nest predation forced late renesting in wheat fields and weather conditions affected timing of harvest differently among years. Nesting success can also be calculated based on the number of hens initially present in spring, but varying rates of spring-to-early summer hen predation influence the results (Table 17). Table 16. Average clutch size and hatching success in relation to nesting season progression, northeastern Colorado, 1979-81. Month Apr May Jun Jul Totals Nests Eggs Young Percent eggs hatching Mean clutch size 4 28 6 6 57 301 62 47 53 259 53 36 93.0 86.0 85.5 76.6 14.2 10.7 10.3 7.8 44 467 401 85.9 10.6 Table 17. Relative nesting success of radio-marked hen pheasants among years based on mid-summer and early spring hen population bases. Variable 1979 1980 1981 Totals Hen monitored to mid-summer Successful nests Nest success, % Hens marked in early spring a Hens of known survival status Nest success of hens of known survival status, % 18 17 94.4 37 31 21 16 76.2 42 37 35 20 57.1 52 45 74 53 54.8 43.2 44.4 Mean 71.6 131 113 aContact was lost with some radio-marked hens for unknown reasons, these hens were excluded from the spring-marked sample. 46.9 �37 Hen pheasant densities progressively increased during the 3-year interval, but there is no basis to suspect that the "inversity principle" (Linder et al. 1960, Wagner and Stokes 1968) caused decreased nest success during successive years of higher population. The population was recovering following a major population crash that occurred because of reproductive failure in 1974 followed by major bl izzard losses in 1975 and 1977 (Snyder 1977). Major snow storms or winter losses were not noted during the 1978 through 1981 interval which allowed consistent population increases in spite of varying nesting success. There is no reason to suspect that nesting success among radio-marked hens was lower than that among the unmarked populat ion. Young per hen, among hens surviving to mid-summer, showed the same trend as nesting success. Approximately 7.9 young/radio-marked hen hatched in 1979, compared to 7.1 in 1980 and 5.5 in 1981. Linder et al. (1960) calculated that about 3.0 chicks per surviving hen were needed to maintain a stable population. Based on that index, production was above average during all 3 years of study. Nest success within wheat averaged 60.3% for the 3-year interval, lower than that found in undisturbed covers (64%) and post-harvest wheat stubble (100%) (Table 18). However, since many more nests were placed in wheat, 75% of' the successful nests occurred there and the proportion approximated 87% in 1979 and 1980 (Table 9). Only 1 successful nest was documented in residual wheat stubble in spring 1981 because of extremely late stubble tillage that spring. The data clearly indicate that 1ittle or no pheasant production can be expected from stubble fields in spring under current summer fallow practices. Nesting success among all cover types averaged slightly over 50% for the 3-year study interval. Table 18. Nesting success by cover type, northeastern Cover type Green wheat Wheat stubble, residualb c Undisturbed vegetation Wheat stubble, post-harvest Totals N nests Colorado, N succe"Ssful 1979-81. Percent successful 63 18 14 3 38 1 9 3 60.3 5.5 64.3 100.0 98 51 52.0 aNests were excluded where human disturbance failure. potentially blncluded residual wheat stubble, sweep-tilled in disced stubble. caused stubble, and 2 nests cUndisturbed vegetation included annual and perennial forbs and grasses and small areas of alfalfa. a �38 An average of 1.4 nests/hen was documented among hens survIving to midsummer. However, this figure was considered low as many nest attempts were suspected but were not confirmed prior to their destruction or abandonment. This was because I decided not to subject the hens to excessive harassment, thereby potentially biasing their nest site selection. Many hens nested or were suspected of nesting, 2, 3, or more times and the highest documented number of nest attempts was 4. An average of 1.8 nests/ surviving hen was reported by Dumke and Pils (1979) in Wisconsin, whereas Linder et al. (1960) reported 2.9 nests/hen and Baxter and Wolfe (1973) recorded 3.4-3.5 nests/hen in south central Nebraska. One hen attempted to renest in 1980 after loss or abandonment of a brood. Dumke and Pils (1979) reported 4 instances of this occurring during their study. Predation, Survival, and Monitoring Variables Predation and Mortality Rates.--Predation was the only documented source of natural mortality among monitored hens although some loss to other factors, subsequently attributed to predation, may have occurred. March through August mortality ranged from 45% in 1979 to 38% in 1981 averaging about 41% for the 5-6 month interval over the 3 years (Table 19, Fig. 13). This rate was probably conservative because it excluded radio-marked hens that could not be located and whose survival status was unknown. Signal loss due to predation was believed to occur more frequently than signal loss due to either egress or transmitter failure. Table 19. Survival (S) and mortality (M) of radio-marked hen pheasants by months and years, northeastern Colorado, 1979-81.a All years M S M S M Percent mor ta 1itJ: 2 10 2 0 1 1 45 39 36 36 35 33 28 6 3 0 1 2 5 113 105 86 80 77 72 66 8 19 6 3 5 6 7. 1c 18. 1 7.0 3.8 6.5 8.3c 1981 1980 1979 Month S M S Mar Apr May Jun Jul Aug Sep 31b 31 25 21 19 17 17 0 6 4 2 2 0 37 35 25 23 23 22 21 Total M 14 16 17 47 M as a % of in it ia1 S 45.2 43.2 37.8 41.6 a Excludes hens that could not be relocated, consequently of unknown survival status, and hens instrumented in Apri 1. bMarked hens alive at start of month. cMost hens were monitored only part of March during the 3 years and only part of August in 1979 and 1980 so morta 1ity data are not directly representative for those months. �39 45 C/) Z w I 1981 0 W ~ c:: -c ~, 0 0 -c c:: u, --_ 0 ..... ----1980 c:: .•.. -- -_ . w OJ -_ 1979 ~ ::::> z MAR Fig. 13. APR MAY JUN JUL AUG SEP Survival of hen ring-necked pheasants at the start of each month, northeastern Colorado, 1979-81. �40 Survival status was confirmed for 41 hens over a 11-12 month interval from March 1981 through late February 1982. Twenty-two of 41 hens died indicating an annual mortal ity rate of 54-60%, much lower than that indicated in the mid-1970's (Snyder 1977). Above average cover conditions for wheat stubble, green wheat, and other vegetation prevailed through the 1981-82 interval and no significant snowfall was received during the fall and early winter. Fall population levels and spring breeding populations increased during the 3 years indicating nesting conditions and winter survival conditions were above average. Winds were not severe enough nor temperatures cold enough to cause high mortality during the only major snowstorm occurring in late March 1980. This was in marked contrast to devastating blizzards that occurred in 1975 and 1977 (Snyder 1977). Hens were lost to predators at proportionately higher rates (p < 0.05) in early spring (X2 = 6.24, 1 df), and from spring through the remainder of monitored intervals (p < 0.05, X2 = 7.60, 1 df) on or near (within 0.4 km) of the Sand Draw property than at more distant monitored areas during the combined 3 years of study. The presence of extensive tree plantings, which provided habitat for avian and mammalian predators was considered the primary factor for increased predation on and near the Sand Draw property. However, pheasants and other prey species were concentrated on the Sand Draw property, especially through the winter and early spring, which probably attracted predators. Most suspected predation on or near the Sand Draw property was attributed to avian predators (19 avian, 6 mammalian, and 12 unclassified predations). In contrast, only 2 predations were attributed to avian predators, 3 to mammalian predators, and 5 were unclassified in locations away from the Sand Draw property. Hens moving away from the Sand Draw property in early spring greatly increased their survival chances. Greatest radio-marked hen loss in 1979 and 1980 occurred in April (after hens had adjusted to their transmitters) when 17 of 29 (58.6%) known spring-summer mortal ities were recorded (Table 19, Fig. 13). Avian predation, primarily by resident great horned owls (Bubo virginianus), was suspected in a majority of the April losses in those years. Cooper's hawks (Accipiter cooperii) and prairie falcons (Falco mexicanus) were frequently observed on or near the Sand Draw property during late winter and spring migration and were suspected of several hen predations. Wheat stubble was of average quality in 1979 when resident great horned owls were feeding young. Delayed growth of wheat that year increased hen vulnerability to predation into May (Table 19). Hens were also vulnerable in early April 1980 when deep snow covered nearly all herbaceous cover. Wheat stubble was short, partially lodged, and offered little protection at a time when hens were forming into harems with increased vulnerability to predation. Great horned owls, although present, did not nest on the Sand Draw property in 1981 and April predation was reduced. Wheat provided excellent concealment cover by 20 April 1981 and little predation occurred after that time. Six radio-marked hens were removed by predators in March 1981 compared with 3 mortalities in April and none in May (Table 19). Migrating prairie falcons and red-tailed hawks (Buteo jamaicensis) were suspected of several hen predations on or near the Sand Draw property in summer 1981, potentially prompting shifts by most surviving hens and broods to a nearby cornfield. �41 The above information strongly supports findings of Petersen (1979) in Wisconsin, who found that great horned owls and red-tailed hawks reduced and held pheasant populations below carrying capacity with peak predation occurring in April. Weigand (1980) likewise found that spring was the major period of gray partridge (Perdix perdix) mortality in Montana. He noted that the 1imiting factor appeared to be the quality and quantity of protective cover, which often was reduced by winter snow, combined with an influx of avian predators. Feral house cats and coyotes (Canis latrans) were suspected as the primary mammal ian predators of radio-marked hens on and near the Sand Draw property. Red fox (Vulpes vulpes), reported as major pheasant predators by Dumke and Pils (1973), were rare to occasionally present within the study area. They seemed to be expanding their range from nearby locations to the south and southeast and could become resident in the study vicinity at any time. The first February was still at study hen radio-mar~ed end released as an adult at study initiation in 1979 was retrapped and reinstrumented in both 1980 and 1981 and alive in April 1982. She was approaching 5 or more years of age termination. Hen Loss to Farming Activities.--Farmers in northeastern Colorado occasionally reported loss of incubating hens during spring stubble tillage. Most hens were flushed immediately prior to tillage during this study to document clutch size and embryo age, so the impact of tillage on hen mortality could not be assessed. One hen,flushed from under a sweep plow,was later found dead where she had landed; this loss could not be confirmed as being caused by the machinery. Several other hens, allowed to remain on their nests, all flushed immediately ahead of the machinery. Hens were not flushed ahead of wheat harvest in wheat fields and loss of 1 hen, nesting in dense wheat, to a combine was suspected but not confirmed. She remained on the nest until harvest occurred but her signal was subsequently lost, only to be found 10 months later when the stubble was disced exposing the transmitter. She had died close to the nest suggesting combine-induced mortality. The data indicate that hen loss to machinery was much less than the 68.8% mortality rate among incubating hens during swathing of alfalfa reported by Galbreath (1973). Transmitter Attachment, Efficiency, and Weight Relationships Solar-powered transmitters, between 150.000 and 151.999 Mhz range, harnessed in back-pack fashion, were used exclusively to monitor hens throughout the 3-year study. Flat 0.64 cm (*-in)-wide nylon elastic material was used for harnesses. In 1979, these elastic straps passed from the transmitter over the front of the wings and fastened together in the center of the breast before coming up behind the wings. This permitted the transmitters to ride too far forward on the hen's back and caused the solar panels to sometimes become partially covered by feathers at the base of the hen's neck resulting in intermittent signal transmission. Harnesses were looped directly under each wing in 1980 and 1981 with greatly improved results. Feathers, at times, partially covered the solar panels on a few hens, possibly because harnesses were too tight, again causing intermittent signal transmission, which greatly increased monitoring difficulty. �42 Transmitters became least functional during incubation in dense stands of wheat, alfalfa, or green annuals. However, persistent monitoring usually verified hen location sometime during the day, frequently when she left the nest to feed. Signal strength also was dampened by tall vegetation such as wheat or corn, and hen locations were difficult to pinpoint under those conditions. Six of 30 single-stage transmitters purchased from Wildl ife Materials Inc. (Carbondale, 111.) in 1979 contained capacitors instead of nickel cadmium batteries. These capacitor units did not produce consistent signals which severely hampered monitoring and collection of data and contact with some hens was lost. All subsequent transmitters were of the single stage nickel cadmium type except for 2 2-stage capacitor units purchased from Telemetry Systems Inc. (Mequon, Wis.) in 1981. These 2 larger units were more light efficient, permitting signal transmission to be more consistent and of greater strength. However, they weighed 24.5 gms (including harness and leg band) which represented 2.7% of the body weight of the hens carrying them. Both hens survived with these heavier transmitters for over 11 months prior to transmitter removal. Units purchased in 1979 averaged 15 gms (including harness and leg band) representing 1.8% of mean hen body weight (849 gms). Twenty-three units purchased in 1980-81 from Telemetry Systems Inc. averaged 18 gms or 2.1% of mean hen body weight. Hens were weighed during late winter instrumentation. Transmitters were recovered from surviving hens in late summer or fall in 1979 and 1980 and reused in subsequent years. Over one-half of the 24 nickel cadmium units placed on hens in 1979 still were in use and providing information through 1981, although signal strength weakened with age. The surface over the solar panels on all units dulled with age and all units to be reused were scraped, polished and recoated to increase light intake after the 1980 season. Signal losses resulted from (a) hen dispersals and movements, (b) transmitter failure, (c) feathers covering the solar panels, or (d) hen mortality. Chances of finding emigrating hens or those making long distance movements diminished with distance, time, and vegetation growth. Three hens were retrapped or recovered wearing malfunctioning transmitters. Numerous transmitters were recovered from predator-killed hens even though many of the transmitters were upside down or were partially shaded in dense vegetation. Persistent monitoring and searches were essential to maintain contact with hens or to relocate hens. The impact of transmitters on hen survival could not be determined, but may have made hens more visible to avian predators. Hens, wearing transmitters, tended to take off more slowly, be unbalanced in fl ight, and to be the last birds flushed within flocks. �43 Age-Weight-Survival Relationships Adult hens weighed more (p < 0.05) (880.59 gms + 19.49) (~+ 1 SO) than subadults (827.74 gms + 15.26) during the late February - mid March trapping interval. Hens were segregated into age classes by the proximal primary method (Wishart 1969, Greenberg et al. 1972) adapted to northeastern Colorado hens (Snyder 1977). Both subadult and adult hens increased in weight with age. For example, adult hen 6502 weighed 860 gms in 1980 and 880 gms 1 year later. Hen 6589 weighed 800 gms as a subadult, 860 gms as a yearl ing, and 890 gms:as an adult. Hens trapped on 20 April 1980 showed no noticeable weight gain (in preparation for egg laying) over previously trapped hens, whereas hens trapped on 14 April 1981 showed markedly heavier weights than those trapped earlier in the year. These data and those in Table 3 indicate that egg laying was late in 1980, probably because of late-March - early April snow-stress conditions. Adult hens survived longer (140.91 days + 21.05) (x + 1 SO) than subadu1ts (106.84 days + 18.09).\ However, survival data were truncated because hens of both age classes were recaptured in late summer and transmitters were removed bringing known survival to an abrupt halt. Therefore, direct statistical comparison based on total days of survival could not be made and the confidence limits placed on days of survival reflect this artificial termination. O. C. Bowden (pers. commun.) found that 55% of the subadu1ts compared to 79% of the adult hens survived over 100 days (~= 1.951, .!:. = 0.051). An analysis of the age-weight-surviva1 relationship, conducted by O. C. Bowden, provided some evidence that weight was a factor in increased survival. However, the results were not conclusive (Fig. 14). Mean weights of subadu1t hens surviving < 100 days and> 100 days were not different (~38 = 1.396, .!:. > 0.05). -Mean weights of adults surviving ~ 100 days and> 100 days approximated significance (~7 = 2.326, .!:. ~ 0.05). Comparative mean weights of adults and subadults surviving < 100 days were not significant (t6 = 0.366, P > 0.6) which would lend strength to the hypothesis that s~rviva1 was partially a function of weight. The sample size of adults in the latter analysis was small, weakening its importance (Fig. 14). In contrast, comparison of mean weights of adults and subadu1ts surviving> 100 days showed significance (t38 = 2.677, P < 0.05) which confuses the issue concerning the influence of weight on survival. Alternate Farming Approaches and Their Impacts on Pheasants Evaluation of Minimum Tillage Fallow With Herbicides.--A 2nd objective of this study was to evaluate minimum tillage fallow with herbicides (Greb 1977) for increasing pheasant nest success in wheat stubble, assuming adequate sample sizes could be obtained. This summer-fallow approach involved app1 ication of a pre-emergent persistent herbicide, usually atrazine, to wheat stubble within 2-4 weeks after wheat harvest singularly, or in combination with a contact herbicide if green vegetation was already present. Subsequent precipitation was needed to place the pre-emergent herbicide into the top soil where it remained over-winter retarding growth of annuals the subsequent spring. Under average conditions the herbicide gradually loses effectiveness so that wheat could be planted in September of the subsequent year, yet need for soil tillage would be delayed until late June or early July - after the primary nesting effort was completed. �44 925 900 800 775 750L-------~------~------~------~~ :5100 >100 :5100 SUBADULTS _ >100 ADULTS DAYS OF SURVIVAL Fig. 14. Mean body weights (vertical bars indicate 95% CI) for subadult and adult ring-necked pheasant hens surviving Colorado, 1979-81. < and> 100 days, northeastern �45 Sweep tillage is recommended as a replacement for discing or plowing and only 2-3 tillage treatments are usually needed prior to fall seeding (Smika 1977). This mini-till approach conserves soil moisture, tillage rep1 ication, fuel, and labor while reducing soil erosion. It has been promoted by personnel of the U.S. Department of Agriculture, Soil Conservation Service and Agriculture Research Service and by Extension personnel in several Great Plains states (Fenster et a1. 1977). Some area offices of the U.S. Department of Agriculture, Agricultural Stabi1 ization and Conservation Service (ASCS) provide limited incentive payments for trial treatments through the Agricultural Conservation Program (ACP) which they administer to farmers. The minimum tillage fallow with herbicides procedure has been tried by many wheat farmers throughout eastern Colorado in recent years, but at the present time, it has not gained wide acceptance as initially expected (Greb 1977). Problems relating to cost of herbicides and their application, their effectiveness under varying moisture conditions, and their varying persistence in different soils and pH levels (Sharman 1980, Wicks and Hergert 1980) within and among fields has tended to discourage dramatic adoption of this new farming approach in Colorado. Minimum tillage fallow with herbicides was applied by the lessee to approximately 16.2-20.2 ha (40-50 acres) of stubble on and adjacent to the Sand Draw property for 4 consecutive years beginning in the 1978-79 summer fallow cycle. Weed control effectiveness of the treatment was rated as fair to good during the first year (Snyder 1980), and tillage was delayed until approximately 1 July 1979 by the lessee. The 1979-80 treatment could not be evaluated because cheatgrass brome (Bromus tectorum) invaded the Sand Draw property fields in fall 1978 and spring 1979 resulting in almost total crop failure in 1979 and stubble too short and sparse to provide nesting cover in spring 1980. Dense stands of cheatgrass brome germinated in spring 1980 after considerable precipitation reduced herbicide concentrations below effective levels. Stubble conditions improved in the 1980-81 summer fallow cycle but the atrazine treatment was considered only marginally effective in controlling weeds. Delayed (1 Sep 1981) app1 ication was necessitated by muddy fields which stimulated dense stands of volunteer wheat in the stubble. This was not completely killed by app1 ication of a contact herbicide during the late summer 1981 treatment so the method again could not be considered effective or satisfactory. Pheasant nesting use of treated stubble was not documented in 1979 and 1981 when stubble was left standing until early July. The lessee was permitted to till the stubble in early May 1980 to remove cheatgrass brome before it matured since stubble cover was deemed inadequate for nest placement. Three factors, spring dispersal of hens (Table 2), excessive hen predation,; and nest predation associated with the Sand Draw property and its extensive tr~e plantings (Table 15) were believed responsible for lack of nest placement in the treated stubble fields. Hen densities also were too low to provide adequate nest samples preventing attainment of the 2nd study objective. �46 As the study progressed, it became evident that hundreds of hectares of wheat stubble would have to be treated along with a sizable increase in the monitored hen sample and rep1 ication for several years would be needed to obtain statistically re1 iab1e data. Such expanses were not available in the vicinity of the Sand Draw property or elsewhere in eastern Colorado. Limited data concerning the potential impact of chemical fallow treatment on nesting pheasants can be derived even though the treatment essentially failed. There was no noticeable difference in cover value for nesting among treated and untreated stubble fields anytime during the study (excluding 1980) although differences could occasionally be expected in years when considerable late summer weed growth occurred. Hens tended to select specific sites usually containing increased straw litter for nest placement rather than sites with greater densities of green vegetation. Green vegetation usually was just starting to grow during late April-early May nest initiation, so it was not a significant factor in nest site selection. Therefore, if all stubble fields in th~ area studied had been chemically treated, and the treatment had effectively delayed need for tillage until late June or early July, many first nest attempts and some renest attempts would have been successful. Two major questions are unanswered and could only be tested after several years of extensive research. First, how many of the initial nests placed in treated stubble would be destroyed by predators, and 2nd, what proportion of the renest efforts would be placed in treated stubble? Renests placed in herbicide-treated stubble fields after mid-May might be subject to subsequent destruction during late June or early July stubble tillage. This would be a major concern in years when stubble quality was high and wheat growth was retarded by drought attracting nesting hens back into stubble. Observations during 1979-81 indicate many renesting hens would probably move to wheat to renest in most years. Thus, nest predation becomes a major concern when contemplating use of chemical fallow as a replacement for conventional summer fallow to benefit nesting success of pheasants. Mid- to late April stubble disCing, forcing hens to move to wheat to nest, may be the best option in many instances, and especially in locations adjacent to trees and unfarmed sites where high nest predation could be expected in stubble. Minimum tillage techniques, employing sweep tillage can increase the amount of straw res ldue on the soi 1_surface through the subsequent winter and year of wheat growth. Excess residue has been a problem during wheat planting in the past but new grain drills that can plant in straw residue are being developed. Straw residue reduces soil erosion over-winter and may be an important factor making green wheat fields increasingly attractive as nesting cover to pheasants, as almost no surface litter is retained after conventional summer fallow operations. �47 Wheat Farming Options and Impacts.--Wheat farming and summer fallow methods vary greatly among farmers, among communities, and among years because farming must take a dynamic approach based on ever-changing weather conditions. New techniques are being developed and tested with varying success, but since farming is based on economics, most farmers are reluctant to change from old and proven methods, especially when new, expensive machinery may be required. Wheat farming options that have been or are being used, or that may have potential in the future, and their potential impacts on pheasants are summarized in Fig. 15. Post-harvest discing, plowing, or sweep-tillage (also called stubble mulching or undercutting) of stubble have been used extensively in the High Plains Region to reduce post-harvest weed growth or to promote volunteer wheat for fall-winter pastures. The practice has been especially common in southeastern Colorado. Such treatments destroy late summer renests and reduce or eliminate stubble essential to pheasants for survival from late summer through early spring, thus the carrying capacity of the land for pheasants is reduced. Other problems associated with post-harvest stubble tillage are promotion of soil drying, reduction in snow and moisture containment, and exposure of the soil to wind. Rodgers (1981) found that up to one-fourth or one-third of pheasant nests escaped tillage destruction when stubble was undercut with large sweep tillage machines (with mulch treader or harrow attachments removed) during the nesting season in north central Kansas. Even nests where eggs were disturbed, but not widely scattered or broken, were often regrouped and nesting was continued. Based on this information and because sweep tillage reduces soil erosion and conserves soil moisture, sweep tillage is being promoted, primarily through the Kansas Extension Service, as an aid to pheasants and other ground nesting birds in Kansas. However, before it can be recommended, or even encouraged for pheasant management in Colorado, 1 important question must be answered. That is, to what extent will sweep-tilled stubble be used for nesting when mulch traders are not attached and most stubble remains standing? Findings from a small sample of hens near the Sand Draw property showed that hens nested in the standing residual from stubble that had been sweep-tilled the previous suqmer, and other hens nested or renested immediately after stubble was sweep-tilled for the 1st time in spring. Hens also displayed a strong tendency to renest close to where their initial nest was located. It takes 5-6 weeks for a nest to be establ ished and hatch (assuming the hen is ready to lay) and farmers seldom wait that long before a 2nd tillage is completed (3-4 weeks between tillage operations is normal). Thus, nearly all pheasant renest efforts in sweep-tilled stubble will be lost, often at such a late date that subsequent renesting may not occur. A question also arises concerning soil and cover differences between the Kansas site and more arid locations in Colorado. Soils in north central Kansas tend to be deeper, more mellow silt-loams containing higher organic matter and tall, more dense stubble than in semi-arid areas further west. Therefore, undercutting might have less disruptive impact on soils, vegetation, and nests there. Inspections of sweep-tilled fields in northeastern Colorado indicate that few nests could escape destruction or abandonment when soils are relatively shallow and dry and stubble is short. Obviously, research is needed in different locations in the High Plains concerning nest survival during undercutting and to determine the extent and success of renesting in sweep-tilled .stubble when mulch treader attachments are not used. �SUMMER FALLOW OPTIONS POST -HARVEST (LATE SUMMER TREATMENT) FALL WINTER STUBBLE VALUE TO PHEASANTS SPRING TREATMENT HERBICIDE TREATMENT STUBBLE SWEEP TILLAGE STUBBLE DISCING I I NONE I I POOR SECOND DISCING NO TREATMENT GOOD GOOD I I I APR DISC ~ . OR SWEEP LATE JUN OR JUL DISC OR SWEEP HERBICIDE & PLANT LATE MAY DISC OR SWEEP r I I .- -. . APR DISCING CORN OR SORGHUM I APR SWEEP TILLAGE - I I LATE MAY DISCING _ -, - LATE MAY CONTACT HERBICIDE I )a J:- ex> NESTING VALUE RECOMMENDED FOR USE IN COLORADO TO MANAGE PHEASANTS NONE NONE f NO NO DESTRUCTIVE IMPACT NO FAIR TO GOOD? POOR TOa YES L1MITEDa APPLICATION FAIR a Ecofallow is marginally adapted to extreme east edge of northeast Colorado and further east. NONE (FORCED ELSEWHERE) YES DESTRUCTIVE TO SUBSEQUENT NESTS NO b Hens will nest in sweep-tilled stubble with probable nest loss to 2nd spring tillage. necked Summer pheasant fallow options survival with respect and reproduction. GOOD YES Some nests are destroyed during spring planting in standing stubble but hens can renest after planting without further disturbance. Fig. 15. NO FAIR TOb to their impacts on ring- �49 Another farming approach with possible application to better dryland areas of extreme northeastern Colorado and to center-pivot irrigated land throughout eastern Colorado is termed "ecofallow". It was developed by agronomists in southwest Nebraska and has been dramatically accepted by farmers in that region (Wicks and Nordquist 1977, Nason 1982). It involves a 3-year wheatrow crop-summer fallow rotation (Fig. 15), although the summer fallow year would be dropped under irrigation. A pre-emergent herbicide (usually atrazine) is applied to wheat stubble after harvest and again the following spring prior to seeding the row crop. Corn or grain sorghums, which are tolerant to the herbicide, are planted directly into the stubble using a "Buffalo" (brand name) slot planter that disturbs only a narrow strip of soil in each row leaving most of the stubble standing. Corn planted in early to mid-May will probably disrupt most nests, but many will still be in the laying stage and hens can immediately renest. Planting of grain sorghum in mid- to late Mayor early June would have a more negative impact on nesting, disrupting many incubated nests. In either case hens can renest in the stubble/row crop complex without being in jeopardy of subsequent machinery disruption of nests. The farmer also has the option that, if soil moisture is deemed inadequate, the 2nd herbicide treatment, applied in spring, can be halted and the approach can be switched to a wheat-summer fallow-wheat rotation. Ecofallow conserves soil, soil moisture, fuel and labor, while changing from a biennial rotation to 2 crops in 3 years. Dramatically increased yields of row crops over conventional farming methods have been reported (Wicks and Nordquist 1977). The primary concern with ecofallow Iies with the effect of extensive use of persistent herbicides on the environment, especially where runoff to streams and reservoirs is common. Population increases of small rodents have also been reported to be a problem at times. However, the approach merits evaluation as to its potential impact on pheasants in higher rainfall areas of northeastern Colorado and to irrigated situations on Division of Wildlife properties. One additional summer fallow approach for use in wheat-fallow-wheat rotations has not been used by farmers, but is recommended for use on Division of Wildlife properties. It would replace initial spring tillage with a low rate (12 oz/acre) application of the non-persistent contact herbicide, Roundup (glyphosate), sprayed with a large boom-type ground sprayer in spring after most green vegetation had made 5-10 cm growth. Tillage of stubble could then be delayed until early July, after the primary nesting season when sweep tillage could be used. The spring application would not conserve soil moisture used by weeds the previous late summer, but would be much more consistently effective than a post-harvest appl ication of atrazine. Roundup is much more expensive than atrazine but can be used effectively on annuals at lower rates. An equally effective herbicide, paraquat, is also less expensive, but is highly toxic (for restricted use by certified appl icators) and should not be substituted. Relating Study Findings to Past Research Controlled Hen Harvest.--A northeastern Colorado study, initiated in 1963 by H. M. Swope and W. W. Sandfort, attempted to evaluate the impact of Iimited hen harvest on a pheasant population. Public and pol itical opposition prevented implementation of hen harvest, but census and environmental data were collected and evaluated for 6 years (1963-68) (Snyder 1970). �50 The hen pheasant has long been cons idered a "sacred cow" protected from hunting because she can be easily distinguished from the rooster. Yet the basis for 1imited hen harvest was not clearly understood or defined at initiation of the hen harvest effort. Numerous studies conducted since that time, including this 3-year study, have provided some insight into hen pheasant population dynamics, but justification for partial hen harvest still cannot be easily explained. Justification for hen harvest can be divided into 2 categories: (a) reproduction, and (b) survival. Most 1imiting factors to reproduction in wheat farming areas of eastern Colorado do not appear to be density dependent, and therefore, it would be difficult to justify hen harvest based on reproduction. To illustrate, vast wheat fields, when available for nesting, provide habitat far in excess of the needs of existing pheasant populations. Likewise, major hinderances to reproduction in the form of stubble tillage and wheat harvest will destroy a certain percentage of nests no matter what hen density or nest density exists. Therefore, the more hens available for nesting, the better, based on these influences. Predation of hens, young, and nests (2 of these are survival factors) are the primary influences that could have density-dependent impact during the reproductive period, but the degree that higher density causes increased percentage losses not considered great among our low density populations. Hen survival from fall through early spring dispersal (to late Apr) is the other variable which must be considered in justification of controlled hen harvest. Bl izzards periodically devastate pheasant populations on the plains of eastern Colorado with 1ittle regard to hen density. Predation, as during reproduction, becomes the only major variable where hen density could influence the percent removed. Snow cover and sparse, short vegetation are factors working with predation during this time. Early spring predation appears to be the primary bottleneck, most years, having densitydependent potential for which controlled hen harvest would be a replacement. It appears that 1imited hen harvest could be conducted, primarily in years of higher than average populations. However, if pheasants were less impacted by bl izzards and by intensive land use influences on nesting, and therefore, were more stable in number, hen harvest would be a much more viable option. Environmental Impacts on Reproduction.--Efforts to relate land use, vegetation, and weather to pheasant reproduction were made, based on 6 years of environmental and census information obtained in the 1960's (Snyder 1970). Some insight concerning environmental influences was obtained at that time. However, a much more precise picture of environmental - pheasant reproduction relationships was obtained during this study, showing impacts of soil moisture and summer fallow, stubble height, wheat growth phenology, and timing of stubble tillage and wheat harvest. Regrettably this study lacked the census base present during the 1960's study, and that study lacked information on some environmental parameters, preventing more complete analyses of the combined data. �51 This study has shown that if environmental and population monitoring could be continued over several years and combined with census information, a strong predictive model could be developed. Insight is also needed on the impacts of severe hailstorms, blizzards, and droughts (none of which occurred during this study) on pheasants and additional data on weather influences, primarily in spring and summer are needed. The CY Index.--Analyses of the 1963-68 census data revealed that 2 indices: BPM (birds-per-mile) obtained during brood counts, and CY (crowing count x young/hen) were strongly correlated (r = 0.97) (Snyder 1970). Either could be used to predict fall population level, but plotting of the 2 on a regression Iine was recommended to compare and potentially strengthen confidence in census findings. A formula incorporating winter or spring sex ratios as a measure of hen density had previously been used to predict fall population level, but was found to be less accurate than either the CY or BPM indices. The reason why the crowing index (C) represented breeding populations more accurately than when combined with sex ratio (CH) to project spring hen density was not clearly understood. Based on telemetry study findings, variable and often rather high hen predation in late March and April may be the primary factor weakening CH as an indicator of breeding population level. Crowing counts were obtained after the major period of spring mortal ltv . : \Jhen used alone, they more accurately represent spring breeding population ,'density than when combined with sex ratio data to project hen density. It should be noted that (Y) (young/hen) was a direct function of percent successful hens surviving to late summer (r = 0.95) (Snyder 1970). It was the most important variable in determining fall population level within the CY index. CONCLUSIONS AND RECOMMENDATIONS 1. Winter wheat, the major cash crop in northeastern Colorado, is the dominant site of successful pheasant nests. Data indicate that 7590% of the annual pheasant production comes from green wheat in most eastern Colorado Tablelands. 2. The amount of precipitation received in any 1 year often indirectly affects pheasant reproduction and survival that year and for 2 subsequent years. For example, soil moisture accumulated during the summer fallow year, influences wheat growth the next year which affects stubble qual ity the 3rd year. This influences early spring nest placement which is primarily determined by the relative qualities of stubble and green wheat. More nests will be placed in high qual ity stubble if green wheat has a low height-density value that spring, and vise versa. 3. Nearly all nests placed in wheat stubble will subsequently be lost during initial stubble tillage which primarily occurs in May. Stubble tillage conducted in April or early May, before hens begin nesting or while they are still laying, forces hens to green wheat to renest. Most will renest immediately to successfully complete clutches there in mid- to late June. �52 4. Indication of a trend toward later (late May-early Jun) stubble tillage was evident during this study. Late tillage destroys many nests which have advanced far into incubation forcing hens to delay 1-2 weeks to physiologically prepare for egg laying before they can renest. Most renest efforts will be in wheat fields where June initiated nests will be in jeopardy of destruction by wheat harvest. 5. Nearly all nests in wheat fields not hatched prior to time of wheat harvest wi 11 either be crushed or abandoned. Most hens losing nests to July harvest will not renest. 6. Among the variables of wheat quality, stubble quality, timing of stubble tillage, and wheat harvest, only time of stubble tillage can be significantly manipulated to increase pheasant production. Stubble tillage preferably should be completed by mid-April, prior to 1 May if possible, and before 10 May at the latest to reduce impact on pheasant production. Discing, undercutting (with mulch treaders attached), or plowing of stubble to flatten or reduce surface residue in early spring will force hens to move to green wheat to renest. Undercutters with mulch treaders attached to flatten the straw should be used where wind or water erosion is a problem. This would reduce its attractiveness for renesting. 7. Use of either late summer or spring applied herbicides and minimum tillage are recommended as alternatives to summer fallow assuming the initial stubble tillage operation can be delayed until at least 25 June or, preferably, later. Only low toxicity herbicides such as atrazine and glyphosate are recommended for use of stubble fields. Paraquat is highly toxic with a restricted-use label and is not recommended for use on stubble. Delay of stubble tillage without use of herbicides is not recommended. 8. Wheat stubble, left standing after harvest, is an essential component of pheasant habitat in eastern Colorado used extensively for feeding, roosting, resting, and escape from fall through early spring. Tall, dense stubble can reduce predation and increase survival during blizzards. Wheat should be cut as high as practical (0.3-m is minimum and 0.5-m is preferred) to increase fall-winter protection to pheasants. Use of herbicides to replace sweep tillage of stubble fields is recommended when post-harvest weed control is essential. 9. Wheat acreage reduction programs, administered by the U.S. Department of Agriculture's Agricultural Stabilization and Conservation Service (ASCS) have a negative impact on pheasants in the Great Plains. Destruction of wheat in June, to comply with acreage quotas, is a common and especially devastating practice to nesting pheasants. Surplus small grain acreages should either be destroyed prior to 1 May, or left standing unharvested through fall and winter. �53 10. Pheasant nesting will be earl ier in years of early wheat growth and later in years of retarded wheat growth. Wheat growth is primarily influenced by the amount of precipitation received during the summer fallow year and the spring when growth occurs. Spring temperatures have much less impact on wheat growth. 11. Grazed or ungrazed mid- and shortgrass sites usually do not provide attractive nesting cover to pheasants in eastern Colorado. Major portions of these if not pastured should be tilled and revegetated to grass-legume mixtures with subsequent periodic renovation to retain more vigorous growth. Strips should be mold-board plowed in February or March and allowed to revegetate with annual weeds. Biennial rotation tillage of these strips is recommended to retain seral vegetation. 12. Roadsides were nearly all farmed to the road shoulder within the study area, as elsewhere in northeastern Colorado, and therefore, were not important for pheasant production. Little opportunity for reclaiming roadsides exists in wheat farming areas, once they are lost. Revegetation to grass-legume mixtures is recommended where barrow ditches now exist or where farmers would permit revegetation in efforts to curtail the continued loss of roadside cover. Limited opportunities exist for revegetation of newly reconstructed county roadways, but only if the Division of Wildlife works closely with county commissioners and local farmers. 13. Sunflowers and other tall, wild annuals provide essential mid- to late summer brood habitat and travel lanes along roadsides even in many cases where field edges have been extended to road shoulders. County commissioners and their road maintenance crews usually defer major roadside mowing efforts until late summer or early fall to reduce the need for mowing repl ications. Late mowing reduces impacts on broods and should be encouraged. 14. Significantly greater than expected hen pheasant predation occurred near trees in early spring based on hen presence at a time when resident raptors were feeding young and migrating raptors were common on the Sand Draw property. Hens were also lost at a higher than expected rate to predation near trees through the late April to late summer interval. Much the same impact could be expected in relation to shelterbelts and riverbottoms throughout eastern Colorado. 15. Significantly greater than expected nest predation occurred in locations on or near the Sand Draw property with its extensive tree plantings than in areas further away. �54 16. Shrubs, such as American plum (Prunus americana) should be used in preference to trees, especially if trees are not already present, to provide winter cover for pheasants while reducing the impacts of avian predation on hens and nests. Shrubs should be planted in short, wide blocks to form thickets and placed in sites protected from inundation by drifting snow when possible. Young trees can be half-cut and older trees can be topped at a height of 1 - 2 m to create and maintain a low, shrubby growth form if only a few trees are present and all can be treated. This will el iminate elevated hunting perchei for raptors at pheasant concentration sites whi le improving the value of the woOdy cover for wintering pheasants. 17. Tree plantings are not a salvation to pheasants during severe winter or early spring bl izzards in eastern Colorado. Pheasants on the Sand· Draw property, with its extensive plantings, were devastated by the March 1975 and 1977 storms just as populations in surrounding localities were affected (Snyder 1977). Woody cover is primarily of value after the storm, assuming food sources are present. Tall wheat stubble, 2nd year growth of sweet clover, and tall weed or sorghum patches provide better bl izzard survival habitat than does woody cover. 18. Hens gained weight with age and average survival increased among older hens that were wearing transmitters. Whether the increased survival was due to age or weight, or a combination of both, could not be accurately assessed. LITERATURE CITED Baxter, W. L., and C. W. Wolfe. 1973. ring-necked pheasant in Nebraska. Publ. 58pp. Brander, R. B. 1968. A radio-package Manage. 32:630-632. Life history and ecology of the Nebr. Game and Parks Comm. Tech. harness for game birds. J. Wildl. Brubacher, J. I., and T. R. Moore. 1969. Soil survey of Sedgwick County, Colorado. U.S. Dep. Agric., Soil Conserv. Serv., Washington, D.C. 61pp. Buss, I. 0., R. K. Meyer, and C. Kabat. 1951. Wisconsin pheasant reproduction studies based on ovulated follicle technique. J. Wildl. Manage. 32:630-632. Carter, A. V. 1971. Season movements and behavior of ring-necked pheasants in eastern South Dakota. M.S. Thesis. S.D. State Univ., Brookings. 44pp. Chesness, R. A., M. M. Nelson, and W. H. Longley. 1968. The effect of predator removal on pheasant reproductive success. J. Wildl. Manage. 32:683-697. �55 Dumke, R. T., and C. M. Pils. 1973. Mortal ity of radio-tagged pheasants on the Waterloo Wildlife Area. Wis. Dep. Nat. Resour. Tech. Bull. 72. 52pp. and -_,--.,1979. by Wisconsin pheasants. Renesting and dynamics of nest site selection J. Wildl. Manage. 43:705-716. Fenster, C. R., H. I. Owens, and R. H. Follett. 1977. Conservation tillage for wheat In the Great Plains. U.S. Dep. Agric., Ext. Servo Publ. PA-1190. Washington, D.C. 32pp. Galbreath, S. S. 1973. Pheasant population studies and pheasant losses from alfalfa mowing operations in the Columbia Basin of central Washington. Proc. West. Assoc. State Game and Fish Comm. 53:326-335. Gates, J. M., and J. B. Hale. 1974. Seasonal movement, winter habitat use, and population distribution of an east central Wisconsin pheasant population. Wis. Dep. Nat. Resour. Tech. Bull. 76. 55pp. Greb, B. W. 1977. Principles of fall weed control and residue management in a fallow - winter wheat rotation. Pages 13-17, ~ A. King, compi ler. U.S. Dep. Agric., Soil Conserv. Servo Agron. Notes 54. 38pp. Greenberg, R. E., S. L. Etter, and W. L. Anderson. 1972. Evaluation of proximal primary feather criteria for aging wild pheasants. J. Wildl. Manage. 36:700-705. Hartman, F. E., and D. E. Sheffer. 1971. Population dynamics and hunter harvest of ring-necked pheasant populations in Pennsylvania's primairy range. Proc. Northeast Sec., The Wildl. Soc., Fish and Wild]. Conf. 28:179-205. Hoffman, D. M. 1975. Pheasant mortality investigation. Pages 5-35 in Colo. Div. Wildl., Fed. Aid Proj~ Rep. W-37-R-2B, Apr. Jones, R. E., and K. E. Hungerford. 1972. Evaluation of nesting cover as protection from magpie predation. J. Wildl. Manage. 36:727-732. Kuck, T. L. 1968. Movements and behavior of pheasants during the breeding cycle as determined by radio-tracking. M.S. Thesis. S.D. State Univ., Brookings. 73pp. Linder, R. L., D. L. Lyon, and C. P. Agee. 1960. An analysis of pheasant nesting in south-central Nebraska. Trans. North Am. Wildl. and Nat. Resour. Conf. 25:214-230. Nason, G. 1982. 44-45. Ecofallow - All to the good. Nebraskaland 60(3):38-39, Petersen, L. 1979. Ecology of great horned owls and red-tailed hawks in southwestern Wisconsin. Wis. Dep. Nat. Resour. Tech. Bull. 111. 63pp. �56 Robel, R. J., J. N. Briggs, A. D. Dayton, and L. C. Hulbert. 1970. Relationships between visual obstruction measurements and weight of grassland vegetation. J. Range Manage. 23:295-297. Rodgers, R. 1981. Saving nests in fallow wheatfie1ds. Conf. 43:94 (Abstr.). Sandfort, W. W. and Parks 1957. Aging pheasant embryos. Game Infor. Leaf. 15. 2pp. Seubert, J. L. 1952. necked pheasant. Midwest Wi1d1. Colo. Dep. Game, Fish, Observations on the renesting behavior of the ringTrans. North Am. Wi1d1. Conf. 17:305-329. Sharman, E. D. 1980. Effects of soil types on the breakdown of chemicals. Not paginated. In Proc. Colo. Annu. Farm Inst. Eco+fa llow Conf. Wray, 12-13 Feb. Smika, D. E. 1977. Residue management, wheat-fallow. Pages 17-25 in A. King, compiler; U.S. Dep. Agric., Soil Conserv. Servo Agroil:"" Notes 54. 38pp. Snyder, W. D. 1970. Pheasant hen harvest investigation. Pages 3-81 ~ Colo. Div. Game, Fish, and Parks. Fed. Aid Proj. Rep. W-37-R-23. Apr. 1974. Pheasant use of roadsides for nesting in northeast Colorado. Colo. Div. Wi1d1. Spec. Rep. 36. 24pp. Pages 1-33 in Colo. 1977. Pheasant mortality investigation. Div. Wi 1d1. Fed. Aid Proj. Rep. \/-37-R-30. Apr. 1980. Evaluation of nesting cover preferences of pheasants in ----r:;lation to wheat farming methods. Pages 3-110 in Colo. Div. Wi1dl. Fed. Aid Proj. Rep. W-37-R-35. Apr. Trautman, C. 1960. Evaluation of pheasant nesting habitat in eastern South Dakota. Trans. North Am. Wi1d1. and Nat. Resour. Conf. 25: 202-213. Wagner, F. H., and A. S. Stokes. 1968. Indices to overwinter survival and productivity with implications for population regulation in pheasants. J. Wi1d1 '.Manage. 32:32-36. Warner, R. E. 1979. Use of cover by pheasant broods in east-central 111 inoi s . J. Wi1d1. Manage. 43:334-346. We igand, J. P. 1980. Ecology of the Hungarian partridge Montana. Wi 1d1;· Monogr. 74. 106pp. in north-central Wicks, G. A., and P. T. Nordquist. 1977. Ecofa110w in wheat-sorghum corn-fallow rotation. Univ. Nebraska, North Platte Exp. Stn. Ext. Rep. 4. 1Opp. or �57 , and G. Hergert. 1930. Soil types and choice of chemicals. ----~N~o-tpaginated in Proc. Colo. Annu. Farm Inst. Eco-fa110w Conf. Wray, 12-13 Feb: Wishart, W. 1969. Age determination of pheasants by measurement primaries. J. Wi1d1. Manage. 33:714-717. of proximal ��59 JOB FINAL REPORT State of Colorado Project No. W-37-R-35 Work Plan No. Job Title: 3 Game Bird Survey Job No. 12 Potential Impacts of Strip Mining on Sage Grouse Movements and Habitat Use Period Covered: Personnel: 1 January 1979 through 31 March 1982 D. Hein, R. A. Ryder, R. G. Walter, Colorado State University; Clait Braun, Steve Emmons, Howard Funk, Ken Giesen, Sue McElderry, Brett Pete~sen, Steve Porter, Tom Schoenberg, John Wagner, Colorado Division of Wildl ife. ABSTRACT Sage grouse (Centrocercus urophasianus) movements and habitat selection were studied in North Park, Colorado during April-August 1979 and FebruaryAugust 1980. Sixteen male (12 adults, 4 juveniles) and 22 female (13 adults, 9 juveniles) sage grouse were captured and fitted with radio transmitters. Mortality of radio-marked sage grouse during the moni.toring period was low (13%). Raptors were the most important predators. Twenty-two of 36 (61%) transmitters were recovered after use on sage grouse. Wildlife Materials transmitters had longer (p < 0.05) average life (209 days) than AVM transmitters (136 days). Sage grouse used 2 major wintering areas in the northeast and southeast quadrats of North Park in 1980. Preferred winter habitat encompassed only 3.7% of the sagebrush (Artemisia spp.)-dominated land in North Park. There was no difference (p > 0.05) between sexes in average daily winter movements or size of winter range areas. Daily movements averaged 1.6 and 1.5 km for males and females, respectively. Winter flock break-up and dispersal to breeding areas began during the 2nd week of April coincident with the onset of the spring thaw. Movements of 3 radio-marked males from the wintering area to leks averaged 27.5 km. Four hens traveled an average of 29.9 km from the wintering area to nests. Daily movements of males from leks to feeding-loafing (FL) sites averaged 0.9 km. Dispersal direction from Raven Lek to FL sites was nonrandom (p < 0.001). Average distance that hens traveled from leks to nest sites was 2.7 km. Adult hens traveled. farther (p < 0.05) than juvenile hens. Preincubation movements from nests to FL sites averaged 0.4 km. �60 Movements of both sexes from breeding areas and nests to meadows along the Michigan and Canadian rivers occurred throughout June, primarily during the latter half of the month. Four of 5 radio-marked males and 5 of 6 radio-marked hens moved to the meadow nearest the lek attended or nest site, respectively. Summer movements were restricted to relatively small areas along the Michigan and Canadian rivers. Few differences in slope and aspect were observed between habitats selected by sage grouse and random sites. Greater differences in habitat selection were seen when topographic features were examined. Sage grouse winter FL sites were primarily in sagebrush-dominated draws and on windswept ridges whereas breeding season FL sites were predominantly ------------GR-O~~~pen-~~pe~e~~~~~+OGd~_P.~e+4~~+aW$-WW~~------------------ little sagebrush and high forb and grass cover. Sage grouse selected winter FL habitats with better (p < 0.05) structural cover (sagebrush clump size, plant dimensions, canopy-cover) than breeding season FL sites. Structural characteristics of nest sites, however, were more similar to winter FL sites. Only leks had poorer (p < 0.05) structural cover than random sites. Except for FL sites of mal.es during the breeding season, sage grouse chose sagebrush with higher (P < 0.05) percent foliation (75-78%) than found at random sites (65%). Males, hens with broods, and unsuccessful hens chose summer meadow habitats with similar (t> 0.05) forb and grass cover and grass height. Males selected breeding season FL sites with higher (p < 0.05) soil organic matter content than did hens or that found at random sites. The higher organic matter content was attributed to 2,4-0 spraying of the area around Raven Lek in 1963 and subsequent decomposition of sagebrush plants. Discriminant function and principal components analyses were also used to investigate sage grouse habitat selection. Three discriminant functions explained 93.6% of the total sample variance whereas 5 principal components explained 94.1% of the sample variance. In both analyses, sagebrush plant size was the most important habitat factor separating different types of sage grouse use and random sites. Degree of microhabitat selection was the 2nd most important factor followed by sagebrush clump size and canopy cover. These analyses revealed differences in habitats selected by sage grouse between and within seasons as well as habitat differences between random and sage grouse use sites. �61 INTRODUCTION Sage grouse are widely distributed American rangelands sagebrush vities dominated by sagebrush. for breeding, nesting, has been extensively Griner 1938, Patterson Aldrich developments Schneegas of sagebrush range by burning, spraying, ing. treated between inated that burning tices on sage (Peterson their 1957. grouse on acti- 1937, Rasmussen and both numbers historic plowing, range lands the previous lands et al. effects 35 years cutting, and beat- in Colorado had been 1976). in Jackson by 2,4-D spraying, 1952, 2-2.5 million hectares chaining, 30%of all sagebrush Detrimental that grazing, and distribution (Patterson during discing, 1975, Beck 1977). for agriculture, (1967) estimated 32%of the sagebrush since of sagebrush 1900 and 1974 (Braun had been disturbed and wintering (Girard had been disturbed Approximately dependence 1974, Wallestad et al. have reduced throughout 1963). North 1952, Klebenow 1969, Eng and Schladweiler Elimination and alteration of sage grouse western Their brood-rearing documented 1972, Wallestad and Schladweiler and other throughout Beck (1975) esti- County, Colorado plowing and seeding, and of sagebrush prac- have been documented 1970~, Wallestad 1975), nesting control for breeding habitat and brood-rearing (Klebenow 1970, Martin 1970), and wintering areas (Higby habitats 1969, Pyrah 1972) . Many important coal deposits. sage grouse With increasing habitats demands are underlain for western by extensive surface-mined coal, �62 a substantial reduction Colorado and other coal accounted in sage grouse western states. for less than habitat In 1968, western 1980). mined areas and federal by state tion practices emphasize introduced establishment of the sagebrush term process, in surface-mined 2%of all coal mined in the United States compared to 20%in 1978 (Slatick is required can be expected Although grasses reclamation laws, current and forbs. community through of rehabilita- Because re- succession mined areas will be lost as sage grouse is a long habitat for decades. In the face of decreasing throughout require the West, management of sage grouse a more complete understanding sage grouse-sagebrush by sage grouse relationship. to discussion or have been too general greater number of habitat used by sage grouse fully understand and winter habitats Klebenow's variables sage grouse habitat selection. blue grouse (Dendragapus obscurus) territories 96%of the time using 10 habitat variables. function nesting, of nesting analysis to identify Martinka data between sites sites will be necessary (1969) study has been the best attempt use Examination of a select breeding, in Idaho using discriminant of habitat canopy cover and and comparative and random or nonuse of the all seasons but have to be of value. factors will description studies of sagebrush how sage grouse habitats. during rangeland populations and detailed Previous have been conducted been limited primarily height amounts of sagebrush to brooding, and brood with 20 habitat critical factors in (1972) was able to distinguish breeding discriminant territories function from non- analysis with �63 It is especially specific habitat preferences mining or other grouse habitat northeast and proposed breeding, disturbance grouse population This study preferences theses tested in Jackson during nesting, critical seasonally County, to identify with mining area in the Consequently, habitat sage specific seasonal habitat North Park, Jackson County, 1979 and February-August movement patterns 1980. and 4) sage grouse habitat habitat Hypo- differ between of the annual cycle (winter, 3) sage grouse for each sex. Colorado and winter habi- 2) sage grouse movement patterns brood-rearing), sexes within seasons, sage grouse County. 1) sage grouse periods by guidelines in this area is locally migratory throughout April-August sexes within seasons, brood-rearing, (Beck 1975). was conducted were: and of sage North Park will impact the entire of sage grouse Colorado during and undisturbed in the principal of the Park in northeastern understanding coal mines in Jackson nesting, concentrations portion seasonal ranges mitigation and rehabilitation The sage grouse population major wintering vary After a better to improve disturbed Existing critical of sage grouse in areas to be disturbed needs is gained, will impact historic tats. to identify developments. can be developed habitats. important for each sex breeding, selection selection differs varies between �64 STUDY AREA The investigation Colorado. North The study Park study terrain of the area varies ridges of the Canadian ranges northern boundary from Pierre of the recent from relatively and benches and It is bounded of the major River north of Cowdrey. River on the Moore Mountain. area are derived of the late Cretaceous coal beds, by tributary Drainage and soils of the study shale sediments of the Canadian sandstones period (Miller and conglomerates (Hail and Leopold 1960). and Michigan river valleys Alluvial are of more origin. Beekly cline district 1). flat to rolling separated into the North Platte Paleocene and Eocene epochs materials (Fig. Total area within these to 2,758 m atop Johnny 1934) and Coalmont shales, of River and on the west and from 2,420 m along the Canadian Geologic formations birds 8 and 9 north, and Michigan rivers. is to the northwest Elevation 2). quarter 250 km2. is approximately with numerous drainages (Fig. County, of North Park (400 38-481 N, 1060 05-15' W). and east by the Canadian Topography all areas on movements of radio-marked by the Michigan River boundaries Jackson in the northeast area lies within townships 78 and 79 west on the north rivers area was centered depending The primary south in North Park, 13 km east of Walden although were included ranges was conducted (1915) considered to be of greater the coal beds of the McCallum anti- commercial importance than all the �65 ~ N ~ 5 0 10 KILOMETERS • DENVER COLORADO Fig. 1. North Park. Jackson County. Colorado. Intensive the stippled area lying between the Canadian and Michigan study rivers. area is �66 t ,ACANUCK N .A PRAGUE ~ o 1 , 234 , I KILOMETERS l{) (\j o o -c a: 9 o o 4 LEKS COAL OUTCROPS Fig. 2. Jackson Sage grouse leks County, Colorado. and coal outcrops in the study area, North Park, �67 remaining coal areas in North Park combined. most of the region between (Fig. 2). by coal. coal outcrops from Perdiz Present selection The vegetative sites, River. tation consists and grasses primarily (Artemisia tridentata) and sedges 5 species bunch spp.), Shrubby occurring species and irrigated Herbaceous in North Park. Approxi- by big sagebrush area, alkali sagebrush occurs vege- and forbs. Within the study big sage- (A. longiloba) et al. in limited areas occurring (Sarcobatus and antelope whortleleaf currant movements 1966). on poorly soils. (Chrysothamnus gooseberry 2 on Raven and alkaline soils (Robertson (A. cana viscidula) black greasewood valleys include 2). 2). grasses of sagebrush (Smith 1966). dry site shrubby species (Fig. along native type is occupied dominates on shallow claypan Other radio-marked and Michigan rivers. dominates on loamy soils whereas nonalkaline (Fig. of sage grouse of birds of perennial mately 90%of the sagebrush Silver sagebrush on the community of the area is dominated by sagebrush (1960) identified include is concentrated region area and an additional and Denmark leks hay meadows of the Canadian drained the entire of the McCallum anticline Studies were primarily Lek but also from Perdiz brush boundary are 5 known leks in the study and habitat Beetle crest, mining activity along the eastern of the Canadian on upland of the anticlinal valleys Lek in the south to Denmark Lek in the north There leks north occupies the Canadian and Michigan river With the exception is underlain The district on the study vermiculatus), bitterbrush on limited moist upland snowberry (Ribes montigenum), rabbitbrush (Purshia sites (Symphoricarpos area tridentata). and river vaccinioides), and willows (Salix spp.). �68 Average study annual herbage area ranges productivity Sagebrush 4 areas study between area operations in the control practices (Terwilliger disturbed to crested were conducted Approximately area were conducted 2 km north wheatgrass in 1958. in 2,250 ha (9%) of the by plowing and reseeding, A 138-ha area sp.) and Smith 1978). in the study 1958 and 1964. (Trifolium and to 62-80 kg /ha on the Coalmont-Claypan types plowed and planted yellow clover types resource have been ing with 2,4-D. rangeland from 129-152 kg /ha on the Sandstone-Gravel Coalmont Shale resource and Shale-Alkali of sagebrush and spray- of Denmark Lek was (Agropyron cristatum) and The most massive spraying in the areas around Raven and Perdiz leks. In 1963, 955 ha of sagebrush were block-sprayed around Raven Lek . In 1964, 777 ha of sagebrush were block-sprayed around Perdiz Lek . A 378-ha block, Most of these although treated 4 km west of Denmark, sprayed areas dead sagebrush have recovered plants was sprayed well after remain standing in 1963. 17-18 years throughout the areas. The climate of North Park is cold and dry with an average frost-free period southwesterly Precipitation weather of 46 days (U.S. and temperature (Table 1). records summer temperatures (Table did not vary was heavy 2). through early were obtained January-June was 26 and 59%above the 30-year years of Commerce 1979). winds are common from winter station There Dep . greatly Winter-early 30-year snowfall and snow accumulation Total snowfall during summer. in 1979 and 1980 respectively. from the Strong from the Walden precipitation average, annual November-March averages. during both was similar �Table 1. January-June temperature, Walden, Colorado, 1979-80. (em) May Jun Totals Jan 0.71 3.53 1. 24 3.45 3.15 13.73 -12.4 -6.3 -3.8 5.74 1. 98 1. 88 1. 47 6.17 0.10 17.34 -7.1 -5.7 -4.6 1. 30 1. 07 1. 27 1. 83 2.59 2.82 10.88 - 9.2 -7.7 -4.6 )an- 1979 1. 65 1980 a and PreciEitation Mar Apr ~Fe~ Year Avg precipitation Mean temEerature Mar Apr Feb (C) May Jun 6.3 11. 2 -0.1 6.6 12.3 1.8 7.1 11. 6 1.6 --aThirty-year Table 2. Year 1978-79 1979- 80 average, November-March 1941-70. snowfall Total snowfall Jan Feb Nov Dec 9.9 56.1 30.7 11. 4 75.7 18.8 and snow depth, Walden, Colorado, (em) 1978-79 and 1979-80. ()'\ '.::J Maximum snow deEth (em) Dec Jan Feb Mar Totals Nov Mar 9.9 32.0 l38.6 5.1 30.5 38.1 38.1 7.6 14.7 22.9 143.5 12.7 10.2 45.7 38.1 20.3 �70 between years 1 month later mid-April. although than snowfall patterns usual in both winters, were different. occurring during Snowmelt was early to �71 METHODS Sage grouse leks throughout on foot using net (Braun were captured the study a spotlight bands Sex, area. Birds Captured size 14 (females) and unnumbered plastic were determined for all birds clips were marked color-coded 1975), weight, 1972) attached with 15-21 g tail-clip 17-g solar-powered collar (Amstrup Transmitter Service radio-collar with and primary molt on or near Raven Lek mounted on tail to the 2 central rectrices. throughout radio packages. the study In addition, and a 26-g battery-powered a radio- 1980) were placed on a female and male, respectively. packages (Denver hoop captured. 1980, 12 males and 17 females trapped area were fitted or to year of capture. with 14-25 g 164 MHz radio transmitters (Bray and Corner During from a truck and size 16 (males) aluminum leg In 1979, 3 males and 4 females trapped were fitted and on with a long-handled grouse bandettes age (Eng 1955, Beck et al. along roads were located and were captured and Beck 1976). serially-numbered while roosting were obtained Research Company (Champaign, Ill.), Center, from the U. S. Fish and Wildlife Denver, Colo.), and Wildlife Materials, AVM Instrument Inc. (Carbondale, Ill. ) . Daily locations of radio-marked feeding-loafing sites, yagi antenna. A snowmobile was used during in areas inaccessible and nests using sage grouse by 4-wheel drive a portable vehicles. were made at leks, receiver winter and 3-element to approach All locations, birds �74 Soil types and capability Geological Survey Package tributions of classification analysis. Multivariate determine if differences ences. functions analysis of the habitat use and random sites. factor rotation analysis principal of variance function to distinguish Principal components analysis sage grouse with varimax on linear functions variables. Whereas discriminant maximizes distances between sites in multi-dimensional were considered significant differ- was used to form linear between sites based describes on habi- for the observed analysis variates analysis dis- chi-square among sites based the relative site in space without maximizing inter site distances All tests Frequency habitat components was used (MANOVA) was used to were accounting was used to ordinate of highl y correlated among all sites. (P < 0.05) existed discriminant (SPSS) programs St u derrt+s t test data were compared using U.S. Colorado. for the Social Sciences variables and what factors Stepwise from 7.S-minute County, 1975) were used in data analysis. to compare means of habitat tat factors, were obtained soil type maps of Jackson The Statistical (Nie et al. classes positions function space, of each (James 1971). at P < 0.05 unless otherwise noted. �75 RESULTS Radiotelemetry During 1979- 80, 36 transmitters adults, 9 juveniles) (Tables 3, 4). were e~uipped 1979. and 16 male (12 adults, Three adult with tail-clip transmitters were fitted and 29 June 1980. with battery- transmitters grouse 2 juveniles) 18 April and 9 May and 17 hens Adult male 8991 and adult and solar-powered sage (2 adults, between 4 juveniles) with tail-clip on 22 female (13 4 juveniles) cocks and 4 hens Twelve males (8 adults, 7 juveniles) were fitted (10 adults, between 1 February hen 5088 were equipped radio-collars, respectively (Tables 3, 4) • Breeding around season captures Raven Lek in 1979 and around leks in 1980 (Fig. along J. C. roads 2). hens Fish and Wildlife Service = (~= 15.7 g) than 20.4 g). transmitters Raven, Winter captures were concentrated Denmark, and Perdiz in 1980 were concentrated 10 and 12. Sage grouse (x of sage grouse were equipped transmitters transmitters Sage grouse (Table 4). primarily (Table obtained 3) which were lighter from Wildlife Materials cocks were equipped Transmitter with AVM and U. S. packages of 1.09 and 0.73% of female and male grouse primarily weighed (WM) with WM an average body weights, respec- tively. Mortality period was low. of radio-marked sage grouse Only 2 of 22 hens (9%) during the monitoring and 3 of 16 cocks (19%) �76 Table North 3. Transmitter Park, Colorado. data and 1979-80. kncwn mortality of radio-marked female sage grouse in Transmitter Age Band number Manufacturer a Weight Percent of bird weight (g) Comments Juveniles 3401 3415 3428 3492 5072 b 5072 5086 5087 5089 5090 USFWS AVM AVM AVM AVM AVM AVM AVM AV1>1 AVM 14.8 15.0 15.9 16.2 15.5 15.7 15.6 15.9 16.3 16.2 0.95 0.91 1. 33 1. 08 1. 01 1. 26 1. 07 1. 05 1. 06 1. 08 AVM AVM AVM AVM AVM AVM AVM USFWS WM AVM AVM AVM WM 15.0 15.9 15.4 16.1 16.1 15.9 15.9 14.5 20.3 16.2 15.9 16.2 17.1 1. 01 0.96 1. 10 0.88 1. 05 1. 06 1. 33 0.97 1. 39 0.99 1. 42 1. 00 1. 17 16.0 1. 09 Shot 13 Sep 1980 Shot 21 Sep 1980 Adults 3411 3427 3429 3430c 3432c 3490~ 4499 4796 5032 5063 5064d 5075 S088e Avg Range 14.5-20.3 aUSFWS = U.S. WilCilife_ Materials. after bTwo different 47 days. Fish and Wildlife transmitters used on transmitter used on 3430 and dSame transmitter used on 3490, radio-collar. 5072 since and owl predation Unknown cause Raptor predation Shot 13 Sep of death 1980 42 the 3432. 4499, horned AVM = AVM Instrument Service, cSarne eSolar-powered 0.88-1. Great 5064. initial transmitter Co., WM = expired �77 Table 4. Transmitter data and known in North "Park, Colorado, 1979-80. mortality of radio-marked male sage grouse Transmitter Age Band number Manufacturer a Weight (g) Percent of bird weight Comments Juveniles 8731 8801 8981 8982 WM WM WM WM 19.7 20.1 20.2 20.4 0.78 1. 00 0.83 0.78 WM WM WM USFWS AVM WM WM WM AVM WM WM WM 20.8 20.5 21. 3 15.3 15.8 20.1 20.6 20.3 15.8 20.7 19.7 26.3 0.70 19.8 0.73 Recaptured 15 Apr 1981 Adults 7016 7149 7181 7327 7367 8745 8747 8802 8829 8879 8882b 8991 Avg Range 15.3-26.3 ~M = Wildlife Materials, AVM Instrument Co. b Battery-powered USFWS radio-collar. = U.S. Golden eagle predation 0.8;; 0.67 0.48 0.53 0.71 0.79 0.80 0.58 0.63 0.65 0.89 0.48-1. Fish Recaptured Recaptured Raptor 7 May 1981 23 Apr 1980 predation Recaptured 23 Apr 1981 . Recaptured 7 May 1981 Raptor predation 00 and Wildlife Service, AVM = �78 were known to have been killed by predators 3430 was killed by a great night she was captured raptor approximately Cause of death frozen into a snowbank and radio-marking. during on the same she had left the nest with no apparent She was at least injuries 5 years cold weather with her She was found 4 days after season. (- 38 C) during to her death. Three while traveling raptor (males 8747, 8882) occurred kills and appeared depending distance rectrices limitations, radio-marked upon transmitter and premature and transmitter Twenty-two during Both other season Five cocks were recaptured and monitoring of 36 (61%) transmitters (Table were recovered. were made after rectrices predation (5), recapture mortality (I). was recovered. premature (4), rectrix 4). equal. and 10 of 15 WMtransmitters recoveries on leks The recov- were approximately Only 1 of 3 USFWS transmitters and unknown receiving or normal molt of the central of AVM and WMtransmitters (6), for varying movements beyond radio package kill (I), 1980 (Aquila the breeding were monitored life, package. Eleven of 18 AVM transmitters recovered. the to summer range. birds i or 2 years following original capture ery rates early to have been by golden eagles. The remaining periods from breeding of hens monitored Male 7016 was killed by a golden eagle chrysaetos) capture old and the trauma 1979 (3401) and 1980 (5063, 5086) were shot during hunting Hen Hen 4796 was killed by a of hen 4499 was unknown. may have contributed 3, 4). (Bubo virginianus) and radio-marked. combined with severely February owl 2 weeks after brood. capture horned (Tables were Tail-clip loss of the central molt (4), The solar-powered hunter radio-collar �79 was recovered after it had either slipped from or had been pulled off by the hen. Transmitter 6) transmitters Average life was calculated used transmitter were recovered was being during 1980. for AVM (Table No data were available life was evaluated and monitored tracked. using or those Transmitters only those which shut on birds since the birds the area while transmitters were still functioning. difficult to maintain maximum line-of-sight WMtransmitters receiving tr'an smit ter s contained mitters contained Average and averaged radios 5, 6). even man ufacturer between greater for battery since the <5 km whereas Differences AVM and WM in power output. whereas WM AVM trans- batteries. life of WMradios than Average contact It was especially as far as 15 km , to differences 73 days longer off while the bird was usually distances 1. 35 V mercury which may have moved out of 3.0 V lithium batteries transmitter (Tables have been distance receiving were attributed radios with AVM transmitters could often be heard in maximum line-of-site transmitters contact for 1979. with which radio was lost were not included radio 5) and WM (Table was greater the average transmitter life of AVM life of WMradios had not 7 transmitters replacement transmitter (~ < 0.05) before would been returned to the the old batteries had expired. Movements Winter Sage grouse to early April winter 1980. movements Sage grouse were studied began from early February moving into the study area �80 Table 5. Colorado, Channel Life of AVM transmitters Pulse Transmitte b life (days) 1 56 2 2 3 3 47 249 207 58 213 42 60 60 47 67 44 64 51 4 5 5 6 6 8 8 65 10 47 64 12 45 9 a used on sage grouse in North Park, 1980. Avg Range aSMl with Comments Radio package " " recovered " and monitored " " " " " 212 days Lost radio contact after Radio package recovered Lost radio contact after " " " and shut " " off 10 days "121" 34 Radio shut off when tracking bird Lost radio contact after 73 days 37 Radio shut off when tracking bird Radio package recovered by hunter Radio shut off when tracking bird Radio package recovered and monitored Lost radio contact after 72 days " 129 47 169 " " "17" 136 34-249 1. 35 V Hg batteries. bOn1y those radios which tracking bird were included. were recovered or which shut off while �81 Table 6. Life of Wildlife Materials in North Park, Colorado, 1980. Channel Pulse 2 5 6 7 8 8 9 9 9 10 11 11 11 Transmitte life (days)b 50 47 51 48 42 43 46 59 60 59 53 66 72 prior Only those recovered " " " " " " " II II II II " II " " " " " " II II " Lost radio contact after Radio package recovered Lost radio contact after Radio package recovered 208+ 145 260+ 193+ 199+ grouse and monitored " " 246 II II II II II Lost radio II contact " " after II 128 days and monitored 11 days and monitored II 87 days 81 " 209 145-260+ aHLP-2750-LD b a used on sage Comments Radio package 199+ 215+ 214+ Avg Range transmitters with radios 3.0 V Li batteries. which were recovered + . 'I'r-an srni tte.r returned to shut off. to manufacturer were included. for battery replacement �82 following a 2-day blizzard had been located and none had been trapped Maximum snow depth 10 em but increased There northeast in Walden before to almost 46 Use of these areas quadrats ridges available of the flat, numerous sage ridges, benches, areas sively 3). used areas respectively. North Park sought . (4,692/125,200) Although quadrat, they distance in the 3). the winters diverse ern) with after covered most the bliz zard on food and cover in areas sage grouse and southeast 6,077 and 5,051 ha, and south 2 preferred areas with of approximately known to have moved back Inten- were 2,282 and 2,410 ha , only 3.7% land in North Park. in the northeast moved into the southeast 12 km. wintering respectively. followed were radio-marked all eventually encompassed quadrat encompassed of the sagebrush-dominated all birds grouse and drainages. in the north These 2). (1975) during deep snow (up to 67 The northeast were approximately (Table Park in 1980 (Fig. are topographically Known movements of all radio-marked 20,692 ha (Fig. time. and sw ales , and on windswept throughout grouse to this used by sage by Beck Both areas Because open areas 28-29 January. of North in deep draws and hillsides. prior Few birds the storm was approximately areas was documented of 1973-74 and 1974-75. on 28-29 January. following the storm ern were 2 major wintering and southeast sagebrush in North Park quadrat, a Adult male 7149 was the only bird and forth between the 2 areas on 2 occasions. Average sexes (Table daily winter 7). movements The greater due to the difference range in receiving were similar (!: > 0.05) between of daily movements by males was distance between WMand AVM �'1.' COWDREY E\ o fS8] OCCASIONAL USE t REGULAR USE [~:,'·:.lINTENSIVE USE N ~ o 1 2 3 4 5 ! KILOMETERS I[) C\J •..... Fig. 3. Jackson Winter use areas of radio-marked Coun t y. Color ado. 1980. sage grouse in North Park. �84 transmitters used on males and females. easily be located daily even after a hen moved a long distance. respectively. Cocks could movements of 8-10 km whereas it often took several if days to relocate her. Table 7. Daily movements and winter Park. Colorado, 1980. (km) Daily movements Sex N x Males 57 1.6 0 Females 34 1.5 0.1- Distance able habitat eagles sexes size of winter ing distance period used x 4 7,212 2 , 564- 10, 692 4 5,314 3,846- Range by preference by golden eagles. and pursuing 7.487 to have for suit- As many as 5-6 flocks of sage grouse range (P > 0.05) for males and females the same wintering areas. for males was probably of AVM radios The greater average due to the shorter receiv- used on hens. movement by adult cocks to the northwest in late February This erratic N but rather sizes were similar A long distance adult (ha) areas. Winter range and both 5.3 in North of daily movements did not appear were seen daily flushing in wintering banded -10.9 by my activities and disturbance of sage grouse Winter range Range and direction been influenced ranges quadrat was not included move was interpreted male 8802 and 3 other during as part as a premature a 2-day warm of his winter range. movement to his �85 breeding area since he later quadrat during Spring Migration attended the breeding Winter flock break-up Bighorn season. and dispersal to breeding during the 2nd week of April 1980 coincident spring thaw. sexes. There Adults was little difference were located in breeding following departure from wintering areas less than was probably cock (7149) and 1 adult Lek In the northwest areas with the onset of the in departure areas areas. began dates between approximately 1 week Tr-avel time to breeding 1 week for most birds. At least 1 adult hen (5032) moved as far as 20 km during a 2-day period. All radio-marked area by the first east quadrat had left the northeast week of April. wintering of hen attendance apparently birds area occurred on leks. also served Peak of departure quadrat area prior wintering from the south- only 2 weeks prior The southeast as a staging quadrat to the peak wintering to dispersal area to breeding areas. Known movements from the southeast were to the northwest quadrats (Fig. to breed and attended center 4). (2 cocks, averaged quadrat Spring area. the northwest 1 hen) and southwest Creek #2 Lek, area (l cock, 4 km southwest Movements of 3 radio-marked to leks in the northwest 27.5 km (range of 29.9 km (range wintering Adult male 8829 remained in the southeast of the wintering the southeast quadrat 20.5-34.7). 25.6-35.1) and southwest from the southeast quadrats. quadrat quadrat of the cocks from and southwest Four hens traveled 3 hens) quadrats an average to nests in �86 ~ N J o 5 10 KILOMETERS m WINTER AREA A LEKS • NESTS Fig. 4. Dispersal quadrat wintering Colorado. 1980. direction of radio-marked sage grouse from the southeast area to leks and nest sites in. North Park. Jackson County, �87 Breeding Season Six adult 1 juvenile and 2 juvenile male from Perdiz males from Raven Lek, were monitored from late April adult all 3 juvenile males but the breeding (P < 0.01) averaged years to early between was attributed (!:_ the years. to a smaller than movements during 95.7% of all movements 2 = Dispersal direction 48.65, = df all FL movements the 5). accurate 1 of 9 1 lek during 3, !:_ locations = 12) than were the end of the breeding to be greater lst 'half of the month. from Raven Approximately Lek were within were within from Raven Lek between In 1980, most locations 2 km of the Lek to FL sites Forty-eight were to the northwest, Thirty-three from Raven The difference from the lek tend < 0.001). (FL) sites sample size in 1980 (N to FL sites 17.7% (23) to the southeast, (Fig. Lek from lek to FL sites 1. 3 km in 1980. 62% (61. 9) of all FL movements (X Only more than Distance 2nd half of May toward when daily < 0.10) 1979-80. Daily movements 1979 (N = "80) and timing of locations. season June to feeding-loafing 0.03-2.4). 0.9 km in 1979 and made during and males from Denmark males attended from all leks 0.9 km (range differed 2 adult 1 adult season. Daily movements averaged and Lek, 1 km and lek. was nonrandom percent (62 of 130) of 28.5% (37) to the southwest, and only 6.2% (8) to the northeast percent were recorded (32 of 96) of all FL sites were within the proposed for which Kerr coal lease. Dispersal more productive movements direction range from Raven may be due to selection sites. Seventy-six percent for sagebrush on (99 of 130) of all Lek were to the west of Williams Draw in the �88 N s Fig. 5. Daily movements of male sage grouse from Raven Lek to feedingloafing sites during 1979-80. Each concentric circle represents 0.5 krn , The proposed Kerr coal lease is outlined. Four locations >2.0 km are not shown. �Sandstone-Gravel herbage and Coalmont Shale resource productivity williger is 129 and 152 kg/ha/year, and Smith 1978). Coalmont-Claypan east of Williams Draw where average lowest (62 kg/ha/yr) deeper, dominates where average respectively productivity Big sagebrush east of Williams Draw (Robertson is the whereas less productive et al. to the dominates soils west of Williams Draw, on the shallow, (Ter- soils predominate herbage in North Park. more productive sagebrush types alkali claypan 1966, Terwilliger on soils and Smith 1978) . Few data were available Perdiz leks to FL sites but random from both leks. on daily movements dispersal Dispersal direction direction appeared (8 of 9) from Denmark Lek , and northwest east (3 of 11) from Perdiz sites The mean distance juvenile km (range 0.8-7.9) juvenile hens. hens. Sixty-four (range 0.3-6.4). to southwest sites northeast breeding hens percent lek-to-nest distance of all hens nested 36%nested nest sites. and southeast on Raven Lek nested of the lek. within 2 hens 1.5 km 3.0 krn , was 2.5 km on Raven Lek The other was 3.8 0.3-1. 4) for beyond from Raven Lek to 7 nests Five of 7 hens breeding to select during from leks to nest and 0.5 km (range The remaining The mean distance were obtained (P < 0.05) farther The average for adult of the lek where bred. (5 of 11) and south- was 2.7 km for all known lek-to-nest Adult hens traveled than north- Lek. Movements of 11 hens from leks to nests movements. and to be non- was predominantly west 1979-80. from Denmark moved south selected Only 1 radio-marked within the proposed Kerr nest hen. coal lease. �90 Nests of adult overburden hens 3411 and 5063, however, piles on areas Preincubation 0.4 km (range movements 0.1-1.8). prior to FL sites were shorter mined south Movements averaged hens to incubation. Preincubation (P < 0.001) than of Raven Lek. averaged 0.3 km for adults 0.05) (~= 100 m of to FL sites Approximately but only 63.9% of juvenile of the nest being of hens from nests (P < 0.05). 0.5 km for juveniles hens currently were within and 89% (88.9) of adult fed within 0.5 km movements from nests daily movements of males from leks to FL sites. Feeding sites Mean distance used during from the nest incubation were located was 160 m and ranged 4 times. from 40 to 420 m. Postbreeding Daily movements range 70-910). during Distance the first distance of 3 hens with broods traveled week after traveled during with each of the broods from the nest hens from 0.3 to 1. 5 km. the first week, however, traveling approximately Although broods respectively. the meadow area the nest. 1.7, 1.8, and uplands and one-half meadow areas the distance Broods nests, to the nearest ha tc hin g, a distance in of had begun, In 1980, hen 5063 and her brood 2 weeks after 1.8 km. for 2 weeks after had only moved 0.8 and 0.9 km from their mately one-fourth area, toward Total (3415, 4796) with broods in sagebrush 12, was more consistent in 1980 by hen 5063 and her brood. movements = (!:::! by each of 3 broods varied 3415 and 4796 remained hatching. these than 320 m hatch Movements to meadows by 2 hens 1979 were later averaged approxi- meadow had arrived at of 3.8 km from �91 In 1980, movements to meadows by unsuccessful occurred primarily during meadows appeared ficient mid- to late June. to be 1- 2 weeks later data to accurately interesting hens, to note, assess however, and males occurred vegetation dessication although in sagebrush between (range sexes averaging 1. 8-7.0) hens nesting Distances near for the summer. to the Canadian Birds to or nest 3.0-7.5) (P > 0.05) for cocks and 4.5 km Perdiz Lek and both Lek moved to the Michigan River meadows from Denmark and Raven leks River meadows. and hen 5086, nesting as a response the lek attended One male attending Perdiz It is unsuccessful moved to meadows were similar 4.6 km (range for hens. years. males and 5 of 6 radio-marked females moved to the meadow area nearest respectively. were insuf- between probably to uplands. In 1980, 4 of 5 radio-marked site, there movements of broods, simultaneously, and males In 1979, movements any difference that hens However, 1. 0 km southwest Michigan River meadows during generally moved male 8991 from Raven Lek of Raven, moved to the the summer. Summer All birds restricted movements to relatively river meadows. overlapped their small areas Summer ranges for birds summer (late Jun through along the Michigan and Canadian along the Canadian from Denmark and Raven leks. ranges along the Michigan River meadows overlapped Perdiz and Raven leks. Approximate the Michigan and Canadian Hen 5063 and her brood, river the Michigan River meadows apart areas River meadows Similarly, for birds of summer ranges meadows were identical however, Aug) remained summer from along at 333 ha each. within a 128-ha area along from all other radio-marked birds. �92 Habitat Sage grouse variables those (Table habitat 8). selection Variables which were thought of habitats. measure Average Selection was analyzed chosen for analysis to be important intercept distance of clump size since it explained of the variation P < 0.001) in clump width. of the variation = 2, 31; ~ < 0.001). grouse (ATINDIS) Approximately 92% (R plant selection ~ < 0.001) 0.868, = in clump width was explained Microhabitat in selection was used as a = 75.4% (r 26 habitat were limited to to sage sion of clump width on ATINDIS and average df using 0.957, by a regres- (!:._ = height was examined 169.73, with ABRDPLEN, ABRDPWID, and ABRDPTHT. Slope There occurrence were few differences within each (0-5, among random and sage sites (Table and leks FL sites was less between sites that use sites except than at all other hens at unsuccessful their sites. = hen FL hen The difference and other of the small sample size (N of 15%) slope class Mean slope at leks and unsuccessful (~ < 0.05) restricted slope or frequency 11-15, and> used by unsuccessful ably the result ful hens 6-10, grouse 9). in average sites was prob- 13) for a few unsuccess- movements to relatively flat areas. Aspect Mean values selection of south-facing ence for east(Table of aspect 10). slopes to south-facing Mean aspect compared among sites by grouse slopes during during at random sites indicated winter the breeding was similar possible and preferseason (P > 0.05) to �93 Table 8. Habitat variablesa used in analysis of sage grouse and random sites in North Park, Colorado, 1979- 80. use Mnemonic Description SLOPE Percent ASPECT Aspect ATINDIS Average transect, APLNLEN Average crown length line transect, em APLNWID Average crown width of sagebrush line transect, cm APLNTHT Average transect, ABRDPLEN Crown length of the plant beside which the bird feeding-loafing or under which nest was located, center of plot at random sites ABRDPWID Crown width of the plant beside which the bird was feeding-loafing or under which nest was located, center of plot at random sites ABRDPTHT Height of the plant beside which the bird was feedingloafing or under which nest was located, center of plot at random sites AFSBCC Foliated ATSBCC Total sagebrush PERFOLCC Percent foliation of sagebrush canopy cover, = AFSBCC .;. ATSBCC at breeding season and random sites, = EFSBCC .;.ETSBCC at winter use sites SBDENS Sagebrush FORBCC Forb GRASSCC Grass ETINDIS Average transect EPLNWID Average crown width of sagebrush plants under line transect exposed above the snow, ern slope at site corrected for declination sagebrush cm height cm of sagebrush canopy canopy density, canopy distance of sagebrush sagebrush canopy intercept cover, cover, plants plants plants cover, cover, under the line under under under the the the line was % % p lan t s Zm+ % % sagebrush intercept distance under exposed above the snow, em the line the �94 Table 8. (Continued) Mnemonic Description EPLNTHT Average transect EBRDPWID Crown width of the plant exposed above the snow beside which the bird was feeding-loafing, em EBRDPTHT Height of the plant exposed above the snow beside which the bird was feeding-loafing, cm EFSBCC Foliated sagebrush snow, % ETSBCC Total sagebrush height of sagebrush plants under exposed above the snow, cm canopy canopy cover exposed cover exposed the line above the above the snow, % TOTCC Total canopy meadows, % TARAXAC Canopy cover of dandelion TRIFOL Canopy cover of clover in meadows, GRASSHT Grass ayariables the text. height cover of forbs in meadows, are presented and grasses in meadows, in summer % % em in the order in which they appear in �95 Table 9. Colorado, Slope at sage grouse use and random sites in North Park, 1979-80. SloEe class ( %) N ~(%) 0-5 6-10 11-15 >15 63 7.5 38.1b 41. 3 12.7 7.9 50 7.6 40.0 38.0 14.0 8.0 93 6.5 49.5 35.5 10.8 4.2 Female PIFL 44 6.8 56.8 25.0 15.9 2.3 Nests 17 7.4 .50.0 22.2 16.7 11.1 Brood FL 23 5.7 60.9 26.1 8.7 4.3 13 2.2 100 5 1.8 100 80 6.2 20.0 13.7 7.5 Sitea Male winter FL Female winter Male spring FL Unsuccessful Leks Random loafing FL hen FL = ~L feedin g-loafing site. b Values are percent site, 58.8 PIFL = preincubation in each slope class. feeding- �96 Table Park, 10. Aspect at sage Colorado, 1979- 80. grouse use and random W-N 25.4 25.4 23.8 20.0 28.0 34.0 18.0 116 55.5 30.0 6.7 7.8 58 143 41. 4 27.6 15.5 15.5 19 157 31. 6 36.9 10.5 21. 0 23 141 43.5 30.4 8.7 17.4 13 106 61. 5 15.4 15.4 7.7 5 129 40.0 20.0 40.0 76 159 35.5 23.7 25.0 :x (0) N-E 63 189 25.4b 50 185 90 Female PIFL Nests a Male winter FL Female winter Male spring Brood FL FL FL Unsuccessful hen FL Leks Random loafing ~L = feeding-loafin site. b Values Aspect E-S in North class S-W N Site sites are percent g site, in each PIFL = preincubation aspect class. feeding- 15.8 �97 breeding season each (N-E, preference sites. E-S, Comparison of the frequency S-W, and W-N) aspect for aspect by males during ence for E-S and S-W aspects Disproportionately grouse during the breeding fairly season sites. prefer- of aspect by sage of the study at random sites However ,- sage grouse N-E and E-S facing slopes to avoid str ong prevailing west winds during no winter. season may be representative distribution selecting and only slight during in indicated use of N-E and E-S aspects area since the frequency similar to breeding however, winter by hens higher class, distribution was may be south- April-June. Topography It is more instructive tion rather than to consider slope or aspect random sites were classified physiographic selected winter swept ridges sites features draws. in draws into 5 topographic categories whereas (Table There were, breeding however, used by broods late spring plants contained Preferential sagebrush in winter contained cover where sagebrush in winter and early great season and those tall stands little sagebrush for forbs on and random The only were in in vegetation found used by broods. Draws with high canopy above deep snow. Draws but had good forb and grass use of draws can be explained summer. based and on wind- sites of sagebrush were available and selection use and ~3. 5%of all locations differences situa- Sage grouse on 0- 5 and 6-10 % open slopes. used by grouse winter 11). in draws and swales, was at brood FL sites where used during cover. Sage grouse FL sites primarily were predominately exception topographic separately. of the habitat and hilltops the overall by selection and insects by broods for in �Table 11. Topographic features of sage sites in North Park, Colorado, 1979-80. grouse use and random To,eograEhic 6-10% >10% open open slopes slopes feature Windswept ridges & hilltops N 0-5% open slopes 66 19.7 18.2 50 18.0 8.0 93 43.0 31. 2 10.8 Female PIFL 59 49.2 20.3 16.9 Nests 19 36.8 26.3 21. 1 Brood FL 23 30.4 13.0 4.4 8.7 43.5 20.0 13.7 7.5 1.3 a Site Male winter FL Female winter Male spring FL FL Unsuccessful hen FL Leks Random "TL site. b 13 100 5 100 80 = loafing feeding-loafing Values b are percent 57.5 site, PIFL 10.6 = 19.7 31. 8 28.0 46.0 15.0 1.7 preincubation in each topographic Draws and swales type. 11. 9 15.8 feeding- �99 Preference be understood accumulated during for windswept and sagebrush areas used during to draws the breeding - Univariate use and random sites (P < 0.05) breeding structural season (P > 0.25) between cant difference average plant sexes distinct open areas were preferred variables (Table were compared 12). Sage grouse FL sites than except nests. There d uring winter, however. (p. < 0.01) between width although from open with deep snow in winter cover at winter use sites used during 1980. Analysis Sample means of 10 habitat grouse that or no snow in the intensively Extensive diversity can also Open slopes and ridgetops season. season were covered Structure since little 3) and were therefore with little topographic the breeding Habitat (Fig. availability in winter was always available. were adjacent used wintering (benches) and hilltops in terms of sagebrush winter slopes ridges values winter among sage selected better at any of the were no differences The only signifi- FL sites and nests of this and other was in habitat variables were lower at nest sites. Due to heavy be expected greater that at winter sagebrush at winter snow accumulation average FL sites was available nonuse values sites during of habitat variables since sage grouse above the snow. and approximately winter sought 1979-80, it would would be much areas Snow depth where averaged 20 ern at sage grouse 40 ern winter FL sites. Among breeding the greatest season sites, (~ < 0.05) average and canopy cover. nesting sagebrush Values of these habitat hens selected clump size, variables habitats plant at nest with dimensions, sites were �100 Table 12. Habitat variables a Site compared ATINOIS Female winter sage grouse use and random ASROPWIO sites in North b Habitat variabJe ASROPTHT AFSSCC Park. ATSSCC Colorado. 1979-80. APLNWID APLNTHT PERFOLCC SSDENS FORSCC 70.1 5.3 49 57.7 2.6 50 42.3 3.5 50 72.3 3.6 50 51. 0 3.9 50 39.1 2.6 49 50.9 3.2 49 77.9 1.7 49 63.8 4.3 .50 57.2 2.5 53 40.6 3.1 53 75.0 3.5 53 50.2 3.6 53 38.9 2.1 50 49.6 2.7 50 77.1 1.3 50 NO 57.6 7.1 19 42.6 3.6 17 34.8 2.7 19 72.1 4.3 17 52.0 2.6 19 32.2 3.0 19 44.0 4.2 19 74.8 3.0 19 3.3 0.3 13 1.7 0.4 19 43.7 3.9 13 42.0 3.3 12 32.5 2.4 13 61. 0 7.1 12 42.1 3.5 12 28.7 3.4 13 38.2 4.5 13 75.9 2.8 13 3.1 0.4 11 5.2 1.8 13 40.2 1.8 92 37.2 1.8 68 29.6 1.1 92 63.9 3.0 67 45.8 1.4 88 24.5 1.6 92 35.0 1.5 92 66.5 2.5 92 3.2 0.4 26 2.6 0.8 92 37.4 4.3 23 36.6 4.3 18 26.3 2.5 23 65.5 4.0 18 38.9 2.3 22 22.2 3.2 23 27.3 3.3 23 77.0 6.7 23 3.2 0.6 9 6.9 2.4 23 35.2 1.7 59 34.5 1.6 41 23.6 1.2 59 58.9 4.3 41 38.2 2.1 50 21. 6 1.3 59 29.1 1.6 59 74.5 2.2 59 3.2 0.3 19 3.2 0.4 59 26.2 1.2 80 28.0 1.1 77 21. 0 1.0 77 34.7 2.2 69 24.8 1.6 69 17.4 1.2 80 26.0 1.6 80 65.2 2.5 80 3.3 0.4 19 2.5 0.3 80 19.6 2.9 20.4 2.7 10.4 2.0 NO ND 10.2 2.2 5 11.2 2.6 5 93.4 4.1 2.1 0.7 4.8 1.8 5 5 FL x SE N Male winter among NO FL x SE N NO Nests x SE N Unsuccessful- hen FL x SE N Male spring FL x SE N Brood FL x SE !:!. Female PlFL x SE N Random x SE N Lek~ x SE N a FL feeding-loafing bMnemonics eND 5 5 of habitat = no data. site. PIFL variables 5 == preincubation described feedin g-loafing in Table 8. site. 5 �101 not greater (~ > 0.10) than at unsuccessful chose spring FL habitat similar not as good (~~0.05) sites were essentially as nest sites. Preincubation There variables variables sage grouse than at random sites. Average tion and brood FL sites Average and brood FL between average sagebrush use sites sagebrush sagebrush height was also similar preincubation FL and random sites. were greater (~ Most habitat (P < 0.05) cover at preincuba- (P > 0.10) to random sites. was similar plant 2 sites. clump size, were greater canopy (P > 0.25) these and canopy cover than random sites. at all other hens but were no differences measured Only leks had lower (~~0.05) plan t dimensions Males also (P > 0.25) to unsuccessful identical. among any of the habitat hen FL sites. (P > 0.10) between Microhabitat < 0.05) at all sage grouse plant dimensions FL and nest sites than at random sites. Sage grouse foliation FL sites, Except for leks, any other foliation site, there (74.5-77.9) defoliated for sprayed regeneration plants sagebrush of sagebrush, 12). PERFOLCC at (P > 0.25) to random sites. of percent sage grouse foliation than sagebrush use sites. did not select higher the breeding areas (Table < 0.05) percent < 0.05) percent was uniform selection during sagebrush percen t foliated (!: among all other (!: with higher was similar male sage grouse sagebrush tolerance however, which had greater The reason their sagebrush (PERFOLCC) than at random sites male spring foliated preferred season may be explained around remained Raven Lek . standing after canopy cover was low. however, percent and foliated Because 17 years, There by many the was good sagebrush canopy �102 cover (AFSBCC) and total sagebrush still much greater canopy cover (P < 0.001) at male spring (ATSBCC) were FL sites than at random sites. There were no differences (~ > 0.05) in sagebrush (SBDENS) among any of the sage grouse 12) . Whereas structural with several differences of the other habitat form among all sage grouse sagebrush density tion between different When considered however, grouse habitat ing season nificant nest 12). sage grouse sites in habitat brush was uni- Taken alone, in habitat selec- or canopy cover, component in understanding sage and unsuccessful to be the result were not measured later Although average forb cover be attributed in the breeding (Table 14) classes the only sig- another selection. useful cover of sagebrush of more forbs (!:_ = -0.456, canopy cover by sage grouse cover and in > 50 ern frequently height = ~ 0.001). (Table 13) and at random means of viewing overlap Whereas sage 3rouse with >60% canopy brood FL and High forb cover at leks may canopy selected breed- and unsuccessful to selection season. of the ranges at brood hen of plant than other (P < 0.05) in forb cover was between due to low sagebrush provided density dimensions use or random sites. can probably Comparison and height 2 sites the greater the diet later plant This did not appear since these hen FL sites be partly and necessary were evident use or random sites. forb cover was found at brood difference sites, of sage grouse (Table selection. (Table phenology sagebrush of differences along with sagebrush The highest FL sites variables, selection use and random sites. types it is a useful use and random sites in habitat gives no indication density and differences selected during sage- winter �103 Table 13. Frequency distribution of sagebrush canopy grouse use and random sites in North Park, Colorado, Canopy N Female winter Male winter 0-10 11-20 so FL 2.0 Sl-60 14.3 20.4 4.1 16.3 36.7 6.0 12.0 10.0 24.0 14.0 32.0 10.S 15.8 21. 1 31. 6 21. 1 17.4 2S.0 18.5 21. 7 10.9 5.4 3S.6 23.7 11. 9 3.4 1. 7 30.4 21.7 8.7 4.3 ,4.3 53.8 7.7 7.7 IS.4 17.5 21. 3 Male spr in g FL 92 1.1 Female PIFL S9 3.420.3 Brood 23 17.4 13.0 13 7.7 7.7 5 40.0 60.0 80 17.5 21. 3 hen Leks Random aFL = FL feeding-loafing site, (%) 41-S0 19 Unsuccessful class 31-40 Nests FL at sage 21-30 49 FL cover cover 1979-80. 22.5 PIFL = pre-incubation >60 feeding-loafing site. b In percent. cUnderlined class. Totals values may only indicate approximate most frequently 100%. used canopy cover c �104 Table 14. Frequency distribution of sagebrush height grouse use and random sites in North Park, Colorado, N Female winter Male winter FL FL 0-10 50 2.0b 53 9.4 11-20 Height class (cm) 21-30 31-40 41-50 Male spring FL 22.0 6.0 10.0 9.4 17.0 18.9 15.1 30.2 52.6 15.8 15.8 15.8 3.3 92 2.2 12.0 44.6 31. 5 6.5 Female PIFL 59 5.1 30.5 49.2 6.8 8.5 Brood 23 8. 7 17.4 43.5 17.4 8.7 7.7 30.8 46.2 15.4 36.4 9.1 2.6 FL Unsuccessful hen FL Leks Random 13 5 60.0 40.0 77 11. 7 40.3 ~L = feeding-loafing loafing site. b In percent. cUnderlined Totals values site. PIFL = preincubation may only approximate indicate >50 22.0 19 Nests at sage 1979- 80. most frequently 4.3 feeding- 100%. used height class. �105 1980, breeding season FL sites were more commonly selected 21-30% canopy cover and 21-30 sites were in canopy winter and breeding sagebrush selected prior height brush season FL sites. No nests < 21 cm. selected of sagebrush selected greater by grouse were in sagebrush > 50 crn , the cover and height that winter the available FL and nesting males, habitat hens prior Sagebrush exposed sagebrush than to incubation, plant to be selecting the only difference height more often than >30 sites. substantially ern Only sites tall. No by This suggests less preferred summer FL habitat unsuccessful plant grouse used most frequently dimensions, hens, 1980 (Table 15). for and broods. of hens and cocks Although structural (P < 0.05) between winter and canopy cover at FL sites sites with better above the snow. sage by random sites. spring-early clump size, where sage- canopy cover> 50%or height classes contains and at random sites in winter cocks, classes averaging above the snow were measured appeared However, 1980 and often as nest habitat hens cover and height 44.9% of the FL and nest had average during than males, of canopy sites. as indicated random sites grouse range at FL and nest 11. 7%of the random sites but selected hens frequently All 5 leks were on sites a broad available between were found where Unsuccessful canopy cover and height what was typically intermediate nest were < 21% and 21 cm, respectively. canopy cover and height classes Preferred canopy cover and height and broods. Sage grouse classes. classes sagebrush to incubation, height cover and height averaged greater ern in the sexes hens cover than was in average �Table 15. Colorado, Habitat 1980. variables Site Female winter among sage grouse winter FL a and random b Habitat variable EBRDPWID EBRDPTHT ETINDIS EPLNWID EPLNTHT 68.1 5.0 46 54.5 2.5 46 34.3 2. 9 46 69.2 3.6 46 56.7 4.4 49 48.9 2.6 49 26.1 2.4 49 15.0 2.4 52 33.5 2.2 33 1l.8 1.3 33 sites in North Park, EFSBCC ETSBCC 40.9 3.1 46 33.7 2.8 46 43.3 3.5 46 67.8 3.5 49 35.7 3. 3 49 28.9 2.4 49 37.5 3.2 49 NDc ND 1.8 0.5 52 2.4 0.6 52 FL x SE N Male winter compared FL x SE N Random x SE N - aFL = feeding-loafing bMnemonies e ND = of variables no data. site. described in Table 8. 0 (1'\ �107 Both male and female sage much greater Exposed (~ < 0.001) sagebrush grouse exposed sagebrush clump size averaged sagebrush plant height Sagebrush was not encountered selected winter cover FL sites than at random 4 times greater was 2- 3 times greater with sites. and exposed at grouse FL sites. on 19 of 52 (36.5%) random winter transects. Differences be attributed There in exposed to selection Small differences canopy (~ < of FL sites o. 05) Selection percent between sexes any month in winter sagebrush clump size, may be attributed with lower than random sites during during used by 1980 (Table 16). plant and width, for lower and March. snow cover and March and April. winter Snow depth ( %) by hens. at sites February (~ < 0.05) percent was apparent cannot to preference Table 16. Snow cover and depth at sage grouse random sites in North Park, Colorado, 1980. Snow cover sexes with lower snow depth snow cover by hens of FL sites snow depth during in exposed cover between height (~ > 0.05) in snow depth were no differences male and female grouse sagebrush FLa and (em) Site Feb Mar Apr Feb Mar Apr Male FL 93.8 84.7 71. 4 30.8 15.7 8.5 Female FL 79.9 73.0 69.4 30.5 19.4 8.8 Random NDb 95.8 86.4 ND 37.5 34.5 aFL b ND = = feeding-loafing no data. site. �108 There measured were no differences during and brood ful hens, Percent swnmer FL sites forb cover hybridum) dandelion and percent were highest lower than at sites and differences 17). overlapped (Taraxacum Summer ranges Percent at brood FL sites, used by males. Although there hens with broods might be selecting areas cover than males or unsuccessful hen, unsuccess- could be expected. cover of alsike clover at brood FL sites. officinale) of males, so few differences were not significant, either variables 1980 in meadows at male, unsuccessful (Table and broods (P > 0.05) in 6 habitat (Trifolium cover of common however, was sample sizes were small is some indication with slightly that greater forb during 1979 at hens. Soils Analysis A soil sample analysis male spring values FL sites, than preincubation of 9 soil variables cent organic matter was done on soils collected were compared was higher in soils from preincubation FL sites copper and manganese among sites concentrations incubation FL sites FL or random sites. Higher organic 18). Per- Soils from male and higher (P < 0.05) soils from random sites. (P < 0.05) at pre- at random sites. matter uted to 2,4-D spraying than were also higher than (Table Mean (~ < 0.05) in soils from male FL sites concentrations Manganese (90.0) and 'random sites. also had lower (~ < 0.01) pH, spring subsequent FL sites, in soils from male FL sites of the area around decomposition can be attrib- Raven Lek in 1963 and of dead sagebrush of the soil samples from male FL sites plants. Ninety percent were collected in the �Table 17. Habitat Colorado. 1980. variables Site Brood compared among sage grouse summer Habitat FORBCC TOTCC GRASSCC 93.0 2.2 10 51. 7 8.1 9 41. 3 9.5 9 96.8 0.9 8 63.4 8.6 7 94.0 1.9 19 53.2 6. 1 19 FL a sites variableb TARAXAC in meadows in North Park. TRIFOL GRASSHT 21. 7 9.1 10 18.5 10.4 10 42.5 3.5 10 33.4 9.0 7 13.6 7.0 8 6.3 5.7 8 40.3 2.0 6 40.8 5.8 19 25.1 5.2 19 6.5 4.7 19 39.9 4.2 18 FL x SE N Unsuccessful hen FL x SE N Male FL - x SE N - ~L = feeding-loafin bMnemonics g site. of variables described in Table 8. 0 \.D �Table 18. Soil variables Colorado, 1979. compared Male spring FL grouse spring FL a and random sites in North Park, b pH OM N03 P K Zn Fe Mn Cu 6. 3 0.1 4.0 0.4 4.1 0.4 9.7 1.7 290 37.3 3.5 0.7 35.8 6.6 21. 9 2.4 3.1 0.3 6. 5 0.2 2.9 0.2 3.8 0.5 6.6 1.0 331 79.5 2.5 0.6 36.2 7.3 15.8 2.1 2.3 O. 3 20 x SE Female PIFL sage Soil variables Sample size Site among 13 x SE 0 Random 20 x SE = feeding-loafing bOM = percent organic aFL 6.7 0.1 site, PIFL matter; 2. 9 0.3 = 3.1 0.5 preincubation all soil nutr-ients 7.9 1.4 269 34.5 feeding-loafing in ppm. 2.2 0.5 site. 23.9 4.2 11. 2 1.2 2.3 0.2 �111 sprayed area whereas and 5.0% of random The pattern nutrients higher (Zn, site soil samples matter. Breakdown The organic acids lower soil pH and cations sites under nitrate matter at either ab Ies . measured between the variations cover and plant sage height (r = 0.425, matter attributes 0.589, There were and several habitat vari- P = 0.001, (~= sites. correlation Significant ~= FL sites sites. FL and random clump length of sagebrush at male spring a high positive growth. (~= among male FL, pre- FL or random matter cover proportions 0.616, 0.001, between of ~ = 0.017, N = 59), and N = 60) can be explained by per- matter. No differences between 1969). 99%of the and canopy matter (~ < 0.05) preincubation in sagebrush cent soil organic of organic Structural in organic (1966) reported N = 12), live plant canopy sites. were greater (P > 0.05) matter become in soils from male FL dimensions, found in organic no differences soil organic plant female preincubation Kononova by micro- (Sauchelli since approximately by breakdown clump size, FL, and random and organic matter of 1973). the same pattern incubation conditions area. acids in the soil. of micronutrients concentration to organic and Troeh Sagebrush reflect nitrogen acidic is a result matter of CO2 and organic slightly in soil is provided (Thompson than these is also related nitrogen production sprayed of micro- of organic increases higher t he concentrations bial activity Slightly in Mn, Cu) in soils from male FL sites organic more available FL site soil samples were collected of lower pH and higher Fe, percent only 38.5% of preincubation grouse (P > 0.25) in soil capability use and random sites. classes were found Only poor soils (classes �112 6, 7, 8) occur 1981). in Jackson Approximately soil class sites where 13.8% from soil class Structure differences < 0.001) analysis difference among sites means of viewing A stepwise function habitat 5). criminating This was similar Since a highly the habitat 6 and in subsequent and principal selection seen than of sage grouse from the analysis Seven habitat groups nant functions (DFs) explained were all highly significnat (Table separation components 19). analyses. using in habitat analysis selec- is used. significant Leks sample size (!:_.::. 0.10) dis- The Lst 3 discrimi- 93.6% of the total sample variance (P < 0.0015). habitat was done for 323 vege- of insufficient (Table 20). offer a several use and random sites. provided Percent so all 11 habi- analyses and differences analysis responsible < 0.001) differences multivariate because variables power between (!:_ when univariate function for of variance variables by sage grouse Similarities to test significant analysis variables 0.019) to group discriminant at 7 types were eliminated were on means among 7 types Highly significant (!:_ = simultaneously. plots variables) a univariate to identify tion can be more readily = (11 habitat were included Discriminant 7. (MANOVA) was used in 10 of 11 habitat slope also contributed (N of variance difference. occurred tat variables use sites Analyses was found, (ANOVA) was performed tation of all sage grouse use and random sites. for the observed Dep. Agriculture 7. in multivariate of sage grouse variables (U.S. 86.2 %of the sample was from soil class - Multivariate Multivariate (!:_ 89% (89.4) Colorado 6 with the remainin g 10.6 %on soil class to random Habitat County, and �113 Table 19. Univariate sites in North Park, a F tests Colorado, among sage 1979-80. variableb Habitat grouse use and random F P 2.43 0.019 ATINDIS 19.66 < 0.001 APLNLEN 26.32 < 0.001 APLNWID 27.82 < 0.001 APLNTHT 11. 16 < 0.001 ABRDPLEN 18.62 < 0.001 ABRDPWID 18.12 < 0.001 ABRDPTHT 12.67 < 0.001 AFSBCC 13.88 < 0.001 ATSBCC 12.50 < 0.001 4.90 < 0.001 SLOPE PERFOLCC adf = 7,311. bMnemonics of habitat variables from Table 8. �114 Table 20. Discriminant function analysis of sage random sites in North Park, Colorado, 1979-80. variablea Habitat grouse use and DF 1 DF 2 DF 3 SLOPE -0.201 0.026 -0.257 ATINDIS -0.036 -0.406 0.629 APLNWID -1. 301b 0.227 -0.127 APLNTHT 0.615 1.213 -0.097 ABRDPTHT -0.043 -1. 737 0.085 ATSBCC -0.063 0.315 -0.921 PERFOLCC -0.316 0.250 0.826 Percentage of variance explained by function 66.8 19.6 7.2 Cumulative 66.8 86.4 93.6 <0.0001 <0.0001 percentage P ~nemonics each bUnderlined function. of habitat coefficients variables indicate from Table key habitat 0.0015 8. variables defining �111) The 1st DF was primarily a function (APLNWID, APLNTHT) and explained Although habitat size), each provided power between groups. plant a similar attribute significant (P < 0.001) A high correlation 0.001) between APLNWID and APLNTHT resulted the coefficients and contrasting effects coefficient for APLNTHT (0.615), the contrasting foliation of sagebrush also an important habitat variable distinguishing !: = signs for Since the for APLNWID (-1. 301) was over twice as large Percent discrimi- in opposite on the function. was diminished. of the 0.848, (~= coefficient the size 66.8% of the sample variance. APLNWID and APLNTHT measured (plant nating of sagebrush effect as the of APLNTHT (PERFOLCC) was between sites in 1st DF. The 2nd and 3rd DFs explained of the discriminating tively. power available selection relative The 3rd DF was a function (ATSBCC, to preferred of sagebrush positions of sage respec- of microhabitat macrohabitat cover grouse examined on the 1st 2 DFaxes 3 distinct Sage grouse groups selected to breeding nesting in terms variables, (APLNTHT). characteristics PERFOLCC) and clump size (ATINDIS). Relative rated 19.6 and 7.2% in the 7 habitat The 2nd DF can be understood (ABRDPTHT) spring an additional (Fig. selected FL sites 6). along a gradient large plants season FL or nest hens use and random sites the largest with greater The 1st DF axis sepaof increasing at FL sites sites. plant during winter Among breeding plants. average were plant Although size. relative season males preferred size than broods incubating hens, the relative position of male spring lower than either due to the influence of low percent sites, FL sites or prewas foliation of �116 1 RANDOM. z 0 MALE WINTER FL ~ a w .....J W C/) ~ I- CD « :c 0 e: N Z G.HEN PIFL z ::> u, o l- ~ Ci5 a w 0 a: • BROOD FL a o « o l- :;! z G UNSUCCESSFUL HEN FL 0 z « z a e. FEMALE WINTER· FL a: • NESTS MALE SPRING FL -1 (J) ~'¥ -2 -1 o DISCRIMINANT FUNCTION INCREASING PLANT SIZE Fig. 6. Habitat relationships of sage grouse Lst and 2nd discriminant function axes. 1 1 use and random sites along the �117 sagebrush. Random sites occupied axis since average than plant size and percent at any of the sage grouse The 2nd DFaxis habitat under selection (i.e., represented that at most breeding during to select habitat selection separated The overall to overlap FL sites. sage grouse sites sites selected habitats plants plant in plants be- season. FL sites cover. as most of the sageability However, of sage grouse micro- use sites use from random sites. of sites was low (42.72%) due were frequently use sites (Table misclassified 21). as male of hens and cocks were similar and misclassified. were correctly large at winter among sage grouse Winter FL sites were also frequently the random at all types selection the average the breeding from available classification FL and nest than were more limited in their sage grouse correct in habitat Preincubation spring distinct Micro- Hens selected With snow covering sage grouse was evident selection. sites. was not as evident season FL sites. winter, Lst DF foliation were lower Males also selected loaf during selection microhabitats and further at nest were much larger side which to feed andlor brush microhabitat on the transect). Microhabitat sagebrush on the use sites. was most evident which to nest the area the lowest position Approximately classified, quite distinct 70% (69.6) however, of indicating from that that generally avail- able. The percentage for a given species for the species sites about misclassification of random sites has been used as an estimate (Titus and Mosher 1981). were misclassified 30%of the habitat as sage grouse is suitable of suitable Overall, use sites as use sites 30.4 % of the random suggesting for sage grouse. habitat that only Random sites �Table 21. Classification results of discriminant in North Park, Colorado, 1979- SO. Actual group membership a Male spring FL N Female PIFL function analysis at sage Predicted group membership Nests Brood FL Unsuccessful hen FL grouse use (%) Female winter FL and random Male winter FL sites Random 67 47. Sb 7.5 4.5 4.5 4.5 6.0 11. 9 13.4 Female PIFL 41 22.0 9. S 7.3 19.5 7.3 7.3 2.4 24.4 Nests 17 35.3 5.9 29.4 17.6 5.9 5.9 IS 11. 1 5.6 5.6 55.6 5.6 11. 1 5.6 12 S.3 S.3 33.3 16.7 S.2 2.0 14.3 S.O 6.0 16.0 2.9 8.7 2.9 Male spring Brood FL FL co Unsuccessful hen FL Female winter Male winter FL FL Random Percent of grouped 6.1 49 50 2.0 69 10.1 cases correctly aFL = feeding-loafing bUnderlined values site, = percent 1.4 classified = 42.72% PIFL = preincubation correctly classified feeding-loafing in each group. site. 16.7 16.7 34.7 30.6 4. 1 24.0 40.0 4.0 2.9 1.4 69.6 �119 were most frequently and brood FL sites has more suitable such as nesting Principal examining sites. and Lohnes pretation nents for these FL sites. components analysis relationships that among sage each of which defined 1971). Varimax factor of the principal were readily sample variance (Table grouse mately 59% (59.4) component. explained of sage and explained selection foliation of sagebrush correlated These with the (Cooley to simplify intercompo- 94.1% of the total plant size habitat (APLNLEN, factor separat- use and random sites. component Approxi- was explained was primarily for another by this a function canopy cover (AFSBCC, habitat and slope were the habitat 4th and 5th principal together and The 10.0% of the total sample vari- as the most important last 2 components sample variance. factor (ABRDPLEN, ABRDPWID, APRDPTHT) sagebrush (ATINDIS) variables 11. 5%of the total sample variance. accounted ance and identified clump size grouse of the total sample variance an additional 3rd component habitat The l st 5 principal component identified The 2nd principal of microhabitat use and random was used APLNWID, APLNTHT) as the most important types method of a common habitat rotation habitat 22). The Lst principal ing different (10.1%) requirements another of correlated components. interpretable for other provided (components) FL sites the total available uses than or winter functions were derived, as male spring (8.7%) suggesting areas habitat Linear misclassified factors. factors components, accounted ATSBCC) and Percent most highly r-es pec tiv ely , for 13.2% of the total �120 Table 22. Varimax rotated principal components analysis of sage grouse use and random sites in North Park, Colorado, 1979-80. Habitat a variable I SLOPE 0.047 0.075 0.017 0.001 0.996 ATINDIS 0.574 0.229 0.702 -0.024 0.007 APLNLEN 0.842b 0.338 0.362 -0.005 0.056 APLNWID 0.824 0.332 0.395 0.017 0.039 APLNTHT 0.846 0.376 0.228 -0.032 0.027 ABRDPLEN 0.273 0.894 0.249 0.070 0.065 ABRDPWID 0.314 0.875 0.219 0.117 0.048 ABRDPTHT 0.528 0.723 0.201 0.021 0.051 AFSBCC 0.268 0.251 0.849 0.346 0.024 ATSBCC 0.310 0.238 0.902 0.011 0.015 -0.038 0.096 0.127· 0.985 -0.000 PERFOLCC Principal II components III IV Percentage explained of variance by function 59.4 11. 5 10.0 Cumulative percentage 59.4 70.9 80. 9 ~nemonics the bUnderlined component. of habitat coefficients variables indicate from Table key habitat V 8.5 4.7 89.4 94.1 8. variables defining �121 Relative plotted axis positions of sage grouse on the Lst 3 principal (plant size) ing season separated component axes from all sage grouse of sage grouse using principal were distinguished differences of sagebrush sites. cessful probably separation separated between during season 3 distinct components analysis. on the basis different selection, selected differences groups secondarily from breeding habitats Nest sites in habitat However, with larger by and finally on the basis selected season which were distinct were the most distinct selection relative hen and br-ood FL sites in 3-dimensional be somewhat different size, were Male and female sage grouse all seasons. sites. groups Furthermore, of plant which were distinct Sage grouse season site but among all breeding breed- The 3rd axis (canopy analysis, cover characteristics. from random sites breeding additional of microhabitat similar winter FL habitats FL and nest use sites. function primarily in degree The 1st use sites. As with discriminant identified 7). The 2nd axis (microhabitat) cover and clump size) provided types (Fig. winter FL sites from all sage grouse and random sites. random sites use and random sites were were evident positions habitat sample sizes. of unsuc- space would �NESTS HIGH CANOPY COVER LARGE CLUMPS MALE SPRING FL I- Z w z o a.. ~ o o ~1cf WINTER FL CANOPY COVER AND CLUMP SIZE UNSUCCESSFUL HEN FL • . -- §t~ - T §J-fl »11 DISTINCT I a.. ~ ~ rS ~~ a.. ~ § ~ ~ If LOW CANOPY COVER SMALL CLUMPS ~ ~ ~ ~ ,.Q 0 .J rJ ....., ~ ~ (} ~ ~ & ~ LARGE SMALL PLANT SIZE PRINCIPAL COMPONENT Fig. 7. Habitat relationships component axes. I among sage grouse use and random sites on the first 3 principal N N �123 DISCUSSION Radiotelemetry Tail-clip There transmitters was no apparent ments or behavior, birds. even interference heavier and laid a fertile this potential clutch the best be solar-powered during high predation transmitter loss of the radio package, are much greater with tail-clip during apparent difficulty for future models, than will used distance. radio-collars The for long- for battery-powered radios. the most common problem could be alleviated Whether or not radio-collars the breeding studies radio-collar receiving of solar-powered Premature male displays without line-of-sight collection radio-collars. 1. 39% She moved approximately package term data powered equaling The solar-powered had excellent life and usefulness encountered carried of 7 eggs. radio-collars. study a WMtransmitter range move- of radio-marked were easily 4 months. to breeding well overall. with daily or seasonal Hen 5032 carried 32 km from winter 1979-80 worked WMtransmitters of her body weight for at least Perhaps during nor was there The slightly by hens. used period using solar- interfere with is unknown. Movements Movements by sage heavy grouse snowfall and subsequent to winter ranges lack of available in response sagebrush to in breeding �124 and summer ranges et al. has been well documented 1963, Beck 1977). depending Distances on how far sage grouse above snow. Dalke et al. traveled, range tances traveled mild winter. concentrated in Idaho. during (1963) reported a severe (1975) during sage grouse northeast quadrat the winters intensive grouse population during precipitation important quadrat averaged all grouse was available shorter during above snow. in the and that the northeast to the entire sage 7 high use sagebrush essential winter the highest rangequadrat habitat even winter use in of 1973-74 and 1974-75 when daily movements and size of winter were restricted by Beck use area in the northeast the winters winter winter. on habitat 26% above and 26%below normal, for males and females during a sage grouse Beck (1975) recognized since it provides during during only 3.7% of the sagebrush Beck (1975) recorded Similar average ing that depend its importance The intensive mild winters. the northeast underlines dis- The fact that only 6.8% of the total available land in North Park. is especially of the winter within the Park. encompassing that of sagebrush of North Park to average used in 1980 were outlined use area comprises in North Park greater of 1973-74 and 1974-75. at least part habitat areas areas from all quadrats quadrat and summer range (1972) reported stands cover flocks travel- winter in North Park than with dense Winter concentration have varied sage grouse Beck (1977) recorded Eng and Schladweiler in areas however, 1952, Dalke have to go to find suitable ing up to 50 miles (80.5 km) from breeding winter (Patterson respectively. range 1980 might be expected to limited areas Eng and Schladweiler daily movements and smaller winter ranges areas consider- where sagebrush (1972) recorded for hens in Montana �125 than was found in North Park. Differences between Colorado may be due to use of 2 widely separated ranges in North Park in 1980 vs. only 1 winter Montana and (12 km) winter range area in Montana. Daily movements by males from leks to FL sites those reported Park. by Emmons (1980) in the Lake John area of North Emmons found that km of the lek compared Schladweiler 90.1% of all daily movements to 95.7% in this (1974) reported that of 1. 8 km from leks to FL sites Nonrandom dispersal by Emmons (1980). to selection features species habitat may also influence of sagebrush appeared study. male sage were within 2.\0 Wallestad and grouse moved a maximum in Montana. from leks has been documented Nonrandom of preferred on the basis were similar to dispersal around structure can most likely be attributed leks although movements. Selection factor topographic of distinct FL habitat Preferred sagebrush was evident. to be an important previously determining dispersal direc- tion from leks to FL sites. There tances is considerable reported variability in the literature. from leks where bred to nest movements were observed and Pyrah (1974) in Montana. average lek-to-nest also observed hens longer in average Average sites by Poley distance (1969) in North Park Petersen (1980), average of 4.4 km from leks to nests. Variability in availability of suitable nesting May (l'91;()) 4 juvenile moved an average in lek-to-nest North Park may be due to small sample sizes but movements probably habitat Compar abde reported Park. hens by hens and Wallestad however, movements in North Park; of 8.2 km while 4 adult dis- traveled in 1979-80 was 2.7 krn , movements of 4.0 km in North moved an average differences lek-to-nest in also reflects close to leks. �126 There is also disagreement ments regarding and Petersen juvenile between (1980) documented hens to select nest juveniles traveling and Pyrah juveniles moving just to quality selected in nesting within summer ranges period of the year. abundant sites. herbaceous During the lek attended with (2.5 km). selected. Adult travel hens suggesting cover nearby. were typically or nest site. than during concentrated dis- 1979-80, adult hens feeding were more restricted vegetation Wallestad and therefore to summer range Sage grouse reported in lek-to-nest habitat habitat nest sites with better the meadow area nearest than age classes site more often than juvenile Movements from breeding This study (2.8 km) than adults age classes move- farther to nest. between of nesting to find suitable fed closer to the nest adults farther between hens may be more selective that of hens. May (1970), however, differences slightly differences distances age classes almost twice as far as adults may be related longer of lek-to-nest adult hens traveling sites. (1974) noted little Observed tances differences among studies to Movements any other in meadow areas so movements to find suitable with FL habita ts were minimal. The only observed occurred during difference the breeding in movement patterns season. Males traveled leks to FL sites than hens moved from nests the hypothesis rejected of differential for winter and summer periods farther to FL sites. movement patterns between between but was accepted sexes from Therefore, sexes was for breeding season movements. Differential were evident movement patterns for. both sexes. between All radio-marked seasons birds within sex class underwent long �127 migrations to wintering were generally restricted Given a mild winter be expected. little areas whereas to movements to the nearest with little snowfall, Before the blizzard snowfall and no observed east quadrat from other areas ments and home ranges the breeding migration to FL areas leks. around ments between seasons their Sage grouse and aspect as that open areas were preferred. winter, encounter FL site periods preferred ridges for west- sage grouse during breeding season movements except move- sagebrush Perhaps more frequently to sampling bias for areas for lek sites 1964, Rotlienmaier differences in topographic of the year, sagebrush . slope at leks where flat, open areas to southeast-facing kept 1980. similar average however. draws and swales or open slopes. Windswept ridges III than of differential 1952, Rogers were distinct different wind and solar irradiation used winter daily move- and males restricted with fairly (Patterson sage grouse winter. had been into the north- were greater Selection of flat, more often than windswept throughout would Selection habitats There during nized a preference 1980, there Hens restricted found at random sites has long been recognized selected migration within sex class was accepted. selected 1979, Dingman 1980). little Average the hypothesis Habitat During winter nests Thus meadow area. of sage grouse of North Park. season for both sexes. around however, in late January used during movements to areas features movements to summer areas Beck (1977) reoog- slopes >5%where virtually free of snow were the 2nd most frequently the reason Beck in sagebrush where sage grouse (1977) did not draws was due could be readily observed. �128 Without radio-marked heavy sagebrush birds, it is difficult to locate sage grouse cover in draws. Use of 0-5 and 6-10% slopes by sage grouse season was fairly however, were used more frequently in the random sample. where herbaceous vegetation forb cover by broods Broods especially was abundant. has frequently grouse use sites a milder winter in structural with winter habitat and Schladweiler (1972) reported at female winter FL sites canopy (1972) during Exposed in 1980 was greater heavy snowfall. In Montana, Eng canopy cover FL sites Sage- FL sites in North Park in by Eng and Schladweiler above the snow at winter found by Beck (1977) during Beck sampled flocks visible may have overlooked would in Montana. of sagebrush than FL sites probably Given (1974) in the same area. than reported milder winters height the most distinct. similar mean sagebrush at winter the at sage (28 %) as was found at male spring cover and height 1980 were much greater in North Park were evident season FL srt es, (32%) by Wallestad and Schladweiler brush draws 1970~, Wallestad 1971). selection winter have been more similar to breeding preferred 1980) and throughout FL sites being in 1980, however, area. than they been encountered 1952, Klebenow 1969, Peterson Differences the breeding Selection for abundant (Gill 1965, Poley 1969, May 1970, Petersen West (Patterson during similar to a random sample of the study Draws and swales, occurred in FL sites a winter with from a snowmobile and flocks which were concealed in heavier sagebrush cover. The only observed tats was in exposed difference sagebrush between height sexes in winter above the snow. FL habi- Hens selected �129 areas with greater found that during exposed plant a winter with heavy flocks chose areas with greater male flocks but that tion of sagebrush in selection but hens there height of exposed selected both winters sagebrush sagebrush were obvious sites. classes Nesting rately preferred hens sought selected by hens prior suggest that Structural to incubation hens continue to select as great as that encountered at preincubation Nest sites had greater sagebrush by Klebenow (1969) in Idaho. Average pared canopy cover ranged to 44%in North Park plant over the nest nest were more comparable cover may be partly in FL habitats the brooding period throughout the was over twice FL sites. cover than previously Martin (1970) and Wallestad 1979-80. to previous FL sites. hens may not accu- (1980) in North Park. from 18.4 to 27%in these and mean sagebrush canopy at unsuccessful however. canopy and Petersen during cover of sagebrush similar habitats Forb cover at brood sites. (1974) in Montana. average and male spring and during season. and Pyrah better similarities breeding reported among breeding Cover attributes nests cover in 1980. a wide range between sexes may not be real and similarities were used. cover. between structural the small sample size for unsuccessful reflect sexes in selec- Differences and again in winter hen FL sites were intermediate However. between better female than predominantly and density with slightly differences Beck (1977) predominantly density height than was found at FL .s'ites although cover and height snowfall. above the snow. FL areas season FL and nest than males. was no difference of Beck+s study There height Average height studies. studies height com- of the in the area around Differences due to small sample sizes for nest sites the In canopy (N = 19) �130 but it is unlikely canopy cover. North Park including that this would account Differences in canopy for a 20% difference cover at nest and Idaho and Montana may be related soils. climate. Sagebrush canopy and species / subspecies sites in between to other factors composition of sage- brush. has been encountered (Klebenow 1971) . sites in North Park However, in North Park. Average (1970), FL sites sagebrush of North Peterson canopy Montana rather however, grouse at brood FL in Idaho and in sagebrush was forb cover of 22- 33% (1970b), and Wallestad summer ranges (1971) in in meadows to 41.3%. reported (1974) found in previous 32%canopy FL studies. cover at breeding cover at male spring FL sites 28% in the Lake John area Park. at male spring tural in Idaho of males in Montana and Emmons (1980) reported canopy than 1970~. Wallestad height to brood ranges ranges cover of 35.0% at male spring than that Wallestad and Schladweiler not, sagebrush reached forb cover increased Wallestad and Schladweiler on brood at brood FL sites once broods sagebrush was also greater 1970. Peterson lower than average was somewhat higher season average forb cover by Martin Montana. sites. FL sites similar periods was comparable Average reported at brood 1969) and Montana (Martin only 6.9%. markedly sites during As with nest Montana. cover FL sites than attributes use sites. was representative a result provide (1974) stated of selection data to support of the habitat as was true that of the study by sage this habitat height area in grouse. statement. are similar between for several sagebrush They did Even if struc- random and sage variables measured �131 at preincubation indication that and brood FL sites in North Park, sage grouse simply be selecting random sites. with higher habitat habitats Other For example, differences and preincubating percent habitats. with similar structural habitat broods selection are not selecting that is not an attributes as may be recognized, however. hens both chose sagebrush foliation than random sites. was evident They may In addition, at all sage grouse micro- use sites compared to random sites. Differences grouse In habitat use sites and between easier to identify neously using discriminant analyses It was evident that fashion and that between that actual sage grouse selected distinct habitats using and nest habitat habitats analyses. to be more similar univariate were selected simulta- components season sites indicated the most valuable was identification best between sage grouse plant and principal FL sites of sage sites analysis, relationships differed. in a nonrandom within as well as seasons. Perhaps analyses breeding types were analyzed and brood FL sites appeared to be similar to winter multivariate different use and random sites were variables function to random sites than other appeared between sage grouse when all habitat Whereas preincubation habitat selection different contribution of habitat types variable components of sage grouse use and random sites. which discriminated of the multivariate use sites and between Sagebrush habitats. clump size were important Finally, variables plant best between dimensions were the 2nd most important of FL and nesting which distinguish sagebrush separating size was the key sites. Microhabitat component in selection canopy cover and sites. Whereas canopy �132 cover has long been deemed of primary importance in distinguishing between and winter habi- tats, sage grouse breeding, plant grouse brooding, dimensions were the most important habitat selection. cover is also important select nesting, habitats cover classes component in sage This does not discount in sage grouse habitat on the basis of both suitable the fact that canopy selection. plant size and canopy of sagebrush. Selection of seasonal habitats canopy cover may be understood with distinct plant dimensions and in terms of cover needs and behavior. During the heavy snowfall winter of 1980, large plants sought since they were available despite hens chose areas from predators selected where relatively and good feeding areas with larger breeding plants season due to larger cealment since they solitary prior large plants cover nearby. concealment Males may have than hens at FL sites during the body size and a need for better in habitats con- hens are used by sage grouse and random sites to selection by grouse what was generally for better structural available and may also be related species and subspecies for Artemisia tridentata of sagebrush sage grouse prefer tridentata vaseyana A. tridentata cover than to preference in the diet. has long been recognized cates that and recent for Preference work indi- wyomingensis over A. (Remington 1981). Similarities in habitat selection between summer) and differences in rejection provided Nesting to incubation. can be attributed (winter, were often deep snow cover. are usually found in flocks whereas The differences different Sage grouse of the hypothesis that during sexes within 2 seasons the breeding sage grouse habitat season resulted selection differs �133 between sexes within seasons acceptance grouse during habitat the breeding selection since FL habitats varies selected from each other. and spring FL habitats seasonally probably large-scale tat. but continued reduce strip quadrat. Park depend on winter of the winter, entire sage grouse and summer were the differences mining activity Approximately on winter eliminate preferred habitat bisect mine daily to reach ferred FL habitat in of North at least part will impact the quadrat may be male segment of the population. of the preferred FL sites of males from Kerr coal lease. FL sites would be indirectly the preferred winter range will in North Park. Raven Lek were within the proposed of preferred habi- in 1980 lease areas during from mining activity winter range from all areas in the northeast to the breeding 33% (33.3) had area of to sage grouse and proposed Since sage grouse population winter 1980. Immediate impacts of mining in the northeast most detrimental between mining in the northeastern loss of habitat sage Impacts of Mining eventually the northeast that would not have been as great mining on current and perhaps spring, to be detrimental Sage grouse tolerated but for each sex was accepted winter, winter Potential North Park would appear The hypothesis However, been less snow during Long-term, and summer periods season. during all distinct there for winter affected Another 12.5% since the mine will FL area and males would have to fly over the these areas. would affect Disturbance of 45.8% of the pre- all males since they all regularly �134 fed in the proposed the entire area unsuitable How great extent lease area. for breeding to preferred a 63% decline in strutting adjacent of strutting in areas sagebrush Braun males 2 years after by 2,4-D spraying adjacent to leks (Rogers a 31%loss of suitable declines in the number of leks have been documented and mechanical treatments of 1964, Higby 1969, Peterson 1970a. and Beck 1976). Immediate impacts of mining on hens during would probably be less detrimental not as dependent on areas of leks where they bred 2 km of the lek , also noted to Raven Lek for nesting 95.7% of males sought Wallestad and Schladweiler of male spring hens often move long distances Coal strip since they are not currently habitat along travel Loss of draws abundant routes and other insects within 2 km FL sites within to leks whereas from leks to find suitable mining is not expected habitat (1974) and Emmons (1980) FL habitat (May 1969. Poley 1970. Wallestad and Pyrah season Hens were Only 64%of all hens nested whereas close association the breeding than impacts on males. adjacent as males were for FL habitats. tats on the Wallestad (1975) reported Substantial males and total abandonment disturbed may make males will be may depend FL habitat. to a lek in Montana. FL areas season use. the impact to breeding of disturbance habitat Loss of preferred nesting 1974. Petersen cover 1980). to impact summer meadow habi- within proposed mining areas. Brood to meadows will be impacted by mining. wet areas with lush herbaceous would be detrimental to broods vegetation dependent on such areas. It is unknown make the areas whether adjacent large-scale. long-term to mines unsuitable coal mining will for use by sage grouse and �135 during any season. It is apparent that areas mining will be removed from sage grouse cover returns. probably be at least several will reduce activity and large-scale recruitment of hens on the lek. disturbed expected eastern decades. available habitat habitat use until suitable to greatly sagebrush for all grouse males. Long-term, to a lek will probably large-scale winter habitats Winter habitat throughout FL Mining may be eliminated altogether preferred portion of the Park. for breeding of preferred grouse to the lek and breeding Production reduce Disturbance loss adjacent of juvenile by mining activity. ally critical by strip How long that period will be is unknown but will habitats reduce disturbed in areas mining can be in the north- in the northeast North Park. activity is especi- �136 RECOMMENDA TIONS FOR MITIGATION AND REHABILITATION Sage grouse select habitats on the basis species and subspecies will move elsewhere to disturbance winter, of suitable and brood-rearing and probably developed suitable mitigation If populations and rehabilitation year-round techniques and mitigating sagebrush It cannot be assumed habitats. mitigation methods of reducing nesting, and maintain the same populations are to be maintained, Several structure composition. of preferred to provide breeding, habitats that present grouse prior of sage grouse practices must be for sage grouse. should be considered. Among impacts of mining are: 1. Maintain or protect preferred habitats 2. Limit disturbance adjacent to and on winter areas to be impacted by mining. 3. Limit disturbance adjacent to leks and on preferred feedingloafing (FL) areas used by males around leks. Avoid road construction and placement of overburden piles adjacent to leks, preferred FL areas, and in flight paths of males moving from the lek to FL sites. 4. Curtail explosions during the mating period (1 hour before to 1 hour after sunrise) from 15 March to 1 June. 5. Reduce or eliminate grazing 6. Fertilization of undisturbed preferred habitat and areas adjacent to coal mines may be useful but needs further documentation. 7. Obtain financial support from coal companies to monitor sage grouse movements and habitat use prior to and throughout the mining period and to develop better techniq ues to re-establish the sagebrush community on reclaimed areas. in areas where possible. around concentration leks. �137 Sage grouse year. Therefore, trate on restoring require a diversity rehabilitation the diverse Possible rehabilitation of habitat of sage grouse habitat techniques structure types habitats present throughout the must concenbefore mining. include: 1. Create topographic diversity in habitat. Flat, open areas «10% slope) are used extensively during the breeding season whereas draws and swales with high sagebrush canopy cover and large plants are important in winters with heavy snowfall. Windswept south-facing ridges and hilltops are also important in winter. Draws with lush herbaceous growth are important for broods in early summer and are also used by unsuccessful hens and cocks. 2. Transplant and/or seed native grasses, forbs, and especially sagebrush. Special consideration should be given to species and subspecies of sagebrush preferred by sage grouse. Big sagebrush (Artemisia tridentata) was preferred over alkali sagebrush (A. longiloba) in the study area. Wyoming big sagebrush (A. !._. wyomingensis) is preferred over mountain big sagebrush (A. t. vaseyana) (Remington 1981). 3. Transplant and/or seed sagebrush throughout reclaimed areas and create "patc hy " areas with dense stands of sagebrush in draws and swales where greater moisture can support better sagebrush cover. Sa~brush density (average) should be at least 3 plants/m . . 4. Fertilization sagebrush, lished. 5. Irrigate reclaimed areas to provide ample moisture during the growing season and build snow-fencing to hold snow on reclaimed areas for additional early spring moisture. 6. Strive to create a diversity in sagebrush structural types to meet sage grouse habitat requirements during all seasons. Preferred FL habitats are those between 25 and 50% average sagebrush canopy cover and 25 to 40 cm sagebrush height. Large plants and high canopy cover are preferred at FL sites during winters with heavy snowfall. Nesting hens also prefer excellent cover and larger plants. Smaller plants and lower canopy cover are preferred at FL sites during the breeding season and low canopy cover 01%) and sagebrush height (10 cm) are found at leks. 7. Provide vigorous stands of sagebrush foliation of sagebrush plants. of reclaimed areas forbs, and grasses should be done annually have become well estab- with at least 75% until �Sage grouse select and within seasons. distinct All of these maintain stable populations out the West. Mining activity abundance and careful requirements how severe scale, maintained, consideration different seasons must be managed properly sage grouse distribution and should be given to all habitat as mining proceeds. While it is not known populations, to be detrimental. with coal and, large- The entire depending over which the area is mined and the success reduced, to in North Park and through- of North Park is underlain and rehabilitation re-establishing nity will affect mining can be expected quadrat Recently, populations habitats during impacts of mining will be to sage grouse on the time interval mitigation types of sage grouse of sage grouse long-term northeast habitat practices, sage grouse populations of will be or lost from the area. there has been increasing sagebrush but little success communities on reclaimed mine spoils. of sage grouse are to be maintained methods of rehabilitation interest and other members of the sagebrush in mining areas throughout must be developed in If commu- the West, better to re-establish sagebrush. 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Winter nutrition of sage grouse in North Park, Colorado. Proc. Bien. West. States Sage Grouse Workshop 12: 10. Richards, L. A., ed. and alkali soils. 1954. Diagnosis and improvement of saline U.S. Dep. Agric. Handb. 60. 160pp. Robertson, D. R., J. R. Nielson, and N. H. Bare. 1966. Vegetation and soils of alkali sagebrush and adjacent big sagebrush ranges in North Park, Colorado. J. Range Manage. 19: 17-20. Rogers, G. E. 1964. Sage grouse investigations in Colorado. Colorado Game, Fish and Parks Dep., Tech. Bull. 16. 132pp. Rothenmaier, D. 1979. Sage grouse reproductive ecology: breeding season movements, strutting ground attendance and site characteristics, and nesting. M.S. Thesis. Univ. Wyoming, Laramie. 97pp. Sauchelli, V. 1969. Trace elements in agriculture. Reinhold Co., New York, N. Y. 248pp. Van Nostrand Schneegas, E. R. 1967. Sage grouse and sagebrush control. Trans. North Am. Wildl. and Nat. Resour. Con£. 32: 270-274. Scott, T. G., and C. H. Wasser. plants for wildlife biologists. D.C. 58pp. 1980. Checklist of North American The Wildl. Soc., Washington, Slat ick , E. R. 1980. Coal data: A reference. Energy U.S. Dep. Energy, Washington, D.C. 53pp. Smith, InL Adrmn , , E. L. 1966. Soil vegetation relationships of some Artemisia types in North Park, Colorado. Ph. D. Diss. Colorado State Univ., Fort Collins. 203pp. Soltanpour, P. N., and A. P. Schwab. 1977. A new soil test for simultaneous extraction of macro- and micronutrients in alkaline soils. Commun. in Soil Sci. and Plant Anal. 8:195-207. ----:- , S. N. Workman, and A. P. Schwab. 1979. Use of inductively-coupled plasma spectrometry for the simultaneous determination of macro- and micronutrients in NH4HC03-DTPA extracts of soils. Soil Sci. Soc. Am. J. 43: 75-78. Terwilliger, C., Jr., in North Park, Dep. Sci. Ser. and E. L. Smith. 1978. Colorado. Colorado State 32. 48pp. Range resource types Univ .• Range Science �143 Titus, K., and J. A. Mosher. 1981. Nest-site habitat by woodland hawks in the central Appalachians. 270-281. selected Auk 98: Thompson, L. M., and F. R. Troeh. 1973. Soils and soil fertility. McGraw-Hill Book Co., New York, N. Y. 495pp. U.S. Department of Agriculture. 1981. Soil survey of Jackson County Area, Colorado. U.S. Dep. Agr-ic , , Soil Conserv. Serv., Washington, D. C. 159pp. U.S. Department of Commerce. 1979. Climatological data: annual summary, Colorado. Natl. Oceanic and Atmospheric Admin. 84( 14). 14pp. Wallestad, R. O. sage grouse 35: 129-136. 1971. Summer movements and habitat use by broods in central Montana. J. Wildl. Manage. 1975. Male sage grouse response J. Wild!. Manage. 39: 482-484. , and D. Pyrah. 1974. hens in central Montana. -----,::- to sagebrush treatment. Movement and nesting of sage J. Wildl. Manage. 38: 630-633. grouse ---- , and P. Schladweiler. and habitat 38: 634-637. , J. ----sage selection 1974. Breedin g season movements of male sage grouse. J. Wild!. Manage. G. Peterson, and R. L. Eng. grouse in central Montana. J. 1975. Foods of adult Wildl. Manage. 39: 628-630. West, P. W., and T. P. Ramachandran. 1966. Spectrophotometric determination of nitrate using chromotropic acid. Anal. Chimica Acta. 350: 317-324. �144 APPENDIX Plant species 1979-80. Scientific identified name in the study area, North Park, Colorado, b Common name ° a SHRUBS AND UNDERSHRUBS Artemisia cana viscid ula A. frigida-A. longiloba A. triden tata Chrysothamnus nauseosus C. viscidiflorus Eurotia lanata Gutierrezia sarothrae Purshia tridentata Ribes montigenum Rosa ar kansana Salix sp. SarCobatus vermiculatus Symphoricarpos vaccinioides Tetradymia canescens silver sagebrush fringed sagebrush alkali sagebrush big sagebrush rubber °rabbitbrush Douglas rabbit brush common win terf at broom snakeweed antelope bitterbrush gooseberry currant Arkansas rose willow bl ack greasewood whortleleaf snow berry gray horsebrush FORBS Achillea lanulosa Androsace septentrionalis Antennaria microphylla Aquilegia caerulea Arenaria congesta Arnica mollis Aster leucanthemifolius Astragalus agrestis A. drummondii A. flexuosus A. gracilis A. kentrophyta A. pectinatus A. spatulatus Atriplex rosea Berberis repens Calochortus gunnisonii Cardamine pennsylvanica western yarrow py gmyflower rockjasmine littleleaf pussy toes Colorado columbine ballhead sandwort hairy arnica daisyleaf aster purple milkvetch Drummond milkvetch flexile milkvetch slender milkvetch Nuttall kentrophyta milkvetch narrowleaf poisonvetch spoonleaf milkvetch tumbling orach creeping barberry Gunnison mariposalily Pennsylvania bittercress �Appendix-Contin Scientific name ued a Castilleja £lava C. integra Chenopodi um sp , Cirsium centaureae Clematis hirsutissima Cleome serrulata Comandra urnbellata Cordylanthus ramosus Cryptantha fendleri C. virgata Dodecatheon pulchellum Epilobium angustifoliurn ~. paniculatum Erigeron nematophyllus Eriogonum ovalifolium E. subalpinum E. umbellatum Erysimum inconspicuum Fragaria americana Gayophytum ramosissimurn Gentiana affinis Geum ciliatum Gilia congesta Gymnosteris parvula Heuchera p arvifolia , Hymenoxys richardsonii Iris missouriensis Lappula redowskii Lepidium ramosissimum Leptodactylon pun gens Lesquerella montana Linanthus harknessll Linaria vulgaris Linum lewisii Lithophragma ten ella Lupinus ammophilus !:_. greenei Lygodesmia juncea Mammillaria vivipara Melilotus officinalis Mertensia humilis Monolepsis nuttalliana Oenothera caespitosa montana ODuntia polyacantha Orthocarpus luteus Oxytropis sericea Paronychia sessiliflora Common name b yellow indianpaintbrush wholeleaf indianpaintbrush goosefoot fringed thistle hairy clematis Rockymountain beeplan t common bastardtoadflax bushy birdbeak Fendler cryptantha miner's candle darkthroat shootingstar fireweed willowherb autumn willowherb mat fleabane cushion wildbuckwheat subalpine wildbuckwheat sulfur wildbuckwheat smallflower wallflower American straw berry branchy ground smoke Rockymountain pleated gentian threeflowered avens ballhead giiia leafless falsephlox littleleaf alumroot Colorado rubberweed Rockymountain iris bluebur stickseed branched pepperweed granite pricklygilia mountain bladderpod Harkness flaxflower butter-and-eggs toad£lax Lewis flax slender woodlandstar sand lupine Greene lupine rush skeletonplant cushion ballcactus yellow sweet clover bluebells Nuttall monolepsis tufted eveningprimrose plains pricklypear yellow owlclover silky locoweed creeping nailwort �146 Appendix-Contin Scientific ued namea Common name Pedicularis crenulata P. groenlandica Penstemon cyathophorus ~. procerus P. strictus Petasites sagittata Phlox bryoides P. multiflora Polygonurn aviculare P. kelloggii Potentilla anserina P. concinna P. diversifolia Pulsatilla ludoviciana Ranunculus glaberrimus Rurnex tz-iang uliv alv is . Salsola kali Saxifraga rhomboidea Sed urn stenopetalum Senecio harbourii S. hydrophilus S. integerrimus Sidalcea neomexicana Sphaeralcea coccinea Taraxacum officinale Thlaspi alpestre Trifolium gymnocarpon T. hybridurn Viola nuttallii ellipticus elliptic us b meadow lousewort elephanthead lousewort Northpark penstemon littleflower penstemon Rockymountain penstemon arrow leaf coltsfoot sq uarestem phlox flowery phlox prostrate knot weed Kellogg knotweed sil v erw eed cinq ue foil elegant cinquefoil varileaf cinquefoil American pasq ueflower sagebrush buttercup Mexican dock common Russianthistle diamondleaf saxifrage wormleaf stonecrop Harbour groundsel water groundsel lambstongue groundsel Newmexican checkermallow scarlet globemallow common dandelion alpine pennycress hollyleaf clover alsike clover yellow prairie violet GRASSES Agropyron dasystachyum A. spicatum Deschampsia caespitosa Hordeum brachyantherum H. jubatum Koeleria cristata Phleurn pratense Poa pratensis P. secunda aScientific names follow Harrington thickspike wheatgrass bluebunch wheatgrass tufted hairgrass meadow barley foxtail bar ley prairie junegrass common timothy Kentucky bluegrass Sandberg bluegrass (1954). bCorrimon names follow Kelsey and Dayton and Scott and Wasser (1980). (1942). Beetle (970), �147 Approved by _~!W~=-='--::7:--. ~~--=-=Clait E. Braun Wildlife Research Leader _ ��149 JOB PROGRESS REPORT State of Colorado ------------------------- Project No. Work Plan No. Job Title: 3 Job No. 13 Responses of Sa~e Grouse to Vegetation Fertilization Period Covered: Personnel: Game Bird Survey W-37-R-35 1 January 1981 through 31 May 1982 R. A. Ryder, Colorado State University; Kevin Berner, Clait Braun, Len Carpenter, Kurt Hundgen, Steve Porter, Tom Remington, Joan Ritchie, Tom Schoenberg, Lynn Stevens, John Wagner, Carol Ann Weinland, Colorado Division of Wildlife. ABSTRACT A study of sage grouse (Centrocercus urophasianus) winter food preference and nutrition was initiated in North Park, Colorado in 1981 and continued in 1982. Sage grouse populations were monitored in 2 study areas and throughout North Park by banding, lek counts, pellet transects, and collection of harvest data. Sagebrush (Artemisia spp.) at feeding and random sites was similar (p > 0.05) in average height (23.56 vs. 24.07 cm, respectively) and percent fol iation (76.88 vs. 73.11%, respectively), but feeding sites had greater (p < 0.05) sagebrush canopy cover (24.20 vs. 19.46%, respectively). Sage grouse selected (P < 0.05) feeding sites with proportionally more Artemisia tridentata wyomi~gensis (ATW) and less A. tridentata vaseyana (ATV) and~. longiloba (AL) than at random sites. ATW comprised 48% of the sagebrush at random sites, 86% of the sagebrush at feeding sites, but 90% of the plants identified as fed upon. Selection (p < 0.10) for food plants with larger or smaller values of height or percent foliation occurred on 52% of the feeding site transects in 1981. ATV contained less (p < 0.05) crude protein (10.64%) than either ATW (14.23%) or AL (15.15%). Crude protein content of ATW from feeding sites (16.42%) was higher (p < 0.05) than ATW samples from random sites (14.23~). Crude protein content-of fed-upon ATW (17.22%) was higher (p < 0.05) than non-fed upon ATW (15.59%). Leaf samples from sage grouse crops and gizzards averaged 15.87 and 13.62% crude protein, respectively. Crops and gizzards differed (p < 0.05) in concentrations of 10 of 13 nutrients assayed. Winter fat-content of adult and juvenile males and females averaged 3.54, 1.20, 2.91, and 1.84%, respectively. A total of 402 sage grouse was trapped and newly banded in 1982. High male counts at 9 leks in the study area totaled 236 and averaged 26/1ek. Hens selected nest sites with higher (p < 0.05) average sagebrush height (43.75 cm) and canopy cover (39.9%) than at random sites. Clutch size in 1981 averaged 7.2, 5.7, and 5.0 eggs for adult 1st and 2nd,and yearl ing 1st clutches, respectively. Pellet �150 transects in the northeast averaged 0.27, 0.33, and 0.12 droppings/day and in the northwest 0.04, 0.06~ and 0.23 after winter 1980-81 and 1981-82 and summer 1981, respectively. Hunters averaged 1.3 birds/hunter in 1981. The northeast quadrat sustained 11.5% of the hunting effort and 10.5% of the harvest. Immatures comprised 47.4% of the harvest, yearlings 22.3%, and adults 30.4%. Apparent nesting success in 1981 based on wing molt patterns was the lowest on record (31.2%). Young/hen (1.3:1) and young/ successful hen (4.1:1) ratios indicated nesting success was only slightly below average in 1981. �151 RESPONSES OF SAGE GROUSE TO VEGETATION FERTILIZATION Thomas E. Remington Coal resources in the western United States are and will become even more important sources of energy derived from fossil fuels. Much of the increased demand for coal will be met by surface mining, an economically attractive method of extraction. Much of the surface-mineable coal in the western states is thinly overlaid with overburden, the top which supports primarily sagebrush-dominated grasslands. Because of state and federal statutes, mining companies are required to revegetate disturbed ar.eas and to mitigate wildlife habitat loss (Surface Mining Control Act, U.S. Congress 1977; Fish and Wildlife Coordination Act of 1934 and amendments). The technology to revegetate disturbed areas with sagebrush is currently lacking. Re-estab1ishment of sagebrush communities by succession is a long term process. Therefore, it is evident that much of the habitat for obligates of the sagebrush community will be lost for extended periods. The need for a suitable mitigation technique for this habitat loss is obvious. The primary dependence of sage grouse upon sagebrush for forage (Rasmussen and Griner 1938, Dargan et a1. 1942, Patterson 1952, Pyrah 1969, Wa1lestad et a1. 1975) and nesting (Keller et a1. 1941, K1ebenow 1969, Autenreith et a1. 1982, Petersen 1980) makes the sage grouse an ideal candidate species to use in evaluating mitigation procedures in sagebrush-dominated grasslands. Fertilization holds promise as a mitigation technique as several studies have lnd i ca t ed animal (grouse) use, productivity, or numbers were increased by fertilization of their food resource (Miller et al. 1970; Watson and OIHare 1973, 1979; \·Jatsonet a1. 1977; Seckerton 1980). A knowledge of pretreatment nutrition and reproductive parameters is essential to evaluate the effects of fertilization on sage grouse populations. This knowledge is lacking for sage grouse. While studies have documented sage grouse food habits in the May through October period, the November through April period has been virtually ignored. Nutrient levels of sage grouse diets are unknown in the wild and have been determined in only 1 study of penned birds (Barber et al. 1969). The ability of sage grouse to select nutritionally superior species, subspecies,or individual plants of sagebrush to feed upon has not been documented, yet this is a basic premise if fertilization is to succeed as a mitigation technique. �152 P. N. OBJECTIVES The objectives of this study during the pre-treatment period are: 1. Determine feeding preferences of sage grouse in the January-late April period within the area to be impacted by mining and mitigation. 2. Determine composition of sagebrush by species and subspecies from feeding and random sites. Quantify nutritional quality of sagebrush leaves from sage grouse crops and gizzards. 3. Quantify habitat use within the area to be impacted by mining and mitigation through reobservations of radio-marked sage grouse and pellet transects. 4. Collect data on sage grouse reproductive parameters in the mining and control areas and for hens moving from the area to be impacted by mitigation. 5. Monitor the sage grouse population through banding, lek counts, and collection of harvest data. SEGMENT OBJECTIVES 1. Review available literature on grouse nutrition, composition of grouse foods, composition of sagebrush, and reproductive parameters of grouse and closely related species of birds (i.e. Galliformes). 2. Review available literature concerning sage grouse and techniques adaptable to studies of galliform birds. 3. Review available literature concerning fertilization of sagebrush rangelands. 4. Capture up to 15 sage grouse within the treated or mining area each year during January and February 1981, 1982 for attachment of tail clip or poncho radios. 5. Relocate radio-marked sage grouse from January into July for observation of patterns of habitat use and location of feeding sites. 6. Collect 2 sage grouse per week within the area to be treated in January-April 1981 and 1982 for analysis of crop and gizzard contents and feeding sites. 7. Analyze crop and gizzard contents and sagebrush collected at feeding and random sites for percent nitrogen, dry matter, calcium, fiber, essential oils, and calories of energy available. 8. Randomly place 10 O.5-km long and 2.5-m wide sage grouse pellet transects in the northwest (control) study area. 9. Search and clear all (10 in the control area, 20 in the mininq area) pellet transects each year in late Mayor early June and late September or early October depending upon snow cover. �153 10. Capture and place tail clip or poncho radios on up to 10 female sage grouse in each of 2 study areas (mining and control) during late March or April in 1981 and 1982. 11. Locate nests of radio-marked female sage grouse within the mining and control areas and those dispersin~ from the study areas in AorilMay-June. Determine egg weights and sizes, clutch size, egg fertility, egg hatching success, nesting success, and nest site characteristics (i .e., vegetation composition, height, cover, etc.). 12. Periodically (3-5 day intervals) locate radio-marked hens after hatching to estimate brood survival from hatching until radio failure or 1 September. 13. Continue counts (at least 4/breeding season) of all sage grouse present on all known leks within North Park. Emphasis will be placed on leks within and adjacent to the study areas. Counts will be in the 04300730 interval from about 20 March to 31 May. Birds present will be classified as males and females. 14. Continue banding samples of male and female sage grouse with a goal of 50 males (necessary for survival estimates) in each of 4 quadrats of North Park. A sample of 20% of the high spring count of males on each lek within the 2 study areas is desired each year. Grouse will be located where they roost 2t night by spotlighting and will be trapped with long-handled nets. Birds trapped will be banded with serially-numbered aluminum bands and color-coded (year) bandettes. 15. Population size will be estimated using counts of birds present on leks, established patterns of lek attendance and known sex ratios, and ratios of banded to unbanded birds in harvest samples. 16. Harvest data will be obtained from check stations and volunteer wing collection barrels. 17. Compile data, analyze results, and prepare progress reports. DESCRIPTION OF STUDY AREAS The investigation was conducted in North Park, Jackson County, Colorado (Fig. 1). Data on winter browse and habitat preference were collected in the northeast study area, centered on the area of current mining activity north and east of Walden. Reproductive data were collected in the mining area and the (control) study area located north and west of Independence Mountain. North Park is a sagebrush-dominated grassland historically used for 1ivestock grazing. Beetle (1960) identified 5 species of sagebrush in North Park; Artemisia argillosa, ~. cana , ~. longiloba,~. ~, and~. tridentata. Subspecies of A. tridentata occupy approximately 90% of the sagebrush type (Smith 1966). �154 ~ N r~ rn ~ 0 ~ ~ 5 10 KILOMETERS "1- • COALMONT f5 oIf ~ !\I s ~ • DENVER COLORADO Fig. 1. North Park, Jackson County, Colorado. Intensive study areas are the stippled areas lying between the Canadian and Michigan rivers and North of Lake John. �155 METHODS AND MATERIALS Telemetry Sage grouse were captured at night while roosting along roads and on leks using long-handled nets and hand-held spotl ights (Pyrah 1959, Giesen et al. 1982). Captured grouse were marked with serially numbered size 14 (females) or size 16 {males} aluminum leg bands and unnumbered plastic bandettes color coded to year of capture. Classification of sex and age followed Beck et al. (1975). Radio transmitters (20-22 g, Wildlife Materials) were attached to the central rectrices using a modified bolt and clamp device similar to the tail cl ip described by Bray and Corner (1972). Radio-marked sage grouse were relocated using a portable receiver and hand-held 3-element yagi antenna. Radio-marked sage grouse were primarily used to locate winter flocks rather than determine movements; thus, daily relocations of individuals were not stressed. Vegetation Vegetatioil measurements (height, canopy cover, percent foliation) at winter feeding, nesting, and random sites were made using a modification of Canfield's (1941) Iine-intercept method. Three or 4 randomly oriented 10-m transects were measured at feeding and random sites. TvJO 10-m North-South transects centered on the nest bowl were measured at nest sites. In addition, slope and aspect were determined with an Abney level and compass, respectively, at all sites. Snow depth was measured, when present,at each plant which intercepted the Iine. There was, however, no significant snow depth in 1981 and most of 1932. Sagebrush leaf samples were collected from each plant that intercepted the line transect~ at feeding and random sites. Sagebrush leaf samples were individually tagged and sealed in air-tight plastic bags, and then frozen until used for plant identification or nutritional analysis. Plants which intercepted the line at feeding sites were examined for evidence of feeding. Sage grouse cut rather than pick leaves from sagebrush plants. The exposed interior of freshly eaten leaves is light green and contrasts sharply with the dark surface of the leaf. Thus, fed-upon plants can be quickly and reliably identified. Sagebrush samples were identified to species by morphological characteristics (Beetle 1960, McArthur et al. 1979, Winward 1980) and to subspecies by the fluorescent pattern of a water extract of crushed leaves observed under ultra-violet light (Stevens and McArthur 1974). A similar method (Young 1965, Winward and Tisdale 1969) using ethanol or methanol leaf extracts gave similar but less satisfactory results. Representative samples of each taxon observed were sent to E. D. McArthur, Principal Research Geneticist at the Shrub Sciences Laboratory, USDA Forest Service, Provo, Utah, for confirmation. All identifications were confirmed. �156 Collections Sage grouse were collected by shooting at, or just prior to, the conclusion of morning or evening feeding periods for crop and gizzard contents and body composition analysis. Nutritional Sagebrush leaf samples from crops and gizzards were freeze-dried and ground to a uniform consistency with a mortar and pestle. Nitrogen was measured by Kjeldahl analysis and protein was estimated by multiplying nitrogen by 6.25. Structural carbohydrates - cell wall constituents (ewe), neutral detergent fiber (NDF), acid detergent fiber (ADF), and lignin were determined using methods described by Van Soest (1963a, b; 1967). Ether extract (crude fat) was extracted with diethyl anhydrous ether as solvent in a Soxhlett apparatus. All analyses of protein, ADF, ewc, lignin, and ether extract were performed in duplicate. Mineral components were extracted by ashing the sample at 550 e for 12 hours. Calcium, sodium, potassium, and magnesium were determined on a Perkin Elmer model 303 atomic absorption spectrophotometer in ppm and converted to percent dry matter (DM). Phosphorous was determined on a Beckman Spectronic 20 spectrophotometer. All analyses were computed on a DM basis. Nitrogen free extract (NFE) , a measure of soluble carbohydrates, was determined by subtraction. Sagebrush leaf samp~es from vegetation transects were pooled by transect, species, subspecies, and feeding status (feeding site transects only). Twenty-gram pooled samples were prepared for analysis by hand-picking equal amounts (wet we lqh t Lof leaves from all plants in a particular group. Samples were immersed in liquid nitrogen in a mortar and ground with a pestle to a uniform powder. Nitrogen and protein were determined as above. Body Composition Data were collected on total body weight, heart, gizzard, crop, M. pectoralis, !!. supracoracoideus,and testes weight, total length, and length of bill, wing, primaries I-X, caeca, small intestine, large intestine, and diameter of the ovary, oviduct, and largest follicle of females. In addition, fat appearance was evaluated on a scale of 1-5 and the caeca and small and large intestines were searched for parasites. Parasites were preserved in a solution of alcohol-formal in-acetic acid fixative for later identification. Sage grouse carcasses were plucked, the feet, outer portion of wings (distal to radiale/ulnare) and the head were removed and the intestinal contents emptied prior to homogenizing with a meat grinder. Two 100 g subsamples of each homogenized carcass were dried at 80 C for 24 hours and ground to a fine powder in a mortar and pestle. Fat was extracted from duplicate 1 g sUbsamples of the dried and crushed material in a Soxhlett apparatus with diethyl anhydrous ether as solvent. �157 Population Monitoring Pellet transects were established and counted using techniques described by Schoenberg (1980). Lek counts were conducted from 0430 to 0630 MST. At least 3 counts, 5 minutes apart, were made of both hens and cocks on a given morning with spotting scope and/or binoculars. One count in each of 4 periods (1-10, 11-20, 21-30 April; 1-15 May) was attempted for each ground. Harvest data were collected from 4 volunteer wing collection barrels (Hoffman and Braun 1975) and 2 check stations in 1981. Wing collection barrels were maintained throughout the 23-day sage grouse season (12 Sep - 4 Oct) at Cameron Pass, Stateline, Willow Creek Pass, and Muddy Pass. Check stations were operated at Willow Creek Pass on Colorado 125 and east of Gould on Colorado 14 on 12, 13, and 20 September. One wing was collected from all sage grouse possible. Data obtained from each party were: county of origin; number of hunters; total hours hunted; birds observed, bagged, and lost; number of banded birds and where shot; and area hunted within North Park~ Reproductive Data Clutch size was recorded from all nests located, and length, width, weight, and volume of all eggs was determined. Egg parameters were measured using a micrometer accurate to 0.1 mm (length and width), a triple beam pan balance accurate to 0.01 g (weight), and a 500 ml graduated cyl inder (volume). Egg volume was determined by water displacement. RESULTS AND DISCUSSION Vegetative Characteristics at Feeding and Random Sites Vegetative characteristics were measured at 41 feeding and 36 random sites. Sagebrush at feeding and random sites was similar (p > 0.05) in average height and percent foliation,but feeding sites had greater (p < 0.05) sagebrush canopy cover (Table 1). Snow cover in 1982 was probably responsible for the larger canopy cover value at feeding sites. There was enough snow in January and February to shift birds into high canopy cover areas to feed. Schoenberg (1982) reported canopy cover of 50.9 and 49.6%, respectively, for females and males at feeding-loafing sites in the same study area during 2 heavy snowfall winters. The average height of sagebrush (23.6 cm) at feeding sites was much less than the total height of 42.3 and 40.6 cm and the above-snow height of 34.3 and 26.1 cm, respectively for female and male feeding-loafing sites in the same study. �158 Table 1. Mean canopy cover, plant height, and percent fol iation of sagebrush on transects at feeding and random sites, North Park, Colorado, January-Apri 1 1981, 1982. Site Canopy cover (%) N Percent fol iated Height (cm) Feeding x SO Range Random 24.20 8.94 10.27-51.10 23.56 9.91 9.81-46.37 76.88 8.02 57-94 b 19.46 7.92 7.98-41.73 24.07 9. 11 8.30-41.93 73.11 10.10 45-95 36 x SO Range aCanopy cover was measured at 40.of 41 sites. bOifferent (~< 0.05) than value obtained at feeding sites. Apparently, shorter, more open stands of sagebrush are selected in winters with relatively little snow. Sage grouse may select such sites because they allow for easy detection of and rapid escape from aerial predators, or because of a preference for small stature food plants as noted by Gill (1965). Although average sagebrush height at feeding and random sites did not differ (p > 0.05), this does not preclude the possibility that sage grouse are selecting feeding sites on the basis of height. They may be selecting for sagebrush close to the average height. Sagebrush at feeding and random sites was identified to species and subspecies. Sage grouse selected (p < 0.05) for feeding sites with proportionally more A. tridentata wyomlngensis (ATW) and less ~. tridentata vaseyana (ATV) and~. longiloba (AL) than at random sites (Table 2). A. cana viscidula and A. argillosa were each encountered on 1 random transec~ut no feeding transects, and were excluded from the analysis. Table 2. Occurrence (%) of 3 sagebrush taxon at random and sage grouse feeding sites, North Park, Colorado, January-Apri 1 1981, 1982. b Site ATW Percent occurrence b ATV a N sagebrush plants ALb encountered Feeding 86 12 2 1,508 Random 48 41 11 1,042 a X 2 -- 433, 2 df, P < 0.005. b~TW = Artemisia ~ridentata wyomingensis, ATV AL - A. 10ngi loba. A. t. vaseyana, �159 Food Plant Selection The species and subspecies composition, and the ·structural characteristics of all plants and fed-upon and non fed-upon plants encountered at feeding sites were determined to test for selection of certain taxon or growth forms as food plants. Within feeding sites, a definite preference (p < 0.05) for ATW as a food plant was observed (Table 3). Six hundred and ten of 676 (90%) fed-upon plants identified at feeding sites were ATW. ATV and AL comprised 7 and 3%, respectively, of the fed-upon plants. Table 3. Frequency (%) of 3 sagebrush taxon as sage grouse food plants, North Park, Colorado, January-April, 1981, 1982. Plant status Percent occurrence b ATW ATVb a N sagebrush plants ALb encountered Food plants 90 7 3 676 All plants 86 12 2 1,508 aX2 AL = 39.32, 2 df, bATW = Artemisia A. long iIoba . t < 0.05 tridentata wyomingensis, ATV = A. t. vaseyana, It is apparent that the selection for ATW at feeding sites (Table 2) was due to the preference for this subspecies as a food plant. ATV was selected against, comprising 12% of the plants at feeding sites but only 7% of the fed-upon plants. In addition, ATV comprised 41% of the sagebrush at random sites (reflecting availability) but only 7% of the fed-upon plants. ATW comprised 48% of the sagebrush at random sites but 90% of the fed-upon plants. AL does not seem to be strongly selected for or against; it seems to be readily eaten when it is encountered at a feeding sjte. These findings do not support the conclusions of Brunner (1972), who felt this species was seldom eaten by sage grouse. Selection (p < 0.10) for food plants with larger or smaller values of height or percent foliation occurred on 11 of 21 (52%) feeding site tra~sects measured in 1981 (Table 4). Grouse usually selected larger, more fol iated plants to feed upon. Larger, more fol iated plants were generally found along road edges, irrigation ditches, fencel ines, or more frequently in small scattered clumps which resemble Mima mounds (Arkley and Brown 1954). Selection for food plants with different structural characteristics than non-food plants seems to be independent of species or SUbspecies differences. �160 Table 4. Mean plant height and percent foliation of fed-upon and non fed-upon sagebrush plants from sage grouse feeding sites, North Park, Colorado, January-April 1981. Ht (cm) Percent foliation Greater N F Feeding site b FO-l FO-2 FO-3 Fo-4,5 FO-7 Col-2 Col-5 Col-12 Co 1-13 3.67 5.45 32.46 17.82 24.86 14.76 27.64 20. 15 24.00 17.53 86.4 95.8 70.0 70.3 79.7 57.2 85.0 71.6 85.56 77.27 Sma 11erb Col-3 Col-9 a 8.06 12.94 11.45 19.32 F = fee-upon plants, N = non fed-upon plants. bGreater = fed-upon plants with mean values great~r (p < 0.10) than non fed-upon plants, Smaller = fed-upon plants with mean values smaller (~< 0.10) than non fed-upon plants. A nutritional analysis of leaf material collected at feeding and random sites was conducted to examine possible reasons for the strong preference for ATW and certain plant growth forms as food plants. Crude protein was the nutrient of choice because of literature documenting grouse selection for protein in their winter diet (Miller 1968, Gurchinoff and Robinson 1972, Moss 1972, Moss et ale 1972, Huff 1973, Doerr et al. 1974). Protein content of the 3 taxon of sagebrush commonly encountered on random transects varied (Table 5). ATW, the most preferred of the 3 taxon, contained the most protein, while the least preferred taxon, ATV, contained the least protein. The 2 samples of AL for which protein determinations were made indicate it is similar to ATW in protein content. This may account for the apparent willingness of sage grouse to feed on this species despite its small stature and relative scarcity. �161 Table 5. Crude protein content in leaf samples of 3 sagebrush taxon collected at random site transects, North Park, Colorado, JanuaryApri 1 1981, 1982. Crude protein, % a ALa 14.23b 1'0. 64 15. 15 1.82 1.40 2.04 SD N a ATV ATW aATW = Artemisia tridentata wyomingensis, vaseyana, AL =~. longiloba. (~< bDifferent 2 13 13 ATV = A. tridentata 0.05) than value for ATV. The crude protein values documented were similar to, but slightly higher than, other publ ished values for midwinter protein levels of sagebrush current growth leaves and stems (Table 6). This difference may represent site specific differences, or it may reflect a nutritional difference between leaves (this study) and leaves and stems analyzed in other studies. ' Table 6. Winter protein levels (%) of leaves and leaves and stems of Artemisia tridentata wyomingensis, A. tridentata vaseyana, and A. tridentata compiled from the literature. Leaves Leaves and stems Leaves Means 14.2 12. 1 10.6 aATW =~. tridentata wyomingensis, ATV =~, Leaves and stems Leaves 13.8 trldentata vaseyana, AT -~. trldentata (no subspecies bHi lchunas ct al. 1978 (Hlddle Park, Colo.). cPrescnt study. dKinney and Sugih~ra eSheehy 1943. 1975 (3 areas in Oreg.). fWelch and HcArthur gPatterson 1979 (location of 1~ accessions; 1952 (Wyoming). Ariz. [1], Colo. [1]. Nevada [IJ. Utah [IIJ). Leaves and stems 8. 1 specified). �162 Protein contents ofATW samples from feeding and random sites were compared to determine if sage grouse select nutritionally superior sites at which to feed (Table 7). Apparently, sage grouse do select feeding sites containing ATW with greater (p < 0.05) protein level s than random sites, although the way in which they do so is unclear. This suggests that there may be nutritional as well as species composition and structural restrictions to potential sage grouse habitat. Table 7. Crude protein content of Artemisia tridentata wyomingensis feeding and random sites, North Park, Colorado, January-April 1981, 1982. Random sites Feeding sites 16.42a Crude protein, % SO 14.23 1.82 2.02 16 N (~< aOifferent at 13 0.05) than value obtained at random sites. A final test for selection was made by comparing protein levels of fed and non fed-upon ATW and ATV (Table 8). Although the results are still incomplete and sample sizes small, particularly for ATV, fed-upon plants appear to contain more protein than non fed-upon plants. Preference for certain growth forms as food plants (principally larger, more foliated plants) may be the means by which high protein plants are selected. Table 8. Crude protein content of fed-upon and non fed-upon plants of Artemisia tridentata at sage grouse feeding sites, North Park, Colorado, January-April 1981. ATWa Fed-liDOn b x - SD N = Non fed-upon Fed-upon Non fed-upon 17.22 15.59 13.37 11.24 1.54 2.24 1.85 1.62 3 6 13 aATW vaseyana. ATVa 13 Artemisia bDifferent (~< tridentata wyomingensis, ATV A. tridentata 0.05) than value obtained from non fed-upon plants. �163 Sage grouse seem to be selecting from available food resources at 3 levels to maximize their protein intake. They stronqly prefer the most nutritious subspecies of sagebrush (ATW), select the most nutritious sites containing this sUbspecies for feeding, and select the most nutritious plants within this subspecies to feed upon. The result of these 3 levels of selection is that sage grouse boost the average crude protein level potentially available from 10.6% for ATV at random sites to 17.2% for fed-upon plants of ATW. The result of this selection perhaps can be better seen in the crude protein content of crop contents of collected birds. The values for crude protein of ATW and ATV fed-upon plants indicate selection. but alone do not tell anything about the protein level in the diet. Birds may feed longer, and hence have a greater intake, on 1 subspecies or on more nutritious plants within a subspecies. For this reason a complete nutritional analysis of crop and gizzard contents was undertaken. Nutritional content of sagebrush leaf samples from sage grouse crops and gizzards varied (Table 9). Crude protein, soluble carbohydrates (NFE), and several minerals apparently undergo some digestion in the gizzard since the values for these nutrients were lower (p < 0.05) for samples from gizzards than from crops. The larger (p <.0.(5) ether extract and structural carbohydrate (NDF, ADF, lignin) values in the leaf samples from gizzards most likely reflect a proportional increase due to the decrease in protein, soluble carbohydrates, and minerals rather than any real increase. The lower (P < 0.05) DM from gizzard leaf samples (or higher moisture content) is-probably due to the addition of digestive enzymes and acids (HCL, Pepsin) to the food material in the proventriculus. Pepsin (a protein) may actually inflate the protein value of gizzard contents and reduce the apparent magnitude of digestion. It would appear that the value of nutritional analyses from gizzard contents is questionable. Table 9. Nutritional analysis of sagebrush leaves from sage grouse crops (N = 14) and gizzards (N = 23), North Park, Colorado, Jan~ary~ Apri I '-9"81. (Expressed as pe-;:centDM). Prota OH Source d NFE EEb NFD 21.8Se 27.22 Crops 32.22d IS.87 40.20 15.233 G izza rds 23.20 13.62 31.36 23.50 LlGc AOF e ASH P d Na K H9 d d 16.86 11.84 1.1S 0.14 0.47 1.60 0.36 21. 15 8.66 lC.37 0.70 0.15 0.44 1.21 0.23 aprot. = Protein. dCrop values bEE = Ether extract. eCrop values smaller cLiG = Lignin. Ca d 6.44e larger (! < 0.05) (! < O.OS) than gizzard than gizzard values. values. �164 The hypothesis that sage grouse feed equally on all plants they feed on, derived from field observations, was tested from these data. If the hypothesis is true, then a pooled mean protein level weighted by the frequency of each of the taxon as food plants (ATW-90, ATV-7, AL-3) should equal the average protein content of crop contents, or 15.87%. The calculation is: 0.9 x 17.22 + 0.07 x 13.37 + 0.03 x 15.15 = 16.89% This value is similar to the 15.87% found in crops, but it may indicate a greater intake of ATV than previously indicated. Winter Fat Levels Body fat content was examined as an index to the nutritional status of sage grouse during winter. Analyses are complete for 1981. Total body fat content varied between sex and age classes (Table 10). Fat levels were within the ranges reported for other tetraonids in winter (West and Meng 1968, Thomas et al. 1975, Grammeltvedt 1978, Thomas and Popko 1981). Apparent differences between groups were not subjected to statistical tests for verification because of small sample sizes. There appears to be real differences between juveniles and adults of both sexes. Juvenile males may have less fat than juvenile females. The large difference in body size may account for this, as juvenile males must synthesize a far greater amount of protein in the same period of time. Substantial variation within sex and age classes was evident. Additional analyses will be used to determine if this variability is real or due to correctable flaws in methodology. Table 10. Total body fat content of sage grouse collected April 1981, North Park, Colorado. Adul t rna1e Total body fat (percent 1ive weight) Juvenile male Adult female in February- Juvenile female 2.58 5.03 2.56 4.87 2.66 1. 08 0.79 2.41 0.75 0.95 2.23 1. 90 3.40 4.15 4.12 2.39 2.21 0.73 2.29 1. 19 3.16 Means 3.54 1.20 2.91 1. 84 Banding Sample Trapping was initiated in mid-February and concluded in late May in both 1981 and 1982. During this period 369 sage grouse (300 males, 69 females) were banded in 1981 and 402 sage grouse (300 males, 102 females) were banded in 1982. The quota of 300 newly banded males was met both years as was the distribution quota of 50 newly banded males/quadrat. The quota of 100 newly banded females was met in 1982 but not in 1981. �165 The larger grounds within the mining study area were trapped extensively. The desired goal of trapping 20% of the high spring count of males on each lek was exceeded on 2 of 8 leks in 1981 and 3 of 9 in 1982. However, 21% and 22% of the high spring count of males on all mining area leks combined were banded in 1981 and 1982, respectively. lek Census All (8 in 1981,9 in 1982) leks in the study area were censused between 1 April and 21 May in 1981 and 1982. Two leks recently abandoned (Pronghorn and Roth) were also checked for activity. Dates of peak lek attendance were similar between years with males peaking in late April to mid-May and females peaking in early to mid-April (Table 11). Much of the censusing in both years occurred after the peak of hen attendance as the total number of hens did not approach twice the male total (expected if a 2:1 ratio of females to males in the spring population is assumed). Table II. High spring lek counts, northeast Males lek Buteo 1981 a Canuck North Park, Colorado, Dates 1982 1981 1982 1981-82. Females 1981 1982 Dates 19S1 1982 13 Apr 22 19 7 May 30 Apr 0 20 All dates 16 33 13 Apr 12 May 5 17 13 Apr 13 Apr 27 Apr 23 50 6 Apr 14 Apr Denmark Hawka 109 71 7 May 7 7 29 Apr 6 Apr 0 13 Perdiz 23 27 7 May 13 Apr 29 35 6 Apr 6 Apr Prague a Ram 26 7 22 Apr 20,22 Apr 11 2 13 Apr 20 Apr 22 6 7 May 6 Apr 8 0 6 Apr A 1I dates Raven 49 44 27 Apr 29 Apr 38 25 14 Apr Turkey b Totals 22 274 236 aNew lek discovered in 1981. bNew lek discovered in 1982. 27 Apr 3 114 All dates 6 Apr 13 Apr 27 Apr 165 Peak lek counts of male sage grouse have remained relatively stable in the 1980-82 interval. Although the total number of males changed only slightly, their distribution within the study area has changed dramatically (Table 12). Such variation in strutting ground counts is not unusual (Beck and Braun 1980). The steady decl ine of Denmark may be of some concern. The extreme decline of Prague is unusual. This decline is not a sampl ing artifact because this ground was counted 6 times in 1982 and did not exceed 7 males. The appearance of a new ground (Buteo) less than 2 km away in the same drainage suggests that Buteo is a new strutting site replacing Prague. Buteo was first discovered in 1981 but it was not a consistent ground until 1982. Birds were not seen on both Prague and Buteo at the same time in 1981. It would appear that Buteo was originally an alternate strutting site for birds from Prague and that some birds switched to or recruited to it as a primary lek. The 7· males remaining on Prague are probably old males; it is likely they will remain at this site as long as they survive. �166 Table 12. Trends in peak lek counts of male sage grouse, northeast Park, Colorado, 1979-81 . Year Lek Buteo Canuck Denmark Hawk Perdiz Prague Pronghorn Ram Raven Roth Turkey Totals Avg/lek 1979 1980 21 136 18 144 16 43 10 8 34 0 Percent change 1981 to 1982 1981 1982 22 16 109 7 23 26 19 33 71 7 27 7 0 6 44 0 22 - 14 +106 - 35 0 + 17 - 73 236 26.2 - 14 0 63 2 43 1 22 49 0 291 41.6 248 41.3 274 34.2 North - 115 - 10 It is interesting to note the steady decline in the average number of males/lek since 1979, even though the total number of males has changed little (at least in the 1980-82 interval). This is due largely to the discovery of 4 new grounds during this period which all contained less than the average number of males. If these grounds were newly established in the year of discovery, it could indicate that the optimum lek size is declining. It may be more advantageous for yearling males to recruit to small grounds or establish new grounds than attempt to compete at large established grounds. There was a marked trend for large grounds to decrease, small grounds to increase, and moderate sized grounds to remain fairly stable in all of North Park during this period (C. E. Braun, unpubl. data). The problem is, if 26 is an optimal size for a ground, why hasn1t the mean revolved around this optimum all along, with grounds being established or created in response to population fluctuations? It may be that optimum lek size varies over time. This impl ies variation in genetically determined behavior, such as has been documented for red grouse (Moss and Watson 1980). Nesting Data Twelve radio transmitters were attached to 17 female sage grouse in 1981. Eleven transmitters were attached to 14 hens in 1982. Several radios were attached to more than 1 bird due to predation or loss of the transmitter by the first hen. Subsequent relocations resulted in finding 9 nests (6 original and 3 renests) in 1981 and 9 nests (7 original and 2 renests) through 31 May 1982. Transmitter failure (6), predation (2 birds), and transmitter loss (3 birds) hindered finding a larger sample of nests. The discussion and data which follow are from 1981 only because data collection and analysis from 1982 nests are not yet complete. One of 8 radio-equipped brought off young. Two 5 were predated during apparently by a coyote hens which initiated a first clutch successfully nests were apparently predated during laying and incubation. One hen was killed while incubating, (Canis latrans). Most nest depredation seemed �167 attributable to ground squirrels (Spermophilus spp.) (3) or badgers (Taxidea taxus) (1). Four hens that lost their first clutch retained functional transmitters long enough to renest. Of these, 2 (both adults) renested and 2 (1 adult, 1 yearling) did not. Both 2nd clutches were successful. Nest site vegetative parameters were measured (Table 13). Nest sites contained significantly taller and denser sagebrush (~< 0.05) than random sites. Sage grouse generally chose plants higher than average unde r wh ich to nes t . Table 13. Average height, canopy cover, and height of sagebrush at sage grouse nests, North Park, Colorado, 1981. Nest site 5092 5121 5122 5123 5124 5127 (1st) 5127 (2nd) 5159 Avg height (cm) Canopy cover (%) x 39.10 32.18 52.96 27.14 39.84 44.48 56.37 57.89 a 43.75 51.0 15.6 38.4 20.6 46.7 5~.7 45.7 50.5 a 39.9 so 11.28 14. 13 Nest plant height (cm) 40.0 41.0 52.0 36.0 41.0 57.0 46.0 61.0 46.75 8.97 a Higher (~ < 0.05) than value obtained at random sites. Data on clutch size and length, width, weight, and volume of eggs were collected and compared to similar data from 1979 and 1980 by age of hen and nest status (Tables 14, 15). First clutch egg length of adults decl ined significantly from 1979 to 1980 and from 1980 to 198i. Adult first clutch egg weight followed a similar trend, decreasing from 1980 to 1981 (no data from 1979). Egg width for this group decl ined significantly from 1979 to 1980 but, contrary to both egg length and egg weight, width increased significantly from 1980 to 198'1. Thus, egg weight seems more correlated with egg length than egg width. Eggs in 2nd clutches of adult hens were significantly longer and heavier in 1981 than 1980. This apparent difference should be interpreted with caution because of a small sample size (1 clutch in 1980 and 3 in 1981). In general, clutch size, egg length, width and weight declined from 1979 to 1981 for first clutches of yearling and adult hens. Yearl ings laid significantly wider and fewer eggs than adults. First clutches of adult hens were significantly lighter and clutch size was larger than 2nd clutches. This increase in weight in 2nd clutches is apparently due to an increase in both egg length and width although the increases were not significant with the sample size available. �Table 14. Clutch size and length, width, weight, and volume of eggs from 1st and 2nd clutches of adult and yearling sage grouse, North Park, Colorado, 1979-81. Age and nest statusa Year 1979 A1 Y1 1980 A1 A2 Y1 x N 58.34c,d 56.19 d 56.97a 54.32 55.94 A1 A2 Y1 1981 Width Len9th N x 38.66c,d 39.24c,d 37.99d 16 13 53 6 23 55.36 58.42 56.32 Weight 38.13 38.35 38.32 38.59 37.84 35 17 5 x N x 16 13 - 53 6 23 44.92d 43.17 43.74 35 17 5 Volume 43.92 46.94 43.36 Clutch size N x N 8.00b,d 6.50 7.62b 53 6 23 3 2 8 6.00 6.67 41.03 43.82 40.40 29 17 5 29 17 5 3 7.20b 5 5.67 5.00 3 1 _. 0' ~Age(A)=Adult (Y)=Yearling. Nest Status (1) 1st clutch, Different (p < 0.05) from yearling values. cDifferent (p < 0.05) from 1980 values. dDifferent (~< 0.05) from 1981 values. ce (2) 2nd clutch. Table 15. Average clutch size and length, width, weight, and volume of eggs from yearling sage grouse 1st clutches and adult 1st and 2nd clutches, North Park, Colorado, 1979-81. Length Width Weight Hen age Nest x N - x N x Yearl ing Adult Adult 1st 1st 2nd 56.07 56.64 57.34 41 104 23 38.57a 38.20 38.47 41 104 23 43.67 44.57 45.96a aDifferent (~< 0.05) from adult 1st clutches. and adult CI size N x 28 82 23 6.33a 7.56 5.75a N 6 16 4 �169 Pellet Transects The 30 randomly selected pellet transects (20 in the northeast, 10 in the northwest) were counted and cl~ared in May 1981 and 1932 and October 1981. The results of these counts and comparative data from summer 1979 through summer 1980 (Schoenberg 1981) have varied (Table 16). The large variability in the data make interpretation difficult. Many transects, particularly in the northwest, contribute...Uttle or no information, since few or no droppings have been found along them. Since no data are available on defecation rates of sage grouse, dropping data of this type cannot be used to estimate population levels. At best they can be used as an index to compare year-to-year variation in populations or concentrations of birds on wintering areas. Transects with few or no droppings do not contribute meaningfully to this index. For instance, transects with 0 droppings/day year after year tell nothing of variation since they donlt vary. Transects with a few droppings/day are misleading since the change of 2 or 3 droppings from 1 year to the next can cause hug~ swings in the number of droppings/day. This can cause the index to change dramatically, even with stable populations and concentrations. It appears that the northeast study area did not support the "normal" concentration of birds in the mild winter of 1981. Dropp!ngs/day declined by 58% from 1980 values. Strutting ground counts and harvest statistics suggest there was little change in the population during this period. The winter 1982 value of 0.33 droppings/day is 20% higher than the 1981 val ue but 49% below the lS80 va lue , This supports. field observa.,.._·· tions that there were somewhat more sage grouse wintering i.nthe northeast in 1982 than in i981, but nowhere near the concentrations of 1980. The pellet data from the northwest are extremely difficult to interpret since 5 of 10 transects have not had a dropping on them when they were cleared. Whether this reflects a much lower density of sage grouse or poorly located transects (relative to use sites) is not known. The numbers of droppings/day in the northwest after the 1981 and 1982 winters were much lower than similar values for the northeast area. It is possible that this is due to relatively few sage grouse wintering in the northwest, even in mild winters, or it may be a sampling artifact. Harvest Data Sage grouse harvest data were collected at Willow Creek Pass and Gould check stations on 12-13 and 20 September, the 1st and 2nd weekends of the sage grouse hunting season. These data, and comparable data from 1974-80 are presented in Table 17. The 1.3 birds per hunter in 1981 was similar to 1980 (1.4) and slightly above the long term (1.2) average although total hunters checked and birds harvested decreased. Hunter distribution and harvest in the mining (Eagle Hill) and control (Independence ~1ountain) areas were similar to 1980 and the long term average with 11.5 and 3.3% of the total hunters and 10.5 and 1.7% of the harvest, respectively (Tab Ie 18). �Table 16. Number of sage grouse droppings/day summer 1979 - winter 1982. Transect Summer 1979 Winter 1979-80 0 0 0.01 0.02 1.27 0.36 0.50 0.79 0.84 0.09 0.56 0.05 1.61 0.71 0.40 0 4.62 0.03 0.39 3 4 48 57 60 74 78 81 83 88 99 115 129 136 166 174 179 194 196 197 O. 11 0.03 2.06 0.76 0.11 0.07 0.03 2.20 0.51 0.59 0.06 3.24 0.48 0.02 0.93 0.29 0 0.41 0.21 1.02 0.08 Avg 0.61 0.65 Northeast Summer 1980 on 30 transects Winter 1980-81 Summer 1981 Winter 1981-82 0.03 0.24 1.43 0.06 1.02 0.08 0.60 0.69 0.02 0.06 0.84 0.25 O. 11 0.52 1.22 0 0.05 0.05 0.21 0.12 0.06 0.04 0.07 0.46 0.39 0.03 0.32 1.47 0.13 0 0 0.27 0.12 0.69 0.03 0 0.55 0.12 0.07 0.59 0.09 0.01 0.01 0.28 0.01 0.05 0.27 0.36 0 0 0.18 0 0.53 0.04 0 0.44 0.01 0.01 0.05 0.21 0.04 0.01 0.81 0.54 0.10 0.71 0.25 0.68 0.01 0.03 0.24 1.68 0.45 0.01 0 0.48 0.02 0.09 0.17 0.38 0.27 0.12 0.33 a in 2 study areas, North Park, Colorado, Transect 12 17 19 40 119 50 63 76 80 95 Northwest Winter Summer 1980-81 1981 0.01 0.27 0 0 0 0.01 0 0 0.06 0 0 1.64 0 0 0 0.02 0 0 0.63 a Winter 1981-82 0 0.15 0 0 0.05 0 0 0 0.38 0 -....J 0 0.04 0.23 0.06 �171 Table 17. a Sage grouse harvest statistics, North Park, Colorado, 1974-81. ,b No. birds observed Crippl ing loss (%) Birds per hunter Year 1974 1975 1976 1977 1978 1979 1980 1981 730 738 595 353 350 521 567 523 6,062 5,735 3,393 3,303 4,922 6,910 5,000 6, 189 785 551 459 385 480 982 794 695 12.9 9.6 13.5 11.7 9.8 14.2 15.9 11.2 5.1 7.1 5.7 10.6 5.7 4.7 4.3 5.4 1.1 0.7 0.8 1.1 1.4 1.9 1.4 1.3 547 5,189 641 12.4 6.1 1.2 Avg No. birds ha rves ted Hunter efficiency (%) No. hunters checked aFrom check station data only. bUnpublished data (C. E. Braun) • Wing Analysis A total of 1,032 sage grouse wings was collected in North Park during the 1981 hunting season, an increase of 10% from the 939 received in 1980. Of the 1,032 wings, 662 (64%) were collected at check stations, 288 (28%) from wing barrels, and 82 (8%) from field checks and miscellaneous sources. An analysis of the age structure derived from these wings (Table 19) indicates a sl ight dec~ease in production (percent immatures) from 1980 (49.1%) and the 8-year average (49.5%). The percent yearlings in the harvest (22.3) was similar to the long term average (22.4%) but substantially below 1980 (26.6%). The adult percentage of 30.4% was higher than both the 8 year average (28.2%) and the 1980 value (24.3%). This increase in the percentage of adults and decrease in percent yearlings represents the large cohort produced in 1979, which entered the adult population in 1981. Table 19. 1974-81.a Age structure of the sage grouse harvest in North Park, Colorado, Immatures Adults Yearl in9s Year N % N % N % 1974 1975 1976 i977 1978 1979 1980 1981 350 212 210 290 385 624 461 489 50.1 42.0 42.3 45.9 53.2 57.7 49. 1 47.4 138 111 117 123 143 256 250 229 19.8 22.0 23.5 19.5 19.8 23.7 26.6 22.3 22.4 210 182 170 219 196 201 228 314 30. 1 36.0 34.2 34.6 27.1 18.6 24.3 30.4 Avg aUnpublished 49.5 data (C. E. Braun). 28.2 �Table lB. Sage grouse harvest and hunting pressure (% of total) within North Park, Colorado, 1974-Bl.a,b,c aOata from check stations only. bTotals may not approximate a particular zone. 100~ as in most years some hunters and birds harvested cUnpubl ished data (C. E. Braun). could not be allocated to �173 Nesting success was determined from molt patterns from collected wings. Successful hens initiate their wing molt later and can be distinguished from unsuccessful hens on this basis. Nesting success for adults (37.4%), yearlings (21.9%), and combined ~roups (31.2%) was the lowest recorded in North Park (Table 20). This low nesting success is difficult to reconcile with average to above average birds per hunter, birds observed by hunters, percent young in the harvest, young per hen, and young per successful hen. It is probable that the number and percent of successful hens was underestimated in 1981 using the wing molt technique. If wing molts and nesting attempts were both earlier than "normal" in 1981 due to the early spring, successful early nesting hens could have progressed far enough in the molt to be classified as unsuccessful. This could account for the high value of young per successful hen in 1981 (the highest on record) since misclassification would lower the denominator (successful hens) and increase the ratio. Thus, 1981 nesting success estimates are undoubtedly minimal. It is safe to conclude that nesting success declined somewhat from 1980 and from the long term average. There are no data available to explain the apparent decline in nesting success. The hens should have been in excellent condition for laying following the mild winter and spring. If harsh winters reproductively stress hens, larger clutch sizes, less abandonment, and more renesting would have been expected in 1981, which should have increased nesting success. Clutch size (and egg size) actually declined significantly from 1979 to 1981 (Tables 14, 1'5) and nesting success declined as well. Sage grouse hens may not be reproductively stressed by harsh winters. The lack of winter and spring moisture in 1981 may have influenced nesting success. Herbaceous growth around nests may provide important concealment for incubating hens. The extent of this growth would depend directly on spring moisture. Table 20. Sage grouse nesting success and production Colorado, 197q-81.a Estimated nesting success Year Adults 1974 1975 1976 1977 1978 1979 1980 1981 65.4 53.2 52.9 59.3 59·7 65.1 55.7 37.4 56.1 Avg aUnpublished Yearl ings 46.1 39.0 26.8 32.7 38.3 55.8 30.6 21.9 36.4 Total 58.7 48.6 43.2 50.3 51.4 60.1 42.7 31.2 50.7 data (C. E. Braun) . rates, North Park, Percent young in harvest Young 2er hen 50. 1 42.0 42.3 45.9 53.2 57.7 49.1 47.4 48.6 1 .4: 1 1. 1 :1 1. 1:1 1.2.:1 1 .8: 1 2.2: 1 1 .4: 1 1 .3: 1 1.4: 1 Young per successful hen 2.3: 1 2.3: 1 2.5: 1 2.0: 1 3.6: 1 3.7: 1 3.3: 1 4. 1:1 3.0: 1 �174 LITERATURE CITED Arkley, J. A., and H. C. Brown. 1954. The ori~in of Mirna Mound (Hogwallow) microrelief in the far Western states. Soil Sci. Soc. Am. Proc. 18: 195-199. Autenrieth, R., W. Mol ini, and C. E. Braun. Editors. 1982. Sage grouse management practices. West. States Sage Grouse Comm. Tech. Bull. 1, Twin Falls, Idaho. 42pp. Barber, T. A., J. G. Nagy, and T. A. May. 1969. Nutrition and dietary preference of penned sage grouse. Proc. West. States Sage Grouse Workshop 6:180-188. Beck, T. D. I., and C. E. Braun. 1980. The strutting ground count: variation, traditionalism, management needs. Proc. West. Assoc. State Game and Fish Comm. 60:558-566. ---...". , R. B. Gill, and C. E. Braun. 1975. Sex and age determination of sage grouse from wing characteristics. Colo. Div. Game, Fish, and Parks Game Inf. Leaf1 . 49. 4pp. Beckerton, P. R. A. 1980. Effects of five dietary protein levels on reproduction of ruffed grouse. M.S. Thesis, Univ. Guelph, Guelph, Orrt . 68pp. Beetle, A. A. 1960. A study of sagebrush (the Section Tridentatae of Artemisia). Univ. Wyoming Agric. Exp. Stn., Bull. 368. 83pp. Bray, O. E., and G. W. Corner. 1972. A tail clip for attaching transmitters to birds. J. Wildl. Manage. 36:640-642. Brunner, J. R. 1972. Observations on Artemisia Manage. 25:205-208. in Nevada. J. Range Canfield, R. H. 1941. Application of the line interception method in sampling range vegetation. J. For. 39:388-394. Dargan, L. M., H. R. Shepherd, and R. N. Randall. 1942. Survey of 1941-42/Food studies, parasite relations data on sharp-tailed grouse in Moffat and Routt counties. Colo. Game and Fish Dep., Sage Grouse Surv., Vol. 4. 20pp. Doerr, P. D., L. B. Keith, D. H. Rusch, and C. A. Fischer. 1974. Characteristics of winter feeding aggregations of ruffed grouse in Alberta. J. Wildl. Manage. 38:601-615. Giesen, K. M., T. J. Schoenberg, and C. E. Braun. 1982. Methods for trapping sage grouse in Colorado. Wildl. Soc. Bull. 10:232-237. Gill, R. B. 1965. Distribution and abundance of a population of sage grouse in North Park, Colorado. M.S. Thesis, Colo. State Univ., Fort Co ll ins. 185pp. �175 Grammeltvedt, R. 1978. Atrophy of a breast muscle with a single fibre type (M. pectoralis) in fasting willow grouse, Lagopus lagopus (L.). J. Exp~ Zool. 205:195-204. Gurchinoff, S., and W. L. Robinson. 1972. Chemical characteristics jackpine needles selected by feeding spruce grouse. J. Wildl. Manage. 36:80-87. of Hoffman, R. W., and C. E. Braun. 1975. A volunteer wing collection station. Colo. Div. Wild1. Game Inf. Leafl. 101. 3pp. Huff, D. E. 1973. A preliminary study of ruffed grouse-aspen nutrient Ph.D. Thesis, Univ. Minnesota, Minneapolis. 160pp. relationships. Keller, R. J., H. R. Shepherd, and R. N. Randall. 1941. Survey of 1941/North Park, Jackson County, Moffat County, including comparative data of previous seasons. Colo. Game and Fish Dep., Sage Grouse Surv., Vol. 3. 31pp. . Kinney, C. R., and J. Sugihara. 1943. Constituents of Artemisia tridentata. (II) J. Org. Chem. 8:290-294. Klebenow, D. A. 1969. Sage grouse nesting and brood habitat in Idaho. J. Wildl. Manage. 33:649-662. McArthur, E. D., A. C. Blauer, A. P. Plummer, and R. Stevens. 1979. Characteristics and hybridization of important intermountain shrubs. III. Sunflower family. U.S. Dep. Agric., For. Servo Res. Pap. INT-220. 82pp. Mi lchunas, D. G., M. I. Dyer, O. C. Wallmo, and D. E. Johnson. 1978. In-vitrq/in-vivo relationships of Colorado mule deer forages. Colo. Div~Wil(ff. Spec. Rep. 43. 44pp. Miller, G. R. 1968. Evidence for selective feeding on fertilized plots by red grouse, hares, and rabbits. J. Wildl. Manage. 32:849-853. , A. Watson, and D. Jenkins. 1970. ----:-lations to experimental improvement of in A. Watson, ed. Animal populations resources. Br. Ecol. Soc. Symp. 10. and Edinburgh, U.K. Responses of red grouse poputheir food. Pages 323-335 in relation to their food Blackwell Sci. Publ., Oxford Moss, R. 1972. Food selection by red grouse (Lagopus lagopus scoticus (Lath.)) in relation to chemical composition. J. Anim. Ecol. 41: 411-428. , and A. Watson. 1980. Inherent changes in the aggressive behavior ----.". of a fluctuating red grouse (Lagopus lagopus scoticus) population. Ardea 68:113-119. , G. R. Miller, and ---captive red grouse in Eco 1. 9: 771 -78 1. S. E. Allen. 1972. Selection of heather by relation to the age of the plant. J. Appl. �176 Patterson, R. L. 1952. Colo. 341pp. The sage grouse in Wyoming. Sage Books, Denver, Petersen, B. E. 1980. Breeding and nesting ecology of female sage grouse in North Park, Colorado. M.S. Thesis, Colo. State Univ., Fort Coll ins. 86pp. Pyrah, D. B. 1959. Sage grouse population trend and trapping study. Wyo. Game and Fish Comm., Job Compl. Rep., Fed. Aid Proj. W-50-R-8, 27pp. J969. Habitat selection and food habits studies of sage grouse in Montana. Proc. West. States Sage Grouse Workshop 6:168-173. Rasmussen, D. I., and L. A. Griner. 1938. Life history and management studies of the sage grouse in Utah, with special reference to nesting and feeding habits. Trans. North Am. Wildl. Conf. 3:852-864. Schoenberg, T. J. 1980. Potential impacts of strip mining on sage grouse movements and habitat use. Colo. Div. Wildl. Prog. Rep., Fed. Aid Proj. W-37-R, Work Plan 3, Job 12. Pp. 245-264. 1981. Potential impacts of strip mining on sage grouse movements and habitat use. Colo. Div. Wildl. Prog. Rep., Fed. Aid Proj. W-37-R, Work Plan 3, Job 12. Pp. 243-264. 1982. Sage grouse movements and habitat selection in North Park, Colorado. M.S. Thesis, Colo. State Univ., Fort Collins. 86pp. Sheehy, D. P. 1975. deer and sheep. Relative palatability of seven Artemisia taxa to mule M.S. Thesis, Oreg. State Univ., Corvallis. 147pp. Smith, E. L. 1966. Soil vegetation relationships of some Artemisia types in North Park, Colorado. Ph.D. Diss., Colo. State Univ., Fort Coll ins. 203pp. Stevens, R., and E. D. McArthur. 1974. tification of some sagebrush taxa. A simple field technique for idenJ. Range Manage. 27:325-326. Thomas, V. G., and R. Popko. 1981. Fat and protein reserves of wintering and prebreeding rock ptarmigan from south Hudson Bay. Can. J. Zool. 59:1205-1211. , H. G. Lumsden, and D. H. Price. ---metabolism of ruffed grouse (Bonasa to energy reserves. 1975. Aspects of the winter umbellus) with special reference Can. J. Zool. 53:434-440. United States Congress. 1977. Public law 95-87, surface mInIng and rehabil itation act. U.S. Stat. at Large, Vol. 91, Stat. 491, Para. 19, Sec. 515. �177 Van Soest, P. J. 1963~. Use of detergents in the analysis of fibrous feeds. II. A rapid method for the determination of fiber and lignin. J. Assoc. Off. Agric. Chern. 46:829-835. 1963b. Use of detergents in the analysis of fibrous feeds. I. Preparation of fiber residues in low nitrogen content. J. Assoc. Off. Agric. Chern. 46:825-829. 1967. Use of detergents in the analysis of fibrous feeds. IV. Determination of plant cell-wall constituents. J. Assoc. Off. Agric. Chern. 50:50-55. Wallestad, R. 0., J. G. Peterson, and R. L. Eng. 1975. Foods of adult sage grouse in central Montana. J. Wildl. Manage. 39:628-630. Watson, A., and P. J. OIHare. 1973. Experiments to increase red grouse stocks and improve the Irish bogland env!ronment. BioI. Conserv. 5:41-44. , and 1979. Red grouse populations on experimentally -----t-reated and untreated Irish bog. J. Appl. Ecol. 16:433-452. , R. Moss, ----:-lizer on red J. Phillips, and R. Parr. 1977. The effect of fertigrouse stocks on Scottish moors grazed by sheep, cattle and deer. Pages 193-212 in P. Pesson, compiler. Ecologie du petit gerbier and amenagement des chasses. Gauthier-Nillars. Paris. Welch, S. L., and E. D. McArthur. 1979. Variation in winter levels of crude protein among Artemisia tridentata subspecies grown in a uniform garden. J. Range Manage. 32:467-469. West, G. C., and M. S. Meng. 1968. Seasonal changes in body weight and fat and the relation of fatty acid composition to diet in the willow ptarmigan. Wilson Bull. 80:426-444. Winward, A. H. 1980. Taxonomy and ecology of sagebrush in Oregon. Oreg. State Univ. Agric. Exp. Stn. Bull. 642. 15pp. , and E. W. Tisdale. 1969. A ---sagebrush identification. Univ. Exp. Stn. Note 11. simplified chemical method for Idaho For., Wildl., and Range 2pp. Young, A. 1965. A chemical study of the taxonomy of section tridentatae of the genus Artemisia. Wyo. Range Manage. 198:2-9 . Prepared by I .J... /1· I fv;z,vt4./1 L. ;~-vu~ ~ Thomas E. Remi ngton I Graduate Research Ass~stant Approved by _~~~~~. 7?-·-~!,...:;!....:::..;._ Clait E. Braun Wildlife Research Leader _ ��Apri I 1982 179 JOB PROGRESS REPORT State of Colorado Project No. W-37-R-35 Work Plan No. Job Ti tIe: 3 Job No. 14 Dispersal and Recruitment of Chick Sage Grouse Period Covered: Personnel: Game Bird Survey 1 April 1981 - 31 March 1982 Clait Braun, Charles Brown, Peter Dunn, Howard Funk, Ken Giesen, Tom Olsen, Colorado Division of Wildlife; Ronald Ryder, Colorado State University. ABSTRACT A study of juvenile sage grouse (Centrocercus urophasianus) dispersal and recruitment was initiated in May 1981 on Cold Spring Mountain, Moffat County, Colorado. This report describes research conducted during April 1981 - March 1982. Brood observations during July indicated that largest numbers of juvenile sage grouse occurred near the eastern end of Cold Spring Mountain and that average brood size (4.5 chicks/hen, N = 51 broods) was only slightly lower than previous averages (4.7 - 5.3 chicks/hen, 197680 range of yearly averages) for Moffat County. Sage grouse bandings totaled 201 birds (158 juveniles). Thirteen solar-powered radio transmitters were placed on 15 juvenile sage grouse (7 females, 8 males) between 14 July and 27 August. Signals from the last transmitter were lost after 21 January 1982. Movements of radio-marked juveniles were only available for 131 locations from 14 July to 3 October because of loss of radios. There were no differences (p > 0.05) between distances or directions moved by males and females. Seventy-four percent of the male relocations and 84% of the female relocations were within 2 km of each bird's capture site. Movements of birds seemed to parallel the northwest to southeast orientation of Cold Spring Mountain. Late summer movements of juvenile grouse on Cold Spring Mountain were not characterized by rapid, synchronous movements by most members of the population. Hunter band recoveries support this conclusion. Mean canopy cover of sagebrush (Artemisia spp.), grasses, and forbs at 41 locations of radio-marked sage grouse from 14 July to 31 August did not differ between males and females (p > 0.05). Sagebrush canopy cover at grouse locations did not increase-from 14 July to 31 August (p > 0.05). Mean height of the plant each radio-marked grouse was found next to was higher than the average height of all plants on corresponding transects. Sage grouse may be selecting taller than average sagebrush. Habitat use by juvenile grouse was similar to other areas in the west. A habitat type map was prepared from aerial photographs. ��181 DISPERSAL AND RECRUITMENT OF CHICK SAGE GROUSE Peter O. Dunn In the western United States, disturbance of sagebrush rangeland by agricultural and energy development may adversely affect species dependent upon sagebrush (Braun et al. 1976). Since sage grouse are dependent upon sagebrush for food and cover, local populations are indicators of sagebrush range quality. Because development and disturbance of western rangelands are accelerating, techniques to mitigate impacts of sagebrush alterations on sage grouse populations will become increasingly useful. Research is being conducted in Colorado and other western states on mitigating adverse impacts of energy-related developments on sage grouse. Although knowledge of sage grouse habitat requirements is generally adequate, there is no information concerning how local populations are formed or maintained. It is not known if young sage grouse produced in 1 area contribute to the maintenance of leks (strutting grounds) in that or some distant area. Without knowledge of dispersal patterns, it will be difficult to predict the impacts of habitat disturbance on breeding populations of sage grouse. Studies of avian dispersal are few, most studies actually investigate fall movements. Among tetraonids, Bendel 1 (1972) noted the paucity of research from fall to spring to assess the importance of dispersal to the number of breeders. No studies have been made of tetraonid dispersal from fall to spring with the exception of red grouse (Lagopus lagopus scoticus) (Moss and ·Watson 1~80, Watson and Moss 1980), spruce grouse (Dendragapus _c~~nsJ? franklini) (Keppie 1979), and greater prairie chickens (Tympanuchus~ pinnatus) (Bowman and Robel 1977). Studies of movements of ruffed grouse (Bonasa umbellus) (Chambers and Sharp 1958, Godfrey and Marshall 1969), sage grouse (Wallestad 1971}, blue grouse (Dendragapus obscurus) (Zwickel et al. 1968), and white-tai led ptarmigan (Lagopus leucurus) (Giesen 1977) have been 1imited to fall and winter. To better understand the importance of dispersal on sage grouse lek formation and maintenance, this research was initiated on Cold Spring Mountain, northwestern Moffat County, Colorado. P. N. OBJECTIVES The goal of this study is the identification of juvenile sage grouse dispersal and recruitment patterns, and distances moved from natal to breeding areas the following spring. SEGMENT OBJECTIVES 1. Review available literature concerning dispersal and recruitment of chick tetraonids and closely related species, especially those that use leks for breeding. Literature will also be reviewed concerning movements or grouse and techniques for investigating bird dispersal. �182 2. Prepare vegetation map of Cold Spring Mountain delineating sagebrush, aspen, wet meadow, and conifer habitats. Habitat type mapping will be done on aerial photographs. 3. Delineate known breeding sites and summer concentration areas used by sage grouse on topographic maps of Cold Spring Mountain. 4. Trap (drive traps, spotlighting and use of long-handled nets) and band (aluminum and color-coded plastic bands) at least 300 chick sage grouse on Cold Spring Mountain. 5. Place and maintain radios (poncho) on at least 12 chick sage grouse during the August-April interval. 6. Periodically relocate radio-marked chick sage grouse to ascertain brood movements to summer use areas and chick dispersal patterns from summer use areas until recruited the following spring on breeding areas. 7. Collect and evaluate band recoveries of marked sage grouse harvested during hunting seasons or found dead. Harvest data will be obtained through field checks of hunters and volunteer wing collection stations. 8. Plot all movement data (reobservation, radio-locations) on overlays of the habitat type map and map of breeding and summer areas to ascertain direction, distance, and patterns of movements in relation to habitat type. 9. Compile data, analyze results, and prepare progress report. DESCRIPTION OF THE STUDY AREA The study area is centered on Cold Spring Mountain, northwestern Moffat County, Colorado (Fig. 1). The sage grouse population on Cold Spring Mountain has been studied from 1978 through 1981. From 150 to over 300 chicks/year have been banded with no recoveries away from the surrounding 160 km2. Counts of male sage grouse attending known leks on and surrounding Cold Spring Mountain have been conducted yearly since 1978. The winter range of sage grouse breeding on Cold Spring Mountain is unknown, therefore, precise study area boundaries have not been established. Cold Spring Mountain is bordered by Diamond Peak and Middle Mountain on the north, on the west by the O-Wi-Yu-Kuts Mountains, on the east by Vermillion Bluff, and on the south by Browns Park. Elevation ranges from approximately 1,800 m in Browns Park to over 3,100 m on Diamond Peak. Cold Spring Mountain, a northward sloping plateau over 2,800 m in elevation, is approximately 24 km long. Vegetation on Cold Spring Mountain is primarily sagebrush rangeland with interspersed meadows and aspen (Populus spp.) stands. Detailed geographic, geologic, vegetational, and climatic features of the study area will be presented in the final report. �IV.) o 5 KILOMETERS Fig. 1. Cold Spring Mountain study area, northwestern Colorado. Contour interval is 1,000 feet. COLORADO Moffat County, �184 METHODS AND MATERIALS Literature was reviewed concerning dispersal and" recruitment of avian species, especially tetraonids. Capture and marking techniques and vegetation analyses were also reviewed for incorporation into a study plan. Juvenile sage grouse were captured for individual marking during July and August 1981. Drive traps (after Tomlinson 1963), a bumper-mounted cannon net, and spot-lighting with the use of long-handled nets (Braun and Beck 1976) were used to capture sage grouse. Classification of sex and age categories of birds captured was by wing molt and primary length (Beck et al. 1975). Birds were weighed on spring scales to the nearest 10 g. All captured sage grouse were marked with serially-numbered aluminum leg bands and colored plastic bands. Color combinations indicated 1 of 4 capture zones on Cold Spring Mountain. Thirteen solar-powered radio transmitters (164 MHz) attached to poncho-type markers (Amstrup 1980) were placed on juvenile sage grouse during July and August 1981~ Radio-marked birds were relocated 3 times weekly during July and August and periodically during September 1981 through February 1982. A 24 channel, 164 MHz radio receiver manufactured by Wildlife Materials, Carbondale, Illinois and a 3-element yagi antenna were used to relocate radio-marked grouse. Two yagi antennas mounted on a Colorado Division of Wildlife aircraft were used to radio-track birds from the air on 3 November 1981. Radio relocations made on the ground were determined by triangulation at close range (approximately 50 m) or by flushing birds. Bird locations were plotted on 7.5 minute series U.S. Geological Survey topographic maps using Universal Transverse Mercator (UTM) coordinates. Vegetation at sites of radio-marked grouse was measured using a modification of Canfield's (1941) line intercept method and Jones' (1968) grouse cover board. Two 10-m transects were measured at each site. Transects were oriented"along North-South and East-West axes with the bird location (hen for broods) as the center of the transect. Vegetation measurements included percent canopy cover of foliated and unfoliated sagebrush, forbs, grasses, and bare ground; height of each sagebrush plant intercepting the transect (weighted by its width perpendicular to the transect, [McDonald 1980]), and 'horizontal cover at transect center from 10-m away. A habitat type map of Cold Spring Mountain was prepared by superimposing cover types (sagebrush, aspen, wet meadow, and pinyon-juniper) on a base map prepared from 7.5 minute series U.S. Geological Survey topographic maps. Cover types were transferred from aerial photographs with a Baush and Lomb zoom transfer scope. During July, distinct broods were counted to identify brood concentration areas and determine average brood size. Brood counts were made during mornings and evenings along roads in the study area. �185 Bands and radio transmitters we~e recovered from harvested sage grouse at the Cold Spring Mountain check station (12-13 Sep 1981) and the Maybell wing barrel (12-27 Sep 1981). Harvest locations of banded birds were determined from hunter reports. Movement data with 2 levels were analyzed with chi-square and the MannWhitney U test, while data with> 2 levels were analyzed with Friedman's 2-way ANOVA. Movement directions were analyzed with Rayleigh's test and Mardia's linear-circular correlation coefficient (Batschelet 1978). RESULTS AND DISCUSSION Literature Review and Study Plan Review of selected literature on dispersal and study methods began 18 May and a completed study plan was submitted on 17 September. There is a paucity of information on the mechanisms and causes of avian dispersal and no information on sage grouse dispersal, except late summer and early fall movements. A complete discussion of the causes and consequences of dispersal will be included in the final report. Brood Counts Brood observations indicated that largest numbers of juvenile sage grouse occurred near the eastern end of Cold Spring Mountain and that average brood size was only slightly lower than previous averages for I'-\offat County. Of 2R8 chicks classified, approximately 74% were east of Willow Spring. Broods were most numerous in the early morning at Arthur's Reservoir and Swede Spring. By 24 July it was not possible to determine distinct broods due to mixing and formation of groups of more than 1 brood. After 5 August, groups of 50 or more grouse regularly occurred at Coyote and" Cold springs, while use of other meadows and springs decl ined. Average brood size on Cold Spring Mountain was 4.5 chicks per hen (N = 51 broods); in Moffat County during 1976-80, average yearly brood-size varied from 4.7 to 5.3 chicks/hen (Braun 1981). Banding Sage grouse bandings totaled 201 birds of which 158 were juveniles (Table 1). Ninety-four birds were banded during July and 107 during August. Most birds (104) were caught at night using long-handled nets and spotlights, however, this was the least efficient capture method (Table 2). Drive traps were most efficient, although they resulted in the fewest grouse being captured. Improper positioning of traps and inexperience driving sage grouse may have resulted in low numbers of birds caught with drive traps. Three mortalities occurred during cannon-netting and 1 during spot-l ighting. �186 a Table 1. Sage grouse bandings , Cold Spring Mountain, Moffat County, Colorado, 13 July - 28 August 1981. Age class Male Sex Female 11+ 2 2+ Totals 85 5 0 6 96 70 17 1 14 102 Unknown Totals 158 22 1 20 201 3 aDoes not include recaptures or mortalities. . a Table 2. Efficiency (birds/hour) of sage grouse capture methods, Cold Spring Mountain, Moffat County, Colorado, 13 July - 28 August 1981. Captures Spot-lighting Cannon-net Drive Trap Totals 104 83 13 200 Hours Efficiency 107.5 53.5 6.0 167.0 0.97 1.55 2.17 1.20 aOoes not include recaptures, mortalities, a noose pole in captures column. or 1 bird caught with Twenty-one birds were recaptured during July and August (Table 3). Twice as many juveniles were recaptured as yearlings and adults; females of all age classes were caught twice as often as males. Juveniles were probably recaptured more often than other age classes because of differential vulnerability to trapping and their larger numbers. Approximately equal numbers of males and females (96 and 102, respectively) were banded, so it is not immediately obvious why more females were recaptured. Only 1 bird banded in 1981 was recaptured farther than 2 km from the original banding site. A juvenile male was banded on Cold Spring Mountain summit 19 July and recaptured 12 August, 4.9 km east-northeast at Swede Spring. �187 Table 3. Sage grouse recaptures, Cold Spring Mountain, Moffat County, Colorado, 13 July - 28 August 1981. Age class at time of recapture 11+ 2 2+ Totals Sex Male Female Totals 8 2 1 3 14 14 2 1 4 21 6 1 7 Radiotelemetry Thirteen solar-powered radio transmitters were placed on 15 sage grouse (7 females, 8 males) between 14 July and 27 August (Table 4). Two radios from birds killed by predators were recovered and placed on male chicks. Transmitters and ponchos were 2.3 and 3.4% of average body weights at time of capture for radio-marked males and females, respectively. During reobservations from 14 July 1981 to 21 January 1982, poncho-mounted transmitters did not noticeably affect behavior. Two radio-marked birds recaptured 1 and 6 days after radio-marking appeared normal. The bird recaptured after 6 days was killed by a predator 12-18 days after radio attachment. Five radios were recovered from birds killed by predators. Maximum signal range was at least 7.5 km. Eleven Wildl ife Materials radios began transmitting 15 minutes before sunrise and stopped 15 minutes after sunset, depending on weather and cover. Two radios manufactured by Advanced Telemetry Systems (Bethel, Minn.) transmitted up to 1 hour after sunset. Sage Grouse Movements Radio-marked grouse Movements of radio-marked grouse were analyzed from 131 locations during 14 July to 3 October 1981 (Table 5). Thirteen relocations between 3 October 1981 and 31 March 1982 were excluded because of the small sample size. No radio-marked grouse (out of a possible 7) were relocated on Cold Spring Mountain during 7-10 February. In addition, no unmarked birds, tracks or droppings were seen on Cold Spring Mountain. The absence of sage grouse was probably due to movement of birds from areas where sagebrush was covered by snow. Analysis of distances and directions moved by male and female chicks from July to October 1981 revealed no differences (Mann-Whitney U, and X2, respectively, P > 0.05). Seventy-four percent of the radio~marked-male relocations and 84% of the female relocations were within 2 km of each bird's capture site during 14 July - 3 October (Fig. 2). There were no differences among distances moved from capture sites by all birds when tested by monthly periods, however, movements increased after 8 August when analyzed by bi-weekly periods (Friedman 2-way ANOVA, P < 0.01) (Table 5, Fig. 3). There were no differences in movement rate (km!day) �Table 4. Sage grouse 28 March 1982. radiotelemetry data, Cold Spring Mountain, Sex Capture date Date of last locat ion Maximum distance moveda(km} 486 785 M M 14 Jul 14 Jul 5 Sep 14 Nov 1.9 8.5 687 438 ( I) 501 820 ( I) 585 712 426 465 636 668 657 820(11) 438(11) M M M 14 16 21 23 26 27 3 5 5 10 12 25 27 3 26 21 11 13 14 12 3 12 12 12 27 Transmitter F F F M F F F F M M Jul Jul Jul Jul Jul Jul Aug Aug Aug Aug Aug Aug Aug aMaximum straightline minimum value. bR epresents . . a mInImum Oct Aug Jan Aug Sep Nov Aug Oct Sep Sep Sep Aug 5 Sep distance va Iue. 2.5 1.1 7.7 1.2 5.7 15.9 4.7 2.3 1.7 1.7 1.5 1.4 0.5 between Date of maximum movement Moffat County, Transmitter 1ife (days)b 12-24 Aug 26 Sep-7 Nov 53 123 11-17 19-24 6-26 4-11 13-19 10-31 6-10 12-19 19-24 19-24 12-16 25-27 27 Aug-5 81 41 184 18 49 110 9 69 38 34 31 2 279 consecutive Aug Aug Sep Aug Aug Oct Aug Sep Aug Aug Aug Aug Sep radio locations. Actually Colorado, 13 July - Remarks Killed by predator Lost signal (intermittent before loss) Lost signal Killed by predator Lost signal Killed by predator Killed by hunter Killed by predator Lost signal Killed by predator Killed by hunter Killed by hunter Lost signal Lost signal Killed by predator (recovered in Apr 82) represents a C» C» �Table 5. Movements (km) of radio-marked juvenile County, Colorado, 14 July - 3 October 1981. Transmitter 14-25 Jul Sex x - 785 M 0.48 501 M 0.54 712 F 687 M 820 (1) F 820 (11) M 438 (1) M 438(11) M 657 F 426 M 486 M 585 F 465 26 JuJ-8 AU9 SO N 0.07 3 1 1 0.67 9-22 Aug x SO N 1. 12 0.52 7 0.67 0.14 0.98 a sites , Cold Spring Mountain, sage grouse from capture x SO N - 1.81 0.83 3 5 1.29 0.34 0.27 5 1.80 1.05 0.53 3 2.22 1. 18 0.26 5 1.25 x 20 Sep-3 Oct 6-19 See 23 AUlr5 Sep x SO N - 0.90 1. 16 2 3 2.26 2.30 2 1.47 1 5. 12 0.31 3 0.75 1 0.94 1 1. 18 1. 18 3 2.08 N - N - 0.88 2 1.40 0.48 1 1.06 0.34 7 2.04 4 0.07 0.90 0.01 0.47 0.71 0.84 1.00 0.46 3 1 5.45 0.16 3 2 _. 1 1..0 co 7 1.84 0.33 3 3.37 0.03 3 1.29 0.43 6 2.60 2.71 3 6.23 0.53 2 5.23 0.02 2 F 0.66 0.22 4 0.98 0.03 2 1.72 1.64 2 636 F 0.65 o. 18 4 0.77 0.40 2 0.81 668 F 1.01 0.114 3 0.98 0.56 2 0.69 0.20 2 0.63 0.92 0.49 3 9 All females aMaximum SO 5.31 0.37 All birds x SO 1.38 A 11 ma Jes Moffat 0.63 0.49 straightline 9 2.78 0.43 2 1 5. 19 0.63 3 1.07 0.43 30 2.42 1.47 19 1.85 1.29 12 1.47 1. 16 0.34 16 1.23 1. 13 21 2.98 2.38 9 2.13 2.05 8 2.25 0.975 3 1.1 0.40 46 1.79 1.42 40 1.95 1.78 21 2.05 1.93 9 3.72 1.77 distance from the respective capture site. Actually represents a minimum 6 value. �190 26 DC MALES II = FEMALES 40 - 30 ~ ~ >(.) Z LJJ :::) 0 20 LJJ 0: u, '. 10 >2-3 KILOMETERS >3-4 >4-5 >5-6 > 6-7 FROM CAPTURE SITES Fig. 2. Frequency (%) of distances moved (km) from capture sites of radio-marked sage grouse on Cold Spring Mountain, Colorado, JulyOctober 1981. Numbers above vertical bars are sample sizes. �191 6 -:e "' D = MALES IIJ = FEMALES 5 ~ C/) LIJ tCf) LIJ 4 a:: :::) ••• Q. <t 0 :e 3 9 19 0 a:: B ~ LIJ (.) z ~ 2 " J2 C/) Q 21 ~ 11- 16 -B II- , 30 9 -B 12-25 JUL 26 JUL- 9-22 8 AUG AUG TIME 23 AUG,5SEP 6 SEP3 OCT PERIODS Fig. 3. Distances (km) from capture sites and time period of radiomarked sage grouse movements on Cold Spring Mountain, Colorado, JulyOctober 1981. Horizontal bars are means; vertical bars are 1 SE above and below the mean. Numbers above vertical bars are sample sizes. �192 among monthly or biweekly periods (Friedman 2-way ANOVA, P > 0.05). All radio-marked grouse moved northwest from capture sites more often than expected and southwest less often than expected (X2, P = 0.03). More northwest movements and fewer southwest movements-may-be due to the northwest to southeast orientation of Cold Spring Mountain and its steep (slope = 18%) southwest face. Among radio-marked sage grouse with 6 or more relocations (N = 11), movements of all but 2 birds had a non-random orientation, or mean direction, from capture sites (Rayleigh's test, P < 0.05). There was no mean direction, however, when individual means of all birds were combined (Mardia's linear-circular correlation coefficient [Batschelet 1978], t > 0.05) (Fig. 4). Band recoveries Sage grouse band recoveries from 1978-80 indicate that birds do not move from Cold Spring Mountain before the September hunting season. Of 156 recoveries of birds banded on Cold Spring Mountain or Gee Flats, only 6 were not recovered on Cold Spring Mountain. These results may, however, be biased by uneven hunter distribution. Late summer movements of juvenile sage grouse on Cold Spring Mountain were not characterized by rapid, synchronized movements by most members of the population, as some studies of ruffed grouse (Godfrey and Marshall 1969) and other populations 0'; sage grouse (Dalke et al. 1963) would indicate. Instead, sage grouse on Cold Spring Mountain may be dispersing at a constant rate throughout September and October, similar to greater prairie chickens (Bowman and Robel 1977). The lack of hunter band recoveries from areas away from Cold Spring Mountain supports this conclusion (Dunn 1982). However, it is also possible that because of few radio relocations in September (N = 15), synchronized, long-distance movements were not detected. Habitat Use Mean canopy cover of sagebrush (both foliated and unfoliated), grass, and forbs at 41 radio-marked sage grouse locations during July and August did not differ between males and females (t test, P > 0.05) (Table 6). Mean sagebrush canopy cover did not increase at grouse locations from 14 July to 31 August (r = -0.02, P > 0.05). Mean height of the plant each radiomarked grouse was found next to was higher than the average height of all plants on corresponding transects (paired t test, P = 0.05). Sage grouse may be selecting for taller than average sagebrush-cover during brood rearing as well as for nest sites as suggested by other investigators (Wallestad and Pyrah 1974, Petersen 1980). Habitat use by juvenile sage grouse on Cold Spring Mountain was similar to other areas in the West (Table 7). The high variability in reported forb cover may be due to sampling vegetation when broods were mainly in either sagebrush or meadows. Some broods may remain in sagebrush throughout the summer, where forb cover is lower, while others may use meadows, where forb cover is likely to be higher (Wallestad 1971). On Cold Spring Mountain, radio-marked grouse were infrequently observed in meadows and vegetation measurements were taken on only 2 of those occasions. �193 N s Fig. 4. Mean angle and distance moved (km) from capture sites of all radio-marked sage grouse on Cold Spring Mountain, Colorado, JulyOctober 1981. Dotted 1ine indicates mean of the 11 vectors (~> 0.05). �Table 6. Canopy cover (%) and sagebrush grouse on Cold Spring Mountain, Colorado, previous studies of brood habitat. height (cm) at relocations of radio-marked juvenile sage 1981, compared with results of 14 July - 31 August Canopy cover Grass Sagebrush - - Forbs - Sagebrush height 25 SO N Source Locat ion 25 SO N 25 SO N 25 SO N This study Colorado 17 12 41 51 27 41 5 11 41 37 16 39 Schoenberg (1982) Colorado 27 16 23 52 24 9 7 46 23 26 12 23 Wallestad (1971) Montana c Martin Montana 12- 21 b 1ge Peterson (1970) Montana 6-12b Klebenow (1969) Idaho (1970) aSeven percent b c 12f in sagebrush June to September 69 47 d 51,47 60 a d b 69 17,27 69 18-25 69 137 40 137 33 47 91 33 91 b 41-51 '.D 91 4 3 areas; 41%, later, when broods moved to meadows. range. No value reported. d1968 and 1969 yearly means for June to September, respectively. eEighty-eight percent of broods 6 weeks old or less that were found were in areas with an average sagebrush canopy cover of 14%. fAll except 3 of 98 broods found were in areas of < 31% shrub cover. Twelve percent figure from addition of canopy cover means for big sagebrush and three-tip sagebrush (Artemisia tripartita). .z::- �195 a Table 7. Canopy cover (%), horizontal cover (%), and height of sagebrush (cm) at relocations of radio-marked juvenile sage grouse on Cold Spring Mountain, Moffat County, Colorado, 14 July-31 August 1981. Males All birds Females x SO N x SO N x SO N 12 11 23 10 10 18 11 11 41 5 6 23 8 10 18 6 8 41 Grass 49 29 23 54 26 18 51 27 41 Forbs 7 14 23 3 6 18 5 11 41 18 22 23 19 19 18 19 20 41 Juniper 3 10 23 0 0 18 3 10 41 Aspen 2 7 23 3 11 18 2 8 41 Rock 2 6 23 3 18 2 5 41 Log 4 7 23 3 18 2 6 41 Hor izonta I cover 36 28 23 22 18 18 30 25 41 Height of fol iated and unfoliated sagebrush 41 16 21 32 14 18 37 16 39 Canopy cover b Sagebrush Fol iated Unfo! iated Bare ground a Measured with Jones' (1968) cover board at center of transects from 10-m away. bSnowberry (Symphoricarpos spp.), rabbitbrush (Chrysothamnus spp.), and gooseberry (Ribes spp.) accounted for < 2% of canopy cover and were not included. �196 Habitat Type Map The habitat map has been cover-typed from aerial photographs and will be completed once cover type areas are filled in with stippling, roads are added, and a legend completed. Sage grouse movements in relation to habitat type will continue to be analyzed from direct measurement at relocation sites, rather than plotting on the habitat type map (Segment objective 8). This change is due to an inherent lack of information in the map concerning height and percent canopy cover of the different vegetation types (sagebrush, grasses, and forbs). The habitat map will be used to place radio relocations on topographic maps and to examine broad trends in movements of radio-marked birds. LITERATURE CITED Amstrup, S. C. 1980. 44:214-217. A radio collar for game birds. J. Wildl. Manage. Batschelet, E. 1978. Second order statistical analysis of directions. Pages 3-24 in K. Schmidt-Koenig and W. T. Keeton, eds. Animal migration, navigation, and homing. Springer-Verlag, Berlin, Germany. Beck, T. D. I., R. B. Gill, and C. E. Braun. 1975. Sex and age determination of sage grouse from wing characteristics. Colo. Div. Wildl. Game Inf. Leafl. 49 (rev ised) . 4pp. Bendell, J. F. 1972. Concluding remarks on the tetraonid symposium. Proc. Int. Ornitho1. Congr. 15:170-177. Bowman, T. J., and R. J. Robel. 1977. Brood break-up, dispersal, mobility, and mortality of juvenile prairie chickens. J. Wildl. Manage. 41:27-34. Braun, C. E. 1981. Vulnerability and population characteristics of sage grouse in Moffat County. Final Rep., Colo. Div. Wildl. Fed. Aid Proj. W-37-R-34, Work Plan 3, Job 11. Pp. 29-73. , and T. D. ---tribution and I. Beck. 1976. Effects of sagebrush control on disabundance of sage grouse. Final Rep., Colo. Div. Wildl. Fed. Aid Proj. W-37-R-29, Work Plan 3, Job 8a. Pp. 21-84. __ ~~' M. F. Baker, R. L. Eng, J. S. Gashweiler, and M. H. Schroeder. 1976. Conservation committee report on effects of alteration of sagebrush communities on associated avifauna. Wilson Bull. 88:165171. Canfield, R. H. 1941. Appl ication of the line interception method in sampling range vegetation. J. For. 39:388-394. Chambers, R. E., and W. M. Sharp. population of ruffed grouse. 1958. Movement and dispersal within a J. Wildl. Manage. 22:231-239. Dalke, P. D., D. B. Pyrah, D. C. Stanton, J. E. Crawford, and E. F. Schlatterer. 1963. Ecology, productivity, and management of sage grouse in Idaho. J. Wildl. Manage. 27:811-841. �197 Dunn, P. O. 1982. Dispersal and recruitment of juvenile sage grouse. Colo. Div. Wildl., Jan. Progress Rep., Fed. Aid Proj. W-37-R-35, Work Plan 3, Job 14. Giesen, K. 1977. Mortality and dispersal of juvenile white-tailed ptarmigan. M.S. Thesis. Colo. State Univ., Fort Coll ins. 55pp. Godfrey, G. A., and W. H. Marshall. 1969. Brood break-up and dispersal of ruffed grouse. J. Wildl. Manage. 33:609-620. Jones, R. E. 1968. A board to measure cover used by prairie grouse. J. Wildl. Manage. 32:28-31. Keppie, D. M. 1979. Dispersal, overwinter mortality, and recruitment of spruce grouse. J. Wildl. Manage. 43:717-727. Klebenow, D. A. 1969. Sage grouse nesting and brood habitat in Idaho. J. Wildl. Manage. 33:649-662. Martin, N. S. 1970. Sagebrush control related to habitat and sage grouse occurrence. J. Wildl. Manage. 34:313-320. McDonald, L. L. 1980. Line-intercept sampling of attributes other than coverage and density. J. Wildl. Manage. 44:530-533. Moss, R., and A. Watson. 1980. Inherent changes in the aggressive behavior of a fluctuating red grouse (Lagopus lagopus scoticus) population. Ardea 68:113-119. Petersen, B. 1980. Breeding and nesting ecology of female sage grouse in North Park, Colorado. M.S. Thesis. Colo. State Univ., Fort Collins. 86pp. Peterson, J. G. 1970. The food habits and summer distribution of juvenile sage grouse in central Montana. J. Wildl. Manage. 34:147155. Schoenberg, T. J. 1982. Sage grouse movements and habitat selection in North Park, Colorado. M.S. Thesis. Colo. State Univ., Fort Co 11 ins. 86pp. Tomlinson, R. E. 1963. A method for drive-trapping Wildl. Manage. 27:563-566. dusky grouse. J. \~allestad, R. O. 1971. Summer movements and habitat use by sage grouse broods in central Montana. J. Wildl. Manage. 35:129-136. , and ---central D. Pyrah. 1974. Movement and nesting of sage grouse hens in Montana. J. Wildl. Manage. 38:630-633. Watson, A., and R. Moss. 1980. Advances in our understanding of the population dynamics of red grouse from a recent fluctuation in numbers. Ardea 68:103-111. �198 Zwickel, F. C., I. O. Buss, and J. H. Brigham. 1968. Autumn movements of blue grouse and their relevance to populations and management. J. Wildl. Manage. 32:456-468. Prepa red by _+.~::-:~:=:....;:.;:;~...:::O,-::-"/2,-"-,,,~~~ _ _-::-:-a~~--=-·--::,::z._.__._~:...=,--Clait E. Braun _ 'Peter O. Dunn Graduate Research Assistant Approved by Wildlife Research Leader �April 1982 199 JOB FINAL REPORT State of Colorado --------------------------- Project No. Work Plan No. Job Title: 9 Job No. 6 Response of Blue Grouse to Aspen Silvicultural Period Covered: Personnel: Game Bird Survey W-37-R-35 Practices 1 April 1980 through 31 March 1982. J. Dickerson, M. Ward, D. Ellis, U.S. Forest Service; B. Wilcox, Colorado State Forest Service; T. Beck, C. Braun, B. Cade, M. Crosby, H. Funk, T. Hobbs, R. Hoffman, and D. Miller, Colorado Division of Wildlife. ABSTRACT A suitable study was located on Terror Creek about 11 km north of Paonia, Colorado. Efforts to obtain the desired level of treatment for development of a study plan failed because the U.S. Forest Service proposal for managing quaking aspen (Populus tremuloides) in t~is area did not conform to research needs. Furthermore, economic restraints created by the lack of a market for aspen wood reduced the acreage that commercial operators were willing to cut to below that which any response from the blue grouse (Dendragapus obscurus) population could be accurately measured. Several alternatives.were considered, the most practical being a combination of services including (1) commercial timber sales to local operators, (2) service contracts for removal of unmarketable timber, and (3) use of the Colorado State Forest Service logging crew to cut the additional acreage necessary to attain the desired level of treatment. Pretreatment information was obtained on population densities, brood use, and habitat types on the study area in 1981. No firm commitment of cooperation was obtained from the U.S.F.S., consequently, field work on the project was terminated after August 1981. ��201 RESPONSE OF BLUE GROUSE TO ASPEN SILVICULTURAL PRACTICES Richard W. Hoffman Blue grouse are widely distributed in the mountainous areas of Colorado occupying varied habitats from shrublands to mature coniferous forests. This species is avidly pursued by sport hunters and the blue grouse is a major upland game resource in Colorado. Studies of blue grouse in Middle Park and near Eagle have focused on population dynamics and habitat use. These studies have identified the aspen type as being of major importance for blue grouse breeding and brood rearing activities. Aspen is a subclimax type and covers hundreds of thousands of hectares in Colorado and adjacent states of the central Rocky Mountains. Management of aspen by public land management agencies could have tremendous impacts upon the blue grouse resource. The intent of this study was to identify blue grouse response to aspen management practices. To successfully accomplish this study, it was necessary to have the cooperation and a firm commitment from public land management agencies, primarily the U.S. Forest Service. P. N. OBJECTIVE Prepare a detailed study plan describing specific hypotheses and/or objectives, procedures, schedules, costs, manpower requirements, etc.; and conduct pretreatment evaluation of breeding densities, brood use, and habitat conditions. METHODS Field personnel of the Colorado Division of Wildlife (CDOW), U.S. Forest Service (USFS), and Colorado State Forest Service (CSFS) were contacted regarding potential study areas. Field surveys were conducted in most recommended areas. Some time was spent in consultation with D. C. Bowden, statistician, concerning design of appropriate sampling and analytical procedures necessary to reliably measure the response of a blue grouse population to aspen manipulation. Efforts were initiated to review and assemble literature pertinent to the subject matter of this study. Grouse were located by systematic search with the aid of a pointing dog and by acoustical census (Hoffman 1981). Once located, birds were first observed with binoculars to count numbers present and ascertain sex. Attempts were made with telescoping noose poles to capture any unmarked grouse encountered (Zwickel and Bendell 1967). All grouse captured were weighed, classified by age (Braun 1971, Zwickel and Lance 1966, Redfield and Zwickel 1976) and sex (Braun 1971, Caswell 1954), and banded. Captured grouse older than 6 weeks were individually marked with color combinations of anodized, aluminum butt-end leg bands (size 14) following the scheme described by Gull ion (1965) . �202 Vegetative classification of the area was done in accordance with procedures outlined by Kuchler (1955). This method of vegetation mapping is a derivation of Braun-Blanquet1s (1951) system of floristic and physiognomic description of plant communities. Vegetation maps have been prepared elsewhere in Colorado using this method (Medin 1962, Braun 1969, Hoffman 1981). RESULTS AND DISCUSSION Study Area Field personnel of the Colorado Division of Wildlife, U.S. Forest Service, and Colorado State Forest Service were contacted for suggestions on study areas. General surveys were conducted on 6 potential study areas including Stoner Mesa, Groundhog Mesa, Black Mesa, Terror Creek, Schaefer Creek, and Slater Creek. Five of the 6 areas are in southwestern Colorado; 2 (Stoner and Groundhog mesas) in the San Juan National Forest and 3 (Terror Creek, Schaefer Creek, and Black Mesa) in the Gunnison National Forest. Southwestern Colorado supports the largest, continuous stands of aspen in the state. Baker (1925) estimated that approximately 695,109 hectares of aspen cover types occur on national forest lands in Colorado of which 65% (451,821 ha) was in the southwest portion of the state. The only area investigated outside of southwestern Colorado was Slater Creek on the Routt National Forest in northwestern Colorado. There are few other areas where large scale aspen manipulation programs are currently in progress or where programs will be implemented in the near future. Most timber sales and wildlife habitat improvement projects involving aspen are done on such a small scale as to preclude any studies to measure impacts on wildl ife populations. The problem stems from the lack of a commercial market for aspen wood accompanied by rising production costs which have reduced the area that can be economically manipulated. As a consequence, commercial operators are only interested in harvesting small areas and economic restraints have limited the amount of noncommercial timber that can be managed solely for wildlife habitat improvement. The Terror Creek management unit was considered the most promising area for conducting this study. Stoner Mesa, Groundhog Mesa, Black Mesa, and Slater Creek have already been subjected to aspen manipulation; consequently, no pretreatment evaluation could be performed on these areas. In addition, Stoner Mesa, Groundhog Mesa, and Black Mesa are not readily accessible for conducting breeding surveys in early spring. Access until late Mayor early June would be by snowmobile. Schaefer Creek fulfilled the study area criterion but the proposal to manage aspen in this area for watershed protection has not progressed past the district level planning stages. It is highly unlikely this project will be implemented in the near future. �203 Terror Creek is about 11 km north of Paonia, Colorado in portions of T125, R91W and T12S, R92W, Delta County, Paonia Ranger District, Gunnison National Forest. The Terror Creek Management Unit is roughly bounded by Big Alder Creek on the north, the Overland ditch on the west, Cunningham Creek on the south, and the Steven's Gulch road on the west. This area encompasses approximately 2,995 hectares and is comprised almost entirely of the following cover types: aspen-tall shrub (2,066 ha), aspen-spruce-fir (450 ha), aspen-tall forb (375 ha), and aspen-riparian (104 ha). More than one-half (58%) of the area has been classified as determinant stands (even-aged), most of which are in the mature and over-mature age classes with virtually no regeneration in the understory. The drainage where treatment was contemplated ranges in elevation from 1,890 to 2,866 m. Access is from the Hubbard Creek and Steven's Gulch roads. Low standard roads exist along east Terror Creek, Cunningham Creek, Betty Park, and along the Curecanti-Hayden Powerline. Four domestic grazing allotments (3 cattle, 1 sheep) are within the proposed unit. USFS Proposed Treatment The Terror Creek Wildlife Habitat Improvement Unit was established during winter 1980. Management plans and an Environmental Assessment Report were prepared by USFS personnel. The plans dealt primarily with aspen and Gambel Oak (Quercus gambelii). Primary objectives for the project in aspen are to regulate the determinant aspen timber to sustained-yield and even distribution of age and size classes and to enhance wildlife habitat by diversification. Photo evaluation and stand mapping was completed before field work began in July 1980. Stage II inventory was completed on the unit during summer 1980. Standard timber information was gathered during this time along with a basic wildlife survey. Aspen cover types were classified as determinant (even-aged) and indeterminant (uneven-aged) and each category was further delineated into productive and nonproductive stands. Productive stands are defined as those capable of producing 1.4 cu m/ha/yr (20 cu ft/acre/yr) of sawtimber material and which are suitable for silvicultural practices. Only the productive, determinant aspen was designated for treatment. This included commercial (net volume/acre> 5,000 bd ft/acre) and cultural (~ 5000 bd ft/acre) treatments. Since the indeterminant aspen supposedly would be self-perpetuating, it was not designated for treatment. - Computed areas of aspen in each category varied (Fig. 1). �204 I\SPEN 2,995 ha Determinant 1,753 ha / Unproductive 305 ha ~ ~ / \ Productive 1,448 ha / Cultural treatment 524 ha\ Indeterminant 1,242 ha Unproductive 149 ha \ Productive 1,093 ha \ Commercial harvest /924 ha Total treatment 1,448 ha Fig. 1. Estimated areas of aspen types based on USFS Stage II Inventory. A separate regulation schedule was established for cultural and commercial treatments. Both treatment types are based on a 60-year rotation with 6-10 year cutting cycles (treatment periods). 1. Cultural Treatment. -- There are 524 hectares designated for cultural treatment. Using a 60-year rotation, the average annual harvest would be 8.7 ha/yr or 87 ha/l0 yrs. All stands designated for cultural treatment were prioritized according to defect percentage. At 1 end of the scale were stands with 0% defect and 0 net volume/ha which were basically seed saps or small poles. These stands were considered least in need of treatment and were placed in the far treatment periods (J.e., 5th and 6th). At the other extreme were stands with 100% defect and o net volume/ha which were severely decadent and in immediate need of treatment. These stands were placed in the first cutting period. The remaining stands were ranked according to defect percentage with the higher percentages rated as higher priority for treatment. Size of the areas were adjusted somewhat to maintain an even flow of cutting at about 87 ha/10 yrs. Aspen Cutting Plan Per iod 1 Hectares to be treated (First 10 years) 94 89 85 85 85 86 2 3 4 5 6 Total 524 �205 Since most of this volume will not be used except possibly as free fuel wood, no attempt will be made to remove the timber. The actual area treated in any 1 year is not binding as long as each unit of treatment is completed within a 10-year period. For instance, the 94 hectares designated for treatment in the first 10 years need not be divided equally among these years, but can be treated in any combination of ways including cutting the entire acreage in 1 year. The only stipulation is that the 94 ha be cut within the 10 years. 2. Commercial Treatments. -- There are 924 hectares designated for commercial treatment. Based on a 60-year rotation, the average annual harvest would be 15.4 ha/yr or 154 ha/l0 yrs. Since the volume in this treatment will be removed for commercial purposes, a transportation system must be developed. The area was divided into 6-10 year cutting periods. Priorities were established on the basis of present wildlife needs, expected transportation system needs, and distribution of treatment units to obtain desired habitat diversity. The area within period 1 would be the most accessible and receive the heaviest wildlife use. Periods 2-6 are increasingly less accessible and of decreasing importance to wildlife. Area to be treated within each period are: Period Hectares 1 2 3 4 5 6 Total 183 135 145 154 151 156 924 For regulation to occur within the 60-year rotation, each 10 year unit must be treated in a 10-year period. However, the order in which each 10-year unit is treated is dynamic and can be changed according to current conditions. For example, if after the first 10-year units (183 ha) are treated, there is a need to change priority area 5 to 2, this can be done without disrupting future age class distribution as long as the integrity of each 10-year unit is maintained. Overcutting in any period would congest the age classes at 1 end of the spectrum and leave a deficiency at the opposite end. Undercutting would extend the time period for regulation beyond a rotation. Unlike the cultural treatments, commercial treatments are dependent upon market conditions and must be imposed on an annual basis to stay even with demand. Pretreatment Habitat Evaluation Investigations Intensive breeding and production surveys were conducted on an area of about 383 ha which was included within that portion of the Terror Creek Management Unit designated for priority treatment. Five major vegetation types were delineated on this area (Fig. 2). Vegetative summary tables (Appendix) were compiled and are presented as supplementary information describing the physiognomy, floristics, and physiography of each vegetation type. �206 TERROR CREEK POPULUS-SYMPHORICARPOSmixed shrub (203 hal QUERCUS-mixed (122 ha) shrub POPULUS-PRUNUSmixed s~27 ha) POPULUS-QUERCUSmixed shrub (23 ha) POPULUS-BETULAr i par i aii(8ii"a) N i SCALE I 0 % Fig. 2. Major vegetation types Creek study area, Colorado. and blue grouse 1 Km territory sites, Terror �207 Breeding Survey The total number of territorial·males recorded during the peak of breeding activities was used as an index to population size. Hens could not be accurately censused during the breeding season because of their greater daily movements, more secretive habits, and lower response rate to recorded calls. Only 11 territorial males (2.9 males/km2) were found within the study area. Densities recorded elsewhere in Colorado have exceeded 10 males/km2 (Hoffman 1981). Location of territories in relation to vegetation types is depicted in Fig. 2. Nine (82%) of 11 territories were associated with the aspenoakbrush interface. The remaining 2 territories were in the oakbrush type. No territories were found solely in association with the aspen types. Of 47 spring observations of blue grouse, only 2 were in continuous aspen types. Both sightings were of females, probably in transit. Lack of vegetative and structural diversity in the nearly 256 ha (67%) of continuous aspen types on the study area is believed responsible for the low density of grouse. Brood Surveys A total of 38 hours was spent searching for broods on the study area between 8-12 July 1981. An additional 42 hours were spent on the study area in midAugust during mapping operations. No broods were observed at either time further suggesting low densities and an avoidance of the continuous aspen types by blue grouse. Study Status and Recommendations Cooperation and a firm commitment of control over the treatments to be imposed were prerequisites to further development of the study plan. However, no commitment was received from the USFS. As of August 1981, their proposal had not progressed beyond the planning stages. Whereas the proposal appeared operational on paper, its practical ity was questionable. There were too many factors which could have interrupted adherence to the cutting schedule; most notably, a continued lull in the aspen market and further restraints on spending by federal agencies. It was emphasized that a study could not be designed around the USFS proposal without control over when, where, and how much aspen was to be cut. Consequently, it was recommended that the USFS abandon their proposal, at least temporarily, and consider a cooperative study. Alternative means of obtaining the desired level of treatment were explored including a combination of (1) commercial timber sales to local operators, (2) service contracts for removal of unmarketable timber, and (3) cooperation of the Colorado State Forest Service in using their logging crew. The USFS was not open to adjusting their proposal. As a result, and because of other economic restraints, field work was discontinued after August 1981 and a recommendation was made to terminate the study. �208 LITERATURE CITED Baker, F. S. 1925. Aspen in the Central Rocky Mountain Region. Dep. Agri. Dep. Bull. 1291. 46pp. U.S. Braun, C. E. 1969. Population dynamics, habitat, and movements of whitetailed ptarmigan in Colorado. Ph.D. Thesis. Colo. State Univ., Fort Co 11 ins. 189pp. 1971. teristics. 4pp. Determination of blue grouse sex and age from wing characColo. Div. Game, Fish and Parks. Game Infor. Leafl. 86. Braun-Blanquet, J. 1951. Germany. pp. 58-66. Caswell, E. B. 18:139. 1954. Pflanzensoziologie. Springer-Verlag, A method for sexing blue grouse. Wien, J. Wildl. Manage. Gull ion, G. W. 1965. Improvements in methods for trapping and marking ruffed grouse. J. Wildl. Manage. 29:109-116. Harrington, H. D. 1964. Inc. Chicago, Ill. Manual of the plants of.Colorado. 666pp. Swallow Press Hoffman, R. W. 1981. Population dynamics and habitat relationships of blue grouse. Colorado Div. Wildl. Job Final Rep. Fed. Aid Proj. W-37-R. April 1981. pp.103-171. Kuchler, A. W. 1955. A comprehensive method of mapping vegetation. Assoc. Am. Geographers 45:404-415. Ann. Medin, D. E. 1962. An ecological investigation of the Cache la Poudre deer herd. Colorado Dep. Game and Fish Job Completion Rep. Fed. Aid Proj. W-105-R. July 1962. Pp. 187-204. Redfield, J. A., and F. C. Zwickel. 1976. Determining the age of young blue grouse: a correction for bias. J. Wildl. Manage. 40:349-351. Zwickel, F. C., and J. F. BendelL J. Wildl. Manage. 31:202-204. 1967. A snare for capturing blue grouse. , and A. N. Lance. 1966. Determining --~J. Wildl. Manage. 30:712-717. Prepared by .: / j;-I' y ,{( I(/({(/ {/ll j/,;/,/ ld./J!u·v~ . Richard W. Hoffman!! Wildlife Researche~ C the age of young blue grouse. �209 APPENDIX Supplementary Tables to Fig. 2 ��211 Physiognomic classification of vegetation. a CAPITAL LETTERS: Herbaceous Woody Vegetation: B: D: E: N: 0: G: H: L: evergreen broadleaf deciduous broadleaf evergreen needleaf deciduous needleleaf without leaves Vegetation: graminoids forbs lichens and mosses SMALL LETTERS: Height: Group I: t: tall; m: medium tall; low; 1: s: z: shrubs; dwarf shrubs; Group II: Group III: e: k: q: Special Features: of vegetation.a Sociability: cover very small Plentiful but less than 1/20 of the area Covering 1/2- 1/4 of the Covering 1/4 - 1/2 of the Covering 1/2- 3/4 of the Covering greater than 3/4 5 a palms bamboos aquatic tree ferns and tuft plants (1955). + Very sparsely present; 4 u: v: w: y: epiphytes lianas succulents cushion plants Cover: 2 3 1 m Maximum height classification 1 25 m 2 m 10-25 m !z-2 m 10 m !z m 1 m Density: aFrom Kuchler Floristic height of trees: height of herbaceous plants: of trees: of herbaceous plants: height of trees: height of herbaceous plants: height: continuous growth interrupted; plants usually do not touch plants scattered singly, or in groves or patches rare, yet conspicuous barren; vegetation largely or entirely absent c: i: p: r: b: j: Minimum Minimum Height Height Maximum Maximum Minimum From Kuchler (1955). 1 2 3 4 area 5 area area of the area Growing singly Grouped or tufted In small patches In extensive patches In great crowds �212 DESCRIPTIVE SUMMARY OF VEGETATION Legal Description: Elevational Range: T12S, R92W, Sections 2500-2866m VEGETATIONAL assification fea tures . and . a 383 ha CLASSIFICATION AND SUPPLEMENTARY Dmc Dsc Hmc ,posure (degrees azimuth) 25, 36; T12S, R91W, Sections Surface Area: Populus-Symphoricarpos mixed shrub YSlognomlc adient TYPE MAP, TERROR CREEK - BLUE GROUSE (%) County: U.S.D.A. Delta 1078-290 FEATURES Populus-Prunus mixed shrub Populus-Quercus mixed shrub Populus-Betula ripa r ian Dsp Dsc Hlp Dmc Dsi Hmc Dli Dsi DzcHmi Dli Dsp Hmc 120-210 175 180 90 4-23 4-25 5-20 5-20 5-15 203 122 27 23 8 FLORISTIC ant species 30, 31 Photo No: Quercus-mixed shrub 90-120 ea (hal Aerial INVESTIGATIONS b I ist Cov Soc CHARACTERISTICS Cov Soc Cov Soc 4 4 3 2 ,+ 3 Cov Soc 3 2 + 2 2 ,ody-Trees: Populus .odv+Sb tremuloides 4-5 4 r ub s : Amelanchier alnifol ia Berberis repens Betula occidental is Chrysothamnus spp. Crataegus spp. Prunus virginiana Quercus gambe II ii Ribes cereum Rosa woods ii Sambucus coerulea Symphoricarpos oreophilus + 2 + 1-2 3 2 3-5 3-4 2 + + 4-5 1 1 4 3-5 2 4 4 3 2 3 + + 3-5 1 2 4 4 5 1 3 3 2 + 3 1 2. + + + 3 3 4 .rbs : Achillea lanulosa Aconitum columbianum Agastache urticifolia Agoseris spp. Allium rub rum Antennaria rosea Agui legia caerulea Arenaria spp. Arnica cordifolia, Artemi s ia spp. Caltha lanceolata Capesella bursa-pastoris Cardamine spp. Ce s t llle j a spp .• Cerastium spp. Chenopodium spp. Cirsium spp. Clatona lanceolata Cleome spp. Crepis spp. Delphinium spp. Epilobium angustifol ia Erigeron spp. Fragaria oval is Frasera speciosa Ga I ium borea Ie Geranium spp. Geum spp. Gil ia aggregata Helenium hoopesii Hel ianthus spp. Heracleum spp. Hydrophyllum capitatum Lathyrus spp. Ligusticum spp. Lupinus spp. Madia glomerata Mentha spp. 1 1 3 1 + + 1 1 2 1 1 2 3 + 2 + 2 1-2 + + 2 1 + I 2 1-2 1-3 1 1 + 1 + 2 + + + + + 1 1 1 2 2 2 4 3 4 + + + 3 + + + 2 2 2 1 1 2 3 1 3 2 I 2 + 1 2 1 3 2 + 1 2 1 3 2 3 + 2 2 �213 DESCRIPTIVE SUMMARY OF VEGETATION Legal Description: Elevational Range: 2500-2866m VEGETATIONAL :Iassification features a ;>hysiognomic oxposure 383 ha CLASSIFICATION AND SUPPLEMENTARY (degrees azimuth) ':'rea(ha) County: U.S.D.A. Delta 1078-290 FEATURES Quercus-mixed shrub Populus-Prunus mixed shrub Populus-Quercus mixed shrub Dsp Osc Hlp Omc Osi Hmc 01 i Osi Ozc Hmi Dmc Osc Hmc ~rad ient (%) INVESTIGATIONS 30, 31 Aerial Photo No: Surface Area: Populus-Symphoricarpos mixed shrub and TYPE MAP, TERROR CREEK - BLUE GROUSE T12S, R92W, Sections 25, 36; T12S, R91W, Sections Populus-~ r ipar ian Dli Dsp Hmc 90-120 120-210 175 180 90 4-23 4-25 5-20 5-20 5-15 203 122 27 23 8 FLORISTIC CHARACTERISTICS ~lant species Fo listb Cov Soc Cov Soc Cov Soc Cov Soc + 2 2 1 1 rbs : (cont inued) Mertensia spp. Mimulus spp. Orthocarpus luteus Osmorhiza spp. Pedicularis spp. Penstemon spp. Polemonium spp. Potentilla spp. Rudbeckia spp. Rumex spp. Saxifraga spp. Senecio spp. Sidalcea candida Smilacina stellata Solidagospp. Taraxacum officinale Thai ictrum spp. Thermopsis divaricarpa Tradescentia spp. Tragopogon dubius Urtica spp. Veratrum californicum Verbascum thapsus Vicia americana Vigulera multiflora Viola spp. Wyethia arizonica 2 + + + + 1 2 + + 2 2 1 2 1 2 1 2 + + 2 2 3 + 1 2 2 + 1 + 2 2 2 2 3 3 1 1 + + + 1 1 2 2 1 1 1 1 2 + 1 + 3 3 4 3 + ;I 1 3 5 3 + 1 2 + 3 3 + 2 1 1 2 1 1 + 1 1 2 2 2 1 2 3 + ~raminoids: Ag~opyron spp. Agrostis spp. Bromus spp. Carex spp. Dactyl is glomerata Elymus spp. Hordeum spp. Mel ica bulbosa Phleum pratense Poa spp. Sitanion hystrix Stipa spp. aAssigned according ~laced at the beginning. blmportant + + 1 1 3 4 1 2 1 3 + 3 4 2 + 2 2 2 + 3 2 2 1 1-2 2 + + 2 + 2 2-3 '-2 2 2 2 2 + 2 + 4 to Kuchler (1955), the physiognomic plant species on the basis of coverage. cCoverage and sociability assigned according 2 2 formula reads left to right with the most conspicuous Plant nomenclature to Kuchler 3 (1955). follows Harrington (1964). type ��215 JOB PROGRESS REPORT State of Colorado --------------------------9 Work Plan No. Job Ti t le : Game Bird Survey W-37-R-35 Project No. Characteristics Job No. 7 and Habitat Preferences of Wintering Populations of Blue Grouse Period Covered: Personnel: 1 April 1981 through 31 March 1982 C. Braun, M. Crosby, J. J. Jeanson, R. Hoffman, S. Steinert, Colorado Division of Wildlife; B. Cade, A. Foster, K. Medve, B. Piske, R. Ryder, Colorado State University~ ABSTRACT Sixty-seven blue grouse (Dendragapus obscurus) were banded and an additional 23 birds were equipped with radio tr-ansmitters at 2 study sites in Middle Park, Colorado. Three banded grouse and 12 instrumented birds provided information on movements between breeding and wintering areas. Mean distance between breeding and wintering areas for 8 grouse migrating from the study areas was 13.3 km (range 3.4-28.0 km). Instrumented birds migrating from Whiteley Peak wintered in the spruce-fir zone in the Rabbit Ears Range. Grouse migrating from Green Mountain wintered in spruce-fir communities adjacent to high peaks (N = 2) or foothills (N = 2) of the Gore Range. Three instrumented females, 1 banded female, and 1 banded male wintered and bred on t~e study areas. Mean winter home range size for 4 instru- . mented females was 7.6 ha (range 1.8-18.7 ha). Sex segregation was not documented during winter. Lone grouse were observed 49 times; 9 were males, 23 were females, and 17 were of unknown sex. Forty flocks were observed ranging in size from 2-12 birds. Both sexes were present in at least 16 flocks. Habitat characteristics were quantified in O.Ol-ha plots at 86 grouse locations on the study areas. Mean tree densities for plots in 3 occupied habitat types at Green Mountain were: 4.7/0.01 ha in Douglas-fir (Pseudotsuga menziesii),5.8/0.01 ha in Douglas-fir/aspen (Populus spp.), and 2.8/0.01 ha for Douglas-fir/juniper (Juniperus spp.). Mean tree densities for plots in 3 occupied habitat types at Whiteley Peak were: 2.7/0.01 ha in open, mixed conifer; 8.1/0.01 ha in dense, mixed conifer; and 3.6/0.01 ha in Douglas-fir/aspen. Tree densities differed among habitat types at Green Mountain and Whiteley Peak, but did not differ between study areas. Dbh of trees at occupied sites on Green Mountain (;~= 25.7 cm) and Whiteley Peak (x = 25.9 em) did not differ, although-canopy heights (GM x = 13.9 m,\.JP x = 17.6 m) did. Occupied sites at both study areas were on moderately steep slopes (16450), at upper elevations on the study area, and occurred on all aspects. Douglas-fir comprised 100% of occupied trees at Green Mountain and 83% at Whiteley Peak. Mean dbh of all occupied trees at Green Mountain and Whiteley Peak were 39.2 and 45.9 cm, respectively. Mean heights were 13.4 m at Green Mountain and 16.7 m at Whiteley Peak. Dbh of occupied trees did not differ between areas but heights did. ��217 CHARACTERISTICS AND HABITAT PREFERENCES OF WINTERING POPULATIONS OF BLUE GROUSE Brian S. Cade P. N. OBJECTIVES The primary objectives of this study are 1) identify the vegetational and structural components of blue grouse winter habitat and 2) determine the spatial relationship between blue grouse wintering and breeding sites. SEGMENT OBJECTIVES 1. Review available literature concerning habitat description and measurement, habitat use by blue grouse, attributes of wintering populations of grouse, use of radio telemetry on birds and other pertinent literature relating to grouse. 2a. Interview individuals with first-hand experience or knowledge of past and present winter occurrence of blue grouse in Middle Park. 2b. Investigate selected habitats for blue grouse occurrence in winter. 2c. Delineate 2 study areas in Middle Park suitable for conducting winter investigations that represent dissimilar habitat types. 3. Band blue grouse on study areas with serially-number anodized aluminum leg bands. and color-coded, 4. Equip 10 grouse at each study area (20 total/year) with radio transmitters. Three adult or yearling males, 3 adult or yearling females, and 4 juveniles will be instrumented at each study area. 5. Relocate radio-equipped grouse to ascertain movements from breeding areas to wintering sites, and to document habitat preferences, ment patterns, and temporal use of wintering areas. 6. move- Conduct weekly searches for grouse on study areas from November through March to determine vegetation types occupied, sex composition and size of flocks, locations of marked grouse, and behavioral activities. 7. Quantify vegetative and physiographic characteristics at grouse wintering sites. 8. Determine area of coniferous, deciduous, and shrub types from aerial photographs of the study areas. 9. Compile data, analyze results, and prepare progress report. �218 METHODS AND MATERIALS Blue grouse were banded on study areas primarily during the breeding and brood rearing periods from early April through mid-August. Grouse were located with tape recorded calls and/or trained pointing dogs (Hoffman 1981) and captured with a telescoping noose pole (Zwickel and Bendell 1967). Grouse were banded with serially-numbered aluminum bands and color-coded, anodized bands for individual recognition (Hoffman 1981). Banding locations were determined to the nearest 50 m as Universal Transverse Mercator (UTM) grid coordinates. Two types of radio transmitters were placed on grouse, a solar capacitorassisted unit (indefinite life) or a lithium battery-powered unit (10 month life). Transmitters were attached to the grouse with a poncho collar(Amstrup 1980) or a backpack harness (Brander 1968). Mounted weights were 20-22 gms for the solar powered transmitters and 29-31 gms for the battery powered units. Transmitters were on 150 and 151 Mhz frequency bands. Radio-equipped grouse were relocated at least once every 2 weeks while they remained on the study areas. Accessibility problems during winter hindered regular relocation of instrumented grouse that wintered off the study areas. An early (Oct-Nov) and late (Feb-Mar) winter location was determined for each grouse that wintered away from the study areas. Radio location was achieved primarily with a hand-held yagi antenna. Locations were determined by visual observation of instrumented grouse and recorded to the nearest 50 m as UTM grid coordinates. One 3-hour period of aerial radio location was conducted in early November to locate missing grouse. Weekly searches of study areas from November through March were conducted on foot, snowshoes, and skiis. Search time was primarily spent in coniferous forests. All coniferous areas on the study sites were searched for grouse. Search time was concentrated at areas where grouse sign (droppings and/or tracks in snow) were observed or where grouse had previously been observed. An attempt was made to capture any unbanded grouse encountered on the study areas during winter. These grouse were instrumented with transmitters not used during the breeding season or with those recovered from birds that died. Habitat characteristics were quantified in O.Ol-ha circular on grouse flushing locations or trees in which grouse were (Stauffer and Peterson 1980). Plot locations were recorded 50 m as UTM grid coordinates and occupied trees were marked flagging and serially numbered aluminum tags. plots centered located to the nearest with colored Physiographic variables recorded at plots included: slope (level, 1-15, 16-30, 31-45, 46-600), aspect (8 cardinal directions), elevation (to nearest 100 m), and relative position on the slope (ridge top, upper, mid, or lower one-third, valley bottom). �219 Variables quantified for each tree species within the plot included: stem density of trees (> 7 cm dbh), stem density of sapplings « 7 cm dbh) , dbh of trees, and canopy height (height of tallest tree). Basal areas calculated from dbh measurements were used to define species dominance. Point centered quarter measurements (Cottam and Curtis 1956) from the plot center tree to the nearest tree in each of 4 quadrats were used to calculate horizontal dispersion (Stauffer and Best 1980) of trees, as well as providing a better estimate of tree density in low stem density stands. Height, dbh, form (normal pyramidal, overmature, deformed), and species were recorded for the plot center tree in which grouse were located. Total area of coniferous, deciduous, and shrub types were measured with a planimeter from enlarged aerial photographs. Coniferous tree stands were classified on the basis of species composition and relative stem density. Deciduous tree stands and shrub dominated areas were not separated into more specific categories. Area corrections were determined for habitat types that occurred on steep slopes. Area corrections were not applied unless they were 10% or greater. Habitat variables measured at occupied grouse locations were analyzed with a 1-way analysis of variance or t tests. DESCRIPTION OF STUDY AREAS Field work was conducted on 2 areas of differing habitat types in northcentral Colorado. Both areas are within Middle Park, about 161 km west of Denver in portions of Grand and Summit counties. These areas are referred to as Green Mountain and Whiteley Peak (Fig. 1). Green Mountain is approximately 19 km south of Kremmling, Colorado in the southwest corner of Middle Park and has been the site of intensive grouse studies since 1975 (Hoffman 1981). The winter study area encompasses 332 ha including the 181-ha area investigated by Hoffman (1981). Coniferous forests, including Douglas-fir, Douglas-fir/aspen, and Douglas-fir/ Rocky Mountain juniper (Juniperus virginiana) associations, cover 138 ha (41%) of the study area. Aspen dominated forests constitute 6 ha (2%) of the area with non-forested areas, including areas with scattered trees, covering the remaining 188 ha (57%) (Fig. 2). Open areas are vegetated primarily with big sagebrush (Artemisia tridentata). Scattered to dense stands of Douglas-fir predominate above 2,700 m on the east slope and extend downward to 2,600 m. Intermediate areas on the east slope consist of a mixed Douglas-fir/aspen/shrub association (Hoffman 1981). On the west slope large tracts of dense Douglas-fir and open Douglas-fir/ Rocky Mountain juniper associations extend from 2,700 m downward to 2,400 m. Maximum reI ief on the study area is 470 m; elevations range from 2,390 to 2,863 m. �220 RIVER • t ./ DENVER N t CONTINENTAL -DIVIDE 12.8 Km COLORADO MIDDLE PARK Fig. 1. Whiteley Peak and Green Mountain study areas, Middle Park, Colorado. .1 �221 GREEN MOUNTAIN n 188 ha (57%) NON-FORESTED r.:-:-7"', 1':':-:':1 I~ .[Ill] ~ '.W....' 6 ha ASPEN DOUGLAS-FI R 67 ha (20%) DOUGLAS-FIR/ASPEN 24 ha DOUGLAS-F I R/ JUN I PER 47 ha (14%) ti-'tl" ).ltU (2%) (7%) .. N i Fig. 2. Habitat types at Green Mountain, Middle Park, Colorado. �222 Whiteley Peak is in the northwest corner of Middle Park about 34 km north of Kremml ing. Total area investigated is 503 ha. Maximum relief is about 575 m with elevations ranging from 2,500 to 3,075 m. Coniferous types at Whiteley Peak include: Douglas-fir; Douglas-fir/aspen mixtures; associations of Douglas-fir with subalpine fir (Abies lasiocarpa), Engelmann spruce (Picea engelmannii), lodgepole pine (Pinus contorta), limber pine (Pinus flexilis), and blue spruce (Picea pungens); and mixed conifer/aspen associations. Coniferous forests cover 122 ha (24%) of the study area. Open areas are primarily dominated by sagebrush communities. Non-forested types, including those with scattered trees, encompass 265 ha (53%) of the area. Aspen dominated stands are present on 116 ha (23%) of area between the sagebrush and coniferous forest communities (Fig. 3). RESULTS AND DISCUSSION Blue Grouse Bandings Sixty-seven grouse were banded at the 2 study sites in Middle Park; 45 at Whiteley Peak and 22 at Green Mountain. Eight males and 14 females were banded at Green Mountain. Twenty-four males and 21 females were banded at Whiteley Peak. An additional 30 juvenile grouse were marked with patagium tags. Radi6~Marked Blue Grouse A total of 23 grouse was instrumented with radio transmitters on the study areas: 3 died almost immediately, 5 died before winter, contact was lost with 3, and 12 were followed to wintering areas (Tables 1, 2). Movements of Males Six instrumented males, 3 each at Green Mountain and Whiteley Peak, survived through summ~r. During the breeding season, the 4 territorial, adult males remained on their territories moving less than 0.3 km from their initial capture location. One non-territorial yearling wandered widely on Green Mountain, moving over 1 km from his initial capture point. Another non-territorial yearling at Green Mountain had more restricted movements remaining within 0.5 km of his initial capture location. Instrumented adults migrated from the study areas by late June and moved to higher elevations in spruce-fir communities. Yearlings left by midJuly and moved to similar areas for the summer. Movements of instrumented males from breeding areas were similar to patterns reported by other workers (Caswell 1954, Bendel I 1955, Zwickel et al. 1968, King 1971). Two adult males made an additional movement in early October of 2.6 and 4.5 km to their respective wintering sites. Similar movements could not be documented for other radio-marked males. Movements of Females Only 4 instrumented females survived the breeding season into fall. An unsuccessful, yearling female at Green Mountain wandered widely off and on the study area until late June, moving as far as 1.9 km from her initial capture location. This grouse moved to spruce-fir habitat at higher elevations where she remained through the winter. An �WHITELEY PEAK o f .-I ASPEN [[II] if ....___J •• rf 't·l'-._"<"1" 'Y-: -:.:':l ~ \:.:.:. :\.,:::",::"":;:::_;,;,:,,,:;:'},:.:.:./ NON-FORESTED DOUGLAS-FIR 265 ha (53%) 116 ha (23%) 6 ha (1%) N N t;~4 MIXED CONIFER 40 ha (8%) ~ MIXED CONIFER/ASPEN 42 ha (8%) ~ DOUGLAS-FIR/ASPEN 34 ha (]%) \. ~ c() d>~ Fig. 3. Habitat types at Whiteley Peak, Middle Park, Colorado. N o i o t %: 1 Krn w �Table 1. Colorado, Characteristics 1981-82. and status of radio-marked Max. distance moved (km) blue grouse at Green Mountain, Total no. of -relocations (winter) Middle Park, Band no. A~e 222 1- F Chick 299 1- F Chick 3.4 9 (7) 25 Aug-22 Mar Wintered off study area 196 1+ F Unsuccessful 8.9 13 (2) 21 Apr -3 1 Ma r Wintered off study area 270 1+ F Successful 0.4 5 21 Jul- 1 Oct Remained in vicinity of study area with brood. Mortality (hunter) 2+ F Successful 0.4 3 22Jul- Remained in vicinity of study area with brood. Mortality (predation) Sex Breeding 306 2+ F Unknown 245 1+ M 243 1+ 244 2+ status 1 Observation per iod Comments 4-15 Sep Mortality 8Aug (unknown) 1.1 12(11) 1 Oct-27 Mar Moved onto study area for winter Non-territorial 12. 1 11 (2) 6 May-31 Mar Wintered M Non-territorial 7.8 11 6 May-20 Sep Moved off study area for summer. Mortality (hunter) M Territorial 10 (1) 6 May- 2 Nov Wintered 12.6 off study area off study area N N -l:" �Table 2. Colorado, Characteristics 1981-82. and status of radio-marked Max. distance moved (km) Band no. Age 295 1- F Chick 302 1- F Chick 201 1+ F Unsuccessful 1.0 203 2+ F Unsuccessful 0.7 207 2+ F Nest attempt 0.3 Sex Breeding status 14.2 blue grouse at Whiteley Total no. of relocations (winter) Comments 21 Aug-10 Oct Moved off study area in fa II. Unable to relocate during winter. 1 26 Aug- 6 Sep Unable to relocate 25 Apr-29 Mar Wintered 5 28 Apr- 2 June Mortality glement) (antenna entan- 5 4 May-17 Jun Mortality nest) (predated on 21 (10) 2+ F Successful 263 2+ F Successful 267 2+ F Successful 308 2+ F Unknown 298 1- M Chick 11.3 202 2+ M Territorial 14. 1 204 2+ M Territorial 241 2+ M Territorial 12.5 13 (1) 242 2+ M Territorial 19.3 8 (2) 28.0 Observation period 3 262 0.5 Peak, Middle Park, on study area 9 26 Jun-19 Sep Unable to relocate winter 1 27 Jun- 6 Jul Mortality 9 (2) 13 Jul-25 Feb Moved off study area for winter; mortality (unknown) 6 (6) 30 Jan-26 Mar Captured during winter on study area 2 21 Aug-13 Sep Moved off study area with brood. Morta 1ity (predation) 28 Apr-30 Mar Wintered 28 Apr- 4 May Mortality 13 (1) 1 during (unknown) off study area (predation) 5May-30Mar Wintered off study area 5 May-30 Mar Wintered off study area N N V1 �226 unsuccessful, yearl in9 female at Whiteley Peak wandered widely on the study area moving 1.6 km from her initial capture location. In mid-October this female moved a short distance uphill to coniferous habitat on Whiteley Peak where she remained during winter. Two instrumented brood hens at Whiteley Peak showed restricted movements during summer,moving less than 0.5 km. One instrumented female began moving her brood away from the study area in mid-August. She moved to higher elevations to spruce-fir habitat for the winter. The other instrumented brood hen remained on the study area until mid-September, when radio contact was lost. Movements of juveniles Five juvenile grouse were radio marked in late August and early September. Two chicks, 1 male and 1 female, at Whiteley Peak were from broods with instrumented females. An additional female chick at Whiteley Peak and 2 at Green Mountain came from broods with unmarked females. One chick at Green Mountain died shortly after being instrumented and the other did not rejoin her broodma:tes. This juvenile female moved away from the study area for winter, moving into coniferous habitat by mid-October. An instrumented, juvenile male at Whiteley Peak moved away from the study area with his instrumented brood hen to spruce-fir habitat at higher elevations. This male was killed by a predator in mid-September approximately mid-way between the study area and the winter location of the instrumented brood hen. Radio contact was lost with the other brood hen in mid-September but her instrumented, female chick moved from the study to higher elevations in the spruce-fir zone. This juvenile could not be relocated during winter. The other instrumented, juvenile female at Whiteley Peak was not relocated after mid-September. Spatial Relationship Between Wintering and Breeding Areas Eight of 11 instrumented grouse that migrated from the study areas were tracked to winter locations. Mean distance to wintering areas for 4 grouse from Whiteley Peak was 17.3 km (range 7.8-28.0 km) and for 4 grouse from Green Mountain was 9.3 km (range 3.4-12.6 km) (Fig. 4). Mean distance for all birds was 13.3 km. Winter locations of instrumented grouse migrating from Whiteley Peak were east of the study area along the Continental bivide in the Rabbit Ears Range (Fig. 4). Two additional instrumented grouse moved into this general area before radio contact was lost. Bands were recovered from 2 juvenile and 1 adult grouse harvested in this area by hunters during the fall grouse season. The Rabbit Ears Range appears to be a major wintering area for many grouse that breed at Whiteley Peak. �227 Instrumented grouse migrating from Green Mountain for winter moved in several directions. The yearling and juvenile female moved into the foothills of the Gore Range. Two males, 1 adult and 1 yearling, moved to sites adjacent to high peaks in the Gore Range. The other yearl ing male moved the opposite direction into the Williams Fork Mountains (Fig. 4). He was shot by a hunter during fall. Whether or not this bird would have remained in this area during winter is not known. Blue grouse are known to winter on the Williams Fork Mountains. One instrumented yearling female and 1 banded adult female remained at Whiteley Peak during winter. The unsuccessful yearl ing wintered 1 km from her initial capture location. The successful adult female wintered less than 0.5 km from where she was captured with her brood in summer. An adult female captured on Whiteley Peak in January 1981 was located on the study area in spring 1982. An adult male banded on the mountain in early fall 1981 and observed on the study area during winter, was recaptured in spring 1982 on his territory 1.0 km downhill from his winter location. Only 1 instrumented grouse wintered on Green Mountain. This adult female was instrumented in late September on Little Green Mountain just west of the study area. Little Green Mountain was determined to be her breeding area since she returned there to nest in spring 1982. Distance moved between her winter location and breeding site was 0.5 km. Zwickel et al. (1968) and King (1971) documented extensive movements of blue grouse from breeding to wintering areas. Hoffmann (1956) and King (1971) found grouse wintering and breeding on the same areas. Hoffmann (1956) also believed that grouse wintering on his study area were the same ones which bred there. Data obtained in Middle Park suggest that a breeding population of grouse may contain individuals that migrate long distances to wintering areas and individuals that winter in coniferous habitat adjacent to their breeding areas. J. Hines (pers. commun.) has indicated that blue grouse at his study area on Vancouver Island also may move long distances (5+ km) to wintering areas or winter and breed at the same location. Winter Home Ranges Home ranges were calculated for 4 instrumented females. Three adults wintered on the study areas and the juvenile female wintered west of Green Mountain. Area corrections were applied to home ranges on steep slopes. Mean home range size for these 4 females was 7.6 ha and ranged from 1.8 to 18.7 ha (Fig. 5). Reobservations of 4 banded grouse (1 male, 3 female) suggest that they also remained in a fairly restricted area during winter. Winter Population Characteristics Green Mountain A total of 306 search hours between mid-October and the end of March resulted in 48 observations of 84 grouse. Grouse were observed in coniferous habitat on Green Mountain as early as 3 October but were not frequently observed until late in the month. Lone grouse were observed 33 times of �N 228 E N E s G MALE o Fig. 4. Winter locations of sites, Middle Park, Colorado. fa 11) . radio-marked (l=Fall FEMALE WINTER WINTER blue grouse migrating from study mortality, 2=Lost contact during LOCAT ION LOCATION �229 ADULT 140 7.1 ha (25% AREA CORRECTION) JUVENILE 108 18.7 ha ~ (3) ADULT 156 2.6 ha (10% AREA CORRECTION) YEARLING 086 1.8 ha (10% AREA CORRECTION) o • 1 ha UTM LOCATIONS (NUMBER OF MULTIPLE Fig. 5. Winter home ranges of 4 instrumented Middle Park, Colorado 1981-82. RELOCATIONS) female blue grouse, �230 which 7 were males, 14 were females, and 12 were of unknown sex. Fifteen flocks of grouse were observed ranging in size from 2 to 6 birds. Roth sexes were present in 3 flocks, males only in 1 flock, females only in 3 flocks, and unknown sexes in 8 flocks. Age class could only be assigned to 1 instrumented female that wintered at the study area. Whiteley Peak A total of 270 search hours produced 41 observations of 91 grouse. Grouse were first observed in coniferous habitat at the end of September. Lone grouse were observed 16 times of which 2 were males, 9 were females, and 5 were of unknown sex. Twenty-five flocks were observed ranging from 2 to 12 birds. The largest flocks observed, 9 and 12 grouse, were located in November. Both sexes were present in 13 flocks, males only in 2 flocks, females only in 8 flocks. and unknown sexes in 2 flocks. Ages were known for 6 marked grouse; 4 adult females, 1 yearling female, and 1 adult male. Sex segregation of blue grouse in winter has been suggested by several authors (Skinner 1927, Marshall 1946, Caswell 1954). Observations in Middle Park do not support this contention. Males and females were observed using identical areas on the study sites and were frequently observed together, often in the same tree. Social organization in winter has been briefly described by several authors. Hoffmann (1956) considered blue grouse to be highly solitary. Our observations conform more to reports of Munro (1919), Beer (1943), and Caswell (1954) who observed the formation of small, loose flocks in winter and to King (1971) who observed lone birds, small groups, and an occasional large flock. Behavior Most grouse observed were perched in conifers. They appeared to be sedentary for the majority of the day. Grouse were observed to go on feeding binges during evening shortly before dark. Feeding grouse often fluttered about on conifer branches as they bit off individual needles and clumps of needles. "Umr rh" calls were frequently heard when grouse were feeding. Caswell (1954) reported similar observations of feeding grouse during the morning in Idaho. It is possible that grouse at our study areas fed early in the morning and that we did not arrive early enough to observe this behavior. Grouse frequently fed in the same trees in which they were roosting. No fl ights to feeding areas, as reported by Caswell (1954), were observed. Grouse tracks in snow were frequently observed although no grouse were actually observed walking in the snow. This behavior would have been missed if it occurred early in the morning. Roosting grouse were observed at all heights in trees except near the top. During inclement weather grouse tended to roost near the tree trunk. During fair weather they were observed at many positions in the canopy. �231 Characteristics of Wintering Sites Species Composition A total of 86 0.01 ha vegetation plots was measured at grouse location on the study areas. Ninety-five percent of the plots (N = 42) at Green Mountain occurred in 3 habitat types: mature Douglas-fir~ Douglas-fir/aspen mixtures,and Douglas-fir/Rocky Mountain juniper associations. In all 3 types, Douglas-fir was the predominant tree in the 0.01 ha plot (Table 3). Ninety-eight percent of the plots (N = 44) at Whiteley Peak occurred in 3 habitat classifications: open mixed conifer stands, dense mixed conifers, and Douglas-fir/aspen associations. Douglas-fir was the dominant species in all 3 types with percent of total basal area varying from 50 to 95% (Table 4). Table 3. Tree species dominance at blue grouse winter locations on Green Mountain, Middle Park, Colorado. Frequency of occurrence \ (%) Mean basa I area (cm2/O.01 hal Percent of total mean basal area Stand type N Species Douglas-fir/ Aspen 12 Douglas-fir Aspen 100 17 2,524 132 95 5 Douglas-fir/ Juniper 16 Douglas-fir Juniper 100 44 '2,903 261 92 8 Douglas-fir 12 Douglar-fir Juniper 100 2,961 44 99 8 1 Table 4. Tree species dominance at blue grouse winter locations on Whiteley Peak, Middle Park, Colorado. Percent of total mean basal area Frequency of occurrence (%) Mean basal area (cm2/O.01 hal 100 33 8 2,980 78 88 95 2 3 Stand type N Species Douglas-fir aspen 12 Douglas-fir Aspen Cottonwood Open mixed conifer 10 Douglas-fir Limber pine Subalpine fir Aspen 70 50 50 10 1,183 880 307 4 50 37 13 <, 1 Dense mixed conifer 20 Douglas-fir Subalpine fir Limber pine Engelmann spruce Aspen 90 70 45 10 20 3,304 1,489 802 104 28 58 26 14 2 < 1 �234 Table 8. Dispersion characteristics of trees at blue grouse winter locations on Whiteley Peak, Middle Park, Colorado. Open mixed conifer(N=6) Habitat types Dense mixed Douglas/fir conifer(~=19) Aspen(~=10) All types (N=36) x distance, m 6.9 4.6 6.6 - bSE 1.4 0.5 1.3 1.1 46.2 47.3 52. 1 5.8 9.4 12.0 48.8 9.4 CV SE 5.5 aMeasured from plot center tree to nearest trees in 4 quadrats. bM ean coe ff" IClent 0f .. variation. Diameters of trees measured by the point-centered quarter method for plots at Green Mountain (Table 9) and Whiteley Peak (Table 10) indicated that mature trees were associated with wintering sites of grouse at both areas. Mean dbh comparisons between all sites at Green Mountain and Whiteley Peak were not significant (~= 0.183, 65 df, P > 0.5). Table 9. Diameter of trees at blue grouse winter locations on Green Mountain, Middle Park, Colorado Habitat types Douglas-fir(DF) DF/aspen DF/juniper (N=10) (N=8) (N=l1) x dbh, cma - bSE CV SE 27.5 3.9 23.6 2.7 56.3 48.1 7.8 11.4 a 27.5 25.7 4.1 51. 7 3.9 53.0 8.8 9.2 Measured at nearest trees to plot center in 4 quadrats. bMean coefficient of variation. All types (N=31) �235 Table 10. Diameter of trees at blue grouse Peak, Middle Park, Colorado. winter locations on Whiteley Habitat types Open mixed Dense mixed Douglar-fir/ conifer(N=19) aspen (N=ll) con ifer (!:!_=6) ~ dbh, cm CV SE b SE a 24.7 25.7 5.8 4. 1 49.9 7.3 49.2 9.2 27.7 6.2 62.9 8.7 All types (!:!_=36) 25.9 4.7 52.4 9.3. Measured at nearest trees to plot center in 4 quadrats. bMean coefficient of variation. Canopy heights of plots at Green Mountain and Whiteley Peak varied (Table 11). Mean canopy height for Green Mountain plots was 13.9 m compared to 17.6 m for plots at Whiteley Peak (t = 3.315, 84 df, P < 0.05). Canopy heights differed between study areas but were indicative of mature trees on both areas. Table 11. Canopy height at blue grouse .'.i; ...;.•.. N . (m) a x HeIght so 12 12 16 42 13 .2 13.4 15.2 13.9 3. 1 3. 1 6.2 4.5 12 10 20 16.5 10.6 22.1 3.3 3.9 3.4 Green Mountaih Douglar-fi r DF/aspen OF/juniper All types Whiteley Peak DF/aspen Open mixed conifer Dense mixed conifer All types aHeight of tallest tree in 0.01 ha plot. Physiographic Characteristics Physiographic variables at Green Mountain (Table 12) and Whiteley Peak (Table 13) indicated that occupied habitat at both areas generally occurred on moderately steep slopes. Eight-three percent of all plots on Green Mountain occurred on slopes in the 16-450 categories; 84% of all plots on Whiteley Peak also occurred in these categories. �236 Table 12. Physiographic characteristics Green Mountain, Middle Park, Colorado. Frequency, % (N=42) at blue grouse winter Level 1-15 16-30 5 12 52 locations on 31-45 31 Aspect Frequency, (N=42) % Level N NE E SE S SW W NW 5 7 41 2 2 2 12 17 12 Vertical position on slope Ridge Frequency, % (N=42) Upper 1/3 12 Mid 1/3 Lower 113 31 7 50 Table 13. Physiographic characteristics at blue grouse winter locations on Whiteley Peak, Middle Park, Colorado. Slope (0) Frequency, % (N=49) Level 1-15 16-30 0 16 61 31-45 23 Aspect Frequency, % (N=49) Level N NE E SE S SW W NW 0 2 2 4 18 12 8 27 27 Vertical position on slope Ridge Frequency, % (N=49) 16 Upper 1/3 Mid 1/3 Lower 1/3 76 4 4 �237 Frequency of plots in aspect categories (Tables 12, 13) were highly variable and should not be construed to suggest a preference for particular exposures. These data suggest that blue grouse winter use sites can occur on any aspect of the 2 study areas. Mean elevation of grouse locations on Green Mountain was 2,645 m and ranged from 2,402 to 2,858 m. Mean elevation of plots at Whiteley Peak was 2,858 m and ranged from 2,614 to 3,040 m. Sixty-two percent of all plots at Green Mountain were on the upper one-third of the slope of the mountain or ridge tops (Table 12). The same 2 categories at Whiteley Peak contained 94% of all plot locations. Occupied Tree Characteristics Variables for occupied trees (trees used) on Green Mountain varied (Table 14). Mean diameters of occupied trees in 3 habitat types ranged from 34.1 to 48.8 cm. Diameters differed for occupied trees among habitat types at Green Mountain (I2_,34 = 4.315; 1: < 0.05). Although differences exist among habitat types, mean diameters were all large. Mean height for all occupied trees on Green Mountain was 13.4 m. Table 14. Characteristics of trees occupied by wintering Green tiountain, Middle Park, Colorado Douglas-fir (N=12) OF/aspen (,!!=10) blue grouse on DF/j un iper All types (,!!= 15) (N=39) x dbh, cm SO Max Min 34.4 6.3 48 26 34.1 12.9 62 10 48.8 19.8 94 27 39.2 16.2 94 10 ~ height, m SO Max Min 12.3 2.7 16 5 13.1 3.5 17 5 14.7 6.9 32 5 13.4 4.9 32 5 Form, % frequency Pyramidal Overmature Deformed 25 58 17 30 60 10 20 47 33 28 51 21 100 100 100 100 Species, % frequency Douglas-fir �237 Frequency of plots in aspect categories (Tables 12, 13) were highly variable and should not be construed to suggest a preference for particular exposures. These data suggest that blue grouse winter use sites can occur on any aspect of the 2 study areas. Mean elevation of grouse locations on Green Mountain was 2,645 m and ranged from 2,402 to 2,858 m. Mean elevation of plots at Whiteley Peak was 2,858 m and ranged from 2,614 to 3,040 m. Sixty-two percent of all plots at Green Mountain were on the upper one-third of the slope of the mountain or ridge tops (Table 12). The same 2 categories at Whiteley Peak contained 94% of all plot locations. Occupied Tree Characteristics Variables for occupied trees (trees used) on Green Mountain varied (Table 14). Mean diameters of occupied trees in 3 habitat types ranged from 34.1 to 48.8 cm. Diameters differed for occupied trees among habitat types at Green Mountain (£2.,34 = 4.315; 1:. < 0.05). Although differences exist among habitat types, mean diameters were all large. Mean height for all occupied trees on Green Mountain was 13.4 m. Table 14. Characteristics of trees occupied by wintering Green t10untain, Middle Park, Colorado blue grouse on DF/j uni per All types Douglas-fir DF/aspen (!!=12) (N=10) x dbh, cm SD Max Min 34.4 6.3 48 26 34. 1 12.9 62 10 48.8 19.8 94 27 39.2 16.2 94 10 x height, m SD Max Min 12.3 2.7 16 5 13.1 3.5 17 5 14.7 6.9 32 5 13.4 4.9 32 5 Form, % frequency Pyramidal Overmature Deformed 25 58 17 30 60 10 20 47 33 28 51 21 100 100 100 100 Species, % frequency Douglas-fir (!:!_=15) (!:!_=39) �Mean diameters for occupied trees on Whiteley Peak in 3 habitat types ranged from 44.2 to 48.0 cm (Table 15). Diameters did not differ among habitat types (£2,37 = 0.209, P > 0.25). Mean heights of occupied trees at Whiteley Peak was 16.7 m. Table 15. Characteristics of trees occupied by wintering Whiteley Peak, Middle Park, Colorado. blue grouse on Open mixed con ifer(N=l 0) Dense mixed con ife r(!::!.= 19) Douglas-fir/ aspen (N=l1) Max Min 44.2 14.3 61 16 45.4 15.9 76 25 48.0 9.5 62 35 45.9 14.4 76 16 ~ height, m SD Max Min 9.4 3.8 14 3 21.2 4.3 28 13 15.9 3.4 20 8 16.7 6.1 28 3 Form, % frequency Pyrami da I Overmature Deformed 10 60 30 47 47 6 27 55 18 34 52 14 Species, % frequency Douglas-fir Limber pine Subalpine fir 60 30 10 84 5 100 83 10 7 x dbh, cm so 11 All types (.!::!_=42) Diameters and heights of occupied trees at both Green Mountain and Whiteley Peak indicated that mature trees were used by blue grouse during winter. Diameters of occupied trees at Green Mountain and Whiteley Peak did not differ (F i.so = 3.81; ~ > 0.05) although heights did (II SO = 6.58; ~ < 0.05T. , , Douglas-fir was the only tree species used by grouse at Green Mountain (Table 14). Blue grouse at Whiteley Peak were observed in limber pine, subalpine fir, and Douglas-fir. Douglas-fir was the species most frequently (83% of all observations) occupied (Table 15). Blue grouse were observed in all 3 forms of trees, however, slightly greater than 50% of all observations at both Green Mountain and Whiteley Peak were in large, overmature trees. �239 Habitat characteristics at winter grouse sites are similar to observations of other workers. Caswell (1954) noted that blue grouse wintered in fairly open stands or dense pockets of Douglas-fir. King (1971) observed that most wintering blue grouse in sUbalpine areas used open parkland and open canopy foreSts. Stauffer and Peterson (1980) observed a similar use of open conifer types in Idaho, although they did observe some grouse sign in dense timber. Observations of blue grouse in Middle Park occurred in both relatively open and dense types. Presence of mature Douglas-fir wasa common characteristic at all sites. Stauffer and Peterson (1980) observed a similar use of large, mature Douglas-fir. Caswell (1954) reported that trees used by blue grouse in his study ranged in size from 15 to 30 cm dbh. King (1971) reported that mature firs with large limbs were used by wintering grouse in subalpine areas of Vancouver Island. Douglas-fir was the dominant species at all wintering areas on the study sites and were used most frequently. Subalpine fir and limber pine were used infrequently. Grouse that migrated from the study areas for winter moved into spruce-fir communities where SUbalpine fir and Engelmann spruce predominate and where Douglas-fir was uncommon. Actual observations of wintering grouse at these sites were too few for analysis. However, instrumented female that wintered west of Green Mountain was repeatedly relocated (N = 7) in lodgepole pine in an area where Douglas-fir was present. Harju (1974) observed blue grouse in limber and lodgepole pine during late winter in Wyoming. Needles from a variety of conifers have been reported in food habits studies of blue grouse (Stewart 1944, Hoffmann 1961, and King 1968). The almost exclusive use of Douglas-fir in Middle Park does not necessarily indicate that blue grouse are dependent on this species during winter. LITERATURE CITED Amstrup, S. C. 1980. 44:214-217. Beer, J. 1943. 32-44. A radio-collar for game birds. Food habits of the blue grouse. J. Wildl. Manage. J. Wildl. Manage. 7: Bendell, J. F. 1955. Age, breeding behaviour, and migration of sooty grouse, Dendragapus obscurus fuliginosus (Ridgway). Can. J. Zool. 33:195-223. Brander, R. B. 1968. A radio-package Manage. 32:630-632. harness for game birds. J. Wildl. Caswell, E. B. 1954. A preliminary study on the life history and ecology of the blue grouse in west central Idaho. M.S. Thesis. Univ. Idaho, Moscow. 105pp. Cottom, G., and J. T. Curtis. 1956. The use of distance measures phytosociological sampling. Ecology 37:451-460. in �240 Harju, H. J. grouse. 1974. An analysis of some aspects of the ecology of dusky Ph.D. Thesis. Univ. Wyoming, Laramie. 142pp. Hoffman, R. W. 1981. Population dynamics and habitat relationships of blue grouse. Colo. Div. Wildl. Job Final Rep., Fed Aid Proj. W-37-R-34. April 1981. Pp. 103-171. Hoffmann, R. S. 1956. Observations on a sooty grouse population at Sage Hen Creek, California. Condor 58:321-337. 1961. The quality of the winter food of blue grouse. Manage. 25:209-210. J. Wildl. King, D. G. 1971. The ecology and population dynamics of blue grouse in the sub-alpine. M.S. Thesis. Univ. British Columbia, Vancouver. 139pp. King, R. D. 1968. Food habits in relation to the ecology and population dynamics of blue grouse. M.S. Thesis. Univ. British Columbia, Vancouver. 62pp. Marshall, W. H. 1946. Cover preferences, seasonal movements, and food habits of Richardson's grouse and ruffed grouse in southern Idaho. Wilson Bull. 58:42-52. Munro, J. A. 1919. Notes on some birds of the Okanagan Valley, British Columbia. Auk 36:64-74. Skinner, M. P. 1927. Bull. 39:208-214. Richardson's grouse in Yellowstone Park. Wilson Stauffer, D. F., and L. B. Best. 1980. Habitat selection by birds of riparian communities: evaluating effects of habitat alterations. J. Wildl. Manage. 44:1-15. , and S. R. Peterson. 1980. Seasonal habitat relationships of blue ----~(~Dendragapus obscurus) and ruffed (Bonasa umbellus) grouse in southeastern Idaho. Annu. Rep., Forest, Wildl., and Range Exp. Stn., Univ. Idaho, Moscow. 37pp. Sakal, R. R., and F. J. Rohlf. 1969. Biometry: the principles and practices of statistics in biological research. W. H. Freeman and Co., San Francisco, Calif. 776pp. Stewart, R. E. 1944. Food habits of the blue grouse. Condor 46:112-120. Zwickel, F. C., and J. F. Bendel1. 1967. A snare for capturing blue grouse. J. Wildl. Manage. 31 :202-204. �241 , I. O. Buss, and J. H. Brigham. 1968. ---grouse and their relevance to populations Manage. 32:456-468. Prepa red by -73-€~~/(,-"",-:HW1~·~+:-,,"j-,--· -'i1o,....::.u:~=/Q_=;;_-Brian S. Cade Graduate Research Assistant _ Autumn movements of blue and management. J. Wildl. ��Apr iI 1982 243 JOB PROGRESS REPORT State of Colorado ----~~~~~------------ Proj ect No. Game Bird Survey W-37-R-35 Work Plan No. 13 Job No. Population Characteristics Job Title: Columbian Sharp-tailed Period Covered: Personnel: 8 and Habitat Requirements Grouse in Northwestern of Colorado 1 January 1981 through 31 December 1981 Mike Bauman, Clait Braun, Ken Giesen, Jim Haskins, Jim Hicks, Rick Hoffman, Gary Miller, Steve Steinert, Chuck Woodward, Colorado Division of Wildlife. ABSTRACT Review of literature, location of 2 study areas, evaluation of field techniques, and development of a detailed study plan were priorities in 1981. Five potential study areas in Routt and Moffat counties were evaluated (California Park, Cedar Hill Gulch, Hayden-North, Trout Creek, Twentymile Park) and Cedar Hill Gulch and Hayden-North were selected for intensive research. Counts of 24 active leks resulted in 241 males, 24 females, and 335 total sharptails being counted. A total of 34 adul"t sharptails (23 males, 11 females) was captured and banded. Four juveniles were captured but were too small to band. A sample of 142 sharptail wings was received from the hunter harvest of which 59 (41.6%) were from juveniles. Harvest centered around 2 locations (California Park, Twentymile Park) with 73.2% of the total harvest occurring the initial weekend. ��245 POPULATION CHARACTERISTICS AND HABITAT REQUIREMENTS OF COLUMBIAN SHARP-TAILED GROUSE IN NORTHWESTERN COLORADO Kenneth M. Giesen The Columbian or Mountain sharp-tailed grouse Wedioecetes.phasianellus) has declined in distribution and abundance throughout its historical range, including Colorado (Aldrich 1963, Miller and Graul 1980). Reasons for this decline are not well documented although changes in land use resulting from agriculture, energy development, and human population growth have coincided with population declines (Hart et al. 1950, Kessler and Bosch 1981) • Although the Columbian sharp-tailed grouse is a game species in Colorado, little information is available upon which to base management decisions. Baseline data on Columbian sharptail populations, habitats, and harvest are lacking not only in Colorado but throughout its range~ Previous studies on distribution of Columbian sharptails in Colorado indicated that the largest populations and apparent best habitats occur in Routt and eastern Moffat counties (Rogers 1969, Giesen and Hoffman 1981) with most of the harvest occurring near Cal ifornia Park and Twentymile Park in central Routt County. Research on the sharptail resource is needed to obtain basic i~formation on breeding densities, nesting success, production and survival of young, fall population numbers, harvest, population turnover, and recruitment to the breeding population. Information on seasonal movements and habitat use is needed to quantify food and cover requirements. Land use changes resulting from human population growth, agriculture, and energy development are expected to further reduce sharptail habitat in the near future while hunting and other recreational uses of sharptails and their habitat are expected to increase. This report covers the initial year of the study. P. N. OBJECTIVES The major objectives of this study are to measure sharptail breeding density, production, harvest, survival and turnover rates, and to obtain qualitative and quantitative measures of Columbian sharptail habitat in western Colorado. The specific objectives during the initial year of planning will be to select 2 or more intensive study areas in Routt County, to evaluate techniques for locating sharptail dancing grounds and broods, to test capture techniques, and to evaluate hunter distribution and harvest rate. �246 SEGMENT OBJECTIVES 1. Review pertinent literature applicable to the objectives of this study. 2. Select 2 or more intensive study areas based on observations of sharptailed grouse, distributions of leks, land use and habitat type, and distribution of huhter harvest. 3. Ascertain distribution of sharptail grouse in Routt and eastern Moffat counties from observations, and contacts with district wildlife managers, landowners, and other interested personnel. 4a. Locate dancing grounds in March, April, and May by systematic search of suitable habitats using binoculars and a parabolic microphone listening device during morning and evening display periods. 4b. Make counts of male and female sharptails on known sharptail during the morning display periods in April and May. leks Sa. Locate sharptail broods by systematic search of suitable habitats and by using a tape-recorded chick distress call. Sb. Ascertain August. brood size from counts of broods located in July and 6a. Trap adult sharptails using cannon nets on leks, drive traps and baited walk-in traps, and vehicle-mounted cannon nets in winter and spring, and mark with numbered aluminum bands and plastic bandettes color-coded to trap site. 6b. Capture sharptail chicks using walk-in drive traps in areas where broods are known to occur. 7. Estimate nesting success from wing molt of hens captured in summer and from wings obtained from hunter harvested birds. 8. Ascertain distribution of hunters and hunter harvest from field checks, check stations, and wing barrels. 9. Obtain number and location of marked birds shot through use of field hunter checks, check stations, and voluntary mail reporting. 10. Food habits will be ascertained from crops of sharptails obtained from hunters and systematic collections. 11. Compile data, analyze results, and prepare progress reports. METHODS AND MATERIALS Field personnel of the Colorado Division of Wildlife (CDOW) in the Northwest region were contacted regarding potential study sites. Field surveys were conducted from March through May using binoculars and parabol ic microphone listening devices to relocate historic lek sites and search for additional leks. Mist nets and walk-in drive traps were evaluated as �247 trapping techniques on leks. Intensive search on foot using a taperecorded chick distress call or a pointing dog were used in locating sharptails in August and September. Distribution of hunters and hunter harvest was measured using 3 check stations and ·10 wing barrels. RESULTS AND DISCUSSION Location and Descri~tion of Study Area Five potential study areas (California Park, Cedar Hill Gulch, HaydenNorth, Trout Creek, Twentymile Park) were evaluated for use as intensive study sites based on distribution of leks, lek counts, habitat type, hunting pressure and harvest, and access. Cedar Hill Gulch and HaydenNorth were selected. Neither area currently receives much hunting pressure, both have good sharptail populations, and both are accessible throughout the year. Vegetatively, the 2 areas are dissimilar with Hayden-North containing numerous wheat fields and Cedar Hill Gulch being primarily native shrub communities and hay meadows. A complete vegetative description will appear in the final report. Lek Counts From mid-March through mid-May, 87 counts of 24 active sharp-tailed grouse leks were obtained in Routt and Moffat counties. No grouse were counted on 3 additional leks, possibly because of late access (Fly Creek) or because the leks had moved or become inactive (Elkhead Road #1, Sage Creek). A minimum of 335 sharptails (14.0 birds/active lek) was counted and a maximum of 241 (12.7/active lek) were classified as males. Only 24 sharptails were classified as females, primarily because of their more secretive habits and the difficulty in separating males from females. Both the number of active leks counted and the number of males per active lek increased from counts of previous years (Table 1). Five dancing grounds were located and counted for the first time this year (Cedar Hill Gulch, Fish Creek, Morgan Creek, Roadside, ViI lards) and small groups of displaying sharptails were observed near Little Buck Mountain and in Cal ifornia Park. The increased number of leks and higher number of males counted per active lek may be the result of increased search effort and may also indicate an increasing population. Table 1. Sharp-tailed grouse dancing ground counts in Routt and Moffat counties, 1964-65 and 1977-81. Year 1964 1965 1977 1978 1979 1980 1981 No. active 9rounds counted 7 15 2 8 15 5 24 No. of males 32 87 16 65 82 24 241 Average no. males per active lek 4.6 5.8 8.0 8. 1 5.5 4.8 12.7 �248 Peak male counts were obtained from 26 March to 8 May on leks where a minimum of 3 counts were obtained. Females were observed on leks only between 9 and 29 April. Because of dense vegetative cover on leks and lack of obvious sexual dimorphism of non-displaying sharptails, it was often difficult to classify birds as to sex or to obtain complete counts of all sharptails attending leks. Trapping and Banding A total of 34 sharp-tailed grouse (23 males, 11 females) was captured and banded in 1981. All but 1 were captured using mist nets or walk-in traps on leks. One yearling hen and 4 chicks (7-12 days old) were captured in late June. Trapping success was not high considering the effort (approximately 50 man-days) involved. Limited efforts to attract sharptails to bait sites in late fall were not successful, possibly because snow cover was insufficient to cover natural foods. Sharp-tailed Grouse Harvest and Wing Analysis The hunting season for Columbian sharp-tailed grouse in 1981 started onehalf hour before sunrise on 12 September and closed on 27 September (Units 14, 16, 18, 20, 22, 24, 26, 28, 54, 56, 58, 60, and 66) and 4 October (Units 10 and 12). Since most huntable populations of sharptails occur in Unit 14, 16~ and 26 they Were exposed to a 16-day season. The sample of wings received in 1981 (142) was more than double that received in 1980 (63 wings) and much greater than that received in any single year since 1976 when we began collecting grouse wings in Routt and Moffat counties. Wings were received from 3 check stations (Cedar Mountain = 1, Cal ifornia Park Road = 35, Twentymile Park Road = 38), 8 wing barrels (Ralph White Reservoir = 5, California Park = 2, Twentymile Park = 6, Steamboat Springs = 0, Oak Creek 12, Moffat Co. Road 3 = 2, Wolcott = 3), and field checks of hunters (24 wings). ~o sharptail wings were received at 1 check station (Steamboat Springs) and 4 wing barrels (Hahns Peak, Black Mountain Road, Milner, Highway 131). Most wings (104, 73.2%) were from the initial weekend of the hunting season, 26 (18.3%) from the 1st week, 7 (4.9%) from the 2nd weekend, 3 (2.1%) from the 2nd week, and 2 (1.4%) from the 3rd weekend. Age was ascertained for all wings examined and gonadal inspection of 60 sharptails provided sex ratios of the harvest (Table 2). The percentage of birds in the yearling age category is a minimum estimate as 75% of the adults and yearlings had molted primary 10. Although samples are small, it appears that the overall sex ratio approximates unity. �249 Table 2. Age composition of harvested sharp-tailed grouse, northwestern Colorado, 1981. Males Age class N Adults Yearlings Ch icks 76 7 59 a N % 14 0 13 53.5 4.9 41.6 a Females % 46.7 0 46.4 N a % 16 2 15 53.3 100.0 53.6 Only those for which gonads were examined. The percentage of juveniles in the fall harvest (41.6%) was less than the previous 3 years indicating production was only moderate in 1981. Data on age and sex composition of the hunter harvest for all years for which data are available are presented for comparison in Table 3. Table 3. Age and sex composition of harvested sharp-tailed grouse, northwestern Colorado, 1976-81. Year Adultsa N % N 1976 1977 1978 1979 1980 1981 10 47 13 32 25 83 4 25 15 47 38 59 71.4 65.3 46.4 40.5 39.7 58.4 Totals 211 6-year average Young % 28.6 34.7 53.6 59.5 60.3 41.6 14 72 28 79 63 142 5 5 1 3 14 397 23 186 53.1 b Adults Males Females Total sample 4 Young b Males Females 6 4 18 7 3 13 10 3 15 27 29 32 46.9 alncludes yearlings. b Known sex only. LITERATURE CITED Aldrich, J. W. 1963. Geographic orientation of American Tetraonidae. J. Wildl. Manage. 24:529-545. Giesen, K. M., and D. M. Hoffman. 1981. Distribution and status of mountain sharp-tailed grouse. Colo. Div. Wildl. Final Rep. Fed. Aid. Proj. W-37-R-34. Apr. 1981. Pp. 183-189. �250 Hart, C. M., O. S. Lee, and J. B. Low. 1950. The sharp-tailed grouse in Utah. Its life history, status, and management. Utah Dep. Fish and Game, Fed. Aid. Diy. Publ. 3. 79pp. Kessler, W. B., and R. P. Bosch. 1981. Sharp-tailed grouse and range management practices. Pages 133-146 in Proc. Wildl./Liyestock Symp., Coeur d'Alene, Idaho. Miller, G. C., and W. D. Graul. 1980. Status of sharp-tailed grouse in North America. Pages 18-28 in P. A. Vohs, Jr., and F. L. Knopf, eds. Proc. Prairie Grouse Symp. Okla. State Uniy., Stillwater. Rogers, G. E. 1969. The sharp-tailed grouse in Colorado. Game, Fish, and Parks, Tech. Publ. 23. 94pp. Prepared by: Kenneth M. Giesen Wildlife Researcher B Colo. Diy. �251 JOB PROGRESS REPORT State of Colorado Project No. W-37-R-35 Work Plan No. Job No. Population Dynamics of White-tailed Job Title: Period Covered: Personnel: 17 Game Bird Survey 7 Ptarmigan 1 January - 31 December 1981 Clait E. Braun and Kenneth M. Giesen, Colorado Division of Wi ldl ife. ABSTRACT Long-term studies of populations of white-tailed ptarmigan (Lagopus leucurus) were continued at hunted (Mt. Evans) and unhunted (Rocky Mountain National Park) areas in Colorado in 1981. Densities of breeding ptarmigan continued to decrease (0.5 birds/km2 at Mt. Evans, 0.2 birds/ km2 at Rocky Mountain National Park) from highs experienced in 1979 (Mt. Evans) and 1976 (Rocky Mountain National Park). These decreases are in the magnitude of 13 and 39%, respectively at the 2 areas from the h~ghs in 1979 and 1976. Nesting success was poor (35.3%) at Mt. Evans in 198.1 and excellent (72.2%) at Rocky Mountain National Park. Average brood size to 1 September at the 2 areas was 3.3 and 4.0 chicks/successful hen, respectively. Hunters at Mt. Evans in fall 1981 removed at least 30% of the estimated fall population of ptarmigan. The increase in harvest from 23% in 1980 to > 30% in 1981 was attributed to the earlier opening date (1 week earlierT in 1981 than in 1980. ��253 POPULATION DYNAMICS OF WHITE-TAILED PTARMIGAN Clait E. Braun and Kenneth 'M. Giesen Long-term studies of trends in population size and investigation of reasons for fluctuations in size of tetraonid populations are lacking. Studies on the population dynamics of unhunted and hunted populations of white-tailed ptarmigan were initiated in Colorado in 1966 and have continued essentially uninterrupted at 2 sites. Studies of the unhunted population (Rocky Mountain National Park) have identified possible short-term cycles of 7-8 years with an amplitude of 25-30% between high and low breeding densities. Conversely, studies of the manipulated population (hunted) at Mt. Evans through 1980 have not indicated any cycl ic pattern and it would appear that controlled hunting may mask any long-term trend that may occur. This report covers the initial year of a 5-year study designed to examine the question whether white-tailed ptarmigan are truly cyclic and whether hunting affects the apparent oscillations. P. N. OBJECTIVES The goals of this investigation are to be able to predict the length and ampl itude of cycles in white-tailed ptarmigan in Colorado, to examine the impact of hunting on cycles, and to clarify underlying causes of the apparent cycles. SEGMENT OBJECTIVES 1. Conduct breeding (May-Jun) and brood (Aug-Sep) censuses of white-tailed ptarmigan using tape-recorded calls of males (breeding) and chicks (brood). 2. Censuses 'will be conducted on previously established, defined study areas at Mt. Evans (hunted) and at Rocky Mountain National Park (unhunted). 3. Capture (noose poles) and band (aluminum and plastic color-coded bands) all unmarked white-tailed ptarmigan encountered on study areas at Mt. Evans and at Rocky Mountain National Park. 4. Individually identify all ptarmigan observed on study areas at Mt. Evans and Rocky Mountain National Park through use of binoculars. 5. Make hunting season and bag 1imit recommendations for Mt. Evans and collect hunting data through use of volunteer wing barrels and hunter field checks. 6. Compile data, analyze results and prepare progress report. �254 STUDY AREA AND METHODS Areas investigated were at Mt. Goliath-Mt. Evans in Clear Creek County and at Tombstone Ridge-Sundance Mountain to Fall River Pass in Rocky Mountain National Park in Larimer County. The physiography, geology, location, and vegetation on these study areas has been previously described (Braun 1969, 1971; Braun and Rogers 1971; Giesen 1977). Ptarmigan were located through use of tape-recorded calls (Braun et al. 1973), captured through use of telescoping noose poles (Zwickel and Bendell 1967) as described by Braun and Rogers (1971), classified as to age and sex and banded following Braun and Rogers (1971). Age of chicks was estimated following Giesen and Braun (1979). Numbered plastic bandettes were not used as in earlier years (Braun and Rogers 1971) as a color-code system using up to'4 different colored plastic bandettes was instituted in 1977-78. A check station was operated on the Mt. Evans highway during the opening weekend (19-20 Sep) of the ptarmigan season in that area. A volunteer wing collection station was available to hunters in the area from 20 September through 4 October when the season closed. RESULTS AND DISCUSSION Breeding Densities Mt. Evans Surveys of breeding white-tailed ptarmigan on the Mt. Evans study area in Spring 1981 (May-Jun) revealed the presence of at least 15 pairs and 6 unmated males. This is a density of 9.0 birds/km2, a slight decrease from 1980 and the high recorded in 1979 (Table 1). Pairing and breeding activities were initiated in early May, earlier than in 1980 because of the lack of snow cover in 1981. Rocky Mountain National Park Surveys of ptarmigan present on the Rocky Mountain National Park study units during May and June 1981 indicated that 16 pair~ and 13 single males were present. This is a density of 8.2 birds/km2, similar to the 8.4 birds/ km2 recorded in 1980 (Table 1). Nesting Success and Brood Size Mt. Evans During the July-early September interval, 6 different hens were identified with broods (average size to 1 Sep = 3.3 chicks/successful hen) while 11 hens were observed without broods. Thus, only 6 of 17 hens (35.3%) were known to be successful in nesting. Estimated hatching dates for 23 chicks for which data are available ranged from 29 June to 13 August with most (17 of 23) hatching between 30 June and 6 July, earl ier than the 19-20 July peak calculated in 1980. �255 Table 1. White-tailed ptarmigan breeding densities, Colorado 1966-81. Studz: Area Year Rocky Mountain National Park (5.5 km2) 1966 1967 1968 1969 1970 1971 1972 1973 1974 1975 1976 1977 1978 1979 1980 1981 11.3 9.8 11.5 12.0 9.6 9.1 8.7 7.8 8.0 11. 1 13.5 12.9 10.7 8.7 8.4 8.2 Mt. Evans (4.0 km2) 3.0 2.7 2.7 2.2 2.0 4.2 7.5 6.2 6.2 6.2 6.7 >6.0 7.5 10.3 9.5 9.0 Rockz: Mountain National Park Thirteen successful hens (13/18 = 72.2%) were observed between 1 July and early September with an average brood size to 1 September of 4.0 chicks. Only 5 hens were observed without broods in 1981 (5/18 = 27.8%). Estimating hatching dates for 32 chicks ranged from 3 July to 10 August with most (29 of 32) hatching between 3 and 16 July. Harvest Mt. Evans In 1981, the ptarmigan season in the Mt. Evans area (Unit 52 south of Interstate 70 and east of the Guanella Pass Road between Georgetown and Grant) was delayed 1 week after opening of the statewide season on 12 September. This was a 7 day shorter delay than that used in the 1977-80 period following the experimentation with season length and limited closure from 1972 through 1976. This experimentation followed the documentation of overharvest in this area in the late 1960's and the 2-year closure in 1970 and 1971. In 1981, the season opened one-half hour before sunrise on 19 September and closed at sunset on 4 October. This was the earliest closure of the statewide ptarmigan season since at least 1977. �256 A wing collection barrel placed along the Mt. Evans highway prior to opening of the season was available until after 4 October. Because of the mild fall, the road remained open until 1 October when it was closed by the state highway department because of an agreement with the CDOW. A check station was operated both days of the opening weekend. During the 2 days of check station operation, 20 hunters with 18 ptarmigan were checked. This compares to 12 hunters with 9 ptarmigan and 9 hunters with 2 ptarmigan checked in the same period (1 week later) in 1980 and 1979, respectively. An additional 7 wings were deposited in the wing barrel on 25 September (6) and 27 September (1). Wings from 7 additional ptarmigan were sent to Fort Collins and 1 banded bird was reported by mail. In all, at least 33 ptarmigan were harvested in 1981 of which at least 10 were banded and 16 (all unbanded) were chicks. The fall popUlation resident in the area was estimated to be about 56 birds (36 adults, 20 chicks). Most unbanded birds shot were not residents of the area studied or were chicks. The total fall population (residents and nonresidents) in the area hunted was estimated at 100 + 10 birds. Thus the harvest of 33 birds represented 30.0 to 36.7% of the fall population. This was the highest estimated removal of the fall population since 1974 when season length and area hunted were experimentally controlled. The increase in harvest in 1981 was higher than during the 2-week delayed season in 1978-80. The harvest experienced in fall 1981. if not actually higher, is near the upper limit that should be allowed if the breeding density is to remain stable. If the 1 week delayed season is maintained, it is anticipated that overharvest of the ptarmigan population along the Mt. Evans highway will occur within 1 or 2 years. The 2-week delayed season concept worked well at Mt. Evans from 1978 through 1980 despite the road being open to vehicular traffic. In practice, closing the road on 1 October has no measurable impact on the magnitude of the pt~rmigan harvest. If the goal is to reduce the breeding density of ptarmigan al6ng the Mt. Evans highway, the 1 week delayed season should be continued. If the goal is to maintain a stable breeding density, the season should be delayed at least 2 weeks. The recommendation is to return to a 2-week delay with the season continuing through the Sunday closest to 10-12 October. This would allow hunting during 1 more weekends on a statewide basis than in 1981. LITERATURE CITED Braun, C. E. 1969. Population dynamics, habitat, and movements of whitetailed ptarmigan in Colorado. Ph.D. Thesis. Colo. State Univ., Fort Collins. 189pp. 1971. Habitat requirements of Colorado white-tailed Proc. West. Assoc. State Game and Fish Comm. 51 :284-292. , and G. -----::-Colo. Div. E. Rogers. 1971. The white-tailed ptarmigan Game, Fish and Parks Tech. Publ. 27. 80pp. ptarmigan. in Colorado. �257 , R. K. Schmidt, Jr., and ---white-tailed ptarmigan with G. E. Rogers. 1973. tape recorded calls. Census of Colorado J. Wildl. Manage. 37:90-93. Giesen, K. M. 1977. Mortality and dispersal of juvenile white-tailed ptarmigan. M.S. Thesis. Colo. State Univ., Fort Collins. 55pp. __ --:-_'and C. E. Braun. juvenile white-tailed 1979. A technique for age determination of ptarmigan. J. Wildl. Manage. 43:508-511. Zwickel, F. C., and J. F. Bendell. 1967. A snare for capturing blue grouse. J. Wildl. Manage. 31:202-204. Prepared by --~~~-=-~-----------------Clait E. Braun Wildlife Research Leader Kenneth M. Giesen Wildlife Researcher ��259 JOB PROGRESS REPORT State of Colorado --------------------------- Project No. Work P Ian No. 22 ---------------------- Job Title: Job No. Upland Game Publications Period Covered: Pe rsonne 1: Game Bird Survey W-37-R-35 1 April 1981 through 30 June 1982 C. E. Braun, B. S. Cade, Hoffman, T. E. Remington, Division of Wildlife; N. Colorado College; and R. sity. P. O. Dunn, K. M. Giesen, R. W. and T. J. Schoenberg, Colorado J. Kitzmiller and R. M. Stabler, A. Ryder, Colorado State Univer- ABSTRACT Publ ications accomplished under this job in Segment 35 are: Autenrleth, R., W. Molini, and C. Braun. Editors. 1982. Sage grouse management practices. West. States Sage Grouse Comm. Tech. Twin Falls, Idaho. Bull. 1. 42pp. Braun, C. E. 1981. Sage grouse population trends and dynamics in Colorado. Proc. Bien. West. States Sage Grouse Workshop. 12:Abstract. 1981. Tapeworms of birds. Pages 77-80 in W. J. Adrian, ed., Manual of common wildlife diseases in Colorad~ Colo. Div. Wildl., Denver. Cade, B. S., R. A. Ryder, and R. W. Hoffman. 1982. Blue grouse movements from breeding to wintering areas. J. Colo.-Wyo. Acad. Sci. 14(1): 54. (Abstract) . Dunn, P.O., and R. A. Ryder. 1982. Summer movements of juvenile sage grouse. J. Colo.-Wyo. Acad. Sci. 14(1):5~. (Abstract). Giesen, K. M., and C. E. Braun. 1982. Dispersal of juvenile white-tafled ptarmigan in Colorado. Cooper Ornithol. Soc. Annu. Mtg. 52:27. (Abstract) . , and T. J. Schoenberg. 1981. Techniques for trapping sage grouse in Colorado. Proc. Bien. West. States Sage Grouse Workshop 12:Abstract. ---- �260 Hoffman, R. W. 1981. Volunteer collection station use for obtaining grouse wing samples. Wi ldl. Soc. Bull. 9: 180-184. , and B. S. Cade. ----:-Colo. Fie 1 d 0 rnit ho 1. 1982. Occurrence of sage grouse above treeline. J. 16: 22- 23 . Remington, T. E. 1981. Winter nutrition of sage grouse in North Park, Colorado. Proc. Bien. West. States Sage Grouse Workshop 12:Abstract. 1982. Winter browse preference and nutrition of sage grouse in ---!i52~. 61-b':.,-------~------------I'IG-F-t-fl--P--a-F-k,--Ge-l-G-FaOO-.----Gee~-o;--trk:-he-h-Se~-t_g:--:-. (Abstract). Schoenberg, T. J. 1981. Sage grouse habitat selection in North Park, Colorado. Proc. Bien. West. States Sage Grouse Workshop 12:Abstract. 1982. Sage grouse habitat selection in North Park, Colorado. Cooper Ornithol. Soc. Annu. Mtg. 52:71. (Abstract). 1982. Sage grouse movementsandhabitat selection in North Park, Colorado. M.S. Thesis, Colo. State Univ., Fort Collins. 86pp. Snyder, W. D. 1981. I 1ike weeds. Colo. Outdoors 30(6): 10-13. 1981. Nest habitat selection by ring-necked pheasants in northeast Colorado. Midwest Fish and Wildl: Conf. 43:93 (Abstract). 1982. Recommended habitat management practices for pheasants in eastern Colorado. Colo. Div. Wildl.,Outdoor Facts 82. 4pp. (Revised). Stabler, R. M., N. J. Kitzmiller, and C. E. Braun. 1981. Redescription of Eimeria centrocerci from sage grouse (Centrocercus urophasianus). Trans. Am. Micros. Soc. 100:86-89. Prepared by _~~a~~:=:...,·::--"z.",--.-",,~==~_ Clait E. Braun Wildlife Research Leader � https://cpw.cvlcollections.org/files/original/1c88208a1bb5ef4ce281602a8a932e38.pdf 73124c7539783ccc65e0fd6dd1eb416f PDF Text Text 1 JOB PROGRESS State of July, 1982 REPORT Colorado Project No. W-126-R-S· Big Game Investigations Work Plan No. Multispecies ---.----J 0 b--No-. Investigations S·--------- ---:--·--Exp·e:r lme:Iital- ImptoVemeIiC·of-Oak.15t"ush--on--------··-Deer, Elk and Cattle Ranges - Hightower Mountain --------. Period Covered: Personnel: 1 - Cervids July 1, 1981 through June 30, 1982 R. Kufeld ABSTRACT Ten year post-treatment vegetation measurements were made in 600 permanent meter square plots, and deer, elk and cattle response to burning, spraying and chaining was measured in 480 pellet group plots. Final data analysis is underway and results will be presented in a future report. - ��3 EXPERIMENTAL IMPROVEMENT OF OAKBRUSH ON DEER, ELK, AND CATTLE RANGES - HIGHTOWER MOUNTAIN Roland C. Kufeld P. N. OBJECTIVE To determine the extent to which the production and quality of deer, elk and cattle forage and livestock and wildlife use can be increased and maintained by chaining, spraying and burning overage Gambel Oak winter ranges. SEGMENT OBJECTIVES To determine the extent of vegetative composition and forage production changes, and the extent of deer and elk use changes which have resulted from implementation of each habitat improvement method tested. Determine if cattle exhibit a preference toward certain habitat improvement units. ACKNOWLEDGEMENTS The following people provided field assistance C. Cupp, E. Hashimoto, and J. Lile. METHODS Post-Treatment on this study: J. Buckholty, AND MATERIALS Vegetation Measurements Ten year post-treatment vegetation measurements were made in 600 permanent meter square plots on the Hightower Mountain Oak study area using the same procedures employed during pre-treatment evaluations (Kufeld 1971). The measurement period was June 30, 1981 through July 30, 1981. Prior to vegetation measurements observers were trained in the use of the technique, and in identification of all plants on the study area. Photo Point Photographs Post-treatment photos were taken at permanent photo points established in 1970, prior to treatment. These show effects of spraying, burning and chaining in the 10th summer following treatment. A black and white photo, and a 35 mm color slide were taken at each photo point with the same cameras used to take pre-treatment photos. �4 Post-Treatment Deer, Elk and Cattle Use Measurements Accumulated deer and elk pellet groups were removed from all pellet plots on August 26, 1981. Groups deposited during the winter of 1981~82 were counted My 14, 1982. When plots were cleared on August 26, 1981, a record was kept of the number of cow chips in each plot in order to provide an index to cattle use. Only those chips which could be identified as having been deposited during the summer of 1981 were counted. RESULTS AND DISCUSSION Vegetation production and deer, elk and cattle use data collected during this segment are currently undergoing analysis. Results of the lO-year post-treatment evaluation will be presented in a Division of Wildlife Technical Bulletin which is presently being prepared. LITERATURE CITED Kufeld, R. C. 1971. Experimental improvement of oakbrush on deer, elk and cattle ranges - Hightower Mountain. Colo. Div. of Game, Fish and Parks, Game Res. Rept. July (1):23-86. Prepared by ~I C. Roland C. Kufeld Wildlife Researcher J!c{~ee/ ' C �5 JOB PROGRESS No. Big Game Investigations W-126-R-5 Work Plan No. ~----Jbb- REPORT Colorado State of Project July, 1982 1 _ Multispecies 7--------:~ Big-Game-------~-------------- No .---- Period Covered: Personnel: - Cervids Investigations Research-Publicati-ons-- July 1, 1981 through June 30, 1982 L. H. Carpenter ABSTRACT During the 1981-82 Segment the Big Game Research Section had 12 publications published, 4 others accepted for publication and 4 manuscripts in the review process. ��7 BIG GAME RESEARCH PUBLICATIONS Len H. Carpenter P. N. OBJECTIVE To publish the results of research conducted under the auspices of Federal Aid Projects W-126-R and W-144-R in a variety of professional journals and other indexed publishing media to insure widespread dissemination and availability of this information to natural resource managers and ecological scientists. SEGMENT OBJECTIVES 1. Federal Aid Job Progress Reports. All studies. 2. Kufeld, R. C., M. Stevens, and D. C. Bowden. Winter variation in nutrient and fiber content and in-vitro digestibility of mountain mahogany (Cercocarpus montanus). and serviceberry (Amelanchier'alnifolia) from diversified sites in Colorado. J. Range Manage. 3. Bartmann, R. M., A. E. Alldredge, and P. H. Neil. 1982. Evaluation of winter food choices by tame mule deer. J. Wildl. Manage. (Accepted for publication in 1982). 4. Bartmann, R. M., and L. H. Carpenter. 1982. Effects of foraging experience on food selectivity of tame mule deer. (Accepted publication in 1982). 5. Bartmann, R. M. mule deer. 6. Bartmann, R. M. and D. C. Bowden. A weather mule deer winter mortality in Colorado. 7. Bartmann, R. M. Mule deer winter diet composition and quality juniper range. Colo. Div. Wildl. Spec. Rept. 8. Freddy, D. J. Predicting Wildl. Manage. 9. Freddy, D. J., and D. C. Bowden. in juniper-pinyon woodland. Accuracy evaluation J. Wildl. Manage. of the helicopter mule deer harvest quadrat for census for severity index for estimating J. Wildl. Manage. in Middle on pinyon- Park, Colorado. J. Sampling mule deer pellet group densities J. Wildl. Manage. 10. Freddy, D. J., and D. C. Bowden. Efficacy of permanent and temporary pellet plots in juniper-pinyon woodland. J. Wildl. Manage. 11. Freddy, D. J. Heart rates for activities Compo Biochem. and Physiol. 12. Milchunas, D. G., and D. L. Baker. 1981. within and between trial variability. for publication in 1981). of mule deer at pasture. J. In vitro digestion sources of J. Range Manage. (accepted �8 1982. Composition J. Wildl. Manage. and quality of elk 13. Baker, D. L., and N. T. Hobbs. summer diets in Colorado. 14. Carpenter, L. H. Twenty-four hour activity pasture. J. Wildl. Manage. 15. Hobbs, N. T., D. L. Baker, and R. B. Gill. Comparative nutritional ecology of montane ungulates in Colorado. J. Wildl. Manage. 16. Gill, R. B., L. H. Carpenter, R. M. Bartmann, and D. L. Baker. Estimating mule deer diets from bitecount and fecal analysis. J. Range Manage. 17. Hobbs, N. T., D. L. Baker, J. E. Ellis, and D. W. Swift. 1981. Energy and nitrogen based estimates of elk winter range carrying capacity. J. Wildl. Manage. 18. Hobbs, N. T., and D. B. Bowden. indexes. J. Wildl. Manage. 19. Carpenter, L. H., R. B. Gill, and D. L. Baker. Tests of a nutritionally based habitat evaluation system. Colo. Div. Wildl. Spec. Rept. Confidence patterns of mule deer at intervals on preference ACKNOWLEDGEMENTS All scientists and graduate students on the Big Game Research commended for their excellent publishing records. PUBLICATION 1. 2. Federal Aid Job Progress Reports. All studies. in: Colorado 1982. of Wildlife. Wildl. Res. Repts. July Part 1 and 2. Kufeld, R. C., M. Stevens, and D. C. Bowden. Winter variation in nutrient and fiber content and in-vitro digestibility of mountain mahogany (Cercocarpus montanus) and serviceberry (Amelanchier alnifolia) from diversified sites in Colorado. J. Range Manage. Data analyses have been completed has not yet begun. 3. PROGRESS These reports have been printed Division Staff are to be for this manuscript, but the narrative Bartmann, R. M., A. W. Alldredge, and P. H. Neil. 1982. Evaluation winter food choices by tame mule deer. J. Wildl. Manage. This manuscript was published of as: Bartmann, R. M., A. W. Alldredge, and P. H. Neil. 1982. Evaluation of winter food choices by tame mule deer. J. Wildl. Manage. 46(3):807-812. �9 4. Bartmann, R. M., and L. H. Carpenter. 1982. Effects of foraging experience on food selectivity of tame mule deer. This manuscript was published as: 1982. Effects of foraging experBartmann, R. M., and L. H. Carpenter. ience on food selectivity of tame mule deer. J. Wildl. Manage. 46(3):813-818. 5. Bartmann, R. M. Accuracy evaluation of the helicopter for mule deer. J. Wildl. Manage. A 1st draft of this manuscript 6. has been initiated. Bartmann, R. M. and D. C. Bowden. A weather mule deer winter mortality in Colorado. This manuscript quadrat census has been submitted severity index for estimating J. Wildl. Manage. to J. Wildl. Manage as: Bartmann, R. M., and D. C. Bowden. Mule deer winter mortality juniper range in Northwest Colorado. 7. Bartmann, R. M. Mule deer winter diet composition and quality on pinyonjuniper range. Colo. Div. Wildl. Spec. Rept. This manuscript 8. has been submitted to J. Range Manage as: Bartmann, R. M. Mule deer diet composition range. J. Range Manage. and quality on pinyon-juniper Freddy, D. J. Predicting Wildl. Manage. in Middle Park, Colorado. This manuscript mule deer harvest has been published 10. Freddy, D. J., and D. C. Bowden. in juniper-pinyon woodland. in Middle Park, Colorado. Sampling mule deer pellet group densities J. Wildl. Manage. Freddy, D. J., and D. C. Bowden. Efficacy of permanent and temporary pellet plots in juniper-pinyon woodland. J. Wildl. Manage. These manuscripts 11. J. as: Freddy, D. J. 1982. Predicting mule deer harvest J. Wildl. Manage. 46(3):802-806. 9. on pinyon- have been accepted for publication Freddy, D. J. Heart rates for activities Compo Biochem. and Physiol. This manuscript has been completed by the J. Wildl. Manage. of mule deer at pasture. J. and is in the internal review process. �10 12. Milchunas, D. G., and D. L. Baker. 1981. within and between trial variability. This manuscript has been published In vitro digestion J. Range Manage. sources of as: Milchunas, D. G. and D. L. Baker. 1982. In vitro digestion-- sources of within - and between - trial variability. J. Range Manage. 35 (1):199-203. 13. Baker, D. L., and N. T. Hobbs. summer diets in Colorado. This manuscript has been published Baker, D. L., and N. T. Hobbs. summer diets in Colorado. 14. of elk as: 1982. Composition and quality of elk J. Wildl. Manage. 46(3):694-703. patterns of mule deer at Hobbs, N. T., D. L. Baker, and R. B. Gill. Comparative nutritional ecology of montane ungulates in Colorado. J. Wildl. Manage. has been accepted for publication by the J. Wildl. Manage. Gill, R. B., L. H. Carpenter, R. M. Bartmann, and D. L. Baker. Estimating mule deer diets from bitecount and fecal analysis. J. Range Manage. This manuscript 17. and quality was made on this manuscript. This manuscript 16. Composition Carpenter, L. H. Twenty-four hour activity pasture. J. Wildl. Manage. No progress 15. 1982. J. Wildl. Manage. to J. Wildl. Manage. has been submitted Hobbs, N. T., D. L. Baker, J. E. Ellis, and D. W. Swift. 1981. Energy and nitrogen based estimates of elk winter range carrying capacity. J. Wildl. Manage. This manuscript has been published as: Hobbs, N. T., D. L. Baker, J. E. Ellis, D. M. Swift, and R. A. Green. 1982. Energy - and nitrogen - based estimates of elk winter range carrying capacity. J. Wildl. Manage. 46(1):12-21. 18. Hobbs, N. T., and D. B. Bowden. indexes. J. Wildl. Manage. This manuscript Confidence has been published Hobbs, N. T., and D. C. Bowden. indices. J. Wildl. Manage. intervals on preference as: 1982. Confidence 46(2):505-507. intervals on preference �11 19. Carpenter, L. H., R. B. Gill, and D. L. Baker. Tests of a nutritionally based habitat evaluation system. Colo. Div. Wildl. Spec. Rept. No progress was made on this manuscript. Additional publications which were not scheduled but whic~ were published accepted for publication during the 1981-82 segment are l~sted below. Geduldig, H. L. 1981. Summer home range of mule deer fawns. Manage. 45(3):726-727. Hobbs, T., and R. Spowart. 1982. Fire. Colo. Outdoors. or J. Wildl. 31(4):26-29. Parkinson, D. E., R. P. Ellis, and L. D. Lewis. 1982. Colostrum deficiency in mule deer fawns: identification, treatment and influence on noenatal mortality. J. Wildl. Dis. 18(1):17-28. Renecker, L. A., R. J. Hudson, and D. J. Freddy. 1982. Heart rate as an index of energy expenditure in moose using two telemetry systems. Intn. Symp on Biotelemetry, Proc. 7: Stanford, Calif. (Abstract). Torbit, S. C., D. M. Swift, A. W. Alldredge, L. H. Carpenter, and J. E. Ellis. 1982. Catabolic dynamics of mule deer energy reserves. AIBS Annual Proc:33 (Abstract). White, G. C., and R. M. Bartmann. 1983. Estimation from band recoveries of mule deer in Colorado. (accepted for publication). Prepared by: Len H. CarpentJr Wildlife Research Leader of survival rates J. wildl. Manage ��13 JOB PROGRESS State of REPORT Colorado Project No. W__ -_1_2_6-_R___5 Work Plan No. ---------Job-No. Period July, 1982 1 _ Big Game Investigations Multispecies - Cervids Investigations 8-------------:--Big--Game-Publication-Edit:ing--and--------------Library Services Covered: Personnel: July 1, 1981 through June 3D, 1982 M. Hershcopf, L. Carpenter ABSTRACT During Segment 5 of W-126-R, 26 books and theses were purchased f'or permanent reference by DOW researchers. Fifty additional publications were located, ordered, and obtained free of charge for use. Twenty theses were obtained on interlibrary loan and 4 computer-aided literature searches were completed for the library files. An additional 785 individual references requested by big game researchers were located by library staff and made available for reference. .-- ��15 BIG GAME PUBLICATION EDITING AND LIBRARY Marian W. Hershcopf SERVICES and Len H. Carpenter P. N. OBJECTIVE To provide a centralized support program for Big Game Research technical editing and library services so that Big Game Research scientists can allocate additional time to the conduct of actual research. SEGMENT OBJECTIVES To provide coordinated, efficient, and economic editing and library services to all Colorado Big Game Research programs (Federal Aid Projects W-126-R and W-144-R). ACKNOWLEDGEMENTS N. McEwen, effort. C. Pankonin, S. Wielgopolan, and C. Doty all contributed to this Silln1ARYOF SERVICES Publications Purchased with W-126-R Funds and Placed in Research Center Library Albrecht, R. L., L. Finkel, and J. R. Brown. 1978. John Wiley & Sons, Inc., New York. 32spp. Banfield, A. W. F. 1981. The mammals Press, Toronto. 438pp. of Canada. BASIC. 2nd edition. University of Toronto Clow, D. J., and N. S. Urquhart. 1974. Mathematics in biology: calculus and related topics. W. W. Norton & Co., Inc., New York. 727pp. Crawford, R. L. 1981. Lignin biodegradation Wiley & Sons, Inc., New York. ls4pp. and transformation. John Donald, A. D., W. H. Southcott, and J. K. Dineen, eds. 1978. The epidemiology and control of gastrointestinal parasites of sheep in Australia. Commonwealth Scientific and Industrial Research Organization. Div. Animal Health, Victoria, Australia. ls3pp. Engdahl, G. R. 1976. Techniques for determining intake by grazing animals. Ph.D. Thesis. Texas A & M Univ., College Station. 72pp. Errington, P. L. Ames, Iowa. 1967. Of predation 277pp. Fowler, C. W., ed. 1981. Dynamics Wiley & Sons, Inc., New York. and life. Iowa State Univ. Press. of large mammal populations. 477pp. John �16 Hartnell, G. F. 1977. Measurement and significance of ingesta turn-over rates in dairy cattle using rare-earth elements. Ph.D. Thesis. Univ. of Wisconsin, Madison. l78pp. Jaeger, E. C. 1978. A source book of biological names and terms. Charles C. Thomas, Publisher, Springfield, Ill. 323pp. 3rd ed., Lascano, C. E. 1979. Determinants of grazed forage voluntary intake in cattle. Ph.D. Thesis. Texas A & M Univ., College Station. 200pp. Lehner, P. N. 1979. Handbook New York. 403pp. of ethological methods. Garland STPM Press, Lentner, C., ed. 1981. Geigy scientific tables. Vol. 1. Units of measurements, body fluids, composition of the body, n.utrition. 8th ed. Ciba-Geigy Corp., W. Caldwell, N.J. 295pp. Moen, A. N. 1982. The biology and management of wild ruminants; a sequence of learning experiences in seven parts and twenty-five chapters. Pt. 2 Behavior of wild ruminants, Pt. 5 - Meteorology and thermal relationships of wild ruminants, Pt. 7 - The management of wild ruminants. Corner Brook Press, Lansing, New York. Moore, T. D. 1974. Identification of the dorsal guard hairs of some mammals of Wyoming. Wyoming Game & Fish Dep. Bull. No. 14. Cheyenne. l77pp. Muller, F. M., J. G. Dijkhuis, and S. Heida. 1973. On the relationship between chemical composition and digestibility in vivo of roughage 2. Centre for Agricultural Publishing and Documentation. National Wildlife Federation. 1979. Bobcat Research Conference Proceedings. Current research on biology and management of Lynx rufus, October 16-18. 1979, Front Royal, Va. National Wildlife Federation, Scientific and Tech. Series No.6. l37pp. Peterson, R. L. 1979. Toronto. 280pp. North American moose. Univ. of Toronto Press, Pieper, R. D. 1978. Measurement techniques for herbaceous and shrubby vegetation. New Mexico State Univ., Dep. of Animal and Range Sciences, Las Cruces. l47pp. Poole, R. W. 1974. An introduction Inc., New York. 532pp. to quantitative ecology. McGraw-Hill, Rogers, L. L. 1977. Social relationships, movements, and population dynamics of black bears in northeastern Minnesota. Ph.D. Thesis. Univ. of Minnesota, Minneapolis. 194pp. Schmidt, G. D., and L. S. Roberts. 1977. C. V. Mosby Co., St,; Louis. 604pp. Foundations of parasitology. �17 Thomas, J. W., and D. E. Toweill. 1982. Elk of North America: and management. Stackpole Books, Harrisburg, Pa. 720pp. Tomek, I. 1981. Introduction to computer organization. Press, Inc., Rockville, Maryland. 456pp. Ecology Computer Van Soest, P. J. 1982. Nutritional ecology of the ruminant. Inc., Corvallis, Oregon. 374pp. Science 0 & B Books, Willms, W. D. 1979. The effects of fall burning or grazing on Agropyron spicatum «Pursh) Schribn. + Smith) and its selection by deer and cattle. Ph.D. Thesis. Univ. of Alberta, Edmonton. l64pp. Publications Obtained. Free or at Low Cost In addition to books purchased with W-126-R funds, about 50 free reports and short publications from state or federal agencies, and from other sources, were located, ordered and obtained for use by big game research personnel. Theses Obtained on Interlibrary Loan for Use by Researchers Abdel-Rasoul, K. S. 1979. The epidemiology of toxascariasis and baylisascariasis in wild carnivores in captivity. Ph.D. Thesis. of California, Davis. 107pp. Berger, J. 1978. Social development and reproductive strategies Univ. of Colorado, Boulder. l57pp. sheep. Ph.D. Thesis. Univ. in bighorn Brooks, J., III. 1981. Comparison of in vivo and in vitro forage digestibility by elk. M.S. Thesis. Utah State Univ., Logan. 115pp. DuBrock, C. W. 1980. An analysis of Virginia black bear population dynamics. M.S. Thesis. Virginia Poly tech. Institute & State Univ., Blacksburg. 115pp. Eager, D. C. 1977. Radioisotope feces tagging as a population estimator black bear (Ursus americanus) density in the Great Smoky Mountains National Park. M.S. Thesis. Univ. of Tennessee, Knoxville. 89pp. Evans, W. 1975. Methods of estimating densities of white-tailed Texas A & M Univ., College Station. 185pp. Ph.D. Thesis. Goodson, N. 1978. Status of bighorn sheep in Rocky Mountain Park. M.S. Thesis. Colorado State Univ., Fort Collins. of deer. National Guynn, D. E. 1979. Management of deer hunters on private land in Colorado-1979. Ph.D. Thesis. Colorado State Univ., Fort Collins. Hamilton, R. J. 1978. Ecology of the black bear in southeastern Carolina. Ph.D. Thesis. Univ. of Georgia, Athens. 2l4pp. North �18 Iskander, F. D. 1973. Factors affecting feeding habits of sheep grazing foothills ranges in northern Utah. Ph.D. Thesis. Utah State Univ., Logan. 82pp. Jobman, W. G. 1972. Consumption of juniper by deer and inhibition of rumen micro-organisms by volatile oils of juniper. M.S. Thesis. Colorado State Univ., Fort Collins. Kirchner, T. B. 1980. Community structure in relation to body size of species. Ph.D. Thesis. Colorado State Univ., Fort Collins. Kvale, C. T. 1981. Mule deer, elk and cattle relationships on the Herd Creek rest-rotation grazing system, East Fork of the Salmon River, Idaho. M.S. Thesis. Univ. of Idaho, Moscow. McKell, C. M. gambe1ii) 1950. A study of plant succession in the oak brush (Quercus zone after fire. M.S. Thesis. Univ. of Utah, Salt Lake City. Marcum, L. C. 1974. An evaluation of radioactive feces tagging as a technique for determining population densities of the black bear (Ursus americanus) in the Great Smoky Mountains National Park. M.S. Thesis. Univ. of Tennessee, Knoxville. 95pp. Narjisse, H. 1981. Big sagebrush acceptability to sheep and goats, its relationship to monoterpene smell, taste and effect on rumen microbial activity. Ph.D. Thesis. Utah State Univ., Logan. Owens, T. E. 1981. Habitat requirements of white-tailed deer in the Palouse range of Idaho. M.S. Thesis. Un Iv , of Idaho, Moscow. Painter, W. W. 1971. Volatile oil content and composition of juniper and its effect on deer rumen microorganisms. M.S. Thesis. Colorado State Univ., Fort Collins. Teeter, R. G. 1981. Indigestible markers: methodology and application in ruminant nutrition. Ph.D. Thesis. Oklahoma State Univ., Stillwater. Vilkitis, J ..R .. 1968. Characteristics of big game violators and extent of their activities in Idaho. M.S. Thesis. Univ. of Idaho, Moscow. 202pp. Computer Literature Searches Obtained for Big Game Researchers Research Center Library Radio telemetry Toxicity, pharmacology Bear/cattle Publications and physiological interactions by Valerius Geist action of Compound 1080 by the �19 Reference Document Location and Delivery The Research Center Library staff also located and delivered about 785 individual references on request for the Big Game Research section during this segment; about 35 of these were not available locally and were obtained through interlibrary loan procedures. Prepared by M c-",,'(;.K>._ tJ. He.-y"'-{J)r~ Marian W. Hershcopf Librarian Wildlife Research Leader ��21 JOB PROGRESS State of __~C~o~l~Q~rwa~d~ou_ Project No. 2 ------------------------ Period Covered: Personnel: Big Game Investigations _ ----------JQb-No:--------l REPORT _ W-126-R-5 Work Plan No. July, 1982 - Cervids Deer Investigations :-Nutritional-Basisfor-Quant-ifying- ----------------Capacity of l..J'inter Ranges to Support Deer July 1, 1981 through June 30, 1982 D. J. Freddy ABSTRACT Plans to study activity patterns and habitat use of wild deer during winter were terminated because of financial and manpower constraints. Thus a study plan was not completed. As an alternative study, cover selection by tame deer at pasture will be investigated. The study plan is currently in initial stages of development and will be completed next segment with the intent of assessing cover choices of deer during winter 1982-83. ��23 NUTRITIONAL BASIS FOR QUANTIFYING CAPACITY WINTER RANGES TO SUPPORT DEER OF David J. Freddy P. N. OBJECTIVE To determine if a system can be developed ranges are capable of supporting. to estimate number of deer winter SEGMENT OBJECTIVES 1. Write a study plan for estimating activity wild mule deer during winter 1982-83. patterns and habitat use of METHODS AND MATERIALS The initial approach to study activity patterns and habitat use (using telemetry and intensive labor) was estimated to be quite costly. During the latter part of this segment it was apparent that level of funding needed to complete the project would not be available in the forthcoming fiscal year. Thus, plans to study activity patterns and habitat use of deer during winter were terminated and no study plan was completed. As an alternative study, it was decided to investigate the use of cover by tame deer at pasture. To date, only a partial literature review and conceptualization of a general study approach has occurred. A study plan will be completed in the forthcoming segment with the intent of initially studying cover preferences of tame deer during winter 1982-83. RESULTS AND DISCUSSION Deer seek cover as a thermal buffer and as an escape mechanism. As a thermal buffer, cover can reduce heat loss, and therefore energy expenditure, by reducing wind flow, radiant heat loss, and by functioning as an energy absorber (Moen 1973). To assess habitat value for deer it is necessary to understand what cover is so that systems can be developed to quantify cover. A habitat evaluation system based on food alone is inadequate, consequently, we must learn more about cover. A basic question involving the use of cover by deer is whether deer prefer different types of cover; that is, given a choice of either reducing wind flow or radiant heat loss, what would a deer select. Cover selection could be monitored with tame deer and mechanically supplying different, albeit artificial, cover choices. If deer show preferences for cover types, such preferences would likely vary with time of day and weather. Assuming selection occurs, specific weather parameters could be monitored simultaneously and possibly "trigger" points could be established as to when �24 and to what types of cover deer select. Furthermore, physiological parameters of deer such as surface, subcutaneous, and deep body temperatures, or heart rate could be monitored to understand patterns of heat flow in an animal using different cover types. These data would be important in developing a system to evaluate deer habitat. The approach proposed to initially study use of cover by deer is as follows: Within a 2-ha pasture, an area approximately 20 x 100 m will be denuded of native sagebrush vegetation. On this area, 5 isolation pens 20 x 20 m will be constructed using woven wire. Within each isolation pen, 1 tame deer will be maintained. Each, animal will serve as a replicate of cover treatments. Treatments will consist of providing 2 types of artificial cover simultaneously to each deer. Deer choices, weather parameters, and possibly physiological parameters will be monitored. This initial approach will, 1) establish whether tame deer will use and select for types of artificial cover and 2) under what atmospheric conditions different cover types are used. LITERATURE Moen, A. N. 1973. Wildlife and Co., San Francisco. Prepared ecology: 458pp. bY/L:£~%// 7 D1.vid 1./Freddy Wildlife Researcher C CITED an analytical approach. W. H. Freeman �29 JOB PROGRESS State of REPORT Colorado Project No. W~-=1=2~6_-~R~-~5~ Work Plan No. _ 2 -=---------------- Job No. 6 Period Covered: Personnel: July, 1982 Big Game Investigations - Cervids Deer Investigations Winter habitat selection and activity patterns of mule deer in Front Range shrubland and forest habitats July 1, 1981 through June 30, 1982 R. Kufeld ABSTRACT Ten habitat types common to the study area were delineated on an aerial photo of the study area and the photo was digitized so it could be recreated by computer. Twelve adult female deer were captured in Lory State Park and fitted with ear tags and radio collars between January 19 and Hay 4, 1982. Eight deer were monitored during 4 6-hour periods representing a 24-hour day during Har ch , 1981. Data are undergoing analysis at this time. Periodic checks showed 10 of the 12 instrumented deer remained on the study area from the time captured through July 13, 1982. In June, 2 deer left the study area and were located on July 13 in a mountain valley 29 km to the west of the study area. ��31 WINTER HABITAT SELECTION AND ACTIVITY PATTERNS OF MULE DEER IN FRONT RANGE SHRUBLAND AND FOREST HABITATS Roland C. Kufeld P. N. OBJECTIVE 1. To test Telonics telemetry equipment to determine its accuracy at various distances in locating a transmitter on the Horsetooth Mountain study area, and the ability of an observer using that equipment to correctly detect deer activity patterns by habitat type. 2. To determine habitat selection and activity patterns of mule deer within habitat types in the Horsetooth Mountain area during winter. SEGMENT OB·JECTIVES 1. Test Telonics telemetry equipment to determine its accuracy at various distances in locating a transmitter on the Horsetooth Mountain study area, and the ability of an observer using that equipment to detect deer activity patterns by habitat type. 2. Classify vegetation 3. Devise a method 4. Capture deer on the study area and instrument activity collars. 5. Monitor radio-collared activity patterns. types on the study area. for computerizing telemetry data. them with Telonics deer to determine habitat selection and ACKNOWLEDGEMENTS K. Risenhoover assisted in all phases of the study. I1ETHODS AND MATERIALS Determination of Telemetry Directional Accuracy, and Observer Ability to Determine Deer Activity Tests conducted during the previous segment (Kufeld 1981) were considered sufficient to determine directional accuracy of the telemetry equipment on the study area, and ability of an observer using that equipment to correctly detect deer activity by habitat type. Therefore, no additional tests were conducted during this segment. �32 Classification of Vegetation Types A color infra-red aerial photo of the Horsetooth Mountain area, taken June 25, 1980 from a U-2 aircraft at 70,000 feet was purchased from NASA and enlarged to 76 x 102 cm. The scale at that size is 71.2 mm/krn. An acetate overlay was taped to the map and boundaries of habitat types were denoted. Habitat types as small as 0.41 ha were marked. Habitat was classified according to the 10 following categories: Ponderosa Pine (+50%)-Douglas Fir (-50%) 10 to 39% canopy coverage 40 to 70% canopy coverage 71 to 100% canopy coverage 1 2 3 Grassland 4 1-fountain~feadow 5 Mountain 6 Mahogany Skunkbush-Wild Riparian 7 Plum-Chokecherry 8 (Cottom-mod-Willow) Rock Outcrops Agricultural 9 10 Fields Computerization of Habitat and Telemetry Data Habitat boundaries were digitized so that the entire map or any portion of it could be recreated by computer. Area within habitat types or portions of habitat types will also be retrievable. A program is also currently being written that will provide error polygons and area of each habitat type within the error polygon when coordinates and signal class designations are entered. Capture and Instrumentation of Deer Twelve adult female deer were captured in Lory State Park and fitted with ear tags and Telonics radio transmitter collars between January 19 and May 4, 1982. Each collar is equipped with a tip switch so it varies in pulse rate when the head is either up or down. Frequency of the pulse is used to determine deer activity. Seven of the 12 does were caught in Clover traps (Clover 1956) and 5 were immobilized with powdered Succynlcholine chloride administered from a dart gun. �33 Monitoring of Radio-Collared Deer A schedule was selected which includes the months of November through March and calls for deer to be monitored during 6-hour periods as follows: sunrise--3 hours before and after sunrise; daytime--3 hours before and after a point midway between sunrise and sunset; sunset--3 hours before and after sunset; nighttime--3 hours before and after a point midway between sunset and sunrise. Beginning November 1, 1982 efforts will be made to schedule 3 sunrise, 3 daytime, 3 sunset and 3 nighttime periods per month. During March, 1982, 8 instrumented deer were monitored for 1 sunrise, 1 daytime, 1 sunset and 1 nighttime period. This was accomplished through triangulation by 2 observers, each one monitoring from inside a camper truck equipped with a preclslon null antenna (Kufeld 1981). Triangulation points were about 1.6 km apart, and observers communicated via 2-way radios. Honitoring procedures were as follows: each observer oriented his antenna to a beacon on Horsetooth Mountain, and recorded his compass rose reading. Both observers then tuned to the radio frequency of the first deer to be sampled. If both observers received a class 1 or class 2 signal (Kufeld 1981), each recorded his signal bearing. If one or both observers received a class 3, 4, or 5 signal both observers selected the frequency of the next deer, and, again compared classes of signal. Upon determing location of a deer each observer oriented his antenna away from the center null and on to a peak, whereupon he locked his antenna in place. His strip chart recorder was then activated for 10 minutes to record activity of that deer. After 10 minutes observers attempted to locate the next deer. After all deer on the study area had been located and activity monitored, or they were passed over because of a poor signal the sequence was repeated beginning with the first deer. This was continued until the 6-hour period ended. RESULTS AND DISCUSSION Monitoring of Radio-Collared Deer Analysis of data on habitat selection collected during 4 6-hour monitoring periods in }1arch 1982, is awaiting completion of a computer program. Therefore, those data will be presented in a future report. Since primary emphasis this winter was on capturing and instrumenting deer. Monitoring time using the large, vehicle-mounted antennas to triangulate was limited. The study area, however, was checked periodically using handheld antennas from the time the first deer was instrumented until mid-July 1982, to determine if instrumented deer remained on the study area. Ten of the 12 remained within approximately 1.6 km of where captured. One deer remained until Hay 27, 1982, but a signal could not be received on June 9, 18, and 21. An aerial search on June 21, within a l6-km radius to the north, west and south of Horsetooth Mountain failed to produce a radio signal from either deer. On July 13, 1982, during another aerial search, both missing deer were located, together, in a mountain valley (2500 m elev.), 29 airline km west (274 ) of the site where instrumented. �34 They were located and observed from the ground the following day. Both appeared in good condition, but neither was accompanied by a fawn. Periodic telemetry checks indicated that instrumented deer remained in a relatively small area throughout the winter and spring, and exhibited regular patterns of movement. Movement was in an east-west direction. None of the deer were recorded moving north and south on the mountain. Generally, deer spent the day on the mountain, moved into the valley and hogback east of the mountain in the evening, and returned to the mountain within 2 hours after sunrise. In some cases, however, individuals would spend several days in the hogback before returning to the mountain. They did not appear to use the open valley between the mountain and hogback during mid-day. LITERATURE Clover, M. R. 1956. 42 (3) :199-201. Single-gate CITED deer trap. Calif. Fish and Game Kufeld, R. C. 1981. Winter habitat selection and activity patterns of mule deer in Front Range shrubland and forest habitats. Colo. Div. Wildl. Game Res. Rep. July (1):97-110. A/) Prepared by f!~/V(/O/ ;; .# t1/ C, Ku_i~c/Ct:~( Roland C. Kufeld Wildlife Researcher Ii C �35 JOB PROGRESS State of July, 1982 REPORT Colorado Project No. W-126-R-5 ~-=~-=-=---------- Work Plan No. 2 Big Game Investigations - Cervids Deer Investigations -----------_~------J ab-No.---- ------~9-----~-~--~- :---P iceance-Deer- S tudy----~----~----~~---------------- Period Covered: Personnel: July 1, 1981 through June 30, 1982 R. Bartmann ABSTRACT The mule deer census in Piceance Basin was flown March 2-4, 1982 with 952 deer counted for a mean density estimate of 7.93 + _2.29 deer/mile2 (90% confidence interval). Total population estimate was 21,094 + 6,091 deer which is considered lower than expected due to extremely poor counting conditions. A manuscript, Mule Deer Winter Mortality on Pinyon-Juniper Range in Northwest Colorado, was submitted to The Journal of Wildlife Management and is being reviewed. Two other manuscripts, Mule Deer Diet Composition and Quality on Pinyon-Juniper Winter Range in Colorado and Mule Deer Winter Foods in Piceance Basin, were submitted for internal review. - ��37 PICEANCE DEER STUDY Richard M. Bartmann P. N. OBJECTIVES 1. To complete transfer of responsibility for conduct of the quadrat deer census in the Piceance Basin from Research to Management. 2. To complete data analysis and publication of results completed under Work Plan 2, Jobs 3 and 4. for field studies SEGMENT OBJECTIVES 1. Assist the Northwest Region designee in conduct of the quadrat deer census in the Piceance Basin to complete transfer of this job from Research to Management. 2. Analyze data and report pertinent and food habits studies conducted results of the mule deer population under Work Plan 2, Jobs 3 and 4. METHODS AND MATERIALS Methods and materials have been previously described (Bartmann 1974). RESULTS The quadrat deer census in Piceance Basin was flown with Northwest Region biologists March 2-4, 1982. For the second consecutive year, snow conditions through out the winter were inadequate for optimum counting conditions. Consequently, all the deer were never forced down to the winter range. Due to the late date of the flights, deer distribution was considerably different than in previous years and counting conditions were poor due to no snow cover on south aspects and little snow on north slopes. Effects of deer distribution on the census are evident as there were 67 quadrats with no deer compared to the previous high of 53 quadrats with no deer, the highest number of deer counted on a quadrat was 93 compared to 87 previously, and confidence limits about the density estimate were the widest ever recorded. There were 952 deer counted for 7.93 + 2.29 deer/quadrat (90% confidence interval). This converts to 31.6 dee~/mile2 and a population estimate of 21,094 ± 6,091 deer. Completion of this work fulfills the Research Section"s involvement in the conduct of the census. A manuscript coauthored with Dr. D. C. Bowden, Mule Deer Winter Mortality on Pinyon-Juniper Range in Northwest Colorado, was submitted to The Journal of Wildlife Management and is currently being reviewed. Another manuscript, �38 Mule Deer Diet Composition and Quality on Pinyon-Juniper Winter Range in Colorado, intended for the Journal of Range Management, has been submitted for internal review. A Game Information Leaflet, Mule Deer Winter Foods in Piceance Basin, was also submitted for internal review. LITERATURE CITED Bartmann, R. M. 1974. Piceance deer study - population density and structure. Colo. Div. Wildl. Game Res. Rep. July. Part 2:363-370. Prepared b(21~===Richard M. Bartmann Wildlife Researcher C. �39 JOB PROGRESS State of July, 1982 REPORT Co] orado Project No. ~WL-_lu2~6L-~R~-_5~ _ Big Game Inyestigations Work Plan No. Deer Investigations ----~-Job--No 2 - Ceryids z:-=====l~O:!=======:-Mule-Deer-Winter-Habitat--Use-in-the---- Piceance Period Covered: Personnel: Basin July 1, 1981 through June 30, 1982 R. Bartmann, and J. Lee ABSTRACT Field work for the first segment of this study is finished and most data analyses completed. A Master's thesis by John Lee is expected by October 1982 and will be on file in the Colorado State University Library. ��41 MULE DEER WINTER HABITAT USE IN THE PICEANCE BASIN Richard M. Bartmann P. N. OBJECTIVE To administer funding and to supervise a graduate student study to assess mule deer winter habitat use in proximity development project using radio-telemetry methods. in conduct of a to an oil shale SEGMENT OBJECTIVES To administer funding and to supervise a graduate student study to assess mule deer winter habitat use in proximity development project using radio-telemetry methods. in conduct of a to an oil shale ACKNOWLEDGEMENTS The following people participated Alldredge and G. C. White. in various aspects of this study: A. W. RESULTS Field work is finished and most data analyses were completed at the Los Alamos National Laboratory in Los Alamos, New Mexico. The graduate student, John Lee, is preparing a Master's thesis which is expected to be complete by October 1982. Prepared bY{~nn Wildlife Researcher C ��43 JOB PROGRESS State of July, 1982 REPORT Colorado Project No. W-126-R-5 ------------------- Work Plan No. 3 Big Game Investigations - Cervids Elk Investigations ---Job-N"o--.------2'------ ---:--Elk-Popu-ration-and--Ecologi-ca-1-Studies--- ------------------------ Period Covered: Personnel: July 1, 1981 through June 30, 1982 G. Bear, and R. Green ABSTRACT All data were gathered and analyses are being made for the 4-year population study in the Rocky Mountain National Park area. Manuscripts will be prepared next segment. Ron Green has completed a thesis on "Elk habitat sel.ecz Lon and activity patterns in Rocky Mountain National Park" and it will be on file in the Colorado State University Library. ��45 ELK POPULATION AND ECOLOGICAL STUDIES George D. Bear and Ronald A. Green P. N. OBJECTIVES 1. Develop techniques population levels. to more accurately 2. Compare means corresponding preclslon levels, and costs of 2 independent systems for estimating numbers of elk in Rocky Mountain National Park. 3. Define natality National Park. and mortality and precisely characteristics estimate elk of elk in Rocky Mountain SEGMENT OBJECTIVES 1. Trap and mark up to 200 elk on the winter 2. Compare means, precision levels, and costs of 2 independent estimating elk numbers in Rocky Mountain National Park. 3. Monitor causes. telemetered elk calves to determine METHODS Methods and materials have been previously described ~~.~ George D. Bear Wildlife Researcher C for rates and probable (Bear, 1979). CITED Bear, G. D. 1979. Elk population and ecology Fed. Aid Prog. Rep. July. Part 2:373-399. by mortality systems AND MATERIALS LITERATURE Prepared range. studies. Colo. Div. Wildl. ��47 JOB PROGRESS State of __~C~o~l~o~r~a~d~o~ July, REPORT _ Big Game Investigations Pro j ec t No. _:_W;_--=1=-=2;..::6;_--=.R::...._ _ Work Plan No. -----job-No. Period 1982 3 - Cervids Elk Investigations --------3-- ~-:--EvaluaEion~of-Factors-Inf-luencing-Elk----------~-Nutritional Status and Population Perfonnance ------------------------ Covered: Personnel: July 1, 1981 through June 30, 1982 D. Baker ABSTRACT Preliminary studies of deer-elk digestive physiology were examined during a 2l-day feeding trial and 7-day complete balance trial. Digestive capabilities of elk were greater than mule deer for dry matter, energy, and structural carbohydrates. In vitro dry matter digestibilities were similar to in vivo values for both deer and elk. Ad libitum intake (g/day/kgo.75) of a grass hay diet ranged from 27 to 45 for deer and 56 to 70 for elk. Comp~rative experiments of deer and elk on native diets were also completed this segment. Chemical evaluation of these diets is currently in progress. ��49 EVALUATION OF FACTORS INFLUENCING ELK NUTRITIONAL STATUS AND POPULATION PERFORMANCE Dan L. Baker P. N. OBJECTIVES 1. To develop and test a system for evaluating to support elk. the potential of habitats 2. To improve the predictive capability of this system by identifying and quantifying variables influencing the range supply-animal demand model of nutritional carrying capacity. SEGMENT OBJECTIVES 1. Begin initial investigations of comparative forage intake of deer and elk. digestive physiology and a. Train and condition experimental animals to confinement in isolation pens, metabolism cages and weighing apparatus. b. Conduct preliminary experiments to (1) define problems associated with experimental equipment and procedures, (2) determine ad libitum intake levels of deer and elk consuming forage under controlled conditions, (3) evaluate both in vivo and in vitro rare earth elements as potential markers of particulate flow through the gastrointestinal tract. 2. Estimate comparative digestive efficiency, mean retention time, rumen fill and fermentation rates of deer and elk fed an array of native forages. 3. Define elk. 4. Develop intake. in vivo/in vitro relationships and/or improve ungulate of native nutrition models forages for deer and for predicting forage ACKNOWLEDGEMENTS P. Neil, J. Ritchie, and L. Stevens assisted in various aspects of the study. �50 Table 1. Mean daily dry matter intake, nitrogen intake and energy intake of deer and elk feed chopped grass hay. Species and no. Elk 90 93 47 98 97 Mean SE CV(%) Deer 150 189 180 165 125 187 Mean SE CV(%) Avg. BW, kg Avg. dry matter intake g/day/ g/day/ kgO.75 kgO.9 g/day Digestible crude protein intake g/day/kgO.75 Digestible energy intake kcal/day/kgO.75 308 312 277 190 194 4,645 4,140 4,727 4,850 3,743 63.3 55.7 69.6 94.7 72.0 26.8 23.5 29.9 43.1 32.7 8.44 5.65 7.01 a a 165.9 130.6 159.1 a a 256 27 23.5 4,421 208.7 10.5 71.1 6.56 20.6 31.2 3.4 24.0 7.03 0.8 12.3 151.8 10.8 12.3 63 58 64 40.5 60.0 67.0 666 941 610 380 480 851 29.7 44.7 27.1 23.6 22.2 36.3 16.0 24.4 14.5 13.6 12.1 19.3 3.8 4.9 3.3 a a a 76.0 87.0 62.4 a a a 58.8 3.9 16.1 655 87.6 32.7 30.6 3.5 27.9 16.7 1.9 27.0 4.0 0.5 20.5 75.1 7.1 16.4 aThese measurements were not made on these animals. In vitro digestible dry matter (IVDDH) coefficients were similar to in vivo values for both elk and deer. Elk IVDDH values were consistently lower than in vivo values. Apparent digestibilities for all constituents were more variable for deer than elk. �Sl METHODS AND MATERIALS Experimental methods and design pertinent to objectives la, 1b(1), and 1b(2) are described in a previous Federal Aid report (Baker and Hobbs 1981). Methods for objective 1b(3) are described in a detailed study plan (Colorado Division of Wildlife files). In vitro evaluation of rare earth elements as particulate markers is currently being conducted at the Animal Science Department, Cornell University, Ithaca, New York", and therefore will not be addressed in this study. Results of these experiments will be summarized in the final report. Detailed methodology and experimental design relative to objective 2 is presented in the study plan. Methods and analysis for evaluating in vivo/in vitro relationships of native forages (objective 3) have also been described in the detailed study proposal. This experiment is currently in progress and results will be presented in the 1983-84 Federal Aid report. Completion of objective 4 is contingent upon results from experiments under objectives 2 and 3. Since these studies were not completed during this segment, nutritional modeling will be addressed in future reports. RESULTS AND DISCUSSION Deer-Elk Comparative Intakes and Digestion: Preliminary Stu.dies Intake Chemical composition of the chopped grass hay and description of the experimental animals used in this study have been previolls1y reported (Baker and Hobbs 1981). Average dry matter intake (g/day/kgO;75) for mule deer ranged from 27.1 to 44.7 and SS.7 to 69.6 for elk (Table 1). Mean voluntary intake for deer (30.6) and elk (71.1) in this study was similar to intakes previously reported for white-,tailed deer (33.1) and elk (67.9) consuming grass hay of similar chemical composition (Mould and Robbins 1982). Dry matter, digestible energy and digestible crude protein intake were greater for elk than deer (P < O.OS) (Table 1). Digestible energy intake (kca1 DE/day/kgO•75) ranged-from 130.6 to 16S.9 for elk and 62.4 to 76 for mule deer; digestible crude protein (g/day/kgO•75) S.7 to 8.4 for elk and 3.3 to 4.9 for mule deer. Mean intake of digestible energy (DE) was SO% greater for elk compared to deer while digestible crude protein intake was 46% greater for elk. Digestibility Elk were more efficient in digesting nutrients in grass hay than mule deer (Table 2). Apparent digestibi1ities for dry matter, digestible energy, neutral detergent fiber and acid detergent fiber were higher for elk than mule deer (f < O.OS). Organic matter digestibility was similar for both species (P > O.OS). While deer were only slightly less efficient in digesting-dry matter than elk (61.9% vs S7.0%), large differences were observed in the digestion of the fiber fraction of grass hay. Elk digested neutral detergent fiber 10% greater than deer and acid detergent fiber 22% greater. �52 Table 2. Apparent digestibilities grass hay. Species and no. Elk 90 93 47 Mean SE CV(%) Deer 150 189 180 Mean SE CV(%) (%) for mule deer and elk fed chopped Dry matter Organic matter Energy 61.8 61.3 62.7 58.8 59.6 62.3 59.3 59.6 60.0 55.8 56.6 60.2 56.3 53.1 57.4 59.1 59.1 61.8 61.9c 0.4 1.1 60.2c 1.1 3.0 59.6c 0.2 0.6 57.5c 1.4 4.1 55.6c 1.3 4.0 60.0 0.9 2.6 55.0 61.6 54.6 53.0 61.0 59.8 52.0 58.9 52.6 47.3 50.0 44.2 43.6 50.0 43.5 61. 7 56.5 55.7 d 57.0 2.3 6.9 c 57.9 2.5 7.5 d 54.5 2.2 7.0 d 47.2 1.7 6.2 d 45.7 2.2 8.2 58.0 1.9 5.62 In vitro dry matter ~DF = neutral detergent fiber. ADF = acid detergent fiber. c,dMeans With different superscripts are significantly different at 0.05 level. Deer-Elk Comparative Digestion of Native Forages Measurements of voluntary forage intake digestive efficiency and rate of passage of native diets was completed during this segment. Complete analysis of these data is awaiting completion of nutritional analysis. LITERATURE CITED Baker, D. L., and N. T. Hobbs. 1981.· Elk investigations--evaluation of factors influencing elk nutritional status and population performance. Colo. Div. Wildl. Game Res. Rep. July, Part 1:145-154. Mould, E. D., and C. T. Robbins. 1982. Digestive capabilities in elk compared to \-lhite-taileddeer. J. Wild1. Manage. 46:22-29. I£~J__;__' Prepared by--!-.:-l__:;____:·· .L i!---"---L-' Dan L. Baker Wildlife Researcher C �53 July, 1982 JOB FINAL REPORT State of Colorado Project No. W-144-R -------------------- Work Plan No. 1 Big Game Investigations Multispecies Investigations ------Job-No-=-.---------1,..---------------:-An4ma-l-and-P-en-SuppoFt-F-ae-i-l-i·t-i-es-Eo-r'------------Big Game Research Period Covered: Personnel: July I, 1981 through June 30, 1982 P. H. Neil ABSTRACT Modifications to the facility and minor construction projects were completed to facilitate several research programs. Intensive training of mule deer and elk for use in digestion cages continued during the early portion of the segment. A study of the comparative digestive physiology of mule deer and elk was undertaken during fall, winter, and spring. Details and objectives of this project are described under W-126-R, WP3, J3, Dan Baker, Principal Investigator. An additional study at the facility investigated the palatability of aspen bark pellets to captive big game species in an effort to determine the feasibility of using these pellets as a feed component or filler. Twenty mule deer fawns, 3 bighorn sheep lambs, and 2 Rocky Mountain goat kids are pres·ently being hand-reared and trained for recruitment into the captive research herd. Including these animals the total captive research herd consists of 34 mule deer, 7 elk, 9 Rocky Mountain goats, 8 bighorn sheep, and 6 pronghorn. ��55 ANIMAL AND PEN SUPPORT FACILITIES FOR BIG GAME RESEARCH Paul H. Neil P. N. OBJECTIVES To provide facilities and maintain populations of captive big game animals and pen to support big game research programs. DESCRIPTION Foothills Wildlife Collins, Colorado. Research OF AREA Support Facility METHODS located northwest of Fort AND MATERIALS Personnel from the CSU Physical Plant were enlisted to construct several modifications to the isolation pens and digestion cages in preparation for intake and digestion trials. The Young-Adult Youth Conservation Corps of the Bureau of Reclamation was enlisted to assist with reconstruction of several sheds and buildings that were severly wind-damaged during the early spring of 1982. They also assisted with construction of a habitat mountain to accommodate Rocky Mountain goats. Preparations were made for fawn rearing at the facility to accommodate up to 20 mule deer fawns. Six female pronghorn were used in isolation pens for a feed comparison intake trial through a cooperative study with Colorado State University. Four mule deer were used in an intake and digestion trial to evaluate the palatability and digestibility of aspen bark pellets in an effort to determine the feasibility of the use of aspen bark pellets as a feed component or filler. Procedures of this study are described in the January-March 1982 Quarterly Report W-144-R, WPl, Jl. Six mule deer and 6 elk were used in isolation pens and digestion cages on a study of comparative digestive physiology of mule deer and elk. Details of this study are described under W-126-R, WP3, J3, Dan Baker, Principal Investigator. RESULTS AND DISCUSSION Reconstruction of all wind-damaged buildings and sheds was completed with the exception of 5 of the 15 isolation pens. All other modifications and construction projects were also completed in support of the various research projects. Preliminary results from the study of comparative digestive physiology are described under W-126-R, WP3, J3. Results of the aspen bark pellet study �56 are pending lab results Quarterly Report. and will be reported in the July through September One bull elk died of apparent "chronic wasting" during the segment. Four mule deer died during the segment. Death resulted from "blue tongue", pneumonia, and "chronic wasting", respectively. One buck pronghorn died of apparent "capture myopathy." Twenty mule deer fawns are presently being hand-reared at the foothills facility for recruitment into the r.esearch herd. Three bighorn sheep and 2 Rocky Mountain goats are also being hand-reared for upcoming research projects. The total big game research herd presently consists of 34 mule deer, 7 elk, 9 Rocky Mountain goats, 8 bighorn sheep and 6 pronghorn. Other activities at the facility Tour of facilities. Tour of facilities. Tour of facilities. Animal Immobilization Manager Trainees. Tour of facilities. Tour of facilities. Prepared by during September September September Training the segment included the following: 1981. Colo. Joint Budget Committee. 1981. Bureau of Reclamation Personnel. Interested public personnel. 1981. Class. March 1982. District Wildlife Colorado State University April 1982. June 1982. Day Camp Science students. \?~-'1~ Paul H. Neil Wildlife Technician III students. �57 July, 1982 JOB PROGRESS REPORT State of Colorado ~--~--~~----------- Project No. W-144-R ------------------- \.J"ork Plan No. ----Job-Nol-::.-----l 2 Personnel: - Noncervids Bighorn Sheep Investigations ------------------------- Period Covered: Big Game Investigations :--Prescribed-Burning-of~Bighorn-Sheep-----------: Mule Deer Winter Ranges July 1, 1981 through May 30, 1982 N. T. Hobbs and R. A. Spowart ABSTRACT Two years following treatment, prescribed burning continued to affect ecological processes in mountain shrub and grassland communities. Mule deer and bighorn sheep ate more rapidly and consumed more graminoids and less shrubs on burned plots compared to controls. Burning increased in vitro digestible organic matter (IVDOt·O content of highorn sheep diets during November in both communities and increased IVDOM levels in deer diets during November in mountain shrub and during January in grassland. Diet overlap between mule deer and bighorn sheep was greater on burned plots than controls in mountain shrub; burning had no effect on diet overlap in grassland. Burning increased the density of IVDOM in the standing crop of herbage in the mountain shrub community. Burned areas had lower concentrations of soil NH4+ than controls; there was no difference in soil N02 + N03' Burning increased the potential for nitrogen fixation in the mountain shrub community. ��59 PRESCRIBED BURNING TO IMPROVE AND ENLARGE BIGHORN SHEEP RANGES N. T. Hobbs and R. A. Spowart P. N. OBJECTIVES 1. Quantify the effects of burning mountain shrub and grassland communities on nutritional status of mule deer and bighorn sheep during winter. 2. Determine the change in nutritional carrying capacity of mountain shrub and grassland winter range for mule deer and bighorn sheep which is brought about by burning. 3. Examine the effect of fire on food niche relations separation of mule deer and bighorn sheep. 4. Explain changes in responses of forage resources, both quantity quality, in terms of processes in the nitrogen cycle. and ecological and SEGMENT OBJECTIVES 1. Estimate nutrient distributions 2. Describe botanical composition and nutritional quality of bighorn and mule deer diets in relation to prescribed burning. 3. Analyze 4. Estimate unburned forage samples soil nitrogen study plots. in burned and unburned for protein fixation METHODS plant communities. sheep content and digestibility. and mineralization rates on burned and AND MATERIALS For a complete description of methods used in experiments during 1981-82 see Hobbs, N. T. and R. A. Spowart (1981); no substantial departures were made from methods described there. RESULTS AND DISCUSSION Prescribed Burning Effects on Diet Composition Prescribed burning exerted small effects on the forage class composition of diets of bighorn sheep in mountain shrub communities 2 years following treatment (Table 1). Bighorn sheep chose diets from burned mountain shrub plots which were similar to diets chosen from control plots during all months except March when bighorn consumed more grass and less forbs on burns compared with controls. This difference can be attributed to increases in the amount of green grass on burned plots during March; �60 during previous months little green forage was available on either burn or control plots. Throughout winter, bighorn diets in the mountain shrub community were dominated by herbaceous plants; they ate little browse. In contrast to the relatively small effects of burning on diet botanical composition, eating rates of bighorn sheep in mountain shrub were substantially higher on burns compared with controls during all winter months (Table 1). It appeared that the dense canopy of shrubs present on control plots interfered with the ability of bighorn sheep to find herbaceous forage. Table l. Dietary botanical composition and biting rates of bighorn in burned and unburned mountain shrub communities during 1981-82. Month Percent grass Burn Control x SE x SE Percent forbs Burn Control x SE x SE Percent shrubs Burn Control x SE x SE sheep Biting rate (bites/min) Burn Control x SE x SE Nov 57 11 54 11 42 12 41 11 4 2 1 0 18 3 10 3 Jan 64 10 59 10 32 11 32 9 4 1 9 3 17 1 11 1 Mar 68 10 40 7 31 9 52 8 1 0 8 2 18 2 8 2 May 83 2 71 3 17 1 26 4 0 0 3 1 37 3 34 7 Bighorn sheep feeding in burned grassland communities chose diets containing more grass and less forbs than diets selected from grassland control plots during November (Table 2). This pattern was reversed during January. There were no differences in the forage class composition of bighorn sheep diets from burned and control grassland plots during March and May. Bite rates of bighorn sheep were higher on burned grassland plots than on controls during November and March; during other months, burning had no effect on biting rates. Table 2. Dietary botanical composition and biting rates of bighorn burned and unburned grassland communities during 1981-82. Month Percent grass Burn Control x SE x SE Percent forbs Control Burn x SE x SE Percent Burn x SE shrubs Control x SE sheep in Biting rate {bites/min} Burn Control x SE x SE Nov 50 12 32 8 49 11 62 7 1 1 6 5 21 2 13 3 Jan 58 11 74 5 39 12 23 4 3 1 3 1 20 3 16 1 Mar 54 10 53 4 45 10 40 5 1 0 7 4 20 3 9 1 May 74 2 81 5 25 2 19 5 1 0 0 0 43 3 41 3 �61 Two years after treatment, prescribed burning continued to affect the composition of mule deer diets chosen from mountain shrub communities (Table 3). Diets of mule deer contained more herbaceous forage, particularly grass, and less browse on burned plots compared with controls. This difference did not appear to be caused by decreased availability of shrubs in burned areas; sprouts of a variety of browses, which were readily eaten on the controls, were also available on the burns. Instead, mule deer foraging on burned plots selected herbs in preference to browse. Biting rates of mule deer were higher on burned mountain shrub plots than on controls during all months except January. Table 3. Dietary botanical burned and control mountain Month Percent grass Burn Control x SE x SE composition and biting rates of mule deer in shrub communities during 1981-82. Percent forbs Burn Control x SE x SE Percent shrubs Burn Control x SE x SE Biting rate (bites/min) Burn Control x SE x SE Nov 31 8 11 4 42 15 43 16 27 6 46 20 10 2 7 1 Jan 27 10 15 9 40 5 41 8 33 10 44 3 7 2 8 1 Mar 74 9 28 15 20 8 41 10 6 1 30 5 13 1 4 1 May 77 7 56 9 17 9 31 2 7 2 13 6 23 2 10 3 Prescribed burning exerted similar effects on composition of mule deer diets in grassland communities (Table 4). Diets chosen by mule deer on burned plots tended to be dominated by herbs; diets from controls contained substantially more leaves and stems of shrubs. However, in contrast to the effect seen in mountain shrub, biting rates of mule deer in grassland appeared to be unaffected by treatment during all months except March. Table 4. Dietary botanical composition and biting rates of mule deer in burned and control grassland communities during 1981-82. Month Percent grass Burn Control x SE x SE Percent forbs Burn Control x SE x SE Percent shrubs Burn Control x SE x SE Biting rate (bites/min) Control Burn x SE x SE Nov 19 9 7 1 63 8 47 7 18 3 45 8 10 1 9 2 Jan 23 1 37 14 56 9 23 2 21 9 40 15 12 2 10 2 Mar 52 2 37 2 37 3 26 3 11 2 37 2 12 2 7 2 May 59 2 69 7 36 2 25 8 5 2 6 2 20 2 17 1 �62 Although effects of prescribed burning on diets of mule deer and bighorn sheep persisted 2 years after treatment, the magnitude of these effects appeared to diminish. During the first year, deer and sheep ate more graminoids and less dicots on burned plots compared with controls in both habitats during all months except May (Hobbs and Spowart 1981). In comparison, the effects of burning on diet composition during year 2 were less consistent, both temporally and spatially. Further, biting rates during the first year following treatment were almost always 2 to 5 times greater on burns than controls. Differences during year 2, although significant, were much smaller in size. Prescribed Burning Effects on Diet Nutritional Quality A rigorous evaluation of effects of prescribed burning on quality of diets of mule deer and bighorn sheep must await completion of laboratory analysis of forage samples. However, some preliminary results on diet quality 2 years following treatment indicate that Durning continues to enhance ungulate nutrition. Diets of bighorn sheep selected from burned plots contained more IVDOM than diets chosen from controls during November and January in the mountain shrub community, and during November in grassland (Table 5). Treatment effects on IVDOM content of mule deer diets were observed during November in mountain shrub and during January in grassland. Table 5. In vitro digestibility (% of organic matter) of diets of mule deer and bighorn sheep feeding in burned and unburned mountain shrub and grassland communities during early winter 1981-82. Species Month Mule deer Bighorn sheep Grassland Burn Control SE SE x x Mountain Burn x SE shrub Control x SE Nov 33 1 33 1 42 2 26 1 Jan 43 1 35 5 42 4 39 3 Nov 44 1 37 2 42 1 29 2 Jan 46 5 44 3 47 1 42 1 Although differences in diet digestibility attributable to treatment were smaller 2 years after burning than 1 year following burning (Hobbs and Spowart 1981), these effects remained biologically and statistically meaningful. Biologically, even small changes in the digestibility of diets within the range of 40-50% digestible dry matter can profoundly influence animal condition (Blaxter et al. 1961). Consequently, the relatively small effects of treatment on dietary IVDOM content we observed in grassland may offer substantially greater benefits for animal condition than the size of �63 these differences might suggest. The much larger effects we observed in mountain shrub during November would, without doubt, enhance the nutritional status of mule deer and bighorn sheep. Rigorous inferences on the statistical significance of treatment effects require a complete analysis of variance; however, the small error associated with our estimates of means (Table 5) suggests that the differences we observed are repeatable. Prescribed Burning Effects on Ecological Separation Physical structure and floristic composition of plant communities influence the extent of ecological separation among sympatric wild herbivores (Lamprey 1963, Jarman 1974, Hirst 1975, Hanley and Hanley 1982). Consequently, treatments like prescribed burning which alter these community properties may affect the niche relations of ungulates who share the same ranges. Two years after treatment, burning substantially increased dietary overlap between mule deer and bighorn sheep during all months in the mountain shrub community and during March in grassland (Table 6). This increase resulted primarily from the shift in mule deer diets to include more grass and less browse on btiIrnedplots compared to controls. Increased similarity of diets mayor may not portend increased competition between mule deer and bighorn sheep. If sympatric populations of deer and sheep are small relative to the total food resources available in burned habitats and adjacent, untreated areas, then the effects of burning on dietary overlap will probably offer negligible effects on the performance of those populations. However, because burning can substantially reduce the biomass of forage on offer (Hobbs and Spowart 1981) and because enhanced nutrition resulting from burning will likely increase ungulate reproductive rates, the potential for burning to intensify competition between mule deer and bighorn sheep is very real. This potential problem requires thorough study in future, more detailed analyses and should be carefully weighed against the potential benefits of burning in formulating fire prescriptions whereever these ungulates cohabit the same ranges. a of bighorn sheep and mule deer on burned Table 6. Percent dietary overlap and control grassland and mountain shrub communities during winter and spring 1981-82. Grassland Control Mountain Burn shrub Control Month Burn Nov 59 60 n 63 Jan 65 63 63 56 Mar 90 70 89 77 May 85 88 94 85 aKulczynski's coefficient, forage class percentages. the sum of the lowest members of each pair of �64 Prescribed Burning Effects on Forage Nutrient Density Prescribed burning continued to increase the density of IVDOM in the mountain shrub community at the end of the growing season 2 years following treatment (Table 7). Biomass of forages containing low concentrations «40%) of IVDOM was substantially greater on control plots compared with burns. The amount of forages of higher quality (>40% IVDOM) on burned plots exceeded the biomass of those forages on controls. In particular, there was 4 times more forage containing 60-70% IVDOM on burns compared to controls. As a result of these biomass relationships, the density of IVDOM in the standing crop of forage was much greater on burned plots than on controls. Unburned areas were dominated by low quality forage; only 21% of the standing crop on control plots contained greater than 40% IVDOM. In contrast, 78% of the total biomass on burns exceeded 40% digestibility. Table 7. Biomass and percentage of the standing crop of forage categorized by content of IVDOM in burned and unburned mountain shrub plots, 2 years after treatment. (g/m2) Control SE x Percent of standing croE Burn Control x SE x SE IVDOM category (%) Biomass Burn x SE > 0-10 1 1 38 14 1 1 9 5 >10-20 2 1 315 141 2 1 56 11 >20-30 6 2 35 2 5 2 8 2 >30-40 18 6 25 6 13 3 6 2 >40-50 23 3 44 9 18 3 9 2 >50-60 61 17 49 6 45 12 11 3 >60-70 21 4 5 1 15 4 1 0 These differences offer important implications for wild ruminants. A deer or bighorn sheep searching an unburned mountain shrub community for nutritious forage must find that meal in the midst of a great mass of undesirable, low quality plant tissue. By comparison, that search is easier in burned mountain shrubs where nutritious forage is much more concentrated. We believe this difference had a large influence on the increased biting rates we observed on burned mountain shrub plots (Tables 1,3); our animals were able to spend less time searching, and more time eating. �65 Prescribed Burning Effects on Nitrogen Cycling We observed no effect of burning on soil mineral N levels 2 years following treatment (Table 8). However, burning appeared to stimulate N fixation in the mountain shrub community (Table 9), an effect opposite of the inhibition of N fixation caused by burning the previous year. Both of these trends can be explained by changes in herbage biomass. We suspect that as plants invaded burned areas and grew, more mineral nitrogen was taken up from the soil than when burned areas had fewer, smaller plants. The rapid growth of these herbaceous species was associated with greater rates of nitrogen immobilization than the slower growth of woody plants in unburned areas. Moreover, as legume biomass increased, the potential for nitrogen fixation on burned plots became greater. Table 8. Concentration (~g/g) of ammonium (NH4+) and nitrate + nitrite (N02 + N03) in top 5 cm of soil in burned and unburned mountain shrub and grassland communities. Mountain Burn x SE Nitrogen form shrub Control x SE Grassland Burn x SE x Control SE NH4+ 4.3 0.5 6.0 0.5 5.2 1.1 7.6 1.3 N02 + N03 2.4 0.3 1.9 0.3 2.0 0.6 1.9 0.2 Table 9. Rate of nitrogen fixation (p moles/g soil dm/hr) based on acetylene reduction in top 5 cm of soil from burned and unburned mountain shrub and grassland communities with soil moisture at field capacity. Plant community Mcuntain Grassland Control Burn x shrub SE x SE 14.0 4.7 10.1 1.2 13.4 3.8 13.3 3.8 �66 Prescribed Burning Effects: Conclusions Two years following treatment, prescribed burning of mountain shrub and grassland communities continued to offer nutritional benefits to mule deer and bighorn sheep. Thorough description of these benefits awaits completion of chemical analyses of forage samples and statistical tests on differences in the responses we described in this report. However, at this point in our research, prescribed burning appears to be an effective means of improving the winter habitat of mule deer and bighorn sheep in Colorado. LITERATURE CITED Blaxter, K. L, F. W. Wainman, and R. S. Wilson. food intake by sheep. Anim. Prod. 3:51-61. 1961. The regulation of Hanley, T. A., and K. A. Hanley. 1982. Food resource partitioning by sympatric ungulates on great basin rangelands. J. Range Manage. 35:152-158. Hirst, S. M. 1975. Ungulate-habitat relationships woodland/sauana. Wildl. Mono. 44. in a South African Hobbs, N. T., and R. A. Spowart. 1981. Big game investigations: prescribed burning to improve and enlarge bighorn sheep ranges. Colo. Div. Wildl. Wildl. Res. Rep. July Part 2:155-181. Jarman, P. J. 1974. The social organization Behavior 48:215-267. their ecology. of antelope in relation to Lamprey, H. F. 1963. Ecological separation of large mammal species in the Tarangire Game Reserve, Tanganyika. E. Afr. Wildl. J. 11:329-38. Prepared by Wildlife Researcher C �July, 1982 67 JOB PROGRESS State of Colorado Project No. ~W_-~1~4~4_-~R~ Work Plan No. _ 3 --~--------------1 Job No. Period Covered: Personnel: REPORT Big Game Investigations Pronghorn - Noncervids Investigations =~:=:==~====::::===-=-=-=-==:=-==-------Pronghorn Population Dynamics Study July 1, 1981 through June 30, 1982 T. M. Pojar A manuscript titled "Quadrat and Strip Sampling to Estimate Pronghorn Population Characteristics" was prepared for submission to the Journal of Wildlife Management reporting on the results of this study. The following is the abstract of the manuscript. ABSTRACT Quadrat and strip sampling designs were used in an aerial helicopter survey of a 72 by 60-km area in east central Colorado to estimate both density and herd structure of a pronghorn antelope (Antilocapra americana) population during the same fly-over in late summer. In a preliminary test it was determined that observed density did not differ between strips 0.8-km and 1.6-km wide, therefore 1.6-~ strips were used. The area to be surveyed was divided into sections 2.59-km by fencZs and roads; because of this and statistical considerations we used 2.59-km quadrats. In 12 hours of flight time we searched 35% of the total with the strip sampling design and it took 16 hours to search 6% of the area with the quadrat design. The population size, buck to doe ratio, and fawn to doe ratio (and 90% confidence intervals) resulting from the strip census was: 3,570 + 18.2%, 46.8:100 + 19.3%, and 63.9:100 ± 10.0%, respectively. The same parameters estimated-from the quadrat census were 6,366 ± 25.0%, 66.9:100 ± 48.2%, and 66.9:100 ± 17.4%, respectively. On the- strip and quadrat censuses 1,203 and 381 pronghorn were classified, respectively. Neither the buck to doe nor the fawn to doe ratios were significantly different between sampling methods (P > 0.10). The population size estimate however, was significantly greater (P < 0.10) from the quadrat sampling method. Potential biases of each sampling method were examined and evaluated, at least intuitively, in seeking an explanation for the large difference in the two estimates. ��69 PRONGHORN POPULATION DYNAMICS STUDY Thomas M. Pojar P. N. OBJECTIVES 1. Test the efficiency and preC1Slon on pronghorn antelope range. of a quadrat and strip census design 2. Test the efficiency and precision of herd structure concurrently with a quadrat and strip census. 3. Test the reliability of doing strip and quadrat summer when it is possible to do herd structure 4. Test the predictability 5. Measure and model greater. estimates when done census during late counts. of the ONEPOP and the POP50 population the effects of reducing a pronghorn simulators. density by 50% or SEGMENT OBJECTIVES 1. Conduct pilot study to test effective rate. 2. Test the efficiency and preC1Slon of quadrat and strip census design on pronghorn antelope range in estimating density and herd structure during the same fly-over in late summer. 3. Establish initial simulation strip width and observability to provide predicted decay values. ACKNOWLEDGEMENTS I would like to express appreciation to Bill Knight, Mark Elkins, Jack Vayhinger, Dr. David C. Bowden,and Bruce Gill for their considerable help in the planning, execution, and reporting of this study. METHODS AND MATERIALS Methods and materials are described in detail in the manuscript titled: 'Quadrat and Strip Sampling to Estimate Pronghorn Population Characteristics." �70 RESULTS AND DISCUSSION The results and discussion of this study have been summarized in manuscript prepared for submission to the Journal of Wildlife Management titled: "Quadrat and Strip Sampling to Estimate Pronghorn Population Characteristics." Wildlife Pojar Researcher C �July, 71 JOB PROGRESS State of Project Colorado No. W-144-R --~~~-----------3 Covered: Personnel: Big Game Investigations Pronghorn --"J~-'o~b~N"-'o"_'_. -=2 Period REPORT ----------------------- Work Plan No. ____ 1982 Investigations _.,_-~Estimation of PronghQ_l:"!!~ Genetj,_~__~ Variability July 1, 1981 through June 30, 1982. T. M. Pojar ABSTRACT Tissue samples (blood, heart, liver, kidney, and muscle) were collected from hunter killed pronghorn during the 1981 season from the Hugo and Saguache study areas. The samples were processed for electrophoresis and frozen. The equipment and supplies for conducting electrophoresis were assembled and the process of electrophoresis was practiced. No electrophoresis runs were made during this segment. ��73 ESTIMATION OF PRONGHORN GENETIC VARIABILITY Thomas M. Pojar P. N. OBJECTIVES 1. To estimate the genetic variability of two pronghorn populations exhibit distinctly different reproductive performance. that 2. To collect genetic variability information on other pronghorn populations as the opportunity of obtaining the necessary biological materials (blood and/or tissue samples) arises from statewide trapping operations. SEGMENT OBJECTIVES 1. To estimate genetic variability of two pronghorn distinctly different reproductive rates. populations that exhibit 2. To collect genetic variability information on other pronghorn populations as the opportunity of obtaining the necessary bioloigcal material (blood and/or tissue) arises from statewide trapping operations. ACKNOWLEDGEMENTS I am indebted and grateful to Leslie Johnson, Dr. Donald Nash, and Mark Elkins for their assistance with this research. METHODS Methods study. and materials AND MATERIALS have been described in the Program Narrative for this RESULTS AND DISCUSSION Additional tissue samples were collected from the Hugo and Saguache study areas during the 1981 hunting season from hunter-killed pronghorn. With this collection it will now be possible to examine all of the 21 isozymes listed in Pojar (1981) for all 4 of the areas sampled (Hugo, Saguache, Craig, and LaJunta). The samples were processed according to Manlove et al. (1975) and frozen. Several practice electrophoresis runs were made to test equipment and gain the skills necessary to successfully execute the electrophoresis process. �74 LITERATURE CITED Manlove, H. N., J. C. Avise, H. O. Hillestad, P. R. Ramsey, H. H. Smith, D. O. Straney. 1975. Starch gel electrophoresis for the study of population genetics in white-tailed deer. Proc. 29th Annu. Conf. SE Game and Fish Commissioners 29:392-403. Pojar, T. M. 1981. Pronghorn investigations: Pronghorn population study. Job Progress Rep. Colo. Div. Wi1d1. Wi1d1. Research Rep. July, Part II: 183-206. Prepared ~ homas M. Poj r Wildlife Researcher G;J;w C �July, 1982 75 JOB PROGRESS State of Project REPORT Colorado --~~~~~----------No. W-144-R --~~~------------ Big Game Investigations Work Plan No. 3 Pronghorn Investigations Job No. 3 Pronghorn Breeding Period Covered: Personnel: Experiment July 1, 1981 through June 30, 1982 T. M. Pojar A manuscript titled "Recurrent Estrus in Pronghorn Antelope" was prepared for submission to the Journal of Wildlife Management. The following is the abstract of the manuscript. ABSTRACT Recurrence of estrus in pronghorn antelope (Anti1ocapra americana) was evaluated with a vasectomized male and 2 females, all 2 years old, in 1980 and with this same group plus 4 yearling females in 1981. All animals were hand-reared and maintained in captivity during the study. The male was fitted with a leather breeding harness that held a bar of colored wax-base marking material between his front legs so he would leave a mark on the doe as he mounted. Date of estrus was estimated by markings and pronghorn behavior. Three and 4 estrous periods were detected for the 2 mature females each year. The 4 yearlings had 1, 2, 2, and 2 estrous periods. Th~ mean (SE, n) cycle length of the mature females was 27.6 (1.21, 5) and 30.4 (2.54, 5) days, respectively in 1980 and 1981. Mean date of first estrus was earlier (P < 0.05) for mature females (30 Sep) than yearlings (22 Oct). ��77 PRONGHORN BREEDING EXPERIMENT Thomas M. Pojar P. N. OBJECTIVES 1. Determine if female pronghorn one breeding season. have more than one estrus period during 2. Compare 3. Compare the number of estrus periods between yearling breeding) and mature females. the date of first estrus of yearling vs. mature females. (age at first SEGMENT OBJECTIVES 1. Compare the date of first estrus of yearling vs. mature 2. Compare the number of estrus periods between yearling breeding) and mature females. females. (age at first ACKNOWLEDGEMENTS I appreciate the assistance of Lisa Lani Wolfe and Paul Neil. Without help this experiment would have been considerably less successful and enjoyable. METHODS their AND MATERIALS The methods and materials employed in this experiment are described in detail in a manuscript by T. M. Pojar and L. L. Wolfe titled "Recurrent Estrus in Pronghorn Antelope" to be published in the Journal of Wildlife Management. RESULTS AND DISCUSSION The results and discussion pertaining above mentioned manuscript. prepared~~ Thomas M. Pojar Wildlife Researcher to this study are covered in the 61J~ C ��79 JOB PROGRESS State of July, 1982 REPORT Colorado Project No. ~W~-~1~4~4~-~R~-~1~ _ Big Game Investigations Work Plan No. Rocky Mountain ... ---~---Job-No 4 - Noncervids Goat Investigations •. -=====:::!It:::=======-:~RockyMount-a-in-Goat---Bighorn-Sheep---Competition Study Period Covered: Personnel: July 1, 1981 through June 30, 1982 D. F. Reed ABSTRACT A detailed study plan (Program Narrative) titled "Seasonal Habitat Selection and Activity of Sympatric Mountain Goat and Bighorn Sheep Populations" was developed, reviewed, and approved. Twenty-five mountain goats were marked with radio telemetry collars, neck bands, or eartags. One bighorn sheep was marked with a radio telemetry collar. Of the 25 habitats used in the study, mountain goats and bighorn sheep occupied 15 and 10 of the habitats during September through May, respectively. The number of mountain goat groups or individuals occupying the 15 habitats was greater (P < 0.005) than the same for bighorn sheep groups or individuals. ��81 ROCKY MOUNTAIN GOAT - BIGHORN SHEEP COMPETITION STUDY Dale F. Reed P. N. OBJECTIVES Evaluate the extent to which mountain goat populations limit seasonal habitat utilization of bighorn sheep in alpine environments and to describe patterns and rates of dispersal of mountain goats from colonization sites. SEGMENT OBJECTIVES 1. Develop a detailed study plan for investigating seasonal habitat sympatric populations of mountain goats and bighorn sheep. 2. Capture and radio collar up to 10 mountain 3. Monitor habitat 4. Map and describe use of radio collared seasonal mountain goats and bighorn mountain sheep. goats and bighorn goat and bighorn use of sheep. sheep habitats. ACKNOWLEDGEMENTS I thank D. McGlasson of the Mount Evans Research Station, University of Denver, for providing a place for a sno-cat and other field equipment. H. E. Wiseman and J. Stone provided field assistance often under arduous conditions. C. E. Braun provided aerial photography and vegetation data that included the study area. DESCRIPTION OF AREA The study area is located in the Colorado Front Range about 50 km westsouthwest of Denver. It includes all of the alpine in the Goliath Peak, Lincoln Lake, and Rogers Peak areas located east of Mount Evans. The elevation ranges from about 3,500 m near the Bristle Cone area to 4,082 m on Rogers Peak summit. The center of the study area is located at about 448 4386 UTM or 390 37.5' N, 1050 36.5' W geographical coordinates. About 8 km of State Highway 5 (from mile post 3 to mile post 8) intersects the study area. The area is composed of gently rolling slopes to steep slopes, rocky ridges, cliffs, and rock slides. Other abiotic factors have been discussed previously (Reed 1980:377). Vegetation in the area has been addressed by Streeter (1969) and Braun (1969). METHODS AND MATERIALS Methods and materials have been described by Reed (1981). Some changes have been made in the study plan (Program Narrative, Appendix A). The route through the study area has been further delineated to systematically cover �82 selected vegetation units (18 to 24) mapped by Braun (1969). RESULTS MiD DISCUSSION Study Plan A detailed study plan (Program Narrative) titled "Seasonal Habitat Selection and Activity of Sympatric Mountain Goat and Bighorn Sheep Populations" was developed, reviewed, and approved (Appendix A). Capture and Marking Twenty-five mountain goats were marked with radio telemetry collars, neck bands, or eartags (Table 1). One bighorn sheep was marked with a radio telemetry collar. This bighorn sheep was a yearling male collared with Red tele (Ch 3) at Rl (Table 1). Capture success of bighorn sheep was ex~ ceedingly low due primarily to their reluctance to enter traps and to habituate to regular apple pulp baiting at dropnet sites. Three of the 25 marked mountain goats were outfitted with radio telemetry collars. These 3 plus the 9 radio collars that are still active (transmitting) from the last segment, provide 12 active radio collars on mountain goats (Table 2). Of these 12 radio collars, 5 are activity (tip switch) collars. The collar outfitted on the bighorn sheep is also an activity collar. Since Blue diag wide, Red tele mountain goat, and Red tele bighorn sheep (Table 2) were captured and collared near the end of this segment, little to no data were collected on them for this reporting period. Description of Habitats The 18 vegetation units used in this study are listed and described in Appendix A. Of these 18 units, 3 were divided into subunits designated by "a" or "b" and 2 of the subunits were further differentiated according to proximate locations within the study area (i.e. E = east, W = west, etc.). The 3 units, subunits, and differentaited subunits are as follows: Unit M-l Subunit Differentiated Subunit �Table 1. Date, age, sex, collar or tag, and selected measuremen ts of mountain goats trapped in the Mt. Evans area. Trap a Location Girth (em) Length (em) Horn length (em) Left Right Wt, (kg) Horn to muzzle (em) Front leg fm scapula Hind foot Date Age Sex 1 28 Aug 81 3 F Black collar 5 TC 96 146 19.3 19.4 20.5 76 30.3 2 30 Aug 81 4 F Black collar 6 R2 101 145 23.0 23.4 22.8 82 29.5 111 159 21. 4 22.1 74 23.5 81 30.5 No Collar/tag (ern) 3 4 Sep 81 5 F Black collar 7 Rl 4 Sep 81 2 M Green eartag R2 92 138 17.8 17.1 45 19.5 73 5 15 Sep 81 6 F Black collar 8 R2 ]05 147 21.9 21. 7 58 23.0 76 30.0 6 6 Oct 81 8 F Black collar S2 133 140 21. 8 21. 2 79 23.8 78 30.0 7 14 Oct 81 K M Black eartag S2 97 114 5.3 5.3 33 15.2 61 26.8 8 20 Oct 81 6 F Green diag (Ch 5) S1 118 153 23.4 22.7 75 24.0 84 32.0 9 20 Oct 81 K H Blue eartag Sl 87 107 4.2 4.1 30 16.0 64 26.5 10 20 Jun 82 Y F Grey eartag DAC 11 20 Jun 82 Y F Black 1 dangle DAC 12 20 Jun 82 Y F Black 2 dangle DAC 13 21 Jun 82 Y M Black 3 dangle DAC 1 14 24 Jun 82 2 F Blue diag wide (Ch 4) BPP 15 25 Jun 82 y M White ear tag 1 DAC 16 25 Jun 82 y White eartag 2 DAC 17 25 Jun 82 y M White eartag 3 DAC 18 25 Jun 82 y F White ear tag 5 DAC 19 25 Jun 82 6 F Red tele (Ch 7) R1 20 26 Jun 82 y F Black 4 dangle DAC 21 26 Jun 82 5 J! White 4 eartag DAC 22 26 Jun 82 y F White 7 eartag DAC 23 26 Jun 82 5 M White 6 ear tag DAC 24 27 Jun 82 4 F White 9 eartag R2 25 27 Jun 82 4 F White 8 eartag Rl a TC = Tumbling Creek at first switchback, Sl and S2 Rl and R2 = = Saddle below curve and mile post 6 (1 and 2, east and west trap, respectively), Rogers east above Lincoln Lake drainage and road between mile post 6 and 7 (1 and 2, north and south trap, respectively, DAC = Data acquisition center site at mile post 7, BPP = Below photo point on road shoulder between Rl-R-2 and mile post 7. co w �84 Table 2. Telemetry collar, channel, frequency, pulse, and activation date for radios outfitted on mountain goats and on a bighorn sheep in the Mt. Evans area. Frequency Telemetry Mountain collar Channel b c d Activation date 1 2 3 2 4 5 6 4 7 148.1300 148.3000 148.3100 172.2375 172.2625 172.2875 172.2625 172.3125 172.3625 172.3875 172.3125 172.4125 78 76 72 11 48 82 65-90c 65-90 65-90 65-90 60 d 90-65 22 18 22 30 29 7 9 16 20 29 24 25 3 172.2875 68-98 17 Jun 82 Aug Sep Sep Sep Sep Oct Jun Jun Oct Jun Jun Jun 80 80 80 80 80 80 81 81 81 81 82 82 sheep Red tele a Pulse per min. (ppm) goats a White tele Blue tele Yellow tele b Yellow diag Green 2 Green 3 Red-White-Blue tele Blue diag Green diag Black tele Blue diag wide Red tele Bighorn (MHz) Tele denotes telemetry Diag denotes diagonal Activity (tip switch) collars; Activity (tip switch) collar; 65 ppm 90 ppm = = head up, 90 ppm = head up, 65 ppm = head down. head down. �85 Unit Subunit Differentiated Subunit M-2a(W) M-2a(E) M-2a(S) M-2 M-2b(E) M-2b M-2b(W) M-8a M-8 < M-8b (N) M-8b M-8b(S) The units and subunits are briefly described in the study plan (Appendix A). Differentiated subunits have the same vegetation types as the subunits from which they were derived. However, they have different locations and often different topographic features. Locations and additional features are as follows: M-2a(W) - Northeast portion of ridge north Rogers Peak, escape terrain north end of ridge. M-2a(E) - South portion of long slope and ridge north and northeast Lincoln Lake, escape terrain in cliff area and 6 distinctive rocky ridges, unbaited lick in west portion. M-2a(S) - Slopes above and below road west Lincoln Lake, escape terrain in nearby subunits, baited saltlick at Mile Post 7 and trapping location. M-2b(E) - Ridge below Mile Post 7 and southwest terrain along top of ridge. Lincoln Lake, escape �86 M-2b(W) - Slopes and rocky point southeast Rogers Peak to northsouth line intersecting Rogers Peak-Mt. Warren saddle, escape terrain on rocky point southeast Rogers Peak. M-8b(N) - Cliff area northwest-north of Lincoln Lake, escape terrain of various quality throughout, includes "cave" occassionally used by mountain goats. M-8b(S) - Steep cliff and large boulder area west to northwest Lake, escape terrain throughout. Lincoln For ease of reference the vegetation units, subunits, and differentiated subunits will be referred to hereafter as "habitats." The total number of habitats considered in the study is 25. Also, for b-revity the "M" which is an abbreviation for Mount Evans (Braun 1969) will be dropped from the habitat designation. Habitat Use Of the 25 habitats used in this study, mountain goats and bighorn sheep occupied 15 and 10 of the habitats during September through May, respectively. Conversely, 10 of the habitats (3, 4, 5, 6, 7, 10, 11, 12, 23, and 24; Appendix A) were not occupied during this period by either species, and 15 were not occupied by bighorn sheep. The number of habitats occupied by both species declined during winter (Fig. 1). The number of habitats occupied by mountain goats was greater than the number of habitats occupied by bighorn sheep for every month except November (Fig. 2). The number of mountain goat groups or individuals occupying the 15 habitats was greater (P < 0.005) than the number of bighorn sheep groups or individuals occupying the 15 habitats (Fig 3). Differences are particularly apparent in habitats lb, 2a(E), and 2a(S) (Fig. 3). These habitats are located above Lincoln Lake and are used frequently by mountain goats throughout the fall, winter, and spring seasons. Habitat lb includes two mineral sites, one where salt was baited and the other where a natural lick occurred. Similar features of 2a(E) and 2a(S), discussed under the previous sub-heading, may account for a relatively high amount of mountain goat use. In theory, the same resources were available to bighorn sheep. These habitats may not have been used frequently by bighorn sheep because; 1) bighorn sheep have different requirements, 2) bighorn sheep are substantially fewer in number, and/or 3) bighorn sheep are excluded by the presence of mountain goats. Although the number of groups or individuals observed was greater for mountain goats, the mean size of groups for the 2 species was not different (P > 0.20; Fig. 4). Selected Animals Of the 12 mountain goats outfitted with radio telemetry collars, Black tele and her cohorts were located and observed more frequently throughout the fall, winter, and spring than any other telemetered animal. Of 42 locations (Table 3), 40 were in alpine habitats and 2 were well into (1,040 m from and 207 m �87 15 o I / I I I o 10 ",, I I ,, I , '0 CIl ~ Po :;:I I I , I , tJ o I , 0 til I " .u cd .u ~ "o/ I .0 Cd ::c 4-1 0 . 0 z 5 Fall Winter Spring Figure 1. Number of habitats occupied by mountain goats (dashed line) and bighorn sheep (solid line) by season (fall ~ Sep-Nov, winter = DecFeb, spring = Mar-May) 1981-82. �88 15 10 0, I I I /\ Z / -, -, / \ / 5 '0 / o . o , I / \\ " '0 '\ I / / \ o / \ \ / / o Sep Oct Nov Dec Jan Feb Mar Apr May Figure 2. Number of habitats occupied by mountain goats (dashed line) and bighorn sheep (solid line) by month 1981-82. �39 I, 0 60 I\ I "\ , I I 50 f I til \ f I \ III ::l oM :> /\ oM ,\ ~ H 30 0 til P::l I I 0 !-o C.!J ~ 0 . 0 , 0 "'d !-o ,, ,, ,I 40 "'d \ \ , ,....; \ \ \ \ I \ I I I , I I I , ,, , , , I 20 ,r 0 \ \ I 0 z I \ 10 0 1\ \ II 0 1b \ I \ \ I 1a I \ I \ I 0 1\ \ I \ ,I 0'\. 0 2a 2a 2a 2b 2b (W) (E) (S) (E) (W) 8a " 8b 8b (N) (S) 9 19 20 21 Habitats Figure 3. Number of mountain goat groups or individuals (dashed line) and bighorn sheep groups or individuals (solid line) observed occupying 15 habitats September through May 1981-82. 22 �90 25 20 o 1\ <I) N J J \ 15 I \ ...-l I { I \ I \ tJ) c, ::l 0 1>1 • C!> 1:>< 10 \ 5 0/ 0_-0/ I ~o ~ \ ", \ I \ I \ I 0 -, \. lb 2a 2a 2a 2b 2b 8a (W) (E) (S) (E) (W) 0 / / \ / 0--0 I '\0 . 0 • • la ,, , 8b 8b (N) (S) 9 • 19 20 • 21 Figure 4. Mean group size of mountain goats (dashed line) and bighorn sheep (solid line) occupying 15 habitats during September through May 1981-82. 22 �91 Table 3. Date, location, and habitat occupied by Black telemetry (Black tele) mountain goat in the Mt. Evans area September through May 1981-82. Location Date 4 8 11 15 18 18 22 22 29 2 6 7 12 17 17 24 25 2 8 16 18 31 20 28 29 18 21 25 2 3 11 13 16 25 1 9 9 10 14 14 15 15 a Sep Sep Sep Sep Sep Sep Sep Sep Sep Oct Oct Oct Nov Nov Nov Nov Nov Dec Dec Dec Dec Dec Jan Jan Jan Feb Feb Feb Mar Mar Mar Mar Mar Mar Apr Apr Apr Apr Apr Apr Apr Apr General 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 82 82 82 82 82 82 82 82 82 82 82 82 82 82 82 82 82 82 82 82 N Lincoln SW Rogers Burn Ridge 1 & 2 New Lick New Lick S Rogers Point SE Rogers Ridge 5 Burn Ridges 1 & 2 NW Lincoln Below Ridge 1 & 2 N Lincoln Slope 1 N Lincoln NW Lincoln Between Ridge 2 & 3 N Rogers Beyond Ridge 4 Ridge 4 Ridge 1 Below rd slope 1 & 2 Beyond slope 2 Below slope 2 East Rogers E Below slope 2 N Rogers NW Lincoln NW Lincoln N Rogers N Rogers N Rogers N Rogers N Rogers Below 1 & 2 Slope 2 Below slope 1 & 2 Near Ridge 2 N Lincoln N Lincoln Below saddle traps a UTM 4850 4690 5014 4910 4820 4808 4720 4775 4930 5018 4902 4800 4896 4845 4849 4840 4800 4908 4695 4933 4933 4892 4836 4815 4835 4800 4825 4694 4795 4794 4691 4700 4695 4673 4692 4843 4836 4828 4909 4902 4850 4875 8562 8482 8423 8555 8567 8540 8450 8495 8555 8425 8560 8554 8547 8556 8569 8561 8577 8547 8655 8545 8555 8552 8566 8580 8565 8582 8563 8530 8523 8525 8643 8660 8657 8580 8663 8563 8578 8559 8557 8550 8553 8563 Habitat Occupied 2a(E) 2b(W) 2a(E) Ib 2a(S) 19 2a(S) 2a(E) 2a(E) 2a(E) 2a(E) 2a(E) 2a(E) 2a(E) Ib 2a(E) la 2a(E) 2a(E) 2a(E) 2a(E) Ib 2a(E) Ib 2a(E) la 2a(S) 2a(S) la la la la la 2a(E) lb 8b(N) 2a(E) 2a(E) 2a(E) Ib First and second UTM coordinates are preceded by 4 and 43, respectively (i.e. 44800 438500). �92 below nearest alpine) a subalpine burn where the "understory" vegetation was luxuriant. She was located in habitat 2a(E) 48 percent of the time (20 locations). Periodically she occupied 6 other habitats (la, lb, 2a(S), 2b(W) , 8b(N) , 19) within the study area. During the last segment Blue tele was located 53 times. During this segment she was located on only 31 occasions (Table 4). Based on these locations she occupied alpine habitats throughout the fall, winter, and spring. She ~as located in habitat 2a(E) 55 percent of the time (17 locations). Periodlcally she occupied 4 other habitats [la, lb, 2b(E), and 8b(N)]. Table 4. Date, location, and habitat occupied by Blue telemetry (Blue tele) mountain goat in the Mt. Evans area September through May 1981-82. Location Date 10 15 13 14 3 24 8 9 16 18 31 8 18 21 25 3 13 16 25 1 9 9 10 14 14 14 15 15 15 25 7 a Sep Sep Oct Oct Nov Nov Dec Dec Dec Dec Dec Jan Feb Feb Feb Mar Mar Mar Mar Apr Apr Apr Apr Apr Apr Apr Apr Apr Apr Apr May UTM General 81 81 81 81 81 81 81 81 81, 81 81 82 82 82 82 82 82 82 82 82 82 82 82 82 82 82 82 82 82 82 82 Ridge 1 NW Lincoln NE Lincoln NE Lincoln N Lincoln Slope 1 Rogers E NW Lincoln Beyond Ridge 4 Ridge 4 Slope 1 N Lincoln Below slope 2 Below slope 2 N Rogers Below Mile Post N Rogers N Rogers N Rogers N Rogers Below slope 1 & E Rogers E Below slope 1 & E Rogers E Near slope 2 NW Lincoln N Lincoln Slope 2 Above slope 1 Below mile post N Lincoln a 7 2 2 7 4900 4795 4885 4885 4872 4848 4780 4820 4933 4933 4858 4842 4800 4825 4672 4812 4700 4695 4673 4692 4843 4818 4828 4815 4820 4830 4830 4830 4845 4838 4825 8560 8555 8535 8535 8545 8569 8580 8565 8545 8555 8568 8563 8582 8563 8534 8468 8660 8657 8580 8663 8563 8589 8559 8590 8586 8568 8555 8575 8590 8464 8555 Habitat Occupied 2a(E) 2a(E) 2a(E) 2a(E) 2a(E) 2a(E) lb 8b(N) 2a(E) 2a(E) 2a(E) 2a(E) lb 2a(E) la 2b(E) la la la la 2a(E) lb 8b(N) lb lb 2a(E) 2a(E) 2a(E) 2a(E) 2b (E) 8b(N) First and second UTM coordinates are preceded by 4 and 43, respectively (Le. 44800 438500). �93 LITERATURE CITED Braun, C. E. 1969. Population dynamics, habitat and movements of whitetailed ptarmigan in Colorado. Ph.D. Diss. Colorado State Univ., Fort Collins. l89pp. Reed, D. F. 1980. Rocky Mountain goat investigations: Rocky Mountain goat ecology study. Colo. Div. Wildl. Game Res. Rep. July, Part 2:353-381. 1981. Rocky Mountain goat ecology study. Game Res. Rep. July, Part 2:209-222. Colo. Div. of Wildl. Streeter, R. G. 1969. Demography of two Rocky Mountain bighorn sheep populations in Colorado. Ph. D. Diss. Colorado State Univ., Fort Collins. 96pp. Prepared 8dlIeS?CJ·· Dale F. Reed Wildlife Researcher C ��95 APPENDIX PROGRfu~ NARRATIVE State: Colorado Project Title: Big Game Investigations Work Plan Title: Hountain Prepared by Selection and Activity Project No. W-144-R-l Work Plan No. Goat Investigations Job Title: Seasonal Habitat Sympatric Hountain (Non-Cervids) of Job No. Goat and Bighorn Sheep Populations :C~6---rc{~a:::i;-{~ec~·· Dale Reed' , J ..._). '.~. /~ C...,.'"""';\~" -i:.'~(/"'--~(__-:_ .. a.:»: Submitted b y : rIo V ~ '? vL· ; =..:: Date: Dale Reed Rev.i.ewe d by: Tom Beck Dan Baker Date: II A. ~t'e/)lOfl(_ Ron Kufeld Date: Approved by: Date: Date: 4- /-?Z /9P/ �96 A. NEED Problem Mountain goats (Oreamnos americanus) They were first introduced additional releases sheep ungulates into the State in 1948 (Denney 1977). This area was within (Ovis canadensis) population. that at some threshold is plausible mountain density The problem that this has already goat and bighorn competitive occurred, sheep populations selection "competition" for mountain goats. It and that the sympatric are already responding to is a logical first step in. examining The degree to which mountain habitats is that as these interactions. This study of habitat problem. advantage a 1961 release grow, it is speculated of one or both species, in a competitive Several the range of an indigenous occupy this range and their populations may occur resulting to Colorado. have been made since then, including in the Mount Evans area. bighorn are not indigenous and how those habitats goats and bighorn this sheep select are used are the subjects addressed by this study. If the findings habitat of this study indicate use or selection be addressed, removal can be predicted reduction" goats are displacing or reduction measurable response of the mountain of the bighorn strategies in which a "removal will be conducted bighorn goat and bighorn and if management future studies will be planned or an "experimental mountain that mountain goat population population. can experiment" to test the hypothesis sheep from choice habitats. sheep Any could result in a that �97 Literature Review There is a substantial habitat selection volume of literature or preference and methods few studies have used experimental procedures types or animal numbers were manipulated Merkt 1981). Most studies have described concerning to evaluate them. of However, where either the habitat (Wecker 1964, Rosenzweig habitat (Telfer 1970, Smith 1977) or have been concerned preferences concepts 1973, use qualitatively with diets and forage (Hjeljord 1973, Peek et al. 1976, Seegmiller and Ohmart 1981). Being able to relate animal locations of abiotic and biotic features of location measures telemetry, of habitat an individual not distributed quantity) preferences. evenly, to their occurrence) is optimal opportunistic Ungulate (Harrington (habitat resources patches differ in 1978) and may select from a mosaic of resources. a X2 test of the hypothesis or avoidance to habitat availability of a given resource of the individual in terms of its that animals use habitats in has been used (Neu et al. 1974). that in comparing to the availability each portion used disproportionately species are, to a greater or lesser degree, availability attributes It has been suggested if the environmental preference et al. (1979) indicated that of both quality and selectively. To determine proportion of using quantitative (where food and cover are i.e. spatial heterogeneity strategy the use Schoen et al. (1979) hypothesized should use that environment (Wiens 1976). often through the importance a patchy environment that a coarse-grained quality to the patchiness of the environment, has increased inhabiting or movements the proportion of those attributes, Schoen of use of various it must be assumed home range is equally available to the that �98 individual animal, so that selection to choices by the individual. there is no difference to the frequency null hypothesis selectivity. criminant of occurrence To determine a rank-order procedure These analytical of that attribute. compared It is noted that if the the type and degree of this Neu et al. (1974) used a method Hudson nominal for preference procedures, based on (1976, 1977) used both multiple scale analyses. are due to be tested is that of use of each attribute X2 does not determine z statistic. and multiple of the attributes Thus the null hypothesis in the frequency is rejected, the Bonferroni or rejection Johnson dis- (1980) developed analysis. as well as others that have been used, are as follows: Affinity index (AI) Cairns and Telfer used with x2 (z) Neu et al. (1974) Bonferroni z statistic Chi-square (X ) Kearney Duncan's 2 multiple Coefficient Electivity (1980) and Gilbert (1976) range test (DMRT) Rounds of association coefficient Multiple classification Multiple discriminant Multiple nominal Multiple regression (C ) Dice 7 (1945), Cole (1949), Singer (1979) (EC) Schoen et al. (1979) analysis analysis scale analysis and partial Rank order (RO) Johnson Schoener's index (MCA) Hudson (MDA) Hudson (MNA) Hudson correlation (1977) (1976), Matthews (MRPC) Hudson (1977) et al. 1981 correlation (1979) (1977) (1980) (SI) Linton Simple and multiple (1981) (SMC) Shannon et al. (1975) �99 Hudson (1977) briefly He indicates discusses that MNA " ... provides data where the dependent dent variables interval, between the application variable are measured flexibility of independent In addition, Hudson variable of habitat variable to interpret to the usual interpretation, evidence either for or against conclusions that the results selection Contrary and resource resource the existence overlap of his study processes. can be used as (Sale 1974). can only be drawn when it is demonstrated and habitat preferences are altered by monitoring or fortuitous most directly changes following follows this study on habitat to habitat use or selection patterns are exhibited. goat activity patterns exhibited activities of salt licking, to habitat selection. Hence, a second selection. is how a given habitat what activity Rideout is used or (1974) reported on mountain during summer and fall periods. feeding, moving, of another (Hudson 1977). removal of one of the species. study logically that in the presence This could be demonstrated Related and among sets partitioning, of competition to productivity generation exist in terms of competitive species and this change is consequential the planned indepen- (nominal, ordinal, of relationships and the dependent (1976, 1977) concludes and that they are difficult distribution categories, at any level of measurement patterns ecological variables." are simply descriptive Definitive for handling is a set of discrete ratio), and where variable any independent of MRPC, MCA, MDA, and MNA. The and resting were not related �100 B. OBJECTIVES The objectives of this study can be stated as hypotheses Hypothesis Mountain 1: goats use alpine habitats to be tested: disproportionately to their availability. Hypothesis 2: Bighorn sheep use alpine habitats disproportionately to their availability. Hypothesis 3: Mountain goats and bighorn sheep use the .. same alpine 4: Mountain goats and bighorn sheep exhibit habitats. Hypothesis C. patterns in alpine habitats. EXPECTED RESULTS a case study of mountain in the Mount Evans area. Any inference strategies D. OR BENEFITS This study is essentially best hazardous. the same activity Specifically of mountain beyond this research goats and bighorn goats and bighorn sheep the Mount Evans area is at should help identify management sheep in the Mount Evans area. APPROACH Experiment a. HI = 1: Mountain Mountain Goat Habitat Selection goats use alpine habitats disproportionately to their availability. b. H o = Mountain goats do not use alpine habitats disproportionately to their availability. c. Methods: Mountain goats will be trapped and marked Evans area at previously other mineral used saltlicks lick locations in the Mount (Baumann undated) that may not be documented. and at �101 Although these locations will not represent an unbiased sample of the experimental population in the study area (Goliath Peak - Lincoln Appendix A) mountain (Reed, unpubl. a random trap distribution, is expected since Lake - Rogers Peak. goats move freely from 1 lick area to another data). Hence, it is estimated has equal susceptibility that each subgroup of being trapped. One to several Clover traps (Clover 1956) will be placed over each lick area. paration, They will be baited with salt, a grain/pellet apple pomace, or alfalfa depending When set, the traps will be checked more frequently in the morning when animals are in the immediate and tagging can be accomplished unpubl. on seasonal promptly. data) indicates mountain or area so that banding Preliminary evidence goats respond vigorously (Reed, to salt traps. Conse- goats will be trapped and banded during these periods. Trapped mountain and placing response. and evening during spring and summer, and are not wary of entering quently, mountain feed pre- goats will be handled a blindfold by tying their horns and legs over their eyes to calm them. A short piece of garden hose will be placed over the horns for safety. Kids will be ear tagged (numbered and/or color coded) or collared with small expandable collars. Yearling vidually numbered and 2-year old females will be collared with indiand/or color coded neckbands. old) will be collared with numbered outfitted with telemetry collars males will only be ear tagged their potential or 1300 West University Drive, in the 148 and 172 MHz ranges. Adult (numbered and/or color coded) because trophy status. collars will be maintained (>2 years and/or color coded neckbands (Telonics, Mesa, AZ 85201) using frequencies Adult females Approximately of 10 active telemetry on adult females during the study. Overall, �102 about 30 collared during or eartagged the study. Marking mountain records goats will be maintained and individual each trapped animal will be maintained characteristics (Appendix B). Radio-telemetry will be used as an aid in obtaining When relatively strong telemetry animals are within versely, within because accurately direction animals it difficult from within finding. telemetered of topographic signals are the result of "signal ridges, making rebound" to triangulate the study area. Additionally, are often visible is possible making visual sightings. sightings or placing will avoid the reported animals sampling features. These weak from nearby peaks and the location of animals of 2-3 km (Reed, unpubl. when triangulation of locating animals are not in the area of this study (alpine), Consequently, method Con- Strong signals are needed for for distances Visual telemetered and can usually be sighted. when weak signals are received, line-of-sight visual sightings. signals are received, line-of-sight of so is the likelihood data). of will be the principal in given habitats. This method errors of radio-triangulation (Springer 1979, Bear and Green 1980:240). The study area will be sampled for mountain a prescribed route and methodically the aid of binoculars, portion of the route spotting searching route for mountain scope, and radio-telemetry. (Mount Evans road from NE Goliath Lake) will be traversed with a vehicle depending goat occupancy on snow conditions) goats with The first to NW Lincoln (pickup, sno-cat, or on foot. by covering or other The second portion of the (road NW of Lincoln Lake to NW end of ridge ~ 1.3 km NE Rogers Peak, then generally along the contour Peak, then back to road NW of Lincoln to the west to ~ 0.7 km N Rogers Lake, then west on the road to �103 SE of Rogers Peak) will be traversed wide expanses The sampling November), Summer for viewing the various periods will be within winter on foot. (1 December-28 The route will provide areas of the study. 3 seasons: February), fall (1 September-30 and spring (1 June-3l August) will not be included because: period is judged to be the least critical in terms of habitat availability, energy expenditures, selection forage nutritional by human recreation. units will be used to sample within be used to sample within weeks, used to sample within days 1200, 1200-1600, seasons, and 1600-1800). conditions. can be made. 3) animals activity are located sample 2-day sample units will 0600-0800, 0800- The point time sample hours since it is unlikely However, nocturnal with Experiment are met: 1) animals cular and nocturnal goat habitat The 2-day sample units will be pancy will be sampled in conjunction when 3 conditions levels, and One-week as follows: units will not be taken during nighttime that visual sightings goats and point time sample units will be (stratified chosen to avoid severe weather May). 1) the summer period for mountain and 2) during this period mountain may be influenced (1 March-3l habitat 4 (see page 13) are located in evening, indicates no movement in early morning. occu- 2) crepus- (Experiment In reference 4), and to the use of point sample units, it is noted that time interval units could be used instead. However, move extensively units represent that mountain in short periods, given intervals The alpine habitats vegetation considering it is likely that most point sample of time spent in given habitats. that will be considered units mapped by Braun types including mineral goats do not normally in this study are 18 (1969) (Appendix C) and 4 non-vegetation lick, roadbed, rock (Shepherd 1975:71), and �104 snowfield. In addition, square quadrats coordinates maps. the study area will be divided based on UTM coordinates are determined Both the habitats and analyzing mountain by overlaying distributional data. coordinates (Appendix E). will be determined extrapolating between The presence color (estimated to the nearest 100 m grid lines, as determined vegetation be overlayed on aerial photographs habitat (Braun, pers. files), type will be determined snowfield from photographs Analysis: The percent occupying each habitat of variance previously and transitory of telemetered (18 vegetation, for each location. locations will mapped with vegetation of each vegetation (hectarage snow cover has blown and marked mountain goats 4 non-vegetation) and each for percent availability will be analyzed across months and across seasons. goats, the frequency by X2. from black taken during late winter after and each quadrat after being adjusted be analyzed a made from the ground. quadrat after being adjusted marked mountain 10 m by by placing types, UTM coordinate The availability areas have developed by analysis Mercator) maps and orienting from these aerial photographs off, and from inspections d. goats [NASA 1:114,323 to 1:116,248], and color-infrared To determine planimetered), and for each quadrat. UTM (Universal Transverse [NASA 1:109,009 to 1:115,950] aerial photographs) types or absence of type of all observed mountain grid on 1:24,000 or 1:62,500 topographic and white, UTM grids on topographic for each habitat date, time, and habitat will be recorded These and the quadrats will be used in recording goats will be recorded Location, (Appendix D). into 500 m For un- of sightings within each habitat for percent availability will �105 Schedule Fiscal Activity Prepare and complete radio collar mountain habitat habitat goats, trap and and monitor 1981-82 use patterns. Trap and radio collar mountain goats, and monitor use patterns. 1982-83 Same as 1982-83 1983-84 Same as 1983-84 1984-85 Data analysis second Experiment a. study plan, Years H2 = and publication. generation 2: Bighorn Bighorn Prepare and complete 1985-86 study plan if appropriate. Sheep Habitat sheep use alpine Selection habitats disproportionately to their availability. b. H o Bighorn sheep do not use alpine habitats disproportionately to their availability. c. Methods: Methods will be the same as those as they apply to bighorn sheep respond to salt with generally very wary vigorously to apple they will be trapped dropnet or with trap response. those of mountain them. sheep. less vigor of entering pomace There and banded than mountain traps. the winter during as is necessary Since bighorn sheep horns Approximately no protection 5 active telemetry goats However, and early these periods other methods goats, 1 except are a few exceptions. Clover during for Experiment and are they respond spring. with depending are not nearly will be necessary collars Bighorn Hence, either a on bait and as sharp as to handle will be maintained �106 on adult females during the study. d. eartagged bighorn Analysis: Analysis Overall, sheep will be maintained about 15 collared or during the study. will be the same as that for Experiment 1. Schedule Fiscal Years Activity Prepare and complete collar bighorn habitat study plan, trap and radio sheep as possible, and monitor 1981-82 use patterns Trap and radio collar bighorn and monitor habitat sheep as possible, 1982-83 use patterns. Same as 1982-83 1983-84 Same as 1983-84 1984-85 Data analysis and publication. second'generation Experiment 3: Prepare and complete study plan if appropriate. ~fountain Goat and Bighorn Sheep Habitat Overlap a. H3 Mountain goats and bighorn sheep use the same alpine habitats. b. H o Mountain goats and bighorn sheep do not use the same alpine habitats. c. Methods: Methods will be the same as those for Experiments except for the following. attributes In addition of slope, aspect, distance and terrain will be calculated for each mountain attributes to the habitat goat location (Appendix F). road northwest of Lincoln at a weather Lake. elevation, on USGS topographic of snow cover and depth, temperature (Appendix F) will be measured types, habitat to escape terrain, or measured 1 and 2 Additionally, maps temporal (OC), and wind speed station located above the Snow cover will be estimated ocu1ar1y �107 for each location. from a photo-point General located west of Lincoln Lake. read from a stake calibrated blown and compacted, estimated in centimeters. depths where with methods Temperature snow cover will be recorded will be recorded station instruments If a given weather estimated d. Analysis: 1. When snows are not wind the animals are located will be on a 7-day hygrothermograph. from an accumulating will be maintained parameter estimates station and photo-point of variance and seasons. anemometer. The 4 times each month. will be derived from the data. will be across species as well as months For unmarked distribution of sightings for independence animals within of both species, each habitat 2 (X , 2 x C contingency the dependent (Appendix F). variable, and 16 stratified Multivariate Nomianal 1977, Hudson 1977) or Schoener's to test multivariate variables, will be used. theta for each test. tion of observations the habitat relationships The multivariate R 2 of relationship at a given site. between between an species, variables (McFetridge (Linton et al. 1981), between such nominal values and a multivariate theta is equal to the propor- that may be correctly parameters In addition, Scale Analysis MNA generates the frequency and independent (1970) index and type will be tested table). analysis will be made testing the relationships strength Average Some of the analysis will be similar to that for Experiment Analysis designed (1971:265). is not taken or can not be reasonably during an observation, average weather Snow depth will be similar to those reported by Geist wind speed will be determined weather photographically R predicted 2 on the basis of values demonstrate the each species and the habitat variables. �108 MNA also generates relationships independent a series of coefficients between various variable independent and the dependent that take into account variables variable and between each (Andrews and Messenger 1973). e. Schedule: Experiment a. H4 = See schedules 4: Mountain Mountain patterns b. H o = Goat and Bighorn goats and bighorn Mountain Sheep Activity Patterns sheep exhibit the same activity goats and bighorn sheep do not exhibit the same activity in alpine habitats. Methods: Telemetry tip-switch activity a remote portable telemetry seasonal in selected habitats. activity and 4 bighorn site (RPTS) As telemetry exceeding collars, a/ in conjunction collars without battery previously tip-switches life, transmitter InitiallY,4 discussed mountain collars. become inactive failure, or investigator's mountain collars. ability activities, These tests are designed (do to goat or collars. to correctly to measure the identify various headup and such as feeding, resting, or moving, by examining ~/ Consists of pole antenna, receiver, digital data processor, chart recorder. These the RPTS will be field tested with animals outfitted with tip-switch headdown goats for each species. sheep death) they will be replaced with activity Initially, with will be used to sample sheep will be outfitted with activity collars will be part of those bighorn 1 and 2. in alpine habitats. patterns c. for Experiments and strip- �109 data from the RPTS's strip~chatt collared an fma l , observers from the strip-chart. and 1600-1800) collection periods. (with binoculars (SCR).~/ For every activity- Hill be trained Ln Lnt er pre t Lng the activity. Stratified 1200-1600, recorder hour periods During these hours, activities or spotting recorded signal strength on a tape recorder interpreting the activity new activity collar outfitted SCR interpretation. angle", tightness collar's data recorded angle"), when the tip-switch changes of mountain to on the strip-chart on the SCR. This of correctly Similarly, each on an animal will be tested to ensure Such variables'as collar "switching and the behavior the period goat and bighorn neck (hence, the of the animal determine (msec between pulses). sheep activity will involve using the RPTS to sample 24-hour periods (four 2-day periods) or related are confident of the collar on the animal's "pendulum Collection behavior period and up and down collar positions. until the observers correct will be observed scope) during every other S-minute another person who will record the observed testing will continue 0900-1200, will be chosen during the general data by 1 person and constantly as it records (0600-0800, each month. The activity-collared pattern data four times animal to be used for sampling during any 2-day period will be chosen at random without replacement procedure until all such animals have been used. will then be repeated. the SCR will be divided moving. Activity patterns into three categores: Each pattern will be considered The transcribed from feeding, resting, and on the basis of dominant ~/ SCR records amplitude or signal strength and periods (msec) between pulses where ~900 msec indicates headup activity (65 pulses per minute) and ~650 msec indicates headdown activity (90 pulses per minute). By examining the change in signal strength and period, feeding, resting, and moving activities can be discriminated. �110 activity (i.e. interspersions interactions, of walking, etc. for periods Each pattern will include Feeding of the following: standing, licking, d. Analysis: intervals Duration of each feeding, (in minutes) goats). will be divided as will be necessary end, and duration into hour, 15-minute, to determine resting, of each of these activities activity with activity will be made within student's feeding, collars. t-test. resting, and moving Comparison the frequency activity the beginning, periods, will be tallied goat and bighorn sheep of the means of each species and between In addition, or and moving activity. for each 24-hour period and for each mountain outfitted position or running The strip-chart 5-minute bedded in head bedded in prostrate (common for mountain - walking drinking bedded and ruminating, curl position, Moving intraspecific 5 minutes will not be delineated). < - grazing, mineral Resting standing, species with the distribution of the both within and between species, will be tested for independence contingency table). species 2 (x , 2 x C Schedule Fiscal Years Activity Prepare and complete study plan, trap and radio collar mountain goats and bighorn sheep with activity Monitor activity Monitor activity patterns. patterns collars. 1981-82 1982-83 �111 Same as 1982-83 1983-84 Same as 1983-84 1984-85 Data analysis second and publication. generation Prepare and complete 1985-86 study plan if appropriate. Personnel D. F. Reed Principal 1 position Seasonal Employees Graduate Research *Vacant (1 position) *Estimated Annual Investigator Costs Person-days (01) Personal Services (21) Operating (28) Travel Costs $ 40,000.00 520 and Services Supplies 7,000.00 Expenses (31) Capital Asst. 1,500.00 Expenditures 1,500.00 (Equipment) $ 50,000.00 *Costs are expected to inflation. E. GEOGRAPHIC and Rogers selected access F. include Peak areas because: all of the alpine located 1) both mountain is reasonable; 3) current per annum of a future mountain RELATED PROJECT FEDERAL Federal Aid Project W-126-R in the Goliath east of Mount Evans. goat and bighorn management the possibility Colorado from 5-10% due LOCATION The study area will Lake, to escalate Lincoln This area was sheep are present; strategies goat removal Peak, may not preclude experiment. 2) �112 LITERATURE Andrews, F. M., and R. C. Messenger. Analysis. Baumann, 1973. Multivariate Inst. Social Res., Univ. Michigan, T. G. Undated. sheep and mountain studies. goat study. 38pp. 1969. 1980. Population tailed ptarmigan Fort Collins. Elk population dynamics, Clover, M. R. 108pp. and ecological July, Part II, 221-313. habitat, in Colorado. Ph.D; Dissert., and movements Colorado of white- State Univ., l89pp. Cairns, A. L., and E. S. Telfer. ungulates Ann Arbor. Typescript. Colo. Div. Wildl. Game Res. Rep. Braun, C. E. Nominal Scale Final report for the 1978 Mount Evans bighorn Bear, G. D., and R. A. Green. Cole, L. C. CITED 1980. in boreal mixedwood 1956. 1949. Habitat use by 4 sympatric forest. J. Wildl. Manage. 44:849-857. Single gate trap. Calif. Fish and Game 42:199-201. The measurement of interspecific association. Ecology 30:411-424. Denney, R. N. 1977. Status and management of mountain Pp. 29-36 in W. Samuel, and W. G. Macgregor the First International Ministry of Recreation Victoria, Dice, L. R. Geist, V. B.C. 1945. between Mountain Measures F. A. and subalpine l64pp. of British Columbia Fish and Wildlife of the amount of ecological Ecology Branch. Mountain 1978. sheep: a study in behavior Ecological communities. association 26:297-302. Univ. Chicago Press, Chicago and London. Harrington, Proceedings 243pp. species. 1971. (eds.). Goat Symposium. and Conservation, goats in Colorado. and evolution. 383pp. segregation Ph.D. Dissert., of ungulates Colorado in alpine State Univ. �113 o. Hjeljord, 1973. Alaska. Hudson, Mountain J. Wildl. R. J. 1976. herbivores. D. H. utilization 1980. on sympatric J. A. 1979. analysis. J. Ecol. R. J. J. R. J. Zool. 1976. range. by wild attribute and availability Ecology Habitat J. Wildl. data. Ecol. measure- 61:65-71. use by white-tailed Manage. 1981. 40:645-657. Resource utilization 50:283-292. of the variability Strategy of some successional using multiple Peromyscus of resource Univ Alberta, An experimental discriminant use by mountain Edmonton. on Mandarte in l48pp. study of habitat maniculatus, goats selection Island, by the B.C. Can. 59:589-597. Neu, C. W., C. R. Byers, and J. M. Peek. of utilization-availability J. M., D. L. Urich, and relationships Wildl. partitioning 67:255-272. Thesis. 1981. deer mouse, Peek, preference. assemblage-types 1977. M.S. of usage J. Anim. A study plant Alberta. of large of nominally-scaled and F. J. Wrona. an assessment. and climax Merkt, analysis resource L. R., R. W. Davies, McFetridge, a community and resource S. R., and F. F. Gilbert. Matthews, within The comparison for evaluating indices: in Sci. 51:101-110. deer and moose Linton, preference 37:353-362. division multivariate Northwest Kearney, Resource Habitat ruminants: ments Manage. and habitat Nat. Can. 103:153-167. ------- ,1977. Johnson, goat forage Monogr. 48. data. 1974. J. Wildl. and R. J. Mackie. to forest management 65pp. A technique 1976. Manage. Moose in northeastern for analysis 38:541-545. habitat selection Minnesota. �114 Rideout, C. B. 1974. of the mountain Lawrence. Rosenzweig, M. L. 1973. Habitat Heteomyid R. C. 1981. on extensive Sale, P. F. goat in western Overlap 1970. habitats. in resource of habitat selectivity of ungulates use and interspecific competition. by location telemetry. 1979. Ungulate Nonsynchronous habitat Pages 178-186 in F. M. Laramie, Wyo. spatial overlap of lizards in patchy Ecology ..51:408-418. feral burros and desert bighorn N. H., R. J. Hudson, Determinants H. R. 1975. in Colorado. Singer, F. J. 1981. Ecological sheep. relationships Wildl. Monogr. V. C. Brink, and W. D. Kitts. of spatial distribution J. Wildl. Manage. habitat 54:111-117. and R. D. Taber. R. F., and R. D. Ohmart. Smith, B. L. Ecology with a pair of Proc. Second Int. Conf. on Wildl. Biotelem. T. W. cervids experiments J. W~.ldl. Manage. 45:187-196. ranges. determined Long, ed. Shepherd, Ph.D. Thesis, Univ. of Kansas, 17:245-256. selection Shannon, selection First approximation Schoen, J. W., G. N. Thornburgh, Seegmiller, Montana. rodent species. winter 1974. Oecologia Schoener, study of the ecology and behavior 146pp. coexisting Rounds, A radio telemetry of Rocky Mountain of 78. 58pp. 1975. bighorn sheep. 39:387-401. Vegetation of two dissimilar bighorn sheep ranges Colo. Div. Wildl. Div Rep. 4. 223pp. 1979. Habitat partitioning in Glacier National 1977. Influence and wildlife Park, Montana. of snow conditions use, and group size of mountain relationships of J. Wildl. Manage. 43:437-444. on winter distribution, goats. Proc. Int. Mountain Goat Symp. 1:174-189. Springer, J. T. 1979. triangulation. Some sources of bias and sampling error in radio- J. Wildl. Manage. 44:381-388. �115 Telfer, E. S. deer. Wecker, Winter habitat selection by moose and white-tailed J. Wi1d1. Manage. 34:553-559. S. C. Weins, J. A. 1970. 1964. 1976. Habitat selection. Population Rev. Eco1. Syst. 7:81-120. responses Sci. Amer. 211:109-116. to patchy environments. Ann. �116 Appendix A. Location of study area. 1 ka - --...._ - -. �117 Init t aLs Collar No./S.Jlor Ear tag No./Color Left/rigr.t ear Trap site (em): He3.surements 1. Girth 2. Body 3. Nuzzle 4. Left horn length 5. Right horn length 6. Hid left f r cr.t hoof to br i ske 7. Hid left front hoof to knee 8. Hid left front hoof to top dew claws 9. Hid length to bas e left horn left front hoof to scapula 10. Le:t hind fOG~ (horn annuli Genitalia Gonads/udder \lei~ht (kg) Comment; t length __ ) (male or female) condition (animal + sling - sling __ ) _ �118 Appendix C. Vegetation units mapped by Braun H-I Ca rex+Ceurn-T'ri Eo Li.urnAlpine H-2 Carex-Geum M-3 Salix M-4 Geum-Poa M-5 Deschampsia M-6 Pinus-Potentilla-Poa-Carex M-7 Potentilla-Poa-Carex M-8 Trifolium-Carex M-9 Potentilla-Poa M-IO Carex-Poa M-II Alpine (1969). Meadmv i'leadmv Wet Meadow He adow Alpine Sno\.]Accumulation Area Dry Krummholz Dry Meadow Alpine Meadow Dry Meadm'7 Cushion Meadow .Salix-Carex-DeschamDsia Conifer Wet Meadow M-12 Carex-Poa Krummholz M-19 Salix-Geum-Carex M-20 Artemisia-Geum M-2l Salix-Carex-Trifolium M-22 Carex-Geum-Kobresia M-23 Salix-Potcntilla-Carex-Poa M-24 Salix-Picea-Carex Wet Meadow Snow Accumulation Alpine Alpine Krummholz Area Headow Mcadow Alpine Meadow �J..J..::1 I J III P I :II -_--_ ..---- ---~----- ~-- , --------- Lake =-: --~ ,_,_.J '834 83 Flats I \ .rc.~. ./-------- Summit / <, ....._ . . _------;----------r>: \ -' -, �120 Ap?endix C. (continued) Description of vegetation units (modified from Braun 1969). maT DESCRIPTIO:'l Ca rex -Ce um-T'ri f ~!-l o li.urn Alpine He adow : Species of Carex cover 25-50 perce~t of the total area of this unit, while Geu:n rossi a"d Trifolium spp. (I. E~ in M-la and !. dasvohvllum in M-lb) each amouets to 5-25 percent of the total coverage. Plant height is less than 1/2 m aed density is mostly continuous, although plants in limited areas occur singly or in small patches. Stones are nu:nerous, and 5-25 percent of the total cover consists of rocks larger than 12 inches in diameter. Black and green lichens ~re common, with black liche~s covering 5-25 perce~t of the exposed surfaces. Slope gradients vary up to 70 perceet, but average between 3S and 40 percent. ::-1- 2 Ca rex-Geum Alpine Hea~;·'.T:Hc,::i:>ers of the Careces cover 25~SO percent of the total area of the M-2a subunit but cover only 5-25 percent of subunit H-2b. Geum rossi accounts for 5-25 percent of the coverage of the entire u~it. Height of the vegetation in this UGit is uniform and is less than 1/2 m. DEEsity is mostly conti~uous, although pla~ts in limited areas are scattered singly. Rocks larger than 12 inches in diameter are abundant and amount to 5-25 percent of the total coverage of subunit M-2a and 25-50 percent of the coverage of subu~it M-2b. Black and green lichens each covers 5-25 percent of the exposed rock surfaces. Areas included in this unit are extremely variable in slope, with some areas aver3gi~g 35-40 percent, others 20-25 perce~t, and still others 50-55 percent. M-3 Salix Wet :leadow: Salix s pp , cover 50-75 p ercen t of this unit, with all bU3hes being less thaG 2 m in height. A few clumps of Ficea eQ~elm~ii are scattered and attaia a height of 2-3 m. Density of plants is continuoGs and few rocks are present. Lichens are uncommon, and the slope averages abo~t 15-17 percent. H-4 Ge urn=Poa Alpine :-leadm.J:Geum rossi and Poa spp. are codorninant3 in chis unit, with ~ach covering 5-25 percent of the total area. The ve~etation of thi3 ~nit is short, less than 1/2 m in height, and the density is contin~ous. Rocks larger than 6 i::lchesin diameter are co~~on, and tho3e larger than 12 inches cover 5-25 percent of the e~tire area. Black and gree~ liche~s �121 Appendix C. (continued) Description of vegetacion units (modified from Braun 1969). U~IT DESCRIPTION dominate. wit~ each covering 5-25 percent of the exposed rock surfaces. Slopes are moderate and average about 20 percent. :1- 5 Deschamosia Snow Accumulation Area: DeschamDsia caesDicosa covers 5-25 percent of this unit, with Sibbaldia Drocumbens and Artemisia spp. being prominent in limieed areas. Height of che vegetation, while taller than that in most alpine areas, is less than 1/2 m. Rocks larger than 12 inches in diameter are abundant and cover 25-50 percent of the total area. Lichens, wh i l,epresent, are unc ommo r;, and their total cover is quite small. Slope gradient is moderately steep and averages about 40-~5 percent. Pinus-?o!:e:1cilla-?oa-Carex Dry KrumrnhoLz : This unit is the transicion becween the subalpine forest and the alpine tundra, with each of the four dominant genera covering 5-25 percent of the total area. The appearance of this unit is ragged, with some Pinus aristata and flexil~s trees approaching 5-6 m in height. Planes occur singly or in small patches, with rocks of all size classes being common. From 25-50 percent of the area is covered with rocks larger than 12 inches in diameter. Black and green lichens each covers 5-25 pe~ cent of the exposed rock surfaces. Slopes are moder: ately steep and average 40-45 percent. R. H-7 Potentilla-Poa-Carex Dry Neadm,,: Species of Potentilla (pri.ncLpa l Ly T". fruticosa and .E. diversifolia), Poa, and Carex dominate this unit, with each genus covering 5-25 percent of the entire area. Scattered conifers which grow to a height of 2-3 m occur in this unit, but most shrubs are less than 1 m tall, and all herbaceous vegetation is less than 1/2 m tall. Plant growth is not continuous, and most occur in sffi311patches or singly. Rocks of all size classes are conspicuous, but to~al rock cover is dominated by stones larger than 12 inches in diameter as this group covers 25-50 percent of the entire area. Green and black lichens are common with each type covering 5-25 percent of the exposed rock surf3c~s. Slope gradienc averages 35-40 percent. ~·1-8 'I'r i f o li u-n-Cc rex Alpine Neadow: Trifolium spp. (I. nanum in SUDu:1it :!-8a and T. dasvnhv llum in subunit ~1-3b) and Carcx spp. each ~ccour.t tor 5-25 percent of the toc31 cover of this unit. Plant height is short, with average height not exceeding 1/2 m. Density is moscly conti~uous in subur.it X-Sa, ~ut is equally �122 Appendix C. from Braun (contiilueJ) J~scription of vegetation units (modified 1969). DESCRTPTI00! UNIT ccn t i.r.uou s Cio.3i:::srnaL_~pa t ch es i.n s ubuni, t M-8b. Rocks larger than 12 iGches in diameter comprise 5-25 percent of the tota~ cover i~ subunit M-8a and 50-75 percent of subu~it M-Sb. Black and green lichens each covers 525 perceo.t of the exposed surfaces of stones in both subunits. SLbu~it M-8a has gentle slopes which average about 20 percent; 5~opes in 3~tu:::itM-8b average about 75 pe r c e r.t., \ H-9 Po t en t Li La-e Po a Dry ~leadc~.J: Pott?':::.s.~s pp , (primarily frucico:;3 and d i.vers i f o li a i and Poa s pp , each covers 5-25 percent c r this unit. Or.e s ma I I area Le s s than I acre ~ con t a Ln i ng a s pr i r.g ar.d d omi.r.a ted by Sa lix _!:. _!:. spp. bLSht?5 ~p to 2-3 m ta:l, is i~cluded in chis uc.it. Avera~e h~igh: of most vegetation is lesE: chan 1/2 m, a Lchough s ca t cered bu she s of Po t en t i lIe. f ru t i c os a may slightly exceed this height. Plane density in this unit is mostly interrupted, as rock classes les2 than 6 inches and larger than 12 inches in diameter each covers 5-25 percent of the total area. Lichens are COmI1l0c.,and those i:-,the b Lack a nd g reen ca t egor Le s each covers 5-25 perc~nt of the avai:2bie rock 3urfaces_ Slope gradi2~t is gectle a~d averages 15-23 perceGt. H-IO Ca rex+Po a Cu sh i.on ?:-12ado",:Carex: s p p , cover 25-50 perc er.t of this uni t, wh i I.e members of the genus Poa com- prise 5-25 percE~t of the total cover. Cushioc plants such a~ TrifoLi~m Da~~m and Are~ari3 obtusiloba are c ons p i cuoi..scut rnake up only srnai l p erce nc age s of the total cover. The vegetation is typically less than 1/2 m tall, arid p lar.ts occur singly or in small patches although in sm~l~ areas continuou~ growth is most prevalent. St.or.e s of a I L d Lame t e r classes are c ornrno n , with those larger than L2 iClches io. diameter ccverir:g 25-50 pe rce r.t of t hc area. Bl..1ckand g re en Li.c hc ns dominate, with each type acco~nting for 5-25 percent of the cover of expos~d·sLrf3ccs. Slopes are moderately steep, and the gradic~t averagc3 3bout JO percent. :-1-11 S,l1i:·:-CJr(C:-:-·Ol'sch:1!!1!)"ia Het ~le:JJo\v: Species of S.31ix C~lr:::: do;;'i_('-:]cc -~hi.; .mi t , w i t h mcrnoers o r cach ger:~s coveri~g 25-50 percel1t of the cot.]l .1rc.1. DcschJmo:ia C:1050itosa i~ an imoort3nt component of t h i s typ;:;---;-nd .irnov n t s ro 5-·25 p~rce.:1tof the t o t a I cover. PIdcr height in this ~~Lt is up to l m, with few bU3hc~ ex:ccediG~ thi3 level) and the Jensity is conti:~;_:oLJ~.Rc'cks la rgcr chan 12 inches in d i arnet er 311d �Appendix C. (continu~d) 123 Description of vegetation units (modified from Sr~un 1969). UNIT DESCRIPTIGt·! are c ornrnon a nd cover 5-25 percent of the total area. Lichens, while present, are not important in terms of cover2ge of rock surfaces. The slope gradient averages 25 perce:-:t. :1-12 Carex-?oa Conifer Krummholz: Conifers such as PinGS ariS[2t~ and Picea enzelmannii are conspicuous in this unit aud combined comprise 5-25 percent of the total cover. Carex spp. and Poa spp. each covers 5-25 percent of the area of this unit. Much of the v ege t a t i.or; of this unit is less than 1/2 m in height, but scattered islands of evergreens reach a height of 2-3 m. Plant density is mostly contin~ous. although stones are nurnerou~. Rocks larger than 12 inches in diameter cover 5-25 ?ercenc of the total area, and 5-25 percent of their sur~ace is covered equally by green and black lichens. Slopes are variable and average 15-20 percent. :1- 19 Sa Li x -Criurn-Cc rcx \-Jet;-leadm.,r: Cc urn r o ss i , Ca rex spp., anJ :.):11i:·: s pp . each comprises 5-25 pc rcen t o t the cover~ge of this area. Plant Jensicy is continuous, and vegecation appearance is irregular as some bushes a f S;J 1i:.:are a b out 1 m tall. S ton c s , w h i 1c comma n , cover less than 25 percent of the unit, with those larger than 12 inches in diameter being most prevalent. Green lichens arc most nurncro~s and cover 5-25 perce~t df the expo3ed rock surfaces. Slopes are moderately steep a~d average 55-60 percent. (-1- 20 Arte~isi3-Ceu~ S~aw Accumu13tion Area: Total vegetation cover of this uni~ is s~all, with both Arte~isia spp . (pric3rily!::_. r..o-r\·~::.ica) and _g=~ rassi cov er i.ng less than 5 percent of the total area. Rocks of all diameter size classes are abucdact, with those in the 0- to 6-ir..chclass covering 25-50 perceGt of the area, while those larger than 12 inches co~?rise 50-75 percent of the total cover. Lichens are inconspicuou8, ~nd slopes are moderate, averaging 30 percent. Sali::-Care:·:-Trifoliu::, .-\lpi:12 ~-:eadO';-_·: Sa l i x spp , and Care:·:S?p. each covers 25-50 percen r or [["11..'3 unit, while Trifolium dasvohvllurn covers 5-25 percent of the total area. l:ierDac~vegetation present is less t ha n 1/2 m tall) while some bushes of Salix attain a height of 1 m. Vegecation density is co;tir..uous, although some planes occur singly in limited 2reas. Stones large:- than 12 inches in diamete!" are COIT::1on a[ld comprise 5-25 percenc of the entire cover. Greec lichens �Appendix C. (continued) 124 Description or vegetation units (modified from Brnun 1969). are dooinant and cover 5-25 percent of exposed rock surfaces. Slope gradients are moderate and average 20-25 percent. H-22 ~12zdm-J: Carex spp. (p r i nc i> pally ~. ely~oiccs) cover 25-50 percent of this unit1 with G2U!":1 Tassi arid Ko bres La _t..--._.___ rnv o su ro i de s e a ch c crno . r i s> ing 5-25 percent of the total cov~r3ge. Pla~t height is mostly less than 1/2 m, although a few scattered bushes of Sa I i x may slightly exceed this level. The density of p La n t s ir'_this un i t is c ort i nuou s , This area appears smooth, but rocks larger th3'- 12 inches ir. diameter do occur a~d cover 5-25 percent of the total area. Lichens arc evenly divided bctwee~ green and black, wit~ each type covering 5-25 perce~t of exposed surfaces. Slopes arc variable and average 25-30 percent. H-23 Sali~-?ot0~till~-C~~0x-?o3 Alpine ~eadow: This unit ii domin~teJ by SJiix spp. which COVEr 25-50 pcrcc,-t of the a rea , .f.0t·.':l~.:U.l2. s pp (p r i ma r i ly _E. frucic:)S2 ar:d P. div~r~i[01i3), C~re~ spp., and Poa spp. ar2 su~doDi~.:!I~r5 a;~3chcov~~S-25 percent-of the unit. This unit appears p~~chy as vc~et3tion ir:-. some are3S exterd3 Ca r ex-Ge cm-Kob re s i a Alpine to a height of 1-2 m, while that in others is less than 1/2 m tall. Plant density is continuous, and rocks are not abundant. Those larger than 12 inches in diameter cover 5-25 percent of the unit and in turn, up to 50 percent of their surfaces are covered equally by green and black lichens. Slope gradients are moderate and average 40-45 percent. H-24 Salix-Picea-Carex Krurrunholz: Species of Salix and Picea engelmannii cover 25-50 percent of this unit. while Carex spp. cover 5-25 percent of the total area. Appearance of this unit is irregular as islands and strips of Picea e~gelmannii extend to a height of 2-3 ~ Salix bushes in this area are less than 2 m tall, while the incersoersed herbaceous vegetation does not exceed 1/2 m in height. Plant density is continuous, and rocks are conspicuous because of their size. Those larger than 12 inches in diameter cover 5-25 percent of the area and in turn are 5-25 percent covered by black lichens. Slopes are relatively uniform and average 17-20 percent. �125 Appendix D. Study area divided into 500 m square quadrats based on UTM coordinates. 500 m �AI;>pendixE. Mountain goat-bighorn sheep location and observation data. No. Date Time Species UTM ---right up Habitat type Activity Group size Air temp Cloud cover Wind spd Hind direction Sn ow cover/ depth Terrain I 2 3 --4 --- 5 6 -- -- Collar or ear tag 1 2 I 3 4 5 6 a/ Group cl~ssification:- K/L YF YM YU 2F 2M 2U 3F 3M 3U AF AH I 2 3 4 5 6 a/ K = kid, L = lamb, Y = yearling, 2 = 2 year old, 3 = 3 year old, and A = adult. ---- AU 'JNC TOTAL t-' N 0'1 �Appendix F. Variables for each mountain goat and bighorn sheep location (after Hudson 1977). Variable name 1 .2 Dependent variable Species Mtn goat Bighorn sheep Stratified variable Weather Clear, warm Windy Cold Snow/ rain " Time Dawn-mid morning Late morning Early afternoon Late afternoondusk " " Month Oct Nov Dec Jan Season Fall Winter Spring " Activity Variable Independent variable Slope (percent) Categories Feeding 0-5 Resting 6-10 :3 4 6 7 8 Feb Mar Apr May -- -- -- -- -- 11-20 21-40 4l-50 51-60 61-70 71+ East Southeast South Southwest West Northwest 601-700 701+ Sali/ Trda/ 5 Moving " Aspect " Distance (m) to escape terrain 0-100 101-200 201-300 301-400 401-500 501-600 " Elevation (m) 3,2003,400 3,4013,600 3,8014,000 4,0014,200 4,201+ " Terrain Slope Rock outcrop 3,6013,800 Cli;ff Couloir Ridge " AI.pl.nea/ Artr/ Gero Carol Deca/ Gero/ Piem/ Posu " Nonvegetation Mineral lick Roadbedshoulder Rock Snowfield North Northeast Posu/ t-' N -....J �Appendix F. (continued) Variable Variable name 1 2 3 Independent variable Snow depth (em) 0 1-10 11-20 21-30 Snow cover (percent) 0-25 26-50 51-75 76-100 0-(-10) (-10)(-20) (-20)(-30) (-30)+ 0-2.0 2.1-4.0 4.1-6.0 6.1-8.0 " " " Temperature (OC) Wind speed (m/sec) 10-0 0 Categories 4 5 6 7 31-40 41-50 51+ 8.1-10.0 10.1-12.0 8 12.1+ !!:_I Abbreviations are first 2 letters of genus and spec~es name. Each of the following dominant vegetation types occur in 1 or more of the 18 vegetation units (Appendix C). Artr/Gero = M-20; Carol = M-1, M-2, M-10, M-12, M-22; Decal = M-5; Gerol = M-4; Pien/posu = M-6; Pasul = M-7, M-9; Sa1il = M-3, M-11, M-19, M-21, M-23, M-24; Trda/ = M-8.-- t--' N 00 �129 JOB PROGRESS State of July, 1982 REPORT Colorado Project No. W-144-R-l Big Game Investigations Work Plan No. 5 Period Covered: July 1, 1981 through June 30, 1982 Personnel: - Noncervids Black Bear Investigations T. D. I. Beck ABSTRACT Twenty black bears were captured during the 1981 season, 9 initial captures and 11 recaptures. Of 32 males tagged in 3 years at least 10 are known dead and possibly 2 others, in. spite of a closed hunting season in the study area. Female age structure in the area is dominated by 2 cohorts, AC 6 and 7, which comprise 33% of the 1982 tagged population. Age of first litters for 3 females was 5 years for 2 and 4 years for 1. Six females have not littered by 5 years and 2 haven't by 4 years. Of 7 yearlings collared in dens, 3 starved to death before emergence. Natural logarithm transformation regression equations were prepared to estimate body weight from chest girth. Denning began the first week of October and was completed by the 3rd week of November. Den emergence began the second week of April, continuing through the 3rd week of May, with most of the bears emerging between 24 April and 15 May. Physical characteristics of 25 densites were described. Mean annual range for 6 adult females in 1981 was4,837 ha, for 5 non-breeding age females was 2,590 ha, and for a single female that makes a long migration for berries was 15,267 ha. A detailed study plan for the population ecology study (Job 1) was approved and a study plan for habitat selection of females (Job 2) was prepared. ��131 BLACK BEAR INVESTIGATIONS Thomas D. I. Beck P. N. OBJECTIVE 1. Describe population dynamics of a selected black bear population for development of a single-species population model. 2. Develop and test hypotheses mechanisms. relative to black bear population to allow regulation SEGMENT OBJECTIVES 1. Determine habitat preferences of selected black bear populations. 2. Describe black bear population dynamics sufficiently various harvest and habitat manipulations. to allow analysis of ACKNOWLEDGEMENTS Assistance with field work was provided by the following Division of Wildlife Personnel: D. Baker, R. Bartmann, G. Bock, L. Carpenter, D. Coven, M. Dege, B. Gill, L. Green, M. Grode, M. Haroldson, D. Miller, R. Parachini, J. Ritchie, B. Stark, and L. Stevens. Their assistance was extremely helpful and deeply appreciated. Additionally, welcome assistance with den work was provided by: F. Hammond and H. Harlow, Univ. of Wyoming; J. Rennicke; and N. Vincent, D.V.M. METHODS AND MATERIALS Capture and Marking Capture efforts lasted from 17 May to 30 June and 1 September to 5 October, 1981 in the Black Mesa Study area (Fig. 1). All captures were made using Aldrich spring-activated snares. All snares sets were out of sight from roads and maintained trails and were baited with fish and molasses. Snares were checked daily. Snared bears were immobilized with a combination of ketamine hydrochloride and xylazine hydrochloride in a mixture of 180 mg ketamine and 90 mg xylazine per milliliter. Drug was administered by use of a 1.8 m jab pole. Denned bears were drugged by use of a jab pole 0.5, 1.2, or 1.8 m in length. Drug dosage was 6.6 mg ketamine per kg of body weight. Bears were ear-tagged in both ears with numbered, plastic Roto-tags. All female bears age class (AC) 3 or older and male bears AC 4 and older were instrumented with a Telonics radio transmitter collar. Yearling bears that were handled in dens with their mother, and thus known to have been born in the study area, were instrumented with small radio-transmitter collars �l32 Figure 1. Black Mesa, black bear study area (outlined with stippled line). �133 designed to operate for 16 months. These small collars were removed from AC 3 bears during the denning period and replaced with the large radio-collars. Numerous physical measurements were taken and a premolar removed for subsequent aging by cementum annuli counts. Blood samples were taken from most bears and urine samples were taken from 6 males. Any bear caught or handled in a den that had been wearing a radio-collar for 18 months or more was fitted with a new radio-collar. Habitat Use Data on habitat use were collected primarily from ground tracking radiocollared bears supplemented by aerial tracking. Location of bears within descrete vegetation communities was not attempted in this segment but a graduate program proposal was developed to investigate habitat use by female black bears. The proposal is still undergoing internal peer review. Den sites were located in October and November 1981 by both ground and aerial tracking and den entrances marked with vinyl flagging. Some dens were not located until March 1982. Dens were visited in January-March 1982 and numerous physical measurements of the den were taken as well as a general description of the site. Home Range Calculations All bear locations described by UTM coordinates will be used to calculate seasonal, annual, and multiple-year ranges. The spatial description of the ranges will employ the minimum polygon procedure and utilize the HOME computer p~ogram described by Harestad (1981). Initially seasonal ranges were established as spring (April IS-June 15), summer (June l6-September 15), and fall (September l6-December 26). Preliminary analyses and knowledge of plant phenology have subsequently shown these dates to be poor choices. Therefore, only annual ranges will be reported in this segment. Future analyses will probably use April IS-June 30 as spring, July I-August 15 as summer, August l6-September 30 as fall, and October 1 to December 31 as pre-denning. These time segments correspond better with food availability and bear movements. RESULTS AND DISCUSSION Capture and Marking Twenty-two black bear captures were made during the 1981 season of which 9 were initial captures, 4 were recaptures of bears marked in 1979, 6 were recaptures of bears marked in 1980, 1 was a recapture of a bear tagged in a den in 1981, and 2 were recaptures of a bear initially caught in 1981. A summary of all bear captures in 1979, 1980, 1981, and den season 1982 is provided in Tables 1 and 2. Estimated ages are generally from cementum annuli counts except in a few instances where the lab estimate was obviously in error. The cementum annuli technique needs a more rigorous testing with known age animals. On several occasions where teeth from recaptured bears were sectioned the estimated age had not increased as much as it should or had decreased. Review of Phillips et al. (1982) points out the uncritical �134 acceptance our profession has generally given to this technique. Two observations are readily apparent from Table 1; the young age structure of the males and the relatively high man-related mortality rate even with the study area co sed to hunting and little or no apparent livestock conflicts. In addition there are 2 males we believe have been illegally killed but can not confirm at present. With the current level of mortality it is doubtful if the resident population will increase nor undergo an upward shift in age structure. Cursory examination of Table 2 indicates an age structure dominated (33%) by AC 6 and 7 if one assumes all the bears are alive and correct ages to June, 1982. As these females are all residents and just becoming of reproductive age the reproductive output of the population should increase. The age at which female black bears first have cubs in the study area is greater than expected based on studies elsewhere (Bunnell and Tait 1981) and the relatively large size of the bears. Interpretation is complicated by a catastrophic berry failure in the fall of 1981 which may have caused a drop in cub production (Rogers 1977). This food failure came in a year when as many as 8 females reached an age when they could have conceived their first litters. As of June 1982 2 females were known to have littered for the first time at 5 years (AC 6) and 1 at 4 years. However, 6 females are known not to have ever had cubs at 5 years and 2 haven't had cubs by 4 years. Ages of first littering for the older females is unknown. Of 6 females checked in March 1982 that were AC 6 or older and not accompanied by yearlings, only 1 had a litter. Only 4 litters of cubs have been recorded from checks of dens in 1981 and 1982 and the litter sizes were 3, 3, 3, and 2. Survival of cubs born in 1981 was poor. Only 2 of 6 that were radio tracked survived to 1 June 1982. One of the cubs disappeared in September 1981, and 3 starved to death in winter dens, thus leaving each of the females with only 1 yearling remaining alive from triplet litters. The fate of the other triplet litter is unknown because the female denned so deeply in the rock cliffs that no radio signal could be detected. Estimation of liv~ body weight from physical measurement has been reported by other workers throughout North America (Cherry and Petton 1976, LeCount 1977a, Payne 1976). Analyses of chest girth, neck girth, and both combined as estimators of weight indicate best results are obtained if separate equations are used for each sex and season and with chest girth as the only variable. When x is the chest girth (cm) , the estimated weight in pounds (y) can be derived from the following equations: Males in June or July Males in September Males in January-March Females in June or July Females in September Females in January-March Iny Iny lny lny lny lny For identical chest girths, western bears studied in Tennessee, Arizona LeCount 1977a, Payne 1976). - - 5.823 4.221 5.912 6.966 7.146 -10.311 + + + + + + 2.458 2.096 2.469 2.452 2.495 3.136 Inx Inx lnx lnx lnx Inx r2 r2 r2 r2 r2 r2 0.958 0.889 0.990 0.972 0.931 0.916 n n n n n n 25 18 10 25 14 13 Colorado bears were heavier than the black and Newfoundland (Cherry and Pelton 1976, �135 Table 1. Estimated age and weight of captured male black bears, Black Mesa study area, Colorado. No. Capture Date Estimated Age Weight (kg) M-l M-l 5-29-79 6-29-79 7 7 141 132 M-Z M-Z M-Z M-3 6-13-79 6-Z7-79 9-15-81 7-25-79 4 4 6 12 68 66 138 121 M-4 M-5 M-5 M-5 M-6 7-30-79 8-0Z-79 6-0Z-80 5-27-81 9-11-79 6 2 3 4 3 107 41 76 86 71 X-7 M-8 M-9 8-11-79 8-27-79 9-Z0-79 5 2 1 84 52 48 9-lZ-80 M-9 5-28-80 M-IO M-dead 6-03-80 M-ll 6-03-80 M-ll 9-29-81 M-IZ 6-04-80 M-12 10-07-80 6-08-80 M-13 2 5 9 4 5 5 6 61 127 159 52 95 91 118 130 M-14 M-14 6-16-80 9-18-80 3 3 48 71 M-15 M-16 M-17 M-17 M-18 M-19 M-ZO M-ZO M-Zl M-ZZ M-Z3 M-24 M-Z5 M-Z6 M-Z7 M-Z8 M-29 M-29 M-30 M-31 M-31 M-31 6-18-80 6-19-80 6-Z5-80 9-09-81 7-01-80 7-03-80 7-12-80 6-Z4-81 8-13-80 8-Z0-80 9-14-80 9-25-80 9-Z5-80 10-06-80 6-10-81 9-01-81 6-2Z-81 6-22-82 9-02-81 9-05-81 9-11-81 9-14-81 3 9 3 4 3 3 3 4 68 139 55 95 57 59 64 66 46 34 82 80 32 80 1.0. 5 2 2 4 3 1 4 1 2 3 4 5 3 3 3 11 52 61 59 102 56 56 56 Remarks Killed 7-lZ-79, 15 km south of study area Illegally killed in study area, 11-80 Killed 7-14-80 approx. 155 km southwest of study area Killed by ADC 9-81, 15 km south of study area Illegally killed while in snare Died from drug reaction in den, 3-81 Killed 6-81, 0.3 km north of stutiy area Illegally killed 10-80 in study area Illegally killed 9-81 in study area Killed 6-82 25 km north of study area �136 Table 2. Estimated age and weight of captured female black bears, Black Nesa study area, Colorado. LD. No. Capture Date Estimated Age Weight (kg) F-1 F-1 F-2 6-13-79 6-08-81 6-17-79 6 8 1 70 82 11 F-3 7-01-79 1 13 F-4 F-4 F-5 F-6 F-7 F-7 F-8 7-18-79 8-16-79 7-19-79 7-20-79 7-27-79 7-09-80 8-01-79 2 2 2 4 6 7 5 27 33 27 48 75 64 43 F-9 F-9 F-10 F-ll F-12 8-01-79 6-10-81 9-10-79 9-14-79 9-17-79 7 9 3 12 3 68 61 59 107 50 F-13 F-14 F-15 F-15 F-16 F-16 F-16 F-17 F-18 F-18 5-25-80 6-07-80 6-08-80 9-22-81 6-19-80 9-05-80 10-03-81 6-21-80 6-24-80 6-20-81 4 4 10 11 4 4 5 6 1 2 57 57 98 86 45 54 71 76 15 27 F-19 F-20 F-21 F-22 F-23 F-24 F-25 F-26 F-27 F-28 F-29 F-30 6-25-80 6-27-80 6-30-80 7-01-80 7-09-80 9-03-80 10-04-80 6-19-81 9-14-81 9-16-81 9-19-81 9-20-81 3 5 2 9 1 5 2 2 2 10 7 2 43 61 30 91 13 52 36 31 39 82 84 32 Remarks Illegally killed 11-79 in study area Illegally killed 11-79 in study area Killed 10-79 1 m north of study area Killed 11-81 0.4 km east of study area Had open bullet wound Killed 11-81 0.8 km north of study area �137 Habitat Use Late August was again the time of significantly increased movements by all bears into vegetation communities dominated by Gambel oak (Quercus gambelii). The longest movements were by bears that normally range in Soap Creek and West Elk Creek, east of the study area. F-24 was caught in the Smith Fork area in September 1980 but moved 26 km southeast to a densite in West Elk Creek. She spent the period from den emergence until late August in the West Elk Creek drainage, then rapidly moved the 26 km back to Smith Fork. Although soft mast was nearly nonexistent, she foraged extensively until the 3rd week of September when she moved back to West Elk Creek to den. This makes for a large annual range (see Table 5). Additionally, 3 additional bears were initially captured in Cow Creek and these collared bears all moved 14-22 km southeast into the Soap Creek area. The Soap Creek drainage and the country east of the study area have very little of the mast producing species that are so dominant in the study area; Gambel oak, chokecherry (Prunus virginianus), serviceberry (Amelanchier alnifolia). All radio collared bears had returned to their traditional summer ranges by 1 October. Movements were limited from 1 October until denning with most bears staying within 0.4 km of their ultimate den site during this 2-5 week period. Den entry times in 1981 exhibited the same trend as in 1980 except that more bears denned in late November in 1980 than in 1981 (Table 3). October 15 through November 15, the period of most denning, coincides exactly with Colorado's fall deer, elk, and black bear hunting season. Table 3. Number of radio-collared black bears entering periods, 1981; Black Mesa, Colorado. Males Females October 1-7 October 8-15 October 16-23 1 0 0 1 1 2 Period October November 24-31 1-7 2 5 1 2 dens by I-week November 8-15 1 3 November 16-23 0 1 Emergence from dens began in mid-April and continued into the 3rd week of May (Table 4). The earliest bear leaving a den was a female that ran from the den as I approached in late March. She then spent 10-15 days alternating between her den and a brushed-over day bed but would not tolerate close approach by humans. She moved out of the area during the second week of April. There doesn't appear to be any clear emergence pattern by age and sex groups. The den emergence times are not as accurate as den entry as access is fairly restricted in late April and early May. Most of the radio tracking is done from airplane or long-range ground tracking and thus some of the times may be delayed by a I-week period. �138 Table 4. Number of radio~collared 1982; Black Mesa, Colorado. April 8-15 Male Female 0 1 April 16-23 2 1 black bears leaving dens by l-week periods, Period April 24-30 2 4 May 1-7 May 8-15 May 16-23 1 1 2 5 0 3 Bears were in poor condition this spring because of the very poor soft mast production in fall of 1981 (brought about by hard freezes on 6-16-81 and 6-17-81) yet did not emerge from dens early. The very poor condition was evidenced by: a) 3 yearlings which starved to death in dens; an AC 3 female (F-30) which declined from 34 kg in September to 22.7 kg by January 26; and M-29 which weighed 2 kg less on 6-22-82 as on 6-22-81 even though he should have been a rapidly growing male (AC 3 to 4). No radio-collared bears emerged after 15 May in 1981 yet 3 did in 1982; a female with 2 cubs, a female with 1 surviving yearling, and a very small female (F-30). The progression of weight loss by denned bears is of interest. I was surprised that a small bear had already lost 33% of predenning weight by January 26 and still survived to emerge 4 months later (F-30). All of the male black bear dens located in 1982 were in rock cavities and 7 of 11 had obviously been used by bears in previous years (Table 5). The genera~ lized impression of den site selection is that the bears either go down into rugged canyons or go to high elevation cliff formations. The rock cavities have small entrances but become quite extensive and large on the inside. In comparison, only 6 of 14 dens used by female black bears were in rock cavities (Table 6). The only natal den was an excavated cavity under an aspen (Populus tremuloides) tree. Thus all 4 natal dens in the study have been ground dens. Three of the 4 dens with family groups (mother and yearlings) were rock dens. Five of the 6 rock dens had evidence of prior bear use while only 2 of the 8 excavated ground dens showed prior use. All dens used by both sexes had extensive amounts of bedding material. �139 Table 5. Site characteristics Den Type. Overstory vegetation Elevation (m) Aspect (degrees) Slope (degrees) Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock b Psme Potr Psme/Abla Psme/Abla Psme Psme Psme Pien/Abla Quga Psme/Pifl Potr/Amal 2286 2728 2880 2926 2926 2553 2704 3239 2713 2804 2743 130 20 67 10 l30 100 270 22 270 250 240 17 40 43 50 30 50 58 32 36 70 36 a b cavity · a cav~ty · a cav~ty · a cav~ty cavity · a cav~ty · cav~ty a · a cav~ty · cav~ty a cavity cavity Evidence Psme Pien of use by bears in previous Douglas fir Quga Engelmanns Spruce Table 6. of dens used by male black bears, 1981-82. Site characteristics Gambel oak Potr Pifl = Limber pine Overs tory vegetation Rock cavitya Excavated-blowdown Rock cavitya Excavated-tree base Excavated-shrub a Excavated-tree base a Excavated-tree base Excavated-shrub Excavated-shrub a Rock cavity Rock cavity Rock cavitya Rock cavity Excavated-blowdown Pien/Abco-Abla Pien/Abla None Potr Quga/Amal Pien/Abla Pien/Abla Quga Quga/Prvi Potr/Pien Quga/Potr Psme Quga/Psme Pien/Abla b Evidence Pien Potr Psme Aspen Amal Abla = Subalpine fir = Serviceberry of dens used by female black bears. Den TYI~e a years. of use by bears in previous Elevation (m) 3063 3231 2957 3124 2316 3292 3444 2362 2621 3322 2926 2804 2499 2758 Aspect (degrees) 7 80 l30 102 5 20 230 355 272 160 169 140 280 60 Slope (degrees) 52 34 44 40 48 58 42 42 44 34 43 52 54 50 years Engelmann Spruce Abco - White fir Abla - Subalpine fir Aspen Quga = Gambel oak Amal = Serviceberry Prri = Chokecherry Douglas fir �140 Home Range Calculations Home range calculations have only been completed for selected females for the 1980 and 1981 seasons. These females were potential candidates for more intensive study of habitat use and thus we needed to plot their ranges in order to develop a vegetation sampling scheme within their composite ranges. No major shifts in range occurred between 1980 and 1981 although ranges were considerably larger in 1981, most likely a result of more extensive radiotracking. The smallest range was that of F-19, an AC 5 bear with no young, and was only 824 ha. Her range is an area containing extensive Gambel oak and aspen forests with numerous streams and ponds. The largest range was tht of F-24, the West Elk Creek resident who moves into the area in the fall, and her range in 1981 was 15,267 ha. The mean annual range of 11 resident females in 1981 was 3,816 ha. If F-24 is included in the mean calculation it becomes 4,770 ha. Mean annual range for the 6 females which have littered in 1981 or before was4,837ha (3 females had litters of 3 each in 1981) while the mean for 5 non-breeding age females (all were AC 4 or 5) was 2,590 ha (F-24 was omitted). This general pattern between age groups agrees with findings in Idaho (Reynolds and Beecham 1977) and Arizona (LeCount 1977b). However, annual ranges in Arizona and Idaho were considerably smaller (Table 7). Table 7. Comparison of annual ranges of female black bears in Rocky Mountains. 2 Annual Range (km ) Adult Female (N) Subadult Female Arizonaa b Idaho Colorado a LeCount b Reynolds Of greater years, and aspects of years have 18 (5) 13 (3) 13 (5) 9 (6) 48.4 (6) (N) 25.9 (5) 1977b and Beecham 1977 interest than total size is the spacing of ranges, shifts between changes as subadult females move into reproductive status. These seasonal and annual range comparisons can not be made until more passed and more data accrued. A detailed study plan for the black bear population ecology study was prepared and approved. Readers interested in more specific methodology should request a copy of the revised Program Narrative. �141 LITERATURE CITED Bunnell, F. L. and D. E. N. Tait. 1981. Population dynamics of bears implications. pp 75-98 in C. W. Fowler and T. D. Smith, eds., Dynamics of large mammal populations. John Wiley and Sons, N.Y. 477pp. Cherry, J. S. and M. R. Pelton. 1976. Relationships between body measurements and weight of the black bear. J. Tenn. Acad. Sci. 51:32-34. Harestad, A. S. 1981. Computer analysis of home range data. Brit. Col. Fish and Wildl. Branch, Fish and Wildl. Bull. No. B-ll, 25pp. LeCount, A. 1977a. Using chest circumference Digest, Ariz. Game and Fish Dep., 2pp. to determine 1977b. Some aspects of black bear ecology Proc. Intl. Conf. Bear Res. and Mgmt. 4:175-180. bear weight. in the Arizona Wildl. chaparral. Payne, N. F. 1976. Estimating live weight of black bears from chest girth measurements. J. Wildl. Manage. 40(1):167-169. Phillips, C. J., B. Steinberg, T. H. Kunz. 1982. determination in bats: a critical evaluation. Dentin, cementum, and age J. Mamm. 63:197-207. Reynolds, D. G. and J. J. Beecham. 1977. Home range activities and reproduction of black bears in west-central Idaho. Proc. Intl. Conf. Bear Res. and Mgmt. 4:181-190. Rogers, L. L. 1977. Social relationships, movements, and population dynamics of black bears in northeastern Minnesota. Ph. D. Diss .. , Univ. of Minnesota, Minneapolis. 194pp. ~ Prepared by ~ ~$~P9&JA(~ T.D.I. Beck Wildlife Researcher C ��143 JOB PROGRESS State of Colorado Project No. W-144-R-l Work Plan No. 6 ------------------ Job No. Personnel: REPORT Big Game Investigations Mountain l-------.~-·Mourrtain-Eion Period Covered: July, 1982 - Noncervids Lion Investigations Populat-ion-Bynamics July 1, 1981 through June 30, 1982 A. E. Anderson ABSTRACT The puma literature synthesis was completed and submitted for review. Three female puma were captured and radio collared in 32 days of hunting; one of these was killed 5 days later. Only 30 of 50 attempts to locate 3 radiocollared puma with aerial telemetry were successful but 15 of the 20 failures were owing to one puma whose linear movements approximated 48 km and 2 of 3 locations were about 2 and 19 km north of the study areas, respectively. ��145 MOUNTAIN LION POPULATION DYNAMICS Allen E. Anderson P. N. OBJECTIVES 1. Investigate seasonal and diel activities, territoriality, population dynamics, habitat use, food habits and bioenergetics of an unhunted mountain lion population. 2. Remove designated numbers of mountain lions from specified sex and age classes and measure individual mountain lion and mountain lion population responses to removal. 3. Simultaneously monitor population dynamics of mountain lions and ungulate prey species to study the effects of mountain lion predation upon ungulate population dynamics. SEGMENT OBJECTIVES 1. Publish puma literature 2. Delineate 3. Capture and radio collar up to 20 puma. 4. Monitor puma and map location habitat review. units within study area. sites. ACKNOWLEDGEMENTS I am grateful for a diverse variety of support services provided by the following people: C. Albright, C. R. Anderson, D. Coven, R. B. Gill, M. Hershcopf, D. Masden, J. Olterman, G. Saville, L. Stevens, G. Tischbein, and D. Usner. Their help made the research enjoyable and successful. METHODS AND MATERIALS Literature Review The bulk of the literature search and statistical analyses for the the review of literature on puma was completed in 1981. Some additional sources were found in early 1982 and had to be incorporated in the ongoing writing begun in 1981. Proofing the voluminous tables and m8ps and preparing the final handwritten text occupied the balance of the segment. �146 Puma Capture and Telemetry Puma were located by searching secondary roads and jeep trails for recent puma tracks beginning at dawn and using two, 4-wheel drive trucks or two snowmobiles with occasional walking or snowshoeing. Daily searches averaged about 43 km with maximum distances about 140 km for each of the two trucks or snowmobile. Trucks were radioequipped with portable "walkie talkies" beginning late January, 1982. Apparently recent puma tracks were tested with trained hounds by the professional hunter and either abandoned as too old to track, or pursued with hounds until the dogs either abandoned the chase or treed the puma. In either case, the pursuit on foot often required several hours. Bayed puma were immobilized with powder-charged, 3 cc capacity syringes equipped with barbed darts, 2 cm in length, fired into the heavy muscles of the hindquarters or shoulder using the Extra Long Range (Powder) Projector with a .22 caliber adaptor using very low power (brown) loads.l The immobilizing drug, prepared by the Department of Pathology, Colorado State University, Fort Collins, was a combination of 100 mg of Rompun and 200 mg of Ketaset per mI. Our target dosage rate was 11 mg/kg of body weight as reported for puma by Hebert (1978:31). If necessary, dosage rates were adjusted by diluting the drug in the syringe dart with distilled water; dart accuracy, unimpressive at best, is lowered if the syringe is only partially filled. Immobilized puma were lowered from trees with rope. Tree climbers and a net to break a puma's fall were available if needed. A special salve was immediately placed in each eye and the eyes covered to prevent dessication of the cornea during immobilization. The puma was weighed, numbered with a tatoo inside each ear, and fitted with a radiocollar. Measurements of the body and dentition were taken as in Appendix 1. Respiratory and pulse rates and rectal temperature were recorded to the nearest 0.1 C. Since there is no accurate method of estimating age in puma, I used a combination of dental chronology and wear (Appendix 2), measurements of gum recession (Appendix 1) from Currier (1979), and a body weight for either sex of about 36 kg is indicated of puma older than 24 months (Robinette et al. 1961). Data were recorded on the form in Appendix 3. Actual hunting began on December 14, 1981 and terminated on February 21, 1982 for a total of 32 days of hunting. Hunting was terminated early because of illness in the family of Chuck Anderson, the professional hunter. Radiocollared puma were tracked from the air with the DOW Southwest Region Cessna 185 equipped with 2-element, strut-mounted, left and right antennae mounted with the long axes parallel to the fuselage. Reception from each antenna is controlled by a manual switch. Flights were made whenever other obligations permitted and the plane was available. Flight times ranged from about 30 to 180 minutes. lAll equipment from Palmer Chemical and Equipment Co., Douglassville, Ga. �147 RESULTS AND DISCUSSION Literature The manuscript was completed Review and submitted Puma Capture for review. and Telemetry Three female puma were captured and radiocollared. An additional puma was treed December 22, 1981, but not radiocollared because of a web of unfortunate circumstances which could have been avoided if 2-way radios had been available. Based on what I learned from puma researchers in the West during my tour of their projects during 1980, our rate of capture of about one puma per 7 days of hunting was about average. The details of the capture and body measurements along with similar information on puma number 1, captured April 16, 1981, are listed in Tables 1 and 2. The erratic dosage rate (Table 2) of the immobilizing drug was at least partly responsible for the highly variable pulse and respiratory rates and rectal temperatures between puma. The 11 mg/kg dosage rate cited for puma reported by Hebert et al. (1978:31) appears inadequate for wild female puma captured with hounds during winter. Rompum/Ketaset was selected as the immobilizing drug mainly because of its reported wide safety margin in dosage rate (Bebert et al. 1978). Given the circumstances and errors committed, a less well tolerated drug may have killed each immobilized puma. Aerial telemetry locations are summarized in Table 4. Only 30 of 50 attempts to locate puma telemetrically from the air were successful but 20 of the failures occurred with puma number 1 which apparently ranged between upper Roubideau Canyon north to the vicinity of Glade Park and Colorado National Monument area; a linear distance of at least 48 km and 19 km north of the GMU 62 study area. Another location was about 2 km north of the study area. This extensive movement is of particular interest because of her relatively limited movements from April through June 1980 (Anderson 1981). �148 Table 1. puma Details on capture and body and dentition measurements of 4 female 1 Item Date of capture Estimated age (months) Legal description of capture site ~S S T R Elevation (m) 4-16-81 21 NW 10 SON 13W Ear tatoo numbera 2 3 1-5-82 mature SE 1-8-82 26+ NE 4 1-21-82 24+ NW 4 4 4 48N llW 48N 11W 47N lOW 2,170 2,146 1,848 1,927 27.2 5 517 12.4 21 30 8389 149.5500 8772 149.7010 8775 149.8005 8773 149.7215 33.1 172.5 74.0 98.5 55.0 31.0 69.0 43.5 187.5 81.3 106.2 68.6 34.9 38.5 201.0 79.0 122.0 9.0 26.5 8.0 58.5 18.7 13.5 9.0 26.0 9.5 42.5 187.0 76.5 110.5 61.0 31.0 66.0 20.5 13.7 9.0 26.5 8.5 3.7 5.5 3.5 4.8 4.5 6.0 4.0 5.5 3.6 5.2 3.5 5.0 3.60 5.50 4.50 1.50 0.72 1.63 2.09 1.58 2.23 1.29 1. 73 1.17 1.29 1. 90 1.03 b Drugs Dosage (mg/kg) Induction time (min.) Immobilization time (min.) Radiocollar serial no. Transmitter frequency MHz Measurements (cm) Body wt (kg) Total body length Tail length Head-body length Chest girth Neck circumference Height at shoulder Head length Zygomatic breadth Ear length Hindfoot length Hind paw length Heel pads; max. diameter Left front: anterior-posterior transverse Left rear: anterior-posterior transverse Teeth Maxillary toothrow (canine-premolar) Upper 2nd premolar Crown length Crown width Lower 1st molar crown length Upper canine anterior-posterior diameter __d c 46.8 79 unknown 21.2 10 165 ------------------------------------------------------------------------------ �149 Table 1. Continued Ear tatoo numbera Item Gumline recession (mm) Upper canine Lower canine 2nd upper premolar 1st lower premolar 1 3 3 1.5 1.5 2 3 4 +2 +2 +1 +1 +1 +1 +1 +1 ~Tatooed inside both ears. Combination of xylazine hydrochloride (100 mg) and ketamine hydrochloride (200 mg) per ml. ~Gross approximation, see Table 2 for explanation. Insufficient drug dosage; puma got up and walked off before measurements completed. This animal was illegally killed by hunter on 1-10-82, radiocollar was returned on 2-8-82. �Table 2. Pulse, respiration rates, and rectal temperature in 3 drug immobilized female puma Pulse rate (beats Eer min.) Puma III Puma 113 Puma 114 Item 10 74.4 6.4 68 84 N Mean SD Min Max Time: Darted Interval First reading Last reading 2 82 1 ----- -80 84 Respiration rate (exhalations Eer min.) Puma III Puma 113 Puma 114 10 21.2 2.7 20 28 1 ----- 2 18.0 -- Rectal temE. (C) Puma III Puma 113 Puma 114 3 37.6 0.6 16 20 37.1 38.3 0923 10S2~171S 1444 1638 09S0 1037-l0S8 0923 10S2-1638 1444 1638 09S0 1037-l0S8 0923 1058-12SS 84 68 64 80 84 20 20 12 -- -- 20 16 38.3 37.1 ---- --- -- -- -- 3 39.4 .06 39.4 39.S 09S0 102S-1058 39.S 39.4 Drug dosage (mg per kg body weight):a Puma 1 3 4 27.2 46.8b 21.2 aCombination of 100 mg xylazine hydrochloride (Rompun) and 200 mg ketamine hydrochloride (Ketaset) per 100 m1. bCross approximation; syringe dart malfunctioned; 2 additional injections with improvised "jab stick" delivered only approximately known amounts to active, aggressive puma in cave. t-' VI o �151 Table 3. Mule deer killed by puma and found without Date Male 12-21 12-22 1- 8 1-22 a 1-24 a 1-24 1-25 2- 6 Female Adult Adult Adult Adult Fawn Adultb Adultb Adult Legal descriEtion R S T ~ NE SW NE SW SW NW NW 15 5 34 5 33 33 25 25 49N 48N 49N 48N 49N 49N 48N 51N llW llW llW llW llW llW llW 14W a systematic Habitat J-P J-P J-P J-P J-P J-P J-P J-P C search Body parts remaining Head, legs, ribcage Head, legs, trunk Almost intact Head, most of skeleton Almost intact Almost intact Head, most of skeleton Almost intact aFrom tracks in snow and telemetry: puma #3 killed deer on same site, dragged both about 11 m into shallow drainage (Piney Creek), carcasses were almost completely covered with snow when found and about 3 m apart. Puma #3 was babout 300 m from carcasses. Incisors obtained for age estimation with counts of dental cementum annuli (Erickson and Seliger 1969). cJ_P is juniper-pinyon woodland of Kuchler (1964). �152 Table 4. Date Summary of aerial telemetry locations for 3 puma ~S S TN RW Elev. (ft) Drainage Puma 1 (caEtured 4-16-81) 12- 4-81 12- 7-81 12-11-81 2-15-82 2-19-82 3-22-82 3-25-82 4- 9-82 4-19-82 4-20-82 5- 2-82 5- 6-82 5-28-82 6- 3-82 6- 3-82 6-12-82 6-18-82 6-28-82 Not Not Not Not Not SW Not Not Not Not Not Not NW Not Not Not located located located - faint signal located - faint signal located l3 50 l3 located, faint signal located located located located, faint signal located 20 14 100 located, faint signal located, Glade Park area located 12 15 103 7 15 102 Not located, poor signal Escalante Criswell 6,000 Cottonwood Roubideau Monitor 7,440 6,800-8,400 Snyder Creek Unaweep Unaweep Roubideau 18 attempts, 3 locations Puma 3 (caEtured 1-8-82) 1-11-82 1-15-82 1-24-82 1-26-82 2-15-82 2-19-82 3-22-82 4- 9-82 4-19-82 4-20-82 5- 2-82 5- 6-82 5-28-82 6- 3-82 6-12-82 6-18-82 6-25-82 6-28-82 located, faint signal 48 11 3 41 11 33 located 11 48 3 11 48 3 located located 48 11 3 11 48 2 11 48 NE 8 11 48 NE 4 11 48 SW 4 48 11 NE 28 Not located, poor signal 11 NE 49 33 11 NW 48 18 11 NW 48 18 Not NW SW Not NW NW Not Not NW 6,240 7,000 Dry Creek a Dry Creek b Piney Creek 6,240 6,240 Dry Creek Dry Creek 6,240 6,800 7,200 6,500 7,100 7,600 Dry Creek Dry Creek Dry Creek Dry Creek Dry Creek Dry Creek Dry Creek Piney Creek Dry Creek Dry Creek 6,500 7,600 7,600 18 attempts, 14 locations ------------------------------------------------------------------------------ �153 Table 4. Date Continued ~S S TN RW Elev. (ft) Drainage Puma 4 (captured 1-21-82) 1-26-82 2-15-82 2-19 82 3-25-82 4- 9-82 4-19-82 4-20-82 5- 2-82 5- 6-82 5-28-82 6- 3-82 6-12-82 6-18-82 6-25-82 6-28-82 Not located NW 10 NW 23 SW 27 NE 9 NW 15 34 NE 8 NE 5 SW 9 SW 19 SE 24 6 31 SW 34 NW 4 NW 34 48 48 48 48 48 48 48 48 48 47 48 48 49 49 48 49 11 10 10 10 11 11 11 11 11 9 12 12 12 12 12 12 7,200 6,600 6,700 7,300 7,200 8,000 7,200 6,500 7,300 7,440 7,600 7,800-8,100 Dry Creek Spring Creek Spring Creeka Spring Creek Dry Creek Dry Creek Dry Creek Dry Creek Dry Creek Dolores Dry Creek Roubideau 6,600 Roubideau 6,700 Roubideau 15 attempts, 14 locations ~ocation verified with ground telemetry and tracking. Located with ground telemetry only. cDivide by 3.2808 to obtain elevation in meters. �154 LITERATURE CITED Anderson, A. E. 1981. Mountain lion investigations--mountain lion population dynamics. Colo. Div. Wildl. Wildl. Res. Rep. July, Part 2:115-325. Ashman, D., and K. Greer. 1976. Age techniques. Pages 199-204 in G. C. Christensen and R. J. Fischer, co-chairmen Trans. mountain lion workshop. u.s. Fish and Wildl. serv., Portland, Oregon and Nevada Fish and Game Dep., Reno. 2l3pp. Currier, M. J. P. 1979. An age estimation technique and some normal blood values for mountain lions (Felis concolor). Ph.D. Thesis. Colorado State Univ., Fort Collins. 8lpp. Erickson, J. A., and W. G. Seliger. 1969. Efficient sectioning of incisors for estimating ages of mule deer. J. Wildl. Manage. 33:384-388. Hebert, D. M., A. R. Maltby, and M. F. A. Nation. 1978. Immobilization of members of the order Carnivora. Wildlife Disease Assn. Conf., S mposium 1. Chemical immobilization of wildlife. Colorado State Univ., Fort Collins. 43pp. + unpaged Appendix. Kuchler, A. W. 1964. Potential natural vegetation of the conterminous United States; map and manual to accompany the map. Amer. Geographical Soc. Spec. Publ. No. 36. New York. l16pp. Robinette, W. L., J. S. Gashwiler, and O. W. Morris. 1961. productivity and life history. J. Mammal. 42:204-217. Notes on cougar Slaughter, B. H., R. H. Pine, and N. E. Pine. 1974. Eruption of cheek teeth in Insectivora and Carnivora. J. Mammal. 55:115-125. Prepared by c~ c:. ?'In ~ Allen E. Anderson Wildlife Researcher C Sh't �155 APPENDIX PUMA MEASUREMENT 1 METHODS Body weight: Live weight, to the nearest kg using a hanging scale (Homs Laboratory Scale) calibrated for the weight of the net used to suspend the puma from the scale. Total body length: Most anterior point of nose, dorsally edge of the last coccygeal vertebra; measured with a along the spine to the nearest cm. The animal is in on a flat surface, with the head, spine, and tail in to the posterior flexible tape lateral recumbency, the same plane. From the articulation of the sacrum with the 1st coccygeal Tail length: vertebra, dorsally, to the posterior edge of the last coccygeal vertebra. Measured with a rigid tape, to the nearest cm. Body length: Length of the tail minus the total body length. Chest girth: Circumference of chest, just posterior to the axilliary border of the scapula with front legs perpendicular to vertebral column. Measured with a flexible tape to the nearest cm. The tape is held snugly without undue pressure. Neck circumference: Measured at the middle of the neck with a flexible steel tape to the nearest cm. The tape is held snugly without undue pressure. Height at shoulder: Vertebral border of the scapula, laterally, to the distal edge of the metacarpals. Measured with a rigid tape to the nearest cm. Animal is in lateral recumbency with front legs extended perpendicular to vertebral column. Head length: Most anterior point of nose, dorsally, to occipital tuberosity (junction of saggital and lambdoidal crests). Measured with a caliper rule to the nearest mm. Zygomatic breadth: Maximum distance between zygomatic Measured with a caliper rule to the nearest mm. Ears: arches across cranium. Base of the incisura, intertragica to the tip of the auricle the hair, measured with a transparent rule to the nearest mm. Hind foot: excluding (2 measurements) Tubercalcanei, along the posterior edge of the hind foot; 1. To the tip of the longest claw; 2. To the distal edge of the metacarpals (not including the paw). Hind paw length: Length of digits from distal edge of metatarsals, ventrally, to tip of longest claw. Measured with a steel tape to the nearest mm. �156 APPENDIX Heel pad: (front Measured 1 (Continued) & rear) length: greatest antero-posterior diameter wi.dth: greatest transverse diameter with a transparent rule to the nearest mm. Maxillary toothrow: From anterior edge of canine to posterior edge of 2nd premolar at alveolar -na r gfn . Measured with a c a Lf.p ez rule to nearest 0.1 mm. Upper carnassial crown length: Antero-posterior diameter of crown, of 2nd premolar at cingulum. Measured with caliper rule to nearest 0.1 mm. Upper carnassial crown width: Transverse diameter of crown of 2nd premolar at widest point anteriorly at cingulum. Measured with a caliper rule to the nearest 0.1 mm. Lower carnassial crown length: Antero-posterior diameter of crown of 1st molar at cingulum. Measured with a caliper rule to the nearest 0.1 mm. Upper canine: Antero-posterior diameter at alveolar margin. with a caliper rule to the nearest 0.1 mm. Measured ,, l Distance from gumline to cingulum on lateral (buccal) Gumline recession:a side of tooth. If the gumline extends over the cingulum the measurement is recorded as positive. If the gumline had receded beyOnd the cementoenamel junction, the measurement is recorded as negative. Measured with a periodontal probe graduated in mm lined up parallel to tooth axis on the following teeth: Upper canine; Lower canine; 2nd upper premolar; 1st lower premolar aFrom Currier (1979). �157 APPENDIX CHRONOLOGY 2 OF TOOTH REPLACEMENT AND WEAR IN PUMA Type of tooth and age at eruption: Temporary Days DaE 1st incisor, upper 1st incisor, lower 2nd incisor, upper 2nd incisor, lower 3rd incisor, llPper 3rd incisor, lower canines, upper canines, lower 1st premolar, upper 2nd premolar, upper 2nd .premolar, lower 3rd premolar, upper· 3rd premolar, lower Permanent 13 15 13 15 13 15 18-20 20-22 18-22 20-21 30-34 27 30-34 30-34 52-53 45 45 48-52 45 Seguence (Slaughter 1974) 1st incisor, upper 1st incisor, lower 2nd incisor, upper 2nd incisor, lower 3rd incisor, upper 3rd incisor, lower canines, upper canines, lower 2nd premolar, upper 3rd premolar, upper 3rd premolar, lower 4th premolar, upper 4th premolar, lower 1st moLar , upper 1st mo l.ar , lower Days 17 17 17 17 17 17 23 23 33 33 30 33 30 Month 5.5 5.5 5.5 5.5 5.5 5.5 8.5a 8 Month 8 1 3 3 2 2 1 1 Canines: eruption, relative wear and staining (Ashman and Greer 1976) About 1/3 erupted . 8 12-13 About 2/3 erupted, enamel-cemention junction absent. 12-24 Fully erupted, enamel-cementum junction absent, upper canine longer Fully erupted, enA.l)1~J-cementumjunction I:'L'e:'ent and 24-30 greater in upper canines, than. in lower, no staining, little or no wear on tips. Stai!1ed light yellow, tips beginning to show wear 36-48 Stained yellow excep!: lower 1/4, tips show definite 49-60 wear with cupping. Stained deep yellow, tips well worn and cupping 61-72 or hooked appearance especially on the inside upper canine. aBoth temporary and permanent for a brief period. �158 APPENDIX 3 PUMA STUDY W-144-R, WP6 ------ ) Puma (Tatto No. Estimated of capture site: ____ Radiocollar Elevation Weather S , T _ R ----- Frequency ------ ft. of capture site ETM --- MHz m. (incl. temp., wind) From to -------------- ---tracks first seen Locatfon Time: \, Serial No. Time in field: Time: Observers Date ----- Age (months) Location Route Sex ------------------ \, S---- of tracks first seen dogs put on tracks (puma treed Estimated puma shot shooter to puma: Behavior before admin. of drug: Behavior after admin. of drug: Drug and ratio: Rompun/Ketamine Atropine/Ketamine Dosage , R =---- , Elev. --------------------- ----- distance: , T--- ------- ft. ____ m. ----_/_------------_/_------- mg/kg Volume drawn Volume administered Injection time Injection site -------------Remarks Start of ataxia Immobilization time --------------- Remarks Pulse rate (beats per minute) Resp. rat e (exhal. per minute) o Rectal temp. ( C) Time of recovery Markings, (animal standing without assistance): pelage Abnormalities ----------------------------------------------------------------- _ �159 APPENDIX Time - Blood Aspiration: Venipuncture site 3 (Continued) Start Finish ------ Total Time ___ min. -------------------------------------------------------------EDTA ---------- cc, Serum ---------- cc Amount of blood aspirated: Comments: Time - Measurements: Body weight Start ------ Finish ----- Total Time min. ----- (kg) Total body length Tail length (cm) Body length (cm) (cm) Chest girth (cm) Neck circumference (cm) Height at shoulder (cm) Head length Zygomatic (cm) breadth (cm) Ears (cm) Hind foot (cm) Hind paw length (cm) Heel pads; maximum diameters (mm): Left front, antero-posterior ----------- Left rear, antero-posterior Maxillary toothrow crown length Upper second premolar crown width Lower first molar crown length Upper canine anterior-posterior transverse ----------- (rnm) (mm) (mm) diameter (mm) ------ (rnm)a: Lower canine + + 2nd upper premolar + 1st lower premolar + Upper canine _ (canine-premolar) Upper second premolar Gumline Recession transverse --------------aIf the gumline extends over the cingulum, the measurement is positive. If the gumline had receded beyond the cingulum, the measurement is negative. � https://cpw.cvlcollections.org/files/original/b20b4a523cecabab6ae7db3b0b010a47.pdf 0fa0df62c9c45332b83b9288eb0da58f PDF Text Text October 1982 JOB PROGRESS REPORT State of Colorado ----~~~~~------------ Project No. Work Plan No. Job Title: Job No. Waterfowl Production Surveys Period Covered: Personnel: Migratory Bird Investigations w-88-R-27 11 May 1981 to 15 June 1982 Those cooperating in the 1981 survey included: M. Nail and staff, Monte Vista National Wildlife Refuge; Jim Creasy and Jim Sellers, Brown's Park National Wildlife Refuge; M. Bauman, J. Corey, J. Dennis, M. DePray, J. Frothingham, J. Gray, J. Haskins, D. Kenvin, J. Lorentzson, D. Masden, T. Rauch, W. Russell, S. Steinert, M. Szymczak, B. Will, Colorado Division of Wildlife. ABSTRACT Water conditions for duck and goose production throughout the state were good in all areas except the San Luis Valley. Surface water continues to decline as more circular sprinklers are installed for irrigation. The total estimated number of breeding pairs increased to 94,199 or 70.3% above the 1980 level and 60.8% above the long-term average. The mallard continues to be the major breeding species. The loss of water on the air-ground transects continues to be a problem in the San Luis Valley. This results in an inflated figure for the Blue-winged and Cinnamon Teal, Redheads, and other divers~ The number of nesting pairs of Canada geese were down 36% from the 1980 count and non-nesting pairs were down 30% from 1980 in the San Luis Valley. The numbers of Canada geese nesting in westcentral Colorado is slowly increasing. The Yampa and Little Snake River populations of Canada geese remain about the same as in 1980. The number of adult Canada geese in the northcentral production area showed an increase of 19.7% from 1980. Gosl ing production was down in some northcentral areas but was still 2.7% above the 1980 count. ��3 WATERFOWL PRODUCTION SURVEYS Gerald Lorentzson Steven Steinert P. N. OBJECTIVES 1. To estimate the number of duck breeding pairs, by species, on selected major waterfowl nesting areas in Colorado. 2. To estimate the number of goose breeding pairs, and in some cases, obtain production data on selected goose nesting areas in Colorado. 3. Compile data and submit reports to appropriate state personnel and the Fish and Wildlife Service for use in monitoring status of the various species and establishing hunting season recommendations. SEGMENT OBJECTIVES 1. To estimate the number of duck breeding pairs, by species, in the San Luis, Cache la Poudre, South Platte and Yampa Valleys, and in North Park and Brown's Park, using procedures presented in the Program Narrative. 2. To estimate the number of goose breeding pairs in the San Luis Valley, on the Yampa, Little Snake and Green rivers in northwest Colorado and in northcentral and westcentral Colorado using procedures presented in the Program Narrative. 3. Evaluate the present boundaries of the area sampled for duck breeding pairs in the San Luis Valley. 4. Alter the duck breeding pair survey in the Cache la Poudre and South Platte areas to compensate for sample sections which can no longer be counted because of changes in land use patterns. METHODS AND MATERIALS The 1981 duck breeding pair surveys were conducted during the period of May 11 to June 15. Surveys in North Park, the Cache la Poudre and South Platte Valleys were conducted exclusively from the air. Brown's Park and counts in the Yampa River Valley were made on the ground. In the San Luis Valley and North Park, aerial counts were adjusted for visib1lity by air-ground comparison studies. Pair estimates for the Monte Vista National Wildlife Refuge in the San Luis Valley were obtained from nesting transects. All survey methods remained the same as in previous years. Six sections were changed in the South Platte River drainage because housing developments and turkey farms made it impossible to fly the existing areas. Three new sections and one river section were changed in the Poudre River Valley to replace those which had to be abandoned because of housing developments. �4 Canada goose surveys were conducted during the period of April 22 through June 16. Estimates of the Colorado, Gunnison, White River, Yampa and Little Snake River populations were obtained by direct counts from a fixed wing airplane. Brown's Park continues to be a ground count conducted by the Brown's Park NWR personnel. Ground counts were not done this year on the Yampa River, Little Snake River or in the Dinosaur National Monument. In the San Luis Vallty a survey of expected breeding pairs was conducted by flying transects in defined sample blocks located in Canada goose production areas throughout the Valley. Population estimates in the northcentral area of Colorado were obtained by direct ground counts of adults and goslings during the period when the birds were fl ightless. All flying was done with Cessna 185 aircraft. Two observers were used when sampling transects, while one observer was used for sampling sections. RESULTS AND DISCUSSION Water conditions for duck production were good in all sections of the state with the exception of the San Luis Valley. Surface water in the Valley continues to decline as more irrigation systems are installed. Most lakes and reservoirs in the state went into the spring at near capacity levels. Spring rains created excellent nesting conditions in most of the state. Low snowpack resulted in a small runoff and little or no flooding. This should have provided good nesting habitat along river systems throughout Colorado. A warm winter and an early spring insured that birds entered the breeding season in excellent condition. The total estimated number of duck breeding pairs increased to 94,199, 70.3% above the 1980 level and 60.8% above the long-term average (Table 1). Mallard breeding population increased from 1980 in total numbers but decreased when compared in percentage of total breeding population. The estimated breeding pairs by species are given in Table 2. The loss of water on the air-ground transects continues to be a problem in the San Luis Valley. As a result, the projected numbers of Blue-winged and Cinnamon teal, Redheads, and other divers are inflated in both the San Luis Valley and North Park (Table 3). The estimated number of Canada geese on the Yampa and Little Snake Rivers remain about the same as last year. This is the second year that the aerial survey has been done, so comparative data is limited. The number of geese observed are recorded by river section in Table 4. The population in Brown's Park appears to be about the same as in 1980 but had an increase in the estimated number of goslings (Table 5). The aerial breeding pair survey of westcentral Colorado was flown on April 23 and 24, 1981 (Table 6). Comparing 1981 results with prior years indicates that the population is increasing slowly in westcentral Colorado (Table 7). �5 Table 1. Summary of Colorado's in selected areas, 1981. duck breeding pair population Total estimated Area 1981 breeding pa irs 1980 Long term average San Luis Valley 26,338 21,571 North Park 40,599 . 13,644 South Platte Valley 14,838 8,318 Cache la Poudre Valley 8,377 9,418 Yampa Valley 1,486 2,572 Brown's Park 1,475 889 Totals 94, 199 Table 2. Species composition population. Species 55,326 of Colorado's estimates Present change From long term average 1980 26,894 16,250 7,452 4,307 2,606 1,110 + 22.1 +197.6 + 78.4 - 12.4 + 73 + 66 2.0 +150 + 99 + 94 1.3 + 33 58,599 + 70.3 + 60.75 1982 duck breeding pa ir Number of breeding pairs 1954-80 average 1981 1980 Percent species composition 1954-80 average 1981 1980 Ma 11 a rd Blue-winged and cinnamon teal Gadwa 11 Pinta i1 Shoveler Green-winged teal Redhead American wigeon Other divers Mergansers 18, 186 15,980 26,664 19.3 28.9 49.0 16,272 12,204 3,556 4,152 3,572 16,243 3,923 15,909 182 7,485 8,805 2,904 6,251 3,501 9, 120 146 1,134 5,988 5,758 3,566 3,619 3,292 2,791 1,097 1,664 17.3 13.0 3.8 4.4 3.8 17.2 4.2 17.0 13.5 15.9 5.2 11.3 6.3 16.5 0.3 2.0 11.0 10.6 6.5 6.6 6.0 5. 1 2.0 3.0 Tota 1s 94, 199 55,326 54,442 �6 Table 3. 1981. Estimated population of duck breeding pairs in Colorado, San Luis Va 11 ey Mallard Blue-winged and cinnamon teal Gadwa 11 Pintail Shoveler Green-winsed teal Redhead Wigeon Other divers Mergansers S.Platte River Yampa Va 11 ey Browns Park N.Park Poudre River 5,192 2,744 3,498 5,339 1,100 313 18, 186 6,682 3, 187 526 542 999 9,136 74 5,589 6,422 1,756 333 1,511 5,333 2,022 14,889 1,618 989 243 877 302 333 177 385 35 1,627 1,315 788 2,264 226 1,252 1,583 422 22 481 151 124 41 385 69 28 69 124 275 220 119 95 149 120 39 144 1 16,272 12,20l1 3,556 4, 152 3,572 16,243 3,923 15,909 182 Total 94, 199 Tota 1 Table 4. Number of Canada geese observed on aerial surveys in Moffat and Routt counties. Nesting ea irs 1981 Area 1980 Non-nesting eairs 1981 1980 Total adults 1981 1980 Yamea River Steamboat Springs-Craig Craig-Juniper Springs Juniper Canyon Juniper Springs-Cross Mtn. Lily Park Totals 7 15 4 27 6 __ 1 15 31 6 59 12 16 5 25 22 80 16 30 22 38 65 18 126 94 341 65 144 94 Li tt 1 e Snake River Cross Mtn.-Powderwash Powderwash-Baggs Br. Totals 1Totals were not censused 21 8 29 21 30 16 46 in previous years. 30 190 142 48 142 �7 Table 5. Number of Canada geese observed on ground counts in Brown's Park NWR. Area Table 6. Total Adults Estimated No. of gos Iin_gs1 1981 1981 1980 1981 1980 277 264 245 170 72 Brown's Park lCalculated Nesting Pairs 1980 60 using number of successful nests x average brood. West central Colorado Canada goose breeding pair survey, 1981. Area Singles White River Meeker-Rio Blanco Lake Rio Blanco Lake-Rangely Rangely-Utah Line Subtotal Colorado River Newcastle-Silt Silt-Rifle Rifle-Parachute Parachute-DeBeque DeBeque-Palisade Palisade-Grand Avenue Bridge Grand Avenue Bridge-Fruita Fruita-Horsethief Canyon Horsethief Canyon-Utah Line Subtota I Roaring Fork River El Jebel-Carbondale Subtotal Gunnison River Hotchkiss-Delta Delta-Mesa Co. Line Mesa Co. L ine-Wh itewa te r Whitewater-Grand Junction Subtotal GRAND TOTAL Pai rs Groups 11 14 23 49 2. 11 33 53 14 .39 74 100 3 6 9 53 130 19 7 18 13 13 5 6 4 1 4 4 40 2 4 2 11 34 20 20 59 1 _3 119 302 ~ 10 11 1 2 7 10 o 4 8 29 o 1 1 o 6 14 32 87 214 444 5 1 o 3 �Table 7. Comparison of the number of Canada goose singles and pairs observed west-central Colorado, spring 1977-81. on aerial surveys in Number observed Area White River Roaring Fork River Colorado 1977 1981 1980 Pairs 1979 1981 1980 Singles 1979 39 30 46 31 15 74 39 50 . 26 32 2 6 3 2 2 7 1 7 0 6 31 47 41 30 13 105 74 77 35 33 9 6 4 4 2 14 11 12 4 7 0 2 0 0 0 4 5 1 0 0 6 3 0 4 0 10 8 5 5 11 87 94 94 71 32 214 138 153 70 89 1978 1978 1977 River Glenwood Springs-5th St.Bridge Bridge - Utah Line 00 Gunnison River Hotchkiss - Delta Delta - Grand Junction TOTAL �9 The estimated number of nesting pairs of Canada geese in the San Luis Valley is down 36% from 1980 (Table 8) and non-nesting pairs were down 30% from 1980. The Canada goose production survey was done on June 15 and 16, 1981 in northcentral Colorado and the results are presented in Table 9. The number of adults has increased 19.7% from the 1980 level and 26.9% from the 1970-80 average (Table 10). Gosling production was down in the Wellington, Fort Collins and Boulder areas but was still 2.7% above 1980 (Table 11). Table 8. survey. Results of San Luis Valley Canada goose breeding pair Year Projected total number Nesting pairs Non-nesting 1975 He Iicopter 59 59 1976 He Iicopter Fixed-wing 92 77 63 69 1977 --He Iicopter Fixed-wing 100 100 101 91 99 96 1980 Fixed-wing 141 226 1981 Fixed-wing 90 159 1978 No counts 1979 --Fixed-wing pairs �Table 9. Results Production area Well ington of northcentral Colorado Water area Terry Lake Deines Reservoir Launer Pond Douglas Lake Stewart Pond Dry Creek Reservoir North Poudre #3 North Poudre 115 Bureau of Standards Reservoir 118 Elder Reservoir Van Sant Pond Cobb Lake Hinkley ReservoIr Wa tson Lake CurtIs Lake Beghtol Lake goose census, No. broods -1 -6 2 -- ---1 2 ------- 1 Subtota I Fort CoIl ins Dixon Reservoir Miller Ponds College Lake Dean Acres Andrijeski Harsh Claymore Lake Sterling r.ravel Pits Larimer County Shop Lindenmeier Lake Grey Lakes Novak's Flatiron Anderson's Parkwood Kitchel Lake Timnath Reservoir Romi Iy Gravel Pits Fossil Creek Schue Ike Lake Horseshoe Lake Wolaver Ponds Subtotal 12 7 4 4 3 6 12 I 7 2 2 10 1 8 12 8 5 5 2 ___! June 15 and 16, 1981. Total no. 20S Iin2s 54 6 23 30 8 25 17 0 4 50 0 0 30 24 35 0 2 308 39 22 14 If, 12 19 "9 2 0 21 6 9 33 5 32 52 32 19 20 6 11 424 Total no. adults and yearlings 92 2 33 45 4 12 18 11 2 135 35 1 31<1 18 101 29 2 864 52 42 229 27 6 89 226 6 57 32 5 7 74 12 45 25 41 10 10 4 (- 1,005 Total birds 146 8 5(, 75 12 42 35 11 6 185 35 1 34? 42 136 29 4 1,174 91 64 243 43 18 108 275 8 57 53 11 16 112 17 77 77 73 29 30 10 _JJ_ 1,42<1 0 �Table 9. Production area Loveland (cont inued) Water area Flatiron Reservoir Boedecker Flatiron Gravel Pits Kauffman Gravel Pits McNeil Reservoir Welch Reservoir Reservoir #12 No. broods 1 6 3 4 5 3 4 2 66 14 20 52 38 20 212 7 10 3 6 9 15 32 0 14 16 9 72 32 0 175 If 1 10 27 36 63 3 12 Subtotal Boulder Ish Lake C rys ta I Lake Terry Lake Faivre Ponds Rest Home Pond Valmont Sawhill & Walden Eddy Pond -- 3 4 3 Ponds Subtotal Denver Ketr in9 Centennial Columbine Country Club Chatfield Golf Course -Bowles Lake Kings Pond Tule Lakes Grant Ponds Marston Lake Pinehurst Country Club Clarefield Reservoir Kendrick Lake Federal Center Pond Sloan's Lake Stanley Lake Denver City Park Colo. Blvd. @ QuIncy Blackmer Reservoir Subtotal GRAND TOTAL Total no. ~os Iin~s ---- -- -- 2 2 10 1 2 15 6 5 10 15 45 1 12 71 25 25 4 8 401 1,520 Total .no. adults and yearl ings 10 107 25 36 114 58 108 458 26 10 100 42 31 138 127 2 476 98 72 53 74 235 156 72 118 39 24 35 46 PI 215 36 230 2 __ 1_0 Total birds 12 173 39 56 166 96 128 670 58 10 1PI 58 40 210 159 2 651 139 82 80 110 298 156 84 118 49 3'3 80 47 26 286 61 255 2 18 1,529 1,930 4,332 5,852 �12 Table 10. production Number of adult Canada geese observed trend areas, 1981. in northcentral Percent change From 1980 1970-80 No. of adults Area Average 1970-80 1981 1980 Wellington Fort Co llins Loveland Boulder Denver 864 1,006 458 476 1,529 773 970 408 407 1,061 722 701 249 535 1,206 + + + + + Totals 4,332 3,619 3,413 + 19.7 Table 11. Number of Canada goose goslings produced Co lorado product ion trend areas, 1981. Area 1981 1980 308 424 212 175 401 340 467 126 220 326 259 308 107 209 284 1,520 1,479 1, 167 Well ington Fort Co lIins Loveland Boulder Denver Totals P repare d b y: J". ./ ..!J£t<.dd '--V. I' ).") ,n/'.~ _::,-",= dIY 461</ Gerald M. Lorentzs . (j.8) Senior Wildlife Biologist Prepared by: d4~:w"_ J Jfl.::t'''k{ Steven F. Steinert (i&) Wildlife Technician III 11.8 3.6 12.3 16.9 44.1 + + + 19.7 43.4 83.~ 11.0 26.7 + 26.9 in northcentral No. of goslin~s Average 1970-80 Colorado Percent change From 1980 1970-80 9.5 9.2 + 68.25 - 20.5 + 23.0 + + + + + + 30.24 2.7 16.0 27.4 98.1 16.3 41.2 �October 1982 13 JOB PROGRESS REPORT State of Colorado ----~~~~~---------- Project No. Migratory W-88-R-27 ---------------------- Work Plan No. ------------------Ecological Job Title: Bird Investigations Job No. 1_4 Studies of the Flightless _ Period of Ducks in Colorado Period Covered: Personnel: 1 April 1981 - 31 March 1982 J. K. Ringelman and M. R. Szymczak, Wildlife. Colorado Division of ABSTRACT A program narrative outlining a 6-year study of flightless prepared, reviewed and approved. adult ducks was ��15 ECOLOGICAL STUDIES OF THE FLIGHTLESS PERIOD OF DUCKS IN COLORADO Michael R. Szymczak James K. Ringelman P. N. OBJECTIVE 1. Prepare a Program Narrative and Segment Narrative for Segment 28 of Project w-88-R describing, in detail, specific studies, their objectives, procedures, schedules and costs. SEGMENT OBJECTIVES 1. Review Ilterature deal ing with all aspects of molt, molt migration and molting habitat in waterfowl. 2. Develop an outline of study objectives. 3. Prepare a draft Program Narrative selected scientists. 4. Prepare a final study plan. to be reviewed by DOW and other METHODS Both published and unpublished literature was reviewed, other researchers were consulted and the program narrative prepared based on the background information obtained. The PN was submitted for review, revised and submitted for approval by DOW and U.S. Fish and Wildlife Service federal aid administrators. RESULTS AND DISCUSSION The Program Narrative and Segment Narrative was approved by DOW and U.S. Fish and Wildlife Service federal aid administrators. The objectives of the study are as follows: 1. Document the species and sex composition and seasonal abundance of ducks molting on selected wetlands in North Park. 2. Identify the physical and biological characteristics by molting ducks. 3. Investigate spatial and temporal differences molting ducks. of wetlands in use of wetlands·by used �16 4. Determine the behavioral molting ducks. time budgets and net energy balance of 5. Investigate differences in duration of the flightless period of selected species of ducks in relation to sex, body condition, habitat quality, and time of molt. 6. Determine the survival rate of molting ducks in relation to sex, body condition, habitat qual ity, and time of molt. 7. Identify and quantify duck molting wetlands in Colorado. The length of the study will be 6 years with objectives and accompanying approaches 1, 2 and 3 active the first year; 1, 3, 4 and 5 the s~cond year; 4 and 5 the third year; 6 the fourth year and 7 the fifth and sixth years. The project will demand about 260 man-days per year for the first 4 years, 80 man-days for the fifth year and 100 man-days for the sixth year. Field work will take place in North Park the first 4 years and expanded to statewide in the final 2 years. �October 1982 17 JOB PROGRESS REPORT State of Colorado ----~~~~~----------W-88-R-27 Project No. Work Plan No. Job Title: 2 Job No. 9 Monitor Banding of the Shortgrass Prairie Canada Goose Population Pefiod Covered: Personnel: Migratory Bird Investigations in Southeastern Colorado January and February 1982 J~nnifer Slater, B. McCloskey, Gerald Lorentzson and others with the Colorado Division of Wildlife. ABSTRACT Because of the shortage of water in certain areas and the denial of landowners to grant permission to trap geese on private land, no birds were banded in 1982. The geese which were present, did not respond to bait because of the open winter and the availability of other foodstuffs. Efforts will be made to trap 1,000 Canada geese in 1983. ��October 1982 19 JOB PROGRESS REPORT State of Colorado --------------------------Migratory W-88-R-27 Project No. Work Plan No. 2 Distributional Job Title: 10 Job No. Characteristics Bird Investigations of Some Populations of Canada Geese Inhabiting Colorado Period Covered: Personnel: 17 June 1981 through 17 February 1982 M. Bauman, L. Budde, H. Burdick, G. Byrne, J. Corey, D. Crawford, M. Creamer, L. Crooks, T. Davis, R. Desilet, K. Dillinger, J. Ellenberger, M. Etl, J. Frothingham, M. Gardner, J. Gray, J. Gumber, D. Homan, R. Kahn, H. Lanning, J. Leslie, G. Lorentzson, R. Moss, S. Porter, C. Reichert, D. Schaefer, S. Steinert, J. Wagner, K. Wagner, P. Will and M. Szymczak, Colorado Division of Wildlife; G. Patten and staff, Arapaho National Wildlife Refuge; and C. Cesar, Bureau of Land Management. ABSTRACT The number of Canada geese (Branta canadensis) banded on production/brood rearing areas exceeded quotas in North Park (363), South Park (220) and northeast Colorado (232). Only 29 geese were banded in west-central Colorado and no birds were captured in northwest Colorado. Analysis of the banding locatjon~ of birds recovered in west-central Colorado since 1968 indicated production areas in Wyoming, particularly those at Ocean Lake in Fremont County and Yellowtail Reservoir on the Big Horn River, were producing birds that have been consistently recovered in west-central Colorado. Most banded birds recovered in west-central Colorado were originally captured and banded on the Wheatland Reservoir molting area in southeast Wyoming •. Trapping efforts post-season in northeast Colorado resulted in 74 "La rqe" Canada geese being banded. Analysis of the banding locations of birds recovered in northeast Colorado since 1972 indicated the birds originated primarily from scattered locations in Alberta, Saskatchewan, North Dakota, South Dakota and Wyoming. �20 RECOMMENDATIONS 1. In view of the poor trapping success encountered during post-season banding efforts in both west-central and northeast Colorado in 1982, it is recommended that the trapping effort begin if possible during the latter part of the goose season in both areas. �21 DISTRIBUTIONAL CHARACTERISTICS OF SOME POPULATIONS OF CANADA GEESE INHABITING COLORADO Michael R. Szymczak Gerald Lorentzson P. N. OBJECTIVES 1. To document the wintering range and harvest distribution of Canada geese nesting in (1) northwest Colorado, (2) west central Colorado, (3) North Park, (4) northeast Colorado and (5) South Park. 2. To ascertain the breeding range of Canada geese wintering central and northeast Colorado. in west 3. To contribute data to the various Central and Pacific Flyway management plans for specific populations of Canada geese. SEGMENT OBJECTIVES la. Trap and band at least 150 Canada geese on production areas in west central Colorado. lb. Trap and band at least 150 Canada geese on production areas in northwest Colorado. lc. Trap and band at least 100 Canada geese on production areas in North Pa rk. ld. Trap and band at least 150 Canada geese on production areas in northeast Colorado. 2. Trap and band 250 Canada geese on wintering areas in west central Colorado. 3. Trap and band at least 250 Canada geese on wintering areas in northwest Colorado and take the measurements meritioned in the Program Narrative. 4. Submit banding schedules and recovery reports to the Bird Banding Laboratory. 5. Plot the recovery location of Canada geese reported that have been recovered or recaptured in west central Colorado or northeast Colorado. 6. Prepare progress report. �22 METHODS AND MATERIALS Canada geese were trapped on production/brood rearing areas in west-central Colorado, northeast Colorado, North Park and South Park during their flightless period utilizing standard drive trapping methods and equipment (Szymczak et al. 1981). All birds captured were banded and released. Winter trapping in west-central and northeast Colorado was accomplished using cannon-nets. Only those birds captured that were judged to be "large" Canada geese were banded in northeast Colorado. The following physical measurements were taken for each bird banded in northeast Colorado: weight, wing chord, tarsus, culmen, bill width (base, posterior nares, posterior nail) and nail length. Annual reports from the U.S. Fish and Wildlife Service's Bird Banding Laboratory containing all birds banded anywhere and reported encountered in Colorado during that specific annual period were examined for the occurrence of banded large Canada geese reported encountered in westcentral and northeast Colorado. In west-central Colorado, only those birds reported taken from 1968 through 1980 in the geographic area covered by latitude 3800 through 39So and longitude 10700 through 108So were recorded. In northeast Colorado, recorded recoveries occurred from 1972 through 1980 in the area from 400 through 40S latitude and 1020 through 1034 longitude. RESULTS AND DISCUSSION Summer Populations Banding quotas were met or exceeded in 3 of the S areas in which summer banding was proposed. In northwest Colorado on the Yampa and White Rivers, concentration of productive adults and accompanying goslings could not be located and therefore no trapping operations were attempted. In west-central Colorado, over lS0 goslings and adults were located but trapping was basically unsuccessful because it was too late in the brood rearing period for drive trapping in that area and most were able to escape the trapping attempts by flying. Banding quotas were exceeded in northeast Colorado, North Park and South Park (Table 1). Winter Population - West central Colorado Trapping and banding Attempts to attract geese to baited cannon-net sites were unsuccessful so no birds were captured during trapping and banding operations postseason in west-central Colorado. �23 Table 1. The number of Canada geese banded on production/brood areas in Colorado, summer 1981. Area/location Adult male rearing Number banded Local Local Adult male female female Total West-central Colorado Colorado R.- 3 2 11 13 29 North Park Walden Res. Pole Mountain Res. 106 8 106 9 59 10 59 6 330 33 Northeast Colorado Guenzi Property Red Lion Wi Id I. Area Johnsons Pond 7 1 21 6 1 21 25 13 44 25 10 58 63 25 144 South Park Antero Res. Eleven Mi Ie Res. 35 10 45 12 51 10 49 8 180 40 Foreign Recoveries Analysis of the location of banding of Canada geese recovered or recaptured in west central Colorado indicates that most banded birds taken had been banded in Wyoming (Figure 1). Extensive banding in southern Alberta and on marshes associated with Great Salt Lake in Utah during the 1970's produced very few recoveries in west-central Colorado. Banding in Montana and eastern Idaho has not been prolific in recent years, yet the paucity of recoveries from production areas in those states indicate they do not produce a substantial number of geese which winter in Colorado. Wyoming banding since 1970 has been consistent although not well distributed through breeding areas. Birds banded at Wheatland Reservoir, which is a major molting area for sub-adults of many segments of the Rocky Mountain Population (Krohn and Bizeau 1979) were well represented in the harvest (Table 2). The other two areas, Ocean Lake and Yellowtail Reservoir, represented by a substantial number of recoveries are both breeding areas. �" ,I I .'0' ,(.) ClOClOO( Ii;·: I ; : I N .J:- 'I" . I r ~ .. 0000 .@: OO(i , •. i ., :/T: " I, :1; . ' , �25 Table 2. Banding areas outside Colorado of Canada geese reported recovered during the hunting season or recaptured during post-season banding operations in west-central Colorado, 1970 through 1980. Number of recoveries Direct Indirect Area Alberta Camrose Area (restoration) Hays Reservoir Scope Reservoir Bantry Reservoir Louisiana Lakes Area Milk River Ridge Reservoir Grassy Lake 2 2 2 3 Sa skatchewa n Cypress Hills Montana Canyon Ferry Fort Peck Reservoir (molting) Wyoming Turbid Lake (molting) Yellowtail Reservoir Ocean Lake Eden Va 11ey Pickett Lake (molting?) Sheridan Co. (Tongue R.) Crook Co. (restoration) Natrona Co. (Platte River) Wheatland Reservoir (molting) Utah Rich County Salt Lake Area Sanpete Co. (restoration?) 3 8 19 11 1 35 3 1 1 1 2 3 31 69 2 4 1 3 Birds have been banded annually at Ocean Lake in Fremont County near Riverton in the Wind River drainage for a substantial number of years and recoveries of locals in west-central Colorado has been consistent over time (Table 3). Recent bandings on production areas near Yellowtail Reservoir on the Big Horn River in north central Wyoming has also produced encounters in Colorado. �26 Table 3. Banding areas outside Colorado of Canada geese reported recovered during the hunting season or recaptured during post-season banding operations in west central Colorado 1970-80 that were classified as wild-trapped locals when banded. Year of recovery Before 1973 1973 1974 1975 1976 1977 1978 1979 1980 Area Alberta Cam rose Area Hays Reservoir Scope Reservoir Bantry Reservoir Louisiana Lakes Area Milk River Ridge Res. Grassy Lake 2 Saskatchewan Cypress Hills Wyomin9 Ocean Lake Ye llowta iI Reservoir 4 3 2 2 3 4 4 3 9 4 5 13 Utah Rich County Salt Lake Area Sanpete Co. Winter Population - Northeast Colorado Trapping and Banding Canada geese were trapped at Prewitt Reservoir located about 15 miles southwest of Sterling and Johnson's Pond located about 10 miles southwest of Julesburg. One cannon-net catch was made at each location. On 28 January 1982 at Prewitt Reservoir, a total of 117 birds were captured of which 111 had not been previously banded. Sixty (51.3%) of the 117 birds were subjectively judged to be large Canada IS. Of the 111 unbanded birds, 55 were judged t6 be large, measured, banded and released, while 56 were judged to be small and released unbanded. Five of the 6 recaptures were considered large geese. Four of the 5 large birds recaptured wore color-marked leg-bands or neck collars. Two of the geese, an adult male and female, were marked in conjunction with an attempt in Alberta to create nesting populations of geese which home to wintering areas in New Mexico. Two adult females were marked in conjunction with Canada goose nesting population restoration in North Dakota. �27 On 17 February 1982 at Johnson's Pond, a total of 23 geese were captured of which 19 had not been previously banded. All birds captured were subjectively judged to be large Canadas. All unbanded birds were measured, banded and released. Three of the 4 banded birds captured had been banded as goslings at Johnson's Pond in June 1981. A summary of some of the descriptive statistics of the birds banded in both areas are presented in Table 4. Foreign Recoveries Analysis of the location of banding of Canada geese recovered or recaptured in northeast Colorado indicates birds originated from scattered locations with Alberta providing most of the banded birds (Figure 2, Tables 5 and 6). Alberta banding locations providing birds to northeast Colorado were widespread and about equally divided between areas designated as Rocky Mountain Population production areas and those considered Hi-Line Population production areas. Banding areas in Saskatchewan were also widespread covering both Hi-Line and Western Prairie production areas. Only one recovery originated in the traditional Hi-Line breeding range in Montana. Birds banded in Wyoming, North Dakota and South Dakota originated primarily from areas in which breeding populations are being restored (Table 5). Birds banded on the Anderson River Delta, Northwest Territories are actually small Canada geese, banded with number eight size bands, which were captured on molting areas. LITERATURE CITED Krohn, W. B., and E. G. Bizeau. 1979. Molt migration of the Rocky Mountain population of the western Canada goose. Pp. 130-140 in Management and Biology of Pacific Flyway geese: a symposium. R. L.Jarvis and J. E. Bartonek (eds.). 346pp. Szymczak, M. R., R. C. Staffon, and J. F. Corey. 1981. Distribution and harvest of Canada geese nesting along the foothills of Colorado. Colo. Div. of Wildl., Spec. Rept. No. 49. 25pp. .' (:) ~... ' L P repare d by}: y __ ~/~£~C~~~(a&C~~. _~~~.~·~J~/~n~Ct~~?l~a:~ _ Michael R. Szy ak (~) Wildlife Researcher C . ( i~~1~dM.'-f~:~enfz;~~·~11;;;~ Sen ior Wi 1d 1ife B i0 Iog ist �Table 4. A summary of some selected descriptive statistics of Canada geese captured and banded in northeast Colorado, January - February 1982. Location Prewitt Res. Age-Sex Ad. Male Imm. Ma le Unk. Female Imm. Female Johnson's Pond Ad. Male Imm. Male Unk. Female Number Banded 26 4 24 1 8 3 8 Weight (k~) S.D. Range Wi ng chord (mm) Mean S.D. Range Culmen len9th (mm) .Mean S.D. Range 3.77 3.77 3.65 3.34 0.67 0.65 2.86-5.00 2.82-4.25 484 0.29 2.96-4.10 476 480 4.57 4.64 0.45 0.22 4.30-5.43 4.40-4.84 4.09 0.32 3.78-4.82 521 492 484 Mean -- 463 33 26 372-530 434-495 52.4 54.3 6. 1 4.6 43.5-62.2 47.4-58.1 19 431-518 53.7 3.3 47.5-59.8 -- -- 59.3 17 4 18 499-534 488-495 57.6 3.4 57.9 460-505 55.7 2.9 1.8 53.2-63.2 56.2-61. 3 53.0-58.6 N 00 �29 er _, l:- ~ oi :2 ~ •.... Wl Q ;a ClJ .,.... s..O ClJCO >1 ON U •...... ClJO'I s....-t �30 Table the 5. Banding hunting season areas outside in northeast Colorado of Canada Colorado, 1972 Area. geese through Direct reported recovered during 1980. Indirect Canada Northwest Territories Anderson River Delta (molting) Alberta Dowl ing lake Shooting lake Gledding lake East of Purple Spring Fincastle lake Grassy lake Hays Reservoir Diamond E Ranch Milk River Ridge Reservoir Saskatchewan Regina Manito lake (rejtoration). Maple lake Fishing lake Quil I lakes Supreme lake United o o 4 o 2 I 2 1 o o o o 2 I o 1 o o 1 2 1 o o o 1 o o 1 o 1 1 States Montana Ph i I lips Co. Wyoming Crook Co. (restoration) Sheridan Co. (restoration) Weston Co. (restoration) Johnson Co. (restoration) laramie Co. (restoration) lincoln Co. Bighorn Co. Goshen Co. Natrona Co. New Mexico San Miguel Co. (experimental) North Dakota Slope Co., Cedar lake (restoration) Adams Co. (restoration) Dunn Co. (restoration) South Dakota Day Co. (restoration) Harding Co. (restoration) Meade Co. (restoration) Jackson Co. (restoration) Nebraska Keith Co. (winter banding) Graden Co. (winter banding) Kansas Rooks Co. (winter banding) Utah Sanpete Co. Wisconsin Dodge Co. (winter banding) aSmal1 Canada geese banded with number o 2 1 o 2 1 o 1 o 1 o o 1 o o 1 o o 2 1 o 2 o o o 1 o o o o 8 bands. o 1 o 1 2 �Table 6. Banding areas outside Colorado of Canada geese reported recovered during the hunting in northeast Colorado 1972-80 that were classified as wild-trapped locals when banded. Area 1972 1973 1974 Year of recovery 1975 1976 1977 1978 1979 season 1980 Canada A.lberta Haus Reservoir Grassy Lake Fincastle Reservoir East of Purple Spring Dowl ing Lake Diamond E Ranch Milk River Ridge Res. Gleddys Lake Shoot ing Lake Saskatchewan Maple Lakes Supreme Lake United States Montana Phillips Co. Utah --Sanpete Co. Wyoming Lincoln Co. Natrona Co. 1 1 2 w ��Oc tobe r 1982 33 JOB FINAL REPORT S ta te of C_o_l_o_r_a_d_o _ P roj ect No. __ W_-_8_8-_R_-_2~7 3 Work Plan No. Central Period Covered: Personnel: Characteristics of Mallards 7 Wintering in West Colorado 1 Apri 1 1981 - 31 March Richard Bird Investigations Job No. Population Job Title: Migratory _ 1982 M. Hopper ABSTRACT A technical report is close to completion but was delayed because of change in responsibilities by the investigator. Publication costs will be covered under Work Plan 6, Job 1, Migratory Bird Publications and the publication wi 11 be completed in Segment 26, 1982-83. »:: P ! I) / '",.' \ j I .~ . repa re d b y __;1.;;:?'S:I~:::<""'&'="~~a;...!;-(.'!7t:t"'_:_:_~r:':_.I..i...1__;:--.A~.C<!-f.~~.~t>~· Richard M. Hopper 'i~} Wildlife Research Chief _ ��October 1982 35 JOB PROGRESS REPORT State of Colorado ----~~~~~----------- Project No. Work Plan No. Job Title: 3 Job No. Bird Investigations 8 Monitor Banding of Eastern Colorado Mallard Populations Period. Covered: Personnel: Migratory W-88-R-27 30 April 1981 - 31 March 1982 M. D. M. T. C. Babler, G. Berlin, L. Budde, L. Childers, J. Corey, Crawford, M. Creamer, L. Crooks, T. Davis, K. Dillinger, Ette, M. Gardner, J. Jackson, R. Kahn, G. Lorentzson, Lynch, F. Marcoux, R. Moore, R. Moss, R. Oehlkers, Pabst, J. Pogorelz, F. Rinella, J. Ringelman, S. Smith, H. Spear, E. Wagner. ABSTRACT A total of 3,485 mallards were banded postseason in six locations of eastern Colorado in segment 27. Some additional work was accomplished on an update of analysis of banding data but changes in personnel assignments out of the project delayed publication of results. ��37 MONITOR BANDING OF EASTERN COLORADO WINTERING MALLARD POPULATIONS Gerald M. Lorentzson P. N. OBJECTIVES 1. To establish monitor banding of wintering mallard populations eastern Colorado as an annual management function. in 2. To continually document, through monitor banding and analysis of recovery data, the annual and long-term status of eastern Colorado wintering mallards to provide a basis for annual hunting season recommendations. SEGMENT OBJECTIVES 1. Band a minimum of 4,000 mallards during the post-season period including a minimum of 500 birds in each of the following general areas of the South Platte Valley and Arkansas Valley: (1) DenverGreeley, (2) Fort Col 1ins-Loveland-Windsor, (3) Greeley-Fort Morgan, (4) Fort Morgan-Sterling, (5) Sterling-Julesburg, (6) Bonny Reservoir, (7) Manzanola-Lamar, and (8) Two Buttes Reservoir area. Divide the banded samples i n each area equally among the four age and sex classes. 2. Submit banding schedules and recapture data to Bird Banding Laboratory. 3. Conduct an updated analysis of band recovery data, including the following major determinations for important population segments of mallards wintering in eastern Colorado: (1) distribution of harvest, (2) recovery rates, and (3) survival rates. 4. Prepare progress report and publish pertinent findings. METHODS AND MATERIALS Procedures and equipment remained essentially the same as in previous years. The research section limited their actual banding to Bonny Reservoir. The remainder of the banding has been turned over to the Southeast and Northeast Regional personnel. �38 RESULTS AND DISCUSSION Nearly 3,500 mallards were banded postseason in January 1982 in eastern Colorado (Table 1). This number was less than the quota of 4,000. Two areas, Two Buttes and Manzanola-Lamar, did not contribute to the sample. The four age and sex classes were well represented in the banded sample. All banding schedules and recovery reports were prepared and sent to the Bird Banding Laboratory. Table 1. Mallards banded postseason by age and sex in 6 eastern Colorado banding areas, ~anuary 1982. Banding area AM Bonny Reservoir Sterling-Julesburg Fort Morgan-Sterling Greeley-Fort Morgan Denver-Greeley Fort Collins-Loveland-Windsor Totals Number of ducks banded A~e and sex SM SF AF Total 517 125 125 146 128 124 150 125 127 149 127 135 180 125 155 95 107 139 144 125 93 108 136 100 991 500 500 498 498 498 1,165 813 801 706 3,485 Updated Analysis of Band Recovery Data A previous segment report indicated the progress made through Segment 24 in regard to the updated analysis of banding data from eastern Colorado wintering mallard populations (Hopper 1979). This same report discussed the steps followed in the analysis, as well as the quantity of data used. Most of the analysis was finalized during Segment 25, and considerable progress was made in writing portions of the final manuscript. Some additional progress was made in Segments 26 and 27. However, preparation of a final report by way of a publication will be completed during the next segment under Work Plan 6, Job 1 and submitted for publication in an appropriate technical series. LITERATURE CITED Hopper, R. M. 1979. Monitor banding of eastern Colorado wintering mallard populations. Colo. Div. of Wildl. Fed. Aid Game Res. Rept:, Oct. Pp. 49-56. / Prepared by: tJ~dC(H.7'[Jr~f;~;t.l/;: Senior Wildlife Biologist �October 1982 39 JOB State of Project Work Job Plan Ti t Personnel: REPORT Colorado ----~~~~------------Migratory No. W-88-R-27 ---------~---------3 No. Job Bird Investigations 9 No. le ; _ __:.M.;...:i....;;gL.:.r..::;a~t~i..::;o.:..:n-.:=.a~n.::.d ....;M:...:;o~r:...t:.:a~l:...i:_;t:...ly:.......;:C:...:;h:..:a;..:.r..::a:...:c:...:t.. ..::.s.:.t.:..i ..::.c s::......:o:..f:.-....::D:..:u:.;c:..:.k.:......:.P.. in the Period FINAL Covered: Michael Inter-mountain 1 April 1981 R. Szymczak, Valleys - 31 March Colorado of Colorado 1982 Division of Wildlife ABSTRACT Work was contlnued toward a technical paper completed. Time and costs were transferred Migratory Bird Publications. The publication Segment 28, 1982~83. for publication, but not to Work Plan 6, Job 1, will be completed in P repa red by ---7}~)-tV""'·,...:;··~~tU~,.;...'·~(~).l...;._'......;L::::;/,.f·~/.."a~I'L,.-;'. "'-"..;.4~L~ Michael R. Szymc~!J Wildlife Researcher C ��October 1982 41 JOB PROGRESS REPORT State of Colorado -------------------------- Project No. Migratory w-88-R-27 Work Plan No. 3 Some Population Job Title: Bird Investigations 10 Job No. Characteristics of Adult Gadwall Molting in North Park Period Covered: Personnel: 21 July 1981 to 21 September 1982 J. Corey, J. Ringelman, S. Steinert and M. Szymczak, Colorado Division of Wildlife. ABSTRACT Trapping of gadwall (Anas strepera) in North Park resulted in the following number of birds being banded: adult male = 338; adult female = 210; immature male = 29; immature female = 19; local male = 52; local female = 41. ��43 SOME POPULATION CHARACTERISTICS OF ADULT GADWALL MOLTING IN NORTH PARK Michael R. Szymczak P. N. OBJECTIVES 1. To document the distribution of harvest of gadwall banded as flightless young or molting adults in North Park, Colorado. 2. To estimate recovery and survival rates of adult gadwall molting in North Park, Colorado. SEGMENT OBJECTIVES 1. Trap and band 400 adult males, 350 adult females and, if available, up to 100 local gadwall in North Park on molting and brood rearing areas in North Park. 2. Submit banding schedules and recapture reports to the U.S. Fish and Wildlife Service's Bird Banding Laboratory. File return information at the Colorado Division of Wildlife Research Center. 3. Prepare progress report. METHODS AND MATERIALS Most birds were captured from an air thrust boat at night using a handheld 12-volt landing light and a long-handled net. The birds captured at MacFarlane Reservoir were taken during drive-trapping operations. All birds captured were banded and released. The band numbers of birds captured that had been previously banded were recorded. Banding schedules and recapture reports were prepared and submitted to the U.S. Fish and Wildlife Service's Bird Banding Laboratory. Information on returning birds recaptured in the same 10-minute grid of banding were filed at the Colorado Division of Wildl ife Research Center. RESULTS AND DISCUSSION Trapping efforts fell short of producing the number of gadwall needed to meet quotas. The 2 major molting areas for gadwall, Walden Reservoir and MacFarlane Reservoir, suffered from adverse water conditions. MacFarlane was completely drained, while Walden had the lowest water levels observed in at least the past 10 years. Gadwall were present in reduced numbers on Walden Reservoir and totally absent from MacFarlane. �44 The molters apparently did not select other marshes in North Park to spend the fl ightless period. No large concentrations of molting adults were found on other ponds and attempts to capture birds on other areas met with limited success. The results of the trapping are presented in Table 1. Table 1. Number of gadwall banded in North Park, July through September 1981. Locat ion Walden Reservoir Pole Mountain Reservoir Case Flats Lake John Annex MacFarlane Reservoir Hebron Ponds Total Prepared by , i LF Tota I 19 0 0 0 0 11 4 36 0 0 1 8 10 22 0 0 1 494 78 58 43 13 3 19 52 41 689 AF 260 25 0 39 13 1 167 39 0 4 0 29 0 0 0 0 338 210 29 tk~.J:._j)1? ~4i_ - Michael R. Szymczak Wildlife Researcher C A£!e and sex 1M IF LM AM �October 1982 45 JOB PROGRESS State of REPORT Colorado ------------------------Migratory Project No. W-88-R-27 Work Plan No. --------------------- 3 Job No. Banding of Preseason Waterfowl Job Title: Bird Investigations 11 Populations in the San Luis Va 11ey Period Covered: Personnel: ·1 July to 15 November 1981 M. Nail and staff, Monte Vista and Alamosa National Wildlife Refuges; J. Corey, G. Lorentzson, J. Ringelman, S. Steinert, Colorado Division of Wildlife. ABSTRACT Totals of 1,080 mallards and 1,651 other ducks were banded in the San Luis Valley during the summer of 1981. Shortages of water with the resulting loss of ducks prohibited us from meeting our quota of all mallards with the exception of immature males. ��47 BANDING OF PRESEASON WATERFOWL POPULATIONS IN THE SAN LUIS VALLEY G. M. Lorentzson P. N. OBJECTIVES The objective of this job is to continually document, through monitor banding and analysis of recovery data, the status of the San Luis Valley preseason mallard population. SEGMENT OBJECTIVES 1. Trap and band at least 300 mallards of each sex and age group, adult males, adult females, immature males and immature females. 2. Band all other waterfowl mallard trapping. 3. Submit banding schedules and recapture reports to the U.S. Fish and Wildlife Services Bird Banding Laboratory, file resulting recovery cards at the Fort Collins Research Center. species captured during the period of 4. Prepare Progress Report. METHODS AND MATERIALS Ducks were trapped in the San Luis Valley from August 12 through September 17, 1981. Mallards were the target species but all other ducks captured were banded. Salt Plains types of duck traps were used in the banding program for capturing the birds. Barley was used as bait. Ducks were banded and recorded according to age and sex. All banding reports were sent to the U.S. Fish and Wildlife Services Bird Banding Laboratory. RESULTS AND DISCUSSION A total of 2,731 ducks were banded in the San Luis Valley of Colorado during the summer months in 1981. The number and composition of each species are presented in Table 1. Except for immature males the quota of 300 mallards of each age and sex group was not reached due to the shortage of water and the absence of birds. �48 Table 1. Number of ducks banded, by species, during the pre-season period, 1981. 1 Seecies Mallard Blue-winged and/or Cinnamon Teal Green winged Teal Pintail Gadwa 11 Redhead Lesser Scaup Shoveler Wigeon Totals in the San Luis Valley Age and sex IF AF LM LF Total 253 13 14 1,080 2 0 6 6 15 0 1 275 62 50 111 20 3 0 0 0 3 1 1 3 11 0 0 0 778 375 213 204 73 5 2 1 379 774 43 33 2,731 AM 1M 262 330 208 142 158 43 1 1 0 0 0 607 281 114 64 81 22 2 75 40 49 2 4 0 0 1 0 895 1 Ilncludes 1,685 ducks banded by Monte Vista National Wildlife Refuge personnel. The water available for waterfowl nesting is declining each year as more irrigation systems are installed. As a result of the change in methods of irrigation, ditches and ponds which were used as a water source for irrigation have been abandoned. This has resulted in the loss of valuable nesting and breeding habitat for waterfowl. Prepared by: Gera 1d M. Lorentzso~ {j3 Senior Wildlife Biologist �October 1982 JOB FINAL REPORT State of Colorado ------~~--~----------Migratory W-88-R-27 Project No. Work Plan No. Job No. Bird Investigations 8 Evaluation of Daily Counts of Band-tailed Job Title: Pigeons as a Census Method Period Covered: Personne 1: January 1979 to 31 March 1982 P. D. Curtis, R. A. Ryder, J. Ell is, and T. E. Olson, Colorado State University; C. E. Braun, H. D. Funk, and K. M. Giesen, Colorado Division of Wildlife; C. J. Curtis. ABSTRACT The objectives of this study have been fully achieved. during 2 field seasons (April-September 1979 and 1980). covering study objectives have either been prepared, or preparation. Those submitted, published, or in progress Curtis, P. D. 1981. Band-tails. Data were collected Manuscripts are currently in are listed below: Colo. Outdoors 30(4}:30-33. 1981. Evaluation of daily counts of band-tailed pigeons as a census method. M. S. Thesis. Colo. State Univ., Fort· Collins. 113pp. 1982. An albinistic West. Birds. In Press. band-tailed pigeon in Evergreen, Colorado. _______ , and C. E. Braun. 1980. Evaluation of daily counts of bandtailed pigeons as a census method. J. Colo.-Wyo. Acad. Sci. 12(1): 36-37. ------,-- , and 1981. bait sites in Colorado. _______ , and tailed pigeons Band-tailed pigeon behavior at artificial J. Colo.-Wyo. Acad. Sci. 13(1):56. 1983. Radio-telemetry location of. nesting bandin Colorado. Wilson Bull. 95:Accepted. �50 , and ---censusing Establishment band-tailed pigeons. and placement of bait sites for Wildl. Soc. Bull. Submitted. , and Band-tailed pigeon --~Colorado.--~~ Behavior. Submitted. ____ , and -:-- _ ta iled pigeons. behavior at bait sites in Wing markers for individual recognition Submitted. of band- J. Field Ornithol. Stabler, R. M., C. E. Braun, and P. D. Curtis. 1981. Bacterial disease in band-tailed pigeons. J. Colo.-Wyo. Acad. Sci. 13(1):59. Prepared by: Paul D. Curtis I~) Graduate Research Assistant Approved by: Clait E. Braun Ip) Wildlife Research Leader �October 1982 51 JOB PROGRESS REPORT S tate of __ __..;C;,_;o;,_;l;,_;o_r_a_d~o _ Project No. Work Plan No. 6 ----------------Migratory Job Ti t le : Bird Investigations Job No. Bird Publications 1 April 1982 - 31 March 1982 Period Covered: Personnel: Migratory w-88-R-27 C. E. Braun, J. F. Corey, P. D. Curtis, W. P. Gorenzel, T. E. Olson, R. A. Ryder, R. M. Stabler, R. ·C. Staffon, and M. R. ~zymczak. ABSTRACT Publications planned for and accomplished are as follows: under this job for Segment 27 Gorenzel, W. P., R. A. Ryder, and C. E. Braun. 1981. American coot distribution and migration in Colorado. Wilson Bull. 93:115-118. , ---teristics , and of American Condor 84:59-65. 1982. Reproduction and nest site characcoots at different altitudes in Colorado. Olson, T. E., and C. E. Braun. 1982. September nesting and crop gland activity of mourning doves in Colorado. Proc. 52nd Annu. Mtg., Cooper Ornithol. Soc., Logan, Utah. Abstract. Stabler, R. M., C. E. Braun, and P. D. Curtis. 1981. Bacterial disease in band-tailed pigeons. J. Colo.-Wyo. Acad. Sci. 13(1):59. Szymczak, M. R., R. C. Staffon, and J. F. Corey. 1981. harvest of Canada geese nesting along the foothills Colo. Div. Wildl. Spec. Rep. No. 49. 25pp. Prepared by ~/aM J. d{.d Howard D. Funk (~~j Wildlife Research L~ader Distribution of Colorado. and ��October 1982 53 JOB PROGRESS REPORT State of Colorado ----~~~~~------------ Project No. Work Plan No. 8 ----~--------------- Job Title: Computerized Period. Covered: Job No. System for Storing and Retrieval of Banding, Recoyery and Recapture Personnel: Migratory Bird Investigations W-88-R-27 ------------------------ Information 30 April 1981 - 31 March 1982 J. Corey, H. Funk, R. Hopper, J. Ringelman, H. Szymczak, Colorado Division of Wildlife; D. Markham, Colorado State University. ABSTRACT Literature reviews and consultations with computer personnel indicated that a system consisting of banding, recovery, and recapture files would provide the best approach to maintaining banding records and accessing data for analyses. Banding and recovery records were obtained from the Bird Banding Laboratory, and need only to be reformated to be fully operational. The Bird Banding Lap can supply annual updates for recoveries, but banding data must be kept current by data input through a Fortran computer program. Data on birds recaptured at the same banding location have been partially coded from field forms and are ready for keypunching. ��55 COMPUTERIZED SYSTEM FOR STORING AND RETRIEVAL OF BANDING, RECOVERY, AND RECAPTURE INFORMATION James K. Ringelman P. N. OBJECTIVE Major objectives of this job are to establish a computer system for banding and recovery data, then to enter all necessary data 'from past ~nd future banding programs into the system for ac~urate ,and efficient record, keeping, analysis, and reporting programs., SEGMENT OBJECTIVES 1. Review literature and consult computer experts regarding development of a computer system for storage and retrieval of banding, recapture, and recovery data~ 2. Develop a computerized system. 3. 'Initiate entry of existing and new data into system., METHODS AND MATERIALS Literature reviews were conducted to determine the other researchers to organize and retrieve banding Systems documentation was obtained for the Colorado Computer to serve as guides to developing retrieval of the U.S. Fish and Wildlife Service Bird Banding sulted for data tapes of bandings and recoveries. methodology used by and recapture data. State University CDC programs. Personnel Laboratory were con- RESULTS AND DISCUSSION A system consisting of three data files was designed to meet the requirements of record keeping and data analysis. The descriptions, formats, and initial and maintenance programming needs of each file are detailed in Table 1. Data for the banding tub file, which consists of banding records for all waterfowl banded in Colorado from 1967-81, was obtained by special request from the Bird Banding Laboratory. The tape has been entered in the tape library, and work has begun on the problem of reformating this IBM System tape to make it compatable with the CDC System. Because annual updates of bandin~ tub data cannoi be obtained from the Bird Banding Laboratory, a fortran computer program (Cowardin and Davenport 1973) was obtained which wil I enable personnel to simultaneously code banding data for generation of banding schedules as well as for direct entry into the tub file records. This program is currently being converted to the CDC System format. ' " �56 Table 1. Description banding data. of the three files to be used in the computerized system for storing and retrieving BANDltlG TUB FILE Description: Contains data on every duck and goose banded in Colorado from 1967 to the present, re9ardless of whether the bird was ever re-encountered. Present tape format: 9 track, unlabeled, epsidic char. set., record length = 53, block s lze record count = 260,695, 615 records/block. Initial setup problem: > 32,595', Data needs to be resorted by band number within hand size. Maintenance problem: As new birds are banded each year, these data must be added to the tub file. It may be desirable to have each sub-file (band size) stored on separate tapes to avoid the need for rewriting the entire tub file with each ,update. RECAPTURE FILE Description: Contains data on banded ducks and geese that were subsequently recaptured by field biologists during banding, operations. In some instances, an individual bird may have several recapture records. Present format: Data on recaptures is of two types. Foreign recaptures (described above) are coded with recoveries in the recovery file. A numeric code identifies foreign recaptures from recovery data. Return data (banded birds recaptured in subsequent years at the same banding site) is presently being coded and keypunched. Initial setup problem: Complete return records must be created by match-merging newl y -coded data with banding tub data using band number as the identifier. Table outlines the present card-image format of the return data. Two important conditions must be met before recapture data can be used. First, only the last (terminal) encounter of a bird is useful in analyses. Thus, birds classified as recoveries must be eliminated from this category if the same bird is represented as a hunting recovery, which constitutes the terminal encounter. Second, recapture data must be compared within itself to eliminate all observations on an individual bird except the terminal recapture. The procedure below is suggested as a means to construct the recovery file: Program needs: User-created editor programs will be used to extract subsets of data according to several criteria. CSU editor "locate" commands, will be used to extract banding information on a single bird based on band number. Once again, to reduce time in extracting data, it may be beneficial to maintain separate tapes for each band size. RECOVERY FILE Description: Contains banding data (as found in the tub file) and recovery information for every duck or goose band reported (usually by hunters) except those encountered during subsequent banding operations. Present tape format: 9 track, unlabeled, epsidic char. set, record length count - approx. 20,000, 1600 BPI. = 81, block size = 4,050, record Initial setup problem: Data needs to be resorted by band number within band size. Additionally, a slight re-formating of records, as shown in Table, wi II be neces sarv ; Data on "foreign recaptures" (birds recaptured at a location different from where they were originally banded), currently combined with recovery data, must be extracted from this data set and recombined with other recapture data in the recapture file. Maintenance problem: The master recovery file must be updated annually with new data, just as the case with the banding tub file. 1. Match-merge return data with banding tub file by band number, then re-format data to conform to the record format shown in Table 2b. 2. Add foreign recapture data to return data to create the basic recovery file. 3. Sort recapture data to create sub-files by band size. Before performing analyses, terminal recoveries must be extracted and placed in a working data set: 1. Compare recovery file with recapture file and temporarily eliminate from recapture file records of birds shot by hunters. 2. Compare recapture file internally and temporarily eliminate all but the terminal recapture record for each bird. Maintenance problem: As new birds are recaptured during annual banding operations, the banding tub file must be accessed to determine the bandin9 history. These data are then merged with the recapture data, and the completed recapture records added to the master recapture file. �57 The master recovery file, consisting of 20,000 entries of Colorado-banded birds recovered since 1967, was received from the Bird Banding Laboratory. The present tape file needs only to be reblocked and reformated to be fully functional. Annual recovery tape updates will be obtained to keep this file current. The recapture file contains data on banded waterfowl that were subsequently re-encountered by banders during banding operations. A new analytical technique (Mardekian and McDonald 1981) will allow these recapture data to be used along with recoveries in the analysis of survival rates. Whereas data on birds re-encountered outside of the area of banding (foreign retraps) are available from the master recovery file, information on birds re-encountered at the same banding location (returns) must be coded and entered from field forms, since these data are usually not included in the Bird Banding Laboratory files. Nearly 1,300 return records during the period 1967-75 have been extracted from field forms, coded~ and are ready for keypunching. Work will continue on recoding returns from 1976-82. LITERATURE CITED Cowardin, L. M., and D. A. Davenport. 1973. Computerized system for organizing and maintaining files of banding data. Bird Banding 44:187-195. . Mardekian, S. Z., and L. McDonald. 1981. Simultaneous analysis of band-recovery and live-recapture data. J. Wildl. Manage. 45:484- 488. Prepared by ~~~~~~~ ~~~~~.~~~'?_~_~_¥ __ __ Ringelman Researcher t~ _ ��October 1982 59 JOB PROGRESS REPORT State of Colorado ------------------------- Project No. Work Plan No. 9 --------------------- Job Title: Migratory Period Covered: Personnel: Migratory W-88-R-27 -------------------- Bird Investigations Job No. Bird Research Planning Apri 1 1981 - 31 March 1982 Howard Funk, James Ringelman and Michael Szymczak. ABSTRACT Progress was made toward long-range research planning for w-88-R in association with the Strategic Plan and Flyway management plans. A change in period of time for long-range plans to 5 years, and alterations in format and procedures caused changes in timing necessary. Thus, an extension of this job was requested to include the 1982-83 segment period. ��61 MIGRATORY BIRD RESEARCH PLANNING Howard Funk James Ringelman Michael Szymczak P. N. OBJECTIVES To prepare long-range research plans for w-88-R for a period of 10 years. SEGMENT OBJECTIVES 1. Review ongoing present research jobs in light of research needs as outlined in the Colorado strategic plan, cooperative Flyway management plans and other ongoing Central or Pacific Flyway cooperative studies and for management efforts which are not in management plan form. 2. Review literature in terms of what the Research Section believes as priority research efforts for in-state needs or for continuity and/or expansion of cooperative efforts among Flyway states, either as a group or in cooperation with the Fish and Wildlife Service and/or Canadian wildlife agencies. 3. Obtain input from other sections of the Division of Wildlife and, where necessary, from other wildlife agencies. 4. Prepare a draft management plan, present the plan for additional input and/or alteration and initial approval to the various Division of Wildlife sections. 5. Prepare final research plan and obtain approval. METHODS AND MATERIALS During the segment, progress was limited to review of literature and examination of research and management needs as presented by the Colorado Strategic Plan, those expressed by the Regions, and other possible programs from Flyway management plans. RESULTS AND DISCUSSION Initial plans for jobs which were believed to be essential or feasible were prepared by Research personnel for a period which would approximate 10 years. Because of management activities which have to remain part of Project W-88-R, all time available by project personnel cannot be allocated strictly to research. Thus, new research jobs to be considered have to be related to time and funds available along with management related functions. �62 Considerable time and effort were expended in meetings with the four Regions in relation to migratory game bird programs, including research and management, which they felt were important for the Division of Wildlife to be involved in from the present to 1988. Those items which were deemed feasible for research, those which involved mainly Region activity, and those which could be cooperatively handled by w-88-R and the Regions were combined and presented to the Denver Staff for inclusion in the 3rd edition of the Strategic Plan scheduled for printing in 1983. Data included for the new plan included updating of harvest, hunter, recreation day and migratory game bird population estimate data. Time was also spent in preparation of narrative information and alteration of data necessitated by changes in plans in regard to the new plan. Other changes in plans in regard to long-range research plans have altered the status of progress. Instead of 10-year plans, the decision was made to prepare 5-year plans and gear these to new operations plans for the Division. These changes include alterations in format. Thus, an extension of time into the next segment (Segment 28) was requested for this job and planning will be completed in the 1982-83 segment. P repa red by J1) - ~f"d Howard D. Funk (-ItS) Wildlife Research Leader ~~atC-r �October 1982 63 JOB PROGRESS REPORT State of Colorado ----~~~~~----------- w-88-R-27 Project No. Work Plan No. 10 -------------------- Job Title: Cooperative Period Covered: Personnel: Migratory Management Bird Investigations Job No. Programs 1 April 1981 - 31 March 1982 John Corey, Gerald Lorentzson, Steve Steinert, Michael Szymczak, Colorado Division of Wildlife. ABSTRACT Work accomplished during this segment consisted of assistance in cooperative management programs with the four regions of Colorado and the States in the Central Flyway. Help was also given to the U.S. Fish and Wildlife Service consistent with the objectives of this work plan. ��65 COOPERATIVE MANAGEMENT PROGRAMS Gerald Lorentzson P. N. OBJECTIVES The major objective of this job is to provide liaison with the various DOW sections in order to assist and work with them in migratory game bird matters, particularly those relating to management responsibilities placed upon the various sections. SEGMENT OBJECTIVES The procedures for this job will be to provide liaison and expertise to the Regions and other DOW entities in assisting them with management activities for which they have been assigned main or partial responsibiIities. This assistance will be given in the way of training and/or help in design of cooperative management programs such as monitor banding and various migratory game bird surveys. Assistance will be given in design of goose transplant programs and monitoring of results. Procedures for federal hunting season 'requirements and limited permit hunts will be explained and assistance given in setting up seasons and monitoring results. As necessary, assistance will be given in sampling schemes, hunter surveys and statistical analyses of results. Cooperative effort will be supplied in preparation of strategic plans for mrg~atory game birds. Project personnel will take part in various technical committees, status and regulations meetings and management plan workshops both instate and out of state as necessary for cooperative research and/or management oriented purposes. METHODS AND MATERIALS Due to the diverse nature of this program, various procedures and material were used to accomplish the objectives. The methods and materials used for trapping, banding, inventory studies, and the compilation of harvest data have been well documented in other segments of this program and will not be repeated in this report. RESULTS Assistance was given to the Regions flying goose breeding pair counts on the Yampa, White, Colorado and Gunnison rivers during April. Revisions were made in the Poudre and South Platte River surveys to replace the sample land and river sections which had been rendered useless by the destruction of habitat by housing developments and changes in farming practices. The initial draft for a goose transplant program was written and submitted for approval. �66 Assfstance was given in flying the breeding pair counts in the San Luis and Cache la Poudre River Valleys, North Park and the South Platte River drainage. Assistance was given in conducting the mourning dove survey program. Numbers of goose nesting structures and their locations were recorded and given to the Northeast Region for their ~se. Gosling production data for the last 10 years was also included in the data supplied the Northeast Region. Time was spent assembling High Plains and Low Plains mallard and other duck harvest data for Colorado and the Central Flyway states. Time was spent in the program for Endrin contamination examination during the waterfowl seasons in Color~do. Harvest data, average bag, numbers of hunters and recreation days (migratory waterfowl and doves) were assembled and submitted for the Strategic Plan. Banding materials and assistance was given to the Regions for the winter banding of ducks and geese. Ducks banded in the west central portion of Colorado are reported in Table 1. Table 1. Mallards Janua ry 1982. banded in the west-central portion of Colorado, AM SM AF SF Total Delta 100 45 47 71 263 Mack (Hi Line Lake) 118 82 99 101 400 281 127 146 172 663 Totals Time was spent instructing DOW personnel in the "reading" of duck wings for age and sex characteristics. This involved several sessions including the pre Wing Bee period when waterfowl collections for the Central Flyway were sorted according to species and states. Assistance was also given to setting up material for the Wing Bee and through actual assistance at the Wing Bee. Colorado hosted the spring Central Flyway Technical Committee in 1982. Time was spent preparing for and participating in this meeting. ,( . ~ Pre pa red by -";'~""G""J;..~~~'""1~(~d~Y""" M:-."'-/)4L.L.~~r-e.z:.fl..t /) ••.. z'/l,Ra,s~o.<!i~ ..•. 'l..:;gtJ'lf~:ct:...t<-'"" 0:;..,.1__ Senior Wildlife Biologist � https://cpw.cvlcollections.org/files/original/cc563f394e01bd845ad64dbb3d8d3af6.pdf 2c776a6b9902c299cd9087f91622c4c5 PDF Text Text 1 JOB PROGRESS REPORTa State of COLORADO Project No. N-1-R (SE-3-S) Work Plan No. 1 (II) Job No. 1 (I) Period Covered: Personnel: Nesting Performance of Peregrine Falcons in Colorado 1 July 1982 - 30 June 1983 D. Berger and G. Craig, Colorado Division of Wildlife; J. Enderson, The Colorado College. ABSTRACT In 1983, the number of occupied peregrine falcon (Falco peregrinus ariatum)breeding territories increased to 13. Released falcons were documented as members of pairs at 3 territories and an adult male released in 1976 was found at a recently discovered nesting site. Eggshell thickness measurements were obtained from 27 wild eggs removed during augmentation efforts and 8 nonviable eggs will be submitted for pesticide analysis. aThis Job Report represents a preliminary analysis and is subject to change. Information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the Director, Colorado Division of Wildlife. ��3 NESTING PERFORMANCE OF PEREGRINE FALCONS IN COLORADO Gerald R. Craig P. N. OBJECTIVE The objectives of this study are to annually monitor breeding numbers and reproduction of Colorado peregrine falcons to document further population declines as well as record the population's responses to recovery efforts. Additionally, hea Lt.h of the population will be monitored indirectly by analyzing pesticide residue levels in the falcons' eggs and principal prey. Information obtained from these investi.gations will be made available through annual reports to the Rocky Mountain/Southwest Peregrine Falcon Recovery Team as well as cooperating agencies to aid in evaluation of recovery efforts. SEGMENT OBJECTIVE 1. Annually monitor the number of breeding pairs of peregrines and reproduction in Colorado. 2. Annually monitor organochlorine pesticide levels in wild breeding peregrines. 3. Monitor recruitment of reintroduced peregrines into the wild breeding population of Colorado. 4. Compile data and submit reports to appropriate state and federal personnel and the Rocky Mountain/Southwest Peregrine Falcon Recovery Team for use in evaluating recovery efforts. (These objectives correspond to Jobs 1113., 1114., '212., 221., 3211., 3212., and 333. of the approved American Peregrine Falcon Recovery Plan for the Rocky Mountain/Southwest Population). METHODS AND MATERIALS Methods and Materials used in this study have been described previously (Craig and Enderson 1981). RESULTS AND DISCUSSION Territory Occupancy The number of occupied breeding territories increased from 11 in 1982 to 13 in 1983 (Table 1). Pairs returned to 6 of the sites occupied in 1982, 2 sites frequented by lone adults in 1982 were vacant, 2 formerly �Table 1. Occupancy and productivity of peregrine falcon eyries in Colorado, 1972-83 .po.. Parameter 1972 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 Known territories 15 23 24 26 27 31 32 33 34 34 34 36 Occupied territories 11 12 9 8 8 12 11 12 13 11 11 13 Adult pairs 8 11 I 6 5 11 7 6 7 7 9 9 Immature pairs a 0 0 0 0 2 0 2 2 3 1 0 3 Lone adults 3 1 2 1 1 1 2 3 3 3 3 1 Successful pairs 0 1 5 2 4 6 5 4 5 6 6 7 No. :yng. fledged No. yng. augmented 0/0 2/0 13/2 5/2 6/4 11/5 16/15 12/12 16/13 15/21 20/26 21/27 Young fledged/ adult pairb 0.00 0.18 1.57 0.83 1.20 1.00 2.29 2.00 2.29 2.14 2.22 2.33 Young fledged/ successful pair 0.00 2.00 2.60 2.50 1.50 1.83 3.20 3.00 3.20 2.50 3.33 3.00 Sites occupied, %c 73 52 38 31 30 39 34 36 38 32 32 36 Sites TN/adult pairs, % 53 49 29 27 26 35 22 18 24 21 26 26 .., aAt least one member of the pair was in juvenile plumage. b1.25 young fledged per pair is considered normal reproduction. cFor a normally reproducing population, particular year. 80-90% of the eyrie sites should normally by occupied in any �5 vacant sites were reoccupied, were discovered. and 2 previously undocumented territories The 2 additional occupied breeding territories located in 1983 increased the total territories recorded for Colorado to 36. It is probable that site 38 was occupied historically since the habitat is ideal for peregrines, but until recentl , the landowner prohibited access to the area. A caretaker encountered the male about 1 mile from the nest cliff, dying of injuries suffered from a powerline collision. The Division was contacted and subsequent investigations located the female on the nest cliff. The 2nd new territory (site 39) was reported late in the breeding season and although adults were observed, no young had been produced. It is possible that this pair relocated from a release site they were frequenting in 1982. Pairs returned to sites 8, 25, 30, 31, and 34 in 1983 and all successfully fledged young. Although both adults and their brood disappeared from site 9 in 1982, an adult male and yearling female reoccupied the territory in 1983. The female at site 34 successfully attracted another adult male when her previous mate failed to return. Territory 27, which had been frequented by a yearling male the previous year, was inhabitated by a pair which successfully fledged young. A pair of yearling peregrines was observed at site 5 which was vacant in 1982. After a 7-year hiatus, site 3 was reoccupied by an adult male (released in 1981) and a yearling female. Sufficient numbers of peregrines were present in the wil.d to replace 1 pair lost at 1 site, a male at another. and to reoccupy 2 vacant territories. Unattached peregrines observed interacting with breeding pairs at sites 30, 34, and 35 are also indicative of population recovery. ~eproduction Reproductive success in 1983 was good (Table 1). In all, 7 pairs were confirmed to have laid eggs and all were manipulated to augment reproduction. It is possible that falcons at sites 38 and 39 may have produced eggs. These sites were located too late in the breeding season to confirm incubation behavior. In all, 27 eggs were removed from the 7 wild nests (3.86 eggs/clutch) for artificial incubation. Several eggs were dented or cracked and others exhibited excessive water loss, all indicative of poor shell condition. A total of 19 young was successfully hatched from wild eggs and returned to the wild. Undoubtedly, reproducti.on would have been less had the eggs been left with the wild adults to be incubated naturally. In 1983, 27 captive-hatched young were placed at the 7 wild sites (3.86 young/pair) and 21 successfully fledged (3.00 young/pair). Golden eagle (Aquila chrysaet~s) predation probably accounted for the death of 1 young at site 8. Five young (1 at site 25, 2 at site 31, and 2 at site 35) disappeared from 3 other broods prior to fledging. figgshell Thickness Shell thickness of the 27 eggs taken from the wild in 1983 have not been measured and will be reported in the next Job Progress Report. �6 The presence of several cracked and dented eggs as well as excessive dehydration of several other eggs in 1983 suggests that shell thinning is still a problem at some sites. Organochlorine Residues in E Contents Pesticide residue analysis was performed by the U.S. Fish and Wildlife Service at the Patuxent Wildlife Research Center on contents of 6 nonviable eggs collected in 1982. DDE residues averaged (arithmetically) 22.06 ppm (range 7.11.-32.09) fresh wet weight basis in the 6 eggs. While this limited sample represents 4 sites (8. 9, 34, and 35), there was an overall increase in residue levels from a sample of 8 eggs collected in 1981 (13.7 ppm arithmetic mean). At the same time, eggshell thickness increased from an average of 0.318 mm in 1981 to 0.323 in 1982. Due to the limited sample of eggs subjected to residue analysis, more credance should be given to the statistically larger sample of thi.cknessmeasurements which continue to demonstrate an improving trend toward thicker eggshells. One sobering note from the 1982 pesticide analysis was that a known-age female hatched in 1979 accumulated critical levels of 32 ppm DDE in her eggs within 3 years. An additional 8 nonviable eggs collected in 1983 will be submitted to the U.s. Fish and Wildlife Service for pestieide residue analysis. Recruitment of Released Peregrines into the .Wi11 A banded male released at a nearby hack site in 1980 returned to site 8 for the 2nd ye r to mate with the wild female which was also present in 1982. The female was sufficiently aggressive to be caught at the eyrie in 1982 and was banded. In 1983, the plastic marker was no longer affixed to her leg, but the U.S. Fish and Wildlife Service band remained attached. A male released at a hack site adjacent to site 11 in 1981 was paired with an unbanded (wild-produced) yearling female at site 3, approximately 48 km west of his release site. The most significant event was discovery of a banded male at site 38. This tiercel appears to have been breeding at that location for several years. In 1976, this male was fostered into a brood at site 7 about 29 km east of site 38. It survived 7 years in the wild before dying from a collision with a power line. Word was also received that a falcon hacked at site 36 in 1982 was observed paired with a wild adult male at an eyrie in New Mexico. A male released from the same site in 1982 returned and fed with the young released there in 1983. A 3rd female falcon released from site 36 in 1982 was trapped at Matamoras, Tamaulipas,Mexico, below Brownsville, Texas. Finally, a female released at site 11 in 1982 was observed interacting with young released at a hack site in Wyoming in June 1983. Photographic Documentation of Wild Adults Photographs were taken of adult peregrines when sites were visited to manipulate reproduction. At present, these photos are being �7 developed and analyzed to compare plumage characteristics of individuals photographed at the sites in previous years. LITERATURE CITED Craig, G. R., and J. H. Enderson. 1981. Nesting performance of peregrine falcons in Colorado. Job Prog. Rep., Colo. Div. Wildl., Wildl. Res. Rep., Jan., pp 13-23. Prepared by G.sz.tut1'd t? .~GH Gerald R. Craig Wildlife Researcher C '; ��9 JOB PROGRESS REPORTa State of COLORADO Project No. N-1-R (SE-3-5) Reintroduction and Augmentation of Work Plan No. 1 (II) Job No. 2 (2) Period Covered: 1 July 1982 - 30 June 1983 Personnel: Peregrine Falcon Production D. Berger, G. Craig, T. Fowler, B. Grebence, R. Meese, and M. Robert, Colorado Division of Wildlife; J. Enderson, Colorado College; J. Hogan and S. Petersburg, U.S. Department of Interior, National Park Service. ABSTRACT Fledging success of 7 wild breeding pairs of peregrine falcons (Falco peregrinus ana tum) was increased to 3.00 young/pair through augmentation efforts in 1983. The 1982 hacking effort was diminished by failure of 2 of the 5 sites which resulted in successful release of only 13 young. A 6th hack site was added in 1983. aThis Job Report represents a preliminary analysis and is subject to change. Information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the Director, Colorado Division of Wildlife. ��11 REINTRODUCTION AND AUGMENTATION OF PEREGRINE FALCON PRODUCTION Gerald R. Craig P. N. OBJECTIVE The objective of this program is to sustain the wild breeding peregrine falconpopulation in Colorado through augmentation of poor natural production and re-establishment of breeding pairs at vacant sites by release of captive-produced falcons. SEGMENT OBJECTIVES 1. Augment poor wild production by placement of captive-hatched wild young and captive-produced young into occupied wild nests. 2. Release captive-hatched wild young and captive-produced young to the wild through "hacking" at-potential and abandoned wild nests. 3. Develop and implement release of adult falcons at potential or abandoned nest sites and at wild nests occupied by lone adults. 4. Monitor results of the efforts, compile data, and submit reports to appropriate state and federal agencies and the Rocky Mountain/ Southwest Peregrine Faleon Recovery Team. (These objectives correspond to Jobs 222., 3133., 321., 322., 331., 332., and 3211. in the approved American Peregrine Falcon Recovery Plan for the Rocky Mountain/Southwest Population). METHODS AND MATERIALS Methods and Materials for this study have been described previously (Craig 1982). RESULTS AND DISCUSSION At~mentation Efforts The 7 wild breeding pairs of peregrines encountered in the 1983 season (sites 8, 25, 27, 30, 31, 34, and 35) were manipulated to increase productivity. One other territory (38) was discovered during incubation, but the male had been killed and the decision was made not to attempt augmentation. No sites were recycled since The Peregrine Fund anticipated that captive production was sufficient to provide young for the augmentation effort. In retrospect, it might have been advantageous to recycle several sites to bring them into synchrony with captive production. Thus, young of the proper age (at least 18 days of age) were not available �12 and sites 31, 34, and 35 had to be delayed with broods of pra1r1e falcon (Falco mexicanus) chicks for 13 days until the captive-hatched peregrines were the proper age for placement in the wild. A total of 27 eggs was removed from 7 breeding pairs (3.86 eggs/pair) for artificial incubation at The Peregrine Fund's Fort Collins facilities and 27 captive-hatched young were replaced in the 7 nests (3.86 young/pair). The 7 pairs fledged 21 young for a success of 3.00 young/pair. Five nestlings were lost from 3 broods from unknown causes. A 6th fledgling disappeared (site 9) at the time of fledging, possibly due to golden eagle (Aquila ~hrysaetos) predation. Manipulation of wild eggs and broods obscures the nesting success the peregrines would have experienced had they been permitted to reproduce naturally. It is possible, however, to develop an estimate of natural reproduction based upon: (1) hatching success of eggs brought into the laboratory, and (2) nestling mortality experienced in the wild during the brood rearing phase. Thus, it can be assumed that in the wild, only 18 of the 27 eggs would have hatched and 6 nestlings would have died, yielding an expected natural fledging success uf 12 young or 1.71 young per successful pair. This estimated natural fledging success is slightly optimistic since the laboratory treatment of eggs probably increased hatching success and nestling exposure to mortality was reduced because young were placed in the wild at 20-24 days of age. Hacking Efforts Six hack sites were operated. Their final success is not known. In the 1982 season, 5 sites were operated with limited success. Twenty-three young were placed in the hack boxes (4.6/site) and 13 (2.6/site) successfully reached independence. One site (#14) failed completely when great horned owls (~_~Q~ virginianus) killed 4 young. Another young was lost at site 7 when it was apparently injured by an adult female peregrine that frequented the Vicinity. The remains of the fledgling were found under a rock at the base of the hack cliff several weeks after it disappeared. It appears that the falcon died of wounds received when the adult attacked. Site 37 was assumed to have failed when the 5 fledglings disappeared during a prolonged storm. A pair of yearling falcons were regularly frequenting the hack site and it is speculated that the young left the site with the yearlings during the storm. Adult Release Based upon the poor results encountered in 1981.and 1982, release of adult falcons was not implemented Ln 1983. LITERAWRE CITED Craig, G. R. 1982. Osprey nesting investigations, Job Prog. Rep., Colo. Div. Wildl., Wildl. Res. Rep., Jan., pp 147-153. �13 Prepared by ~ ~._6r · Gerald R. Craig ~ Wildlife Researcher C ��15 JOB PROGRESS REPORT a State of COLORADO Project No. N~I~R (SE~3~5) Work Plan No. 1 (II) Job No. 3 (3) Period Covered: Personnel: 1 March 1981 Peregrine Falcon Captive Maintenance 30 June 1982 W. Burnham, D. Konkel, C. Sandfort, E. Levine, G. Eitemiller, The Peregrine Fund, Inc.; G. Craig, Colorado Division of Wildlife. ABSTRACT In-1983, 35 adult peregrine falcons (Falco peregrinus anatum) were maintained at The Peregrine Fund, Inc's facilities at Fort Collins, Colorado. No losses or injury were incurred during the contract period. All falcons were maintained in robust condition and good health. aThis Job Report represents a preliminary analysis and is subject to change. Information presented herein 11AYNOT BE PUBLISHED OR QUOTED without permission of the Director, Colorado Division of Wildlife. ��17 PEREGRINE FALCON CAPTIVE MAINTENANCE William Burnham and Gerald R. Craig P. N. OBJECTIVE The objective of this program is to maintain a colony of adult peregrine falcons (Falco peregrinus ana tum) which is genetically representative of the wild population, free of disease, and capable of reproductive activity. SEG~NT OBJECTIVES 1. Annually contract with The Peregrine Fund, Inc. to maintain 35 adult anatum peregrine falcons in captivity. 2. Prepare an annual report of maintenance efforts associated with the contract. METHODS AND MATERIALS Methods and materials for this study have been described previously (Burnham and Craig 1981). RESULTS AND DISCUSSION In 1983, 35 adult ana tum peregrine falcons were maintained at the Fort Collins, Colorado facility of the Peregrine Fund, Inc. All falcons were fed a daily diet of Coturnix quail (Cotourix sp.) and 5-6 week-old cockerels. No adverse health conditions or disease problems were encountered and no losses or injury were incurred during the period. The security of the facility was not compromised nor did unusual events occur during this period. LITERATURE CITED Burnham, W., and G. R. Craig. 1981. Peregrine falcon captive maintenance. Job Prog. Rep ,, Colo. Div. Wildl., Wildl. Res. Rep., Jan., pp 34-37. Prepared by &4.hJd 11. ~ Gerald R. Craig ~ Wildlife Researcher C �Table 2. Osprey reproduction by site in Colorado, 1973-83. I-' 00 Site JA-1 JA-2 JA-3 JA-4 JA-5 JA-6 JA-7 LA-I LA-2 LA-3 GR-1 GR-2 GR-3 GR-4 GR-5 GR-6 GR-7 GR-8 GR-9 GR-10 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1/3/3a 1/7/0 1/1/0 1/1/0 1/1/2 1/1/2 1/1/0 1/1/0 1/7/1+ TO 2+/0/0 7/?/0 7/7/0 uo 1/1/1+ AB Uori AB TD AB UO ?/7/2 OC uo ?/1/2 ac ?/1/1 UO AB 0/0/0 7/2/1 AB UO 1/1/0 TO 1/0/0 UO 2/0/0 2/0/0 3/0/0 AB TO TO AB 2/1/1 UO OC TD 3/0/0 AB AB AB AB 2/0/0 ?/O/O 7/7/0 1+/0/0 3/0/0 3/1/1 uo 3/0/0 3/0/0 1/0/0 TO AB 3/0/0 UO AB AB 3/0/0 uo 2+/0/0 2+/0/0 TD OC AB AB 3/010 lJO UO 3/010 AB UO AB TD 3/2/2 AS 3/2/2 AB AB uo 3/0/0 AB 3/3/3 AB Uo TD 7/2/1 TO TO 5/0/0 AB 4/010 AB AB 3/0/0 4/0/0 AB 3/1/1 AB 3/0/0 UO 3/2/1 AB UO TO TO UO OC ua 1/0/0 2/0/0 TO UO AB 3/1/1 AB LA AB AB UO LA 3/0/0 3/3/3 AB 3/0/0 AB AB 2/a/fFUO AB 3/0/0 AB 3/0/0 3/3/3 TO TO TD AB AB TO 7/0/0c uo uo ac oc 1/1/2 1/1/0 1/0/0 2/0/0 2/0/0 ART 7/7/0 ART ART UO 1/0/0 1/0/0 OC 117/0 1/0/0 1/0/0 lJO UO 1/1/1 1/0/0 OC 7/0/0 UO 7/0/0 ua ART uo 1/0/0 ca-rt oc 7/0/0 TD 7/0/0 uo oc 1/7/0 TO GR-12 GR-13 GR-14 GR-15 GR-16 GR-17 GR-18 GR-19 GR-20 GR-21 GR-22 GR-23 sproduction AB 2+/0/0 1/0/0 0/0/0 2+/0/0 UO 0/0/0 uo code: bStatus codes: 1/3/3 OC a UO AB ART" TO • LA • G 1 eggs, 3 young hatched, 3 young fledged occupied breeding territory. unoccupied breeding territory. abandoned. artificial nest constructed. tree or nest blown down. lone adult present. cGR-20 failed and recycled at GR-7. UO 1/0/Q:,-2/0/0 2/010 �Table 2. Site JA-l JA-2 JA-3 JA-4 JA-5 JA-6 JA-7 LA-1 LA-2 LA-3 GR-l GR-2 GR-3 GR-4 GR-5 GR-6 GR-7 GR-8 GR-9 GR-10 GR-ll GR-12 GR-13 GR-14 GR-15 GR-16 GR-17 GR-18 GR-19 GR-20 GR-21 GR-22 GR-23 Osprey reproduction 1973 by site in Colorado, 1?74 1975 1976 1977 1978 7/3/3a 7/1/0 1/7/0 7/7/0 7/1/2 UO 7/1/2 7/1/0 7/1/0 1/?/1+ TD 2+/0/0 1/1/0 ?/?/O UO 1/1/1+ AB OC 7/7/0 aproduction ,UO OC 7/7/2 7/0/0 2/0/0 2/0/0 ART ?/?/O 1973-83. uo ART ART UO 1/0/0 7/0/0 OC 1/?/0 7/0/0 7/0/0 UO 1/?/1 7/0/0 OC 7/0/0 UO 1/0/0 UO ART uo 1/0/0 1979 UOIl AB TD AB UO 7/?/2 OC UO ?/?/2 OC 7/7/1 7/0/0 ?/7/0 1+/0/0 OC ?/O/O TD 7/0/0 Uo OC 7/?/0 TO 3/0/0 3/1/1 VO 3/0/0 3/0/0 ?/O/O TD AB 3/0/0 UO AB AB 3/0/0 UO 3/0/0 AB UO AB 1980 1981 1982 1983 UO AB 0/0/0 7/2/1 AB UO 717/0 TD 7/0/0 UO 2/0/0 2/0/0 3/0/0 AB TD TD AB 2/1/1 UO OC TD 3/0/0 AB AB AB AB 2/0/0 2+/0/0 TD 3/2/2 AB 3/2/2 AB AB uo 3/0/0 AB 3/3/3 AB Uo TO ?/2/l TO TD 5/0/0 AB 4/0/0 AB AB 3/0/0 4/0/0 AB 3/1/1 AB 3/0/0 UO . 3/2/1 AB UO TO TO UO OC UO 1/0/0 2/0/0 TD UO AB 3/1/1 AB LA AB AB UO LA 3/0/0 3/3/3 AB 3/0/0 AB AB 2+/0/0 TD OC AB AB 3/0/0 110 UO AB 2+/0/0 ?/O/O 0/0/0 2+/0/0 UO 0/0/0 uo code: bStatus codes: ?/3/3 Q ? eggs, 3 young hatched, 2/0/rFUO AB 3/0/0 AB 3/0/0 3/3/3 TO TO TO AB AB TO 7/0/0c UO 1/0/0;:-2/0/0 2/0/0 3 young fledged OC occupied breeding territory. UO - unoccupied breeding territory. AB abandoned. ART • artificial nest constructed. TO tree or nest blown down. LA • lone adult present. Q Q Q cGR-20 failed and recycled at GR-7. •.... \0 �Table 2. Osprey reproduction by site in Colorado, 1973-83. N 0 Site JA-l JA-2 JA-3 JA-4 JA-5 JA-6 JA-7 LA-l LA-2 LA-3 GR-1 GR-2 GR-3 GR-4 GR-5 GR-6 GR-7 GR-8 GR-9 GR-I0 GR-ll GR-12 GR-13 GR-14 GR-IS GR-16 OR-17 OR-IS GR-19 OR-20 GR-21 GR-22 GR-23 1973 1974 1975 1976 1977 1978 1/3/3a 7/7/0 7/7/0 1/7/0 1/1/2 7/7/2 ?/7/0 1/7/0 7/7/1+ TD 2+/0/0 7/7/0 7/7/0 1/7/1+ AB 1979 UOt> AB TD AB uo uo OC 7/7/0 7/7/2 7/0/0 2/0/0 2/0/0 ART 7/?/0 aproduction .OC uo ART ART UO 1/0/0 1/0/0 OC 7/?/0 7/0/0 7/0/0 UO 7/1/1 7/0/0 OC 1/0/0 UO ?/O/O UO ART uo 1/0/0 uo uo 7/7/2 OC UO 1/7/2 OC 7/1/1 7/0/0 7/7/0 1+/0/0 OC ?/O/O TD ?/O/O UO OC 7/7/0 TD 3/0/0 3/1/1 uo 3/0/0 3/0/0 7/0/0 TD AB 3/0/0 UO All AB 3/0/0 UO 3/0/0 AB UO AB 1980 1981 1982 1983 UO AB 0/0/0 ?/2/1 AB UO 7/7/0 TD 7/0/0 UO 2/0/0 2/0/0 3/0/0 AB TD TD AB 2/1/1 UO OC TD 3/0/0 AB AB AB AB 2/0/0 2+/0/0 2+/0/0 TO OC AB AB 3/0/0 UO UO TO 3/2/2 AB 3/2/2 AB AB UO 3/0/0 AB 3/3/3 AB TD 5/0/0 AB 4/0/0 AB AB 3iO/0 4/0/0 AB 3/1/1 AB 3/0/0 UO 3/2/1 AB UO TD TO DO OC TO UO AB 3/1/1 AB LA AB AB UO LA 3/0/0 3/3/3 AB 3/0/0 AB AB 2/0/OC UO AB 3/0/0 AB 3/0/0 3/3/3 TD TO TO AB AB TO 7/0/0c UO 1/0/O;c2/0/0 2/0/0 AB 2+/0/0 7/0/0 0/0/0 2+/0/0 uo TO 7/2/1 TD UO 0/0/0 uo UO 7/0/0 2/0/0 code: bStatus codes: 7/3/3 u 7 eggs, 3 young hatched, 3 young fledged OC = occupied breeding territory. UO = unoccupied breeding territory. AB '" abandoned. ART = artificial nest constructed. TD tree or nest blown down. LA ~ lone adult present. 0 cOR-20 failed and recycled at GR-7. �21 JOB PROGRESS REPORTa State of COLORADO Project No. N-I-R (W-124-R) Work Plan No. 2 (II) Job No. 1 (1) Period Covered: Personnel: Osprey Nesting Studies 1 July 1982 through 30 June 1983 E. Bowden, G. Claassen, G. Craig, G. Rosendale, S. Vana, and J. Wagner, Colorado Division of Wildlife. ABSTRACT Nest site occupancy by osprey (Pandion halieetus) in Colorado increased from 12 in 1982 to 14 in 1983. Eggs were laid at 8 sites but young were fledged at only 3 sites. Shell fragments were collected from 15 eggs and contents of 9 eggs were preserved for pesticide analysis. Observations from blinds continued, in an interrupted fashion, at 2 low disturbance and 2 high disturbance sites to determine effects of human activities upon reproduction. aThis Job Report represents a preliminary analysis and is subject to change. Information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the Director, Colorado Division of Wildlife. ��23 Osprey Nesting Studies Gerald R. Craig P. N. OBJECTIVES The objectives of this study are: (1) locate previously unknown nesting ospreys and monitor productivity of all sites, (2) determine causes of reproductive failure and develop methods to restore normal reproduction, .(3) determine nesting habitat requirements and implement protective measures to assure continued occupancy, and (4) compile data and prepare annual and final reports. SEGMENT OBJECTIVES 1a. Locate and map previously Colorado. unknown osprey nesting lb. All known nest sites will be visited reproductive success. 2a. Observe nesting pairs from concealment to identify behavioral abnormalities, weather conditions, predation, or human disturbance which might impact reproduction. Sites in remote localities will be compared with those adjacent to public use areas to determine responses to human activities. 2b. All unhatched eggs and shell fragments encountered during nest visitations described in 1b will be collected and submitted for pesticide analysis. Shell fragments will be measured and compared with pre-DDT era eggs for possible thinning. 3. Habitat features such as hunting areas, topography, vegetative type, climate, and geology will be recorded for each nest site to. establish habitat requirements. 4. Analyze annually sites throughout to establish data and prepare a report of the findings. METHODS AND MATERIALS Methods and Materials (Craig 1982). for this study have been described previoBsly RESULTS AND DISCUSSION In 1983, 14 sites were occupied, 2 of which were frequented by lone adults (Tables 1, 2). Eggs were produced at 8 sites occupied by adult pairs, but only 3 pairs (GR-2, GR-13, and JA-4) successfully produced �N ~ Table 1. Osprey reproduction in Colorado, 1973-83. Parameter 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 Total Total nests 5 8 16 16 17 17 15 16 16 17 16 159 Occupied nests 4 6 6 8 13 6 8 13 11 12 14 101 Active nests 4 5 6 8 13 6 8 10 9 10 12 91 Successful nests 0 3 0 2 2 3 2 1 5 2 3 23 Young hatched 0 7 0 3 3 4 2 2 10 4 7 42 Young fledged a 7 a 3 3 4 2 1 9 3 7 39 Young/occupied site 0.00 1.17 0.00 0.38 0.23 0.67 0.25 0.08 0.82 0.25 0.50 0.39 Young/active site 0.00 1.40 0.00 0.38 0.23 0.67 0.25 0.10 1.00 0.30 0.58 0.43 Youngisuccessful pair 0.00 2.33 0.00 1.50 1.50 1.33 1.00 1.00 1.80 1.50 2.33 1.70 �Table 2. Site JA-1 JA-2 JA-3 JA-4 JA-5 JA-6 JA-7 LA-l LA-2 LA-3 GR-1 GR-2 GR-3 GR-4 GR-5 GR-6 GR-7 GR-8 GR-9 GR-10 GR-ll GR-12 GR-13 GR-14 GR-15 GR-16 GR-17 GR-18 GR-19 GR-20 GR-21 GR-22 GR-23 Osprey reproduction 1973 7/7/0 1974 1975 1976 1977 1978 7/3/3a 7/7/0 7/7/0 7/7/0 7/7/2 uo 1/7/2 7/7/0 7/7/0 7/7/1+ AB OC uo OC 7/7/1+ TD 2+/0/0 7/7/0 7/7/0 Uo 7/7/2 7/0/0 2/0/0 2/0/0 ART 7/7/0 aproduction by site in Colorado, 1973-83. ART ART UO 1/0/0 7/0/0 OC 7/7/0 7/0/0 7/0/0 UO uo 7/7/1 7/0/0 OC 7/0/0 UO 7/0/0 UO ART uo 1/0/0 -, 1979 uob AB TD AB Uo 7/7/2 OC TIO 7/7/2 OC 7/7/1 7/0/0 7/7/0 1+/0/0 OC 7/0/0 TD 7/0/0 uo OC 7/7/0 TD 3/0/0 3/1/1 uo 3/0/0 3/0/0 7/0/0 TD AB 3/0/0 UO AB AB 3/0/0 UO 3/0/0 AB uo AB 1980 1981 1982 1983 uo AB 0/0/0 7/2/1 AB UO 7/7/0 TD 7/0/0 uo 2/0/0 2/0/0 3/0/0 AB TD TD AB 2/1/1 uo OC TD 3/0/0 AB AB AB AB 2/0/0 2+/0/0 2+/0/0 TD OC AB AB 3/0/0 uO' UO TD 3/2/2 AB 3/2/2 AB AB uo 3/0/0 AB 3/3/3 AB uo TD 7/2/1 TD TD 5/0/0 AB 4/0/0 AB AB 3/0/0 4/0/0 AB 3/1/1 AB 3/0/0 UO 3/2/1 AB UO TD TD UO OC UO 7/0/0 2/0/0 TD uo AB 3/1/1 AB LA AB AB UO LA 3/0/0 3/3/3 AB 3/0/0 AB AB 2/0/OC UO AB 3/0/0 AB 3/0/0 3/3/3 TD TD TD AB AB TD ? /O/Oc UO 1/0/0-2/0/0 2/0/0 AB 2+/0/0 7/0/0 0/0/0 2+/0/0 UO 0/0/0 uo code: bStatus codes: 7/3/3 = ---- 7 eggs, 3 young hatched, 3 young fledged OC = occupied breeding territory UO = unoccupied breeding territory AB = abandoned ART = artificial nest constructed TD = tree or nest blown down LA = lone adult present cGR-20 failed and recycled at GR-7 N V1 �26 young for an average of 0.30 young/active site. As in previous years, all nesting failures occurred during the incubation phase. Of the nests that failed, 4 were due to infertile or dead eggs, 2 nests blew down during high winds, and 2 were abandoned for undetel~ined causes. Pairs of ospreys at 2 sites recycled after their 1st nest failed. The pair nesting at GR-22 failed when the nest blew down on 13 May, prior to completion of the clutch. The pair rebuilt the nest in the same tree and initiated incubation of 2 eggs on 28 May. The 2nd clutch was abandoned between 9 and 11 June. The pair nesting at GR-20 abandoned their eggs on 19 May after 11 days of incubation. Cause of abandonment could not be ascertained, but human disturbance was unlikely since it was in a remote locality. The pair relocated 2 km to GR-7, which is considered a high disturbance location. and began incubating 2 eggs on 18 June. The renest attempt failed and 1 nearly full term dead egg was collected after 42 days of incubation. In 1983, 15 fractured or intact, nonviable eggs were collected and will be submitted to optical measurement for shell thickness. The 9 intact eggs were encountered in clutches at GR-1, GR-4. GR-5. GR-7, GR-10, and JA-4 and will be submitted for chlorinated hydrocarbon insecticide analysis. The Colorado Epidemiological Pesticide Studies Center at Colorado State University analyzed samples from 7 eggs collected between 1978 and 1982 and reported DDE values (uncorrected) from 0.10 to 36.07 ppm. These samples will be corrected for water loss for comparison with values obtained from the 1983 samples. Four sites were selected in 1983 for intensive observation from blinds to document the impact of human activities upon reproduction (Table 3). Sites GR-l and GR-2 were chosen as representative of high disturbance nests and were observed throughout incubation (160 hrs) until it was determined from prolonged incubation that the eggs were nonviable. Cause for failure could not be attributed to human disturbance. Low disturbance nests were represented by GR-10 and GR-20, but due to premature failures, GR-22 was substituted for GR-10 and GR-2 was substituted for GR-20. GR-2 was subsequently observed through hatch (40 hrs) , but GR-22 failed after only 16 hours of observation. Observation was then switched to GR-7 which was considered a high disturbance location and 40 hours of observation were accrued until the eggs failed to hatch. In no instances could direct human disturbance be attribu.ted to nesting failures. In anticipation of the final report, site specific information was recorded for each nest, meteorological data and lake ice conditions were collected, and information on recreational use was obtained. LITERATURE CITED Craig, G. R. 1982. Osprey nesting investigations. Job Prog. Rep., Colo. Div. Wildl., Wildl. Res. ep., Jan., pp 147-153. Prepared by ~d R. e..•~ Gerald R. Craig Wildli.fe Researcher C �Table 3. Intensive blind observation of nesting osprey, 1983. Disturbance level Initiation of observation Termination of observation Total hours GR-l High 14 May 19 lun 80 Observed throughout incubation; ended after collection of nonviable eggs. GR-4 High 15 May ·18 .Iun 80 Observed throughout incubation; ended after collection of nonviable eggs. GR-I0 Low 03 May 27 May 34 Observation ended with nest failure on 27 May. GR-20 Low 08 May 19 May 18 Observation ended with nest failure on !9 May. GR-22 Low 29 May 04 Jun 16 Observation ended with nest failure on 8 Jun. GR-2 Low 21 May 1:5Jun 40 Observed through incubation to hatch. GR-7 High 21 Jul 24 Jul 40 Observed until nesting failure. Total hours 308 .Site Comments N -....I ��29 JOB PROGRESS State of Project COLORADO No. N-1-R (W-136-R) Work Plan No. 3 (1) Job No. 1 (1) Period REPORT Covered: Personnel: 01 January Population and Mammal 1982 through Surveys Species 31 December of Selected Bird in Colorado 1982. R. Calderon, W. D. Graul, M. Kandel, G. C. Miller, T. E. Olson, and T. J. Schoenberg, Colorado Division of Wildlife. ABSTRACT Population and habitat data were collected from all known active and inactive great blue heron (Ardea herodias) and double-crested cormorant (Phalacrocorax auritus) nesting colonies in Colorado. The human-heron interaction study field work was completed during this segment. A final statewide survey of colony sites in 1983 is suggested. ��31 POPULATION SURVEYS OF SELECTED BIRD AND MAMMAL SPECIES IN COLORADO G. C. Miller P. N. OBJECTIVES 1. For all great blue heron and double-crested cormorant nesting colonies in Colorado, both active and inactive, document the following: (1) colony size and past population trends, and (2) distances to different forms of human development and activity. 2. For all active great blue heron and double-crested cormorant nesting colonies in Colorado document the anticipated future human development and activity patterns near the sites. and the number and condition of the trees being used as nest sites. 3. Document the reactions of nesting great blue herons and doublecrested cormorants to different types and quantities of human activities at nesting colonies. 4. Determine the availability of future alternate nest trees for the active great blue heron and double-crested cormorant colonies in Colorado. SEGMENT OBJECTIVES 1. For great blue heron and double-crested cormorant nesting colonies in Colorado, both active and inactive. document colony size and past population trends, and the distances to different forms of human development and activity. 2. For active great blue heron and double-crested·cormorant nesting colonies in Colorado, document the anticipated future human development and activity patterns near the sites. and the number and condition of the trees being used as nest sites. 3. Document the reactions of the nesting great blue herons and doublecrested cormorants to different types and quantities of human activities at 2 nesting colonies. METHODS AND MATERIALS Methods and materials have been described previously (Miller and Vos 1982). By the end of the reporting period, all known colony locations in Colorado had been visited. DESCRIPTION OF STUDY AREA Graul (1981) gave detailed description of study areas for the humanheron interaction portion of the study (P.N.O. #3). The statewide portion �32 of the study (P.N.O. fI's 1, 2, and 4) was conducted in the riparian and reservoir-riparian zones of the major drainages of Colorado and their tributaries. RESULTS During the reporting period, the status of most colonies in northeastern and northwestern Colorado were reported (Miller and Vos 1982). Between August and November 1982, habitat and population data were collected from 18 sites in Kit Carson, Prowers, Pueblo, Delta, Gunnison, Mesa, and Yuma counties. Reduction an.danalysis of these data were not accomplished during the reporting period. DISCUSSION Population and habitat data collection was accomplished for the statewide survey somewhat faster than originally estimated. In addition, there now exist for most colony sites in the state 2 or more population estimates separated in time from which some trends may be inferred. Present plans are to conduct 1 final population survey of colony sites just prior to the time when nest trees (primarily Populus. spp.) have enough green leaves to significantly obscure adult herons and cormorants. This activity is suggested for April-May, 1983. The human-heron interaction study field work was completed and pre Hrrdnary results reported during this reporting period (Miller and Vos 1982). The final results should be reported during the next (final) segment of this study. LITERATURE CITED Graul, W. D. 1981. Population surveys of selected bird and mammal species in Colorado. Job Prog. Rep., Colo. Div. Wildl., Wildl. Res. Rep. Jan. pp 83-129. Miller, G. C •• and D. Vas. 1982. Population surveys of bird and mammal species in Colorado. Job Prog. Rep ,, Colo. Di v , Wildl.. Wildl. Res. Rep. Jan. pp 73-107. Prepared by ~c-~~ caryc~ Miller Wildlife Researcher C �33 JOB PROGRESS State of COLORADO Project No. N-1-R Work Plan No. 4 Job No. 1 Period Covered: Personnel: REPORT (FW-145-R) 1 January Development of a Preservation For Insular Populations Program of Prairie Grouse 1982 through 31 December 1982 R. Calderon, W. D. Graul, S. B. Lustig, G. C. Miller, T. E. Olson, D. A. Opperman, T. J. Schoenberg, and B. van Sant, Colorado Division of Wildlife. ABSTRACT The study area was selected in January 1982. The area encompasses 650 km2 of Yuma County, Colorado, and contains areas inhabited and uninhabited by greater prairie-chickens (Tympanuchus cupido pinnatus). During calendar year 1982, data were collected relative to distribution, population trends, and habitat relationships. Assistance in the restoration of Tamarack Ranch as prairie-chicken habitat was given to the management branch of the Colorado Division of Wildlife. Data analyses are proceeding, and all objectives were addressed in this or previous reports. ��35 DEVELOPMENT OF A PRESERVATION PROGRAM FOR INSUJ~ POPULATIONS OF PRAIRIE GROUSE G. C. Miller P. N. OBJECTIVES 1. By 1986, ascertain for 1 species of prairie grouse in Colorado the combination(s) of habitat island size, condition, inter island distance, and number (by sex and age-class) of grouse needed to establish and maintain populations with a persistence probability of R_.::. 0.5 and extinction time of .!~ 100 years. SEGMENT OBJECTIVES 01 January-3D June 1982 1. By July 1982 locate, select, and describe occupied (by praar i,e grouse) and unoccupied prairie habitat blocks ("islands") most suitable for study of the biogeography and other aspects of prairie grouse ecology. 2. By July 1982, ascertain the nature of occupancy of selected pra1r1e islands by grouse and, where applicable, an index of abundance. 3. Begin to ascertain population characteristics of grouse in selected areas. 01 July-31 December 1982 4. Ascertain 1 (natality) , j.l(mortality),and.!:.values (intrinsic rates of growth) for study populations of greater prairie-chickens using radiotelemetry (20 radio-marked birds) and line-transect sampling. 5. Ascertain relationships between habitat conditions, pra1r1e relict size and arrangement, and greater prairie-chicken densities using line-transect sampling and Landsat vegetation classification. 6. Assist the management branch of the Colorado Division of Wildlife in applying research results to the greater prairie-chicken restoration efforts on the South Platte Wildlife Management Area (Tamarack Ranch). 7. Publish research results. METHODS AND MATERIALS Surveys to find leks in the intensive study area were conducted as described previously (Miller 1982). Surveys for leks in other portions of Yuma County were conducted by Northeast Region, CDOW personnel in 1981 and 1982. Methods and locations were given in an unpublished report �36 by B. van Sant in 1982, in the files of Northeast Region, CDOW. Prairie-chicken distribution in 1982 was delineated using lek presence as the primary criterion. Areas were classified as unoccupied by prairiechickens if leks were not present and if winter flocks or other evidence of prairie-chicken use was not found. Nearest-lek distances (NLD) were calculated for a random sample of 20 leks and 2 NLDmax was initially used to classify leks as belonging to the same or different populations. Data from this investigation were compared to the findings of Swope (1953), Evans (1964), and unpublished surveys from<>CDOW files (Appendices A, B. C). Population indices were computed from previous studies and compared to the 1981-82 lek surveys. Between December 1981 and continuing into this reporting period, techniques of trapping and radio-fitting greater prairie chickens were evaluated. The techniques included night--lighting, walk-in funnel traps. and cannon nets over bait and on leks. All birds captured were weighed, inspected for external characteristics of sex and age, and banded with aluminum leg bands. Birds not fitted with radio transmitters were banded with colored leg bands. Radio transmitters used- (model SPCB-1250-3X) were solarpowered units manufactured by Wildlife Materials, Inc. and weighed approximately 16 g without the attachment apparatus. Locations of radio-marked greater prairie--chickens were ascertained from directional readings of radio signals at predetermined "locator points" which were marked on the ground with vinyl flagging and recorded on 7.5 minute USGS topographic maps. Directional readings, taken with hand-held compasses (liquid-filled Suunto or Silva models), were recorded from a minimum of 3 locator points aridplotted on the topographic maps. The resultant polygon (usually a triangle) was considered usable for analysis if directional signals were recorded within a 1 hour period. DESCRIPTION OF AREA General study area descriptions have been provided previously (Miller 1981). The intensive study area for greater prairie-chickens, selected in January 1982, was in Yuma County in northeastern Colorado (Fig. 1). The study area encompasses that portion of Yuma County lying north of the Arikaree River, bounded on the west by State Highway 59 and on the north by County Road 26. The eastern boundary is irregular, delineated by County roads AA, Z, 20, Y, 23, and X. The study area is 650 km2 in area and has as its potential natural vegetation KUchler's (1964) sandsagebluestem (Artemisia-Androp~~!!_) prairie type. RESULTS Greater Prairie-chicken Distribution A distribution map for grea t er+p'r a Lr Le chickens in Yuma. County was generated using the maximum nearest-lek distance observed, 4 km, as the criterion for population continuity (Fig. 2). The intensive study area �37 I J I YUMA Vernon Hea r ts t rong CO. J# STUDY AREA LOCATION MAP I I I I * 0 I Abarr I (j) I() * W l- => 0 a:: w le:( ICI) U.S. ROUTE 36 Joes YUMA o I I I I I L- I 2 3 4 511,IILES I I II II I I 0 I :5 4 5 KILOMETERS 2 COUNTY COLORADO ....l__j --------- ------ Fig. 1. Greater prairie-chicken intensive study area, Yuma County, Colorado. Stars indicate active lek locations in 1981. �38 GREATER DISTRI BUTION COLORADO 1982 PRAIRI E~CHICKEN YUMA COUNTY, -@-j 1--- 0' LOCATION MAP t2a 1982 D 1952 , a4 '1 - r I I J /"' I Sandhills boundary J U.S. 34 (J I viI . I I J I I I I I I W I fo,<{ f0r./) I I ·1 J U. S. 36 u S. 36 Joes I Bonny Res. I '------_._._--Fig. 2. Colorado, Greater 1952-82. pra Lr Le=ch Lcken distribution, --- Yuma County, -~- J �39 was 100% surveyed and negative as well as positive data were recorded. Most of the remaining sandhill areas were also surveyed (B. van Sant, unpubl. rep., Colo. Div. Wildl., NE Reg. off., Fort Col~ins, 1982). The distribution of greater prairie-chicken in 1982 differed from that shown by Swope (1953). He identified 20 areas which, in 1952, contained " ••• some of the best prairie chicken habitat ..." in Yuma County (Swope 1953: Fig. 3). The distribution map generated in 1982 showed 6 (30%) of those areas as no longer within the occupied range of prairie-chickens. Another 6 '(30%)had less than 50% of their former extent classified as occupied by prairie-chickens (Fig. 2). The greater prairie-chicken distribution in 1981-82 also differed from that observed by Evans (1964) in 1962-63. Classification of 44 lek locations from 1962-63 revealed that 20 (45%) were not within areas classified as occupied in 1981-82 (Table 1). Twenty-four locations were within 1981-82 occupied range. Of the 24 locations (quarter sections) still within occupied range, 23 were evaluated with respect to 1981-82 lek locations. Leks were in approximately the same locations in 1981-82 as they were in 1962-63 in 7 of the-23 quarter sections (See Appendix D). Population indices based upon number of leks per linear unit of search were computed for 1962-63, 1972-73, 1975-77, and 1981 (Table 2). Data from 1982 surveys were not analyzed in this fashion during the reporting period. A total of 42 sections (l09 km2) of occupied range in northern Yuma County, selected in a stratified random sampling plan, was 100% surveyed for leks in 1976-77 and 1981-82. A summary of the raw data is provided in Table 3, but a critical evaluation of accuracy was not performed. In the intensive study area, in southern Yuma County, lek densities were computed fot 2 apparently disjunct populations. Preliminary computations of lek densities in 1982 were 0.40 leks/mi2 (0.15 leks/ km2) and 0.25 leks/mi2 (0.10 leks/km2). Habit?_~?-nd Prairie-Chicken Relationships Vegetation characteristics for 6 nest sites located in 1982 were summarized Table 4) as were brood-rearing sites (Table 5). Reduction of macrohabitat data from 38 brood sites was not accomplished during the reporting period. Tamarack Ranch Activities Restoration of the Tamarack grassland was begun by NE Region, cnow personnel in April 1982. Two grass seed mixture recommendations were provided at that time for interseeding 2 types of areas, based on topgraphic features (Table 6). Seeding rates were modified from previous recommendations (Miller 1980) to accommodate altered species composition and possible reduction of seed viability from prolonged storage. Evaluation of seeding success in autumn 1982 was performed by Regional personnel and are not presented here. More complete descriptions of techniques w re reported previously (Miller 1980). �40 GREATER PRAIRIE-CHICKEN YUMA COUNTY, ,--- LJ'I~ LOCATION I MAP I I Sandhills boundary I I ~ LV U.S. 34 I I I ,I I en I I{) I I I I I u S. I 36 U.S. 36 Joes I ..__ - --- ----- I Bonny Res. ---- J Fig. 3. Greater prairie-chicken dist:r:ibutions. Yuma County, Colorado (after Swope 1953). Negative data were not reported; lack of stippling does not necessarily indicate unoccupied habitat. �41 Table 1. Greater prairie-chicken distribution in 1981-82 and 1962-63 (Evans 1964) .s! N % 1962-63 lek locations within 1981-82 range 24 54.5 1962-63 lek locations outside 1981-82 range 20 45.5 ~/ Data are in Appendix D. Table 2. Number of greater prairie-chicken leks per mile (per km) of search, Yuma and Phillips counties, Colorado, 1962-81. Year N leks found Miles (km) searched Leks/mile (km) Source 1962 25 128 (205) 0.20 (0.13) Evans (1964) 1963 27 128 (205) 0.21 (0.13) Evans (1964) 1972 39 186 (298) 0.21 (0.13) Tullya/ Appendix A 1973 28 179 (286) 0.16 (0.10) Tullya/ Appendix A 1975 10 131 (210) 0.08 (0.05) Graulb/ Appendix B 1976 11 25.5 (41) 0.43 (0.27) GraulS' Appendix C 1977 2 25.5 (41) 0.08 (0.05) Nongame Research files 1981-~/ 38 186 (298) 0.20 (0.13) van Sant, unpubl. ~/ R. J. Tully, unpubl. rep., Colo. Div. Wildl., Nongame Res. Off., Fort Collins, 1974. E_/ W. D. Graul, unpubl. rep., Colo. Div. Hildl., Nongame Res. Off., Fort Collins, 1975. s:./ W. D. Graul, unpubl. rep., Colo. Div. Wildl., Nongame Res. Off. ~ Fort Collins, 1976. E._/ Computed on the bas:is of leks found wihtin 3.2 km (2 miles) of routes surveyed in 1972. �42 Table 3. Comparison of 100% survey of 42 sections of occupied greater prairie-chicken range in northern Yuma County, Colorado, 1976-77 and 1981-82. 1976-77 1981-82 Sections with leks 17 (40.5%) 15 (35.7%) Number of leks detected 22 16 Number of leks reported in 1976-77 that were not present in 1981-82 16 11 Number of leks not reported in 1976-77 that were present in 1981-82 Table 4. Vegetation characteristics of 6 greater prairie-chicken nest sites, Yuma County, Colorado, 1982. Nest sites (N=6) range x Tallgrass & midgrass Ht, cm 111m2 Shrub Ht, cm 111m2 Bare Ground, % Alternate sites (_!!=24) x 66.9 16.7 30-104 13-20 68.0 15.3 45.5 1.8 26-63 1-4 44.4 0.7 15.0 5-30 32.9 �43 Table 5. Characteristics of greater prairie-chicken brood-rearing sites, (roosts), Yuma County, Colorado, 1982. x SD (N=25) 58.1 6.1 24.5 5.8 69.0 13.0 22.4 6.6 45.6 0.8 14.0 1.3 Bare Ground, % 31.1 17.8 Aspect, Azo 199.6 102.7 Slope 2.0 1.9 Topography, II sites Valley Slope Ridge Unclassified 20 2 Characteristics Tallgrass Ht, cm 111m2 Tallgrass-midgrass combined Ht, em 111m2 Shrub Ht, cm 111m2 2 1 �.p. .p. Table 6. Seeding recommendations provided to NE Region, April, 1982..ill Topographic site Lowest; valleys, basins Lower side slopes and "flats" cnow personnel for the Tamarack Ranch~ Pure live seed rate (lbs/Ac ) Bulk rate (lbs/Ac) Percent of full rate Total lbs. to be seeded (est.) Indiangrass (Sorghastrum nutans) 4.0 6,8 40 320 Big bluestem (Andropogon _gerardi) 2.0 4.0 18 150 Sand bluestem (~. hallii) 1.3 4.7 8 106 Switchgrass (Panicum virgatum) 1.6 1.8 33 128 Indiangrass Switchgrass Sand Bluestem 3.0 3.8 3.5 5.1 4.1 12.6 30 85 23 232 301 270 Species a/ The Lnterseeder was cali.brated to deposit 80 seeds per foot to ensure at least 40 live seecis--perfoot (1.3 live seeds!cm) , Rates expressed in English units for convenience in purchase and seeder calibrations. �45 DISCUSSION Distribution of greater prairie-chicken has changed since 1952. The lack of negative data from 1952 (Swope 1953) and 1962-63 (Evans 1964) allows no unequivocal statement about net changes in distribution. However, the loss of prairie-chickens from many areas earlier considered as some of the best habitat in the state must be viewed with concern. Within presently occupied range, indices of abundance are lower than similar indices in other states. The leks/mile indices in Colorado ranged be.tween 0.43 and 0.. 08 while similar indices in Ka.nsaswere 0.73 and 0.76 leks/mile in 1978 and 1979 (Wells 1980). Preliminary computations of lek densities in the intensive study area of 0.25 and 0.40 leks/mi2 were similar to those in Nebraska. (0.24 leks/mi2) (Sisson 1976), B. van Sant (unpubl. rep., Colo. Div. WHdl., NE Reg. off., Fort Collins, 1981) reported an overall index for Yuma County as 0.11 leks/mi2. LITERATURE CITED Evans, K. E. 1964. Habitat evaluation of the greater prairie chicken in Colorado. M. S. Thesis, Colo. State Univ., Fort Collins, 98pp. Kuchler, A. W. 1964. Potential natural vegetation of the conterminous United States. Am. Geogr. Soc. Spec. Publ. 36. 39pp. Miller, G. C. 1980. Development of a preservation program for three species of praf.ri e grouse, Job Frog. Rep ,, Colo. Div. Wildl., Wildl. Res. Rep. Jan. pp. 76-94. 1981. Development of a preservation program for three species of prairie ·grouse. Job Pr'og, Rep., Colo. Div. Wildl., Wildl. Res. Rep. Jan. pp. 38-52. 1982. Development of a preservation program for insular populations of pra i.r ae grouse . Job Prog. Rep. ~ Colo. Div , Wildl., Wildl. Res. Rep. Jan. pp. 63-71. Sisson, L. 1976. The sharp·-tailedgrouse in Nebraska. Parks Comm ,, Lincoln. 88pp. Nebr. Game and Swope, H. M. 1953. Surveys to determine the population status of the pralrle chicken. Job Completion Rep., Colo. Dep. Game and Fish, Proj. W-37-R-6. pp. T/-80. Wells, R. 1980. Kansas praa.r r.echicken annual report, 1979. Kans. Fish and Game Comm , Fed. Aid Proj. W-23-·R-lS, 36pp. ~lXl~~_ Prepared by ~~ ~ ~Miller vJildlife Researcher C ��47 APPENDIX A SPRING STATUS OF THE GREATER PRAIRIE CHICKEN AND OTHER. OBSERVATIONS IN THE NORTHER...~ SANDHILLS OF YUMA COUNTY, COLORADO by Robert J Tully 0 Wildlife Management Section (Colorado Division of Wildlife) March, 197~ ", Results of pra~r~e chicken counts on booming grounds conducted during mid-April of 1973 compared to previous studies indicate the spring breeding population was similar to that of 1972 but significantly lower than in either 1962 or 19520 1952 1962 1963 1967 1971 1972 1973 Ave. No. Birds/ Ground 26.5 6.1 604 408 4.0 5~9 6.7 Range of Birds/ Ground 8-73 2-28 1-50 1-10 1-7 1-18 2-23 20 21 26 9 13 39 28 600 600 No. of Grounds Counted Total Population Estimate 2,000 700-800 200-300 The loss of tal1 grass is probably the most important factor in the long term population decline. Habitat research in the 1960's "indicated the birds became scarce as the percentage of tall grass and shrubs decreased and the percentage of bare ground increased. The birds remain most abundant where about two thirds or more of the area is in native grass. Changes in land use since the peak of population abundance are refl.ectedin the following 50 year summary for Yuma county. Noo Irrigated Acres Ave. Acres/Farm No. Irrigation Wells No. of Cattle 1930 1940 1950 1960 1970 3,620 1,550 1,550 11,130 110,560 560 816 927 1,286 2,500 0 3 7 40 882 24,600 15,500 30,000 58,700 137,000 Research and occasional studies over approxiw2tely 660 square miles of sandhillssandsage grasslands interspersed with cropland, indicate that without intensive habitat management the greater prairie chicken may ~ecome extirpated from its only Colorado stronghold during this centurj. The prairie chicken does occur elsewhere in the county and in smaller numbers in several other counties. Primary needs for populaticil enhancement are: (1) Improved management of about 18 square miles of state school lands, (2) modi.fication of federal farm program �48 practices, (3) habitat improvement of federal, state and private lands, (4) acquisition of key lands which can be managed primarily for wildlife production, (5) development of vigorous subclimax grass-forb communities on retired croplands and native prairies interspersed with grain fields and shrub communities and (6) continuing population and habitat inventory and analysis. Breeding ground and habitat research related to the pra~r~e chicken ~ms accomplished in 1952, 1962 and 1963 by the Colorado Division of Wildlife, Harold Swope and Keith Evans, through Federal Aid in Wildlife Res t ora t Lon Projects and is reported in various progress reports, technical papers and a Master of Science thesis from Colorado State University. Occassional studies "Were conducted from the early 1950 I s through 1971 by various Division of Wildlife and Co Lorado State University personnel however no special reports were prepared. Historical information on the abundance of prairie chickens and changing habitat conditions was reviewed by the author in 1971 when photographs were taken at various locations in.northern Yuma. county. Results were r.epor.tedthrough slide presentation at the 9th Prairie Grouse Technical Council, 1971 and at other meetings. With the assistance of several conservation organizati.ons and many individuals, distribution and density counts were completed during April 15 and 16, 1972 and April 14 and 15, 1973. Following an informal introduction and instruction meeting at the Yuma County Courthouse, lvray the eveni.ng before each years census ac t Lvi.t g participants took to the field before daylight to seek pleasure. in observing various wildlife within native gra sland and farmland habitats. The census was carried out during the early mOn1ing and late afternoon activity periods by slowly driving assigned mapped routes which showed previously recorded prairie grouse booming grounds. Participants stopped about every quarter mile and at all marked locations to observe and listen. Most grouse were located by the booming sound which sometimes carries over one mile. Pra.irie grouse on both historic and newly located grounds were recorded as well as were all wildlife species observed within an area of nearly 700 square miles, generally north of Wray. At approximately the peak of courtship activity during 197~233 prairie chicken were observed on 39 booming grounds by 54 persons. Sign or sound indicated birds were also present at 19 additional sites. During 1973, 36 ob~ervers located 186 birds on 28 booming grounds and heard prairie grouse at 14 addi.tional sites. Birds were also seen flying at several locations in both years. Through careful observation at various locations it was estimated that not over 17 percent of all birds were females. All duplicate observati.ons were eliminated by mapping the location and number of birds seen at various times. During the two years recorded. Observers and 57 bird species, mammals, totaling 94 of investigations 71 bird species and 9 mammal species were in 1972 documented 7 mammal species totaling 148 individuals totaling 4,679 individuals. Observers in 1973 recorded 8 individuals and 51 bird species, totaling 2,515 individuals. - 2 - �49 Contributors: Persons who assisted represented the Colorado Division of Wildlife, Aiken Audubon Society, Boulder Audubon Society, Colorado Field Ornithologists, Colorado State University, Denver Audubon Society, Denver Field Ornithologists and Foothills Audubon Club. Hy sincere appreciation is extended to all who participated in the field or otherwise assisted in making the counts possible and my apologies to those who are not named. April 15 and 16, 1972: Mr. and Mrs. E. Ausfahl, Del Benson, Steve Brock, Joe Brailey, Christine Bonny, Merle Barbour, Henry and Vi Bossman, ~~rtha Bildstein, Sophia Bogart, Dave Clippinger, Jerry Craig, Allegra Collister, Camille Cummings. John and Joyce Cooper, Lori Chappell, Ron Desilet, Don Dominick, Glen Eyre. Wm Ewing, Tony and Al Esposito, Bill and Patty Echelmeyer, Dale Hein, Bob and Yvonne Gibbons. Thomas and Mrs. Henry. Gayle Ireland, Al Koewing, Ron Les t Lna , Thompson and Susan Harsh. Fran Marcoux, Sue Merrick, Marge Hdlilliams, Susan Reese, Jim Roscoe, Warren Rupke, Ron Ryder, Frank Scarpella, Warren and Mrs. Snyder, Dale Stahlecker, Joe and St eve Tully, Helen Thurlow, Harry and Elinor Wills, Jack and Grace Ivelsh and Jerry Wolfe. Weather -- On Saturday the minimum temperature was 34° and maximum temperature 0 was 56 • The wind was O-lOmph and the skies were clear to 10 percent partly cloudy. It was an excellent observation day. Sunday was completely overcast and windy, 10-30mph, with a trace of precipitation between 7:30 and 9:30am. Minimum temperature of 360 and maximum of 760 and counting conditions were poor. April 14 and 15, 1973: Ed and Irene Arenson, Christine Bonney, Lou Brevard. Bonnie Butzman, Bob Brunson, Marilyn Cook, Jerry Craig, Dennis Davis, Jim Dennis. Gay Eckes, Heather Flanagan, Bill Gillespi, Bill Goslin, Dale Hein, Francis Hames , Stephan Henry. Dave Ha t t an , Rick Krasa , Al and Jane Koewi.ng , Hike Lf.ns haw , Dan NcKeon, Dave NcCargo, Al Morgan, Bruce McCloskey, Bill Reneau, Dale Stahlecker, Warren and Mrs. Snyder, Frank Scarpella, Charlie Summ.ers, John Torres, Joe and Steve Tully, Jack Wasserbach and John Heins. Weather -- On Saturday it was clear with less than 20 percent cloud cover and wind very slight, never exceeding l5mph. Minimum temperature of sf and a maximum temperature of 80° . Observation conditions were excellent throughout the day. On Sunday it was completely overcast, very windy and raining heavy in most locations. Counts were not attempted by most observers. - 3 - �50 Birds and Hammals Observed in the Vicinity of 'Wray, Colorado, April, Number Recorded ....... Pied-billed Grieb Night Heron Black-crowned Canada Goose Snow Goose Mallard Gadwall 7 6 1 250 16 24 6 Pintail Greeno·winged Teal Blue~winged Teal Cinnamon Tea 1 Ame ri can Hidgeon Shoveler Lesser Scaup Red-tailed Hawk Swainson s Hawk Rough-legged Hawk Ferruginous Hawk Golden Eagle f Mar.sh Hawk Sparrow Hawk Grea.ter Prairie Chicken Cambe 1 f 1/ . 1972- ~eci.es Qua i 1 R~ng-necked Pheasant S 37 14 20 5 14 5 2 22 3 1 2 3 48 233 69 7 2 2 6 69 100+ 5 6 4 1 25 99 186 20 2 348 American Coot 14 Ki.l1.deer Common Snipe 19 11 1 Long-bi lled Cur Lew 11 4 12 Up Land Plover Baird's Sandpiper Western Sandpiper Marbled Codwi.t Wilson's Phalarope Frank lin s Gull 1 10 1 3 1 I Rock Dove 12 12 42 6 7 19 Hourning Dove Barn ()wl Grea.t Horned Owl Burrowing Owl Short-eared 2 O-wl 16 1 Belted Kingfisher Yellow-shafted Flicker 6 Red-shafted Flicker 9 Hairy Woodpecker Downy Woodpecker Say I s Phoebe Horned Lark 1 3 13 1 1 893 11 375+ 1972 and 1973. �51 Birds and Ma~~a1s Observed in the Vicinity of Wray, CO., April, 1972 and 1973. (Cont.) Number Recorded " , ~cies Barn Swallow Cliff Swallow Black-billed Magpie White-necked Raven Common Crow Robin Townsend's Solitaire Northern Shrike Loggerhead Shrike Starling Audubon's Warbler House Sparrow Western Meadowlark Yellow-headed Blackbird Red-winged Blackbird Brewer's Blackbird Common Grackle Brown-headed Cowbird Vesper Sparrow Gray-headed Junco Chipping Sparrow Brewer's Sparrow White-crowued Sparrow ~h~sEn~t:c~l!a~e~ ~o~g~p~r_ Unidentified Small Rodents Kangaroo Rat Unidentified Vole White-tailed Ja~krabbit Black-tailed Jackrabbit Cottontail Rabbit Striped Skunk Badger Coyote Pronghorn Antelope Mule Deer , 'll' 1972- 1973'!:/ 3 8 114 2 69 11 35 9 22 22 2 18 86 44 11 10~ 1 60 2»001 4 182 217 87 22 68 - ~O+ 675+ 8 75 8 10 9 28 1 11 1 3 24 8 45 2 11 75 3 1 1 5 5 1 13 35 11 31 12 1/ Total includes 1 unknown junco, 2 unknown sandpiper and 2 unknown broadwinged hawks. 2) Total includes 2 unknown ducks and 2 unknown falcons (probably prairie fa lcon) . Those who participated in both 1972 and 1973 agreed that Horned Larks were less abundant and Western }!eadowlarks more abundant than in 1972. ��53 APPENDIX B SUMMARY OF 1975 Greater Prairie Chicken Census - Yuma County On AprilS and 6, 1975, thirty-two people (members of The Division of Wildlife, Foothills Audubon Society, C.S.U., U.N.C.) participated in the annual census of the Greater Prairie Chicken. Ten cars were used on Saturday morning and each car was assigned a mapped route. The census was conducted by stopping at one mile intervals to listen for birds. Booming grounds were mapped from these sound records (they can be heard for over one mile). Additionally, birds were counted on some grounds. The morning count went from daybreak (5:45 am) to about 8:30 am - Daylight Savings Time. Selected routes (6) were checked Saturday evening from about 5:30 pm until 7:45 pm (dark) with the emphasis placed on counting birds on known grounds. On Sunday morning, two groups were formed and each group visited a readily accessible ground so that everyone could see birw"booming. On Saturday, the temperature started in the low thirties and rose into the sixties. The wind was 0 - 10 m.p.h. and skies were clear - an excellent observation day. On Sunday, the temperature started in the low thirties and rose into the fifties, but s me areas had dense ground fog in the morning, making observati n difficult. Ninety-five birds were observed with 69 of these b~ing on grounds (6.9 birds per ground). An additional 19 grounds were mapped from hearing the birds. Assuming the 6.9 average for all grounds, we have a total estimate of 200. By assuming that 80% of the birds on the grounds were males and assuming a 50-50 sex ratio, we estimate 360 birds in the area covered by our routes. By assuming that we could hear birds for l~ miles, our routes covered 75% of the total nesting area. The total estimate for the Yuma County population, therefore, comes to 480 birds. The same population was estimated to be about 600 birds in'1972 and 1973. Since the 1975 census was conducted relatively early, it is highly possible that 600 birds are still present. I will he checking known grounds later this month to verify this. Thus, we can summarize the overall population trend for this species in Colorado. At one time, the Greater Prairie Chicken nested over a 15 county area in Colorado. In 1952, the "population was limited to Yuma County - about 2000 birds. As the land has steadily been plowed, the population has declined. In 1962 the population estimate was 700-800 birds. By 1967, it had dropped to 200-300 birds, but by 1972 it had ,climbed back up to around 600 birds and it appears to be remaining relatively stable. All of these later figures are probably conservative. Walter D. Graul Nongame Bird Specialist �54 Birds and Mammals Observed Near Wray. Colorado. April. 1972. 1973. 1975 Species 1972 Horned Grebe Eared Grebe Pied-billed Grebe White Pelican Double-crested Cormorant Great Blue Heron Black-crowned Night Heron Canada Goose Snow Goose Mallard Gadwall Pintail Green-winged Teal Blue-winged Teal Cinnamon Teal American Widgeon Northern Shoveler Wood Duck Redhead Canvasback Lesser Scaup Common Goldeneye Bufflehead Ruddy Duck Common Merg8;nser Red-breasted Merganser Turkey Vulture Cooper's Hawk Red-tailed Hawk Swainson s Hawk Rough-legged Hawk Ferruginous Hawk Marsh Hawk Golden Eagle Prairie Falcon American Kestrel Greater Prairie Chicken Bobwhite Gambel 's Quail Ring-necked Pheasant American Coot Killdeer Common Snipe Long-billed Curlew Upland Sandpiper I Number Recorded 1973 1975 4 1 4 7 11 12 9 6 1 16 6 14 20 5 14 5 250 24 37 7 2 2 6 69 10(}t- 2 5 22 3 1 4 3 25 2 1 1+8 99 186 20 233 69 2 348 14 19 6 4'+ 11 76 310 19 44 46 10 1 0 36 100 6 40 31 96 31 36 12 50 3 6 1 10 2 2 5 22 1 56 95 63 13 19 1 11 12 4 1 2 �55 Birds and Mammals Observed Near Wray. Colorado, April. 1972. 1973. 1975 Species Baird's Sandpiper Western Sandpiper Marbled Godwit Wilson's Phalarope Herring Gull Ring-billed Gull Franklin's Gull Rock Dove Mourning Dove Barn Owl Great Horned Owl Burrowing Owl Short-eared Owl Belted Kingfisher Common Flicker Hairy Woodpecker DOwny Woodpecker Say I s Phoebe Horned Lark Barn Swallow Cliff Swallow Black-billed Magpie White-necked Raven Connnon Crow Black-capped Chickadee American Robin Townsend's Solitaire Mountain Bluebird Ruby-crowned Kinglet Northern Shrike Loggerhead Shrike Starling Audubon's Warbler House Sparrow Western Meadowlard Yellow-headed Blackbird Red-wi.nged Blackbird Brewer's Blackbird Common Grackle Brown-headed Cowbird American Goldfinch Vesper Sparrow Gray-headed Junco Chipping Sparrow Brewer's Sparrow Harris' Sparrow White-crowned Sparrow 1972 Number Recorded 1973 1975 1 10 1 3 12 6 7 6 9 1 12 42 2 19 16 1 16 10 300 20 4 30 8 3 30 1 1 893 3 8 114 2 69 11 18 86 44 11 375+ 5 585+ 35 9 22 25 22 2 11 100+ 3 16 8 250 6 1 1 8 11 1 60 2,001 4 182 217 87 22 68 50+ 675+ 8 75 8 10 9 22 1,085 88 2 8 10 1 28 1 11 4 1 3 3 4 �56 <. Birds and Mammals Observed Near Wray. Colorado. April, 1972. 1973, 1975 SJ2_ecies 9~~~~~~~:~~!!~E~~_~~~8~e~E Unidentified Small Rodents Kangaroo Rat Vole White-tailed Jackrabbit Black-tailed Jackrabbit Cottontail Rabbit Striped Skunk Badger Coyote Pronghorn Antelope Mule Deer Fox Squirrel 1972 ?~ 8 2 Number Recorded 1973 : 1975 ~~ 45 1 1 11 75 3 1 1 5 5 1 1 1 13 11 35 31 13 13 12 3 �57 APPENDIX C STATE OF COLORADO IIcMrd D. !.amlil. Governor DEPARTMENT OF NATURAL RESOURCES DIVISION OF WILDLIFE .Jack R. Gr'.b, D'rector 6060 Broadway Denver. Colorado 80216 (825-1192) Greater Prairie Chicken 1976 Booming Ground Survey Historically, we have relied on roadside counts, utilizing volunteers, to estimate the Colorado spring population of Greater Prairie Chickens. Such counts have documented a long-term decline in the population, but have not been adequate for calculating an accurate total population figure. Rough estimates, however, have been made. For instance, the total population in 1952 was estimated to be 2,000 birds. From 1972-75 the estimate has remained around 600 birds - all located in the northern segment of Yuma County. In 1976 the census procedure was changed. The overall known breeding range was divided into two strata. The interior strata. (300 sections) covered the area where most birds occur. The exterior strata (268 sections) represent the peripheral boundary of the known population and undoubtably does not have as many birds as the interior strata. This year 15 study areas, each consisting of two square sections, w re randomly chosen in the interior strata. Five WCO's then made on the ground counts of the 15 areas - 3 areas per WCO. All counts were conducted between April 13 and April 25. On each count, a route was followed such that the observer was at some point within one-quarter mile of·any part of the study area. Counts were not conducted in inclement weather and all were made with the wind less than 15 m.p.h. The raw data are shown on the attached chart. As expected, the chickens are not randomly distributed over the area. In fact, 6 or the 15 areas accounted for all the chickens recorded. One interesting aspect of the surVey is that 11 of the 30 sections chosen randomly had irrigation systems on them 36.6%. This is a sur risingly high figure considering the rough terrain of this sandhill region. Clayton Wetherill wins the prize fo!'censusing the most irrigation systems - 8 irrigation circles. Using this survey to estimate the total population must be done cautiously. There are several variables that could drastically influence such an estimate. This survey, however, can yield a much better estimate than.we have had previously. On a gross level, 74 chickens were counted. Since the survey covered 10% of the 300 sections in the interior strata, this would yield a minimum estimate for the interior strata of 740 birds. Such an estimate, however, must be considered extremely conservative. For instance, we know that for a given booming ground there can be from 30% - 100% of the males using the ground present at any given time. Additionally, we know that on a given DEPARTMENT OF NATURAL RESOURCES.Harris Sherman. Executive Director Thomas Far! Y. Vice Chairman • S m Caudill, ecretary 0 WILDLIFE COMMISSION, Vernon C. Williams, Chairman • Jean K. Tool, Member'" Roger Clark, Member �58 - 2 - ground at a given time there will not be nearly as many females as males. Furthermore, we cannot assume a 50-50 sex ratio. For instance, in Sage Grouse there are about 70 females for every 30 males in the population the result apparently of high male mortality. If we assume that: (1) 80% of the males in the area were present on the grounds, (2) 80% of the birds on the grounds were males, and (3) there are 4 females for every 3 males, we would have a population estimate of 1657 chickens for the interior strata. Of course, we also know that at least some chickens are in the exterior strata. So, about all we can say at this time is that we have at least 740 birds, but we may have 2000+ birds. I realize that such an estimate range is vague, but I do not want to mislead anyone with unjustified "exact" figures. We will need more data to refine our estimates. One thing is certai.n - our past estimates have been low. Another figure that can provide valuable trend information is the number of birds per ground. This year it is 6.55 birds per ground with a range of 2-15. The following table shows how this compares to past years. Yuma Co. Greater Prairie Chicken Booming Ground Counts 1952 1962 1963 1967 1971 1972 1975 1976 BirdS/Ground 26.5 6.1 6.4 4.8 4.0 5.9 6.9 6.5 Range/Ground 8-73 2-28 1-50 1-10 1-7 1-18 3-23 2-15 In conclusion, I would like to thank everyone that participated in the census this year. With this type of cooperation, I am hopeful that we can get a handle on the chicken situation and start designing some solutions for the overall problem. Sincerely, Walter D. Graul Nongame Bird Specialist WDG/kd �., -, 59 : Greater Prairie Chicken Survey Route 11= Observer - 1976 Birds on _:SoomingGrounds Birds flushed-Not on Grounds Other Wildlife Recorded 1 Rupke 0 0 0 2 Wetherill 0 0 0 3 Rupke 0 0 0 4 Wetherill 0 0 0 5 Rupke 0 0 0 6 Wetherill 5 0 0 7 Budde 0 0 8 Richardson 6,15; Heard on area 0 0 9 Scarpella 6;8; Heard on area 0 0 10 Scarpella 4', Heard on area 0 0 11 Scarpella 9', 7', 5 0 0 12 Budde 0 0 0 13 Richardson 0 0 14 Budde 15 Richardson Totals 2, 5 2 0 0 72 on 11 grounds 3 grounds not located 2 8 Antelope 11 Antelope 2 Coyotes 0 5 Coyotes 2 Jackrabbits �61 APPENDIX D Greater prairie-chicken lek locations in 1962-63 (Evans 1964) and 1981-82, Northeast Colorado. 1962-63 lek location Range Township Section Quarter 42W 3N 2N 43W 5N 4N 3N 2N 44W 4N 3N 2N 45W 4N Within (+) or outside (-) 198182 occupied range Lek present in 1962-63 and 1981-82 Yes 7 31 33 4 4 21 SW SW SE NE SE SE 33 34 1 11 33 13 17 18 21 21 23 23 24 25 25 26 31 5 NW SW NW SE NW NE SW SW SE SW NE SW SW SE SE SE NW SW not classified + + + + + + not classified + + Yes Yes 32 1 2 13 21 28 31 1 7 16 18 SW NE SW SE unk. NW NW NW unk. SW unk. + + + No Yes Yes 18 19 27 unk. NW NW + not classified Yes + + Yes + No No No No No No No �62 APPENDIX D - cont. 1962-63 lek location Range Township Section Quarter 3N 2N 27 12 15 15 16 18 19 2 12 12 unk. NW SW SY-l sw NW NE SW NW SE Within (+) or outside(-) 198182 occupied range not classified + + + + + + + + + Lek present in 1962-63 and 1981-82 No No No No No No No Yes No �63 JOB PROGRESS COLORADO State of Project No. Mountain N-I-R Work Plan No. Job No. REPORT Plover Transplant-Experimental 5 ------------------ 1 ------------------------- Period Covered: Personnel: 1 January 1982 through 31 December 1982 B. Cordova, L. Fisher, W. D. Graul, G. C. Miller, Sedgwick, Colorado Division of Wildlife. and J. ABSTRACT Fifty mountain plover (Charadrius montanus) chicks were trapped in Colorado and transported to Kansas as part of an agreement between Kansas Fish and Game Commission and the Colorado Division of Wildlife. No chicks died during captive maintenance prior to release. The first 2 groups of chicks were held at the Kansas release site for 1 to 2.5 days; the rest were released immediately upon arrival at the site. Reports by Kansas personnel are included. ��65 MOUNTAIN PLOVER TRANSPLANT-EXPERIMENTAL Gary C. Miller Historically, mountain plovers nested throughout western Kansas. Early records exist (Allen 1872, Goss 1891, Menke 1894), but records since that time have been scarce (Long 1940, Goodrich 1946, Rising and Kilgore 1964). Graul (1973) did not find evidence of recent nesting in western Kansas. Graul and Webster (1976) believed that the Pawnee National Grassland in Weld County, Colorado, represented the stronghold of the species' breeding range. Restoration of mountain plovers to Kansas was identified as a desirable objective for that state's nongame program (M. Schwilling, pers. comm.). A letter from B. Hanzlick, Director, Kansas Fish and Game Commission, to J. Grieb, Director, Colorado Division of Wildlife, dated 21 April 1982, requested 40 mountain plover/year over a 3-year period in a trade for greater prairie-chickens (~anuchus cupido pinnatus) (Appendix A). Thus, a possible restoration technique for mountain plover could be tested at little expense to Colorado. P. N. OBJECTIVES This project, funded through nongame checkoff funds, was initiated prior to the implementation of Administrative Directive I-I, Research Planning and Project Implementation (03 Aug 1982). The documents that exist (Appendices B, C) stipulate the general objective of providing mountain plovers to Kansas in return for greater prairie-chickens. SEGMENT OBJECTIVES Documents generated at the onset of this proposal included: 1. Formal correspondence from the Director, Kansas Fish and Game Commission to the Director, Colorado Division of Wildlife, requesting mountain plovers in exchange for greater prairie--chickens. 2. Survey of potential habitat and selection of a release site by Kansas personnel. 3. Provision of 75+ mountain plovers to Kansas between 1982 and 1984. DESCRIPTION OF STUDY AREA Graul (1975) described the area from which mountain plovers were taken (Pawnee National Grassland, Weld County, Colq.). The potential release areas in Kansas were described by L. Fisher in a report submitted to the �66 Kansas Fish and Game Commission on 8 June 1982. The site where the plovers were released was in Wallace County. 32 km northeast of Sharon Springs, Kansas. Details of the release were provided in a report by J. Ptacek, Kansas Fish and Game Commission, dated 04 August 1982 (Appendix E). METHODS Mountain plover chicks were captured by hand or with long-handled nets. Only chicks that were within approximately 2 weeks of fledging were taken from the area. Suitable chicks were those with feathering of the humeral tracts and at least some feathering of the capital tract, and with the longest primary feather partially exposed. Captured chicks were banded and held in captivity at the CDOW's Wildlife Research Station northeast Fort Collins. Prior to placing the chicks in the facility, it was cleaned and disinfected, and a clean sand over cement substrate was provided. Different techniques of captive maintenance were used. The chicks were transported to Kansas approximately 1-4 days before anticipated fledging. RESULTS Fifty mountain plover chicks were trapped, maintained, and delivered to Kansas personnel between 23 June and 02 July 1982. The first 2 groups were held in an enclosed pen at the release site from 1 to 2.S days. After consulation with CDOW personnel, the release technique was changed, and plovers were released in groups of 3-S immediately upon arrival at the release site. Bands from 2 chicks were found at a swift fox (VuJpes velox) den shortly after releases were made. Appendices D and E are reports written at the conclusion of the 1st field season. None of the SO chicks captured for release in Kansas died during the captive maintenance period in Colorado. Analysis of maintenance techniques has not been accomplished. DISCUSSION AND RECOMMENDATIONS It was originally believed that as many as 120 chicks would have to be captured to provide 7S+ chicks for transplanting. This original estimate (120 chicks) was a "worst case" scenario based upon chick mortalities observed elsewhere (W. D. Graul, pers. commun.). Because of the high survivorship of chicks during the captive maintenance phase, it probably will not be necessary to take as many chicks from the Pawnee Grassland as originally proposed. A second cohort of plovers will be captured and transplanted to Kansas in 1982, beeause it is not known if mountain plovers in the wild will breed or return to a suitable breeding area during their first year. In the spring of 1984. Kansas Fish and Game Commission personnel plan to evaluate the desirability of continuing the transplanting effort. �67 LITERATURE CITED Allen, J. A. 1872. Ornithological notes from the West. Goss, N. S. 1891. History of the birds of Kansas. Topeka, Kans. 692pp. Goodrich, A. L. 340pp. 1946. Birds in Kansas. Am. Nat. 6:263-275. G. W. Crane and Co., Kans. State Bd. Agric., Topeka. Graul, W. D. 1973. Adaptive aspects of the mountain plover social system. Living Bird 12:69-94. 1975. 87:6-31. Breeding biology of the mountain plover. , and L. Webster. --- Condor 78:265-267. 1976. Wilson Bull. Breeding status of the mountain plover. Long, W. S. 1940. Check-li.st of the Kansas birds. Sci. 43:433-441. Trans. Kansas Acad. Menke, H. W. 1894. List of birds of Finney County. Kansas. Univ. Q. 3:129-135. Kans. Rising, J. D., and D. L. Kilgore, Jr. 1964. Notes on birds from southwestern Kansas. Bull. Kansas Ornith. Soc. 15:23-25. Prepared by ~ e· TIL_\lc_ ~. Miller Wildlife Researcher C ��,. ·,~!,;~';-", APPENDIX A REGIONAL OFFICES: Northw~.t R'/llonar Offla RI. 2, 183 Ryp", •• Hav», Kan,OJJ 67601 tnc en trol Re,Ionol Offlc. Box 489,611 Cedar ConcordIa. Kan,,,,, 6690 I Nor BOX 54A, RURAL ROUTE 2, PRATT, KANSAS 67124 Northealll Regional Offla 3300 S. W. 29th Street 7'opeRa. Kana". 66614 (316) 672·5911 69 SouChwa,' R~,'onal Otrlctl 808 HI,hwo)j ,';6 Dod,. Clly, Ko""08 67801 Soulhcentrol Rrl{lonal (Jfflce Box 764,204 Newlan, We,t SIxth Kansa 67114 Sauth.lUt llettfonal Offler 222 We.1 Mal" lIuildln, Suite C &. D Chanute, Ka,,,u. 66720 April 21, 1982 Mr. Jack R. Grieb, Director Colorado Division of Wildlife 6060 Broadway Denver, CO 80216 Dear Jack: The Kansas Fish and Game Corrunission is interested :inpursuing a potential wildlife trade with the Colorado Divi~ion of Wildlife. . ,.~~ We would be interested in obtaining mountain plovers for a reintroduction project and feel that Colorado offers the b~st choice of transplant stock. Our nongame project leader has di~c~ss4. ~N-,s ~JsibPity with Dr. Walter Graul and concludes that a mutually beneflc1N: ~J3de can ?c made. . , .' ..,..,--..... -~ 'iJ. ", ,I". .,..~.,,':.,. of ••• " .' Our program is to ob ta ina t~tal ,of 140 pre-flight age plovers over a three year period (40, per year). Co.lorado would ·receive greater prairie chicken at a rate nrutually agreeabl~,beth'een pW'fwo agencies. Details can he· worked out once the basic trade conCept ,has ~een finalized. If agreeable, we would like to receive the first contingent of plovers yet this summer. , .. ,: '. 'i. '. . . \ , '! : ., ' \ '1):,., ; ~ , J] {Si~l~e.rely• /, .. I ; j 'W.!i.... :.... , Bill Hanz l i ck , Director Kansas Fish and Game COTllmission t : BDH/ddd ��1, Correspondence ;Di~isionQI -: ::F, APPENDIX B Only 71 STATE OF COLORADO .:. DIVISION ; DEPARTMENT Of WILDLIFE OF NATURAL l, I RESOURCES DATE: , 12 January John Torres rift' !?: \ ~.'FROM: . Dick Hopper I RE: Mountain plover transplant to Kansas The Kansas nongame person (Marvin Schwilling) and Walt Graul have been pursuing the idea of a mountain plover transplant into historical range in Kansas. If we get the necessary approvals from both ends, we would like to initiate the project this summer. My concern at this point is that since the mountain plover has always been viewed as a species of special interest to many people in the general public, it seems to me that we should inform our Citizens Advisory Council of this project. We certainly would not want to do something of this nature if it met with public disapproval. I would, therefore, like to see this project discussed at an Advisory Council meeting -- hopefully prior to April 1. I suggest that the enclosed narrative should be sent to Council members prior to the meeting and Walt will be happy to attend the meeting to answer questions. If a timely meeting is not possible. I would hope that we could obtain written connnents from Council members by April 1. RH/ds cc: _,--- ---.~.~ DOW-A-F-8 -~ Walt Graql Wayne Sandfort 1982 ��73 APPENDIX C PROPOSAL MOUNTAIN PLOVER TRANSPLANT TO KANSAS Justification: The mountain plover historically nested throughout western Kansas. It still nests in southwestern Oklahoma -- due south of the former Kansas range. It is speculated that the breeding population was lost from Kansas during the Dust Bowl days as the main breeding range contracted westward. Although there now appears to be suitable habitat in Kansas, it is felt that the species has no way of re-establishing itself since birds return to the general area where they were raised to breed and the main migration is westward and southward. Someone might que.stion why we are proposing Kansas for the transplant rather than some area of former range in eastern Colorado. First, it should be recognized that we view this as an experiment -- something of this nature has never been tried with shorebirds. Although such a transplant is not considered a high priority in Colorado it is in Kansas (we have a substantial population; Kansas does not). Kansas is willing to cover all essential costs and we are hopeful that an agreement can be worked out whe reby we receive greater prairie chickens in return -- for a transplant to our Tamarack grassland. Thus, by conducting the transplant in Kansas we have an opportunity to evaluate an experiment for a minimal cost that may benefit us in the future, and we may obtain greater prairie chickens to help meet our endangered species program goals. Procedures: 1. 2. 3. Kansas will formally request the plovers via correspondence between the Kansan and Colorado agency Directors. Kansas will conduct a survey of potential habitat 'and select the best site in the spring of 1982. A total of 75+ birds will be provided between 1982-84. This is the estimated number needed to establish a breeding population of 50. This will require approximately 120 birds to be taken to ac ccunt :for pra-rr e Lea se mortality. The details of providing the birds are as follows: (a) Kansas will pay someone to capture the birds ~- 30-40 per year. (b) Chicks will be captured that are within two weeks of fledging and they will be released in Kansas so that they can fledge on the transplant site. Ce) Birds will be taken from the Pawnee National Grassland area -- the stronghold of the population. (1) �74 (d) I will obtain the necessary collec.tion permits and I will notify the administrators of the Grassland. also supervise the collection so that: birds are taken from a widespread area to avoid hurting a local population. (e) The chicks will be held temporarily here at our research station so that they can be delivered in groups to minimize travel. I will care for them. (f) The birds will be delivered by car or plane -- the latter would be best. Kansas will cover transportation costs. All birds will be color-banded plus receive U.S. Fish. & Wildlife Service bands. (g) The transplant site will be monitored intensively by Kansas in the following few years to determine success. Overview of technique: Mountain plover chicks are highly precocial and previous research has documented that they are quite capable of surviving without parental care once they are a couple weeks old -- I have held them in captivity previously. In fact. most chick mortality in the wild occurs the first three days after hatching. We hope that if the chicks fledge on a Kansas site, they will return there to breed. Hopefully, they will move south to winter (some birds do winter in the grasslands of Texas). Although this has never been tried with shorebirds, the basic procedure has worked well with waterfowl. Waterfowl are noteworthy in this respect since they have several features in common with mountain plovers -- precocial young, young return to where they obtain their first flight experience~ etc. Environmental Impact: Graul and Webster (Condor 78:265-267) estimated the total mountain plover population to be in the range of a few hu.ndred thousand birds. The Pawnee National Grassland population was estimated to be in the range of 20,000 birds. Thus, removing approximately 120 chicks represents about one+ha l.f of one percent of this population. The impact is actually less, since a substantial portion of the chicks removed 'i,1Quld not have made it through their first year anyway. Additionally, it should be recognized that habitat in the private segment of the Pawnee National Grasslan.d area continues to be converted to agricultural lands. Since suitable habitat appears to be at full carrying capacity in terms of mountain plover breeding density, the loss of habitat indicates a local surplus of mountain plovers is being created relative to rema i.m.ng habitat. The best use of this surplus is to attempt to establish new populations. Prepared (2) By: Walter D. Graul Nongame Research Leader Colorado Division of Wildlife 317 W. Prospect Street Fort Collins, CO 80526 �. / APPENDIX D . 75 STATE OF COLORADO Richard D. L8mm. GOY mer DEPARTMENT OF NATURAL RESOURCES DIVISION ..Jack R. erleb, F WIL LI Director 6060 Broadway Denver, Colorado 90216 (925-1192) RRPORT MOUNTAI_N P.LOVER TRANSPLANT TO KANSAS - 1982 The project design called for delivering approximately 100 mountain plover chicks to Kansas between 1982 and 1984. Fifty chicks were delivered between 23 June and 2 July 1982. The fifty chicks were captured on the Pawnee National Grassland and held briefly in captivity at the Fort Collins "7ildlife Research Station (48 were held from 1-7 days; 2 were held 9 days). Four shipments of chicks were made such that all chicks reached Kansas within a few days of anticipated fledging. The release sites were located between Goodland arid Sharon Springs~ Kansas. There were 4 release sites, all within 4 miles of each other, within an overall shortgrass prairie area. Bands from 2 chicks were found at a nearby swift fox den. Other birds seemed to exhibit normal behav:lor--see attached report. After some initial experimentation, a suitable release method was developed. Prepared By: Walter D. Graul Nongame Research Leader DEPA TMENT OF NATURAL RESOURCES, Monte Pascoe, Executive Director Donald Fernandez, Vice Chairman •..•.!_L __ I U:_L __ e WILDLIFE COMMISSION, Wilbur Redden, ~ James Smith, Secretary" Jean K. Tool, Member OJ Vernon C. Williams, .l! __ L_ ..•. <: __ r_ .. ...J:tI li6 •.•..h.o.r _ Pi,.h"r"" niVAlhic.c. MAmh@r Chairman Member ��•. r 77 APPENDIX E MOUNTAIN PLOVER REINTRODUCTION IN KANSAS Jim Ptacek Kansas Fish & Game Commission 832 East 6th Emporia, Kansas 66801 August 4, 1982 �78 Introduction The mountain plover (Eupoda montana) is a common summer resident in the short grass prairies of eastern Colorado. The stronghold of this population during the summer is found at the Pawnee National Grasslands of northeastern Colorado (Graul, 1976). Up to and including the early 1900's, wintering populations in Texas and California extended their spring migration to include the prairies of western Kansas (Allen 1872, Goss 1891. Menke 1894, Snow 1903, Long 1940, Rising 1974). By the mid 19000s though, due to changing agricultural practices and climatic hardship, populations ceased their migration into Kansas (Tordoff, 1956)0 Breeding populations remained adequate in Colorado throughout the 1900's for expansion into Kansas. but the plovers failed to relocate (Graul, per. comm. p 1982). On June 8, 1982. a report was submitted to the Kansas Fish and Game Commission on habitat assessment for the reintroduction of mountain plovers into western Kansas (Fisher, 1982). Short grass prairie northeast of Sharon Springs in Wallace County, Kansas was selected to Initiate this experimental program. Mountain plovers would be reintroduced into this area through cooperative efforts between Kansas and Colorado Fish and Game Commissions. From June 22 to July 59 1982. field work on Mrs. George Harrison's ranch~ (T-l1, R-38, S-34), was conducted which ultimately led to the reintroduction of 50 mountain plovers. During this period daily surveys were conducted to monitor the birds' movement patterns. possible predationv Observations were also made to note inter and intraspecific competition. feeding habits behavior, and general welfare of the plovers. p �79 -2- Materials and Methods Two procedures were utilized for the reintroduction. The first method involved confining the plovers in a five foot high, ~" mesh wire fence. This structure enclosed a40 ft square area of pasture with less than 20 slope. A double wire electric fence was placed around the holding pen to discourage predators, while the author kept a nightly surveillance for additional protection. Lean-tos were constructed in two areas of the pen for shade and protection from storms. Water was furnished in metal pans. Mea1worms plus catfood, supplemented the diet of invertebrates obtainable inside the pen. After a brief acclimation period within the pen (24 hours up to 2.5 days) the fence was opened and all birds released. After releasing 24 birds complications were noted and a new technique was devised. The second procedure abandoned the method of retaining the plovers for any length of time. Instead, new arrivals were divided into smaller groups of three or four birds, based on similar primary and secondary feather development. These groups were then released in various habitats in the study area on the day they were received. The remaining 26 plovers were released in this manner. Daily surveys were run from 6:00 a.m. - 9:00 a.m. and again from 6:00 p.m. - 10:00 p.m. Surveys began at the.release sites and included walking and driving to locate flocks. Particular attention was paid to flat, sparsely vegetated areas such as those found around cow paths, stock tanks, and roads. All of the mountain plovers transplanted into Kansas were captured and banded from the Pawnee National Grasslands by the Colorado Fish and Game Commission. The plovers were then transported into Kansas after their �80 -3- feather tracts had developed sufficiently to allow for normal thermoregulation and near flight capabilities. All release sites were within a three mile radius of the Harrison's ranch (Table I, Figure 1). Study Area The 1982 release sites are encompassed by 580 sq. km. of grassland 20 miles northeast of Sharon Springs, Kansas. Blue grama and buffalo grass are the dominant plants in this area, averaging 4 cm in height (Fisher, 1982). Varying amounts of cropland reverted to midgrass, averaging 8 to 20 cm in height, are also present (Fisher, 1982). Wheatgrass, yucca, pricklypear cactus, barrel cactus, globmallow, western wallflower, and thistle are 'common.l.y found associated with both of these grasslands. Twenty-four plovers were released in an 8 sq. km. area with less than 20 slope and vegetation less than 5 cm tall. The sLte was bordered by midgrass pasture 2000 ft. to the north and east. a road 4000 ft. to the south, and rolling hills to the west. The remaining 26 plovers were 0 released in various habitats with slightly more than 2 slope. Vegetation in these areas was a mixture between shortgrass and clumps of midgrass. Vegetation height ranged from 4 cm to 20 em. These areas better typified mountain plover brood rearing habitat than the original 8 sq. km. release site. Discussion Retaining the plovers at the release site until fledging occurred, proved unsatisfactory in several ways. Food consumption rates exceeded the supply of invertebrates found within the pen. This factor limited the �81 -4Table 1. Legal description of mountain plover release sites in Wallace County, Kansas. Dates released # of birds released Legal description of release site 6/23/82 19 T-11, R-38 , S-26 6/25/82 5 T-11, R-38, S-26 6/30/82 4 s 4 T-11p R-38. S-27 6/30/82 4 T-11, R-38, S-16 6/30/82 5 T-ll, R-38. S-18 7/2/82 3 T-Up 7/2/82 3 T-11, R-38 , S-17 7/2/82 3 T-12, R-38, S-9 R-38, S-16 �-5- Figure 1. Release site locations with number of plovers released at- each site. N WALLACE-IOO �83 -6acclimation time. Thus the initial 19 plovers were maintained within the pen only 24 hours, while a second group of 5 plovers was contained 36 hours longer. In both cases, the inability to provide an adequate food supply led to the release of the plovers prior to fledging. could fly when freed. Only three plovers The remaining birds fledged 10-48 hours later. This lowered the rate of survival by increasing the chance of predation on the f Lf.ght Less birds. A government band was found at a swift fox den within one day after releasing a group of 19 plovers. Another band was discovered at the same den five days after a similar release of plovers. Both times feathers found around the den showed primaries and secondaries in an early stage of development. The captive plovers tended to develop an attraction to the release site which delayed their rate of dispersal and caused an additional problem. After being released, 10 plovers occupied an area within 500 ft. of the pen for 48 hours before abandoning it completely. A group of nine plovers foraged six days within 1000 ft. of the pen, while four others remained nearby for 13 days before dispersion occurred. During this time many of the plovers periodically returned to the holding pen and demonstrated a lack of wariness. These birds could easily be approached within 2 ft. without becoming alarmed. Plovers released without a period of confinement demonstrated quick adaptability to the area. More wariness was evident and these plovers were immediately reluctant of being approached. They would forage within 1000 ft. of the release site for one day and then completely desert the area. At this point they would wander for considerable distances, always remaining in a group. Four plovers were relocated two days after being released .3 �84 -7miles from the original release site, while another flock of three plovers moved .75 miles. One plover moved 2.5 miles in five days, the longest distance recorded. The majority of these plovers tended to migrate toward areas barren of vegetation. Wheel tracks were most notably inhabited. Mountain plovers were often observed foraging along these tracks in the mornings and evenings. They were also seen at night sleeping in groups of five within the deepest tracks. to sit on during the midday. Old prairie dog mounds were often used Roads and barren areas around windmills attracted plovers during evening hours from 8:30 p.m. until 10:30 p.m. These plovers were also quick to demonstrate defenses against possible predation. They were quite adept at hiding in the grass. Several times Swainson's hawks were seen flying over mountain plovers without locating them. One plover was even seen flattening its body to the ground as a dove passed overhead. unsuccessfully. Skunks were also noted searching areas for plovers No predation was evident upon examination of four burrowing owl nesting holes. Very little intraspecific competition occurre~ between any of the plovers. The only observed aggression occurred during confinement, at which time the plovers would compete for space at the water trough. When unconfined no competition was observed, though a distance from 2 to 6 ft. was maintained between foraging birds •. Only once was interspecific competition noted. A horned lark attacked one mountain plover three times before the plover moved out of its territory. Summary Habitat assessment was conducted in Kansas for the reintroduction of mountain plovers. Fifty mountain plovers were reintroduced into shortgrass �85 -8prairie northeast of Sharon Springs in Wallace County, Kansas. Two methods were used for their release, with an immediate release technique proving to be the best method. The plovers were released in nest and brood habitat within a three mile radius of the Harrison's ranch (T-II, R-38, S-34), with both habitats utilized similarly. Two government bands found at the same swift fox den, and one plover which died before release were the only mortalities observed. Movement patterns were sketchy but indicate plovers moved in flocks .3 to 2.5 miles within the first week after being released. Preferred areas inhabited by the plovers lacked vegetation such as: wheel tracks, roads, and areas around windmills. Two other areas have been assessed for further reintroductions of mountain plovers: White Woman Creek south of Sharon Springs, Kansas, and the Coolidge area north of Coolidge in Hamilton County, Kansas. �'. ( , 86 -9- LITERATURE CITED Allen, J.A. 1872. 263-275. Ornithological notes from the West. Amer. Nat. 6(5): Fisher, L.E. 1982. Habitat assessment for peregrine falcons and mountain plovers in western Kansas. Unpublished 34pp. Goss, N.S. 1891. History of the birds of Kansas. Topeka, Kansas. 692pp. G.W. Crane and Co., Graul, W.o. 1976. Breeding status of the mountain plover. 78(2):265-267. 1982. The Condor. Personal Communication. Long, W.S. 1940. Checklist of the Kansas birds. Science, Vol. 43:433-441. Trans. Kansas Acad. Menke, H.W. 1894. List of birds of Finney County, Kansas. Quar. 3:129-135. Kansas Univ. Rising, J.D. 1974. The status and faunal affinities of the summer birds of western Kansas. Univ. of Kansas Sci. Bull. 50(8):347-388. Snow, F.H. 1903. Catalog of the birds of Kansas. of Science. Vol. 3:13. Trans. Kansas Acad. Tordoff, H.B. 1956. Check-list of the birds of Kansas. PubIs. Mus. Nat. Hist. 8:307-359. Univ. Kansas �87 JOB PROGRESS REPORT State of COLORADO Project No. N-2-R·· Fishes of Colorado: Work Plan No. 1 Job No. 1 Period Covered: 1 January 1981 to 31 December 1982 Personnel: An analysis of Distributional Change J. Bennett, K. Galat, C. Haynes, J. Woodling, Colorado Division of Wildlife. ABSTRACT Information retrieval/storage systems required to evaluate the historic and present distribution of the fishes of Colorado are described. Annotated bibliographies of literature, reports and records pertaining to the Rio Grande, San Juan, and Colorado rivers are provided. ��89 FISHES OF COLORADO: AN ANALYSIS OF DISTRIBUTIONAL CHANGE Charles M. Haynes and Karen H. Galat P. N. OBJECTIVES The long-range objective of this project is a comprehensive publication detailing the historical distribution and current status of the fishes of Colorado. Intermediate goals are two fold: (1) the preparation of a bibliography of information sources concerned with fishes in each of the major river drainages of Colorado, and (2) a data bank containing a summary of the information in the bibliographic sources. To achieve these goals a data collection and storage system was designed that would answer 2 objective questions: 1. Where in Colorado has this fish species been recorded? 2. What fish species have been found in this water (may be 1 of 7 designated drainages, a subdrainage , or a specific location in Colorado). In -the 1981-82 seasons these goals were narrowed to the San Juan, Rio Grande and Colorado river drainages in Colorado. Approaching the collection of data on a drainage-by-drainage basis permitted a more systematic search and a finished product, i.e., a bibliography of these regions. A method of data collection was required that would effectively summarize a source of information in terms of the objectives. Relevant data were specifically defined so that a document was efficiently and consistently evaluated. Two methods of-information storage were desired. A manual filing system at the Fort Collins Nongame Research facility would permit quick access to the data and would help answer relatively simple questions. Storage of the data in a computer system would permit more sophisticated queries and data evaluation. We tried to keep both storage and retrieval systems simple so that researchers interested in the information would not need an elaborate set of instructions to retrieve the information. SEGMENT OBJECTIVES 1. Document the historical and present distribution of both native and non-native fishes in the Colorado River drainage, with particular emphasis upon nonharvested and threatened and/or endangered species. 2. Provide a data-base which may serve as a predictive tool for recognizing future potential negative impacts upon segments of the fish fauna as a consequence of energy development. stream and river modification (e.g., dams, withdrawals), agricultural development (e.g., irrigation returns, livestock damage, etc), acid-precipitation, etc. �90 METHODS Data Summarization The basic criterion which established the usefulness of a document was whether it cited the observation of a fish species in Colorado. Once the document was judged to be useful, the data in it were summarized. The information had to specify the fish at least to the genus. The following data were delineated as those to be sought and placed on a recording form (Fig. 1): Document Accession number (assigned as document was discovered) Citation (author, publication date, title, etc.) Publication type Keywords Abstract Data Species Description of location Hydrologic unit Longitude Latitude \\latercode Elevation Location name Habitat type Relative abundance Stocking number Year of observation A list of pre-defined keywords was used to describe each document for the purposes of a bibliographic index and to permit sorting on the computerusing' these words (Table 1). The abstract further informs the user of the specific relevancy the information source may have to his/her query. Species Species names used are those outlined by Bayly (1980). Where common names were used in recent publications, the associated specific name was that listed by Bayly (1980). In older documents, personal communication with J. R. Behnke or W. Wiltzius was sought to establish the correct taxonomic identification. In this way, a list of species with associated common names and past scientific names was prepared to maintain consistency in nomenclature (Table 2). Description of Locations Hydrologic Unit The first location descriptor is a hydrologic unit as defined by the U.S. Geological Survey Hydrologic Unit map. This is a hierarchical classification of the drainages of the United States and consists of 8 �." ->. to c: -s (J) ~~cetiAl(J\\ I'age _~.__ ItUlIlluar of _ Dcation _----_._---------~blicatio" Type: Survey, Environ. Creelcensus, _. Il' MIl' Stock. Record, Thes1.s, iiYliROI,OGIC'!"".""].i:>.n,,"""I"ATE'C'A: 9'ECIES o Study, UNIT - ..-.-- .... --.. '. .. I--.---.- ....j ..•................ tCODE ---{ ..... -- Vl --_ Il' -s ....•• . .•. _- RevieH, NAHE Guide, NeHspaper, ABUN~ __ - c: ~ l IWATERTY"I "',"D,--I --.---. l==t_l==t= I'llb., ScI"nl·. ,,"',., NO. pe r s . comm , n:AR SUCCESS .. - ..•...... - ... ---- t-----t---t--t----+-- .. j .._.--------1- N Il' M- _.. ·-·----···----·-··1···----·--4-·--·· o :::l ..•. --1·----1----1 ---------------1--'----1------+--_·· +--- f--f--- 1------.-... -----. -t; o ~ -----------1-----+---+---·1--~--1 -- ..- ... --._--.... -.- I-- 1----1--·--1----1 1--·1 ----{··---I-·----I--- ---.-- +--I---·~ I I I I--- l--l~ 1 II+--- 1----1----1-----1---+ ------- ~ .---- 1----1--·'· .--.--t-.--. ..------ -----------_ -.--- .. ---t---+----, .-----_. -+---·-1---1--+ ------ 1 I-----t---t---t----t--- 1--1-- ..------ ~---1.--+---I----+-l---I---I---·I I---+---l +-- • I--I·-··-·---j------·EE--' E .... __.. == _~=~._ I_--= =--= === -l==I===t== . -.--~ ... --.----.--.-- _______________ I- I-----~-- I I .. ~_·-f--I----I I I--- .-1 ~:~~~:;--.--_._---._._._-_-_ ---.--.. .._.._._---_ . -. ---.--=--==~~:~.-==_._..__ K:. . 1.--._._---- ...•.. -xc-··- ..-- ... I _ .. .•. -1_ .•.. L. ._-----_._ ...... -.------.--.. - , 1 ,__• ---I---.~ ....-----_._._-_. ·'-0' t. ~1.. .~--.-.I .-"'"\ 1.0 I-' �92 Tab1e 1. List of keywords. Abundance Age/Growth Alteration Anguillidae Archaeology Behavior Catostomidae Centrarchidae Cichlidae Clupeidae Competition Cottidae Cyprinidae Cyprinodontidae Description Disease Distribution Ecology, i.nteractions Ecology, trophic ~elationships Esocidae Fishery, commercial Fishery, sport Food Habits Genetics Habitat History Hybridization Ictaluridae Inventory Invertebra tes, benthos Invertebrates, plankton Limnology Management Migration Movement Natura1 Hi story Percichthyidae Percidae Physiology Poecillidae Pollution Predation Production Propagation Recruitment Reproduction Salmonidae Sciaenidae Spawning Species List Standing Crop Status Stream Flow Taxonomy Toxicant Water Quality, chemica1 Water Qual ity, physical �93 Table 2. List of synonymous Scientific species and common names. Common name name Anguillidae AnguiZla rostrata AnguiZZa chrysypa American ee19 freshwater eel Clupeidae Dorosoma cepedianum AZosa sapidissima Dorosoma petenense Gizzard shad Threadfin shad Salmonidae Coregonus clupeaformis Salmo clarki. l.eoiei. SaZmo cZarki virginaZis SaZmo cZarki spiZurus Salmo clarki pZeuriticus SaZmo mykiss pZeuriticus Salmo clarki stomias SaZmo clarki macdonaZdi SaZma gairdneri Salmo irideus Salmo t-ioulax-ie SaZmo saZar SaZmo sebago Salmo trutta Sa'lmo fario Salma fario levenensis Salmo aguabonita Oncorhynchus nerki Oncorhynchus kisutch Oncorhynchus tsawytscha Salve linus fontinaZis SalveZinus namaycush Cristivomer namaycush Prosopium wilZiamsoni Coregonus wiZZiamsoni ThymaZZus arcticus ThymaZZus signifer ThymaZZus montanus Lake whitefish Yellowstone cutthroat, Rio Grande cutthroat Colorado cutthroat Greenback cutthroat Yellowfin cutthroat Rainbow trout, coastrange California brooktrout trout, Lake Atlantic salmon, landlocked salmon. Sebago salmon, Schoodic salmon Brown trout, Lochleven, Von Behr Golden trout. Kokanee, Little redfish, landlocked sockeye. blueback Coho Chinook,quinnat9 California salmon Brook trout, speckled trout, saibling Lake trout, mackinaw Mounta in wh itefi sh, Wi 11 iamson IS whitefish Arctic grayling, American grayling, Montana grayling Esocidae Esox americanus vermicuZatus Esoc vermicuZatus Esox lucius Esox masquinongy native trout Grass pickerel Northern pike Muskellunge �94 Table 2. (cont.) Scientific name Common name Cyprinidae Campostoma anomalum Campostoma aikenii Tutilus anomalus caraeeiue auratue Carassius carassius Cyprinus auratus Cyprinus carpio couesius plumbeus Hybopsis plumbea Gila cupha Gi La e Leqane Gila emoyri Gila robusta elegans Gil.a nigrescens GiZa pandorae Gila pulchel.Lus Tigoma nigrescens Leuciseus pulcher Leuciscus pulcher Ceinostomus pandora Gila robusta Gila emorii Gila emoyri Gila grahami Ptychocheilus vorax Hybo~thus hankinsoni Hybognathus nuchalis Hybognathus placitus Hybopsis aestivalis Hybopsis dissimilus Couesius dissimilus Hybopsis graciZis gracilis Ceratichthys physignathus Platygobio physignathus Pogonichthys physignathus Hybopsis gracilis gulona Hybopsis storeriana Nocomis biguttatus Hybopsis biguttata Hybopsis kentukiensis Notemigonus cr.ysoZeucus auratus Notropis blennius Notropis cornutus Notropis mega lops Notropis doreal ie doreal ie Central stoneroller, greased chub Goldfish Carp, German carp Lake chub Humpback chub Bony tail chub Chihuahua chub Rio Grande chub, Pescadito Roundtail chub Brassy minnow S i lvery mi nnow Northern plalns minnow Speckled chub, Arkansas chub Streamlined dace. mountain dace Plains flathead chub. thick-jawed chub Southern flathead chub Silver chub Hornyhead chub, Indian chub, jerker Western go1den shiner River shiner Common shiner Central bigmouth shiner �95 Tab 1e 2 (cont.) Common name Scientific name Cyprinidae (cont.) Notropis Notropis Notropis Notropis dorsaZis piptoZepis heteroZepis Zutrensis Zutrensis stramineus Nota-op-ie de l-icioeue Notxoopie ecul.la Notropis texanus Notropis voZuaeZZus Phenoaobius mirabiZis Phoxinus eos Chrosomus eos Phoxinus erythrogaster Chrosomus erythrogaster Phoxinus naeogaeus Chrosomus naeogaeus PimephaZes promeZas PZagopterus argentissimus PtyahocheiZus Zucius Rhinichthys cataractae Rhinichthys henshawi Rhinichthys duZais Rhinichthys oscuZus Agosia yarrotJi Rhinichthys nubiZus yarrotJi Richardsonius baZteatus GiZa bal.teatue SemotiZus atromacuZatus Tinea tinca Western bigmouth shiner Blacknose shiner Plains red shiner, redfin Sand shiner Mimic shiner Suckermouth minnow Northern redbelly dace. Southern redbelly redbelly dace Finescale dace dace, western Fathead minnow, blackheaded minnow Woundfin Colorado squawfish Longnose dace, dulcis minnow Colorado speckled dace Redsided shiner Northern Tench cre'ek chub, horned dace Catostomidae Carpiodes carpio Carpiodes cyprinus Carpoides velifer Catostomus aatostomus Catostomus griseus Catostomus commersoni Catostomus aZticoZus Cyprinus commersoni Cyprinus teres Catostomus discoboZus Pantosteus deZphinus Pantosteus virescens Minomus delphinus Minomus bardue Catostomus Zatipinnis Acomus latipinnis River carpsucker Quillback, sailfish Highfin Longnose sucker White sucker Bluehead Mountain mouthed sucker Flannelmouth sucker, sucker chisel- �96 Tab 1e 2 (cont.) Common name Scientific name Catostomi ae (cont.) Catostomus platyrhynahus Pantosteus jordani Catostomus pZebeius Pantosteus plebeius Moxostoma macroZepidotum Moxostoma aureolum Moxostoma breviceps Xyrauchen texanus Catostomus aypho Catostomus texanus Xyrauohen aypho Xyrauahen uncompaghre Mountain sucker Rio Grande mountain sucker Short head redhores,northern redhorse Razorback sucker, humpback sucker Ictaluridae Iatalurus furcatus Ictalurus punctatus Iatalupus melas Ameiurus mel-as Iatalurus nebulosus Ameiurus nebuZosus Noturus [laoue PyZodiotus olivarus Blue catfish Channel catfish, white catfish Black bullhead Brown bullhead, common bullhead, horned pout Stonecat Flathead catfish Cyprinodontidae Fundulus zebrinus Fun,dulus kansae Fundulus saiadicus Fundulus floripinnis Zygonec°t;esJZoripinnis Plains killifish, zebra fish, zebra topminnow, dogfish Plains topminnow, littl~ redfin, little green topminnow Poeci 11 idae Gambusia affinis Mosquitofish. gambusia Percichthyidae Morone ahrysops Lepibema chrysops Roccue chrueope Marone s~xatiZis White bass Striped bass Centrarchidae Arahoplites rupestris Ambloplites rupestris Lepomis cyanellus Apamotis cyaneZlus Lepomis gibbosus Sacramento perch, rock bass Green sunfish, bluespotted Pumpkinseed sunfish �97 Table 2 (cont.) Conunon name Scientific name Centrarchidae (cont.) Lepomie gu Loeus Lepomis humi Us Lepomis macrochirus Lepomis paLLidus Chaenobryttus guLosus Lepomis microLophus Lepomis megaLotis Mieropterus doLomieui Mieropterus saLmoides Pomoxis annuLaris Pomoxis nigromaeuLatus Pomoxis sparoides Warmouth Orangespotted sunfish Bluegill, bream Redear sunfish Longear sunfish Smallmouth bass, tiger bass Largemouth bass, straw bass White crappie Black crappie. calico bass, strawberry bass Percidae Etheostoma exiLe Etheostoma iowae Etheostoma eragini Etheostoma nigrum BoLeosoma nigrum Etheostoma radiosum Etheostoma spectabiLe Perea f1avescens Pereina caprodes Stizostedion eanadense Stizostedion vitreum vitreum Iowa darter Arkansas darter, Cragin's darter Johnny darter Orangebelly darter Plains orangethroat Yellow perch, ringed perch Log perch Sauger Walleye .' Sciaenidae ApLodinotus grunniens Freshwa ter drum Cottidae Cottus biardi Cottus punctuLatus Cot~us semiseaber Cottopsis semiseaber Mottled sculpin. Rocky Mountain sculpin, blob, Miller's thumb Cichlidae Ti Lapia aurea Blue tilapia �98 numbers. The first 2 numbers described major drainages. In Colorado there are 3: th Platte (10), the Rio Grande (13), and the Colorado (14). The next 2 numbers described subdrainages within these areas. In Colorado there are 3 subdrainages to the Platte River: the North Platte (1018), the Sough Platte (1019), and the Republican (1025). These subdrainages are further divided and identified by the remaining numbers. Thus, Cache la Poudre River in the North Platte drainage is 10190007. Such a hierarchical classification easily permits queries on whole drainages or parts of a drainage. Use of this system also encourages consistency with a biologically meaningful scheme already organized on a nationwide basis. Longitude and Latitude We felt that longitude and latitude could be biologically useful in evaluation of fish distribution so each site summarized from a document was described to the nearest southern 8 minutes (latitude) or nearest eastern 8 minutes (longitude). Thus, if a site was between 37'000 and 37'050 it was described as 37'000• A map of Colorado divided into 8 minute longitude and latitude intervals was prepared and used to locate the sites. Occasionally a site was not described well enough in the document to precisely locate it .. Water Code The Colorado Division of Wildlife (CDOW) has identified 13 major drainages in Colorado: North Platte, South Platte. Arkansas, Rio Grande, San Juan, Colorado, Dolores. Gunnison, Yampa, White, Republican, Larimer and Green. Under the CDOW water code system, each water body surveyed is assigned a water code number. The water body is located within one of these drainages and a county as well as the range, section, and township. Biological, chemical, and physical information of the water body is stored on a computer system under the water code number. Habitat Type The water in which the fish was observed was described as 1 of 7 types if there was sufficient information in the document to classify it: Reservoir (RES) Permanent river (RPERM) Intermittent river (RINT) Lake (LAKE) Pond (POND) Irrigation ditch (CAN) Ground water spring (SPR) Relative Abundance If the data were "quantitative" (creel census, electro shocking , netting, etc.), an indication of the relative abundance of t.heobserved species was given by using the foLl.owi.ngcode: �99 - 100% - 90% - 75% - 50% - 25% 10% 5 5% 91 76 51 26 11 1 2 3 4 5 6 7 This approach, although somewhat arbitrary, does provide a higher degree of resolution for comparing relative abundance than subjective density classes. This was not always possible, however, and observations made by the author is which he gave his/her opinion of relative abundance were summarized by the following code: abundant common unco on rare A C U R Stocking Number and Year of Observation If the document contained stocking data, the numbers of each species stocked were included. Finally, the year of observation was recorded. Data Storage Manual System Data Summarization File The field summarization forms are stored numerically by accession number. Card Files Author File. Each document has a card with the citation information on it in bibliographic form. The accession number is included but the cards are arranged alphabetically by author. Geographic file. The purpose of this file is to answer the question, "What fish species have been found in this drainage?" Seven major drainages were chosen as main divisions and number- and color-coded: North Platte (1) - white South Platte (2) - blue Arkansas (3) - yellow Rio Grande (4) - orange San Juan (5) - purple Dolores, Gunnison, and Colorado River mainstem (6) - green White and Yampa (7) - red These drainages were subdivided into 3 - 10 subdrainages based on the divisions or consolidations of subdivisions on the U.S.G.S. Hydrologic �100 Unit map of Colorado. The file is arranged numerically by drainage and subdrainage. The correlation between the U.S.G.S. drainages and the manual system are outlined in Table 3. Within subdrainages is a card(s) for each fish species found within that subdrainage. These are arranged alphabetically by species. The information on the card is observation year, specific location, and accession number to the document from which the information came. For example, Fig. 2 illustrates a card for rainbow trout from the LaPlata River drainage. Species file. Thi file contains information identical to that of the geographic file. The cards, however, are arranged to answer the question, "Where in Colorado has a particular fish species been observed?" The file is, therefore, arranged alphabetically by species. Within a species division are cards for each drainage/subdrainage arranged numerically as in the geographic file. Because the cards are also color-coded, a quick look reveals in which major drainages a species has been observed. Map file. For each fish species observed in Colorado are one or more drainage maps of Colorado. Specific locations of observation are indicated on the map accompanied by t.heassociated accession number for that observation. Maps are further designated as pre-1900, 1900-1939, or 1940 - present. Maps are filed alphabetically by family and species within a family. Computer Storage and Retrieval System The information summarized on the data collection forms will be on the Gold system of the Colorado State University computer in called MANAGE. This is an interactive program. Conversational queries can be demanded which prompt an answer in the form of a instance, stored a program type list. For "List a1·1 fish species observed in 14080202 (McElmo drainage)," "List all Rio Grande subdrainages (1301) in which Tinca tinca (tench) have been observed," HList all publications by Lemon, D. L. on file," "List all documents with the following keywords in common: (Salmonidae, Habitat, Water quality, chemical." Information stored in the MANAGE program not in the manual file include the CDOW stream survey and lake and stream creel census/stocking records after 1973. These data bases are interfaced with this data base by the water code number. RESULTS An annotated bibliography pertaining to fish observations in the Rio Grande, San Juan and Colorado river drainages was compiled. Each bibliography is indexed by subject (keywords) and listed alphabetically by the author. Each reference is accompanied by the accession number in parentheses which locates the summarization form in the manual file and the �101 Table 3. Relation between organization references to Colorado regions based on drainages. Hydrologic unit 10 Name 18 0001 0010 Platte River N. Platte N. Fork N. Platte Laramie River 19 0001 0002 0003 0004 0005 0006 0007 0008 0009 0010 0011 0012 0013 0014 25 Manual system 1 Division of Wildlife 37 NP 1-1 1-1 36 L S. Platte Upper S. Platte N. Fork S. Platte Cherry Creek Clear Creek St. Vrain Big Thompson Cache la Poudre Owl Creek Crow Creek Kiowa W. Bijou Lower S. Platte Beaver Pawnee 2 2-1 2-2 2-3 2-4 2-5 2-6 2-7 2-8 2-8 2-8 2-8 2-9 2-9 2-9 41 SP Republican Arikaree N. Fork Republican S. Fork Republican Frenchman Sand 2-10 2-10 2-10 2-10 2-10 '2-10 38 REP 0001 0002 0003 0005 0006 26 0001 0002 0004 Smoky Hill S. Fork Smoky Hill N. Fork Smoky Hill Ladder 2-10 2-10 2-10 2-10 Arkansas River }fiddle Arkansas Upper Arkansas Cripple Creek Fountain Chico Chicosa Huerfana Apishapa Horse Fort Lyon Purgatoire Big Sandy 3 11 02 0001 0002 0003 0004 0005 0006 0007 0008 0009 0010 0011 3-1 3-2 3-2 3-3 3-3 3-3 3-3 3-3 3-4 3-4 3-5 31 A �102 Tabel 3 (cont.) Hydrologic unit Name Manual system 0012 0013 Rush Two Buttes 3-5 3-5 0002 Arkansas River Kiowa Lake Albert 3-5 3-5 3-5 11 03 04 0001 0002 0003 0004 0005 Cimarron Carrizo Ute No Fork Cimarron Arroyo Bear 0001 0002 0003 0004 0005 Rio Grande Sangre de Cristo Upper Rio Grande Alamosa San Luis Saguache Conejos 4-1 4-2 4-3 4-4 4-5 Rio Chama Chama 4-6 13 01 02 0102 14 01 4 0001 0002 0003 0004 0005 0006 6-1 6-2 6-3 6-4 6-5 6-6 0001 0002 0003 0004 0005 0006 Gunnison Taylor Blue Mesa Tomichi N. Fork Gunnison Escalante Uncompahgre 6-7 6-8 6-7 6-8 6-9 6-9 0002 0003 0004 0005 Dolores W. Dolores San Miguel Unaweep Little Dolores 6-10 6-10 6-10 6-10 03 31 A 3-5 3-5 3-5 3-5 3-5 3-5 Colorado River Upper Colorado Kawuneeche Blue Eagle Roaring Fork Grand Mesa Carr 02 Division of Wildlife 39 RG 6 32 C 34 G 33 D �103 Table 3 (cont,) Hydrologic unit Name Manual system 0106 0109 Green Vermillion Lodore 0001 0002 0003 0005 0006 0007 Colorado River Yampa Upper Yampa Milk Little Snake White Piceance Douglas 7-2 7-2 7-1 7-3 7-4 7-4 0101 0102 0104 0105 0107 0201 0202 0203 San Juan Upper San Juan Piedra Animas LaPlata Mancos Cowboy McElmo Monument 5 5-1 5-1 5-2 5-2 5-3 5-3 5-3 5-3 04 14 05 08 Division of Wildlife 35 GR 7-1 43 Y 42 W 40 SJ �104 ,....••• ; •.•..,_,' "~~1'~~ .- ~•.'•.. ~" ....."•.. ":";ti#".~.,... 5' .-_.3 ..... Salm..o.;t7. a...Lr-dne ri.. _.Q ~•.~ •••••. '••••4_ •. ....,.... .••• _""" .. "... ••.~ ••.~---. .,.-.i·":.-~·'..•. , :!t.... .' .Q.. 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('(j(KljuIIlNP ,.~ / ~W50/1(1r: (r/ ~I}flc; r;.s, 3u 1)v;.,lhd ~ "JR/)/l<j (,.1 /PII'dJCJII1'11J(' jt _S;C..Kl)//);J.iJ::~ ~tnn:ir~.s)j ~\ Z}:ltusor:(I', ]tn/,') i. l'Qrr~ otn nPp RaJ )()mfY'o ~ Ib~ ~ ";()I7vI{/~1 ]rt:1W;®~ (i; 5:i"'6/lS 't,rc,,_' J)P/J/1) ~ ..5:0 Cpr, ;V}c<fl/w~ '!)a(pS(ylcr, J c., !;:/)/'j.!..../ . .'9.;.3 fbrhm:;1/ '!J/Of;..F/ 'iJawsoJ? tb.s; 30 (}:JrrClJo//)/l'P -------..:.. ~almo 19.54 HIS ga1cdneri 5-3 Jackson Gl}l,ChJ. 13 c. jV/Q;}(,Oj,'t, 19~.2 Ih>r'r,'tll'J !;rOIA) __,C:, -' 3/ lc) Ibfl(l,)"{///.:I)</) J /'1:....1 19jC/ fblhnJl1 LYa~ /(j(/;I}C ,/11'}I"jJ 8:; 1 j JrovJ) Jb/b~itNj ,'~i'-',!. /f1/11r.1l11'vcli'.1,~'dnnL'':'· 0'1 ,.;Jr'/ / ]}(1(,tJ 1>:..ias-an . .. ,; I tfl1/!{05, 31 ,) ()J, I'ItM(I,..-, No//[oJ,3/ 3/ E I1tln(~ (.()/ttllKoJ I ~aft(~ rr /(p-: i/O; 1Ya11CO.r; 3/ Y/1/!<" (. 'U/)'I)'\,./IIJ) fb/ / .~J)' W~ -o/Tyn /0.110 ,s'liI'J;; /('/"Y; . _ ;}v ..5iJ.; /}J, ttl/n{o.) Po/./nI2/1/(l~(: ,(),.·(I·.:'v~r0~Y//JIIi:,-l:'::?J j';;;.r SInOn I ~'~:J/NJr/'" /..i ,...,; 1!..b/lc/<~j//,/,/1;:L)9·J /v, r K( / /' 30 J I -'~)}/J1/:;j,r , _j"o . J/d.;u /)f~:)f{jl)(r lY/,}/' c:. thiZI'O~/''7/'~-I}J-' _!//JIi11/i!.:p.l30 . I . , J ) ; . )51 .Ik ~ !{//~h /.. Jlcll-i_, ·.1-0Jo/1 r~ W7y~. ~ d~! ~() c;/ 0 I 'a I f-,.",tl)) Ut1(.) -:id:.IJ.;)" (/ TY/)/!. c.!... J.bc'/0J' ~ ,j d rJ If J J(J/r;:! ·ftI ]6 . / II.tJ f7'.. / .I) / J I 'i_-;,.r Jh/!lrd,l1 /{.A. .» 0ra (~ /?.; j f}a I hv-an J),a.vJ ~Ul.V;, .' /?b).._ ''1;;,'-: ~ 'J$/}/1'1 J- (/ J f1{f'ii7tJ (_~ 'or!};{;" Ja,j" ~);\ j!o/rin/./···,. '_', :..-' - '(//u' l..f-' •.. ~1-.-/,I~·/; - .,..,;,- ,/.. (M/rCl""iull1/' ': &j ;;~flIf.lf let) 3) / (~j//}1(.)IJJ7Jla4~ '/. / ;Y'/7/'), '-/ (1-)(>/::' ..:::_~·.) . J --'It.h;/q'J:'.u.,',·;,, / .. .',.;. J~ 30 • Figure 2. Example of a region card and species card for SaZma in the LaPlata River drainage. gairdneri �105 same data in the computer file, an abstract, and the keywords which describe the document. A complete annotated bibliography will be included in the job final report and will be provided upon request prior to the final'report. The documents often mentioned observations of fish in other drainages as well. The site was listed on the summarization form in enough detail so that at a later date the remaining location descriptors may be filled in. The document doe not have to be repeatedly searched and summarized. When a compilation of literature for another drainage is begun, those references already obtained would be a logical starting point. Several cards in the manual file per salmonid were required to list all observations. It was attempted to put the years in chronological order, but this was not always possible. Furthermore, to find all observations for a species in a particular year, especially after 1962, it is necessary to look at all the cards in the subdrainage of interest. It is possible that observations occurring in that year are listed on more than 1 or all the cards. This may be up to 5 cards. However, the process is still quick and easy considering the information that is obtained. Likewise several maps for the time period 1940-present were required for each salmonid species to avoid overlapping and illegibility. Published bibliographic sources searched for potentially useful documents are listed in Table 4. Non-published material found to be useful included in-house memos, scientific collecting permits, and creel census and stocking data of the Colorado Division of Wildlife (CDOW) in Fort Collins, Denver, Montrose and Grand Junction. Stream and lake survey data of CDOW are stored in a computer data base in Denver accessible by water code number of the water body. These data are not in the manual file. Prepared by Wildlife Haynes Researcher C �106 Table 4. Bibliographic sources that were searched. Aller, B. B. 1958. Publications of the United States Bureau of Fisheries, 1871 - 1940. U.S. Fish and Wildl. Servo Spec. Sci. Rep.--Fish. 284. Aquatic Biology Abstracts Aquatic Science and Fisheries Abstracts 1972-78 Bibliography for aquatic resource management of the upper Colorado River. U.S. Fish and Wildl. Servo Resour. Publi. 135. Bibliography of research publications of U.S. Bureau Sport Fisheries and Wildlife 1928-72. Bibliography of the upper Colorado River system. Servo Off. BioI. Servo U.S. Fish and Wildl. Bibliography on reservoir fishery biology in North America. U.S. Fish and Wildl. Servo Bur. Sport Fish. and Wildl. Res. Rep. 68. CLASS literature search at CSU. Biological Abstracts 1969-80. Current bibliography for aquatic sciences Index of Fishery Technological Publications 1918-55. Wildl. Servo Circ. 96. U.S. Fish and Sport Fisheries Abstracts U.S. Dep Inter. , Fish and Wild I. Servo Publi. 27 - 125. U.S. Fish and Wildl. Servo BioI. Servo Publi. 1976 - present. U. S. Fish and Wildl. Servo Circ. U. S. Fish and Wildl. Servo Tech. Paps. 1 - present. �107 JOB PROGRESS REPORT State of COLORADO Project No . N-2-R (SE-3) Identification of Habitat Requirements Work Plan No. 2 and Limiting Factors for Colorado Job No. 1 Squawfish and Humpback Chubs Period Covered: Per sonnel: 1 July 1982 to 30 June 1983 Charles M. Haynes and Gary T. Skiba, Colorado Division of Wildlife; Clarance A. Carlson and Robert T. Muth, Colorado State University. ABSTRACT A total of 694 seine and ichthyoplankton samples was collected in both the Colorado River and Yampa River study areas. A total of 16 young-of-year (YOY) Colorado squawfish (Ptychochielus lucius) was collected in the Colorado study area while 20 YOY squawfish were collected in the Yampa River study area. Estimated spawning dates were 10 July - 15 August and 6 - 28 July for the Colorado and Yampa River sites, respectively. During 1979-82, spawning was, with 1 exception, closely correlated with decreasing flows and water temperatures of 22 C. Activities in 1983 will include increased emphasis on ichthyoplankton sampling in the Yampa River study area . A tentative 1983 field schedule is presented . . ... 1ij~~1 1 ij'1l1~1 11~rn11i~m~~r1ij111r~i1l1'~1 BDOW027830 ��109 IDENTIFICATION OF HABITAT REQUIREMENTS AND LIMITING FACTORS FOR COLORADO SQUAWFISH AND HUMPBACK CHUBS Charles M. Haynes, Robert T. Muth, and Gary T. Skiba As a consequence of documented declines in numbers and ranges, the Colorado squawf ish and humpback chub (Gila cypha) have been listed as endangered by both the federal government and the State of Colorado. Suspected causes of decline have been discussed by Haynes and Muth (1982), Holden and Wick (1982), Valdez and Clemmer (1982), Haynes et al. (in press), and others. Since 1977, the Colorado Division of Wildlife has been investigating the distribution, ecology, and status of these rare fishes via systematic sampling in selected reaches of the Colorado, Gunnison, White, and Yampa rivers. Aspects of study have been conducted cooperatively with the U.S. Fish and Wildlife Service (Colorado River Fisher.ies.Project). Nongame Research (Colorado Division of Wildlife, Fort Collins) investigations have been directed toward evaluating reproductive success for both species via collection of young-of-the-year (YOY) and juveniles, evaluating distribution and timing of spawningiriselected reaches of the Colorado and Yampa rivers, and identifying habitat electivity for early life-history stages. P. N. OBJECTIVES The overall objective of this investigation is to identify the physical and biotic factors which limit the distribution and reproduction of Colorado squawfish and humpback chubs in Colorado. Similarly, this project is designed to develop field and laboratory methods for evaluating squawfish and humpback chub habitat, and reproductive success which may be used by m~nagement personnel for future habitat enhancement and/or reintroduction programs in accordance with overall ~ecovery efforts. Additionally, information derived from this and other state and federal studies may be used to evaluate the impacts of several upper Colorado River water development projects and enhance the probability of meaningful mitigation where such development projects are believed to have deleterious effects upon rare fish populations. Aspects of this investigation relating to the humpback chub will be incorporated into a doctoral dissertation in 1984-85 (R. T. Muth, Dep. Fish. and Wild!. Biol., Colo. State Univ.). SEGMENT OBJECTIVES 1. Identify, measure, and analyze habitat parameters which limit the distribution and abundance of Colorado squawf ish (Ptychochielus lucius) and humpback chubs (Gila cypha) in the upper Colorado River. 2. Identify macro- and microhabitat features (e.g., flow, temperature, substrate, depth) which are associated with presence - absence of young-of-the-year (YOY) squawfish, humpback chubs, and other associated species. �110 3. Determine probable time of spawning of squawfish and humpback chubs and correlate with data on adult fishes developed by the Northwest Region, Colorado Division of Wildlife and the U.S. Fish and Wildlife Service. 4. All collections will be made according to a computer-compatible (FWS MANAGE), detailed physical stratification system which reflects the geomorphological, hydrological, and ecological variables of the drainages. 5. Study areas will include the lower Yampa River, Moffat County from Cross Mountain Canyon to the Green River confluence in Dinosaur National Monument and the Colorado River, Mesa County from Loma to the state line. 6. Devise methods for the identification and differentiation of larval and juvenile humpback chubs, roundtail chubs (G. robusta), bonytail (Q.. elegans), and hybrids and/or intergrades. - 7. Morphomeristic counts and chub larvae and juveniles with known-age pure stock Beach and Dexter National measurements will be made on field-collected from the 2 study areas for comparison and hybrid crosses provided by Willow Fish hatcheries. STUDY AREA AND METHODS The Colorado and Yampa River study areas have been previously described by Haynes and Muth (1982) and Haynes et al. (in press) as have been the field and laboratory methods used. In 1982, in addition to methods previously described, drift-net sampling for YOY fishes was conducted in the Colorado River study area (stratum K) on 6 dates between 13 July and 26 August. Drift-net sampling was conducted in the' Dinosaur National Monument section of the Yampa River study area (strata 1 and 2) on 22-26 August. The goals of this additional survey were to evaluate techniques for sampling YOY fish in swift, deepwater habitats ineffectively sampled by seines, to determine ichthyoplankton abundance and species composition, and to evaluate drift as ··an agent of squawfish and/or humpback dispersal. Most species occurring in the Colorado and Yampa study areas exhibit peak spawning activity during early June through mid-August (Vanicek and Kramer 1969, Carlson et al. 1979, Snyder 1981, Tyus et al. 1981, Haynes and Muth 1982). Reports by Harrow et al. (1975), Gallagher ~nd Conner '{1980), and Lathrop (1982) indicated that larval fish drifting densities were greater along shoreline areas than mid-channel. This information was used to design the present drift sampling program. . 2 Near-shore surface drift net collections were made using 0.5-m diameter conical plankton nets (Wildlife Supply Co., Saginaw, Michigan) mounted on 0.5 x 0.3-m rectangular steel frames and fitted with 33-cm long removable �111 PVC collection buckets (10-cm diameter). Each net had a 560 µmesh, a length of 4 m and an open-mesh to net mouth ratio of 11:1. This design enhances self-cleaning and filtration efficiency of the net theoretically approaches 100% (Faber 1968, Tranter and Smith 1968). Attached to each net frame was a removable 4-point, steel cable bridle assembly terminating in a spring-loaded carabiner. Nets were deployed either by staking the net frames to the substrate or by fastening the bridle carabiner to a polypropylene line fixed to either an instream boulder or a metal post driven into the shore. Drift-net (ichthyoplankton) sampling in the Colorado River study area was conducted in the Black Rocks area (km 219.8 to 218.2) at dawn, dusk, and midnight using 3 nets during each collection period. Sampling duration ranged from 30 minutes to 1 hour/set depending upon accumulation of sand and organic debris in the collection buckets. A Marsh-McBirney flowmeter (Model 201) was used to measure water volume filtered by each net. An attempt to attach nets to a cable stretched the full width of the river was abandoned for safety reasons. In the Yampa River study area, only 1 net was used/site due to restricted storage space in the inflatable raft used to travel within the area. Drift sampling was conducted in conjunction with regularly-scheduled random sampling and sampling times per. date varied; however, approximately equal numbers of nocturnal and diurnal' collections were taken over the period. Sampling duration varied from 30 minutes to 7 hours depending upon the amount of organic debris and sand accumulation. Water volume filtered by the net was measured with a pygmy flowmeter (Gurley Model 625 F). Samples were preserved in 10% formalin and returned to the laboratory for sorting and analysis. Specimens were stored in 3% buffered formalin. Specimens were identified to the lowest feasible taxon, counted, and measured to the nearest 0.1-mm total length (TL). Since reliable criteria for the differentiation of YOY and juvenile chubs (including hybrids and/ or intergrades) are not yet available, all cht.tbs were listed as Gila spp. Drift rates were computed as number/1,000 m3. A nonparametric~~ permutation technique (Mielke and Iyer 1982) was performed on Colorado River data to compare drift densities (response variable), collectively, and for predominant taxa, among the 3 sampling times (treatments) over the entire sampling period. Lack of replication and sampling time variability did not permit a similar analysis of Yampa River drift data. RESULTS AND DISCUSSION A total of 694 individual collections (seine and drift net) .was made in both study areas combined (Table 1). In the Colorado River, 384 samples were collected between 5 March and 27 August, 1982. In the Yampa River, 310 samples were collected between 18 April and 23 October, 1982. Due to a rafting accident, 46 Yampa samples collected on 8-11 June, were lost. Preliminary sorting and analysis of 1982 samples are complete. Data have been stored on the Fish and Wildlife Service MANAGE database program at Colorado State University, Fort Collins. The current status of MANAGE data files from 17 March, 1981 to present are shown in Table 2. �Table 1. Distribution of 1982 Colorado and Yampa River study area field collections as number per stratum by date. ..... ..... N Field Dates Colorado River strata H K Total 5-7 May 1982 27 24 51 16-18 Jun 1982 32 31 63 12-16 Jul 1982 83 83 26-30 Jul 1982 70 70 79 38 117 138 246 384 23-27 Aug 1982 Totals 1 YamEa River strata 3 4 5 2 Total 384 18-22 Apr 1982 22 2 8-11 Jun 1982 30 16 46 25-29 Jul 1982 27 20 47 5-8 Aug 1982 54 23 77 22-26 Aug 1982 87 19 106 22-23 Oct 1982 Totals 193 80 Study Total ) Totals 27 3 4 2 1 7 7 2 1 310 310 694 ) ) �113 .~ Table 2. Data files in storage on FWS MANAGE program, Colorado State University, Fort Collins, 17 March 1981 - 30 June 1983. File name N Data included records YM 1 Yampa River study area, 17 Mar 1981, stratum 5 YM 2 Yampa River study area, 22 Apr 1981, stata 5 and 6 22 YM 3 Yampa River study area, 16-19 Jun 1981, strata 1 and 2 32 YM 4 Yampa YM 5 Yampa River study area, 11-18 Aug 1981, strata 1,2,3,4, and 5 YM 6a Yampa River study area, 18 Apr - 23 Oct 1982, strata 1,2,3,4, and 5. Number of records = 310. Colorado River study area; 5 May - 27 Aug 1982, strata H, K Rive~ 1 study area, 22-26 Jul 1981, strata 1 and 2 210 159 384 CO 6 Colorado River study area, 27 Apr - 1 May 1981, strata H and K co 7 Colorado River study area, 26-28 May 1981, strata H and K 31 co 8 Colorado River study area, 8-10 Jul 1981, strata H and K co 9 Colorado River study area, 27-31 Jul 1981, strata H and K 119 ' be separated by study a File to area and verification of records. sampl~ng 24 22 periods following Analysis of habitat associations for species other than Colorado squawf ish are incomplete and will be deferred until the final report. Similarly, morphomeristic analysis of field-collected Gila spp., while underway, are incomplete and will be def erred until the final report and the doctoral dissertation (R. T. Muth, 1984-85). Colorado Sguawf ish A total of 16 YOY squawf ish was collected in the Colorado River study area in 1982 (Table 3)~ Of these, 2 were collected by drift net on 27 July in the Black Rocks area and 14 were collected by seine on 23 August at 8 sites from km 216.1 to 244.4. All seine specimens were captured in low-velocity habitats, i.e., backwaters (10), embayments (2), and a single concavity (2). In the Yampa River study area, 20 YOY squawfish �Young-of-year Colorado squawfish collections, Colorado Table 3. a~d ........ Yampa River study areas, 1982 ,c.. Study area Date a Colorado River 27 Jul Total and range 23 Aug Total and range Yampa River 7 Aug 21 Aug 22 Aug 24 Aug 25 Aug Total and range b River Km N TL (mm) Temp {C) 219.6 219.0 1 1 2 9.3 7.8 7.8-9.3 22 22 4 3 2 1 1 1 9.8, 13.1, 13.3, 13.6 12.2, 12 .. 8, 13.9 13.2, 13.8 12 .. 3 13.4 20.1 16.8 9.3 9.3-20.1 28 9.9 10 14-16 26 20 25 11 24 244.4 244.3 243.9 243.1 238.9 224.1 218.0 216.1 18.0 17.5 15.9 5.1 0.5 19.6 19.5 16.6 13.2 3.9 1 1 T4 1 1 2 1 8 1 1 3 1 1 20 13-21 11.2 10.7 15.1, 17.1, 19.8 16.9 11.8 9.9-21 29 25 26 24 24 24 20 25 26 26 26 22 25 Habitatc Estimated a.ge 20 10 (Days) MC MC MC MC MC MC MC MC MC MC MC MC SC MC SC MC MC SC SC SC Estimated spawning period (Dates) SH SH 5-15 13-23 Jul 9-45 10 Jul-15 Aug BA BA co EM BA BA BA EM BA EM. PO co IP BA BA IP IP IP 11-48 6-28 Jul a Specimens collected by drift-net. b Samples collected by Northwest Region personnel. c Habitat descriptors as given by Haynes and Muth (1982) ) ) ) �115 were collected between 7 and 25 August at 10 sites between km 0.5 to 19.6 (stratum 1) in low-velocity habitats, i.e., backwaters (3), embayment (1), pool (2), concavity (1), and 13 in isolated pools. Age at time of capture (using methods described in Haynes and Muth (1982) and Haynes et al., (in press)) was 5-15 days post-spawning for drift collections (27 Jul) and 9-45 days for seine collections in the Colorado River study area. Estimated spawning dates in the Colorado River are 13-23 July for drift-caught animals and 10 July - 15 August for seine specimens. In the Yampa River, spawning is estimated to have occurred from 6 to 28 July 1982. Analysis of YOY Squawfish, 1979-82 Since 1979, 102 YOY squawfish have been collected in the Colorado River study area (Table 4), of which 89 were collected in Stratum Hand 13 in Stratum K. In the Yampa, only Stratum 3 (Deerlodge Park area) was sampled in 1979 while Cross Mountain Canyon (Stratum S), Lily Park (Stratum 4), and the downstream Dinosaur National Monument reach (Strata 1 and 2) were added in 1980. A total of 89 YOY squawfish has been collected in the Yampa during 1980-82 (Table 4), all of which were collected in the lowermost 30 km of Dinosaur National Monument (Stratum 1). In the Colorado River, estimated spawning dates during 1979-82 based upon maicimum and minimum size per collection and estimated growth rate ranged from 18 June (1981) to as late as 26 August (1980) (Fig. 1). In the Yampa, spawning is estimated to have occurred as early as 16 June (1980) and as late as 3 August ·(1981) (Fig. 2). In 1981, we estimated that spawning occurred in the lower Yampa as early as 25 June, which correlates closely with the dates and locations of ripe radio-marked adults observed over apparent spawning gravels in the same area by Tyus et al. (1981), i.e., 26 June - 10 July km 19.6 - 0.2. Comparative data for 1982 are not available. Fish growth-rate is temperature-dependent (De Vlaming 1972), and lower late-August water temperatures can be expected to reduce rate of squawfish growth. Therefore estimated late-summer spawning dates should probably be adjusted forward. Since growth rate-temperature data for young squawfish are lacking, it is not possible, however, to determine to what extent late-season spawning periods should be corrected. Our estimated spawning periods indicate that squawf ish spawning occurred as water levels were decreasing and water temperatures were rising (Fig. 1, 2). The possible interrelationships between spawning, flows, and water temperature have been the subject of:intense discussion and speculation and has been summarized by Haynes et al. (in press). At this time, the influence of temperature upon spawning is-apparently based upon observations of adult ripeness by Vanicek and Kramer (1969), hatchery observations by Toney (1974) and Hanunan (1981), and this report. �116 Table 4. Young-of-year Colorado squawfish collections, Colorado and Yampa river study areas, 1979-82a. Study area 1979 1980 1981 1982 Total Stratum H 8 66 1 14 89 Stratum K 0 11 0 2 13 Totals 8 77 1 16 102 Stratum 1 46 23 20 89 Stratum 2 Stratum 3b 0 0 0 0 0 0 0 0 Stratum 4 0 0 0 0 Stratum 5 0 0 0 0 46 23 20 89 Colorado River Yampa River 0 Totals aCombined collections by Nongame Research and Northwest Regional personnel. b Only stratum 3 sampled in 1979. For the purposes of this investigation, the earliest dates during 1979-82 when water temperatures reached or exceeded 18, 20; and 22 C - along with our earliest estimated spawning dates are summarized in Table 5. For those years for which temperature records are available, except 1979, spawning is closely correlated with the earliest date of record when the water temperature reached or exceeded 22 C. In 1979, spawning in the Colorado River possibly occurred as early as 29 June, coinciding with a temperature of 18 C but nearly a month prior to 22 C. This suggests that water temperature at the spawning site may differ from the site at which the temperature was measured or that 1 or more fish spawned at the lower temperature. The relationship between decreasing flows and the timing of the spawning is apparent; however, a quantifiable relationship between relative level and duration of flow and spawning success is less obvious. Further analysis will await 1983 collections and more detailed data review, including interpretation of capture-per-unit-of-effort values, and will be deferred for the final report. �) ) ) 30--~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~--30 ~ - 26 22 18 14 14 I- 10 6 10 6 2 2 0 - 0 1000 --• 0 . <1> (/) J A S AMJ J AS 26 22 18 A M J J AS lOOO 900 900 800 800 700 700 ,aoo 600 500 500 400 400 (") -wE CJ 0: <( I 0 300 (fJ 0 200 200 100 100 0 AMJJAS AMJJAS AMJJAS AMJJAS 1979 1980 1981 1982 (8) 29 June-2 Aug (77) 28 Jun-26 Aug (1) 18-24 Jun (16) 10 Jul-15 Aug ..... ..... ....... �118 30 26 30 22 22 18 14 10 6 2 18 14 10 500 - 26 6 ,., ~. A M J J A S AMJJAS A M J J A S 5()0 450 450 400 400 350 350 300 300 I 0· Q) en (") E ..__. w "' 2SO a: 250 J: 200 200 150 150 100 100 50 50 <( (.) Cf) Ci AMJJAS AMJJAS 1980 1981 (46) 16 J~ Aug (23) 25 Ju;.:3 Aug A M J. J A S 1982 (20) 6-28-Jul Figure 2. Thermographs, hydrographs, and estimated spawning dates for Colorado squawfish, Yampa River study area (1980-82). Broken thermograph line indicates missing data. Numbers in parentheses = N. ~ · ·-· · · �119 Table 5. Earliest dates of spawning-related water temperatures, Colorado and Yampa river study area, 1979-83 (U.S.G.S. Data). Records were unavailable for 17 Apr - 21 Jul 1980. Colorado River a Yampa River 1979 1980 1981 1982 Maximum >18 C 29 Jun 25 Jun .10 Apr 18 C + 30 days 28 Jul 24 Jul c 12 Jul Maximum >22 c Earliest estimated spawning date Maximum >20 b 1981 1982 27 Jun 19 Jun 28 Jun 10 May 26 Jul 18 Jul 28 Jul 26 Jun 28 May 9 Jul 20 Jun 9 Jul 22 Jul 30 Jun 20 Jun 18 Jul 24 Jun 15 Jul 29 Jun 28 Jun 18 Jun 10 Jul 25 Jun 6 Jul 1980 16 Jun ~ecords at Colorado-Utah line. bRecords at Maybell, Colorado (Moffat Co.) with exception of 1982 which were recorded at Deerlodge Park, Dinosaur National Monument. Drift Net Collections The 258 specimens collected in 61 drift samples (48 in the Colorado River and 13 in the Yampa River) were predominantly larvae (juvenile and adult forms comprised <3%) and represented 12 species and 4 families (Table 6). Of the 12 species, 6 were native and comprised 77% of the total ichthyoplankton sampled in 1982. A total of 118 specimens was collected in the Colorado River study area, representing 10 species and 4 families. Bluehead suckers (Catostomus discobolus), speckled dace (Rhinichthys osculus), and unidentified chubs (Gila spp) were the predominant native species, constituting 29, 27, and 25% of the total catch, respectively. The channel catfish (Ictalurus punctatus) was the most abundant non-native species, comprising 6% of the total. Notably, 2 larval Colorado squawfish were collected in the Colorado River study site on 27 July (Table 7). Although larvae of this endangered species have been taken on a number of occasions with fine-mesh seines and dipnets (Wick et al. 1981, Haynes and Muth 1982) this was the 1st drift-net collection of larval Colorado squawf ish throughout the entire upper Colorado River basin. �120 Table 6. Numbers, percentages of the total, and frequency of occurrence of fish collected during 1982 ichthyoplankton sampling in the Black Rocks area of Ruby Canyon (river kilometer 219.8> 218.2), Colorado River, Colorado and in Dinosaur National Monument (river kilometer 72.4 ~O.O), Yampa River, Colorado. N fish sampled Fish taxa Percent of Frequency of total number occurrence sampled (%)a Colorado River Cyprinidae Speckled dace,Rhinichthys osculus* Gila spp·. * Colorado squawfish,Ptychocheilus lucius* Fathead minnow,Pimephales promelas Common carp,Cyprinus carpio Roundtail chub, Gila robusta* 67 30 2 1 1 1 Catostomidae Bluehead sucke~ Catostomus discobolus* Flannelmouth sucker, C. latipinnis* 57 27 25 2 1 34 1 1 2 2 43 35 8 36 29 29 7 15 Ictaluridae Channel catfish,Ictalurus punctatus 7 7 6 6 10 Centrachidae Green sunfish,Lepomis cyanellus 1 1 1 1 2 86 75 61 53 54 9 6 38 1 1 1 1 8 7 4 3 5 3 2 23 47 34 34 46 46 Total 32 25 21 4 2 33 10 2 118 Yampa River Cyprinidae Gila spp.* Speckled dace,Rhinichthys osculus* Red shine~ Notropis lutrensis Sand shiner, N. stramineus Catostomidae Bluehead sucker,Catostomus discobolus* Flannelmouth sucker, C. latipinnis* Ictaluridae Channel catfish,Ictalurus punctatus Total 47 69 8 23 15 140 a Based on 48 samples for the Colorado River and 13 samples for the Yampa River. * Native species �121 For the Yampa River study area, 140 specimens were collected, representing 7 species and 3 families. The most abundant native was Gila spp., comprising 53% of the total. Channel ·catfish were the predominant non-natives, constituting 34% o_f the total catch. Collectively, ichthy9plankton density-drift rates in the Colorado River tended to be higher. in the dawn samples with a dawn: dusk: midnight abundance density ratio of almost 3:1:1 (P = 0.1476) (Table 7). Speckled dace were slightly more abundant in dawn samples with a dawn: dusk: midnight ratio .of 4:1:2 (P = 0.1774). Gila spp. exhibited higher drifting densities in the dawn samples with a dawn: dusk: midnight abundance ratio of 11:1:1 (P = 0.0064). Differences in drifting densities among the 3 sampling times were not evident for bluehead suckers (P = 0.8562). Sampling in the Yampa River was inadequate for similar analysis (Table 8). Razorback Sucker No YOY or juvenile razorback suckers (Xyrauchen texanus) were identified in either study area during 1982. Third-Year (1983) Investigations A tentative 1983 field schedule for the Colorado and Yampa River study areas is presented in Table 9. Deviations from this schedule may result from unpredictable problems in the study areas, or with gear and/or personnel. During this 3rd year, intensive drift sampling will be conducted in the Yampa River in addition to regularly-scheduled YOY sampling in both study areas. One drift net station will be installed above Harding Hole at the lower end of stratum 2 and a 2nd station will be installed in the vicinity of Boxelder Campground near the confluence (stratum 1). A maximum of 2 days will be spent at each site at times presented in Table 9 with 1-2 hour samples collected at dawn, noon, dusk, and midnight. Aknown-age larval series of roundtail chubs will be prepared using either upper Yampa or White River brood stock. Adults will be collected in mid-June with the assistance of Northwest Regional electrof ishing personnel and injected on-site with acetone-dried carp pituitary extract. Eggs will be stripped, fertilized, and water-hardened at the site. Eggs and larvae will be reared at Colorado State University, Fort Collins. LITERATURE CITED Carlson, C. A., c. G. Prewitt, D. W. Snyder, E. J. Wick, E. L. Ames, and W. D. Fronk. 1979. Fish and macroinvertebrates of.the White and Yampa rivers, Colorado. U.S. Dep. Inter., Bur. Land Manage. Colo. Biol. Sci. Ser. 1. 276pp. De Vlaming, V. L. 1972. Environmental control of teleost reproductive cycles: a brief review. J. Fish. Biol. 4:131-140. �122 Faber, D. J. 1968. Soc. 97:61-63. A net for catching limnetic fry. Trans. Am. Fish. Gallagher, R. P., and J. v. Conner. 1980. Spatio-temporal distribution of ichthyoplankton in the lower Mississippi River, Louisiana. Proc. 4th Annual Larval Fish Conf. U.S. Dep. Inter., Fish and Wildl. Serv. Biol. Serv. Prog. FWS/OBS-80/43:101-115. Hamman, R. L. 1981. Spawning and culture of Colorado squawfish in raceways. Prog. Fish-Cult. 43:173-177. Harrow, L. G., I. Cherkov, and A. B. Schlesinger. 1975. Seasonal and distributional patterns of ichthyoplankton in the Missouri River. Omaha (Nebr.) Public Power District, Environ. Ser. Bull. Haynes, c. M., and R. T. Muth. 1982. Identification of habitat requirements and limiting factors for Colorado squawfish and humpback chubs. Colo. Div. Wildl. Endangered Wildl. Inv. SE-3-4. Prog. Rep. 43pp. , T. A. Lytle, E. J. Wick, and ---squawfish Ptychochielus lucius basin, Colorado, 1979-1981. R. T. Muth. 1983. Larval Colorado Girard in the Upper Colorado River Southwest Nat. In Press. Holden, P. B., and E. J. Wick. 1982. Life history and prospects for recovery of Colorado squawfish. Pages 98-108 in W. H. Miller, H. M. Tyus, and c. A. Carlson, eds. Fishes of~he upper Colorado River system: present and future. Proc. Symp. Annu. Mtg. Am. Fish. Soc. 1981, Albuquerque, N. M. Lathrop, B. F. 1981. Ichthyoplankton density fluctuations in the lower Susquehana River, Pennsylvania from 1976 through 1980. Pages 28-38 in c. F. Bryan, J. V. Conner, and F. M. Truesdale, eds. The 5th Annu. Larval Fish Conf., La. State Univ., Baton Rouge. Mielke, P. W., and H.K. Iyer. 1982. Permutation techniques for analyzing multi-response data from randomized block experiments. Conun. Stat. Theory and Methods. 11:1427-1437. Snyder, D. E. 1981. Contributions to a guide to the cypriniform fish larval of the upper Colorado River system in Colorado. U.S. Dep. Inter., Bur. Land Manage. Colo. Biol. Sci. Ser. 3. 8lpp. Toney, D. P. 1974. Observations on the propagation and rearing of two endangered fish species in a hatchery environment. Proc. West. Assoc. State Game and Fish Comm. 54:252-259. Tranter, D. J., and P. E. Smith. 27-56 in D. J. Tranter, ed. Paris, France. 1968. Filtration performance. Pages Zooplankton sampling. UNESCO Press, Tyus, H. M., E. J. Wick, and D. L. Skates. 1982. A spawning migration of Colorado squawf ish (Ptychochielus lucius in the Yampa and Green �123 rivers, Colorado and Utah, 1981. Annu. Symp., Desert Fishes Counc., Death Valley Natl. Monument., Calif. 13:3. Valdez, R. A., and G. H. Clemmer. 1982. Life history and prospects for recovery of the humpback and bonytail chub. Pages 109-119 in W. H. Miller, H. M. Tyus, and C. A. Carlson, eds. Fishes o"f""the upper Colorado River system: present and future. Proc. Symp. Annu. Mtg. Fish. Soc., 1981, Albuquerque, N. M. Vanicek, C. D., and R.H. Kramer. 1969. Life history of the Colorado squawfish, Ptychocheilus lucius and the Colorado chub, Gila robusta, in the Green River in Dinosaur National Monument, 1964-1966. Trans. Am. Fish. Soc. 98:193-208. Wick, E. J., T. A. Lytle, and C. M. Haynes. 1981. Colorado sqauwfish arid humpback chub population and habitat monitoring, 1979-1980. Colo. Div. Wildl. Endangered Wildl. Invest., SE-3-3. Prog. Rep. 156pp. aynes Wildlife Researcher �Table 7. Ichthyoplankton drifting densities Q!/1,000 m3) by species in dawn, dusk and midnight drift samples collected during 1982 sampling in the Black Rocks area of Ruby Canyon (river kilometer 219.8~218.2), Colorado River, Colorado. Size ranges are cumulative for the entire sampling period. 26 Aug Date 13 Jul 15 Jul 27 Jul 29 Jul 25 Aug Timea DN/DK/MN DN/DK/MN DN/DK/MN DN/DK/MN DN/DK/MN Sample volume (m3) Total DN/DK/MN DN/DK/MN 444/395/412 461/280/280 333/189/296 362/3i0/412 346/346/346 362/346/346 2,30811,926/l,092 219.8, 219.0, 218.6 219.8, 218.9, 218.6 219.6, 219.3, 219.0 219.6, 219.3, 219.0 219.4, 219.2 219.4, 219.2 19 19 22 21 23 18/10/ 0 9/ 0/ 0 9/ 0/ 0 3/ 3/12 6/ 0/ 3 0/ 0/ 6 16/ 0/ 0 11/ 0/ 0 21/ 0/ 0 11/ 3/ 5 6/ 0/ 0 0/ 3/ 0 11/ 1/ 0/ 0/ 0 0/ 0/ 0 0/ 5/ 3 0/ 0/ 0 0/ 0/ 0 0/ 0/ 0 0/ 1/ 0/ 0/ 0 .~/ 0/ 0 0/ 0/ 0 3/ 0/ 0 0/ 0/ 0 0/ 0/ 0 1/ 0/ 0 0/ 0/ 0 0/ 0/ 0 3/ 0/ 0 0/ 0/ 0 0/ 0/ 0 0/ 0/ 0 1/ 0/ 0 0/ 0/ 0 0/ 0/ 0 0/ 0/ 0 0/ 0/ 0 0/ 0/ 3 0/ 0/ 0 0/ 0/ 13/ 6/ 0 7/ 4/ 4 6/11/ 0 8/ 5/19 0/ 3/ 3 0/ 0/ 3 6/ 5/ 4 .£. latipinnis (17.5-28.0) 2/ 3/ 0 0/ 0/ 0 3/ 0/ 0 3/ 3/ 7 0/ 0/ 0 0/ 0/ 0 1/ 1/ 1 Ictalurus punctatus 0/ 0/ 0 0/ 0/ 0 0/ 0/ 0 0/ 0/ 2 6/ 3/ 6 0/ 3/ 0 1/ 1/ 1 0/ 0/ 0 0/ 0/ 0 0/ 0/ 0 0/ 0/ 0 0/ 0/ 0 3/ 0/ 0 1/ 0/ 0 49/19/ 0 27/ 4/ 4 42/16/ 3 28114/45 18/ 6/15 3/ 61 9 30/11/13 River kilometer x water temp. , C 219.8-218.6 21 23 Fish taxa (total length range in nun) Rhinichthys osculus Gila spp. ( 6. 5-53. 0) (8.5-114.5) Ptychocheilus lucius Pimephales promelas Cyprinus carpio Gila robusta (9.0-9.5) (12.5) (28.0) (176.0) Catostomus discobolus Lepomis cyanellus (11.0-33.5) (11.5-74.0) (17.5) Total aThree sampling times: ) " DN a Dawn, DK a 8/ 2/ 4 Dusk, MN= Midnight. ) 111 ) �) ) ) Ill 3 Table 8. Ichthyoplankton drifting densities (N/1,000 m ) by species in drift samples collected during 1982 sampling in Dinosaur National Monument (river kilometer 27.4>0.0), Yampa River, Colorado. Size ranges are cumulative for the entire sampling period. Date Sample volume (m3 ) River kilometer 22 Aug 23 Aug 24 Aug 25 Aug 26 Aug Total 129 411 615 1,331 1,096 3,582 54.6 54.6, 25.8 25.8, 14.5 14.5, 2.7 217 54.6, 25.8, 14.5, 2.7 23 23 21 22 20 22 Gila spp. (13.5-32.5) 7 58 24 23 5 23 Rhinichthys osculus (8.0-22.5) 7 5 3 2 3 x water temp., C - I Fish taxa (Total length, range in mm) Notropis lutrensis (18.5) 1 3 1 N. stramineus (45.5) Catostomus discobolus (17.5-27.5) 3 1 .f_. latipinnis (18.5-39.5) 3 2 Ictalurus punctatus (13.5-30.0) Totals 14 ~ess than 0.5/1,000 m3 of water sampled. a 2 1 1 15 11 15 13 11 87 35 45 22 40 �126 Table 9. Tentative field schedule for 3rd year (1983) studies (Nongame Research, Fort Collins) Colorado River Study Area Jun-Jul 6-10; strata H, K; stratified random sampling 27-1; strata H, K; stratified random sampling Jul 11-15; strata H, K; stratified random sampling Aug 8-12; strata H, K; stratified random sampling 22-26; strata H, K, stratified random samping Sep-Oct 12-17; strata H, K; stratified random sampling 26-1; strata H, K; stratified random sampling Oct 12-17; strata H, K; stratified random sampling Nov 14-19; strata H, K; stratified random sampling Yampa River Study Area May 16-20; stratum 1, inspect suspected spawning bars Jun 11-18; strata 1 and 2, install drift net stations, YOY sampling Jul 4-10; strata 1 and 2; stratified random sampling, drift-net sampling 18-22; strata 1 and 2; drift net. YOY sampling Aug-Sep 1-5; strata 1-5; stratified random sampling, drift-net sampl~ng 15-20; strata 1 and 2; drift net, YOY sampling 29-2; strata 1-4; stratified random sampling, drift-net sampling Sep 19-21; strata 3 and 4, YOY sampling �127 JOB PROGRESS REPORT State of COLORADO Project No. __~N_-~3_-=R~ Work Plan No. Job No. Personnel: 1 ~~-------------1 Period Covered: _ _ Natural Historv of Brazilian Free-tailed Bats in the San Luis Valley, Colorado 1 January 1982 through 31 December 1982 S. Bissell, W. Graul, C. Haynes, and P. Svoboda, Colorado Division of Wildlife; J. Choate, Fort Hays State University, Hays, Kansas. ABSTRACT Arrival, departur~, sex composition, and estimated population size are reported from field work conducted during June through November 1982. Breeding was documented by capture of a pregnant female (29 Jun) and young-of-the-year (10 Aug - 16 Oct). Population estimates (~= 6) between 19 June and 5 September averaged 52,112 (range 11,396154,242). Arrival and departure of the colony in 1982 was from <12 June to between 16 October and 13 November. Of 22 loci examined in preliminary electrophoretic analysis, 17 were monomorphic and 5 were variable. ��129 NATURAL HISTORY OF BRAZILIAN FREE-TAILED SAN LUIS VALLEY, COLORADO BATS IN THE Peggy L. Svoboda " l The Brazilian free-tailed bat (Tadarida brasiliensis) initially interested researchers in the United States because of its possible link to the spread of rabies (Courter 1954, Davis et al. h962, Constantine 1967). Recently, interest was stimulated by declines in numbers of these bats in 2 of the large maternity colonies in the southwest-Carlsbad Caverns (Allison 1937, Altenbach et al. 1979, cited by Geluso et al. 1981) and Eagle Creek (Cockrum 1969, Reidinger 1972, cited by Geluso et al. 1981). This decline has focused attention on possibie causes of mortality, especially those resulting from pesticide poisoning because these bats regularly feed and drink in intensively-farmed areas where pesticides are used (Geluso et al. 1981). ' . I . I A little-known colony of Brazilian free-tailed bats lives in Colorado during the summer. Meacham (1975) _reported that, in August of 1968, he found about 9,000 or more ~. brasiliensis occupying a mine which had been abandoned since 1932. The most recent estim?te of the size of the population prior to this study was 75,000 to 100,000 individuals in 1981. This colony seemed to be predominately male with few fem~les and juveniles present (J. Freeman and L. Wunder, in prep.). This colony represents a major nongame wildlife resource in Colorado, with unique characteristics: (1) the colony was only recently discovered (Meacham 1975) and could be increasing in size; (2) it is unusually large for a primarily male summer roost as described by Cockrum (1969); (3) it is at the northern margin of the range of T. brasiliensis and in between the 2 easternmost populations of ~. ~. mexfcana as outlined by Cockrum (1969). Because of these characteristics, the Colorado Division of Wildlife initiated a 2-year study in 1982 to better understand the biology and ecology of this little known colony. P. N. OBJECTIVES The goal of this study is to investigate the natural history of a large and apparently isolated colony of ~. brasiliensis in Colorado. This study includes field observation of seasonal dnd daily chronology, sexual composition, reproduction, and feeding activities of the colony and laboratory analysis of genetic relat'ionships with other populations in the southwest. SEGMENT OBJECTIVES 1. Describe seasonal and daily chronology of T. brasiliensis at the �130 Orient Mine by regular in their roost. observation of the outflight arid of the bats 2. Document sexual composition of the colony by regularly outflight with 9.l-m mist nets. 3. Document reproduction in the colony by observing the condition caught in mist nets and of individuals that are collected. 4. Ascertain colony size by use of a photoestimation technique and measuring the amount of guano left in the mine by T. brasiliensis during occupation of the roost for 1 season. - 5. List food items eaten by T. brasiliensis in the San Luis Valley by analyzing stomach and guano samples taken from individuals caught in the valley or returning to the mine. 6. Locate feeding areas of T. brasiliensis by capturing in mist nets set at locations in the San Luis Valley Orient Mine. 7. Investigate the genetic relationship of the San Luis Valley colony to other colonies in southwestern United States through electrophoretic analysis of heart, kidney, and liver tissue taken from bats of the Orient Mine, Oklahoma, and Arizona. DESCRIPTION sampling the of bats individuals away from the OF STUDY AREA The colony is on the edge of the San Luis Valley, in Saguache County about 37 km southeast of Poncha Pass. The San Luis Valley is in southcentral Colorado and forms the northern part of the Rio Grande River Valley. The valley is about 160 km long and 80 km wide; its center has an elevation of about 2,300 m .. It extends into New Mexico, where there is a slight change in the topography as the valley merges with the Rio Grande depression. Poncha Pass, at the northern end of the valley, is at the junction of the San Juan and Sangre de Cristo Mountains Mld separates the Sawatch Range from the Sangre de Cristos. The Sangre de Cristos form the eastern edge of the valley, whereas the San Juan massif lies along the western edge. The colony roosts in a large, open slope created by iron m1n1ng activities in the late 1800's and early 1900's. The stope is in a hillside at an elevation of 2,880 m, about 580 m above the valley floor. The hillside has a generally southwest facing slope and is covered with Gambel oak (Quercus gambelii). North-facing slopes in the area are more mesic than south-facing slopes and are covered by coniferous forests and patches of quaking aspen q(Populus tremuloides). Vegetation in the valley is desert scrub with some areas of sagebrush (Artemisia spp .) pii'(on-juniper (Pinus edulis-Juniperous spp), and grassland. The climate is generally sunny with cool short summers and cold dry winters. This area is climatically the driest part of Colorado but is irrigated for agriculture because numerous wells and canal systems have been developed. �131 Land use patterns in the valley have been influenced by availability of water for irrigation. Chief crops are barley, wheat, alfalfa, potatoes, and cool-climate vegetables such as lettuce, cauliflower, and peas for commercial and domestic use (Ramaley 1942). The northern end and edges of the valley are used for grazing. METHODS AND MATERIALS Mist nets (4.6 and 9.l-m) were set at 10 sites in the vicinity of the Orient Mine for a total of 39.0 net nights, May through October 1982 (Table 1). Nets that were not left up for'an entire night are considered to comprise one-half a net night. Table 1. Free-tailed bat trapping effort (number of net nights), Orient Mine, Colorado, May - October, 1982. Net sites 1 2 May Jun Jul Aug Sep 2.0 0.5 4.0 1.0 3.0 2.0 0.5 1.0 2.5 1.0 3.0 3.5 2.0 1.0 0.5 1.0 2.0 1.5 3 1.5 3 4 5 6 1.0 7 8 9 10 Totals 2.5 11.5 1.0 3.0 1.0 0.5 0.5 10 10.5 Oct Totals 10.5 1.5 13.0 -;, '~,4~ 0 1.5 1.5 5.0 1.0 0.5 0.5 39.0 :.. Bats caught in nets were identified to species, sex, and age. ,Each bat was weighed to the nearest 0.5 mm with a 50-g pesola scale, and its forearm was measured to the nearest 1 mm with a metric ruler. Young-ofthe-year were identified by trans-illuminating the proximal metacarpal joint on the 4th digit of the right wing and inspecting it for presence of cartilaginous zones. Elongate joints not totally ossified on both or either side of the joint indicated the bat was young. Colony size was estimated by a modification of a photoestimation technique described by Humphrey (1971). A pentax camera with a flash attachment was placed on a tripod at the edge of the mine. As the emerging bats flew between the camera and a vertical rock wall, photographs were taken every 2 minutes; with 400 ASA Tri-X black-and-white film. Speed of the bats was measured at every 2 minute interval to the nearest 0.01 second with a digital stop watch. Any gaps in the out-flight were measured to the nearest second. Bats in each frame were counted by inspecting �132 the photographs and negatives on a light table with a magnifying glass. The number of bats per photograph was multiplied by the quantity obtained when 2 minutes minus the length of any gaps was divided by the speed of the bats across the frame. This calculation gave an estimate of the number of bats passing in front of the camera every 2 minutes. The numbers of bats in each interval were summed to obtain the number of bats passing the photographic site during the main out-flight. The main pile of guano near the roo.st was covered with black plastic on 28 June. The dimensions of the pile are approximately 16 by 6 m. The amount of guano accumulated on the plastic was measured in November 1982 after the bats had left the roost. Width and depth were measured to the nearest 2.54 cm every 1.5 m down the chute. A contour map of the new guano was obtained and amount of guano was estimated to the nearest 0.1 m3• Fifty samples of guano were obtained from free-tailed bats by placing individual bats in styrofoam cups covered with aluminum foil. Bats were held in the cups until they defecated. Stomachs were taken from·18freetailed bates. All samples were preserved in 10% formalin. Bats from which samples were obtained were either caught later in the night at the mine, presumably returning to the roost after a night of feeding, or in the valley while feeding or drinking water. Samples were inspected with a 15- to 90X dissecting microscope. Heart, kidney, and liver tissues were collected from free-tailed bats at the Orient Mine (~= 50); Merrihew Cave, Oklahoma (~= 9); and Picacho Peak, Arizona (N = 27) in August and September. Samples were placed in Nunc cryotubes and frozen in liquid nitrogen; Samples were prepared for starch gel electrophoresis at Texas Tech University, Lubbock. A subsample of 8 tissues from each of the 3 localities was analyzed for variability in 22 loci. Frequency of occurrence of electromorphs in variable loci will be statistically analyzed through the BIOSYS program. RESULTS AND DISCUSSION Seasonal Chronology Arrival and departure of the colony was investigated from 1978 to 1981 by J.' Freeman (unpubl. data). From these records and my observations for 1982, timing of arrival at this roost is variable (Table 2). In 1982, a large number of bats was present in the roost on 12 June and numbers doubled by the beginning of July. The main part of the colony departed by mid-September with reduced numbers present through mid-October. No bats were observed at the mine on 13 November. �133 Table 2. Approximate arrival and departure times of the Tadarida brasiliensis colony, Orient Hine, 1978-82. Year Arrival Departure ,.j 1978 <17 Jul Mid-to lat~ Sep Reference \ j ~ 1979 >27 Jul Hid-to late Sep 1980 Between 30 Jun and 3 Jul Between 2 Jul and 26 Jul <12 Jun Hid-to late Sep 1981 1982 i Hid-to late Sep ! Reduced nu~bers by early Sep, completely gone by ~l 13 Nov I I " J. Freeman (unpubl. data) J. Freeman (unpubl. data) J. Freeman (unpubl. data) J. Freeman (unpubl. data) Svoboda (this report) Composition' Composition estimates of the colony at.e based upon total catch at the mine: 86.3% adult males, 11.1% adult females, and 2 .. 6% young-of-theyear (Table 3). Adult males comprised 88.6% of all adult captures, lower than the 94.5% reported by J. Freeman and L~ Wunder (unpubl. data) for previous years. Host of the young were caught in September and October when the proportion of males caught in the initial out-flight decreased. Increased proportions of adult females and young in the outflight occurred coincidentally with an overall reduction in colony size and a more dispersed flight pattern. Dispersed flights common at the beginning of the summer contained few adult f ema.Las • Seven other species of bats were caught at the mine (Table 4). However, I caught only ~. brasiliensis during the main out-flight. I Table 3. Age and sex composition of free-tailed bats captured at the Orient Hine, Colorado, 1982. N captured 707 91 17 l:. Total 819 Age Sex Percent 'of total Adult Male Female Male Female 86.3 11.1 2.1 0.5 Young 100 �134 Table 4. Other species of bats caught at the Orient Mine, 16 June - 15 October 1982 and net sites 3, 4, 5, and 10. ; Adult male Species Myotis lucifugus (Little Brown Bat) M. evotis (Long-eared Myotis) M. volans (Long-legged Myotis) M. leibii (Small-footed Myotis) Eptesicus fuscus (Big Brown Bat) Lasiurus cinereus (Hoary Bat) Plecotus townsendii (Townsend's Big-eared Totals N individuals Adult female Totals 2 2 1 1 4 3 l ,I f i' 1 5 1 4 1 1 3 3 10 10 Bat) 24 2 26 Reproduction At the Orient Mine on 29 June, the female in the sample superficially appeared to be pregnant. On 10 August, a lactating female and a juvenile were caught and a 2nd juvenile was caught ou 18 August. At least some females in this colony probably are bearing young about mid-July; this has been documented by J. Freeman who caught pregnant females on 3 July 1980 (unpubl. data). If true, by mid-to late-September most young bats 'Would be about 2 months old. AJ:. that time it is difficult to distinguish young-of-the-year from adults. If a space existed on either side of the joint, I considered the bat to be young. The 2 young bats caught in mid-October weighed less than the rest of the sample, i.e., 8.5 and 9 g as compared to an average of 11.9 g for adult females and 12.3 g for adult males caught on the same night. Colony Size Six photographic estimates of population size were made between 19 June and 5 September 1982. Approximately the same number of bats appeared to emerge from the roost in early September as in mid-June. Maximum numbers of bats in the main out-flight occurred in late July and August. Average number of bats was 52,112 (range 11,396 - 154,242). 3 The amount of guano left beneath the entrance to the roost was 3.3 m after 3 51 days (28 Jun - 18 Aug) and 10.4 m after 139 days (28 Jun - 14 Nov). �135 Food Habits Samples of guano (N = 50) and stomachs (N = 18) from free-tailed bats await final analysis. In preliminary processing, lepidopteran scales are present, which is to be expected according to other studies (Ross 1961, Freeman 1981) (Table 4). Table 4. Number of free-tailed bat.stomach and guano samples in the San Luis Valley, Colorado, summer 1982. N sam12les Stomach collected Totals Locality Guano Orient Mine Valley View Hot Springs Mineral Hot Springs Baca Grande Golf Course Kerber Creek Freel's Ranch Raybe's irrigation pond 22 15 4 3 1 1 1 1 4 4 2 1 Totals 50 18 68 9 5 2 4 31 20 6 Although several feeding areas for free-tailed bats were identified~ a primary feeding area for the colony has yet to be found. Adult male T. brasiliensis (N= 56) were caught at 7 locations in the San Luis Valley from 2.4 to 25.7 km from the mine (Table 5). Nine other locations were netted in the valley from 11.7 to 85.7 km from the mine, but no T. brasiliensis were caught (Table 6). T~ble 5~ 'Tadarida brasiliensis caught away from the Orient Mine, Saguache County, Colorado, 1982.. (All individuals were adult males) • Date Location N Distance (km) from colony Direction 15~ 21 Jul 22 Jul Valley Vie~7 Hot Springs Mineral Hot Springs Kerber Creek Baca Grande Golf Course Raybe's Irrigation Pond Freel's Ranch Bunker's Pond 24 10 10 7 3 1 1 2.4 10.5 14.5 25.7 7.6 8.0 4.0 S WSW WNW SSE WSW WN1.J SSE 1 Aug 27 Jul 16 Jul 31 Jul 3 Aug Totals 56 �136 Table 6. Locations netted in the San Luis Valley, Colorado, 1982, where Tadarida brasiliensis was not captured Distance (km) from colony Direction Alamosa National Wildlife Refuge . Denton Spring, G.S.D.N.M.ab Wetherill Property, D.O.W. 85.7 S 61.9 38.6 SE SW Lazy VK Estates 12.9 SW Maxwell's pond Bosarge Ranch Clayton's irrigation pond Angell Ranch Iridian Springs 14.5 11.7 23.3 29.8 61.9 SSW NW SSW NW SSE Date Locality 1 Jun 10 Jun 8 Jul 11 Aug 22 Jul 3 Aug 3 Aug 29 Jul 3 Aug 8 Aug 12 Aug a b Great Sand Dunes National Monument Division of Wildlife Electrophoresis I obtained distinct electrograms on 22 loci. Of these loci, 17 were monomorphic and 5 were variable; 2 other loci require further testing at a different pH level to obtain more distinct electrograms. Variable loci were esterase-2, phosphoglucomutase-l (PGM-l), PGM-2, 6-phosphogluconate dehydrogenase, and tetrazolium oxidase. In July 1983, I will test all 'tissue samples for all variable loci to discern whether there is more gene flow between the Orient Mine colony and 1 or the other of the southern populations as a clue to the geographic affinity of the Colorado colony. This work will also test the hypothesis that populations with different migrational patterns are genetically, as well as behaviorally, distinct as suggested by Cockrum (1969). LITERATURE CITED Allison, V. C. 18:80-82. 1937. Evening flight from Carlsbad Caverns. J. Mawnal Altenbach, J. S., K. N. Geluso, and D. E. Wilson. 1979. Population size of Tadarida brasiliensis at Carlsbad Caverns in 1973. Pages 341-348, in H. H. Genoways and R. J. Baker, eds. Biological investigations in the Guadalupe Mountains National Park, Texas. U.S. Dep. Inter., Natl. Park Servo Proc. and Trans. Ser. 4. �137 Cockrum, E. L. 1969. Migration in the guano bat, Tadarida brasiliensis. Univ. Kansas, Mus. Nat. Hist., Misc. Publ. 51:303-336. Constantine, D. G. 1967. Activity patterns of the Mexican free-tailed bat. Univ. New Mexico Pub1. BioI. 7. 79pp. Courter, R. D. 1954. Bat rabies, Public Health Rep. 69:9-16. Davis, R. B" C. F. Herreid II, and H. L. Short. 1962. tailed bats in Texas. Ecol. Monogr. 32:311-346. Mexican free- Freeman, P. W. 1981. Correspondance of food habits and morphology in insectivorous bats. J. Mamma~ 62:166-173. Geluso, K. N., J. S. Altenbach, and D. E. Wilson. 1981. Organochlorine residues in young Mexican free-tailed b.a ts from several roosts. Am. Midland Nat. 105:249-257. Humphrey; S. R. 1971. Photographic estimation of population size of -the Mexican free-tailed bat, Tadarida brasiliensis. Am. Midland Nat. 86:220-223. Meacham, J. W. 1975. A Colorado colony of Tadarida brasiliensis. Bat Res. News 15:8-9. Rama1ey, F·. 1942. Vegetation of the San Luis Valley in southern Colorado. Univ.' Colorado Studies, Ser. D 1:231-277. Reidinger, R, P .• J:r .•• 1972. rectors influencing Arizona bat population levels. Ph.D. Thesis. Univ. Arizona, Tucson. 172pp. Ross, A. J.- 1961. 42 :66"':'71. Prepared bY~~ Appr-oved by Notes on the food habits of bats. _ J. Mammal. � https://cpw.cvlcollections.org/files/original/80461f0a0e0255db644be1bc7a0c8db3.pdf b4ea2742f51b0da21455372da13e1c12 PDF Text Text JOB FINAL REPORT State of Colorad~ ------~~~~------------- Project No. W-37-R-36 Game Bird Survey Work Plan No. Job No. Job Title: Evaluation 22 --------~~---------- of Nesting Cover Preferences of Pheasants in Relation to Wheat Farming Methods Per iod Covered: April 1982 through 30 June 1983 Clait E. Braun and Warren D. Snyder, Colorado Division of Wi ldl ife. Personne 1: ABSTRACT The objectives of this study have been fully attained and are reported the series of reports listed below. in Snyder, \1. D. 1981. Nest habitat selection by ring-necked pheasants in northeast Colorado. Midwest Fish and Wildl. Conf., Wichita, Kans. 43:Abstract. 1982. Evaluation of nesting cover preferences of pheasants in relation to wheat farming methods. Final Rep., Colo. Div. Wildl. Fed. Aid Proj. W-37-R-35, Pp. 1-57. 1982. Recommended habitat management practices for pheasants eastern Colorado. Colo. Div. Wildl. Game Inf. Leafl. 82. 4pp. 1983. Review. 1983. in A second crop from wheatfields. Colo. Outdoors. Under Phea san t ; Co lor ado l s ring-necks. Colo. Country Life 30(7) :4,9-10. 1983. Pheasant nesting ecology in relation to wheat farming. Gray Partridge/Ring-necked P~easant Workshop. Campbellsport, Wis. (Synopsis to be published.) 1984. Ring-necked pheasant nesting ecology and wheat farming on the High Plains. J. \.Jildl.Manage. 48:ln Review. 1984. Survival of radio-marked Wildl. Manage. In Prep. Prepared by: Warren D. Snyder Wildlife Researcher hen ring-necked pheasants. J. ��3 JOB PROGRESS REPORT State of Colorado --------------------------- Project No. Work Plan No. Job Title: 3 Job No. 13 Responses of Sage Grouse to Vegetation Fertilization Per~od Covered: Personnel: Game Bird Survey W-37-R-36 1 July 1982 through 30 July 1983 R. A. Ryder, Colorado State University; Kevin Berner, Clait Braun, len Carpenter, Kurt Hundgen, Steve Porter, Tom Remington, Joan Ritchie, Tom Schoenberg, lynn Stevens, John Wagner, Carol Ann Weinland, Colorado Division of Wildlife. ABSTRACT Sage grouse (Centrocercus urophasianus) food selection, diet quality, and energy reserves were studied in North Park, Colorado during January-April 1981-82. Radiotelemetry was used to locate flocks of feeding birds for observations and locations of feeding sites. Big sagebrush (Artemisia tridentata spp.) comprised 97% of the shrubs at feeding sites and 87% of the shrubs at random sites. Canopy cover ranged from 19% at random sites to 22 and 27% at feeding sites in 1981 and 1982, respectively. Sagebrush height averaged 24 cm at random sites and 17 and 30 cm at feeding sites in 1981 and 1982, respectively. Variation in cover and height of sagebrush at feeding sites between years was related to snow depth. Wyoming big sagebrush (A. t. wyomingensis) comprised 86% of the sagebrush at feeding sites and 48% at random sites. Mountain big sagebrush (A. t. vaseyana) comprised 12 and 41% of the sagebrush at feeding and random sites, respectively. Ninety percent of the plants identified as fed-upon by sage grouse were Wyoming big sagebrush. Mountain big sagebrush (7%) and alkali sagebrush (A. longiloba) (3%) were the only other plants fed-upon. Wyoming big sagebrush (ATW) contained more (p < 0.05) protein (14.1 vs. 10.8%) and a lower (~< 0.05) level of monoterpenes (1.2 vs. 2.8%) than mountain big sagebrush (ATV). There was significant (p < 0.05) variation in protein content within sub-species, fed-upon plants> non fed-upon plants> random plants. Monoterpene content did not vary (p > 0.05) between fed-upon, non fed-upon, and random plant samples, at least within ATW. Ether extract levels were not related to sage grouse food preferences. Discriminant �4 function analyses were used to identify variables discriminating between fed-upon, non fed-upon, and random plants within sub-species of big sagebrush, and to quantify the magnitude of group differences. Plant vigor and protein content distinguished between fed-upon, non fed-upon, and random ATW plants. Group distinctions were weak, as only 31% of the variation in plant vigor and protein content was related to feeding status (fed-upon, non fed-upon, random). Within ATV an oxygenated monoterpene, protein, and plant vigor weakly (38% of the variation explained) distinguished between fed-upon, non fed-upon, and random plant samples. Good separation (81% of the variation explained) of fed~upon and non fed-upon plant samples was evident after random samples were el iminated. All (17 of 17) ATV cases were correctly assigned to fed-upon or non fed-upon categories based on their content of 3 oxygenated monoterpenes and protein. Crop and gizzard contents differed (p < 0.01) in chemical content. Gizzard contents contained less protein, cell contents, ash, and minerals, and more ether extract, neutral and acid detergent fiber, and lignin than crop contents, apparently because of partial digestion of leaf material in the gizzard. Fat content varied among and within sex and age-classes of sage grouse. Adults had higher (p < 0.05) fat contents than juveniles (4.67 vs. 2.87%), and birds collected in 1982 had more (p < 0.05) fat than birds collected in 1981 (4.03 vs. 3.43%). Fat content increased (p = 0.06) from early to late winter. Estimated energy reserves of sage grouse would last from 3 to 4.5 days for fasting juveniles, 4-6 days for fasting adult females, and 5-8 days for fasting adult males. Petroleum ethet extracted less (p < 0.001) fat than diethyl ether (3.60 vs. 4.04%). Diethyl ether extracts were predictable (p < 0.001, r2 = 98.4%) from petroleum ether extracts. Nutritional quality of-the winter diet, and by inference, requirements of sage grouse were higher than those of other grouse. The lack of a grinding gizzard may explain how sage grouse can eat a diet of 100% sagebrush in winter and why they must. Abstract of M.S. Thesis. Prepa red by _'--=fk.,.--_-....:..;4=-'~6:::;::.-::~~~·~t:J~tnv~~Thomas E. Remi ngton 1Tt.t-) Approved by !!Iad:5:~ Clait E. Braun �5 JOB PROGRESS REPORT State of Colorado Project No. W-37-R-36 Work Plan No. Job Title: 3 Job No. Dispersal and Recruitment Period Covered: Personnel: Game Bird Survey 14 of Chick Sage Grouse 1 Apri I 1982 - 30 June 1983 Clait Braun, Peter Dunn, Ken Giesen, Jerry Hupp, Colorado Division of Wildlife, Ronald Ryder, Colorado State University. ABSTRACT Natal dispersal, lek fidelity, and recruitment of sage grouse (Centrocercus urophasianus) were studied on Cold Spring Mountain, Moffat County, Colorado, from July 1981 through June 1983. There was no difference between male and female natal dispersal distances from juvenile capture to breeding sites, although the sample size was small (N = 25). Sixty percent of yearling grouse (birds 8-10 months old) were observed displaying on the lek closest to their juvenile banding location. Sixteen percent of all individuallymarked juveniles (25/157) were known to have recruited to leks as yearlings. There was no difference between yearling and adult female lek attendance rates, however, yearling males attended leks less often than adult males. All yearlings visited 2 or more leks more often than adults. These differences may be related to yearlings' inexperience with breeding. Fall dispersal movements were analyzed from 331 relocations of radio-marked juveniles. Grouse steadily moved away from capture sites until November each year. Movements to wintering areas occurred in late November and were related to snowfall and subsequent availability of sagebrush (Artemisia spp.). There were no differences between male and female fall movements. Sage grouse movements generally followed topographic features, although they were capable of long-distance (23 km) movements over areas without sageb rush cove r. M.S. Thesis prepared as a series of papers. Prepared by Q~ tP.~ --~~~~~~~~~~-----------Peter O. Dunn (1~) Approved by Clait E. Braun ��7 LATE SUMMER - SPRING MOVEMENTS OF JUVENILE SAGE GROUSE PETER O. DUNN, Department of Fishery and Wildlife Biology, Colorado State University, Abstract: Fort Collins, CO 80523 Late summer to early spring movements of radio-marked juvenile sage grouse (Centrocercus urophasianus) were studied on Cold Spring Mountain, Moffat County, Colorado from July to February 1981-82 and August 20 May 1982-83. Movements were analyzed from 118 locations (N during July-November August-November 1981 and 213 locations (N 1982. = = 8 grouse) 10 grouse) during Grouse steadily moved away from capture sites until November each year when they moved to winter-use sites. Movements to wintering areas in late November were related to snowfall and subsequent availability of sagebrush (Artemisia spp.). wintering areas was 30.3 km (N differences = 4 Maximum 1-way distance to radio-marked grouse). in fall movements between males and females. There were no Sage grouse movements generally followed topographic features, although they were capable of long-distance (23 km) movements over areas without sagebrush cover. Key words: movements, Centrocercus urophasianus, Colorado, dispersal, emigration, sage grouse. Spacing behaviors may produce changes in birth, death, and movement rates which subsequently limit population size (reviewed in Watson and Moss 1970, Krebs 1978). Red grouse (Lagopus lagopus scoticus) are a classic example of the importance of summer and fall movements the breeding population (Watson and Moss 1980). to recruitment Although the natal into �8 movements of some species of grouse have been examined Marshall (Godfrey and 1969, Bowman and Robel 1977, Keppie 1979, Jamieson and Zwickel 1983), movements of juvenile sage grouse from summer ranges to breeding areas have not been studied. important for understanding Knowledge of sage grouse movements may be grouse population dynamics and for mitigating adverse human impacts on habitat. This paper describes the late summer to early spring movements of juvenile sage grouse in northwestern Colorado. STUDY AREA AND METHODS The study was conducted on Cold Spring Mountain in northwestern Moffat County, Colorado and in adjacent Wyoming and Utah from July 1981 through June 1983. The study area is semi-arid sagebrush rangeland with inter- spersed quaking aspen (Populus tremuloides) and pinyon pine (Pinus edulis)Rocky Mountain juniper (Juniperus scopulorum) stands, and meadows. Lodgepole pine (Pinus contorta) and Douglas-fir occur above 2,620 m on Middle Mountain (2,904 m) and Diamond Peak (2,909 m) on the northern edge of the study area. receives 46-51 cm of precipitation (U.S. Dep. Inter. 1978). (Pseudotsuga menziesii) Cold Spring Mountain (2,622 m) annually, of which 11% falls in August From 120 to 300+ juvenile sage grouse were banded on Cold Spring Mountain during July and August each year from 1978 through 1982. In 1981 and 1982, juvenile grouse were captured and individually-marked with numbered aluminum bands and unique combinations of colored plastic bands. Drive traps, a bumper-mounted cannon net, and spotlights and long-handled nets were used to capture grouse (Giesen et al. 1982). Captured birds were classified to sex and age by wing molt and primary �9 length (Beck et al. 1975). poncho-type markers Forty-two radio transmitters attached to (Amstrup 1980) were placed on juvenile grouse during July and August 1981 and July through October 1982. Thirteen solar-powered radio transmitters were used in 1981, while 23 battery-powered solar-powered radio transmitters were used in 1982. and 6 Radio-marked birds were relocated at least 3 times weekly from 15 July to 31 August 1981 and from 1 August to 11 September 1982. At least monthly made after 11 September to 15 November each year. mid-April 1981, 4 radio-marked 1 time/year. (~4 During January to juvenile birds were relocated at least Weekly radio relocations were evenly divided hours after sunrise), mid-day before sunset), and evening « into morning (> 4 hours after sunrise to > 4 hours 4 hours before sunset) time periods. time periods coincide with activities data). relocations were such as feeding and roosting (unpubl. A 3-element yagi antenna was used to relocate radio-marked on the ground. An aircraft with 2 strut-mounted the ground were determined by triangulation U.S. Geological grouse yagi antennas was used to locate birds on 3 November 1981 and 15 January 1983. 200 m) or by flushing birds. These Radio-locations on at close range (approximately Bird locations were plotted on 7.5-minute Survey topographic maps using Universal Transverse Mercator coordinates. Movement data with 2 categories were analyzed with Mann-Whitney tests. ~ Distances moved by grouse from their capture sites were first analyzed with 2-way analysis of variance individual birds. (ANOVA) for interactions among If the interaction was significant, capture site for each bird was individually distance from regressed on other variables �10 (age of bird, date, or movement non-parametric rate). Directions were analyzed with tests (Batschelet 1978). significant at the 0.05 probability Statistical tests were considered level. RESULTS Fifteen and 43 juvenile sage grouse were radiomarked respectively. in 1981 and 1982, Locations of 8 birds in 1981 and 10 birds in 1982 were used for analysis because of loss of radio signals or transmitters, hunter harvest, or incomplete data. predation, Movements were analyzed from 118 locations in 1981 and 213 locations in 1982. During both years grouse steadily moved away from their capture sites during August to November (Fig. 1). Because of unequal sample sizes and interaction effects (2-way ANOVA, ~ = 0.01) between individual birds and time periods, it was not possible to determine when increases in distances from capture sites occurred. However, the data (Fig. 2) indicate that increases were gradual during both years. vary with date in either year (Table 1). Movement rates (km/day) did not Distance from capture sites was most highly correlated with date for 13 of 18 birds (Table 1). Three birds showed no relationship between distance from capture sites and date, age of bird, or movement rate. 1 August to 4 September 1982,82 During 15 July to 4 September 1981 and and 69% of all locations (~= 135, 106), respectively, were ~ 2 km from capture sites, while during 3 October to 14 November, were> 77 and 78%, respectively, of all radiolocations 2 km from capture sites. (~= 13, 36) �11 To determine covariance if females move farther than males, analysis of (ANCOVA) was used which allows control of the effect of date on distance moved while testing for a difference females. However, the assumption be made for either year (Partial of parallel f. between males and regression test, ~ > 0.05). lines could not In 1981, females were farther from their capture sites than males on any date after 10 August, while in 1982 males were farther from capture sites than females on any date after 1 August. Because ANCOVA is inappropriate lines and males and females alternately each year, the most parsimonious of no difference without parallel moved farther than the other sex action was to accept the null hypothesis in distances moved between males and females during August to Novembe r. Movements of all but 3 (! random orientation, 18) birds in 1981 and 1982 had a non- or mean direction, P < 0.00) (Fig.2). from capture sites (Rayleigh's test, There was no mean direction when individual means of all birds were combined However, = (Mardia's correlation the restriction that! coefficient, ~ > 0.05). should be equal for all birds was violated, although "a sl ight variation of ! would hardly damage the statistical analysis" (Batschelet range 15-28, ~ = 1978:6) (1981: range 8-16, ~ = 13, CV = 0.23; 1982: 21, CV = 0.24). Males and females did not differ in mean direction moved in either year (Mardia-Watson-Wheeler text, P > 0.05). Except for 1 male grouse (#501) found on Cold Spring Mountain on 21 January 1982, the last 2 radiolocations 1981 were on 14 November. of juvenile grouse marked in Male #501 and 35 other grouse were located in �12 exposed sagebrush on 21 January 1982, but no grouse were seen after 2 February following a snowstorm which covered all exposed sagebrush. Fall 1982, 7 grouse were radiotracked until 14 November. In One radio-marked adult hen was located on its wintering area 5 km northeast of Cold Spring Mountain on 15 December 1982. A juvenile male (#734, Fig. 3) which was missing from 14 November 1982 to 14 January 1983 moved to its subsequent wintering area during 23 to 30 October 1982. The number of radio-marked grouse with known locations in 1982 decreased from 17 on 30 October to 7 on 14 November and to 0 on 25 November. During this period snow levels on Cold Spring Mountain increased from 0-5 cm to 20-50 cm. snow depth was> covered. By 14 December, 1 m and all sagebrush on sage grouse summer range was Some birds appeared to leave summer range in late October-November; 9 of 58 radio-marked grouse were lost prior to 3 October. None of these birds was later relocated, so I do not know if they were dispersing earlier than other grouse or whether there were other reasons for loss of radio signals. However, 4 of the 9 birds had moved relatively far (> 3 km) when they were lost. Four juveniles (3 males, 1 female) radiomarked in Summer 1982 were located in January and February 1983 after being lost during late-November to mid-January. January-April Maximum distance from capture sites to their locations in 1983 averaged 18.2 km for these 4 birds. The female grouse moved a maximum of 30.3 km from her capture site, while the 3 males moved from 11.4 to 17.3 km from their capture sites (Fig. 3). radio-marked grouse were found wintering in 2 areas: Color-banded and area was a 200 km2 �13 sagebrush flat 5-30 km northeast of Cold Spring Mountain; the other area (25 km2) was 7 km southwest of Cold Spring Mountain along the Utah-Colorado boundary (Fig. 4). A radio-marked male (#694) in the latter wintering area moved 23 km to the wintering area northeast of Cold Spring Mountain between 2 February and 26 March 1983 and attended Gee Flats Lek from 4 April to 19 May (Fig . 3). DISCUSSION Late summer movements of juvenile sage grouse on Cold Spring Mountain were not characterized the population, by rapid, synchronized movements by most members of as Godfrey and Marshall (Bonasa umbellus). (1969) found for ruffed grouse Instead, male and female sage grouse movements on Cold Spring Mountain are sporadic throughout September and October, similar to greater prairie-chickens (Tympanuchus cupido) (Bowman and Robel 1977) and sage grouse in Idaho (Dakle et al. 1963). Movement rates on Cold Spring Mountain did not change during September-November because movements> 2 km were quick (about 1 day) and separated by periods of up to 20 days when birds did not move> 0.3 km/day. The lack of difference in movements between male and female sage grouse suggests that if there are differences between the movements of the sexes that affect recruitment, they probably occur in Spring similar to the greater emigration of female than male spruce grouse (Dendragapus canadensis) follow topographic (Keppie 1979). Sage grouse seem to features since mean angles of sage grouse movements were generally along the northwest to southeast orientation of Cold Spring Mountain and away from its steep (80% slope) southwest face. However, sage grouse may cross large areas without cover as did radio-marked male #694 which crossed Cold Spring Mountain when all sagebrush on the mountain was snow covered. �14 Synchronized, long-distance movements during mid-November to mid-January porarily lost during that time. to winter range may have occurred since most radio-marked birds were tem- Field observations indicated that sage grouse generally moved north to sagebrush flats and lower valleys as snow levels increased in November. Schoenberg Patterson (1952), Dalke et al. (1960), and (1982) reported that sage grouse movements between wintering and breeding (nesting areas and leks) areas were related to snow level and its effect on the availability of sagebrush. Movement distances probably varied depending on the distance to suitable cover above snow. (1952) and Dalke et al. (1960) reported 1-way movements Patterson from summer to winter ranges of up to 160 km in Wyoming and Idaho, respectively. In Colorado, winter to breeding range distances averaged 28-30 km for 4 radio-marked grouse (4 females, 3 males) (Schoenberg 1982) and 8-12 km for color-banded grouse (68 males, 10 females) traveling from breeding to winter range (Beck 1977). The 1 other study of fall movements of radio-marked sage grouse (both adults and juveniles were tracked until 30 Nov) found that grouse generally leave summer range in October and November (Connelly and Markham 1983). From 10 July to 7 September, 95% of all radiolocations < (N = 131) were 2 km from the general capture area, while 82% of all locations (N were> 2 km during October to November. results from Cold Spring Mountain. = 22) These findings are consistent with Three of 14 radio-marked grouse in Connelly and Markham's study moved from summer range prior to mid-September, while most birds did not leave until after 1 October. or post-hatching emigration may increase a yearling's if there is a lower probability of establishing in other areas (Watson and Moss 1980). This early dispersal chances of recruitment a territory in natal than �15 During Summer 1982, 1 yearling female and at least 2 chicks may have exhibited post-hatching emigration as described by Watson and Moss (1980). From hatching during 25-27 June until 7 August, the hen and chicks were < 2.5 km from the nest site. On 7 August the female and chicks (45-47 days old) were 5.2 km from the nest site and the next day were 9.6 km from the nest. The female was last seen with a chick on 17 August and started moving back towards the nest site on 12 September. last located 2.2 km from the nest site on 3 October. radio-marked The hen was In contrast, 1 other female with a brood remained within 2 km of her nest site until loss of the radio transmitter on 8 August and 1 of her chicks, which was radio-marked, September. did not move> Among 7 radio-marked 2.1 km from the nest site until 12 females which were unsuccessful nesters, maximum distance from nests averaged 3.8 km prior to 20 August (range 1.3- 6.5 km). Juvenile sage grouse produced on Cold Spring Mountain dispersed an area covering at least 874 km2 in northwestern Utah and Wyoming. Understanding Colorado and adjacent movements of grouse in such a large area is important for proper management tats. of populations In this case, protection of summer brood-rearing directly benefit wintering during February-late into and seasonal habiareas would not grouse as none was found on summer ranges April each year. It is important to know if grouse on Cold Spring Mountain can be managed as 1 population or whether birds are recruited into other nearby populations. From the results of radio- tracking, recaptures of banded birds, and hunter band recoveries throughout �16 Moffat County (C. E. Braun, unpubl. data), it appears that grouse produced on Cold Spring Mountain return to breed in the study area. However, hunting pressure and trapping effort within 30-50 km of Cold Spring Mountain during the study interval were light or lacking. LITERATURE Amstrup, CITED S. C. 1980. A radio collar for game birds. 1978. Second order statistical J. Wildl. Manage. 44:214-217. Batschelet, E. Pages 3-24 in K. Schmidt-Koenig migration, analysis of directions. and W. T. Keeton, eds. navigation and homing. Spring-Verlag, Animal Berlin, West Germany. Beck, T. D. I. selection ---- 1977. Sage grouse flock characteristics in winter. J. Wildl. Manage. , R. B. Gill, and C. E. Braun. 1975. sage grouse from wint characteristics. Leafl. 49 (revised). 41 :18-26. Sex and age determination Colo. Div. Wildl. Game Inf. 4pp. Bowman, T. J., and R. J. Robel. mobility, and habi tat 1977. Brood break-up, dispersal, and mortality of juvenile prairie chickens. J. Wildl. Manage. 41 :27-34. Connelly, J. W., and O. D. Markham. concentrations 1983. of sage grouse in Idaho. Movements and radionuclide J. Wildl. Manage. 47:169- 177 . Dalke, P. D., D. B. Pyrah, D. C. Stanton, J. E. Crawford, and E. F. Schlatterer. 1960. sage grouse in Idaho. Conf. 25:396-407. Seasonal movements and breeding behavior of Trans. North Am. Wildl. and Nat. Resour. of �17 and ------- 1963. tivity, and management of sage grouse in Idaho. Ecology, produc- J. Wildl. Manage. 27:811-841. Giesen, K. M., T. J. Schoenberg, and C. E. Braun. trapping sage grouse in Colorado. Godfrey, G. A., and W. H. Marshall. of ruffed grouse. Jamieson, Keppie, D. M. J. Wildl. Manage. 33:609-620. regulation. J. Wildl. Manage. 43:717-727. A review of the Chitty hypothesis of population 1952. The sage grouse in Wyoming. 1982. Sage grouse movements and habitat selection in North Park, Colorado. M.S. Thesis, Colo. State Univ., Fort 86pp. u.S. Department of Interior. 1978. A supplement to the Northwest Colorado coal regional environmental Bur. Land Manage. DES 76-21. Watson, A., and R. Moss. aggression 1970. u.S. Dep. Inter., 555pp. Do~inance, spacing behavior, and A. Watson, ed. to their food resources. and statement. in relation to population limitation in vertebrates. Pages 167-220 ~ ------- , Sage Books, Denver. 341pp. Schoenberg, T. J. Collins. Dispersal and site fidelity Can. J. Zool. 56:2463-2480. Patterson, R. L. Colorado. 1983. Dispersal, overwinter mortality, and recruitment of spruce grouse. 1978. Brood break-up and dispersal Can. J. Zool. 61:570-573. 1979. Krebs, C. J. Methods for Wildl. Soc. Bull. 10:224-231. 1969. I. G., and F. C. Zwickel. in blue grouse. 1982. ------- 1980. Animal populations in relation Blackwell Sci. Publ., Oxford, U.K. Advances in our understanding of the population dynamics of red grouse from a recent fluctuation numbers. Ardea 68:103-111. in �Table 1. R2 and probability (p)a of simple and multiple regression models of sage grouse distances from capture sites (km) with movement rate from previous locations (m/day), date, and age of bird on Cold Spring Mountain, 15 July 14 November 1981 and 1 August - 14 November 1982. ...• 00 Distance from capture sites R2 (P) Year Bird 1981 438 465 486 501 585 687 712 785 Sex M F M M . F M F M AIJ. b..l:r.ds 1982 520 562 576 669 694 702 712 734 757 797 l\ 11 birds N locations 14 10 16 18 13 11 18 18 Movement rate 0.01 0.01 0.01 0.11 0.00 0.00 0.12 0.00 (>0.5) (>0.5) (>0.5) ( 0.30) (>0.5) (>0.5) ( 0.4) (>0.5) Date 0.22 0.77 0.76 0.38 0.66 0.56 0.75 0.57 (0.25) (0.00) (0.00) (0.01) (0.00) (0.01) (0.00) (0.00) Age 0.22 (0.25) 0.77 (0.00) O.6I! (0.00) 0.38 0.66 0.56 0.75 0.57 (0.01) (0.00) (0.01) (0.00) (0.00) 118 M F F M M M F M M M 28 18 27 15 21 22 17 15 28 22 213 0.00 0.48 0.08 0.27 0.06 0.14 0.02 0.04 0.02 0.02 (>0.5) ( 0.00) ( 0.15) ( 0.04) ( 0.27) ( 0.09) (>0.5) ( 0.47) (>0.5) (>0.5) 0.59 0.10 0.34 0.37 0.00 0.74 0.67 0.56 0.56 0.10 (0.00) (0.20) (0.00) (0.02) (>0.5) (0.00) (0.00) (0.00) (0.00) (0.16) 0.60 0.08 0.34 0.45 0.00 0.73 0.67 0.56 0.56 0.09 (0.00) (0.25) (0.00) (0.00) (>0.5) (0.00) (0.00) (0.00) (0.00) (0.16) Best mult~ple model None 0.77 0.76 0.38 0.66 0.56 0.75 0.57 (0.00) (0.00) (0.01) (0.00) (0.01) (0.00) (0.00) Va riab 1es in multiple model None Date or age Date Date Date Date Date Date 0.44 (0.00) Date 0.60 0.69 0.34 0.78 None 0.74 0.67 0.56 0.56 None Age Movement rate, age Date or age Age, movement rate None bate Date or age Date or age Date or age None (0.00) (0.00) (0.00) (0.00) (0.00) (0.00) (0.00) (0.00) 0.59 (0.00) Date and movement rate �19 Fig. 1. Distance (km) moved from capture sites by 8 radio-marked sage grouse in 1981 and 10 grouse in 1982 on Cold Spring Mountain, Moffat County, Colorado. and below the mean. locations. Horizontal line is the mean. The box is 1 SE above Number above the box is the number of radio �20 N 4 KM w E s Fig. 2. Mean angles and mean distances moved (km) from capture sites of radio-marked juvenile sage grouse (~= Colorado, July-November 18) on Cold Spring Mountain, 1981 and August-November are males; dashed lines are females. 1982. Solid lines �21 WYOMING WHISKEY D.RAW COLORADO * & MIDDLE W MTN J: « •.... ;:, .L!:'::I_ DIAMOND W PEAK GEE N ~ Fi~. 3. o CAPTURE * LEK SITE *~T~~*734 o I SWEDE 234 FLA..(:::_> 5 KM Summer to spring movements of 4 radio-marked juvenile sage grouse (3 males; 1 female, #452) on Cold Spring Mountain, Moffat County, Colorado 9 August-28 May 1982-83. �22 NATAL DISPERSAL AND LEK FIDELITY OF SAGE GROUSE PETER O. DUNN, Department of Fishery and Wildlife Biology, Colorado State University, Abstract: Fort Collins, CO 80523. Natal dispersal and lek fidelity (attendance within and between years) of sage grouse (Centrocercus urophasianus) were studied on Cold Spring Mountain, Moffat County, Colorado, from July 1981 through June 1983. There was no difference between male and female natal dispersal distances, although the sample size was small (~= 25). Sixty percent of the marked yearling grouse (birds 8-10 months old) re-identified attended the lek closest to their juvenile banding location. percent of all individually-marked have attended leks as yearlings. and adult female lek attendance (25/157 birds) were known to rates, however, yearling males attended All yearlings visited 2 or more leks These differences may be related to yearlings' inexperience with breeding. attended Sixteen There was no difference between yearling leks less often than adult males. more often than adults. juveniles in spring More females and possibly more yearlings leks with higher maximum numbers of males. Key Words: Colorado, sage grouse, Centrocercus lek fidelity, recruitment. urophasianus, dispersal, �23 Dispersal has a major role in population Krebs et al. 1976) and distribution regulation (Taylor and Taylor 1977). also have a role in the evolution of song dialects 1978), sex (Shields 1982), mating systems of local populations importance, (Lidicker 1962, It may (Baker and Mewaldt (Greenwood 1980), and the stability (Reddingius and den Boer 1970). Despite its potential studies of avian dispersal are few, most research has investi- gated fall movements, not movement from natal to initial breeding areas. Dispersal of species with lek mating systems has been examined few birds and to a limited extent, even though polygamous show exceptions in only a species might to the general pattern of greater female dispersal birds (Greenwood 1980). in Sage grouse are an ideal species to study the effect of a lek mating system on dispersal because they have 1 of the most highly developed dominance hierarchies 1978, Stiles and Wolf 1979). lek fidelity among lekking species (Wiley This paper describes natal dispersal and (attendance within and between years, and moves between of a population of sage grouse in northwestern Colorado. leks) tested the hypothesis that female sage grouse follow the typical avian pattern of dispersing farther than males. I also predicted that yearling grouse (birds 8-10 months old) would show natal philopatry and that among males, yearlings would have a low rate of lek fidelity because they are firsttime breeders. STUDY AREA AND METHODS The study was conducted on Cold Spring Mountain in northwestern Moffat County, Colorado and adjacent parts of Wyoming and Utah from July 1981 �24 through June 1983. The study area is semi-arid sagebrush (Artemisia spp.) rangeland with interspersed quaking aspen (Populus tremuloides) pinyon pine (Pinus edulis)-Rocky stands, and meadows. Mountain juniper (Juniperus scopulorum) Lodgepole pine (Pinus contorta) and Douglas-fir (Pseudotsuga menziesii) occur above 2,620 m on Middle Mountain (2,904 m) and Diamond Peak (2,909 m) on the northern edge of the study area. Spring Mountain and (2,622 m) receives 46-51 cm of precipitation which 11% falls in August (U.S. Dep. Inter. 1978). have been conducted on the area since 1978. Cold annually, of Sage grouse studies From 120 to 300+ juvenile sage grouse have been banded during July and August each year. In 1981 and 1982, juvenile grouse were captured and individuallymarked with numbered aluminum bands and unique combinations plastic bands. Drive traps, a bumper-mounted with long-handled of colored cannon net, and spot-l ighting nets were used to capture grouse (Giesen et al. 1982). Captured birds were classified length (Beck et al. 1975). to sex and age by wing molt and primary During March through May 1982 and 1983, searches were made of leks with spotting scopes for birds individually banded during that spring and previous summers. birds on leks were recorded and straight-line Observations of marked dispersal distances between the juvenile banding location and the lek on which the bird was observed displaying (attended) as a yearling were calculated. w-re used as an approximation marked before long-distance unpubl. data, this study). of a juvenile's (> 2 km) movements Banding locations natal area since most were took place (Wallestad 1971; For birds which attended more than 1 lek, the lek attended the most or where mating occurred was used in calculating dispersal distances. �25 Lek attendance was calculated as the percentage of days that a bird was observed on a lek divided by the total days that males or females, depending on the sex of the particular bird, were observed on the lek. Attendance was corrected for birds captured during each spring breeding season (26 adult males, 5 adult females, 5 yearling males) by subtracting the number of days of the respective sex's days of lek attendance to the bird's capture, from the denominator. prior For example, adult male #9947 was captured and banded on Gee Flats Lek on 11 April 1983 after 6 days of lek observations during which males were seen. Therefore, the total days that males were seen on Gee Flats (37) minus the days prior to banding equals the number of days (31) in the denominator. bird #9947 = 18 days seen on the lek + 31 = 58%. Lek attendance Recruitment, as the number of birds entering the breeding population, dividing the number of yearlings number of juveniles for defined was estimated by seen on leks each spring by the total banded the previous summer. for known mortality by subtracting at the 0.1 probability Recruitment was adjusted hunter harvest and known predation from the number of banded juveniles. significant (6) Statistical tests were considered level. RESULTS Natal Dispersal There were no differences in dispersal distances from natal to breeding areas between 1982 and 1983 for either males or females (Kruskal - Wallis test, ~ > 0.1); consequently data from both years were combined. The rate of resighting of males and females was probably equal since the sex ratio of juveniles at banding was nearly equal (0.94 in 1981 and 0.87 in 1982; X2 = 2.76, ~ = 0.09) and almost equal numbers of marked individuals of each sex were seen each spring on leks (3 females and 2 males in 1982, 10 females and 10 males in 1983). �26 Dispersal distances of yearling males and females did not differ (Mann-Whitney ~ test, ~ > 0.1), although the sample size was small (N = 25). The median distances moved by males (7.3 km) and females (10.3 km) in 1983 suggest that females may actually disperse farther. Male and female yearling sage grouse also did not differ in tendency (Fisher's exact test, ~ > ~.1) to attend the closest lek to the location of their banding as juveniles. Sixty percent of all yearling grouse with known recruitment (15/25) attended the lek closest to their natal area. Recruitment Eight known leks were on the study area. on 2 previously-used leks in 1982 or 1983. Grouse were not observed Observations were made on 2 active leks in 1982 and 6 active leks in 1983 (2 leks were found late in May 1983 and only observed on 2 days) (Table 1). Direct recruitment rates of yearlings were 16% (25/157 individually-marked juveniles) for both years, and 12 (5/41) and 17% (20/116) for 1982 and 1983, respectively. data for 1982 and 1983 did not differ {! test, P Lek attendance the data were combined (Table 2). 0.1) so > Females had a lower attendance rate than males for both yearlings and adults (ANOVA, least significant test LSD, attendance ~ < 0.1). There was no difference between yearling and adult rates for females, but the difference was significant (ANOVA, LSD, ~ < 0.1). difference for males Because the mean number of days is not corrected for birds which were banded during the breeding season, and because lek attendance rates are less biased, the most parsimonious use results from lek attendance rates. action was to only �27 Most yearling grouse were recruited at Gee Flats Lek (56%), Beaver Basin Lek had the next most (24%), and Sugarloaf Lek was 3rd (16%). This sequence of leks is also the same for highest to lowest maximum number of attending males (Table 1). Although the sample size is small, these data suggest that yearlings selectively trend is not evident, however, recruit to larger leks. if one compares recruitment numbers of males on leks (Table 1). attending This same rates to mean Maximum numbers of males and females leks in North Park, Jackson County, Colorado during 1974-79 (C. E. Braun, unpubl. data) were analyzed to determine if there was a trend among females to recruit to leks with larger maximum numbers of males; these data from Cold Spring Mountain were not adequate for testing. Research methods in North Park were similar to those used in this study: intensive counts (> 8 counts/lek/year) Park. were made at 21 leks in North Using these data, maximum number of attending females was positively correlated with maximum number of attending males (r2 = 0.44, P = 0.00, N = 57 lek counts). Ten adult sage grouse (6 males, 4 females) were seen on leks during Spring of both 1982 and 1983. Of these, 1 male (17%) and changed leks attended between years. female (25%) No interlek movements were known to have been made in 1982, while 14 of 91 (15%) individually-identified were seen at more than 1 lek in Spring 1983. Forty-eight grouse percent of all interlek movements were from Sugarloaf Lek to Gee Flats Lek (7.5 km). Thirty percent of all movements were' in the opposite direction to Sugarloaf. Thirteen percent of all movements were from Gee Flats to Beaver Basin (13.1 km); there was non in the opposite direction. There was only movement each (4%) from Sugarloaf to Whiskey Draw Lek (12.2 km) and Whiskey Draw to Beaver Basin Lek (10.9 km), both by 1 yearling male. �28 Ye~rling g~ouse made more (33% interlek movements age-classes however, than adults. 3/10 for males, 20% 2/10 There were no differences in the number of interlek movements for females) among sex and per grouse (ANOVA, P > 0.1), the samp 1e size was sma 11 (N = 25). DISCUSSION In birds, females generally move greater distances than males between natal areas and initial breeding sites (reviewed by Greenwood Explanations 1980). for the sex bias in dispersal have been based on the type of mating system (resource defense [monogamy] generally female dispersal and mate defense [polygamy] generally male dispersal) leads to greater (Greenwood 1980), the promotion of inbreeding and subse- quently the maintenance (Dobson 1982). leads to greater of sex (Shields 1982), and competition I could not falsify the hypothesis for mates that female sage grouse do not disperse farther than males, however, the small sample size and the slight tendency for females to disperse farther than males lead me to refrain from accepting or rejecting the null hypothesis. Because 60% of all yearling grouse attended their natal area lek, sage grouse may be philopatric; both of these hypotheses will require larger sample sizes for adequate testing. Unfortunately, research to acquire adequate dispersal studies require long-term sample sizes: Jamieson and Zwickel (1983) (see also Zwickel 1983) recorded dispersal distances of 99 blue grouse (Dendragapus obscurus) after 10 years of study, and Keppie (1979) observed movements of 25 yearling spruce grouse (Dendragapus juveniles after 3 years of study. canadensis) banded as �29 After natal dispersal, birds are generally faithful to the initial breeding area in successive years (Greenwood 1980). have discussed the establishment territories. technique of and fidelity of sage grouse to lek Studies by Jenni and Hartzler examined the effect of variable (Patterson 1952). Few investigators (1978) and Emmons (1980) lek attendance on a commonly used lek count Wiley (1978) suggested that most females breed their first year, while yearling males rarely mate, even though they are physiologically mature. Almost all matings are performed by adult males which return yearly to the same lek and territory within the lek (Wiley 19 These findings suggest that adult males may have a greater reproductive investment in their lek territories than yearling males. greater adult than yearling male lek attendance of yearling males making interlek movements The reproductive In this study, and the greater percentage support this hypothesis. success of male grouse which occupy territories at or near the mating center may be high (Davies 1978), although maintenance these territories may be energetically expensive Males which regularly defend territories center as opportunities peripheral territories (Beck and Braun 1978). and move them toward the mating arise may have greater reproductive males which are non-territorial of fitness than or visit several leks and only establish (Davies 1978). Average male reproductive success may be greatest when yearlings visit several leks to find 1 suitable for territory establishment and then retain territories reaching the mating center as an adult. on that lek until ). �30 The only other study of male lek attendance rates of 85% for 13 radio-marked yearlings and 92% for 17 radio-marked adults; these rates did not differ. The difference studies may be due to Emmons' determination lation of radio signals. (Emmons 1980) reported in results between of lek attendance In this study, grouse at the periphery of a lek and in heavy cover may not have been recorded as attending. radio-telemetry, near leks. Emmons, using would have been able to identify grouse hidden in sagebrush This may explain the large differences attendance. by triangu- between studies in lek However, physical presence near a lek does not necessarily mean that a bird held a territory on a lek. After mid-May in 1982 and 1983, grouse were regularly seen sitting within 200 m of the periphery of Gee Flats Lek but they were not counted as attending the lek because they were not seen displaying. Yearling males may be moving among leks more frequently grouse because they are inexperienced investigate than other at breeding and may need to several leks before establishing a territory. Other studies have also indicated that yearling males make most interlek movements. Dalke et al. (1960, 1963) found that yearling males account for most interlek movements. C. E. Braun and T. D. I. Beck (unpubl. rep., Colo. Div. Wildl. Fed. Aid Proj. W-37-R-29, 1976) found 14 of 27 (52%) of year- ling and 4 of 48 (15%) adult males made interlek movements Colorado. movements in North Park, Emmons (1980) reported that male yearlings made more interlek than adult males (all 13 radio-marked yearlings visited 2 or more leks). The lack of difference in lek attendance females and the greater percentage of yearling ing 2 leks suggests that yearlings adults spend on 1 lek. between yearling and adult than adult females attend- spend as much total time on 2 leks as Yearling attendance and interlek movements may be �31 greater than that of adults if yearlings need to examine several leks prior to mating because of inexperience or if they need to receive behavioral cues for mating by watching older females (Wiley 1978). there were no differences Because between yearling and adult females in lek atten- dance in this study. or a study with radio-marked females (Petersen 1980). need for behavioral cues does not appear to lead to a lengthening of lek attendance by yearlings. The percentage of yearling females (7%. 2/27) engaging in interlek movements (20%. 2/10) vs. adult suggests that yearling females may actually be examining more leks than adults prior to mating. Petersen (1980) observed 2 of 14 (14%) yearling and 1 of 11 (9%} adult females attending 2 leks. Lek observations from Cold Spring Mountain and North Park suggest that more females and possibly more yearlings maximum numbers of males. recruit to leks with higher Numbers of females attending a lek may be a com- promise between a female preference for more males. since larger numbers of males allow greater choice among mates with relatively expenditure less energy (Emlen and Oring 1977). and an opposing male selective force against clustering on leks which. wiht increasing numbers of competing males. probably reduces mean male reproductive fitness (Bradbury 1981). The low percentage of variation explained by the correlation between maxi- mum numbers of females and males on leks may be due to many factors influencing lek attendance. For example. male attendance related to life history parameters is probably (life expectancy and rate of turnover on leks) which affect lifetime fitness. the probability of mating using �32 alternative mating strategies, mate choice (Bradbury 1981). and the degree of uniformity Nevertheless, in female there appears to be a rela- tionship between numbers of males and females on leks which deserves further study. LITERATURE CITED Baker, M. C., and L. R. Mewaldt. dispersal in white-crowned Evolution 32:712-722. 1978. Song dialects as barriers to sparrows Beck, T. D. I., and C. E. Braun. 1978. (Zonotrichia leucophrys nuttalli). Weights of Colorado sage grouse. Condor 80:241-243. ---- , R. B. Gill, and C. E. Braun. 1975. Sex and age determination of sage grouse from wing characteristics. Inf. Leaflet 49 (revised). Bradbury, J. W. Alexander 1981. Colo. Div. Wildl. Game 4pp. The evolution of leks. and D. Tinkle, eds. Pages 138-169 ~ R. Natural selection and social behavior. Chi ron Press, New York, N.Y. Dalke, P. D., D. B. Pyrah, D. C. Stanton, J. E. Crawford, and E. F. Schlatterer. 1960. sage grouse in Idaho. Conf. Seasonal movements and breeding behavior of Trans. North Am. Wildl. and Nat. Resour. 25:396-407. ---productivity, and management , and ----- 1963. Ecology, of sage grouse in Idaho. J. Wildl. Manage. 27:811-841. Davies, N. B. 1978. Pages 317-350 ~ ecology. Ecological questions about territorial J. R. Krebs and N. B. Davies, eds. Blackwell Sci. Publ., Oxford, U.K. behavior. Behavioral �33 Dobson, F. S. 1982. male dispersal Competition in mammals. Emlen, S. T., and L. W. Oring. for mates and predominant Anim. Behav. 30:1183-1192. 1977. evolution of mating systems. Emmons, S. R. 1980. Colorado. Ecology, sexual selection and the Science 197:215-223. Lek attendance of male sage grouse in North Park, M.S. Thesis, Colo. State Univ., Fort Collins. Giesen, K. M., T. J. Schoenberg, and C. E. Braun. trapping sage grouse in Colorado. Greenwood, P. J. 1980. Wildl. Soc. Bull. 10:224-231. in 1983. Dispersal and site fidelity Can. J. Zool. 61 :570-573. Janni, D. A., and J. E. Hartzler. 1978. impl ications for spring censuses. Keppie, D. M. Methods for Anim. Behav. 28:1140-1162. I. G., and F. C. Zwickel. in blue grouse. 1982. 69pp. Mating systems, philopatry and dispersal birds and mammals. Jamieson, juvenile 1979. Attendance at a sage grouse lek: J. Wildl. Manage. 42:46-52. Dispersal, overwinter mortality, and recruitment of spruce grouse. J. Wildl. Manage. 43:717-727. Krebs, C. J., I. Wingate, J. LeDuc, J. A. Redfield, M. Taitt, and R. Hilborn. 1976. Microtus population biology: populations of M. townsendii. Lidicker, W. Z. 1962. dispersal in fluctuating Can. J. Zool. 54:79-95. Emmigration as a possible mechanism permitting the regulation of population density below carrying capacity. Am. Nat. 96:29-33. Patterson, Colo. R. L. 1952. The sage grouse in Wyoming. Sage Books, Denver, 341pp. Petersen, B. E. 1980. Breeding and nesting ecology of female sage grouse in North Park, Colorado. Fort Collins. 86pp. M.S. Thesis, Colo. State Univ., �34 Reddingius, J., and P. J. den Boer. illustrating Oecologia stabilization 1970. of animal numbers by spreading of risk. 1982. Philopatry, inbreeding, and the evoluation State Univ. New York Press, Albany. Stiles, F. G., and L. L. Wolf. mating behavior Monogr. 27: 1979. population Ecology and evolution of lek Ornithol. 78pp. mechanics. u.S. Department 1977. of Interior. 1978. Bur. Land Manage. DES 76-21. o. 1971. A supplement migration and to the Northwest statement. U.S. Dep. Inter., 555pp. Summer movements broods in central Montana. 1978. Aggregation, Nature 265:415-421. Colorado coal regional environmental Wiley, R. H. of sex. 245pp. in the long-tailed hermit hummingbird. Taylor, L. R., and R. A. J. Taylor. R. experiment 5:240-284. Shields, W. G. Wallestad_ Simulation and habitat use by sage grouse J. Wildl. Manage. 35:129-136. The lek mating system of the sage grouse. Sci. Am. 238:114-125. Zwickel, F. C. to breeding 1983. range. Factors affecting the return of young blue grouse Can. J. Zool. 61 :1128-1132. �Table 1. Sage grouse lek observations, Gee Flats Cold Spring Mountain, Beaver Basin Moffat County, Sugarloafa Colorado, March-May Whiskey Orawa Swede Flatsa 12(7) 16 (7) 20(20 May) 21 (25 May) 1982-83. Cold . a 5 pring 1982 Male attendance ~ 13 (7) (SO) Max N (date) Total days b 26(9) 39(26 Apr) 30 (13 Apr) 27 23 Female attendance ~ (SO) Max ~ (date) Total days Observation days (N)c 20(22) R2 (21 Apr) 20 23(23) 66(08 Apr) 21 32 27 1983 Male attendance 30 (8) 22 (15) ~ (SO) Max N (date) Total days 42(23 Apr) 15 (R) 26(10 Apr) 39(07 May) 9 37 31 4 (1) 5(25 May) 13 2 2 1 (2) 3 ( 1) o 3(25 May) Female attendance ~ 9(11) (so) Max ~ (date) Total days Observation days (N)c 10(18) 7(5) 14(07 May) 42(23 Apr) 35 45 88(10 Apr) 26 8 10 35 4 2 o o 17 2 2 8(28 Apr) W \J1 aSugarloaf, Whiskey Draw, and Swede Flats leks were found Two other leks had no birds in 1982 or 1983. bOnly includes days when observers cine Iudes dav s whpn hi rrlc: IAIP rp nnt were present ",-pcpnt r." at leks. 1 AI"", in 1983. Cold Spring lek had no birds in 1982. �36 Table 2. Sage grouse lek attendance (%, x days), Cold Spring Mountain, Moffat County, Colorado, March-May 1982-83. Age (yrs)a <1 2 3 4 All adults -x 33 46 51 63 43 SO 28 31 42 26 2-50 3-74 16-96 3-96 Sex and parameter Male attendance Range N x 19 number of days b 7.0 6 16.2 3 52 9.0 10.8 Female attendance x 11 5 SO 6 3 Range 3-25 N x 13 number of days a Unknow~age b 2.2 6 8 6 3-9 3-25 12 30 1.7 1.9 birds were excluded from the age columns, but included under all adults. bThe mean number of days a bird was seen on a lek is an uncorrected values for birds banded during the Spring. corrected. Percent lek attendance is �37 JOB PROGRESS REPORT State of Colorado ---------------------------Game Bird Survey W-37-R-36 Project No. 9 Work Plan No. Characteristics Job Title: Job No. 7 and Habitat Preferences of Wintering Populations of Blue Grouse Period Covered: Personnel: 1 April 1982 through 31 March 1983 C. E. Braun, D. J. Freddy, J. J. Jeanson, R. W. Hoffman, S. F. Steinert, Colorado Division of Wildlife; A. L. Cade, B. S. Cade, K. A. Medve, R. A. Ryder, Colorado State University. ABSTRACT Twenty-four of 64 banded blue grouse (Dendragapus obscurus) were equipped with radio transmitters at 2 study sites (Green Mountain and Whiteley Peak) in Middle Park, Colorado during 1982-R3. In addition, 10 radio-marked grouse surviving from 1981-82 were monitored in 1982-83. Movements of 5 banded and 32 radio-marked grouse during 1980-1983 were analyzed. Males arrived on breeding territories in late March-early April. Territory abandonment was initiated in late June and continued through July. Males localized in coniferous habitat during summer before moving to wintering areas in October and November. Females arrived at breeding areas in late April and early May. Brood hens stayed within 2.0 km ·of their nest sites until early August. Some females migrated from the study areas in August, others did not leave until September, and some remained through winter. Females arrived at wintering areas in October. Resident grouse, those remaining within 3.0 km of their breeding sites year-round, moved 0.1 to 2.8 km (median = 0.6 km, N = 16). Median distance moved by migrant grouse, those moving> 3.0 km from their breeding sites for winter, was 11.5 km (range 5.0-29.5 km, N = 13). Adult and yearling males moved greater distances (median = 11.1 km, range 1.0-29.5 km, N = 10) than adult and yearling females (median = 0.7 km, range = 0.1-28~0 km, N = 19), resident and migrant grouse combined (p = 0.009). Winter home ranges of 10 adults (median = 3.0 ha, range =-1.3-11.4 ha) were larger (p < 0.01) than those of 3 juveniles (median = 18.7 ha, range 9.2-42.2 ha)~ Lone grouse were observed 23 times; 8 were males, 12 were females and 3 were of unknown sex. �38 Forty-nine flocks were observed ranging in size from 2-21 birds (x = 5.3). Both sexes were present in at least 19 flocks. Habitat characteristics were quantified in O.Ol-ha plots for 118 grouse locations on the study sites in 1982-83 and 86 locations in 1981-82. Douglas-fir (Pseudotsuga menziesii) was the predominant species in 8 occupied stands at Green Mountain and 3 at Whiteley Peak. Grouse sites at Green Mountain ranged from low density (1.7 trees/O.Ol ha) stands of large conifers (X Dbh = 50 cm) to high density (7.7 trees/O.Ol ha) stands of large(x Dbh = 23-30 cm) or small (x Dbh = 14-19 cm) conifers. Large diameter Dbh = 2231 cm) conifers were-present in low (2.8 trees/O.Ol ha) to high density (7.1 trees/O.Ol ha) conifer stands occupied by blue grouse at Whiteley Peak. Occupied sites at both study areas were on moderately steep to steep slopes (16-45°), at upper elevations, and occurred on all aspects. Douglas-fir comprised 100% of trees occupied at Green Mountain and 84% at Whiteley Peak. Mean dbh of all trees occupied at Green Mountain was 36 cm and differed (p < 0.001) among occupied stands. Mean dbh of aJl trees occupied at Whiteley Peak was 47 cm and did not differ (p = 0.94) among stands occupied. Mean age of trees occupied at Green Mountain ranged from 63 to 268 years and from 153 to 315 years at Whiteley Peak. Habitats occupied by blue grouse wintering off the study areas included spruce/fir (Picea engelmannii/Abies lasiocarpa), lodgepole pine (Pinus contorta), and lodgepole pine-spruce/fir mixes. ex �39 CHARACTERISTICS AND HABITAT PREFERENCES OF WINTERING POPULATIONS OF BLUE GROUSE Brian S. Cade P. N. OBJECTIVES The primary objectives of this study are to: (1) identify the vegetational and structural components of blue grouse winter use sites and (2) determine the spatial relationship between blue grouse wintering and breeding sites. SEGMENT OBJECTIVES 1. Review available literature concerning habitat description and measurement, habitat use by blue grouse, winter ecology of grouse, use of radio telemetry on birds, and other pertinent literature relating to grouse. 2. Conduct breeding and production surveys of blue grouse on the study areas from late April through mid-August using established techniques. 3. Band samples of male and female grouse on the study areas. 4. Equip 10 grouse at each study area with radio transmitters. 5. Relocate radio-equipped grouse to ascertain movement patterns from breeding and brood areas to winter use sites and to document habitat use preferences, movements, social organization, and period of use while on wintering areas. 6. Conduct weekly searches of the study areas from November through March keeping records as to: (a) date, location, and vegetation type where grouse are observed, (b) sex composition and size of winter flocks, including the sex and total number of encounters with lone birds, (c) location of feeding and/or roosting trees, (d) position of the grouse in the tree and its specific behavioral activities at the time of observation, and (e) birds captured and previously marked birds identified. 7. Equip wintering grouse on the study areas with transmitters not used during the spring and summer or with transmitters instrumented birds that died. recovered from �40 8. Document pattern of winter use on the study areas in relation to vegetative and structural features and measure habitat variables at grouse locations. 9. Identify major habitat types on the study areas and calculate the area encompassed by each type. 10. Measure habitat characteristics occupied and unoccupied types. at randomly located plots within 11. Compile data, analyze results, and prepare progress report. DESCRIPTION OF STUDY AREAS Field work was conducted on 2 areas of differing habitat types in northcentral Colorado. Both areas, Green Mountain and Whiteley Peak, are within Middle Park, about 161 km west of Denver. The study areas were described by Cade (1982). METHODS Blue grouse were banded on the study areas throughout the year with most captures during the breeding and brood-rearing periods (Apr-Aug). Grouse were located with tape-recorded calls and/or trained pointing dogs (Hoffman 1981), captured with a telescoping noose pole (Zwickel and Bendell 1967), and banded with serially-numbered aluminum bands and colorcoded, anodized bands for individual recognition (Hoffman 1981). Banding locations were determined to the nearest 50 m as Universal Transverse Mercator (UTM) grid coordinates. Two types of 150-151 MHz radio transmitters were placed on grouse, a solar capacitor-assisted unit (indefinite life) or a lithium battery-powered unit (10 month life). Transmitters were attached with a poncho collar (Amstrup 1980) or a backpack harness (Brander 1968). Mounted weights were 20-22 gms for the solar units and 29-31 gms for the battery-powered units. Radio-marked grouse were relocated at least once every 2 weeks while they remained on the study areas. Lack of accessibility during winter hindered regular relocation of radio-marked grouse off of the study areas. An attempt was made to visually locate all birds wintering off the study areas at least once during winter (Nov-Mar). Radiolocation was achieved with a 3-element, hand-held yagi antenna. Locations were determined by visual observation of radio-marked grouse and recorded to the nearest 50 m as UTM grid coordinates. �41 Weekly searches of the study areas were conducted from November through March. Search time was primarily spent in coniferous habitats. All coniferous areas on the study sites were searched for grouse and/or grouse sign. Searches were concentrated in areas where grouse sign, dropping and/or tracks were observed or where grouse had been previously observed. Habitat characteristics were quantified on 0.01-ha circular plots centered on trees in which grouse were observed (Stauffer 1983). Plot locations were recorded to the nearest 50 m as UTM grid coordinates and occupied trees were marked with colored flagging and serially numbered, aluminum tags. Physiographic variables recorded at plots included: slope (level, 1-15, 16-30, 31-45, 46-60°), aspect (8 cardinal directions), elevations (to nearest 30 m), and relative position on the slope (ridge top, upper, mid, or lower one-third of slope, valley bottom). Variables quantified for each tree species within the plot included: stem density of trees (~7 cm dbh), stem density of saplings « 7 cm dbh), dbh of trees, and canopy height (height of tallest tree). Basal areas calculated from dbh measures were used to define species dominance. Point-centered quarter measurements (Cottam and Curtis 1956) from the plot-centered tree to the nearest tree in each of 4 quadrants were used to calculate mean tree spacing. The coefficient of variation of mean tree distances was used as an index of spatial heterogeneity of trees (Roth 1976). Height of trees sampled by the point-centered quarter method was recorded in 1982-83 but not in 1981-82. Height, dbh, and species were recorded for the plot center tree in which grouse were observed. Trees were assigned to 1 of 3 form categories: pyramidal, overmature, or deformed. In 1982-83 the ages of occupied trees were ascertained from cores taken with an increment borer. When possible, height, aspect, and position of the grouse in the tree were recorded. Total area of coniferous, deciduous, and shrub types were measured with a planimeter from enlarged aerial photographs. Coniferous tree stands were classified on the basis of species composition and relative stem density. Quaking aspen (Populus tremuloides) stands and shrub types were not classified into more specific categories. Area corrections were determined for habitat types that occurred on steep slopes. Area corrections were not applied unless they were 10% or greater. Coniferous forest stands were sampled with randomly located 0.01-ha circular plots. Sample sizes were 10 plots for stands < 10 ha and 20 plots for stands> 10 ha. The same physiographic and habitat variables recorded at grouse locations were recorded at random plots. Random plots were used to ordinate conifer stands based on forest stand characteristics. �42 Conifer stands were classified as occupied or unoccupied based on winter observations of grouse and grouse sign. Nineteen habitat and 4 physiographic variables (Table 1) were analyzed for plots at grouse locations. One-way analysis of variance (Nie et al. 1975) was used to test for differences among habitat variable means of the different tree stands. Bartlett's test for homogeneity of variances was performed on each variable. Variables that departed significantly from the assumption of homogeneous variances were analyzed with a 1-way, nonparametric, multiple response permutation procedure (MRPP). Program MRPP computes a test statistic (delta) based on the average between point distances within a priori disjoint subgroups of the given data points in a multi or univariate context (Berry and Mielke 1983). One-way ANOVA was also performed on 3 variables for occupied trees: tree age, dbh, and height. The MRPP program was used if variables departed from the assumption of homogeneous variances. Table 1. Habitat variables measured at blue grouse winter use and random sites in Middle Park, Colorado. Mnemonic Variable description DECO DEOT OT06 C006 C015 C023 C038 CO>39 CODBH CVCODBH CAHT OTDBH CVOTDBH DBH Density of conifers> 7 cm dbh in O.Ol-ha circular plot. Density of non-conifers> 7 cm dbh in plot. Density of non-conifers < 7 cm dbh in plot. Density of conifers < 7 cm dbh in plot. Density of conifers 7-15 cm dbh in plot. Density of conifers 16-23 cm dbh in plot. Density of conifers 24-38 cm dbh in plot. Density of conifers> 39 cm dbh in plot. Mean dbh (cm) of conifers in plot. Coefficient of variation (%) of conifer dbh in plot. Canopy height (m); height of tallest tree in plot. Mean dbh of non-conifers in plot. Coefficient of variation of non-conifer dbh in plot. Mean dbh of 4 nearest trees sampled by the point-centered quarter method. Coefficient of variation of tree dbh sampled by the pointcentered quarter method. Mean distance (m) to nearest 4 trees sampled by the pointcentered quarter method. Coefficient of variation of distances to nearest trees sampled by the point-centered quarter method. Mean height (m) of 4 nearest trees sampled by the pointcentered quarter method. Coefficient of variation of heights of 4 nearest trees sampled by the point-centered quarter method. Slope (degrees) across plot by categories: 0, 1-15, 16-30, 31-45, 46-60. Aspect corrected for declination by categories: N, NE, E, SE, S, SW, W, NW. Vertical position on slope by categories: valley bottom, lower, mid, and upper one-third, ridgetop. Elevation to,nearest 30 m CVDBH DIST CVDIST HTTR CVHTTR SLOPE ASPECT VERTPOS ELEV �43 Tree species importance at grouse observation sites was determined from mean basal area, relative dominance, relative frequency, and relative density (Mueller-Dombois and Ellenberg 1974) of trees measured in O.Ol-ha plots. The importance value (sum of relative dominance, frequency, and density) of Curtis (1959) was divided by 3 to obtain an importance percentage (Risser and Rice 1971). Breeding to wintering site distances were determined from UTM grid coordinates. Distances are minimum straight line distances between the closest UTM grid points at breeding and wintering sites. Breeding sites were defined as male territories, May-June home ranges of non-territorial males and unsuccessful females, and nest locations of successful females. Wintering sites were defined as coniferous habitat occupied during November through March. Distances between wintering and breeding sites for male and female grouse were compared with 1-way MRPP. Home ranges were measured by the minimum area method of Mohr (1947). and juvenile home ranges were compared with the 1-way MRPP program. Adult RESULTS AND DISCUSSION Bandings Forty grouse were banded between 1 April 1982 and 30 March 1983 at the 2 study sites including 34 at Whiteley Peak and 6 at Green Mountain. Thirty-seven (92%) were banded during the breeding and brood rearing period (Apr-Aug). Only 3 (8%) were banded during fall or winter. Twenty females and 14 males were banded at Whiteley Peak. Only 1 male and 5 females were banded at Green Mountain. An additional 31 juvenile grouse were marked with patagial tags. Radiomarking Twenty-four grouse on the study areas were instrumented with radio transmitters; 12 in April-August 1982, 9 in September-December 1982, and 3 in January-March 1983 (Tables 2 and 3). Six grouse died before they moved to wintering sites, 1 slipped the radiocollar, and 17 were followed to wintering areas. In addition, 10 radio-marked grouse surviving from 1981-82 were monitored during 1982-83 of which 4 died before winter, radio contact was lost with 2, and 4 were followed to winter locations. Two transmitters were recovered from grouse that apparently died during fall or winter 198182. �Table 2. Colorado, Band # b Characteristics 1982-83. Sex 245 M 319 355 M M 162 196b F F 266 306c F 317 F 359 F F F 299b 313 F Breeding status 2+ Territorial 06 May 81 1+ 12+ 2+ Non-territorial Chick Successful Unsuccessful 19 03 20 21 2+ 2+ Successful Successful 30 Jun 82 01 Oct 81 6.3 0.5 2+ 2+ Successful Unknown Unsuccessful Successful 11 24 25 07 May Oct Aug May 82 82 81 82 1.4 Successful Chick Chick Chick 26 31 24 04 Jun Aug Oct Dec 82 82 82 82 1+ 1+ F 1+ 357 F F F 111- 358 aMinimum distance (km) between bWintered off GM in 1981-82. cWintered on GM in 1981-82. Date marked blue grouse at Green Mountain Age 337 363 and status of radio-marked May Sep Jun Apr breeding Distance to . . a winter site 82 82 82 81 11 .5 Park, Comments Recaptured 19 May 82, wintered off GM at same location as in 1981-82 Mortality 20 Jun 82, predation Mortality 26 Sep 82, predation Wintered on GM Returned to GM 29 Apr 82, mortality (unknown) Wintered off GM Moved off GM to breed, returned to GM 26 Sep 82, mortality (unknown) Wintered on GM Wintered on GM Remained west of GM, wintered off GM Remained at GM into winter, slipped transmitter Nov 82 Slipped transmitter Aug 82 Mortality 29 Sep 82, predation Remained on GM during winter Mortal ity 19 Dec 82, unknown O. 1 1.2 0.1 and wintering (GM), Middle site. J:" J:" �Table 3. Colorado, Characteristics and status of radio-marked blue grouse Age Breeding status M 2+ Territorial 28 Apr 81 241b M 2+ Territorial 05 May 81 242b M 2+ Territorial 05 May 81 305 M 2+ Te rr ito ria 1 19 Apr 82 1.0 310 2+ 2+ 2+ Territorial Territorial Unknown 27 Apr 82 06 May 82 14 Sep 82 10.6 29.5 000 M M M 366 M F 2+ 2+ Unknown Successful 09 Mar 83 25 Apr 81 0.4 F 2+ Unknown 26 Jun 81 304c F 308c F 2+ 2+ Successful Successful 15 Sep 82 30 Jan 82 0.5 344 F 2+ Successful 01 Sep 82 15. 1 295b F 1+ Unsuccessful 21 Aug 81 302 F 1+ Unknown 26 Aug 81 309 F 316 341 F Unsuccessful Unsuccessful Successful 10 Apr 82 10 May 82 F 1+ 1+ 1+ 13 Jul 82 360 F 365 F 11- 367 F 1- Chick Chick Chick 26 Oct 82 15 Jan 83 16 Mar 8~ Band # Sex 202b c 312 201c 262 c at Whiteley Peak (WP), Middle Park, 1982-83. :Minimum distance (km) between Wintered off WP in 1981-82. CWintered on WP in 1981-82. Date rnarked breeding Distance . . to w ln t e r s It e a Recaptured 28 Apr 82, mortality Sep 82, hunting Recaptured 21 Jun 82, returned to same winter location as 1981-82, mortality 24 Oct 82, predation Returned to WP spring 82, transmitter non-functional Recaptured 19 Apr 82, wintered on WP at same location as 1981-82 Wintered off \-IP Wintered off WP Wintered on WP, mortal ity 30 Dec 82, predation Wintering on WP Remained at WP, lost radio contact during summer Recovered transmitter spring 82, probable fa 11 morta 1 ity Mortality 25 Sep 82, predation Returned to same winter location on WP as 1981-82, mortality 07 Nov 82, predation Moved off WP for winter, mortality fall 82, predation Returned to WP spring 82, lost radio contact Recovered transmitter summer 82, probable fa 11 81, morta 1 ity Wintered on WP Wintered off \vP Remained at WP, mortal ity 26 Sep 82, rredation Wintered on WP Wintering on WP Wintering on WP 6.2 1 .8 5.0 and winteri~' Comments site. _J:I.J1 �46 Seasonal Movements Timing Males radio-marked in spring 1981 returned to their breeding territories at the study areas in late March-early Apri 1 1982 (Table 4). The earl iest arrival date was 30 March and the latest was 7 April 1982. Radio-marked territorial males (N = 6) remained localized on their territories through the breeding season~ Territory abandonment was initiated in late June and continued through July (Table 4). The median departure date for males in 1982 was 18 July (Table 4) compared with a median departure date of 7 July in 1981 (Table 5). Males remained localized on summering areas until early October when they began moving to winter locations. Median arrival dates on wintering sites was 16 October (Table 4) in 1982 and 23 October in 1981 (Table 5). Table 4. Arrival and departure Park, Colorado, 1982-83. Band # Sex 245 202 241 305 310 312 366 000 306 162 196 266 317 359 308 201 344 225 299 313 309 316 358 360 365 367 M M M M M M M M F F F F F F F F F F F F F F F F F F Age 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 1+ 1+ 1+ 1+ 1111- dates of radio-marked Status Location Territorial Terri torial Territorial Territorial Territorial Territorial Territorial Unknown Successful Successful d Successful Successful Unknown Successful Successful Succes sfu I Unknown Unsuccessful Successful Unsuccessful Unsuccessful Chick Chick Chick Chick GM WP WP WP WP WP WP WP GM GM GM GM GM GM WP WP WP WP GM GM WP WP GM WP WP WP aGreen Mountain (GM) or Whiteley bW·Inter morta I·Ity. cFall mortal ity. dS . p r i ng morta llty. Peak (WP) . blue grouse a from seasonal Breeding Arrive site Depart 07 Apr 30 Mar 30 Mar 27 02 26 29 09 24 12 May Jul Jul Jul Jul Jul Jun 25 Sep 05 Aug use sites, Middle Wintering Arrive site Depa rt b ? Oct 11 10 21 ? c OctC Oct Oct Oct 21 Nov c 05 Nov b b 30 Apr c b 29 Apr 10 Oct 09 Oct 05 Nov ? Oct 26 OctC c c 29 Jun 01 23 24 24 05 Aug 30 Oct 07 May 29 Apr 28 Apr 06 May Oct Oct Sep Oct ? Oct ? 03 May 20 May 12 May 13 May 13 05 15 13 May May May May �47 Table 5. Arrival and departure Park, Colorado, 1981-82. dates of radio-marked Band # Sex 245 243 244 202 241 242 306 267 308 196 2D1 298 295 299 M M M M M M F F F F F M F F Age 1+ 1+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 1+ 1+ 111- Status Locat ion Non-territorial Non-territorial Territorial Territorial Territorial Territorial Unknown Successful Unknown Unsuccessful Unsuccessful Chick Chick Chick GM GM GM WP WP WP GM WP WP GM WP WP WP GM aGreen Mountain (GM) or Whiteley b Fal1 mortality. ~inter blue grouse from seasonal a Breeding Arrive use sites, Middle site Deeart Wintering Arrive site Depart 23 22 05 27 10 26 13 Oct b 03 Nov 10 Oct ? Oct 03 Nov 21 Oct 11 Oct 27 Mar Jul Jul Jul Jun Jul Jun 28 Aug 25 May 28 Aug 22 Sep 28 Sep c 10 May c 29 Apr 19 Oct 26 Oct b 21 Oct. 29 Apr Peak (WP) . mortality. Radio-marked females did not move from coniferous areas where they wintered to breeding sites until late April-early May. The earliestarrival date was 28 April and the latest 12 May. Four of 5 radio-marked brood hens at Whiteley Peak remained on the study area into fall and 1 migrated from the area by 10 October (Table 4). Two of 4 radio-marked brood hens at Green Mountain left the study area by 5 August and the other 2 remained on the area until winter. Median arrival dates on wintering sites for females was 23 October (Table 4) in 1982 and 21 October in 1981 (Table 5). Summer Locations Five radio-marked males that wintered off the study areas, moved to higher elevations in spruce-fir habitat for summer (Jul-Sep). Median distance between breeding and summering areas for these 5 males was 11.5 km (range 10.6-29.8 km). A male that wintered and bred at Whiteley Peak moved 2.9 km to a summer location in mixed conifer/aspen adjacent to the study area. Accurate summer and winter locations were determined for 4 males; all occupied winter sites that were spatially distinct from their summer sites. Median distance between summer and winter locations of these 4 males was 1.3 km (range 0.9-4.5 km). Separate summering and wintering sites for radio-marked males was also observed in 1981-82 (Cade 1982). �48 Radio-marked brood hens remained at Whiteley Peak through summer moving in and out of aspens and aspen/shrub edge habitat within 2.0 km of their nest sites. One unsuccessful, yearling female moved between an aspen/ conifer forest and sagebrush (Artemisia spp.) type within 2.0 km of her winter location. Another unsuccessful yearling localized during summer in a willow (Salix spp.)/riparian/hay meadow area 4.0 km from where she attempted to nest. Brood hens that remained on Green Mountain during summer localized in sagebrush on conifer/shrub types within 2.0 km of their nest locations. A female that nested west of Green Mountain had a similar movement pattern. One radio-marked brood hen that left Green Mountain in August moved 1.0 km to a willow/riparian area west of the study area where she remained until returning to Green Mountain for winter. The other brood hen that left Green Mountain moved into spruce-fir habitat during summer. An unsuccessful yearling female spent the summer in shrub and aspen/shrub habitat within 1.2 km of her wintering site. Winter Location Breeding to wintering site movements of 37 blue grouse observed between 1980 and 1983 varied (Table 6). Eighteen radio-marked grouse were tracked in 1982-83, 9 in 1981-82, and 5 in both years. Four banded females and 1 banded male also provided breeding to wintering site information. Breeding to wintering site distances of yearlings and adults for both study areas and all years were combined for analysis (Table 7). Movements of juveniles were treated separately. Movements of juvenile males and females could not be compared because of small sample sizes (Table 8). Adult and yearling males moved greater distances (MRPP, P = 0.009) between breeding and wintering sites than adult and yearling females (Table 7). The shortest distance moved by a male was 1.0 km. Eleven females traveled < 1.0 km between breeding and winterlng sites; 3 females moved only 0.1 km between nest sites and winter locations (Table 6). Table 7. Breeding to wintering site distances blue grouse, Middle Park, Colorado, 1980-83. Descriptive x Median Max r1in N statistic Male 10.9 11. 1 29.5 1.0 10 (km) for adult and yearling Female 3.8 0.7 (.!: = 0.009) 28.0 0.1 19 �49 Table 6. Breeding to wintering site distances for banded and radio-marked blue grouse, Middle Park, Colorado, 1980-83. Band # 184c 245 244 243 202 241 242 305 310 312 366 298 162 266 306 317 359 299 196 313 201 265c 267 342c 308 304c 344 225 322c 309 316 358 360 365 224 367 295 Sex Age M M M M M M M M M M M M F F F F F F F F F F F F F F F F F F F F F F F F F 2+ 2+ 2+ 1+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 12+ 2+ 2+ 2+ 2+ 2+ 1+ 1+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 1+ 1+ 1+ 111111- Status Territor al Territor al Terri tor al Non-terr torial Territor al Territor al Territor al Territorial Territorial Territorial Territorial Chick Successful Successful Successful Successful Successful Unsuccessful Unsuccessful Successful Successful Successful Successful Successful Successful Successful Successful Successful Unsuccessful Unsuccessful Unsuccessful Unsuccessful Successful Unsuccessful Unsuccessful Unsuccessful Unsuccessful Breeding sitea Distance to b winter site(km) SC GM GM GM WP WP WP WP WP WP WP WP GM GM LGM GM GM SC GM GM 2.3 11.5d 11.8 7.8e 14. 1 \oJP 0.4 0.4d 28.0 .0.7 0.5d 1.2 15. 1e 0.3 0.7 d 1.8 5.0 0.8 0.5 0.0 1.3 1.3 14.2g WP WP WP WP WP WP WP WP WP WP GM WP WP WP WP WP 7.6d 17.9 1.0d 10.6 29.5 2.8 11.3e O. 1 6.7 0.5 1.2 0.3 f 0.1 (1.2) 8.7 O. 1 Years observed 1980-81 1981-83 1981-82 1981 1981-82 1981-82 1981-82 1981-83 1982-83 1982-83 1983 1981 1980-83 1981-83 1981-82 1982-83 1982-83 1981-83 1981-82 1982-83 1981-82 1981-83 1981-82 1982-83 1981-82 1981-82 1982-83 1983 1982-83 1982-83 1982-83 1982-83 1982-83 1983 1983 1983 1981-82 aSpring Creek Ranch (SC), Green Mountain (GM), Little Green Mountain (LGM), Whiteley Peak (WP). bMinimum straight line distance between breeding and wintering site. cBanded grouse without a radio transmitter. dWintered in same location in consecutive years. eFa 11 morta 1 ity. fWinter locations differed in consecutive years. 9Minimum distance moved before radio contact was lost. �50 Table 8. Breeding to wintering site distances grouse, Middle Park, Colorado, 1981-83. Descriptive x Median Max Min N statistic (km) for juvenile blue Male Female 11.3 11.3 3. 1 1.3 14.2 0.0 7 Eleven radio-marked grouse, 9 adult and yearling females and 2 adult males, were year-round residents of the study areas, remaining within 2.0 km of the study areas at all times. Five (4 females, 1 male) banded but uninstrumented birds were also documented as year-round residents. Median distance between breeding and wintering sites for the 16 resident grouse was 0.6 km (range 0.1-2.8 km). Five juvenile females may have been residents of the study areas, although they had not been tracked for an entire year as of the end of this segment. Breeding locations for 9 of the 11 radio-marked grouse were in aspen/shrub types that differed from the coniferous habitat they occupied in winter. Two females spent the winter in conifer stands adjacent to the conifer stands they used as nest sites. Another female, #299 (Table 6), captured as a juvenile in 1981 established and became a year-round resident on an area west of Green Mountain in 198283. As an unsuccessful yearling in 1982, this female moved 1.2 km between her spring and winter locations. In 1983 she attempted to nest only 0.1 km from her winter location. Thirteen adult and yearling radio-marked blue grouse, 5 females and 8 males, migrated from the study areas for wlnter. All those surviving the winter (N = 8) returned to the study areas to breed the following spring. Median distance between breeding and wintering sites of these migrant grouse was 11.5 km (range 5.0-29.5 km). A juvenile male and a juvenile female moved a minimum of 11.3 and 14.2 km, respectively, from Whiteley Peak to winter sites. Grouse migrating from Green Mountain moved southsouthwest into the Gore Range; 1 male moved northeast into the Williams Fork Mountains (Fig. 1). Winter locations of radio-marked grouse migrating from Whiteley Peak were east along the Rabbit Ears Range (Fig. 2). �51 N E 5 • MALE o FEMALE Fig. 1. Winter locations of radio-marked blue grouse migrating from Green Mountain, Middle Park, Colorado, 1981-83. (1 = fall mortality) �52 N s • MALE . o FEMALE Fig. 2. Winter locations of radio-marked blue grouse migrating from Whiteley Peak, MIddle Park, Colorado, 1981-83. (1 = fall mortality; 2 = lost radio contact during fall) �53 Zwickel et al. (1968) and King (1971) reported extensive movements (range 1.6-49.6 km) of blue grouse from breeding to fall use areas in Washington and British Columbia, respectively. Bendel 1 and Elliott (1967) documented movements of blue grouse in British Columbia ranging from 1.6 to 16.0 km based on band recoveries. Hoffmann (1956) working in California and King (1971) in British Columbia found blue grouse wintering and breeding on the same areas. Hoffmann (1956) suggested that blue grouse wintering on his study areas were those which bred there. Data obtained in Middle Park indicate that populations of blue grouse contain individuals that migrate long distances between breeding and wintering sites and individuals that winter in coniferous habitat adjacent to their breeding areas. Herzog and Keppie (1980) observed similar variation in movements of spruce grouse (Dendragapus canadensis) in Alberta. Fidelity to Winter and Summer Locations Three radio-marked males (#IS 245, 241, 305), 2 radi~marked females (#IS 308, 309) and 2 banded females used the same wintering sites in 1982-83 that they used in 1981-82. All were adults. One yearling female used a different winter area in 1982-83 than occupied as a juvenile during winter 1981-82. The distance between the 2 winter sites was 0.5 km. Three radio-marked males (#IS 202, 241, 245) also used the same summer locations in 1982 as used in 1981. Breeding site fidelity for territorial male blue grouse has been well documented (Bendell and Elliott 1967, Lewis 1979, among others). Breeding site fidel ity for females is less well established (Jamieson and Zwickel 1983). The limited data from Middle Park suggests that adult blue grouse also have fidelity to winter use sites. Winter Home Ranges Home ranges were calculated for 9 radio-marked blue grouse (1 male, 8 females) located from November 1982 through March 1983. Mean home range size was 7.8 ha and ranged from 1.6 to 42.2 ha. Mean home range size for 4 radio-marked females during winter 1981-82 was 8.7 ha (range 1.3-19.4 ha). Home ranges for 1981-82 and 1982-83 combined of 10 adults were smaller (median = 3.0 ha, range 1.3-11.4 ha) than those of 3 juveniles (median = 18.7 ha, range 9.2-42.2 ha) (MRPP, < 0.01) t Winter Population Characteristics Green Mountain Forty-four observations of 154 grouse on Green Mountain during winter (Nov-Mar) 1982-83 included 14 lone birds (11 females, 3 unknowns) and 30 flocks. Mean flock size was 4.7 birds (range 2-21). Sex composition of 16 flocks was only partially determined; these flocks contained females �54 and unknown birds. Eight flocks contained only females and 6 flocks contained both sexes. When males were observed, females were present. Marked adult and juvenile females were together 6 times. Adult males were the only sex and age-class not identified on Green Mountain; marked birds identified included 5 adult females, 2 juvenile females, and 1 juveni le male. Whiteley Peak Twenty-eight observations of 128 grouse on Whiteley Peak during winter 1982-83 included 9 lone birds (8 males, 1 female) and 19 flocks. Flock size ranged from 2 to 14 birds (x = 6.3). Both sexes were present in 13 flocks, females only in 2 flocks~ and unidentified sexes in 4 flocks. All sex and age-classes were observed on Whiteley Peak; marked birds identified included 3 adult males, 1 juvenile male, 5 adult females, and 3 juvenile females. Sex segregation of blue grouse in winter has been suggested by several authors (Skinner 1927, Marshall 1946, Caswell 1954, King 1971). Observations in Middle Park do not support this contention. Marked adult males were observed with marked adult and juvenile females at Whiteley Peak and occupied the same forest stands and stand types. Hoffmann (1956) considered blue grouse to be highly solitary during winter. Our observations conform more to those of Caswell (1954) who observed the formation of small, loose flocks in winter, and King (1971) who observed lone birds, small groups, and an occasional large flock. Winter Behavior Most blue grouse observed during winter were in conifers. Grouse were observed on the snow or ground at 6 of 61 (10%) grouse observation sites at Whiteley Peak and 3 of 62 (5%) sites at Green Mountain in 1982-83. During winter 1981-82 the corresponding figures were 7 of 43 (16%) sites for Whiteley Peak and 3 of 39 (8%) sites at Green Mountain. Blue grouse were sedentary during most of the day with feeding binges occurring shortly before dark. At this time, birds fluttered about on the branches as they bit off individual needles and clumps of needles. "Umrrh" calls were heard when grouse were feeding. Caswell (1954) reported similar observations during early morning in Idaho. Grouse occasionally were observed feeding at mid-day, although they were not as active as during the evening hour. Grouse often fed in the same trees in which they were roosting. Movements to adjacent trees were common during evening feeding, but no long distance (> 100 m) flights to feeding areas, as reported by Caswell (1954) in Idaho, were observed. �55 Grouse tracks in the snow were commonly observed during winter. Following a sub-zero period in mid-December 1982, old snow roosts were also frequently found. Caswell (1954) observed blue grouse using snow roosts during winter in Idaho. The large number of tracks and snow roosts observed during winter in Middle Park suggests that blue grouse may spend more time on the snow than the recorded observations indicate. Grouse tracks, but no snow roosts, were also observed during winter 1981-82 (Cade 1982). Heights of grouse in trees was measured for 33 birds at Green Mountain and 62 birds at Whiteley Peak during winter 1981-82 and 1982-83. Height was expressed as the percentage of the tree height (Table 9). While grouse were observed in all height classes, 56% of the observations (Table 9) were between 31 and 60% tree height categories. Lateral branch positions for 47 of 93 (51%) grouse observations were within 1 m of the tree trunk, 45% were> 1 m from the trunk but < 1 m from the branch tip, while 4% were on the outer 1 m of the branch. Table 9. Height of blue grouse in relation to total tree height, Middle Park, Colorado, 1981-83. Percent of tree height Lower Statistic 1-10 Upper Middle 11-20 21-30 31-40 41-50 51-60 61-70 71-80 81-90 91-100 N 4 8 11 16 26 11 7 7 2 3 Frequency, % 4 8 12 17 27 12 7 7 2 3 Forest Stand Characteristics Random plots were measured in 16 conifer stands at Green Mountain and 13 at Whiteley Peak. Data are currently being tabulated for analysis. Habitat Characteristics of Wintering Sites Species Composition A total of 204 O.Ol-ha vegetation plots was measured at locations where blue grouse were observed on the study areas during winters 1981-82 and 1982-83. Ninety-four percent of the plots (N = 100) at Green Mountain were in 8 tree stands. Douglas-fir was the dominant species in all stands; importance percentages ranged from 75 to 100% (Table 10). Subdominant species included aspen in 2 stands, Rocky Mountain juniper (Juniperus scopulorum) in 3 stands, and limber pine (Pinus flexilis) in 1 stand (Table 10). �Table 10. 1981-83. Stand # Tree species dominance Stand size(ha) 20.5 at blue grouse winter Mean basal area (m2/O.01 hal locations on Green Mountain, Relative dominance(%) Relative frequency(%) Middle Park, Colorado, Relative density(%) Importance percentage N Tree species 27 Douglas-fir Aspen 0.30 0.00 < 100 1 96 4 99 1 98 2 2.9 12 Douglas-fir 0.32 100 100 100 100 3.7 14 Douglas-fir 0.14 100 100 100 100 IV 40.8 14 Douglas-fir Juniper 0.32 0.02 93 7 70 30 67 33 77 23 V 5.3 7 Douglas-fir Juniper 0.15 0.01 94 6 54 46 77 23 75 25 VI 4.2 3 Douglas-fir Juniper 0.23 0.03 87 13 75 25 69 31 77 23 VII 23.3 6 Douglas-fir Limber pine 0.34 0.01 97 3 85 15 95 5 92 8 VIII 6.8 11 Douglas-fir Aspen 0.27 0.00 100 1 92 8 .99 1 97 3 II III almportance percentage = sum of relative dominance, < frequency, density f 3. \J1 0'> a �Table 11. 1981-83. Stand # II III Tree species dominance at blue grouse winter Mean basal area (m2/O.01 ha) locations on Whiteley Relative dominance(%) Stand size(ha) N Tree species 3.5 36 Douglas-fir Subalpine fir Limber pine Aspen Engelmann spruce 0.33 0.16 0.05 0.01 0.03 58 28 8 1 5 19.3 25 Douglas-fir Aspen Cottonwood Limber pine 0.26 0.02 0.01 0.01 12.8 35 Douglas-fir Subalpine fir Limber pine Aspen 0.25 0.02 0.06 0.00 almportance - percentage = sum of relative dominance, Peak, Middle Park, Colorado, Relative dens ity (%) .. Importance percentage 36 33 14 10 7 32 53 7 6 2 42 38 9 6 5 91 6 2 1 64 31 3 2 58 40 2 < 1 71 26 2 1 75 6 18 < 1 53 23 23 1 52 34 13 1 60 21 18 1 frequency, Relative frequency(%) a density + 3. V1 '" �Table 12. Vegetation characteristics at blue grouse winter locations on Green Mountain, Middle Park, Colorado, 1981-83. Stand numbera II (~=27)c Variableb IV (tJ.=14) x SE III (tJ.=14) x SE (t:1.=12) x SE ~ SE DECO OEOT 5.0 0.0 0.7 0.0 4.1 o 0 o 0 0.9 0.3 OTf/l6 COf/l6 COl5 0.1 1.6 1.6 0.1 0.6 0.4 O. I O. I 0.2 o o C023 C038 CO>39 COOBH 1.2 1.6 0.6 0.3 1.6 0.3 0.2 0.4 0.3 2..I 0.2 1.7 0.6 0.4 0.2 0.2 3.2 0.7 0.7 1.4 1.2 30 0.4 9.4 5. I 2.0 0.6 29 CVCOOBH CAHT OTDBH CVOTDBH DBH 23 CVDBH 51 01ST CVOIST HTTR CVHTTR 38 15 12 o tl.3 0.1 I.Il 5.9 0.7 0.0 0 43 16 o o 0.5 }.2 0.6 o o 7.7 o 14 37 10 o o 0 0.6 3.0 0.4 o o 1.3 (!!_=24) 25 1.9 14 0.9 5.7 (!!_=24) 7.8 0.8 (!!=24) 57 4.2 (!!_=24 ) 10 0.7 (!!_=16) 49 6.4 (!!_=16) 64 7.3 37 4. I 7.5 52 10 52 0.7 6.6 1.0 (!!_=8) 8.2 (!!_=8) 3.4 53 0.3 9.8 0.5 (!!_=1 1) 20 3.0 (!!_=11) 8 . (t:1.=7) SE ~ 4.9 1.4 0.8 0.4 3.7 1.7 0.9 1.7 0 3.7 1.4 0 1.9 0.8 0.5 0.6 5.7 0.7 0.7 3.8 0.7 0.3 D 0 0.9 0.4 0 0.2 0.2 1.9 1.1 o 0.7 1.0 o o 1.7 0.6 V1 VI (tJ.=3) x SE V 50 7 16 18 16 29 0.2 5.3 2.5 1.7 2.7 5.0 2.9 (!!_=9) 50 5.3 (!!_=9) 11.2 1.5 (!!_=9) 63 11.0 (!!_=9) 9 ·1.5 (!!_=2) 59 7.5 (!!_=2) 19 42 1.1 2.7 0.9 0.3 0.3 29 . 3.2 7.3 0.1l 31 12 4.3 2.4 13 o o o 15 0.7 2.6 1.0 24 1.5 42 5.7 45 4.4 12 9 6 4.4 0.5 4. I 0.1 38 7.4 84 8 0.5 o o 42 7. I o o 17.9 VII (1i=6) SE ~ 00 VIII (tJ.;;'l j) pd ~ SE 6.6 0.1 1.4 < 0.1 < o o < 6.5 3.6 1.5 1.0 0.5 2.6 < < < 0.003* 0.225'\ 0.001* 23 1.3 0.6 0.4 0.2 2.2 0.001* 0.001* 0.001 t, 0.001'\ 0.001* 3.5 1.2 o o o o 1.7 0.7 1.3 0.4 0.5 1.7 0.7 42 0.3 0.7 0.2 7.2: 32 16 7.2 2.2 53 12 9.5 0.7 < < O.OOP 0 10 < 0.001* 0 o o o 19 1.8 < 0.011 " 0.001 55 1l.3 o o 27 1.5 (!!_=5) 56 7.7 (!!_=5) 12.5 2.9 (!!_=5) 51 2.9 (!!_=5) 6 0 (!!_=I) 49 0 (!!_=I) 5.7 0.7 < 0.001" 0.001* 0.185 < 0.001 39 3.4 0.101 9 0.7 0.131 30 4.3 0.003 aSee Table 6. bMnemonics of habitat variables described in Table I. cSample size unless otherwise denoted. dSignificance of a I-way ANOVA to test for differences among means of the different stands; * denotes I-way analysis with nonparametrlc MRPP. �59 Ninety-two percent of all plots (N = 104) at Whiteley Peak were in 3 stands. Douglas-fir was the dominant species in all stands. Importance percentages ranged from 42 to 71% (Table 11). Subalpine fir was 2nd in importance in 2 stands while aspen was 2nd an 1 stand. Structural Characteristics Habitat structure was defined with 19 variables (Table 1) describing conifer and non-conifer density, density by size categories of conifers, mean dbh, tree spacing, dispersion characteristics, and heights. At Green Mountain 4 variables, density of conifers 24-38 cm dbh (C038), coefficient of variation of dbh determined by point-quarter sampling (CUDBH), coefficient of variation of distances to trees sampled by point-quarter method (CUDIST), and mean heights of trees sampled by point-quarter sampling, (HTTR) did not differ among the 8 stands (Table 12). Three variables at Whiteley Peak did not differ among plots in the 3 occupied stands. They were conifer dbh (CODBH), coefficient of variation of conifer dbh (CVCODBH), and coefficient of variation of mean tree heights (CUHTTR) (Table 13). Analysis of habitat variables indicated that while some structural similarities exist among grouse sites, marked differences were common. Grouse sites at Green Mountain ranged from low density stands of large conifers to high density stands of large or small conifers. Large diameter conifers were common at all grouse sites at Whiteley Peak but density varied. Physiographic Characteristics Physiographic variables for occupied habitat on Green Mountain and Whiteley Peak varied (Tables 14 and 15). Seventy-seven percent of the plots on Green Mountain and 88% on Whiteley Peak occurred on moderate to steep slopes between 16 and 45°. Occupied sites occurred on all aspects, 88% of the plots at Whiteley Peak occurred on south to northwest aspects while 81% of the plots on Green Mountain were on northwest to northeast aspects. Mean elevation at grouse locations on Green Mountain was 2,640 m and ranged from 2,400 to 2,850 m. Mean elevation of plots at Whiteley Peak was 2,850 m and ranged from 2,610 to 3,030 m. Seventy-four percent of the plots at Green Mountain were on the mid- to upper one-third of the slope of the mountain (Table 14). Ninety-seven percent of the plots at Whiteley Peak were on the upper one-third of the slope or ridgetop (Table 15). Characteristics of Occupied Trees Variables of occupied trees on Green Mountain differed (MRPP, P < 0.001) among the 8 occupied stands. Mean diameters of occupied Douglas-firs ranged from 22 to 52 cm. Mean heights ranged from 10 to 16 m and ages ranged from 63 to 268 years (Table 16). �60 Table 13. Vegetation characteristics Whiteley Peak, Middle Park, Colorado, I {.M.=3b~ Variable b x' - at blue grouse winter 1981-83. Tree stand number II (N=25} SE x - SE locations on a III {.!'i- 35} x - ·SE P DECO 7. 1 0.6 2.8 0.3 3.5 0.4 < 0.001'" DEOT 0.4 0.2 2.0 0.5 0.0 0.0 < 0.001 ,', OT9)6 0 0 5.7 1.1 1.5 1.1 < 0.001 ,', C09)6 3.9 1.1 0.5 0.2 2.0 0.6 < 0.001", C015 1.9 0.3 0.6 0.2 1.3 0.4 = 0.040 C023 1.2 0.2 0.3 0.1 0.2 0.1 < O.OOP C038 2.6 0.3 1.0 0.3 0.8 O. 1 < 0.001", CO>39 1.4 0.1 0.8 0.1 1.1 O. 1 = 0.005'" CODBH 31 1.7 33 1.9 35 2. 1 = 0.341 CVCODBH 49 3.2 43 7.7 40 5.4 0.445 CAHT 23 0.7 16 0.6 14 0.7 < 0.001 OTDBH 13 1.1 11 1.2 7 0 < 0.001 7 1.9 19 3.7 0 0 < 0.001 CVOTDBH DBH 24 1.2 (N=34) 22 2.3 (N=23) 31 2.3 (N=31) = 0.007 CVDBH 3.0 49 (N=34) 63 3.6 (!:!_=23) 46 3.9 (N=31) 0.003 DIST 0.2 4.5 (!:!_=34) 5.8 0.4 (N=23) 0.4 7.0 (N=31) CVDIST 3.4 39 (N=34 ) 3.3 39 (!:!_=23) 4.0 54 (N=31 ) 0.004 HTTR 14 0.9 (!:!_=15) 10 1.0 (N= 11 ) 11 0.8 (!:!_=24 ) = 0.015 CVHTTR 5.0 36 (N=15) 42 6.5 (N= 11) 3. 1 38 (!:!_=24 ) = 0.689 < 0.001 aSee Table 7. bMnemonics cSample of habitat variables size unless otherwise described in Table 1. denoted. dSignificance of a l-way ANOVA to test for differences the different stands; * denotes l-way analysis with MRPP. among means of �61 Table 14. Physiographic characteristics at blue grouse winter locations on Green Mountain, Middle Park, Colorado, 1981-83. 51ope (0) Frequency, % (N=100) Level 1-15 2 21 16-30 31-45 61 16 Aspect Frequency, % (_!!=100) Level N NE 2 21 42 E 5E 5 5W W NW 7 7 18 Vertical position on slopes Ridge Frequency, % (N=100) Upper 1/3 14 Mid 1/3 35 Lowe r 1/3 12 39 Table 15. Physiographic characteristics at blue grouse winter on Whiteley Peak, Middle Park, Colorado, 1981-83. Frequency, % (N=104) Level 1-15 0 9 510 e (0) 16-30 locations 46-60 31-45 54 34 3 Aspect Frequency, % (_!!= 104) Level N NE E 5E 5 5W W NW 0 2 7 2 9 24 13 16 27 Vertical eosition on sloees Rid~e Frequency, % (N=104) 10 Upper 1/3 Mid 1/3 87 2 Lower 1/3 �0' N 16. Table Characteristics of trees occupied by blue grouse during winter, Green Mountain, Tree stand number II (~':.27) b ------- Variable SE ~ Dbh, cm Height, m Age, years 2.4 37 15 182 0.7 19.7 -~ III (N=10) 39 16 212 IV (N=14) V ~ 1.7 0.6 34.0 22 1.2 52 5.11 10 0.4 2.8 15 268 1.9 32.0 63 SE ~ 2.6 25 11 68 0.6 1.4 (!!.=19) (!!.=o) (!!=9) Pyramidal 18 Round 67 15 0 80 20 100 0 0 61 31 86 14 0 100 100 100 100 100 (!!=2) 1981-83. VII VI SE ~ Park, Colorado, a (N=7) (!!.=13) sE' SE Middle (N=3) SE ~ 32 11 1.7 3.2 (!!=7) VII SE ~ 6.3 2.7 0 39 15 199 ALL (N=II) (N=5) SE ~ 5.1 0.8 25.0 37 11 139 (N=89) ~ SE 36 1.6 13 149 0.5 12.1 (!!.=1) (!!.=10) (!!.=56) 40 60 0 27 46 36 48 27 16 100 100 100 Form, % top Deformed Species, 8 33 o 67 % Doug Ias- fir aSee Table bSample 6. size unless otherwise denoted. 100 . �Mean dbh of occupied trees at Whiteley Peak did not differ (MRPP, P = 0.94) among the 3 occupied stands. Mean diameters of occupied trees ranged from 45 to 48 cm. Mean heights and ages of occupied trees at Whiteley Peak differed (MRPP, P < 0.001). Mean heights ranged from 13 to 21 m and ages ranged from 153 to 315 years (Table 17). Table 17. Characteristics of trees occupied by blue grouse during winter, Whiteley Peak, Middle Park, Colorado, 1981-83. Tree stand numbera Variable Dbh, cm Height, m Age, years Form, % Pyramidal Round top Deformed Species, % Douglas-fir Subalpine fir Limber pine I II (N=33) (N=22) SE x SE x c ALL (N=89) III (N=34) SE x x SE 47 17 237 1.7 0.6 20.2 48 3.1 21 0.8 180 13. 1 (N=14) 47 3.0 16 0.6 153 73.8 (N=11) 45 £.6 13 0.8 315 159.2 (N=22) 30 55 15 32 59 9 15 65 20 25 59 16 76 100 15 9 o 82 6 12 84 8 8 o (~=47) aSee Table 7. b Sample size unless otherwise denoted. Douglas-fir was the only tree species used by grouse at Green Mountain (Table 16). Blue grouse at Whiteley Peak were most frequently observed in Douglas-fir (84% of observations), but subalpine fir and limber pine were also used (8% each) (Table 17). Blue grouse were observed in all 3 tree forms. Forty-eight percent of grouse observations at Green Mountain and 59% of those at Whiteley Peak were in overmature, rounded-top trees. �64 Characteristics of Wintering Sites Off the Study Areas Plots at grouse sites off the study areas are too few for detailed analysis. Grouse moving from Whiteley Peak into the Rabbit Ears Range used spruce-fir forests and lodgepole pine mixed with spruce-fir. Large, mature conifers (> 23 cm dbh), as well as, smaller (7-15 cm dbh) conifers were present in these stands. Female #316 moved into a Douglas-fir/spruce-fir type east of Whiteley Peak. This stand was similar to stand I (Table 7) on Whiteley Peak with large Douglas-firs (> 39 cm) mixed with mature spruce-fir. Grouse that moved from Green Mountain into the Gore Range for winter were in mature spruce-fir types or mature lodgepole pine mixed with subalpine fir. Areas were dominated by large (> 23 cm dbh) conifers with smaller (7-15 cm) conifers also present. Female #299 (Table 6) occupied a lodgepole pine stand with large (> 39 cm dbh) Douglas-firs and a scattering of subalpine firs. Lodgepole pines were used while she remained in this stand. However, female #299 moved to a Douglas-fir in late January and apparently remained in a single tree until the end of winter. Habitat characteristics at blue grouse winter sites in Middle Park cover a broader range of conifer types than has been reported in any other single study. Caswell (1954) noted that blue grouse in Idaho wintered in open stands or dense pockets of Douglas-fir. King (1971) observed that most wintering blue grouse in subalpine forests (Tsuga spp., Abies spp., Pinus spp.) in British Columbia used large, mature firs in open parkland and open canopy forests. Stauffer (1983) observed a similar use of open canopy Douglas-fir forests in Idaho, although some winter grouse sign was observed in dense timber (no species given). Stauffer (1983) observed blue grouse using large (> 40 cm dbh) Douglas-firs; Caswell (1954) reported use of 15-30 cm dbh Douglas-firs in Idaho. Winter observations of blue grouse in Middle Park, Colorado occurred in open to dense Douglas-fir dominated forests, as well as in dense sprucefir and lodgepole pine forests. Presence of large (> 23 cm dbh), mature (> 150 years age) Douglas-firs was characteristic of 8 winter sites on the study areas. Two areas of smaller (7-15 cm dbh), younger (60-70 years) Douglas-fir were also heavily used. Lodgepole pine types that were occupied off the study areas were comprised of large (16-38 cm dbh), mature (90-100 years age) trees. Spruce-fir types used as winter sites off the study areas contained mixed size (age-) classes of trees. Species composition varied greatly among winter sites on and off the study areas. Douglas-fir was the dominant species at all wintering sites on the study areas (Tables 6 and 7) and were used most frequently (Tables 14 and 15). Grouse that wintered off the study areas occupied spruce-fir communities where subalpine fir, Engelmann spruce, and lodgepole pine predominated. Harju (1974) observed blue grouse in limber and lodgepole pine during late winter in Wyoming. Boag (1963) indicated that blue grouse in Alberta used lodgepole pine during fall. Data from Middle Park indicate that species composition at blue grouse wintering sites vary even among contiguous areas. �65 LITERATURE CITED Amstrup, S. C. 1980. 44:214-217. A radio-collar for game birds. J. Wildl. Manage. Bendell, J. F. 1955. Age, breeding behavior and migration of sooty grouse, Dendragapus obscurus fuliginosus (Ridgway). Can. J. Zool. 33: 195-223. -----:-- , and P. W. Elliott. in blue grouse. 1967. Behavior and the regulation of numbers Can. Wild1. Servo Rep. Ser. 4. 76pp. Berry, K. J., and P. W. Mielke. 1983. Computation of finite population parameters and approximate probability values for multi-response permutation procedures (MRPP). Commun. Vtat. Simulation Computation B12:83-107. Boag, D. A. 1963. Significance of location, year, sex and age to the autumn diet of blue grouse. J. Wi1d1. Manage. 27:555-562. Brander, R. B. 1968. A radio-package Manage. 32:630-632. harness for game birds. J. Wi1dl. Cade, B. S. 1982. Characteristics and habitat preferences of wintering populations of blue grouse. Job Progress Rep., Colo. Div. Wi1d1. Fed. Aid Proj. W-37-R-35. Pp. 215-241. Caswell, E. B. 1954. A preliminary study on the life history and ecology of the blue grouse in west central Idaho. M.S. Thesis, Univ. Idaho, Mo scow. 105 pp . Cottam, G., and J. T. Curtis. 1956. The use of distance measures phytosociologica1 sampling. Ecology 37:451-460. Curtis, J. T. 1959. The vegetation of Wisconsin. communities. Univ. Wisconsin Press, Madison. An ordination 657pp. in of plant Harju, H. J. 1974. An analysis of some aspects of the ecology of the dusky grouse. Ph.D. Thesis, Univ. Wyoming, Laramie. 142pp. Herzog, P. W., and D. M. Keppie. 1980. Migration of spruce grouse. Condor 82:366-372. in a local population Hoffman, R. W. 1981. Population dynamics and habitat relationships of blue grouse. Final Rep. Colo. Div. Wildl. Fed. Aid Proj. W-37-R-34. Pp. 103-171. Hoffmann, R. S. 1956. Observations on a sooty grouse population Hen Creek, Cal ifornia. Condor 58:321-337. at Sage �66 Jamieson, I. G., and F. C. Zwickel. 1983. Dispersal and site fidelity in blue grouse. Can. J. Zool. 61:570-573. King, D. G. 1971. The ecology and population dynamics of blue grouse in the sub-alpine. M.S. Thesis, Univ. British Columbia, Vancouver. 62pp. Lewis, R~ A. 1979. Suitability and selection of territorial sites used by male blue grouse. M.S. thesis, Univ. Alberta, Edmonton. Marshall, W. H. 1946. Cover preferences, seasonal movements, and food habits of Richardson's grouse and ruffed grouse in southern Idaho. Wi lson Bull. 58:42-52. Mohr, C. o. 1947. Table of equivalent populations small mammals. Am. Midl. Nat. 37:223-249. of North American Mueller-Dumbois, D., and H. Ellenberg. 1974. Aims and methods of vegetation ecology. John Wiley & Sons, New York, N.Y. 547pp. Nie, N. H., C. H. Hull, J. G. Jenkins, K. Steinbrenner, and D. H. Bent. 1975. Statistical package for the social sciences. McGraw-Hill Book Co., New York, N.Y. 675pp. Risser, P. G., and E. L. Rice. 1971. Phytosociological analysis of Oklahoma upland forest species. Ecology 52:940-945. Roth, R. R. 1976. Spatial heterogeneity Ecology 57:773-782. Skinner, M. P. 1927. Bull. 39:208-214. Richardson's and bird species diversity. grouse in Yellowstone Park. Wilson Stauffer, D. F. 1983. Seasonal habitat relationships of ruffed and blue grouse in southeastern Idaho. Ph.D. Diss., Univ. Idaho, Moscow. 108pp. Zwickel, F. C., and J. F. Bendell. J. Wildl. Manage. 31:202-204. 1967. A snare for capturing blue grouse. , I. O. Buss, and J. H. Brigham. 1968. ---grouse and their relevance to populations Manage. 32:456-468. Prepared by Approved by Autumn movements of blue and management. J. Wildl. �67 JQ]3 State of Project PROGRESS REPORT Colorado ----~~~~~---------No. W-37-R-36 -----------------------13 Work Plan No. Population Job Title: Personne 1: 8 Job No. Characteristics Columbian Sharp-tailed Period Covered: Game Bird Survey and Habitat Requirements Grouse in Northwestern 1 January 1982 through 31 December of Colorado 1982 Mike Bauman, Clait Braun, Ken Giesen, Jim Haskins, Jim Hicks, Rick Hoffman, Mike Middleton, Steve Steinert, Chuck Woodward, Colorado Division of Wildlife. ABSTRACT Counts of 26 active sharp-tailed grouse leks in Routt and Moffat counties in 1982 resulted in 182 males, 38 females, and 317 total sharptails being counted. There was no apparent peak of male attendance while female attendance at leks peaked during the last week of April and the 1st week of May. A total of 34 sharp-tailed grouse was trapped and banded on leks (30 males, 4 females) and 6 (5 males, 1 female) were fitted with ponchomounted solar radio t~ansmitters. A sample of 178 sharptail wings was received from the hunter harvest of which 117 (66.1%) were from juveniles. Most wings (N = 90, 50.6%) were from the initial weekend of the hunting season. Major harvest areas were California Park, Hayden Gulch, Twentymile Park, and Steamboat Springs-South. Radiotelemetry data indicated that sharptails summered within 1.0 km of leks but dispersed up to 4.5 km during winter. ��69 POPULATION CHARACTERISTICS AND HABITAT REQUIREMENTS OF COLUMBIAN SHARP-TAILED GROUSE IN NORTHWESTERN COLORADO Kenneth M. Giesen The Columbian or Mountain sharp-tailed grouse (Tympanuchus phasianellus columbianus) has declined in distribution and abundance throughout its historical range, including Colorado (Aldrich 1963, Miller and Graul 1980). Reasons for this decline are not well documented although changes in land use resulting from agriculture, energy development, and human population growth have coincided with population decl ines (Hart et al. 1950, Kessler and Bosch 1981). Although the Columbian sharp-tailed grouse is a game species in Colorado, little information is available upon which to base management decisions. Basel ine data on Columbian sharptail populations, habitats, and harvest are lacking not only in Colorado but throughout its range. Previous studies on distribution of Columbian sharptails in Colorado indicated that the largest populations and apparent best habitats occur in Routt and eastern Moffat counties (Rogers 1969, Giesen and Hoffman 1981) with most of the harvest occurring near California Park and Twentymile Park in central Routt County. Research on the sharptail resource is needed to obtain basic information on breeding densities, nesting success, production and survival of young, fall population numbers, harvest, population turnover, and recruitment to the breeding population. Information on seasonal movements and habitat use is needed to quantify food and cover requirements. Land use changes resulting from human population growth, agriculture, and energy development are expected to further reduce sharptail habitat in the near future while hunting and other recreational uses of sharptails and their habitat are expected to increase. This report covers the 2nd year of the planned 5-year study. P. N. OBJECTIVES The major objectives of this study are to measure sharptail breeding density, production, harvest, survival and turnover rates, and to obtain qualitative and quantitative measures of Columbian sharptail habitat in western Colorado. Segment Objectives: 1. Review pertinent literature applicable to the objectives of this study. 2a. Locate dancing grounds in March, April, and May by systematic search of suitable habitats using binoculars, spotting scopes, and a parabol ic microphone listening device during morning and evening display periods. �70 2b. Count male and female sharptails on known sharptail leks in the 2 study areas twice weekly from mid-March through May during morning display periods. Count male and female sharptails on leks adjacent to study areas at bi-weekly periods during April and May. 2c. Identify individual male and female sharptails on leks using binoculars and spotting scopes to observe color band combinations. 3a. Locate sharptail broods by systematic search of suitable habitats and by using a tape-recorded chick distress call. 3b. Ascertain brood size from counts of broods located in July and August. 4a. Trap adult sharptails using walk-in traps on leks and brood areas and baited walk-in traps in fall and winter, and individually mark with aluminum bands and colored plastic bands. Place solar or batterypowered radios on 2 males and all females captured on the study leks. 4b. Capture sharptail chicks using walk-in and/or drive traps in areas where sharptails are known to occur. 5. Locate all radio-marked sharptails weekly for documentation range size and seasonal habitat use. of home 6. Habitat at sharptail use sites and random sites will be quantified using cover boards, vertical intercept, and point center quarter measurements. 7. Estimate nesting success from wing molt of hens captured in summer and from wings obtained from hunter-harvested birds. 8. Ascertain distribution of hunters and hunter harvest from field hunter checks, check stations, and wing barrels. 9. Obtain number and location of marked birds shot through use of field hunter checks, check stations, and voluntary mail reporting. 10. Food habits will be ascertained from crops of sharptails from hunters and systematic collections. 11. Compile data, analyze results, and prepare progress obtained reports. METHODS Field surveys were conducted on both study areas (Cedar Hill Gulch, Hayden-North) from mid-March through May using binoculars and a parabolic microphone listening device to locate leks within the study areas. Binoculars and spotting scopes were used to obtain counts of male and female sharptails on leks and flush counts were used to obtain complete counts of sharptails on leks. Walk-in drive traps were used to capture sharptails on leks. Intensive searches on foot using a tape-recorded chick distress call or a trained pointing dog were used to locate sharptail �71 broods in July and August. Six solar-powered radios were attached to ponchos (Amstrup 1980) and placed on sharptails to facilitate periodic relocation. Vegetative structure and species composition was measured on leks, sharptail use sites, and random sites using a cover board (Jones 1968), vegetative intercepts (Wiens and Rotenberry 1981), and the pointcentered quarter methods (Cottam and Curtis 1956). Distribution of hunters and hunter harvest was measured using 2 check stations, field checks, and 10 wing barrels. DESCRIPTION OF STUDY AREA Two areas (Cedar Hill Gulch, Hayden-North) were selected for intensive study. Cedar Hill Gulch is in Moffat County (T9N R89W, parts of Sections 31, 32, and 33; T8N R89W, parts of Sections 4-10 and 15-17) and is primarily dominated by native shrub communities and irrigated hay meadows. Hayden-North is in Routt County (T8N R88w, parts of Sections 3-5, and T9N R88W, parts of Sections 19-21 and 28-33). Hayden-North is dominated by native shrub communities, pastures, and wheat fields. A complete vegetative description will be included in the final report. RESULTS AND DISCUSSION Lek Counts From mid-March through late May, 127 counts of 26 active sharp-tailed grouse leks were obtained in Routt and Moffat counties. Twenty-two counts were made on an additional 7 historic leks (California Park Rd. #1, Energy, Fly Gulch, Green Acres, Hicks, Pinnacle) on which no sharptails were seen. Of these inactive leks, only 2 (Energy, Hicks) had more than 2 sharptails in 1981. A minimum of 317 sharptails (12.2/active lek) was counted of which 182 were classified as males. Only 38 sharptails were classified as females, primarily because of their more secretive habits and the difficulty in separating males from females. Because harvest data suggest a 1:1 sex ratio, the counts of females should be used to indicate peak of mating activities and not as an indication of the sex ratio within the population. The number of active leks counted in 1982 increased from previous years while the total number of birds counted and the average count per active lek decreased slightly from 1981 (Table 1). Three dancing grounds were located and counted for the 1st time in 1982 (Barnes, Masiarell i, Scott Place) and several additional grounds originally located by Glenn Rogers in the 1960·s were relocated and counted this year. Although we are counting more active dancing grounds than in previous years, I estimate we are aware of less than 10% of the sharptail leks in Routt and Moffat counties. �72 Table 1. Sharp-tailed 1964-65 and 1977-82. grouse lek counts in Routt and Moffat counties, Year No. active 1eks counted Total no. of sharptails counted Average no. of sharptails/active lek 1964 1965 1977 1978 1979 1980 1981 1982 7 15 2 6 15 5 24 26 38 91 16 54 123 36 335 317 5.4 6.7 8.0 9.0 8.2 7.2 14.0 12.2 Leks in the Cedar Hill Gulch study area had high counts of 10 males and 2 females (Cedar Hill Gulch) and 13 males and 2 females (Pelly1s). Smith1s lek in the Hayden-North study area had a high count of 20 males and 1 female. No apparent trend in timing of peak male counts was observed although female attendance at leks peaked the last week of April and the 1st week of May. Trapping and Banding A total of 34 sharp-tailed grouse (30 males, 4 females) was captured and banded in 1982. All were captured using walk-in drive traps on leks. Between 40 and 60% of the males attending leks on the study areas were captured and banded (Table 2). Five radios were placed on males (1 at Pelley1s lek, 2 each on Cedar Hill Gulch and Smith1s leks) and 1 radio was placed on a female (Pelly1s lek). Only 5 of 30 males trapped on leks were yearl ings (16.7%) suggesting that adult males were easier to trap because of their dominant behavior or that yearlings do not generally recruit to the breeding population. Two of 4 females (50%) trapped were yearl ings. Table 2. Trapping success of male sharp-tailed Hill Gulch, Pe llv 's , and Sm lth s Leks, 1982. grouse on leks at Cedar t High count of males Lek Cedar Hi 11 Gulch Pelly1s Smith1s All 1eks No. males banded Percent males banded 10 13 20 4 7 12 40.0 53.8 60.0 43 23 53.5 �73 Dispersal from Leks Weekly locations of 5 radio-marked sharptails (4 males, 1 female) produced data on movements and habitat use. All radio-marked sharptails establ ished summer home ranges within 1.0 km of the lek they were originally trapped. Four of these sharptails selected irrigated hay meadows or grass pastures for their summer range and 1 male used a mixed-shrub community. Movements of 3 radio-marked males to winter sites ranged from 1.0 to 4.5 km from their lek site. Sharp-tailed Grouse Harvest and Wing Analysis The hunting season for Columbian (mountain) sharp-tailed grouse in 1982 started one-half hour before sunrise on 11 September and closed on 26 September (Units 14, 16, 18, 20, 22, 24, 26, 28, 54, 58, 60, and 66) and 10 October (Units 10 and 12). Since most huntable populations of Columbian sharptails occur in Units 14, 16, and 26, they were exposed to a 16-day season. This was the same season length as in 1981 but shorter than the 1980 season in Units 14 and 16 (25 days). Bag and possession limits were 3 and 6 and separate from the bag limits of sage grouse (Centrocercus urophasianus). This is the 1st season in modern times having separate ba-g-and possession 1imits for these 2 species. The sample of wings received in 1982 (178) was greater than that received in 1981 (142) and more than twice that received in any single year since 1976 when we began collecting grouse wings in Routt and Moffat counties. The increased sample of wings received in the last 2 years was primarily due to the establ ishment of 2 check stations and 10 wing barrels. Wings were received from 2 check stations (California Park = 34, Twentymile Park = 24), 9 wing barrels (100 wings total), and field checks of hunters (20 wings). Of this sample, 1 was from Unit 54 (0.6%), 63 from Unit 14 (35.4%), and 114 from Unit 26 (64.0%). Over half of the wings were collected during the initial weekend of the season in spite of extremely wet weather and poor road conditions. Harvest pressure then stabilized at a lower and nearly constant level for the remainder of the season. Age and Sex Composition of the Harvest Age was ascertained for 177 of 178 wings examined and gonadal inspection of 59 sharptails provided sex ratios of the harvest (Table 3). The percentage of birds in the yearl ing age category is a minimum estimate as most adults and yearl ings (62%) had molted primary 10. Three of 23 (13.0%) birds retaining P10 were yearl ings. Sex was ascertained from 7 adults and 2 yearl ings retaining P10; all but 1 were females. Thus, it is likely that most sharptails retaining P10 during the hunting season are late or renesting hens. �74 Table 3. Age composition of harvested sharp-tailed grouse, northwestern Colorado, 1982. Males Age-class N % N Adults Yearlings Chicks 57 3 117 32.2 1.7 66. 1 9 0 24 a Females % N % 64.3 0 55.8 5 2 19 a 35.7 100.0 44.2 aOnly those for which gonads were examined. While we cannot assume a random sample was obtained from the sharptail population, we can draw some conclusions from our data. 1. Production of young was excellent in 1982 (66.1% chicks) and was the highest measured in 7 years of wing collections. 2. Since the percentage of juveniles in a stable population is a measure of total year-to-year turnover, we can be confident that even in years of poor production (i.e., 1981) more chicks are produced than can be recruited into the following years· breeding population. 3. The sex ratio in the adult and young-of-the-year age-classes is sl ightly skewed towards males although the 1976-82 average indicates an even sex ratio. Other studies of this species have also indicated an even sex ratio in all age-classes. Data on age and sex composition of the hunter harvest. from all years for which data are available are presented for comparison in Table 4. Table 4. Age and sex composition of harvested sharp-tailed grouse, northwestern Colorado, 1976-82. Adults Year N 1976 1977 1978 1979 1980 1981 1982 10 47 13 32 25 83 60 Totals 7-year average a b Adults Females Males % N % Total sample 71.4 65.3 46.4 40.5 39.7 58.4 33.9 4 25 15 47 38 59 117 28.6 34.7 53.6 59.5 60.3 41.6 66. 1 14 72 28 79 63 142 177 5 5 1 3 14 9 574 32 271 Young 303 47.2 alncludes yearl ings. b Known sex only. 52.8 4 Young b Males Females 6 4 18 7 7 3 13 24 10 3 15 19 34 53 51 �75 LITERATURE CITED Aldrich, J. w. 1963. Geographic orientation J. Wi1d1. Manage. 24:529-545. Amstrup, S. C. 1980. 44:214-217. of American Tetraonidae. A radio-collar for game birds. J. Wildl. Manage. Cottam, G., and J. T. Curtis. 1956. The use of distance measures phytosocio10gica1 sampling. Ecology 37:451-460. in Giesen, K. M., and D. M. Hoffman. 1981. Distribution and status of mountain sharp-tailed grouse. Colo. Div. Wi1dl., Final Rep ., Fed. Aid Proj. W-37-R-34. Apr. 1981. Pp. 183-189. Hart, C. M., O. S. Lee, and J. B. Low. 1950. The sharp-tailed grouse in Utah. Its life history, status, and management. Utah Dep. Fish and Game, Fed. Aid Div. Pub1. 3. 79pp. Jones, R. E. 1968. A board to measure cover used by prairie grouse. J. Wi 1d1. Manage. 32: 28-31. Kessler, W. B., and R. P. Bosch. 1981. Sharp-tailed grouse and range management practices. Pages 133-146 in Proc. Wi1d1./Livestock Symp., Univ. Idaho, Moscow. Mi 11er, G. C., and W. D. Graul. 1980. Status of sharp-ta iled grouse in North America. Pages 18-28 in P. A. Vohs, Jr., and F. L. Knopf, eds. Proc. Prairie Grouse Symp., Okla. State Univ., Stillwater. Rogers, G. E. 1969. The sharp-tailed grouse in Colorado. Game, Fish, and Parks, Tech. Pub1. 23. 94pp. Prepared by ~ ~ --~K~e~n~n~e~t~h~M~.~G~i~e~s~e=n~----------Wildlife Researcher B Colo. Div. ��77 JOB PROGRESS REPORT State of Colorado --------------------------- Project No. W- Work Plan No. Job Title: Game Bird Survey Job No ,: 17 Population Period Covered: Personnel: 37- R- 36 Dynamics of White-tailed 7 Ptarmigan 1 January - 31 December 1982 Clait E. Braun and Kenneth M. Giesen, Colorado Division of Wildlife. ABSTRACT Long-term studies of populations of white-tailed ptarmigan (Lagopus leucurus) were continued at hunted (Mt. Evans) and unhunted (Rocky Mountain National Park) areas in Colorado in 1982. Densities of breeding ptarmigan continued to decrease from hli qhs experienced in 1979 (Mt. Evans) and 1976 (Rocky Mountain National Park). Nesting success was poor (25%) at Mt. Evans in 1982 and good (67%) at Rocky Mountain National Park. Average brood size to 1 September at the 2 areas was 3.0 and 3.2 chicks! successful hen, respectively. Hunters at Mt. Evans in fall 1982 removed at least 16% of the estimated fall population of ptarmigan. ��79 POPULATION DYNAMICS OF WHITE-TAILED PTARMIGAN Clait E. Braun and Kenneth M. Giesen Long-term studies of trends in population size and investigation of reasons for fluctuations in size of tetraonid populations are lacking. Studies on the population dynamics of unhunted and hunted populations of white-tailed· ptarmigan were initiated in Colorado in 1966 and have continued essentially uninterrupted at 2 sites. Studies of the unhunted population (Rocky Mountain National Park) have identified possible short-term cycles of 7-8 years with an amplitude of 25-30% between high and low breeding densities. Conversely, studies of the manipulated population (hunted) at Mt. Evans through 1980·have not indicated any cyclic pattern and it would appear that controlled hunting may mask any long-term trend that may occur. This report covers the 2nd year of a 5-year study designed to examIne the questIon whether white-tailed ptarmigan are truly cyclic and whether hunting affects the apparent oscillations. . P. N. OBJECTIVES The goals of this investigation are to be able to predict the length and amplitude of cycles in white-tailed ptarmigan in Colorado, to examine the impact of hunting on cycles, and to clarify underlying causes of the apparent cycles. SEGMENT OBJECTIVES 1. Conduct breeding (May-Jun) and brood (Aug-Sep) censuses of white-tailed ptarmigan using tape-recorded calls of males (breeding) and chicks (broods) . i 2. Censuses will be conducted on previously established, defined study areas at Mt. Evans (hunted) and at Rocky Mountain National Park (unhunted). 3. Capture (noose poles) and band (aluminum and plastic color-coded bands) all unmarked white-tailed ptarmigan encountered on study areas at Mt. Evans and at Rocky Mountain National Park. 4. Individually identify all ptarmigan observed on study areas at Mt. Evans and Rocky Mountain National Park through use of binoculars. 5. Make hunting season and bag limit recommendations for Mt. Evans and collect hunting data through use of volunteer wing barrels and hunter field checks. 6. Compile data, analyze results and prepare progress report. �80 STUDY AREA AND METHODS Areas investigated were at Mt. Goliath-Mt. Evans in Clear Creek County and at Tombstone Ridge-Sundance Mountain to Fall River Pass in Rocky Mountain National Park in Larimer County. The physiography, geology, location, and vegetation on these study areas has been previously described (Braun 1969, 1971; Braun and Rogers 1971; Giesen 1977). Ptarmigan were located through use of tape-recorded calls (Braun et al. 1973), captured through use of telescoping noose poles (Zwickel and Bendell 1967) as described by Braun and Rogers (1971), and classified as to age and sex and banded following Braun and Rogers (1971). Age of chicks was estimated following Giesen and Braun (1979). Numbered plastic bandettes were not used as in earlier years (Braun and Rogers 1971) as a color-code system using up to 4 different colored plastic bandettes was instituted in 1977-78. A check station was operated on the Mt. Evans highway during the opening weekend (25-26 Sep) of the ptarmigan season in that area. A volunteer wing collection station was available to hunters in the area from 26 September through 10 October when the season closed. RESULTS AND DISCUSSION Breeding Densities Mt. Evans Surveys of breeding white-tailed ptarmigan on the Mt. Evans study area in Spring 1~8~ (May-Jun) revealed the presence of at least 10 pairs and 6 unmated males. This is a density of 6.5 birds/km2, a decrease from 1981 (Table 1). Pairing and breeding activities were initiated in late May, later than in 1981 because of the extensive snow cover in 1982. Rocky Mountain National Park Surveys of ptarmigan present on the Rocky Mountain National Park study units during May and June 1982 indicated that 15 pairs and 13 single males were present. This is a density of 7.8 birds/km2, lower than the 8.2 birds/km2 recorded in 1981 (Table 1). Nesting Success and Brood Size Mt. Evans During the July-early September interval, only 2 different hens were identified with broods (average size to 1 Sep = 3.0 chicks/successful hen) while 6 hens were observed without broods. Thus, only 2 of 8 hens (25.0%) were known to be successful in nesting. Estimated hatching dates for 7 chicks for which data are available ranged from 5 July to 14 August with most (5 of 7) hatching between 5 and 12 July, later than the 30 June6 July peak calculated in 1981. �81 Table 1. White-tailed ptarmigan breeding densities, Colorado 1966-82. Study Area Year Rocky Mountain National Park (5.5 km2) Mt. Evans (4.0 km2) 1966 1967 1968 1969 1970 1971 1972 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 11.3 9.8 11.5 12.0 9.6 9.1 8.7 7.8 8.0 11. 1 13.5 12.9 10.7 8.7 8.4 8.2 7.8 3.0 2.7 2.7 2.2 2.0 4.2 7.5 6.2 6.2 6.2 6.7 >6.0 7.5 10.3 9.5 9.0 6.5 Rocky Mountain National Park Fourteen successful hens (14/21 = 66.7%) were observed between 1 August and early September with an average brood size to 1 September of 3.2 chicks. Only 7 hens were observed without broods in 1982 (7/21 = 33.3%). The peak of hatch occurred during the 3rd week of July, about 1 week later than the long-term average. Harvest Mt. Evans In 1982, the ptarmigan season in the Mt. Evans area (Unit 52 south of Interstate 70 and east of the Guanella Pass Road between Georgetown and Grant) was delayed 2 weeks after opening of the statewide season on 11 September. This delay was similar to that in the 1977-80 period following the experimentation with season length and limited closure from 1972 through 1976. This experimentation followed the documentation of overharvest in this area in the late 1960's and the 2-year closure in 1970 and 1971. In 1982, the season opened one-half hour before sunrise on 25 September and closed at sunset on 10 October. �82 A wing collection barrel placed along the Mt. Evans highway prior to opening of the season was available until after 10 October. Because of the mild fall, the road remained open until 1 October when it was closed by the state highway department because of an agreement with the CDOW. A check station was operated both days of the opening weekend. During the 2 days of check station operation, 16 hunters with 5 ptarmigan were checked. This compares to 20 hunters with 18 ptarmigan and 12 hunters with 9 ptarmigan checked in the same period in 1981 and 1980, respectively. No additional wings were deposited in the wing barrel. Two banded birds were reported by mail. In all, at least 8 ptarmigan were harvested in 1982 of which all were banded. The fall population resident in the area was estimated to be about 50 birds. The total fall population (residents and nonresidents) in the area hunted was estimated at 50-60 birds. Thus the harvest of 8 birds represented about 16% of the fall population. LITERATURE CITED Braun, C. E. 1969. Population dynamics, habitat, and movements of whitetailed ptarmigan in Colorado. Ph.D. Thesis, Colo. State Univ., Fort Collins. 189pp. 1971. Habitat requirements of Colorado white-tailed Proc. West. Assoc. State Game and Fish Comm. 51:284-292. , and G. --~Colo. Div. E. Rogers. 1971. The white-tailed ptarmigan Game, Fish and Parks Tech. Publ. 27. 80pp. ---.,- , R. K. Schmidt, .Jr., and G. E. Rogers. white-tailed 37:90-93. 1973. ptarmigan with tape recorded calls. ptarmigan. in Colorado. Census of Colorado Manage. J. Wildl. Giesen, K. M. 1977. Mortality and dispersal of juvenile white-tailed ptarmigan. M.S. Thesis, Colo. State Univ., Fort Collins. 55pp. ----:-- , and C. E. Braun. juvenile white-tailed 1979. A technique for age determination of ptarmigan. J. Wildl. Manage. 43:508-511. Zwickel, F. C., and J. F. Bendell. 1967. A snare for capturing grouse. J. Wildl. Manage. 31:202-204. Prepared by Clait E. Braun Wildlife Research Leader Kenneth M. Giesen Wildlife Researcher blue �83 JOB PROGRESS REPORT State of Colorado -------------------------- Project No. Game Bird Survey W-37-R-36 22 Work Plan: Job Title: Job No.: Upland Game Publications Period Covered: Personnel: 1 April 1982 through 30 June 1983 C. E. Braun, K. M. Giesen, R. M. Hoffman, T. J. Schoenberg, and W. D. Snyder, Colorado Division of Wildlife ABSTRACT Publications accomplished under this job in Segment 36 are: Braun, C. E., and K. M. Giesen. 1983. Winter home range size of whitetailed ptarmigan. Wilson Ornithol. Soc. Annu. Meet. 64:16 (Abstract). Giesen, K. M., and C. E. Braun. 1983. Reproductive performance of female white-tailed ptarmigan in Colorado. Cooper Ornithol. Soc. Annu. Meeting. Abstract. , T. J. Schoenberg, and ------sage grouse in Colorado. Hoffman, R. W. 32-35. 1983. C. E. Braun. 1982. Methods for trapping Wildl. Soc. Bull. 10:224-231. The wings the thing. II. Colo. Outdoors 32(2): Snyder, W. D. 1982. Minimum tillage techniques for establishing shrubs in clump plantings. Colo. Div. Wildl. Spec. Rep. 53. 17pp. 1983. Colorado's ring-necks. Colo. Country Life 30(7) :4, 9-10. 1983. Developing a PATREC model for pheasant habitat evaluation in the High Plains. Perdix III Workshop. Abstract. 1983. 18-20. Every farm needs a plum thicket. 1983. Pheasant nesting ecology Perdix III Workshop. Abstract. Colo. Outdoors in relation to wheat farming. 1983. Shrub thicket establishment in Colorado's Colo. Div. Wi ldl. Game Inf. Leafl. 108. 4pp. Prepared by Clait E. Braun Wildlife Research Leader 32(2): High Plains. � https://cpw.cvlcollections.org/files/original/2816c4ae4abdebf359da230c2a8a5f20.pdf 5c39decd3a1eb9486af0c4f51a312a76 PDF Text Text Colorado Division of Wildl ife Wildlife Research Report July 1983 JOB PROGRESS REPORT State of Colorado --~~~~--------------- Project No. 45-01-502-15050 Work Plan No. Author: Personnel: - Cervids Mu l.t ispec ies Inves t igat ions Job No. 5 Period Covered: Big Game Investigations 7/1/82-6/30/83 Experimental Improvement of Oakbrush on Deer, Elk and Cattle Ranges Hightower Mountain R. C. Kufeld R. C. Kufeld ABSTRACT A manuscript entitled "Responses of elk, mule deer. cattle, and vegetation to burning, spraying and anchor chaining of Gambel Oak Rangeland" was completed and sent through the review process and will be published as a Division of Wildlife technical publication. ��3 EXPERIMENTAL IMPROVEMENT OF OAKBRUSH ON DEER, ELK, AND CATTLE RANGES - HIGHTOWER MOUNTAIN Roland C· Kufeld P. N. OBJECTIVE To determine the extent to which the production and quality of deer, elk, and cattle forage and livestock and wildlife use can be increased and maintained by chaining, spraying, and burning overage Gambel oak winter ranges. SEGMENT OBJECTIVES Complete analysis of data and write a manuscript describing effects of burning, spraying,and chaining of Gambel oak on vegetation production and responses of deer, elk, and cattle to treated areas 2, 5 and 10 years after treatment. METHODS AND MATERIALS Methods are described by Kufeld (1971, 1972). RESULTS AND!DISCUSSION A manuscript entitled "Responses of elk, mule deer, cattle and vegetation to burning, spraying and anchor chaininq of Gambel oak ranaeland" was completed and sent through the review proc~ss. Upon completi;n of recommended revisions it will be published in the Colorado Division of Wildlife Technical Publication Series. LITERATURE CITED Kufeld, R. C. 1971. Experimental improvement of oakbrush on deer, elk and cattle ranges - Hightower Mountain. Colo. Div. of Game, Fish and Parks, Game Res. Rep. July (1):23-86. Kufeld, R. C. 1972. Experimental improvement of oakbrush on deer, elk and cattle ranges - Hightower Mountain. Colo. Div. of Wildl. Game Res. Re p . J u 1y ( 2) :8 1- I05 . Prepared by ~c~ Roland C. Kufeld Wildlife Researcher C 1 ( I ��Colorado Division of Wildlife Wildlife Research Report July 1983 5 JOB PROGRESS REPORT State of Colorado Project No. 45-01-502-15050 Work Plan No. Job No. Author: Personnel: Muftispecies 7 Period Covered: Big Game Investigations - Cervids Investigations Big Game Research Publications 7/1/82-6/30/83 L. H. Carpenter R. B. Gill, L. Lovett, N. McEwen and All Big Game Researchers and Graduate Students. ABSTRACT During the 1982-83 Segment, the Big Game Research Section had 15 manuscripts published, 13 others accepted for publication and 7 manuscripts in the review process. ��7 BIG GAME RESEARCH PUBLICATIONS L. H. Carpenter P. N. OBJECTIVE To publish the results of research conducted under the auspices of Federal Aid Projects 45-01-502-15050 and 45-01-503-15050 in a v~riety of professional journals and other indexed publishing media to insure widespread dissemination and availability of this information to natural resource managers and ecological scientists. SEGMENT OBJECTIVES 1. Job Progress Reports. All studies. 2. Anderson, A. E. 1983. A synthesis of the literature Div. of Wildl. Spec. Rep. 3. Baker, D. L., and N. T. Hobbs. summer diets in Colorado. 4. Bartmann, R. M. 1983. Mule deer winter diet composition pinyon-juniper range in Colorado. J. Range Manage. 5. Bartmann, R. M. 1983. Preliminary evaluation of the helicopter quadrat census for mule deer on pinyon-juniper winter range. Colo. Div. of Wildl. Spec. Rep. 6. Bartmann, R. M., and D. C. Bowden. 1983. Mule deer winter mortality pinyon-juniper range in Northwest Colorado. J. Wi Idl. Manage. 7. Beane, R. D., and N. T. Hobbs. 1982. The Baerman technique for determining Protostrongylus infection in bighorn sheep. Effect of laboratory procedures. J. Wildl. Diseases. 8. Bear, G. D. 1983. Mark-recapture technique for deriving population estimates as applied to an elk population. J. Wildl. Manage. 9. Bear, G. D. 1983. Population dynamics of the Estes Valley elk population. Colo. Div. Wildl. Spec. Rep. 1982. Composition J. Wildl. Manage. on puma. Colo. and quality of elk and quality on 10. Bear, G. D., and L. H. Carpenter. 1983. Elk production and calf survival as related to range carrying capacity. J. Ecology or J. Wildl. Manage. 11. Bear, G. D., and R. L. Green. 1983. Seasonal distribution Northcentral Colorado as related to vegetation types. Manage. 12. Beck, T. D. I. 1982. Colorado black bear harvest data, 1979-81. Colo. Div. Wildl. Spec. Rep. of elk in J. Wildl. on �8 13. Beck, T. D. I. Colorado. 1982. A specimen of Ursus arctos horribi 1 is from J. Mamm. 14. Beck, T. D. I., and M. Haroldson. 1982. Black bear weight estimation from chest girth measurements. Colo. Div. Wildl. Outdoor Facts. 15. Beck, T. D. I., R. A. Nelson, and D. S. Well ik. 1982. Natural history of seasonal changes in behavior and biochemistry in wild black bears. Science. 16. Bowden, D. C., A. E. Anderson, and D. E. Medin. 1982. Sampl ing plans for sex and age ratios of mule deer. J. Wildl. Manage. 17. Bowden, D. C., and N. T. Hobbs. indices. J. Wildl. Manage. 18. Carpenter, L. H. 1983. Twenty-four hour activity patterns of mule deer at pasture. J. Wildl. Manage. 19. Carpenter, L. H., R. B. Gi II, and D. L. Baker. 1983. Tests of a nutritionally based habitat evaluation system. Colo. Div. Wildl. Spec. Rep. 20. Dailey, T. V., and N. T. Hobbs. 1983. Comparative of mountain goats and bighorn sheep. 21. Freddy, D. J. Colorado. 22. Freddy, D. J. 1982. Sampling mule deer pellet grup densities juniper-pinyon woodland. J. Wildl. Manage. 23. Freddy, D. J. 1982. Efficacy of permanent and temporary in juniper-pinyon woodland. J. Wi ld 1. Manage. 24. Freddy, D. J. 1983. Reactions of mule deer to human harrassment. J. Wi 1d 1. Manage. 25. Freddy, D. J. 1983. Heart rates for activities J. Compo Biochem. and Physiol. 26. Freddy, D. J., R. M. Bartmann, L. H. Carpenter, and R. B. Gill. 1983. Measuring mule deer sex and age ratios in juniper-pinyon woodland. J. ~ildl. Manage. 27. Gill, R. B., L. H. Carpenter, R. M. Bartmann, and D. L. Baker. 1983. Estimating mule deer diets from bitecount and fecal analysis. J. Range Manage. ~ 28. Green, R. L., and G. D. Bear. 1983. Daily activity patterns of elk in Northcentral Colorado as related to vegetation types. J. Wildl. Manage. 29. Hobbs, N. T. 1983. The nutrient density hypothesis: Fire effects on a relative shortage of food for herbivores. Oecologia. 1982. Confidence intervals on overlap 1982. Predicting mule deer harvest J. Wildl. Manage. nutritional ecology in Middle Park, in pellet plots of mule deer at pasture. �9 30. Hobbs, N. T. 1983. mineralization. Fire effects on soil nitrogen Oecologia. 31. Hobbs, N. T., and R. A. Spowart. 1983. Fire effects on nutritional quality of ungulate diets during winter. J. Wildl. Manage. 32. Kufeld, R. C. 1983. Deer, elk, cattle and vegetation responses to burning, spraying and chaining,of Gambel Oak Rangeland. Colo. Div. Wildl. Spec. Rep. 33. Lance, W. 1982. Diseases of pronghorn Division of Wildlife publication. 34. Pojar, T. M., and L. L. Wolfe. 1982. antelope. J. Wildl. Manage. 35. Pojar, T. M., D. C. Bowden, R. B. Gill, and M. P. Elkins. 1982. Quadrat and strip sampling to estimate pronghorn population characteristics. J. Wildl. Manage. 36. Spowart, R. A., and N. T. Hobbs. 1983. Fire effects on diet selection by mule deer and bighorn sheep. J. Wildl. Manage. 37. Torbit, S., L. H. Carpenter, D. M. Swift. and A. W. Alldredge. Dynamics of mule deer body composition. J. Wildl. Manage. 38. Torbit, S., L. H. Carpenter, A. W. Alldredge, and D. M. Swift. 1983. Mule deer body composition - A comparison of methods. J. Wildl. Manage. 39. White, G. C., and R. M. Bartmann. 1983. Banding analysis of a White River, Colorado mule deer herd. J. Wildl. Manage. antelope, Recurrent fixation and a literature estrus review. in pronghorn 1983. ACKNOWLEDGMENTS All scientists and graduate students on the Big Game Research Staff are to be commended for their continued excellence in publishing results of their work. PUBLICATION 1. Job Progress Reports. in: Colorado Div. of Wi ldl. All studies. 1983. PROGRESS These reports have been printed Wi ldl. Res. Rep. July Part 1. �10 : i 2. Anderson, A. E. 1983. A synthesis of the literature on puma. Div. of Wildl. Spec. Rep. This manuscript was published Anderson, A. E. 1983. (Felis concolor). 3. as: A critical review of the literature on puma Co 10 ..D iv. of Wi 1d 1. Spec. Rep. 54: ·.91pp. Baker, D. L., and N. T. Hobbs. summer diets in Colorado. This manuscript Co~o. was published 1982. Composition J. Wildl. Manage. and quality of elk as: Baker, D. L., and N. T. Hobbs. 1982. Composition and quality of elk summer diets in Colorado. J. Wildl. Manage. 46(3) :694-703. 4. Bartmann, R. M. 1983. Mule deer winter diet composition on pinyon-juniper range in Colorado. J. Range Manage. This manuscript was accepted for publication and quality by the J. Range Manage. as: Bartmann, R. M. 1983. Composition and quality of mule deer diets on pinyon-juniper winter range, Colorado. J. Range Manage. 36(4): 534-541. 5. Bartmann, R. M .. 1983. Preliminary evaluation of the helicopter quadrat census for mule deer on pinyon-juniper winter range. Colo. Div. of Wildl. Spec. Rep. This manuscript 6. is still in the review process. Bartmann, R. M., and D. C. Bowden. 1983~ Mule deer winter mortality on pinyon-juniper range in Northwest Colorado .. J. Wildl. Manage. This manuscript was accepted for publication by the J. Wildl. Manage as: Ba rtmann, R. M. 1983(?). Estimating mule deer winter mortality Colorado. J. Wildl. Manage (in press). 7. Beane, R. D., and N. T. Hobbs. 1982. The Baerman technique for determining Protostrongylus infection in bighorn sheep. Effect of laboratory procedures. J. Wildl. Diseases. This manuscript 8. in was published in J. Wildl. Diseases. 19(1) :7-9. Bear, G. D. 1983. Mark-recapture technique of deriving population estimates as applied to an elk population. J. Wildl. Manage. A draft of this manuscript to the review process. has been prepared and is ready for submission �11 9. Bear, G. D. 1983. Population dynamics of the Estes Valley elk population. Colo. Div. Wildl. Spec. Rep. Data analyses, tables and graphs have been completed Work on the first draft will begin soon. 10. for this manuscript. Bear, G. D., and L. H. Carpenter. 1983. Elk production and calf survival as related to range carrying capacity. J. Ecology or J. Wildl. Manage. No progress was made on this manuscript. 11. Bear, G. D., and R. L. Green. 1983. Seasonal distribution Northcentral Colorado as related to vegetation types. Manage. Data for this manuscript are being incorporated population dynamics by Bear. of elk in J. Wildl. into the manuscript 12. Beck, T. D. I. 1982. Colorado black bear harvest data, 1979-81. Colo. Div. Wi Idl. Spec. Rep. 13. Beck, T. D. I. Colorado. 1982. on A specimen of Ursus arctos horri b i l i s from J. Mamm. No progress was made on these manuscripts. 14. Beck, T. D. I., and M. Haroldson. 1982. Black bear weight estimation from chest girth measurements. Colo. Div. Wildl. Outdoor Facts. This manuscript has been accepted by Southwest Naturalist as: Beck, T. D. I., and M. Haroldson. 1983. Black bear weight estimation from chest girth measurements. S.W. Nat. (in press). 15. Beck, T. D. I., R. A. Nelson, and D. S. Well ik. 1982. Natural history of season changes in behavior and biochemistry in wild black bears. Science. This manuscript is in the review process. made in authors: The following change was R. A. Nelson, T. D. I. Beck, and D. S. Wellik. 16. 198{?). Bowden, D. C., A. E. Anderson, and D. E. Medin. 1982. Sampl ing plans for sex and age ratios of mule deer. J. Wildl. Manage. This manuscript was accepted by J. Wildl. Manage as: Bowden, D. C., A. E. Anderson, and D. E. Medin. 198{?). Sampl ing plans for sex and age ratios of mule deer. J. Wild1. Manage. (in press). 17. Bowden, D. C., and N. T. Hobbs. 1982. Confidence overlap indices. J. Wildl. Manage. This manuscript is still in preparation. intervals on �12 18. Carpenter, L. H. 1983. Twenty-four hour activity patterns of mule deer at pasture. J. Wildl. Manage. 19. Carpenter, L. H., R. B. Gi 11, and D. L. Baker. 1983. Tests of a nutritionally based habitat evaluation system. Colo. Div. Wild1. Spec. Rep. no progress was made on these manuscripts. 20. ·Dailey, T. V., and N. T. Hobbs. 1983. Comparative ecology of mountain goats and bighorn sheep. This manuscript nutritional was accepted by J. Wild1. Manage. as: Dailey, T. V., N. T. Hobbs, and T. W. Woodard. 198(?). Experimental comparisons of diet selection by mountain sheep and mountain goats. J. Wildl. Manage. (in press). 21. Freddy, D. J. Colorado. This manuscript 22. 1982. Predicting mule deer harvest in Middle Park, J. Wildl. Manage. was published as: Freddy, D. J. Colorado. 1982. Predicting mule deer harvest J. Wildl. Manage. 46(3):801-806. in Middle Park, Freddy, D. J. juniper-pinyon 1982. Sampling mule' deer pellet group densities woodland. J. Wildl. Manage. in This manuscript was published as: Freddy, D. J. 1983. Sampling mule deer pellet group densities in juniper-pinyon woodland. J. Wild1. Manage. 47(2) :476-485. 23. Freddy, D. J. 1982. Efficacy of permanent and temporary pellet plots .I.njuniper-pinyon woodland. J. Wild1. Manage. This manuscript.was published as: Freddy, D. J. 1983. Efficacy of permanent and temporary pellet plots in juniper-pinyon woodland. J. Wi ld l, ~1anage. 47(2) :512-516. 24. Freddy, D. J. 1983. Reactions of mule deer to human harrassment. J. Wildl. Manage. This manuscript 25. Freddy, D. J. pasture. This manuscript is s t l11 in draft stages. 1983. Heart rates for activities of mule deer at J. Compo Biochem. and Physio1. has been submitted to J. Wildl. Manage. �13 ,i 26. Freddy, D. J., R. M. Bartmann, L. H. Carpenter, and R. B. Gill. 1983~ Measuring mule deer sex and age ratios in juniper-pinyon woodland. J. Wild1. Manage. No progress was made on this manuscript. 27. Gill, R. B., L. H. Carpenter, R. M. Bartmann, and D. L. Baker. 1983. Estimating mule deer diets fr~m qitecount and fecal analysis. J. Range Manage. This manuscript as: has been accepted for publication Gill, R. G., L. H. Carpenter, P. M. Bartmann, G. G. Schoonveld. 1983. Fecal analysis J. Wi ldl. Manage. 47(4): (in press). 28. in J. Wi1dl. Manage. D. L. Baker, and to estimate deer diets. Green, R. L., and G. D. Bear. 1983. Daily activity patterns of elk in Northcentral Colorado as related to vegetation types. J. Wi ldl. Manage. No progress was made on this manuscript. 29. Hobbs, N. T. 1983. The nutrient density hypothesis: Fire effects on a relative shortage of food for herbivores. Oeco10gia. First draft of this paper has been prepared review. 30. Hobbs, N. T. 1983. Fire effects on soil nitrogen mineral ization. Oecologia. This manuscript 31. to submLt to internal has been submitted fixation and to J. Range Manage. Hobbs, N. T., and R. A. Spowart. 1983. Fire effects on nutritional quality of ungulate diets during winter. J. Wildl. Manage. This manuscript has been accepted by J. Wi1dl. Manage. as: Hobbs, N. T., and R. A. Spowart. 1983. Fire effects on nutritional quality of ungulate diets during winter. J. Wildl. Manage. (in press). 32. Kufeld, R. C. 1983. Deer, elk, cattle and vegetation responses to burning, spraying and chaining of Gambel Oak Rangeland. Colo. Div. Wildl. Spec. Rep. This manuscript is being published as: Kufeld, R. C. 1983. Effects of burning, spraying, and anchor chaining ~n elk, mule deer and cattle use, and vegetative yields of Gambel oak rangeland. Colo. Div. Wildl. Tech. Pub. 34 (in press). �14 33. Lance, W. 1982. Diseases of pronghorn antelope, a literature Division of Wi ld l. publ ication. review. This manuscript with the addition of T. Pojar as a junior author, has been drafted and is ready for internal -review. 34. Pojar, T. M., and L. L. Wolfe. 1982. antelope. J. Wildl. Manage. This manuscript Recurrent estrus in pronghorn has been accepted by the J. Wildl. Manage. as: Pojar, T. ~1., and L. L. Wolfe. 198{?). Recurrent estrus in pronghorn antelope. J. Wi ldl. Manage. (in press). 35. Pojar, T. M., D. C. Bowden, R. B. Gill, and M. P. Elkins. 1982. Quadrat and strip sampling to estimate pronghorn population characteristics. J. Wildl. Manage. No progress made on this manuscript. 36. Spowart, R. A., and N. T. Hobbs. 1983. Fire effects on diet selection by mule deer and bighorn sheep. J. Wildl. Manage. The first draft of this manuscript is being prepared. 37. Torbit, S., L. H. Carpenter, D. M. Swift, and A. W. Alldredge. Dynamics of mule deer body composition. J. Wild1., Manage. 1983. 38. Torbit, S., L. H. Carpenter, A. W. Alldredge, and D. M. Swift. 1983. Mule deer body composition - A comparison of methods. J. Wildl. Manage. Both of these manuscripts have been submitted to J. Wildl. Manage. and are in the review process. 39. White, G. C., and R. M. Bartmann. 1983. Banding analysis of a White River, Colorado mule deer herd. J. Wildl. Manage. This manuscript was published in J. Wildl. Manage as: White, G. C., and R. M. Bartmann. 1983. Banding analysis of a White River, Colorado mule deer herd. J. Wildl. Manage. 47:506-511. Additional publications which were not scheduled but which were published or accepted for publication during the 1982-83 segment are listed below: Anderson, A. E., and O. C. Wallmo. 198{?). Mule and black-tailed Mammalian Species No. (?). 8pp. (in press). deer. Bartmann, R. M. 1983. Winter foods of mule deer in Piceance Basin. Div. Wi ldl. Game Inform. Leafl. 107. 4pp. Colo. �15 Gill, R. B., L. H. Carpenter, and D. C. Bowden. 1983. Monitoring large animal populations: The Colorado experience. Trans. N. Amer. Wildl. Conf. 48: (in press). Freddy, D. J. 1983. A "But t e!' of an investment. Colo. Outdoors (in press). Garrott, R. A., and R. M. Bartmann. 198(?). Evaluation of vaginal implants for mule deer. J. Wildl. Manage. (in press). Hobbs, N. T., and R. Spowart. 1982. Karrow, K. K., and D. J. Freddy. Outdoors (in press). Kufeld, R. C. 1983. (in press). Pojar, T. Reed, D. F. 40-42. 1983. 1982. Fire. 1983. Colo. Outdoors 31 (4) :26-29. Mountain whitetails. Preferred deer habitat: Speedster of the plains. What is it? Colo. Colo. Outdoors Colo. Outdoors 32(2):40-44. When sheep and goats compete. Colo. Outdoors 31(6): Reed, D. F., T. D. I. Beck, and T. N. Woodard. 1982. Methods of reducing deer-vehicle accidents: benefit--cost analysis .. Wild. Soc. Bull. 10(4):349-354. Swift, D. M. 1983. A simulation model of energy and nitrogen balance for free-ranging ruminants. J. Wildl. Manage. 47(3): (in press). Prepared by )~&J ~a:& Len H. Carpenter Wildlife Research Leader ��Colorado Division of Wildlife Wildlife Research Report July 1983 17 JOB PROGRESS REPORT State of Colorado Project No. 45-01-502-15050 Work Plan No. 1 ------------------- Job No. Multispecies 8 Period Covered: Author. Big Game Investigations - Cervids Investigations Big Game Publication Library Services Editing and 7/1/82-6/30/83 M. Hershcopf and L. Carpenter Personnel: M. Hershcopf, L. Carpenter, R. B. Gill, N. McEwen, C. Doty, A. Morekill, J. Asbed, J. Dudley ABSTRACT During the 1982-83 Segment, 26 books were purchased for permanent reference by DOW personne1. Forty-five additional publications were located, ordered, and obtained free of charge for use. Sixteen theses were purchased, obtained on interlibrary loan or given to the library and 4 computer-aided literature searches were completed for the library files. An additional 975 individual references requested by Big Game Researchers were located by 'library staff and made available for reference. About 35 of these requests were not available locally and were obtained through interlibrary loan. ��19 BIG GAME PUBLICATION EDITING AND LIBRARY SERVICES Marian W. Hershcopf and Len H. Carpenter P. N ..OBJECTIVE To provide a centralized support pro~ram for big game research technical editing and library services so that Big Game Research Scientists can allocate additional time to the conduct of actual research. SEGMENT OBJECTIVES To provide coordinated, efficient, and economic editing and library services to all Colorado Big Game Research programs (Federal Aid Projects 45-01-50215050 and 45-01-503-15050). SUMMARY OF SERVICES Publ ications Purchased with 45-01-502-15050 Funds and Placed in Research Center Library Amlaner, C. J., Jr., and D. W. MacDonald, eds. 1980. A handbook on biotelemetry and radio tracking. Permagon Press, Oxford, England. 804pp. I Armstrong, D. M. 1975. Rocky Mountain mammals. Inc. and USDI, Natl. Park Servo 174pp. Campbell, G. S. 1977. An introduction Springer-Verlag, Inc., NY. 159pp. Rocky Mtn. Nature Assoc., to environmental Chapman, J. A., and G. A. Feldhamer, eds. 1982. America--biology, management and economics. Baltimore. 1147pp. biophysics. Wild mammals of North John Hopkins Univ. Press, Clutton-Brock, T. H., et al. 1982. Red deer; behavior and ecology of two sexes. Univ. of Chicago Press, Chicago. 378pp. Conservation Foundation. 1982. The state of the environment. Foundation, Washington, DC. 439pp. Dasmann, W. 1981. Deer range: Improvement McFarland, Jefferson, NC. l68pp. and management. Conservation 2nd ed. Ffolliott, P. F., and S. Gallina. 1981. Deer biology, habitat requirements and management in western North America. Instituto de Ecologra, A. C. Mexico, DF. A binational Mexico-US Man and Biosphere (MAB) Prog. Invest. Instituto de Ecologfa Publ. No.9. 238pp. �20 Food and Agriculture Organization of the U.N. and the International Atomic Energy Agency. 1979. Laboratory training manual on the use of nuclear techniques in animal research. Internatl. Atomic Energy Agency, Vienna. 299pp. Fowler, C. W., et a l. 1980. Comparative population dynamics of large mammals: A search for management c rlt er la , US Marine Mammal Comm., Washington, DC. 330pp. Hainsworth, F. R. 1981. Animal physiology; Addison-Wesley Publ. Co., Reading, MA. adaptations 669pp. Hall, E. R. 1981. The mammals of North America. Sons, NY. 2 vols., 1260pp. Hartl, D. L .. 1981. A primer of population Sunderland, MA. 191pp. Leopold, A. S., et al. Charles Scribner's 2nd ed. genetics. Houston, D. B. 1982. The northern Yellowstone MacMillan Publ. Co., Inc., NY. 474pp. in function. elk: John Wiley and Sinauer Assoc., Inc., Ecology and management. 1981. North American game birds and mammals. Sons, NY. 198pp. Lyman, C. P., et al. 1982. Hibernation Academic Press, NY. 317pp. and torpor in mammals and birds. Margaris, N., et al., eds. 1982. Aromatic plants: Basic and applied aspects. Martinus Nijhoff Publ., Boston. 283pp. Meijs, J. A. C. 1981. Herbage intake by grazing dairy cows. Centre for Agricultural Publ. and Documentation, Pudoc, Wagenigen, The Netherlands. 264pp. Murie, A. 1981. The grizzlies Washington, DC. 251pp. of Mt. McKinley. USDI, Natl. Park Serv., Masacchia, K. J., and L. Jansky. eds. 1981. International symposium for survival in the cold--Survival in the cold: Hibernation and other adaptations. Elsevier, NY. 225pp. Nielson, L. 1982. Chemical immobilization in urban animal control work. The Wisc. Humane Soc., Inc., Milwaukee. 93pp. Nielson, L., et al. 1982. Chemical immobilization of North American wildlife. The Wisc. Humane Soc., Inc., Milwaukee. 447pp. Peek, J. M., and P. D. Dalke, eds. 1982. Wildlife-livestock relationships symposium: Proc. 10. Univ. of Idaho, For., Wildl. and Range Exp. Sta., Moscow. 614pp. Proc. of the 2nd Internatl. Symp. on Protein Metabolism and Nutrition. 1977. Pudoc, Centre for Agric. Publ. and Doc., Wagenigen, The Netherlands. 178pp. �21 Thomas, J. W., and D. E. Toweill. 1982. Elk of North America, ecology and management. Stackpole Books, Harrisburg, PA. 698pp. Wallmo, O. C., ed. 1981. Mule and blacktailed Univ. of Nebraska Press, Lincoln. 605pp. deer of North America. Publications Obtained Free or at Low Cost In addition to books purchased with Federal Aid Funds, about 45 free reports and short publications from state or federal agencies or from private sources, were located, ordered and obtained for use by Big Game Research personnel. Theses Purchased, Obtained on Interlibrary Loan or as Gifts for Use by Researchers Ackerman, B. B. 1982. Cougar predation and ecological energetics southern Utah. MS Thesis, Utah State Univ., Logan. 95pp. in Adams, L. G. 1981. Ecology and population dynamics of mountain goats, Sheep Mountain - Gladstone Ridge, Colorado. MS Thesis, Colo. State Univ., Ft. ColI ins. 189pp. Anderson, D. A. 1981. Response of the Columbian black-tailed deer to fertilization of Douglas fir forests with municipal sewage sludge. PhD Thesis, Univ. of Washington, Seattle. 186pp. Cart, T. W. 1971. The struggle for wildlife protection in the United States, 1870-1900: Attitudes and events leading to the Lacey Act. PhD Thesis, Univ. of North Carolina, Chapel Hill. 218pp. Currier, M. J. P. 1979. An age estimation technique and some normal blood values for mountain lions (Felis concolor). PhD Thesis, Colo. State Univ., Ft. Co llins. 81pp. Eicher, T. A. 1978. Some life history characteristics and habitat use of mule deer in the Sacramento Mountains. MS Thesis, New Mexico State Univ., Las Cruces. 73pp. Gibbs, H. D. 1978. Nutritional quality of mule deer foods, Piceance Basin, Colorado. MS Thesis, Colo. State Univ., Ft. Collins. 179pp. Green, R. A. 1982. Elk habitat selection and activity patterns in Rocky Mountain National Park, Colorado. MS Thesis, Colo. State Univ., Ft. Co llins. 165pp. Hemker, T. P. 1982. Population characteristics and movement patterns of cougars in southern Utah. MS Thesis, Utah State Univ., Logan. 59pp. Hier, R. H. 1981. Mule deer and rabbit use of natural and cabled pinyonjuniper woodland at Fort Stanton, New Mexico. MS Thesis, New Mexico State Univ., Las Cruces. 43pp. �22 McColl oug h, S. A. 1982. Mountain, Colorado. Impact of cattle grazing on bighorn sheep, Trickle MS Thesis, Colo. Stat~ Univ., Ft. Collins. 119pp. Morrison, B. G. 1961. Some aspects of the histology and growth of the horns of Antilocapra americana. MS Thesis, Univ. of Wyo., Laramie. 48pp. Owen, R. 1981. Classification of mule deer habitat using biophysical MS Thesis, Univ. of Nevada, Reno. 121pp. variable. Oyer, E. R. 1939. A study of the structure of hair as a means of mammal identification. MS Thesis, Ft. Hays State College, Hays, Kansas. 39pp. Shackelton, D. M. 1973. Population quality and bighorn sheep (Ovis canadensis canadensis Shaw). PhD Thesis, Univ. of Calgary,~erta. 227pp. Simmons, B. W. 1982. Summer-fall ecology and behavior of bighorn sheep, Waterton Canyon, Colorado. MS Thesis, Colo. State Univ., Ft. Collins. 211 pp. Computer Literature Searches Obtained for Big Game Researchers Hunting wounding loss, crippling loss, deer, elk, pronghorn. Effects of external telemetry packages on behavior. Pellet group census technique. Trapping deer in summer. Reference Document Location and De 1ivery The Research Center Library staff also located and delivered about 975 individual articles on request for the Big Game Research Section during this segment; about 35 were not available locally and were obtained through interlibrary loan procedures. Prepared by M 0--vI' Q:IA-- tJ. ikx-s-" Cot Marian W. Hershcopf Librarian Len H. Carpenter Wildlife Research Leader F �Colorado Division of Wildlife Wildlife Research Report July 1983 JOB PROGRESS REPORT State of Colorado Project No. 45-01-502-15050 Multispecies Work Plan No. 9 Job No. Period Covered: Author: Big Game Investigations - tervids Investigations Cervid Research Administration 7/1/82-6/30/83 L. Carpenter Personnel: L. Carpenter, R. B. Gill, L. Lovett, and N. McEwen ABSTRACT Documents that were prepared for Five-Year Program Plans, 3 new Study Plans, a new employee evaluation system and the research audit highlighted Cervid Research administration during the 1982-83 segment. ��25 CERVID RESEARCH ADMINISTRATION Len H. Carpenter P. N. OBJECTIVE To supervise and administer 15050. research on ~¢er, and elk in Project 45-01-502- SEGMENT OBJECTIVE Supervise and administer 45-01-502-15050. all deer and elk research studies in Project METHODS AND MATERIALS The Wildlife Research Leader position exists to provide detailed planning, budgeting, and coordination for the individual research jobs within a research project or projects. During the 1982-83 segment, considerable effort was expended by the Wildlife Research Leader in long-range planning (Program Plans) and project administration as dictated by the research audit. RESULTS AND DISCUSSION A Five-Year Program Plan detailing the thrust of future Cervid Research in Colorado was prepared by project personnel and drafted into DOW Program Plan format. These plans were circulated to all sections within the Division for comment. The Program Plan identifies products, deadlines and relevance to the DOW Strategic Plan of each scheduled activity. Three new Study problems were selected and new, detailed Program Narratives (Study Plans) were drafted and approved by the contract biometrician and the work implemented during the 1982-83 segment. Detailed budget plans were prepared and executed and all stated objectives were achieved. Project budgets in 1982-83 were reduced from previous years. Project scientists were innovative in their approaches to becoming more efficient with less money. Despite reductions in funding and frustrations with the research audit, project scientists published at a record rate (see Job Progress Report for Work Plan 1,Job 7). Fifteen manuscripts authored or co-authored by project personnel, were either publ ished or accepted for publication by a variety of publishing media. The research audit consumed a large portion of the Wildlife Research Leader's time after 1 January 1983. It is estimated that 75% of my time since January 1 was involved in audit activities. Much of this time was spent in drafting responses to various requests by the audit committee chairman. An attempt was made by all research personnel to be as thorough and open as possible with regard to the audit. Unfortunately, much of the material that was obtained, summarized and submitted to the audit committee was not �26 used in the final committee report. Research personnel have maintained a complete record of these transactions on file at the Research Center. As the segment ended, a second "mini audit" was being conducted by the Director. This audit also required considerable time and effort to re-summarize the material we prepared for the first audit. To minimize impacts on the research process, the Wildlife Research Leader prepared most of the audit materials. During the 1982-83 segment, the Colorado Division of Wildlife changed systems for employee evaluation. Evaluations were changed from the Performance Planning and Review System (PPR) to the Factor Anchored Performance Appraisal System (FAPAS). FAPAS was to be implemented on January 1983 for all Division employees for a 6-month trial. Due to numerous delays in printing forms, etc., the Research Section did not receive pertinent instructions for FAPAS until late March. At this time a 2~ month test of a 6 month FAPAS had to be prepared. All supervisors in the Wildlife Research Section prepare detailed Annual Work Plans for their employees. These work plans specify products and time frameworks for each segment. Considerable effort was expended to convert each employee's Annual Work Plan from the PPR system to FAPAS. At the beginning of the new segment this conversion was nearly complete and all research employees in Cervid Research have an Annual Work Plan tied to the new system. Considerable time was spent by the Wildlife Research Leader coordinating development of a computer model to simulate impacts of oil shale development in Northwest Colorado on deer and elk for the Northwest Colorado Wildlife Consortium. The model is based on the nutritional contributions of the forage and the corresponding nutritional requirements of the big game animals. Conceptual knowledge and specific data for this model :resulted from past and current DOW big game: research. Development of the model was primarily done by coordinating all this work. By the end of the segment, initial runs of the model had been made. u- ~ Co.r~tv Prepare d b y~ __~~~ ~~~ Len H. Carpenter Wildlife Research Leader _ �Colorado Division of Wildlife Wildlife Research Report July 1983 27 JOB FINAL REPORT State of Colorado Project No. 45-01-502-15050 Work Plan No. Multispecies 10 Job No. Period Covered: Author: Personnel: Big Game Investigations Nutritional Ruminants - Cervids Investigations Value of Big Sagebrush to 7/1/82-6/30/83 L. Carpenter L. Carpenter, P. Neil ABSTRACT A literature review was conducted and a study plan prepared but availability of research animals was insufficient to properly conduct the experiment during the 1982-83 segment. ��29 NUTRITIONAL VALUE OF BIG SAGEBRUSH TO RUMINANTS Len H. Carpenter P. N. OBJECTIVE In vitro trials Hypothesis: Sagebrush inhibits activity of cellulolytic bacteria .. In vivo trials Hypothesis: Energy assimilation by mule deer is inversely related to the proportion of sagebrush in a sagebrush-grass diet. The proportion ME/DE is inversely related to the proportion of sagebrush in the diet. SEGMENT OBJECTIVE Review pertinent literature, develop appropriate hypotheses, prepare a study plan, and begin testing effects of different levels of big sagebrush on digestive physiology of mule deer, both in vivo and in vitro. METHODS AND MATERIALS I The literature was reviewed and a study plan was prepared that described in vitro and in vivo experiments necessary to test the stated hypotheses. Other priority research required use of the experimental animals that were planned for this work. As a result, it was decided that this job could not be conducted this segment. The study plan will be filed away at this time and when money and resources become available, the work will be continued. P repa red by -:--~~---:::--~_-:-Cc_c;,.."r-,--4-~ Len H. Carpenter Wildl ife Research Leader _ ��Colorado Division of Wildlife Wildlife Research Report July 1983 31 JOB PROGRESS REPORT State of Colorado Project No. Big Game Investigations - Cervids 45-01-502-15050 Work Plan No. Deer IQvestigations 2 ------------------- Nutritional Basis for Quantifying Capacity of Winter Ranges to Support Deer--Evaluation of thermal cover used by Deer Job. No. Period Covered: Author: 7/1/82-6/30/83 D. J. Freddy Personnel: D. J. Freddy, K. K. Karrow ABSTRACT Use of simulated thermal cover by tame mule deer was monitored during 4 trials at monthly intervals between December, 1982 and March, 1983. Eight deer were individually confined to large isolation pens where they could selectively use 4 treatments: 1) a tree, 2) windbreak, 3) conical mound of dark soil, or 4) a control area providing no thermal cover. Use of treatments was analyzed only when deer were bedded. Comparisons among 4 treatments indicated that differential use of tr~atments (p < 0.08) occurred in February and March, and for all trials pooled. Soil was the most frequently used treatment. Comparisons indicated deer used cover treatments collectively over controls during each trial and for all trials pooled (~< 006). Measurements of thermal characteristics of the soil, tree, and windbreak suggested deer could obtain thermal advantages from all these treatments at low wind velocities. Adjustment by deer to confinement and their positive use of simulated cover offers potential to further experimentally assess effects of cover on deer energetics. ( J ,1 ��33 NUTRITIONAL BASIS FOR QUANTIFYING CAPACITY OF WINTER RANGES TO SUPPORT DEER David J. Freddy P. N. OBJECTIVE To determine if a system can be developed to estimate number of deer winter ranges are capable of supporting. SEGMENT OBJECTIVES 1. Complete study plan (Program Narrative) uti lized by mule deer during winter. for evaluating thermal cover 2. Measure behavioral responses of tame mule deer to simulated cover within enclosures during winter. ACKNOWLEDGMENTS The U. S. Forest Service, Rocky Mountain Forest and Range Experiment Station, kindly loaned instrumentation for measuring weather conditions. Karl K. Karrow provided di 1igent assistance in collecting and summarizing data. METHODS AND MATERIALS See Program Narrative (Appendix A). Exceptions to the Program Narrative are as follows: 1. Eight deer (4 M, 4 F) were used as planned in trials 1 and 2. Unfortunately, 1 female (182) became ill (eventually died) after trial 2 so an additional male (190) was substituted for trials 3 and 4. Most data summaries herejn are based on 7 deer common to 4 trials. 2. Dry matter digestibility (in vitro) of pelleted ration was 74% (Pers. comm. D. Baker) which was considerably higher than anticipated. During trials, deer were thus fed at approximately 81% of maintenance instead of the desired 60% maintenance level. 3. Numbers of observations of deer lying on each of 4 treatments were transformed according to the proportion of usable area (m2) within each pen that each treatment comprised. To transform data, total observations of deer lying on each treatment were divided by the following values: 1) tree, 0.033, 2) windbrea~ 0.076, 3) soil, 0.033, and 4) control, 0.858. For comparisons between lion any cover treatment" and lion contro1" data were transformed using 0.142 for on cover and 0.858 for on control. Transformation accounted for differing probabilities of deer lying on each treatment. �34 RESULTS AND DISCUSSION Use of simulated thermal cover by deer was monitored for 6 days during each of 4 trials conducted at monthly intervals between December, 1982, and March, 1983 (Table 1). Weather conditions were generally mild with minimum temperatures of only -19 C. Wind was of significant velocity and duration only in December and coldest temperatures occurred in January (Table 2). Deer were probably not metabo1 ica11y stressed during any trial assuming a lower critical temperature of -23 C (Mautz et a1., in press). I Deer adjusted well to confinement in isolation pens (Fig 1). Time spent among 4 activities (Table 3) was consistent among trials. Percent time lying (6570%) exceeded values (57%) reported by Carpenter (1976) for free-ranging tame deer subsisting on native forage during winter within an adjacent pasture. Nervous behaviors, such as pacing, were evident only during the hour preceeding the evening feeding. All deer used all cover treatments during the study (Figs 2a, b, c). I IAll deer lost weight during the experimental period (Fig 3) even though deer were, for the most part, fed a high qual ity pel1eted ration ad libitum and ! also had access to native forage. From 10 December 1982 to 20 March 1983 (101 days), average weight loss for males was 9.4% (73 g/day) and for females 11.3% (62 g/day) (n = 7). Deer continued to lose weight when fed only the pelleted ration during trials. On average, weights decreased 1% for males and 2% for females each trial. Only 2 deer (203 M, 191 F) did not consistently consume all the pelleted ration offered (Table 4). Treatment Use Conclusions regarding teatment preference by deer varied with analysis used. An analysis of variance test among the 4 treatments (tree, windbreak, soil, control) indicated that differential use of treatments occurred (p < 0.08) during February and t1arch and in all trials pooled (Table 5). For all trials pooled, deer use of each cover treatment exceeded (p = 0.05, Tukey's Q) the cont ro.l treatment in the following decreasing order: soi 1, tree, windbreak. In February, treatments in decreasing order of use were soil, tree, control-wind and in March, soil, tree, wind-control. However, this analysis using 7 blocks (deer) and 4 treatments/block produced large error mean squares which suggested a deer x treatment interaction. An analysis of variance test between 2 treatments (on any cover or on control) indicated significant (p < 0.06) use of cover treatments during each trial, and all trials poo1ed-(Table 5). Neither AN OVA indicated significant (p > 0.40) main effects associated with deer (blocks). Frequency of lying activity for all trials was not affected by individual isolation pens (p > 0.50). However, deer exhibited a preference (p < 0.02) for lying in quadrants 2 and 4 (see diagram of pen in Program Narrative). Although there was little elevational change within pens, quadrants 2 and 4 contained the elevationally higher one-half of each pen and quadrant 4 contained the entrance gate. Fortunately, treatments occurred equally in all quadrants to minimize systematic effects of quadrant use. Control quadrants ranked third or fourth in frequency of lying activity in 7 of 8 pens. Infrequent use of the control quadrant suggests: 1) deer may be reluctant to bed in open areas and prefer to be near some cover, or 2) effects of cover treatments on thermal conditions extend beyond defined limits and measurements. �35 Use of cover treatments by the deer tended to increase from days 1-3 to days 4-6 of each trail as evidenced by increasing ratios of cover use:control use (Fig 4). However, a regression between cover:contro1 ratios and individual days was significant (r = 0.85, P < 0.05) only in January when temperatures were coldest. This tendency ·of increasing use may reflect a cumulative effect of a restricted diet, deer learning to use cover as they become famil iar with an isolation pen, o~ (least likely) unfavorable ambient conditions during latter days of each trial. Cover Qualities Wind profiles on leeward sides of windbreaks were measured to demonstrate effects of the windbreak on wind velocity. At a height of 30 cm (approx. height of a deer1s spine when lying down), wind velocities at 0 m and 1 m from windbreak averaged 36% and 61%, respectively, of the velocity measured 2 m from windbreak ( Fig 5). When wind velocity at height of 2 m increased to 7.1 m/sec (Fig 5, top), velocity at height of 30 cm and 1 m from wind~ break sharply increased. Therefore, as wind velocity increases deer must move closer to the windbreak to receive maximum benefit. The definition that deer are on the treatment when they are lying at or within 1 m of the windbreak appears reasonable as effects on wind velocity are most pronounced within this distance. Thermal advantages provided by the windbreak and artificial tree were monitored during daylight hours (Fig 6). Black globe temperatures at a height of 50 cm averaged 1.7 C warmer at 1/2 m than at 1 1/2 m from the windbreak (p < 0.01, paired t-test, n = 11). Globe temperatures averaged 0.5 C warmer at 1/2 m than at 1 1/2 m from base of tree (p < 0.02, paired t-test, n = 12). Although this difference may be real, the-0.5 C resolution approached accuracy levels of 2 thermometers used. Radiant thermal advantages near the windbreak and tree decreased as wind velocity increased above 3 m/sec. Both the windbreak and tree provided thermal conditions warmer than ambient temperatures (Fig 6). . Surface temperatures of the soil treatment when dry were 3 C warmer than surface temperatures of adjacent snow (p < 0.01, paired t-test, n = 20) (measured with Mikron infrared thermometer). Measurements represent differences during daylight hours as the infrared thermometer would not function during colder night temperatures. The soil treatment thus provided a thermal advantage over lying on snow. This thermal advantage likely decreases as wind velocity increases. Cover Use and Ambient Conditions Conclusions regarding relationships between cover use and ambient conditions await considerable data analysis. Compounding this problem is the influence of individual deer behavior on treatments used. Thermal characteristics of cover treatments suggested deer could obtain advantages from either the soil, windbreak, or tree when wind velocities were low. Differences among cover treatments may therefore be negligible and thus preferences of individual deer may guide cover use. It may be better to collectively treat the 3 cover treatments as 1 and assess relationships between cover use and ambient conditions. However, when wind velocities reach a threshold value that negate conductive or radiant thermal advantages of cover treatments, deer �36 may more predictably seek or abstain from using a particular treatment. This response is suggested by use of treatments in relation to wind velocity (Table 6). Use of the windbreak first exceeded other treatments at wind velocities of 5-6 m/sec. CONCLUSIONS Deer adjusted well to isolation pens and used c~ver collectively more than controls. This experimental approach therefore offers the opportunity to test effects of presence and absence of cover on deer energetics. A major problem with further experimentation involves developing a pelleted ration lower in digestible energy to more reasonably approximate a natural diet. Further experimentation may compare differences in activity patterns and loss of body weight between deer having access to cover structures and deer having no natural or artificial cover. LITERATURE CITED Carpenter, L. H. evaluation. 1976. Middle Park cooperative deer study--deer habitat Colo. Div. Wildl. Game. Res. Rep. July, Part 2:284-406. Mautz, W. W., P. J. Pekins, and J. A. Warren. Cold temperature effects on metabol ic rate of white-tailed, mule, and black-tailed deer in winter coat. Proc. Int. Conf. on Biology of Deer Production. Dunedin, New Zealand. Feb., 1983. (in press). Ullrey, D. E., W. G. Youatt, H. E. Johnson, L! D. Fay, B. L. Schoepke, and W. T. Magee. 1969. Digestible energy requirements for winter maintenance of Michigan white-tailed does. J. Wildl. Manage. 33:482-490. Prepared by (~.;1 Y:f Davi J. e dy I Wildlife Researcher C �37 Table 1. Experimental Dates Trial (M/D/Y) 12/11/8212/20/82 2 1/10/831/20/83 conditions Exper imenta 1 deer useda 4 d" castrates, 4 ~ , all aged H v rs+ 4 d' cas trates, 4 ~ , a 11 aged 2~ v rs+ 3 4 2/ 7/832117/83 3/ 9/833/19/83 for 1982-83 cover trials. 5 d' cas trates, 3 ~ , all aged H yrs+ 5 r:J" castrates, 3~, all aged H v r s+ Trial conditions Isolation pens devoid of snow; highest wind velocity and prolonged periods of wind. Isolation pens covered by snow 3 of 6 days when data collected; coldest average amb ient temperatu re; essentially no wind during trial. Isolation pens covered by snow all 6 days of data collection; wind infrequent and of low velocity. Isolation pens covered by snow 5 of 6 days of data collection; warmest average ambient temperature; wind erratic in direction and moderate in velocity; series of snowstorms throughout trial; persistent low elevation clouds. aBeginning with trial 3, an additional for a sick female. male castrate was substituted �w Table 2. Ambient conditions 0 Temperature Ambient SE Min Month x Dec -3.2 0.3 2 Jan -9.4 3 Feb 4 Mar Trial co during 1982-83 cover trials. C Black Globe SE Min Max Solar radiation (watts/m2) x SE Radiant heat load (watts/m2) x SE Total solar radiation (cal/cm2/day) x SE - Max x - -18 9 -1.7 0.4 -19 19 225 10 322 4 164 20 0.3 -19 1 -4.5 0.5 -20 19 291 10 312 4 225 17 -3.6 0.3 -19 8 1.6 0.6 -20 26 352 12 363 6 296 19 -1.2 0.2 -9 8 4. 1 0.5 -13 29 403 13 395 7 344 31 - - ------------------------------------------------------------------------------------------------------ Tri a I Month Wind veloci t/ .(m/sec) 0 ~3 0-99 b Wind di rection (degrees) 90-179 180-269 270-359 Cloud cover c (%) 0-25 > 50 d Cloud elevation High Low Snowing Dec 50 20 5 13 35 47 48 42 39 61 2 2 Jan 92 0 0 0 8 92 51 37 38 62 10 3 Feb 69 I 8 22 7 63 36 48 35 65 9 4 Mar 45 8 24 22 18 36 9 87 86 14 33 aData bData ~Data Data eData are are are are are percentages percentages percentages percentages percentages of of of of of 432 samples of wind velocity/trial. samples with velocity ~1 m/sec, 432 samples of percent cloud coverage/trial. samples with clouds present. 432 samples of active snowfall/trial. e �39 Table 3. Percent of total observations during 1982-83 cover trials. Trial 2 Trial I December Activity by activity .Janua ry for 8 adult mule deer Tri a I 3 February Trial 4 March Lying 65 65 70 70 Standing 14 12 12 14 Feeding 14 16 12 11 Othera 7 7 6 5 3,456 3,456 3,456 3,456 Total (N)b alnclusive of walking, pacing, bTotal number of observations running. at 10-minute intervals. �40 Table 4. trials. Body weights and food consumption TRIAL I --DECEMBER a Feed Feed Body weight (kg) offered consumed (g) (%) Beg in End % chng Deer of deer during 1982-83 cover TRIAL 1 -- JANUARY Feed Feed Body weight (kg) offered consumed Begin End % chng. (g) (%) 85 88 +4 10,750 100 86 87 +1 9,648 100 180cf' 81 81 a 10,370 100 80 78 -3 9,306 100 203 c:i' 79 75 -5 9,980 75 80 72 -10 9,306 56 227 d'1 60 60 a 8,450 100 60 61 +2 7,587 100 182 ~ 59 56 -5 8,060 75 54 54 a 6,894 100 225 ~ 54 53 -2 7,680 100 52 51 -2 6,552 100 191 ~ 56 53 -5 7,680 35 52 52 0 6,552 78 000 ~ 60 60 a 8,450 100 59 58 -2 7,236 100 144 6" TRIAL 3 --FEBRUARY Feed Feed Body weight (kg) offered consumed (g) (%) Beg in End % chng Deer TRIAL 4 -- MARCH Feed Feed offered consumed Body weight (kg) (g) (%) Beg in End % chng 144 ~ 84 83 -1 9,648 100 79 79 a 8,964 180 (j'f 76 75 -1 8,955 100 72 70 -3 8,613 203 @ 72 70 -3 8,613 94 68 68 o 8,271 100 227 6" 58 58 a 7,236 100 57 58 +2 7,236 100 190 r? 72 69 -4 8,613 100 70 70 o 8,271 100 225 ~ 52 48 -8 6,552 78 47 48 +2 6,201 100 191 ~ 50 49 -2 6,552 66 50 49 -2 6,552 100 000 ~ 58 55 -5 7,236 100 55 54 -2 6,894 100 100 aDeer fed daily a restricted diet estimated to provide 81% of maintenance digestible energy (160 kcal DE/kg BWO.75 = maintenance; Ullrey et al. 1969); restricted diet fed for 10 days in trial 1 and for 9 days in trials 2-4. Pelleted ration was 74% digestible (in vitro, dry matter basis) and estimated to contain 3.37 kcal digestible energy/gram. Amount of feed offered was adjusted to pre-trial body weights. �41 Table 5. Analysis of variance for the 4 trials. Design is a randomized complete block. Left column is based on 4 treatments per block (tree, soil, wind, control) ~nd right column is based on 2 treatments per block (on any cover, control). Blocks represent deer (n = 7). Source df F F-Prob. Source df F F-prob. 5.722 0.054 0.550 0.757 9.553 0.021 0.594 0.729 5.730 0.054 0.557 0.753 14.769 0.009 0.520 0.777 20.292 0.004 0.556 0.753 TRIAL 1 -- DECEMBER Treatments 3 1:098 0.376 Treatments Blocks 6 1.038 0.434 Blocks 6 Error 18 Error 6 TRIAL 2 -- JANUARY Treatments 3 1.206 0.336 Treatmen ts Blocks 6 0.401 0.869 Blocks 6 Error 18 Error 6 TRIAL 3 -- FEBRUARY Treatmen ts 3 3.110 0.052 Treatments Blocks 6 0.650 0.690 Blocks 6 Error 18 Error 6 TRIAL 4 -- MARCH Treatments 3 2.724 0.075 Treatments Blocks 6 0.422 0.855 Blocks 6 Error 18 Error 6 ALL TRIALS Treatments 3 3.628 0.033 Treatments Blocks 6 0.603 0.725 Blocks 6 Error 18 Error 6 �42 Table 6. Weighted frequency of use of individual cover treatments by deer in relation to wind velocity during 1982-83 trials. Values are transformed and represent all trials pooled. Wind velocity, Treatment Tree Windbreak Soi 1 Control 0 1-2 27,333 8,848 8,263 m/sec 5-6 7-8 545 394 121 3,000 1 ,132 961 224 29,848 17,061 2,091 485 30 4,075 1 ,821 309 113 16 3-4 �43 Fig. 1. Isolation pen with cover treatments: 1) upper left, windbreak; 2) lower left, artificial tree; 3) upper right, conical mound of soil; 4) lower right, control quadrant having no cover structures. In the center is a feed box for pelleted ration. Fig. 2a. Deer lying next to windbreak structure. �44 Fig. 2b. Deer lying under artificial Ffg. 2c. Deer lying on mound of soil. tree. �45 _. H H H H f-! -f-! n n <: rx: 80 -e c<) N 90 H <: <: H p:; p:; E-< n "...... eo ~ '-' 70 144 f-! ::c 0 ~ H W ::;: 60 _,~~-t 190 • 180 203 :>< Q 0 p::) 50 40 \ 'a 182 OCT NOV DEC JAN FEB MAR APR MAY MONTH Fig. 3. Body weights of adult male (--) and female (---) mule deer in cover experiments, October 1982 ·to April 1983. Deer identification numbers are shown at right and timing of cover trials 1-4 is shown at top of graph. �46 r»; <r -ot r-l r-l "....,. rr-; II') r-l r-l r-l II') N N r-l <;» r-, 0 6 N r-l ....:l 0 0::: '-.../ "....,. <;» <:» 5 E-< Z 0 u <-; 0::: ~ ::> 0 u 4 3 ~ 0 0 2 H E-< < 1 0:: 1 DEC 2 3 4 JAN FEB MAR TRIAL Fig. 4. Ratios of weighted observations of deer lying on cover treatments (soil, tree, and windbreak) to weighted observations of deer lying on control areas during days 1-3 ([]) and days 4-6 (.) for each trial. Ratios based on transformed data while actual sample sizes of lying observations are shown in parentheses. Data based on 8 deer per trial. �47 5 4 r»: C) ()) C/J E: 3 WIND VELOCITY 2 AT 2 m HEIGHT 1 = 0 <:» 4 ~ 3 • H u 0 .....:I 2 t:.Ll :> 1 C! Z :::;:: 7.1m/sec WIND VELOCITY AT 2 m HEIGHT = 3.8m/sec 0 H 4 • 3 • 2 1 • • 0 2 1 • --- WIND VELOCITY AT 2 m HEIGHT = 4.0m/sec ··0 DISTANCE FROM WINDBREAK (m) Fig. 5. Wind velocity profiles on leeward side of windbreak at heights of 30 cm (II) and 60 cm (4t). Velocities measured with a Casella anemometer having sensitivity of 0.002 ft/sec . ••_:. ; '0 • �48 CLOUD COVER 0<25% 81 26-50% • >50% ~,.. ....~ ..•.. :. .,; 1.2 z 0 H E--< <C H Q <C p:; p:; <C ....:l 0 o: • ~ 6 5 0.8 • r>: o Q) (fJ 4 e:; <;» 'M e:; 0.6 3 :>-< E--< e:; 0.4 2 U 0 ro 0.2 o 1 w - N o H ....:l r-l ::> o 0.0 Q Z H ::s: 25 r»; U 0 20 '-' 15 w p:; ::J 10 <C p:; W 5 ~ 0 E--< -5 E--< W w p:; E--< W 25 r-- 20 OU '-' 15 w p:; .: ~ w p:; ::J I'Q W Q Z H ~ E--< <C p:; ~ W E--< 10 5 0 -5 7- 8 9 10 11 12 13 14 15 16 17 18 TIME (hrs) Fig. 6. Hourly solar radiation, cloud cover, .and black globe temperatures measured during May 1983. Black globe temperatures measured simultaneously at t m (.) and H m (A) distance from a windbreak panel with a southern exposure and at the base of an artificial tree. Ambient temperature ( ,lower graphs) and solar radiation measured at nearby weather station. Wind velocity measured at height of 2 m near cover structures with Casella anemometer and cloud cover occularly estimated each hour. �49 APPENDIX A PROGRAM NARRATIVE State of Project No. Colorado 5502X --~~~-------------- Work Plan No. Job No. A. 2 ----~------------1 Big Game Investigations - Cervids Deer Investigations Nutritional Basis for Quantifying Capacity of Winter Range-Evaluation of Thermal Cover Used by Deer NEED Maintaining homeothermy is critical to survival of mule deer (Odocoi1eus hemionus) during winter. To maintain body temperature, deer must balance heat input with heat loss. During winter, deer obtain energy through food consumption and reduce energy loss by minimizing physical activity and utilizing cover to reduce thermal gradients (Jacobsen 1973, Moen 1973, Short 1981). Behavioral studies of free-ranging ungulates during winter have shown animals to use certain plant communities apparently for cover (Verme 1965, Moen 1966, Loveless 1967, Rongstad and Tester 1969, Ozoga and Gysel 1972, Beall 1976, Peek et al. 1976, Leckenby 1977). However, these studies failed to elucidate whether cover was used for escape from predators or for thermal regulation, or that use may have been independent of cover attributes and more correctly reflected responses by animals to areas having reduced snow depths (Gilbert et a1. 1970). Studies of thermal cover are limited. Verme (1965) and Ozoga (1968) showed that conifer stands used by white-tailed deer (Odocoi1eus virginianus) in.winter provided energetically favorable microclimates, but these authors could not separate effects on deer of microclimate and reduced snow depth. Beall (1976) presented limited information suggesting bedding sites used by elk (Cervus canadensis) were related to their favorable thermal characteristics. Robinson (1960) attempted to test the effect of cover on body condition of captive white-tailed deer during winter by placing deer in conifer stands having different canopy densities. As body condition of deer did not differ between treatments, he concluded that deer could select and use sparse cover to create as favorable microclimate as in heavy cover. Moen (1966) suggested white-tailed deer on a highly nutritious diet could survive winter without cover. He suspected that cover became physiologically important only when the diet did not provide sufficient metabolizable energy. This viewpoint was supported by Swift et al. (1980) who concluded that active thermoregulation by mule deer and elk during winter �50 placed minimal demands on these animals' energy reserves. Furthermore, these authors felt that heat increment of feeding would normally be sufficient to offset the need for thermoregulation. Unlike Moen (1966), Swift et a1. (1980) based their conclusions on deer and elk subject to a typical winter diet low in digestibility and protein. Stevens (1972) approached evaluation of cover from a perspective of thermodynamics and processes of heat exchange between white-tailed deer and their winter environment. Her measurements and simulations provided a good basis for understanding mechanisms of heat transfer that affect energy balance of deer. Modes of heat transfer between deer and their environment are conduction, convection, radiant exchange and evaporation (Stevens 1972, Campbell 1977). Evaporation is usually unimportant to deer during winter (Stevens 1972). These modes of heat transfer generally interact. During winter, mule deer commonly associate with southern aspects having steep slopes and reduced snow depth. Deer could be using these areas because of reduced snow (Gilbert et al. 1970) or because of enhanced microclimates. During a sunny day on a southern exposure, a deer could decrease heat loss by lying on bare, warm soil to reduce conduction, by absorbing direct solar radiation, and by being sheltered from northerly winds to reduce convection. At night, this deer could lie under trees, if present, to reduce radiant heat loss to a cold night sky (Moen 1966) and additionally, continue to reduce heat lost due to conduction and convection. Observations of free-ranging deer do not necessarily elucidate heat loss mechanisms of paramount importance to deer, although reducing the effects of wind have been considered extremely important (Moen 1966, Stevens 1972, Staines 1976). As heat transfer mechanisms potentially differ in importance from day to night, types of cover utilized by deer could differ provided deer had the opportunity to select from among different cover types. Importance of cover types may also change during winter as the insulative value of deer pelage changes as winter progresses (Jacobsen 1980). If deer were provided cover types that reduced heat loss via specific mechanisms of heat transfer and their use of these types monitored, our ~nderstanding of when and why deer use cover would be enhanced. However, Moen (1966) commented that discerning between preferences of deer and their physiological needs is a difficult distinction. Wal1mo and Schoen (1981) aptly summarized our knowledge about cover: "The general term 'cover' refers to various animal-environment relationships that have been widely discussed but seldom measured." Clarifying the use of thermal cover by deer is necessary if land managers are to properly evaluate habitats used by deer. Proper integration of cover requirements into habitat manipulation programs involving herbicides, burning, or chaining depends on a more thorough understanding of what types of cover are important to deer. �51 B. OBJECTIVES The objective of this research is to assess use by tame mule deer of cover types providing different modes for deer to reduce heat loss during winter. Conceptually, this research will progress in 2 phases. The first phase (this study plan) will be an observational study of deer using cover types under controlled conditions and will focus upon cover used when deer are bedded. Phase 2, assuming phase 1 is successful, will assess cover selection by deer in relation to nutritional stress. C. EXPECTED RESULTS OR BENEFITS Completion of this research will aid in developing habitat evaluation system by establishing the role of cover in maintaining energy balance of deer during winter. The habitat evaluation system will provide a quantitative basis for the Division of Wildlife to assess and improve winter ranges used by deer. D. APPROACH The approach of this study will be to monitor selection of artificial cover by tame deer when bedded and confined in isolation pens during winter. Articicial cover will be provided to offer deer the simultaneous choice of reducing heat loss by minimizing heat transfer due to conduction, convection, or radiation. Rationale: Providing deer with different types of artificial cover, each designed to reduce heat loss primarily via a specific mode of heat transfer, will enable deer to reveal through behavioral choice what modes of heat transfer are important to maintaining heat balance. Hypotheses l. 2. 3. 4. that will he tested: Deer will exhibit Deer will choose Deer will choose Deer will choose (December-March). no preference for cover types. cover types independen t of time of day. cover types independent of ambient conditions. cover types independent of time of year Observations of deer responses to cover will be made during 4 trials beginning in mid-December and ending in mid-March (see Schedule). Data collection periods will consist of 6 days during each trial (Appendix I) whereby deer are observed during 4 3-hour shifts: 0300-0600, 0900-1200, 1400-1700, and 2000-2300 hours (MST) (Fig. 1). These observation shifts will include the most energetically favorable and most detrimental ambient conditions. �52 Experimental Deer and Their Maintenance Eight adult (2 1/2 yrs+) tame deer (4 male castrates, 4 females) will be used for this experiment. An additional adult male castrate will be held in reserve. Body weights of these deer range from 57 to 94 kg (Table 1). Body size can influence rate of heat loss (Stevens 1972) so this range in body weights m~y provide insight into its effects on cover use by deer. Table 1. Body weights (kg) of experimental deer. ~leighta Deer Sex 144 180 203 227 M M M M 94 86 85 66 191 000 182 225 F F F F 66 63 60 57 ~eights as of 15 November 1982. Deer will be maintained on a deer-elk pelleted ration having approximately 15% protein, 55% digestibility, and 2.5 kcal/gm of digestible energy (D. Baker, pers. commun.). This diet is l~ss digestible than either the commercial dairy or Pawnee Special rations used previously which should help to reduce the effect of diet quality on the need for deer to use cover. Deer will be conditioned~to the new ration for 2 weeks during October at the Foothills facility and then ad libitum intake will be determined for a 7-day period. Additionally, a complete in vivo digestion trial using 3 deer will be conducted to determine digestible dry matter of the ration (D. Baker, pers. commun.). While at the Foothills facility, deer w±ll be weighed at 2-week intervals to monitor adaptation to the ration. After deer have been transported to the Junction Butte facility in mid-November, they will be fed the ration at an ad libitum rate and have access to native forage. However, during experimental trials, deer will have access only to pelleted ration fed individually to each deer at a rate of 60% maintenance (maintenance equals 160 kcal DE/kg BwO·75, Ullrey et al. 1969). If these deer were fed at the 60% maintenance level for 120 days (Dec.-Mar.), they would lose 10-17% of their body weight as predicted from the regression equation of Ullrey et al. (1969) for adult white-tailed does. This level of feeding �53 would not appear to be overly restrictive as Baker et al. (1979) found mule deer fawns tolerated weight loss of 14-16% during winter. As a fully fed deer may not need to seek cover (Moen 1965) this reduced diet may stimulate their need to use cover during trials and more closely mimic nutritional status of wild deer. During trials, deer will be fed one-half their daily ration in the early morning and again in late afternoon (Fig. 1). This feeding schedule attempts to simulate peak grazing periods of adult mule deer which occurred at 5-6 PM, 12 PM, and S AM (L., H" Carpenter, unpubl. Job Progress Rep. W-3S-R-3l, Colo., 1976). Observation of cover used by deer will not commence until 2 hours after feeding to reduce the effect of an initial spike in heat of fermentation on body heat balance and possibly cover use. By feeding twice instead of once per day, effects of heat of fermentation on animal heat balance will be more evenly distributed throughout a 24-hour period. Deer will be weighed before and after each trial and at 2-week intervals between trials. Before the initial trial, deer will be encouraged to voluntarily enter and adjust to isolation pens by placing pelleted ration in pens. If the voluntary approach fails, deer will be preconditioned to pens and cover types by restricting deer to pens. Isolation Pens Eight isolation pens 10 x 10 x 2.2 m high will be constructed within a 2.2-ha pasture of sagebrush (Artemisia tridentata wyomingensis) enclosed by a 2.4 m high, l5-cm mesh woven wire fence (Fig. 2). All pens will consist of l5-cm mesh woven wire supported by steel fence posts. Gates into pens will be 1 m wide x 2 m high and have 1.9-cm steel rebar for a frame covered by l5-cm mesh woven wire (Fig. 3). At the center of each pen will be a feeder consisting of a wooden box supported by a 1.S-m steel post (Fig. 3). This feeder will be capable of holding 3 kg of pelleted ration for deer. Centering the feeder within the pen will reduce potential bias of the feeder on deer use of cover types. Pens will be located within the pasture to: (1) minimize potential topographic differences between pens, (2) be at least 10 m from any permanent fence, structure, or other isolation pen, and (3) within an SO-m viewing radius from an observation tower (Fig. 2). All sagebrush within pens and within an area 5 m around the boundaries of pens will be mechanically chopped and reduced to a height <6 cm to remove potential effects of natural cover in and around pe;s. Additionally, all debris and sagebrush stubble within pens will be removed to reduce potential effects of this material on selection of artificial cover by deer. Vegetation remaining in pens shall consist only of native bunchgrasses and low growing forbs. �54 Deer will be randomly assigned to an isolation pen each trial. Additionally, deer will occupy a specific pen only once during the 4 trials. Artificial Cover Deer will be provided 4 choices of cover': (1) an artificial tree to reduce radiant heat loss to the cold night sky, (2) a windbreak to reduce convective heat loss, (3) a mound of black colored soil to reduce heat lost via conduction when lying on snow or frozen ground, and (4) no artificial cover, which will consist of an area covered by snow. The 3 sources of artificial cover will encompass approximately equal areas (3.2 m2) and will be centered in 1 of 4 quadrants within each isolation pen (Fig. 3). Cover types will be randomly assigned to each of 4 quadrants in each of 8 isolation pens (Fig. 4). To balance distribution of cover types among quadrants, each cover type will occur twice in each of the 4 quadrants throughout the 8 pens. This will minimize systematic effects of unknown factors that might make 1 quadrant more or less hospitable to deer. Artificial Tree The artificial tree will consist of a circular piece of 1.O-cm plywood supported in the center by a 10 x 10 cm post (Fig. 5).' The plywood surface will be solid to maximize po t en.tLaL effect of reducing radiant heat loss. The plywood surface will be 1.2 m above the ground, or lower if deer will tolerate a lower height. The support post will be painted white to reduce absorption and emission of radiation while the top and bottom surfaces of the plywood will be dark green. Artificial Windbreak The artificial windbreak will consist of 4 pieces of 1.0-cm plywood 1.80 x 0.61 m high attached together to form a square (Fig. 6). Support braces will prevent deer from entering and using the area inside the square for cover (Fig. 6). Plywood will be painted white to reduce absorption and emission of solar radiation. Additionally, holes 1.3 cm in diameter will be drilled to remove approximately 0.4% of the surface area to allow some air to flow through the windbreak. This air flow will further reduce the reflective heat that may be afforded to the deer by the plywood surface. Sides of the windbreak will face northwest, southeast, northeast, and southwest. Prevailing winds are from the H-NW and S-SW, but the windbreak will offer protection to deer from winds in all directions. �55 Dark Soil A conical mound of black colored, finely fragmented and weathered shale soil 2 m in diameter and 0.17 m high at the center will be provided as an absorbing surface upon which deer may lie down (Fig. 6). This mound will be kept free of snow by manually removing snow on a daily basis. Control The control will be the lack of artificial cover and will encompass entire quadrant. The control type will also occur in each quadrant those areas not considered part of another cover type (Fig. 3). an in Snow Management The presence of snow is important to this study, primarily to insure the dark soil surface offers an advantage to deer over lying on snow. However, large amounts of snow would be detrimental as deep snow could provide thermal cover and negate the effectiveness of the artificial cover. I will attempt to keep snow depth 210 em ~.,rithin isolation pens and in the 5~m area around pens by packing snow with snowmobiles and snowshoes. Snow will be added to pens if small areas of natural soil become exposed. Data Collected Animal Behavior Behavior of all 8 deer will be monitored simultaneously during the 4 observation shifts. Time (minutes) spent by deer within each cover treatment will be the primary response measured. Although continuous measurement of time spent in a cover type is preferred, in reality continuous observation of deer is not possible, especially at night. Thus, a sampling scheme must be used to estimate total time deer spend in a cover treatmen t. Deer use of cover types will be sampled atlO-minute intervals during all shifts. At the 10-minute intervals, status of the 8 deer will be quickly observed as a form of scan sampling (Lehner 1979). At night, a high intensity spotlight will aid observation. Jacobsen and ~Viggins (1982) found thelO-minute interval provided an unbiased estimate of time in activity for deer during both diurnal and nocturnal periods. Estimates of time in a cover treatment (TIC) will be computed as TIC (x/n) t where (x) is the number of observations of each cover treatment used, (n) the total number of observations obtained, and (t) the total time over which the observations were obtained. In 1 3-hour sampling shift, (n) = 18 and (t) = 180 minutes per deer and for I 6-day trial, (n) = 432 and (t) = 4,320 minutes. �56 Deer behavior will be classified l. 2. 3. 4. 5. 6. as follows: Bedded Bedded, head up Bedded, head down Standing Feeding Other Once deer are assigned to a behavioral category, they will be assigned to 1 of 4 quadrants in each isolation pen and classified as either on or off treatment. Criteria for being on a cover treatment will be: 1. 2. 3. 4. Artificial Tree: At least 50% of the deer's body must be underneath the downward vertical projection of the tree surface. Artificial Windbreak: At least 50% of the deer's body must be within 1 m of the windbreak surface. Dark Soil: At least 50% of the deer's body must be within the vertical projection of the soil surface. Control: A deer anywhere within the control quadrant will be considered on the control. Also, if deer are within quadrants other than the control and are considered to be "off" treatment, they will be considered to be on the control. Additionally, intervals at which deer change behavioral states or cover treatments will be specifically noted as these occurrences may provide reliable estimates of ambient conditions that trigger behavioral responses to different modes of heat transfer. Ambient Conditions The following 1. measurements of ambient conditions will be made: Black globe temperature provides an index of the combined effects of radiant energy, air temperature, and air velocity (Bedford and Warner' 1934, Bond and Kelly 1955) and can be used as an index to compare ambient conditions that affect the comfort of animals between days and ti~e of day. Provided accurate measurements of wind speed are obtained, radiation intensity (positive and negative) can be calculated from globe temperature. A globe thermometer will be constructed using a 10-cm diameter copper toilet-float painted flat black with a sensing element of a thermometer (-50 to +50 C) placed at the center of the float (Bond and Kelly 1955). During winter, black globe temperature is typically higher than ambient temperature during the day and lower at night (L. Renecker~ pers. commun.). Globe temperature at 1/2 m above the ground will be read manually each half-hour beginning each sampling shift. �57 2. Ambient temperature will be measured continuously using a selfrecording thermograph (Belfort, -35 to +110 F). Additionally, a min-max thermometer (-45 to +50 C) will be monitored daily. Temperature instruments will be housed in a standard weather shelter placed 1/2 m above the ground. 3. Wind speed and direction will .be .mon Lt ored continuously using a self-recording 3-cup anemometer (0 to 50 m/sec) and wind vane (WeatherMeasure). Measurements will be made a 2 m above the ground. Wind chill indexes will be calculated according to Ames and Insley (1975). 4. Direct and diffuse solar radiation will be monitored continuously (daylight hours) using a self-recording pyranograph (Belfort) placed 1/2 m above the ground. 5. Percentage cloud cover will be visually estimated at one-half hour intervals beginning each sampling shift. Percentage classes will be 0%, 1-25%, 26-50%, 51-75%, and 76-100%. Clouds will be classified as high in elevation (cirrus, cirrostatus, cirrocumulus, altostratus, altocumulus) and low in elevation (nimbostratus, stratocumulus, stratus), or fog (Ordway 1966). Weather instruments will be placed in an area 10 x 15 m (Fig. 2) where sagebrush will be reduced to a height <6 cm. Data Analysis; This experiment will be a randomized complete block design with deer treated as blocks (random effect) and each block subject to 4 cover treatments (fixed effect). Analyses will assess differences in amount of time deer spend in cover due to treatments, blocks, sampling shift, and trials. Use of cover treatments in relation to ambient parameters will be assessed using a multivariate approach. It is anticipated that use of cover types will be described within "limits" of ambient parameters, either acting singularly or in combination. Schedule Activity Period July-November September October November 1982 1982 1982 1982 Literature review and development of study plan. Construct isolation Deer conditioned pens. onto ration. Deer transported to Junction Butte facility; begin behavioral adjustment to pens. • �58 December January 1982 1983 February March Trial l--December Data summary Trial 2--January Data summary 1983 April-June 10-21 Trial 3--February iData summary 1983 Trial 4--March Data summary 1983 10-21 10-21 13-24 Data analysis, prepare yearly reports. Personnel David J. Freddy Wildlife Tech. I-A Clerk-Typist Principal Investigator Field Assistance Secretarial support Estimated Annual Costs Person-Days (01) Personal Services 264 llO 22 David J. Freddy Wildlife Tech. I-A Clerk Typist (21) Operating (28) Travel (31) Capital Costs Supplies $ 30,348 5,830 1,334 and Services 9,8ll 1,040 Expenses 0 Total E. $ 52,815 LOCATION Field work will be conducted at the Division of Wildlife's Junction Butte Research Center located 5 km south of Kremmling, Colorado. F. RELATED FEDERAL AID PROJECTS 5502-X, Work Plan 2, Jobs 6 and 10 5503-X �59 LITERATURE Ames, D. R., and L. W. Insley. 1975. sheep. J. Anim. Sci. 40:161-165. CITED Wind-chill effect for cattle and Baker, D. L., D. E. Johnson, L. H. Carpenter, O. C. Wallmo, and R. B. Gill. 1979. Energy requiremp.nts of mule deer fawns during winter. J. Wildl. Manage. 43:162-169. Beall, R. C. 1976. Elk habitat selection in relation to thermal radiation. Pages 97-100 in Hieb, S. R., ed. Proceedings of the Elk-Logging-Roads Forest, Wildl., and Range Exp. Sta., Univ. Idaho. 142pp. Bedford, T., and C. G. Warner. 1934. The globe thermometer heat and ventilation. J. Hygiene 34:458-473. in studies of Bond, T. E., and C. F. Kelly. 1955. The globe thermometer research. Agric. Engr. 36:251-255, 260. in agricultural Campbell, G. S. 1977. An introduction Springer-Verlag, New York. to environmental biophysics. Gilbert, P. F., O. C. Wallmo, and R. B. Gill. 1970. Effect of snow depth on mule deer in Middle Park, Colorado. J. Wildl. Manage. 34:15-22. Jacobsen, N. K. 1973. Physiology, behavior, and thermal transactions 346pp. white-tailed deer. PhD. Thesis, Cornell Univ. of 1980. Differences of thermal properties of white-tailed deer pelage between seasons and body regions. J. Therm. Bio.l. 5:151-158. , and A. -----activity O. Wiggins. 1982. Temporal estimated by time-sampling. and procedural influences on J. Wildl. Manage. 46:313-324. Leckenby, D. A. 1977. Management of mule deer and their habitat: applying concepts of behavior, physiology, and microclimate. Proc. Western Assoc. State Game and Fish Comm. 57:206-217. Lehner, P. N. 1979. Handbook New York. 403pp. of ethological methods. Garland STPM Press, Loveless, C. M. 1967. Ecological characteristics of a mule deer winter range. Colo. Game, Fish and Parks Dep. Tech. Bull. 20. l24pp. Moen, A. N. 1966. Factors affecting the energy exchange and movements of white-tailed deer, western Minnesota. PhD. Thesis, Univ. of Minnesota. l2lpp. 1968. Energy exchange Ecology 49:676-682. of white-tailed deer, western Minnesota. �60 1973. Wildlife ecology: an analytical and Co., San Francisco. 458pp. Ordway, R. J. 1966. Earth Science. Princeton, New Jersey. 705pp. Ozoga, J. J. 1968. Variations yard in northern Michigan. approach. D. Van Nostrand Co., Inc. in microclimate in a conifer swamp deerJ. Wildl. Manage. 32:574-585. --- , and L. W. Gysel. weather. W. H. Freeman 1972. Response of white-tailed J. Wildl. Manage. 36:892-896. deer to winter Peek, J. M., D. L. Ulrich, and R. J. Mackie. 1976. Moose habitat selection and relationships to forest management in northeastern Minnesota. Wildl. Mono. 48. 65pp. Robinson, W. L. 1960. Test of shelter requirements deer. J. Wildl. Manage. 24:364-371. of penned white-tailed Rongstad, O. J., and J. R. Tester. 1969. Movements and habitat use of white-tailed deer in Minnesota. J. Wildl. Manage. 33:366-379. Short, H. L. 1981. Nutrition and metabolism. Pages 99-127 in O. C. Wallmo, ed. Mule and black-tailed deer of North America. Univ. of Nebraska Press, Lincoln. 605pp. Staines, B. W. 1976. The use of natural shelter by red deer (Cervus elaphus) in relation to weather in northeast Scotland. J. Zool. Lond. 180:1-8. Stevens, D. S. 1972. Thermal energy exchange and the maintenance of homeothermy in white-tailed deer. PhD. Thesis, Cornell Univ. 23lpp. 1980. Nitrogen and energy Swift, D. M., J. E. Ellis, and N. T. Hobbs. requirements of North American cervids in winter--a simulation study. Pages 244-251 in E. Reimers, E. Gaare, and S. Skjenneberg, eds. Proc. 2nd Int.~eindeer/Caribou Symp., Roros, Norway. Ullrey, D. E., W. G. Youatt, H. E. Johnson, L. D. Fay, B. L. Schoepke, and W. T. Magee. 1969. Digestible energy requirements for winter maintenance of Michigan white-tailed does. J. Wildl. Manage. 33:482-490. Verme, L. J. 1965. Swamp conifer deeryards in northern ecology and management. J. Forestry 63:523-529. Michigan: their Wallmo, O. C., and J. W. Schoen. 1981. Part 2. Forest management for deer. Pages 434-448 in o. C. Wallmo, ed. Mule and black-tailed deer of North America. Univ. Nebraska Press, Lincoln. 605pp. �61 APPENDIX I Dailey Schedule for Each Trial Day 0: Feed removed from bulk-feeder mid-day. in holding pen. Deer weighed and placed Day 1: Deer placed in isolation pens using pelleted ration as a reward. Begin feeding at 60% maintenance and behavioral adjustment to pens. Day 2: Continue as day 1. Day 3: Continue as day 1. Day 4-9: Observation of cover selection, data collection. Day 10: Deer remain in isolation pens, fed 80% maintenance diet. Day 11: Deer removed from isolation pens, weighed, and returned to ad libitum ration fed in bulk. �62 1200 I· 0900 0300 fEED ~M \,\ n"''''\\'~ . 0600 Fig. 1. Clock times of 4 observation shifts (heavy lines) and times at which deer will be fed pelleted r~tion. �63 170m 30m :. . ~'- ~ ~ .' ""-- Weather Instrumentation N 00 E approx. Fig. 2 • Location of 8 10m x 10m isolation pens (numbered anrl crosshatched) within a 2.2 ha pasture of sagebursh. Areas 20m x 20m (hatched) had sagebrush reduced to a heif,ht less than 6cm and aueas within pe~s had sagebrush stubble remov~d. Dotted line denotes area containin~ 54 potential 10m x 10m pens of which 8 were chosen. �64 STEEL POSTS~ GATE • • CONTROL TREE COVER WIND COVER 00 E approx • DARK SOIL COVER •1 m .•.... Fig. 3 • Design of 10m x 10m x 2.2m high isolation pen constructed of 15 em mesh woven wire. Cover types centered within 4 quadrants and feeder located at center of pen. �65 PEN 1 PEN 2 PEN 3 I ~ A 1 D I B C __ ._._ . B -- I I D ~_-t-- B I PEN 5 I D C • 1---.--A , I ,B .A -'I - - 1-- B C '-' I PEN 6 I I ,.. I I PEN 4 -- -1- f- I A C A D I C I 1 A - - l- - - I D D • B I PEN 7 PEN 8 , I C •... I I B - -.I ...• D A B I- l.c ---'-~L A I D .......... / / . . N 00 E approx. A = B C = :D = = Tree cover Wind cover Control, no artificial cover Black Dirt cover ~Fig. 4. Assignments of cover types within 1 of 4 quadrants in each of 8 10m x 10m woven wire isolatinn pens. �66 2.0 m t 50 em 2 120 10 x POST 10 x em em GROUND SIDE VIEW t 10 x 10 / em POST TOP Fig. 5. The artificial tree cover treatment. VIEW 10 em SUPPORT �67 SUPPORT 2 x 10 , • • 1 0.61 m J . • • • • • • • • •• • • • • • • • • VENTILATION HOLES • 1.8 m 17cmt~ 2.0 m 6 • TOP. The artificial windbreak cover treatment. Ventilation holes will be in all 4 panels. BOTTOM. Cross-section of the conical mound of dark soil cover treatment. Fig. ��Colorado Division of Wildlife Wildlife Research Report July 1983 69 JOB PROGRESS REPORT State of Colorado Project No. 45-01-502-15050 Work Plan No. 2 ------------------- Job No. 6 Period Covered: Author: 7/1/82-6/30/83 Big Game Investigations - Cervids Deer Investigations Winter habitat selection and activity patterns of mule deer in Front 'Range shrubland and forest habitats R. C. Kufeld Personnel: R. C. Kufeld ABSTRACT Fourteen adult female deer, 21 fawns and 2 bucks were captured in late January and early February, 1983, at Lory State Park. Does were fitted with radio collars and ear tags. Fawns and bucks were ear tagged. Telemetry triangulation data showing approximately 1500 deer locations and 250 hours of recorded~around the clock, deer activity inv91ving 25 radiocollared deer were collected during the 15 November 1982 through March, 1983, period. These data are currently unaergoing analysis. Two of 12 radio-collared deer migrated from Lory State Park in June, 1982, and returned in late October. They spent the summer in a mountain valley 26 km west of the study area. The other 10 deer plus the 14 deer captured during the winter of 1983 have remained within 1.6 km of their capture point as of June 1, 1983, when last checked. A population of 128 ± 13 deer (x ± t.025SE) deer was projected for Lory State Park based on 10 counts of tagged and un tagged deer. Ten such counts are sufficient to estimate the deer population within 10 percent. of. the mean with' 95 .percent confi dence. ��2 71 WINTER HABITAT SELECTION AND ACTIVITY PATTERNS OF MULE DEER IN FRONT RANGE SHRUBLAND AND FOREST HABITATS Roland C. Kufeld P. N. OBJECTIVE 1. To test Telonics telemetry equipment to determine its accuracy at various distances in locating a transmitter on the Horsetooth Mountain study area, and the ability of an observer using that equipment to correctly detect deer activity patterns by habitat type. 2. To determine habitat selection and activity patterns of mule deer within habitat types in the Horsetooth Mountain area during winter. SEGMENT OBJECTIVES 1. Capture up to 10 additional deer on the study area and instrument with Telonics activity collars. 2. Monitor radio-collared mule deer to determine habitat selection activity patterns throughout 24-hour periods. 3. Develop computer methodology and activities for analysis of telemetry them and data on location of mule deer. ACKNOWLEDGEMENTS S. McHugh assisted in deer trapping and collection and tabulation of field data. D. Schrupp coordinated creation of a system for computerizing habitat and telemetry data. METHODS AND MATERIALS Capture and Instrumentation of Deer Fourteen adult female deer were caught in Clover traps (Clover, 1956) in Lory State Park, west of Fort Collins, and fitted with ear tags and Telonics Radio collars between 31 January and 3 February 1983. During this period 19 fawns were also captured and fitted with 1 yellow ear tag. Each radio transmitter has a tip switch which causes pulse rate to vary when the animal~ head is either up or down. Pulse frequency is used to indicate deer activity. �72 Monitoring of Radio-Collared Deer Radio-collared deer were periodically monitored from 2 established triangulation points from 15 November 1982 through March, 1983, to determine habitat selection and activity. Equipment, procedures, and monitoring schedules were described by Kufeld (1981,1982). Eleven deer were monitored during November, December, and January. These~ plus the additional 14 deer instrumented in early February, 1983, were monitored during February and March. The monitoring schedule included 3 sunrise, 3 daytime, 3 sunset, and 3 nighttime periods during Novembe~ January, February, and March, and 2 of each period during December. Each period lasted 6 hours. During the rest of the year, periodic checks using hand-held telemetry equipment were made at intervals of 1 week to 10 days to determine location of all radio-collared deer. Computerization of Habitat and Telemetry Data A computer program was written which will overlay error polygons for each triangulation location on a computerized aerial photo of the study area (Kufeld, 1982), and determine area of each habitat type within polygons when telemetry signal coordinates and signal class designations are entered. Deer Population Estimates Upon completion of trapping on 3 February 1983, 48 tagged deer were present on the study area including 25 adult does with radio collars and orange ear tags, and ..21 fawns and 2 bucks with yellow ear tags. Since this constituted a relatively large sample of tagged deer, the opportunity was taken to estimate deer population density based on counts of tagged and un tagged deer. Ten counts were made between 16 February and 5 May 1983. Each count was made by vehicle and foot, along the same route, which began at the entrance to Lory State Park and ended about 100 m south of Arth~r·s Rock trailhead. Each count began at sunrise and lasted for about 1.5 hours. Deer were counted and classified according to tagged or un tagged using a spotting scope and binoculars. Population size was estimated using a modified "Ll nco ln lndex" technique (Seber, 1973). RESULTS AND DISCUSSION Monitoring of Radio-Collared Deer Telemetry data showing approximately 1,500 deer locations, and 250 hours of recorded deer activity involving 25 radio collared deer were collected during the 15 November 1982 through March, 1983, period. These data are currently undergoing analysis and will be presented in a future report. Triangulation data assembled from 15 November 1982 through March, 1983, and periodic checks during the rest of the year using hand-held telemetry equipment showed that 10 of 12 adult does instrumented during the winter and spririg of 1982 remained within 1.6 km of where captured until 1 June 1983, �73 when this report was written. One deer left the area between 27 May and 9 June 1982 and another left between 9 and 18 June 1982. Both were located on 13 July 1982 together in a mountain valley (2,500 m elev.) 29 airline km west (2740) of the site where instrumented. These deer remained together in this valley during the summer and early fall. Bot~ left the valley and returned to Lory State Park between 25 October and 1 November 1982. One adult doe instrumented 31 January through 3 February 1983 remained within 1.6 km of where captured until 1 June 1983. ' Deer Population Estimates Frequent monitoring of radio-collared does showed all 25 remained within the area where counts were made during the entire 16 February through 5 May 1983 period. Since 21 of the 23 ear tagged deer were fawns and many were offspring of collared does, it is reasonable to assume they were also present during the entire counting period. Based on these 10 counts a population (; ± t.025SE) of 128 ± 13 or from 115 to 141 deer is projected for Lory State Park (Table 1). Upon completion of analysis of 15 November 1983 through March 1983 triangulation data, an aggregate home range area can be computed for all radio-collared deer. That area estimate can be combined with the population projection to determine deer density per km2 in Lory State Park. Ten counts were sufficient to estimate the deer population within of the mean with 95 percent confidence (Table 2). 10 percent LITERATURE CITED Clover, M. R. 199-201. 1956. Single-gate deer trap. Calif. Fish and Game. 42(3): Kufeld, R. C. 1981. Winter habitat selection and activity patterns of mule deer in Front Range shrubland and forest habitats. Colo. Div. Wild]. Game Res. Rep. July (1):97-110. Kufeld, R. C. 1982. Winter habitat selection and activity patterns of mule deer in Front Range shrubland and forest habitats. Colo. Div. Wildl. Game Res. Rep. July:29-34. Seber, G. A. F. parameters. 1973. The estimation of animal abundance and related Hafner Press, New York. 506pp. Prepared by ..:.....:&~-=-~~~C--'~;......__i7'-~_=-=4__ Ro 1and C. Kufe 1d Wildlife Researcher C �74 Table 1. Deer population estimates at Lory State Park based on. 10 counts of tagged and untagged deer conducted between February 16 and May 5, 1983.1/ Date Total Deer Seen Tagged Deer Seen Untagged Deer Seen 2-16-83 2-23-83 28 44 11 17 17 27 2-24-83 3- 3-83 3- 9-83 3-10-83 4-28-83 5- 3-83 5- 4-83 5- 5-83 Total 34 42 30 44 50 46 54 46 11 11 10 18 16 17 20 16 23 31 20 26 34 29 34 30 Estimated21 Popu1a t i.on-> Variance of Estimated2 Population_! 117.4 121.5 477.462 299.803 779.629 141.9 1290.988 174.6 795.069 137.1 238.213 115.1 522.667 146.0 336.322 126.9 264.444 127.3 425.006 134.5 128,±13(;± ·t.025SE)l! 41. 729i1 l/There were 48 tagged deer in the area. 2/Based on an a formula for an unbiased estimate from pg. 105 of: Seber, G.A.F. 1973. 3/The total or pooled estimate was obtained by weighting each individual estimate inversely proportional to its variance. 4/variance for pooled population estimate. �75 Table 2. Number of counts needed to estimate the deer population size within a given percentage of the mean at 2 confidence levels. Percentage of the Mean 5 10 20 Counts Needed per confidence level 95% 90% 39 10 3 28 7 2 ��77 JOB,FINAL REPORT State of Colorado Project No. 45-01-502-15050 Work Plan No.2. -------------------- Job No. 9 Period Covered: Author: Big Game Investigations - Cervids Deer Investigations Piceance Deer Study 7/1/82-6/30/83 R. M. Bartmann Personnel: R. M. Bartmann, D. C. Bowden, G. C. White ABSTRACT Two reports were published, 2 accepted for publication, and 2 were submitted for review. Status of each is in the report for Project 45-01-502-15050, Work Plan 1, Job 7. ��79 PICEANCE DEER STUDY Richard M. Bartmann P. N. OBJECTIVE To complete data analysis and publicatioh of results for field studies completed under Work Plan 2, Jobs 3 and 4. SEGMENT OBJECTIVE Analyze data and report pertinent results of deer population habits studies conducted under Work Plan 2, Jobs 3 and 4. and food RESULTS During the final segment of 45-01-502-15050 for Work Plan 2, Job 9, 2 reports were published, 2 were accepted for publication, and 2 were submitted for review. Titles, status, and publication outlet for each report is in the report for Project 45-01-502-15050, Work Plan 1, Job 7. Prepared By g~~ Richard M. Bartmann Wildlife Researcher i ��Colorado Division of Wildlife Wildlife Research Report July 1983 81 JOB PROGRESS REPORT State of Colorado --~~------------------ Project No. 45-01-502-15050 Work Plan No. 2 ------------------- Job No. 10 Period Covered: Author: Bi~ Game Investigations - Cervids De~r investigations Mule Deer Winter Habitat Use in. Piceance Basin 7/1/82-6/30/83 R. M. Bartmann Personnel: R. M. Bartmann, A. W. Alldredge, L. H. Carpenter, R. A. Garrott, J. E. Lee, G. C. White. D. M. Collins, ABSTRACT Two manuscripts,one on assessing accuracy of a radio-telemetry system for mule deer and the other an evaluation of winter habitat use by mule deer, are being written by M.S. graduate student John Lee. A study of summer habitat use by mule deer and of the effects of oil shale clisturbance on summer habitat use by mule deer in upper P~rachute Creek drainage was begun by M.S. graduate student David Collins. A paper entitled "Evaluation of vaginal implants for mule deer" co-authored with R. A. Garrott, Los Alamos National Laboratory, was accepted for publication as a Short Communication in The Journal of Wildlife Management. ��83 MULE DEER WINTER HABITAT USE IN PICEANCE BASIN Richard M. Bartmann P. N. OBJECTIVE To administer funding and to supervise a gr~duate student in conduct of a study to assess mule deer winter habitat use in proximity to an oil shale development project using radio-telemetry methods. SEGMENT OBJECTIVE Same as P. N. Objective. RESULTS John Lee, M. S. candidate at CSU, completed analysis of most data collected to assess winter habitat use by mule deer in Piceance Basin. A rough draft of a paper on assessing accuracy of a radio-telemetry for mule deer is being reviewed. A second paper covering winter habitat use by mule deer is still being written. Both papers will be published in professional journals and included as part of the M. S. thesis. A new M. S. graduate project was started by David ColI ins to·evaluate summer habitat use by mule deer in upper Para~hute Creek drainage and to evaluate effects of oil shale-related disturbance on deer habitat use and activity in the same area. A study plan is nearly completed and work was begun to set up equipment on the study area. A paper entitled "Evaluation of vaginal implants for mule deer" was co-authored with R. A. Garrott of the Los Alamos National Laboratory based on cooperative research conducted at the Little Hills Wildlife Area. The paper was accepted for publication as a Short Communication in The Journal of Wildlife Management. Prepared by ~ ~~, R~rd.Bartmann Wildlife Researcher ��Colorado Division of Wildlife Wildlife Research Report July 1983 85 JOB PROGRESS REPORT State of Colorado Project No. 45-01-502-15050 Work Plan No. 2 ------------------11 Job No. Period Covered: Author: Bi~ Game Investigations - Cervids De~r lnvestigations Testing of Deer Census Methodology 7/1/82-6/30/83 R. M. Bartmann Personnel: R. M. Bartmann, D. C. Bowden, R. A. Garrott, D. A. Garrott, M. Sovada, A. E. Acheson. ABSTRACT Preliminary tests were made to evaluate using radio-collared deer to estimate counting bias in the quadrat deer census. This approach was rejected due to the time needed to verify the number of deer on a quadrat and the few deer present on most quadrats. A defecation monitor implant was developed and tested in a domestic sheep. Problems of short transmitting range and spurious artifact signals were identified. A third implant to incorporate changes to alleviate these problems is to be developed. A study plan was prepared detailing objectives and procedures for the first year of a 5-year study to test deer census methodology. ��87 TESTING OF DEER CENSUS METHODOLOGY Richard M. Bartmann P. N. OBJECTIVE See Segment Objectives. SEGMENT OBJECTIVES 1. Select a specific research project which will include the processes of reviewing pertinent literature, developing hypotheses for testing, selecting a specific study area, and writing a detailed study plan for further testing of mule deer census methodology in pinyon-juniper habitat. 2. Develop a fecal monitoring system for mule deer, conduct preliminary tests with tame deer in small pens, and write a detailed study plan to test hypotheses relative to defecation rates of mule deer. RESULTS AND DISCUSSION Quadrat Census In January 1983, I explored the possibility of using radio-collared deer to assess counting bias. Through the cooperative deer study in Piceance Basin with Los Alamos National Laboratory and Colorado State University, about 100 radio-collared deer were available in the south part of the Basin near the C-b oil shale tract. Radio-collars, 5-cm wide on adult females and 2.5-cm wide on female fawns, were white and easily seen from the air. Twelve 0.648-km2 quadrats were delineated in areas where marked deer were thought concentrated. Number of radio-collared deer on each quadrat was determined by aerial tracking from a fixed-wing aircraft and with some help from 2 people on the ground. Immediately after a quadrat was checked for number of radio-collared deer, it was searched from a helicopter. The usual quadrat counting procedure was used and marked animals noted. Several problems were identified that limited practicality of the method. One was length of time required to determine how many radio-collared deer were on a quadrat. Even with help from a ground crew, it took 8-32 minutes to check a quadrat. This prolonged flying over a quadrat may have spooked some deer and partially contributed to a second problem; the low number of marked animals of quadrats. One quadrat had 5 deer, 3 others had 1 deer each, and 6 had 0 deer. On 2 other quadrats, deer near boundaries could not definitely be distinguished as on or off the quadrat. These low numbers occurred in spite of efforts to select quadrats where radio-collared deer were thought concentrated. Thus, relatively little information would probably be gained concerning counting bias for a large investment of time and money. �88 Pellet Group Counts A defecation monitor implant was developed in cooperation with Wyoming Biotelemetry, Inc., Longmont, Colorado. Due to delays in design and manufacturing, an implant was not available for testing unti 1 March 1983. The implant was surgically placed in a domestic sheep (avis aires) but failed 10 days later due to battery corrosion. The sheep was observed for 13.5 hours,while the implant was working. There were copious artifact signals and nearly every movement of the sheep triggered a signal change. A second implant functioned properly but failed afier 40 days due to leakage. Defecation patterns on the strip-chart were distinct and no other activities of the sheep caused readings that would be interpreted as defecations. Thirty-four defecations were observed and accurately recorded during 19 hours of intensive monitoring. The main problem now is the short transmitting distance. received much beyond 200 m. Also, artifact signals which defecation patterns increase rapidly beyond about 20 m. being assembled will incorporate changes to remedy these Signals cannot be could camouflage A third implant problems. Study Plan A study plan was prepared for deer census research to be done in 1983-84. The study duration is scheduled for 5 years, but objectives and procedures will be prepared in I-year segments with annual updates for each succeeding year. This approach is considered necessary due to the "b lu i ld i nq-b lock" characteristics of each study phase. Development of some methods must still be done and information gained from one year1s work will influence the design of studies the next year. Prepared by ---J.._.:fC~.:;....h±!~~ /..t:E.~~::::2!:!.=:::::!+::::===:::::::/ __ Richard~rtmann Wildlife Researcher �Colorado Division of Wildlife Wildlife Research Report July 1983 89 JOB PROGRESS REPORT State of Colorado Project No. Work Plan No. 2 ------------------ Job. No. Personnel: - Cervids Deer Investigations 12 Period Covered: Authors: Big Game Investigations 45-01-502-15050 Determination Mule Deer of Body Composition of 7/1/82-6/30/83 S. C. Torbit and L. H. Carpenter S. C. Torbit, L. H. Carpenter, R. L. Bartmann, A. Wm. Alldredge, S. Bai ly, E. Winicov P. H. Neil, ABSTRACT Body composition was determined for 24 free-ranging, adult, female mule deer inhabiting the Piceance Basin during the 1982-83 winter. Six deer were collected during October, December, February and April. The fat component was largest in December S.D. =11.44%± 2.30%) and smallest in April (x S.D. = 3.66% ± 1.90%). Protein reserves also decreased during winter but much less than fat reserves. Left kidney fat index was found to be positively correlated with total fat reserves. Multivariate analysis is proceeding to test relationships between measurements of body size, (live weight, chest girth, total body length, left hind foo~ length) kidney weights, and size of fat and protein reserves. ex - .. __ ... ~"-.'~" "_ :',: ��91 DETERMINATION OF MULE DEER BODY COMPOSITION Stephen C. Torbit P. N. OBJECTIVES 1. Determine body composition and describe the relationship between body water, body fat and lean body tissue of mature female mule deer inhabiting the Piceance Basin winter range of western Colorado. 2. Compare chemical estimates of total body fat to kidney fat and bone marrow fat to determine validity of these indices as indicators of animal condition. 3. Evaluate variances of measured parameters of deer and if results seem feasible, develop a detailed study plan to investigate winter dynamics of body composition of wintering mule deer. SEGMENT OBJECTIVES 1. Review pertinent literature, develop appropriate hypotheses, and prepare a study plan on the most efficient approach to determine body composition of wild mule deer over winter. 2. Collect from 10 to 20 road-killed mule deer to establish the relationship between body water, body fat, and lean body tissue. METHODS AND MATERIALS Twenty-four adult female mule deer were collected from the Piceance Basin during the 1982-83 winter. Six deer were collected in October, December, February and April. Deer were either trapped in Clover traps (Clover, 1956) and euthanized with a drug overdose or shot. All six deer were shot in October and 1 deer was shot in April (April 6). Shooting was necessary in that deer were not attracted to Clover traps at those times. Immediately after death, deer were weighed to the nearest 0.5 kg and the following measurements taken; total body length, chest girth, and left hind foot length. Animals were wrapped in plastic bags, frozen and transported to Fort Collins, where they remained frozen until processed for chemical analysis. Carcasses were reweighed, thawed and prepared for analysis as described elsewhere (Torbit, 1981). Kidney fat index was determined for the left kidney on all 24 deer and for the right kidney for 18 deer (December, February, and April collections) and bone marrow was removed from the left femur and analyzed for fat content from all deer. Samples from the minced carcass were collected to assay for water, fat, protein and ash (Trobit, 1981). An additional aliquot was taken for determination of total and radioactive potassium (K-40). Fetuses from pregnant does were collected and body composition analyzed separately from the dam. �92 RESULTS AND DISCUSSION All chemical and radiological analyses are not yet complete, however, chemical estimates of the 4 body composition components have been completed (Table 1). Fat was the most variable body component measured, ranging from 15.10 to 1.56 percent of empty body weight .• Highest percentages of body fat occurred in deer collected in December and lowest values were found from deer collected in Apri 1. Body water content: increased during winter as expected since deer were catabol izing fat. However, percent carcass water of Apri 1 samples increased not only due to loss of fat, but also because the amniotic fluid (not measured) recovered from the placenta was mixed with the dam's empty body. Percent protein content of the carcass apparently remained constant across winter (Table 1), however, this energy reserve appeared dynamic when a ratio of protein to ash was calculated (Table 2). Kidney fat indices and femur marrow fat were measured to test the relationship between these often used indices and total fat reserves. All bone marrow samples have not been analyzed, however, a significant relationship was detected between the left kidney fat index (LKFI) and total body fat (Table 3). This relationship (Fig 1) [% Total Body Fat = 6.84 + 4.30 log LKFI, r2 = 0.85J was found to be significant at the 5% probability 1eve 1. Multiple regression procedures are currently being implemented to establish relationships between measurements of skeletal size, (body weight, chest girth, left hind foot length and total body length), kidney ~eights and chemical estimates of fat and protein reserves. These analytical procedures are not yet completed, however, preliminary analyses suggest that other significant relationships will be detected. Reproductive status of the deer was determined beginning with collections in February (Table 3). October collections occurred before breeding began and December collections occurred immediately after breeding and before fetal development. All 12 deer collected in February and April were pregnant with 24 fetuses (2.0 fetuses/doe). Thirteen of the fetuses were male and 11 were female. Age, as determined by tooth replacement and wear, varied from 1 year to 10 years with a mean age of 6.0 ± 2.96 (x ± s). Incisors were taken from each deer and submitted to the Colorado Division of Wildlife Laboratory for analysis. At this writing these data are just being completed and will be presented next segment. \. �93 Table 1. Chemical composition of female mule deer collected from Piceance Basin October, 1982 - Ap rIl, 1983 % % % % Water Fat Protein Ash 1 2 3 4 5 6 63.60 65.57 65.34 61.07 64.06 63.70 10.38 10.01 9.61 11.82 11.74 10.96 19.88 19.88 19.55 20.77 19.74 20.08 6. 15 4.58 5.50 5.51 4.45 4.90 x 52 5 63.89 2.17 1.47 10.75 0.69 0.83 19.98 0.15 0.39 5.18 0.35 0.59 December 1 2 3 4 5 6 64.20 60.15 65.39 55.63 63.67 63.48 9.02 14.06 9.80 15.10 9.91 10.76 20.59 19.64 19.96 21.90 19.91 19.90 6. 15 5.60 4.76 7.24 6.48 5.49 52 5 62.10 10.66 3.27 11.44 5.27 2.30 20.32 0.58 0.76 5.95 0.62 0.79 February 1 2 3 4 5 6 66.13 65.61 69.39 67.33 63.43 67.10 9. 15 10.34 6.32 7.75 11.28 8.38 19.29 18.06 18.44 19.44 19.67 18.80 5.40 5.92 5.86 5.50 5.69 5.74 66.50 3.29 1.81 8.87 2.67 1.64 18.95 0.32 0.57 5.69 0.03 0.18 1 2 3 4 5 6 70.96 75.09 73.48 71 .16 78.10 71.28 5.77 1.56 2. 18 5.47 1.58 5.38 18.06 18.30 19.40 19.02 16.64 19.23 5.21 5.06 4.95 4.36 3.69 5. 12 x 73.18 7.56 2.75 3.66 3.60 1.90 18.44 0.88 0.94 4.73 0.29 0.54 Deer Number October x x 52 5 Apri 1 52 5 �94 Table 2. Ratio of total protein to total ash for female mule deer coalected from the Piceance Basin October, 1982 - April, 1983. Kg Protein/ Kg Prote in/ Deer Deer Kg Ash Kg Ash December October 3.23 4.33 3.55 3.77 4.43 4.10 1 2 3 4 5 6 x = 5 6 x = 3.46 Apri 1 3.57 3.05 3.15 3.53 3.46 3.28 x .. : 3.35 3.50 4.19 3.02 3.08 3.63 4 3.90 February 1 2 3 4 5 6 ' 1 2 3 = 3.34 1 2 3 4 5 6 3.47 3.63 3.92 4.36 4.51 3.75 x = 3.94 �-s __ Age_?_reprodu_c_~_i_v~_~_~~~_L!s_L b09),~ ize J'!leasuremen!s_L chem ica 1_e_~t_i_mate_oL~ody_f9.!_,_~nd_1e f t Tab 1e___3_. and right kidney fat indices for female mule deer collected from the Piceance Basin October, 1982 - April, 1983. Total Left Hind Chest Deer Body No. Foot a/ Number Agefetuses (sex) Weight(kg) Girth(cm) Length(cm) Length(cm) Body Fat(%) RKF I(%) LKFI(%) October 1 2 3 4 5 6 December 1 2 3 4 5 6 Februa ry_ 3 6 9 1 1 2 9 9 9 8 5 1 2 1 8 2 2 5 2 3 9 2 4 8 2 5 7 2 6 3 Apr i1 1 2 8 2 4 3 2 3 7 2 4 4 2 10 5 1 6 8 a/ -Age determined-by E!Right kidney was - b/ - - 56.4 54.4 59.5 46.8 50.8 58.0 88.0 90.0 91.0 87.0 89.0 95.0 149.0 143.0 142.0 130.0 140.0 144.0 46.0 46.0 46.0 45.0 46.0 45.0 10.38 10.01 9.61 11.82 11.74 10.96 - 66.8 64.6 67.2 69.9 57.1 56.7 94.5 91.5 90.5 97.4 90.5_ 87.0 157.0 154.5 155.5 151.0 147.0 139.0 49.0 48.3 47.0 45.0 48.0 48.0 9.02 14.06 9.80 15.10 9.91 10.76 28.57 65.05 57.73 103.00 - 63.54 -34.03 34.55 71.98 55.45 123.40 74.09 55.14 - - - - 87.65 44.03 34.19 78.16 72.61 88.24 (2 (1 (1 (1 (1 (2 F) M, M, M, M, M) F) F) F) F) 73.3 52.9 72.6 69.7 68.8 66.4 98.0 89.5 93.0 91.0 90.0 87.5 155.0 141.0 153.0 147.5 147.5 149.5 47.0 46.0 47.0 47.0 44.8 46.5 9.17 10.34 6.32 7.75 11.28 8.38 41.38 57.32 16.25 31. 47 44.69 61.97 31.58 42.56 14.63 31.58 35.52 51.95 (1 (2 (2 (2 (2 (1 M, 1 F) M, 1 F) F) M) M) F) 57.3 62.5 59.2 60.5 54.9 62.0 87.0 90.5 89.5 88.5 88.0 93.0 148.0 160.0 146.0 153.0 159.0 159.0 45.0 47.0 45.5 48.0 46.0 46.5 5.77 1.56 2.17 5.46 1.58 5.38 24.28 20.21 9.30 19.27 10.98 25.00 22.03 8.98 10.80 23.16 6.13 26.75 1 1 1 1 tooth replacement and wear not collected in October. ~ U1 �96 16 x 14 x 12 10 .--.. IN' 4-1 co LL. >- -c 8 0 a:l co 4-1 0 I- Left Kidney Fat Index(%) Fig 1. Relationship between left kidney fat index(%) and total body fat(%) for 24 female mule deer collected from the Piceance Basin during 1982-83 winter. �97 LITERATURE Clover, M. R. 199-201. 1956. CITED Single gate deer trap. Calif. Fish and Game 42(3): Torbit, S. C. 1981. In vivo estimation of mule deer body composition. Ph.D. Diss., Colorado State Univ., Fort Collins. 98pp. c-I-LL Prepared by _)... ~ ~~~'~·~.~i90~'~;;~~~(_~·_~~/~?:~·~~~ _ Stephen c. Torb it Re sear ch Associate Len H. Carpentef Wildlife Research Leader ��Colorado Division of Wildlife Wildlife Research Report July 1983 99 JOB PROGRESS REPORT State of Colorado Project No. Work Plan No. 3 ------~----------- Job No. 2 Period Covered: Author: Big Game Investigations 45-01-502-15050 - Cervids Elk Investigations Elk Population and Ecological Studies 7/1/82-6/30/83 George D. Bear Personnel: George D.-Bear ABSTRACT All data were gathered and analyses completed for the 4-year population study in the Rocky Mountain National Park area in north central Colorado. Titles, status and publication outlet for the 3 manuscripts summarizing this work are presented in the Progress Report for Project 45-01-502-15050, Work Plan 1, Job 7. ��101 ELK POPULATION AND ECOLOGICAL STUDIES George D. Bear P. N. OBJECTIVES 1. Develop techniques to more accurately population levels. and precisely estimate elk 2. Define natality and mortality 3. Determine seasonal movements, daily activity patterns, and habitat preferences of elk in Rocky Mountain National Park and adjacent seasonal ranges. problems of selected elk populations. SEGMENT OBJECTIVES 1. Attempt to capture and weigh 30 2. Data analysis and preparation elk calves as near birth as possible. of final manuscripts. METHODS AND MATERIALS Methods and materials have been previously described (Bear, 1979). RESULTS AND DISCUSSION Data analysis is completed and manuscripts LITERATURE are being prepared. CITED Bear, G. D. 1979. Elk population and ecology Fed. Aid Prog. ~ep. July Part 2:373-399. Prepa red by ~.t) Geo rgeo:Bar Wildlife Researcher C studies. Colo. Div. Wildl. ��103 Colorado Division of Wildlife Wildl ife Research Report July 1983 JOB PROGRESS REPORT State of Colorado Project No. 45-01-502-15050 Work Plan No. 3 ------~---------- 3 Job No. Period Covered: Author: 7/1/82-6/30/83 Big Game Investigations - Cervids Elk Investigations Evaluation of Factors Influencing Nutritional Status and Population Performance Elk D. L. Baker ABSTRACT Comparative voluntary intake, digestion, and rate of passage were studied for mule deer and elk consuming native diets. Mean voluntary intake (g/kg BW) was greater for elk than mule deer when fed low quality grass diets but less than mule deer for grass-browse combination diets. Apparent digestibility of dry matter and fiber constituents was greater for elk than deer on both forages. Turnover time for mule deer was less than that for elk regardless of diet. This difference was most apparent for the grassbrowse diet where elk retained this forage 25% longer tfian deer. Turnover time for both deer and elk was greater for low'qual ity grass than grassbrowse diets. Different rare earth elements appeared to give similar estimates of rate of passage and turnover time. In vitro rates of dry matter digestion appeared to be related to the proportion of browse in the diet. As browse in the diet increased, initial rates of digestion increased while extent of digestion decreased. In vitro predictions appear to overestimate in vivo estimates for both deer and elk. This may be related to the shorter retention time of residue in the rumen than in the in vitro tube~ '_ .,_ .•.....•.. , - , ..-, .••.., -,--.-.~ . ,. -,-- .. ,,: _;". ,- ", < .. ,."'.:_;·A ..·· - ., ~ ..,,:,, ..•....•. . ��105 EVALUATION ELK NUTRITIONAL OF FACTORS INFLUENCING STATUS AND POPULATfON PERFORMANCE Dan L. Baker P. N. OBJECTIVES 1. To develop and test a system for evaluating to support elk. the potential of habitats 2. To improve the predictive capability of this system by identifying and quantifying variables influencing the range-supply-animal demand model of nutritional carrying capacity. . SEGMENT OBJECTIVES 1. 2. Begin initial investigations of comparative forage intake of deer and 'elk. digestive physiology and a. Train and condition experimental animals to confinement pens, metabolism cages, and weighing apparatus. in isolation b. Conduct preliminary experiments to (1) define probl~ms associated with experimental equipment and procedures, (2) determine ad libitum intake levels of deer and elk consuming forage under controlled conditions, (3) evaluate both in vivo and in vitro rare earth elements as potential markers of particulate flow through the gastrointestinal tract. Estimate comparative digestive efficiency, mean retention time, rumen fill, and fermentation rates of deer and elk fed an array of native forages. 3. Define in vivo/in vitro relationships of native forages for deer and elk. 4. Develop and/or improve ungulate nutrition models for predicting forage intake. ACKNOWLEDGMENTS J. Ritchie, L. Stevens, and P. Neil assisted in various aspects of the study. �106 METHODS AND MATERIALS Experimental methods pertinent to objectives la, lb(l) and lb(2) are described previously (Baker and Hobbs 1981 :50-S1). Evaluation of rare earth elements as particulate markers lb(3) is currently in progress and will be reported in future reports. Detailed methodology and experimental design relevant to objectives 2 and 3 are presented in Appendices A and B respectively. Completion of objective 4 i~ contingent upon results from these experiments. Since these studies were not completed during this segment, nutritional modeling will be addressed in the final report. RESULTS AND DISCUSSION Deer-Elk Comparative Intake and Digestion Pre 1imlnary Studi es Studies: Initial results for deer and elk consuming a chopped grass hay indicated that mean voluntary intake and digestive efficiency was significantly greater for elk relative to mule deer (p < O.OS). Differences in digestibility were most pronounced with regard to digestion of fiber constituents of grass hay. Detailed results of this trial have been previously reported (Baker 1982:49-S2). Voluntary Chemfcal Composition Intake and Comparative Digestion of Native Diets of Diets The array of experimental diets composited for in vivo and in vitro trials provided a gradient of chemical constituents (Table 1). As the proportion of browse in the diet increased, neutral detergent fiber (NDF) , acid detergent fiber (ADF) , cellulose, gross energy and in vitro digestible dry matter (IVDDM) decreased while cell solubles, lignin, and crude protein increased. All 5 diets were used to evaluate in vftro relationships among rates of digestion, volatile fatty acid CVFA) and gas production. Three diets (75% browse:2S% grass (Diet 1), 50% browse:SO% grass (Diet 2), and 100% grass (Diet 3) were fed to deer and elk to examine in vivo digestion parameters. Consumption of the 100% browse diet by deer and elk resulted in either food refusal, depressed intake, and/or minor digestive upsets. Diluting the concentration of browse by 2S% with grass appeared to improve palatability and reduce digestive problems. Voluntary Intake of Grass-Browse Diets Measurements of voluntary intake of browse-grass diets were made on 17 elk and 16 deer out of a possible 18 observations for each species. At least 5 replicate observations were made for each diet for both deer and elk during the study. Summarization of intake data is currently in progress. �107 Preliminary observations of intake for 1 elk and 1 deer consuming Diets 2 and 3 showed elk intake to be greater (25%) than mule deer for Diet 3 but less than deer for Diet 2 (Table 2). This trend was similar whether intake was expressed as a linear function of body weight or expressed as a function of metabolic body weight. Table 1. Chemical composition of native diets used for in vivo and in vitro expe riments (100% dry-matter basis). Diet Browse-Grass 100% 75% 50% 25% 0% - 0% - 25% - 50% - 75% -100% Table 2. NDF Ce 11 solubles 49.3 60.0 65.4 70.2 73.2 50.7 40.0 34.6 29.8 26.8 Dry-matter Average dry-matter intake .'ADF Lignin Cell u lose CP GE 38.1 19.3 17.7 12.3 9.5 4.5 18.8 25.1 28.8 34.3 37. 1 7.5 6.8 6.2 5.0 5.7 4248 4370 4466 4644 4804 112.8 42.0 43.8 41.6 intake for mule deer and elk used in digestion trials. 100% grass Diet 50% grass-50% browse 4151 634 3353 1203 14.5 10.8 10.8 16.0 59.6 30.0 43.8 47.2 g/day: Elk Deer g/day/kg BW: Elk Deer g/day/Kg BWO.75: Elk Deer Apparent Digestion of Grass-Browse Diets Laboratory analyses of feed, feces, and orts for 28 total balance trials 06 elk, 12 deer} were completed durfng thrs segment. Drgestive measurements for elght animals (~ elk, 6 deerl were unavarlable due to feed refus.a1, d iqes.tive upsets, or excess ive we lqht losses. Most feed refusa 1s occurred when deer were forced to consume Dtet 1. Excessive weight loss was most apparent for mule deer. This was due in part to the length of the study (Oct to Jun), low nutritional quality of test diets, and apparent rnability of mule deer to maintain or increase body weight between digestion �109 trials. Deer lost an average of 25% body weight over the study period while elk experienced a 14% loss over the same period. Thus, elk appeared to be better adapted physiologically to the rigors of intensive digestive experiments when forced to consume diets of low quality for extended periods. Prel iminary estimates of comparative digestibility for 1 deer and 1 elk consuming Diet 2 and 3 suggested that elk were more efficient than mule deer in utilizing nutrients in these diets (Table 3J. Elk digested dry matter, NDF, ADF, and cellulose of both diets more efficiently than mule deer. Magnitude of these differences was greater for the grass-browse diet. Table 3. Apparent in vivo and in vitro digestibi1ities Apparent digestibi 1ity 100% grass for elk and deer. Diet 50% grass-50% browse Dry matter Elk Deer 46.8 34.8 43.0 30. 1 Elk Deer 45.8 37.3 34.0 15.6 Elk Deer 42.3 36.4 28.4 6.9 48.8 43.9 58.1 25.8 56.2 41.5 NDF ADF Cellulose Elk Deer In vitro dry matter Rate of Passage of Grass-Browse Diets Measurements of rate of passage and rumen turnover time were determined from neutron activation of sequentially marked feces for 14 elk and 14 deer. At least 4 replicate estimates were made on each diet for each animal species during this study. Marker concentration curves generally followed a first order model of digestive kenetics (~rovum and Williams 1973). Examples of these curves are shown in Figs. 1 and 2. The ascending portion of these curves (K2) is assumed to represent passage of materials through the lower tract whi Ie the descending phases are suggested to describe passage of undigested residue from the rumeno-reticulum. Slope of the descending curve theoretically represents rate of passage of undigested residue and turnover times are obtained by taking the reciprocal of these slopes. �109 60 z ELK 50% Browse-500/0 Grass •• • • • • • • • • • 50 Rate of passage: 2.4S%/hr. Turnover time: 40.7 hrs. 0 I- <C a:: 40 I- Z woE za. 30 a:: 20 00. 0- w ~ a:: <C :2 10 • o o~--~----_.----_.----_.----~----~----~ 20 60 40 80 100 120 140 TIME (hrs.) Fig. 1. Marker-excretion curve for elk consuming a grass-browse diet. DEER . 500/0 Browse-500/0 Grass • 60 z 0 50 i= a:: Iz ui •• • <C Rate of passage: 3.32%/hr. Turnover time: 30.1 hrs. •• -e- 40 oE zo. oS 30 0 a:: •• w ~ a:: <C • • 20 :2 10 0 0 20 • 40 60 80 100 120 TIME (hrs.) Fig. 2. Marker-excretion curve for mule deer consuming a grass-browse diet. 140 �110 Neutron activation analysis has been completed on 4 of 28 animals. Initial results for I elk and I deer consuming Diets 2 and 3 suggest differences in turnover time between species and diets (Table 4). Turnover time for mule deer was less than that for elk regardless pf diet. This difference was most apparent for Diet 2, where elk retained this diet 25% longer than deer. Turnover time for both deer and elk was greater for grass than grass-browse combination. Different rare earth elements used to mark different fractions of the same diet were found to provide similar estimates of rate of passage and turnover time CTable 4). This was particularly interesting for Diet 2 where both browse and grass were observed to have similar patterns of movement. Table 4. Rate of passage and turnover time for mule deer and elk. % Grass Marker Kl (%/h r) Turnover time (hrs) 100 Vb - Grass Ce - Grass 0.0396 0.0323 25.2 30.9 50 Vb - Both Ce - Grass Sm - Browse 0.0245 0.0256 0.0252 40.8 39. 1 39.7 Diet Species % Browse Elk Elk 0 50 ----------------------------------------------------------_._-------------0 Deer Deer 50 100 Vb - Grass Ce - Grass 0.0383 0.0408 26.5 24.5 50 Vb - Both Ce - Grass Sm - Browse 0.0341 0.D333 0.0349 29.3 30.0 28.7 In Vivo-In Vitro Relationships of Native Diets Rate of Digestion Measurements of rates of dry matter and fiber digestion and VFA and gas production were estimated for 5 grass-browse diets during this segment (Table 1). Analysis and interpretation of results of these measurements are currently in progress. Initial estimates of rate of dry-matter digestion for these diets indicate differences in relative rates of digestion (Fig. 3). Rapid initial rates appeared to be related to the proportion of browse in the diet. The all-browse diet showed the highest rate at 3 and 12 hours but the lowest rate at 48 hours (Table 5). This response was also observed for the 75% browse diet but below this concentration there appeared to be little influence of browse on initial rates. In contrast, while diets containing up to 50% grass showed the lowest rates at 3 hours, they had the highest rates at 48, 72, and 96 hours. �lJ] 100 GRASS 0 w ..... 50 so W (!J {75 GRASS 25 BROWSE 0 40 a: w ..... ..... <!: {50 GRASS 50 BROWSE 30 ~ {25 GRASS 75 BROWSE >- a: 0 20 u. 0 100 BROWSE ..... z w o 10 a: w a. 0 036 24 12 36 48 60 72 TIME (hrs) Fig. 3. Table 5. Percent of forage in dry matter digested Rates of in vitro dry-matter digestion Diet % Browse % Grass 3 100 75 50 25 0 0 25 50 75 100 6.87 4.30 2.74 2.90 2.49 (in vitro) over time. for 5 grass-browse Hours 12 3.06 2.74 2.09 1.90 1.95 diets. 48 0.12 0.45 0.71 0.75 0.74 Rapid fermentation of high browse diets is likely related to the relatively high proportion of readily available cell solub1es (Table 1). Once these cell solub1es are fermented, little microbial digestion can occur due to the highly lignified cell wall of browse tissue. Thus, most of the nutritional value of browse is obtained during the first 24 hours of digestion. Fermentation of grass or grass-browse mixes continues over longer periods of time due to a relatively greater proportion of potentially digestible cell wall and its lower lignin content. �11.2 In Vivo-In Vitro Relationships Preliminary comparisons of in vivo and in vitro digestible dry matter coefficients are shown in Table 3. In vivo estimates for 1 deer and 1 elk consuming Diets 2 and 3 were compared to in vitro values derived from a cow fed a grass hay diet. In vitro procedures followed those of Tilley and Terry (1963). In vitro digestible dry matter for Diet 3 overestimated deer and elk in vivo values by 21 and 10 units of digestibility, respectively. For Diet 2 in vitro overestimated deer in vivo values by 10 units and was similar to elk in vivo digestion coefficients. Similar descrepencies between in vivo-in vitro estimates have been reported for deer and elk consuming low quality native forages (Milchunas 1978, Mould and Robbins 1982). Several reasons for these differences can be hypothesized. One, the difference in physical form of the diet fed in vivo and that used in vitro may contribute to this difference. Second, and most probable is the length of time the substrate remains in the rumen compared to the in vitro system. Food residues are subjected to at least 48 hours of digestion in vitro while rumen turnover for deer and elk is much shorter. Thus, opportunity for increased extent of digestion is greater for in vitro estimates. LITERATURE C ITED Baker, D. L. 1982. Elk investlgations--evaluation of factors influencing elk nutritional status and population performance. Colo. Div. Wildl. Wi ldl. Res. Rep. July:47-52. , and N. T. --- influencing Hobbs. 1981. Elk investigations--evaluation of factors elk nutritional status and population performance. Colo. Div. Wild1. \..Jild1. Res. Rep. July, Part 1 :145-154. Grovum, W. L., and V. J. Williams. 1973. Rate of passage of digesta in sheep. 4. Passage of marker through the alimentary tract and the biological relevance of rate constants derived ffom the changes in concentration of marker in feces. Br. J. Nutr. 30:313-329. Milchunas, D. G., M. I. Dyer, O. C. Wallmo, and D. E. Johnson. 1978. In vivo/in vitro relationships of Colorado mule deer forages. Colo. Dfv~Wfldl. Spec. Rep. No. 43. 43pp. Mould, E. D., and C. T. Robbins. 1982. Digestive capabilities in elk compared to white-tailed deer. J. Wildl. Manage. 46:22-29. Tilley, J. M. A., and R. A._Terry. 1963. A two-stage technique for the in vitro digestion of forage crops. J. Brit. Grassl. Soc. 18:104-111. Prepared by ~ r;f &L ~D-a-n~L-.~B-a7k-e~r---------Wildlife Researcher C �APPENDIX A STUDY PLAN COMPARATIVE DIGESTIVE OF MULE DEER AND by D. L. Baker October 1981 PHYSIOLOGY ELK 113 �114 PROGRAM NARRATIVE for Research and Survey State: Colorado Project Title: I. Study Title: Project No. W-126-R Elk Investigations 3 --~-- Evaluation of Factors Influencing Elk and Mule Deer Nutritional Status and Population Performance 1. A. Work Plan Forage Consumption Utilization and Nutrient Job No. 3 NEED Efficient management of ungulates and the habitats they occupy depends on understanding the influence of forage supplies on animal performance and, conversely, the effects of large herbivores on the productivity of their food plants. Despite this dependence, the process controlling the capacity of native rangelands to support wild and domestic herbivores remain> vaguely described. In particular, maximizing the productivity of ungulates requires knowledge of factors which influence individual animal condition and range nutrient supply. Such factors include (1) quantity and quality of selected forages, (2) seasonal nutritional requirements, and (3) nutrient intake and utilization. Realistic allocation of seasonal food supplies is contingent on quantification of these variables for each ungulate species influencing the forageanimal grazing system. Knowledge of these nutritional parameters for wild ungulates in general and elk (Cervus elaphus nelsoni) and mule deer (Odocoileus hemionus hemionus) in particular is limited. Information pertinent to factor (1) is relatively abundant. Seasonal food habits for mule deer and elk - have been studied extensively (reviewed by Kufeld 1972, reviewed by Crouch 1981, reviewed by Urness 1981, reviewed by Wallmo and Regelin 198D. Although measurements of relative abundance and nutritional quality of selected forages are more rare (Regelin et al. 1974, Wallmo et al. 1977, Hobbs 1979) methods for estimating these parameters are widely available (Harris et al. 1952, Tilley and Terry 1963, Van Soest 1964, Harris et al. 1967). Chemical methods have been developed to measure the quantity of available nutrients in forages and effectively predict the value of those forages to the animal. Several in vitro techniques have been shown to be useful for evaluating big game forages (Tilley and Terry 1963, Van Soest 1967, Pearson 1970); a limited number of direct comparisons for deer and elk suggest a strong relationship between in vitro rumen digestion and in vivo digestibility (Robbins et al. 1975, Ruggiero and ~fuelan 1976, Milchunas et al. 1977, Urness et al. 1977, Mould and Robbins 1981). �115 Although description of factor (2) is incomplete for mule deer and elk, a considerable amount of basic metabolic research for closely related species has accumulated (Maloiy et al. 1968, Moen 1973:333-364, Gates and Hudson 1978, Mautz 1978, Simpson et al. 1978, Robbins et al. 1979, Kautz et al. 1981). Most poorly understood of these 3 factors is food consumption and nutrient utilization of native forages. Productivity of grazing ruminants is largely dependent on the amount of a given forage that they eat and efficiency of forage digestion and metabolism. It is anomalous that the process of voluntary forage intake (VFI), which profoundly influences animal performance, has received scant scientific study. Nutrient Utilization and Rate of Passage Few experiments have compared the ability of wild ruminants to digest forage diets. Despite this paucity of information considerable theoretical evidence suggests that patterns of nutrient utilization are predictable. Hoffman (1968, 1973) suggested that wild ruminants can be classified relative to their feeding strategies and digestive morphology as roughage feeders (eating mostly grass), transition forms (eating grasses and browse) and concentrate selections (eating browse tips, forbs and succulent grasses). An important influence on the quantity and quality of forage a ruminant eats is its need for energy. Maintenance energy requirements of a mature, non-productive ruminant at rest are related to a logarithmic function of its body weight (in kilograms) raised to the 0.75 power (Kleiber 1975). Consequently, a small ruminant must consume more digestible energy per unit of body weight than a large one (Janis 1976, Kay et al. 1980). This can be achieved by consuming more forage or selecting highly digestible forages. If rumen fermentation is to supply a significant portion of nutrient requirements for these animals, then a relatively rapid rate of passage is necessary. By selecting diets containing little fiber and high cell solubles, only small residues of undigestible fiber remain in the rumen; consequently intake is unimpeded. In contrast, roughage and transitional feeders such as elk (Church and Hines 1978, Kay et al. 1980: Table 1) are characterized by capacious rumens with well developed pilliars which increase retehtion time and maximize cellulosis. These characteristics allow large ruminants to meet nutritional requirements from relatively poor quality forages by thoroughly digesting forage fiber. A conceptual model illustrating theoretical digestion kinetics for a large ruminant (elk) and a small ruminant (deer) is shown in Figure 1 (Huston 1978). This model illustrates digestion rates that are influenced by retention time of digesta particles in the rumino-reticulum. Campling (1964) hypothesized that since most ~oughage digestion occurs in the rumino-reticulum, there should exist a close relationship between retention time, extent of digestion and voluntary intake. If deer are assumed to have a retention time of RT2 that is less than elk (RT1), then deer would have an increased passage of undigestible particles and produce a smaller proportion of end products per unit of DM.passed (R2). �116 /\ , 0 " 10 0 .~ leo/ I , 0 . RRO -.-. --- .••.....•. ~ RS ./ 0 ! ! \ Figure 1. Conceptual model of different digestion rates associ.ated with feed digestion in the ruminant and the influence of rumen volume. Rl = feed intake, RZ = ruminal digestion, R3 = passage of undigested particles, R4 = lower tract digestion, RS = fecal excretion, RRO = ruminoreticular oriface, RTI = elk, RTZ = deer. Retention times related to different rumen volumes. �117 Since the more fermentable cell constituents would be fermented first, R3 would be increased more than R2 would be decreased. This would increase Rl, voluntary feed intake. Digestion in the lower tract (R4) and fecal excretion (RS) would be in close relationship to R3. Thus, for an animal with a small rumen, food-particles would be expected to pass through the rumen more quickly and food consumption would be greater and more frequent. As a result, rate of voluntary intake would be relatively greater, and percent digestibility less, for a deer compared with elk. ' I Given these assumptions, one would predict elk to be better adapted to digest fibrous diets than mule deer. Unfortunately, experimental evidence to support this hypothesis is limited. Baker and Hobbs (1981) reported that mule deer digested cellulose in native grass hay 18% less efficiently than elk. Mould and Robbins (1981) found elk more efficient digesters of cell walls than white-tailed deer. Comparisons of fiber digestion between deer and published reports for domestic ruminants also suggest that deer cannot effectively utilize un1ignified fiber (Short 1963, Short et al. 1965). These observations on digestive physiology are consistent with findings of diet selection studies. Elk have been observed to consume diets higher in unlignified cell wall and lower in cell solubles than either mule deer or bighorn sheep; however, in contrast to predictions of ecological theory, mule deer were less selective and consumed diets consistently less digestible than those eaten by elk and bighorn sheep (N. T. Hobbs, D. L. Baker, R. B. Gill unpublished dat~). Deer fawns have also been reported to choose diets which were less digestible than diets of larger bodied mature domestic sheep and goats (Bryant et al. 1980). Deer diets are often digested poorly because they are dominated by highly lignified leaves and stems of shrubs (reviewed by Wallmo and Regelin 1981). These findings suggest measurements of digestibility alone may not be an accurate predictor of forage quality without considering intake. Furthermore, the nutrient contribution of less digestible forages is probably related to attributes of the animal eating the forage as well as characteristics of the forage itself. For mule deer consuming predominantly browse diets, turnover time of ingesta in rumen is an important determinant of nutrient intake. Voluntary intake by ruminants consuming most natural diets is controlled by the rate of turnover of ingesta in the rumen. Rumen turnover depends on the rate of digestion and rate of excretion of ingesta. However, because browse diets consumed by mule deer are digested poorly, excretion of ingesta is probably the dominant influence on rumen turn~ over, and, hence, intake. Rate of excretion of forage is a function of the rate of breakdown of forage particles (Mertens 1973). These rates are dependent because the rumino-reticular oriface (RRO) acts as a filter allowing only particles of a given size to pass (Fig. 1) (Balch and Campling 1962, 1965, Hungate 1966, Van Soest 1966, Mertens and Ely 1979). Voluntary intake by mule deer, then is strongly influenced by rate of passage as mediated by the rapidity of particle size reduction. �118 The chemical and physical structure of lignin and its relationship to rate of passage may be particularly important for understanding deer digestive physiology and diet selection. Several lines of evidence suggest lignin makes plant cell walls brittle which increases their tendency to shatter during mastication-and rumination (Van Soest 1966, Smith 1968, Mertens 1973, Milchunas et al. 1978:28). Furthermore, these shattered particles may be of size and shape more suitable for passing out of the rumen than unlignified particles, typical of grasses (Troelson and Campbell 1968, 'Mertens 1973:28). Consequently the relatively greater lignin concentration of browse would theoretically allow these plants to be passed from the rumen more quickly. A limited number of feeding experiments suggest this may be a plausible hypothesis for deer consuming browse diets (Nagy et al. 1974, Milchunas et al. 1978, Kay et al. 1980). a The overall effect of consuming lignified forage may be a reduction in volume of ingesta in the rumen resulting from the breakdown of ligni~ fied plant cells to a small efficient size, and consequent acceleration of rates of passage. This reduced volume would allow increased voluntary intake as a result of diminished fill limitation. These relationships could explain why mule deer consuming predominantly browse diets high in lignin and low in digestibility may depend more on rapid excretion than rapid fermentations. Rapid excretion would decrease efficiency of fiber digestion. However, this inefficiency may be more than compensated by elevated voluntary intake of high1y_ digestible cell solub1es. Browse diets, high in lignin and cell solubles and lo~ in holocellulose, may be nutritionally important to deer'in winter. Studies of domestic ruminants have reported an increase in rate of passage with a decrease in ambient temperature (reviewed by Young 1981). This increase results in decreased rumen fermentation of fiber and an increase in soluble constituents reaching the lower tract. If liquid flow rates were increased beyond that of division rate of microorganims, microbes could be washed out of the rumen and their numbers reduced. Theoretically, more dietary protein would then escape degradation in the rumen and thus increase the amount digested in the small intestine (Christiansen et al. 1964, Hemsley 1975). Yield of microbial protein in the rumen is a function of available energy to the animal and the outflow rate of rumen contents (Bergen et a1. 1978). Therefore, as rate of rumina 1 outflow increases, extent of organic matter digestion is decreased but efficiency of utilization of energy (ATP) by microorganisms is increased. Thus, there is a higher protein yield per rate of available ATP. Conversely, if ruminal outflow is decreased, extent of organic matter digestion is increased, but efficiency of energy utilization by microorganisms for growth is decreased (Bergen and Yokoyama 1977). Thus, an increase in fluid turnover rate would benefit deer in two ways; by increasing the abomasal supply of bacterial protein and by increasing the amount of feed protein escaping fermentation. This could provide more efficient use of protein for animals consuming low protein diets in winter. �119 Nutrient Intake Voluntary intake of low density forages by domestic ruminants has been clearly demonstrated to be a function of rumen fill and transit time (Campling and Balch 1961, Weston 1966, Egan 1972). The bulk limitation theory suggests animals eat to a constant level of dry matter in the rumen (Freer and Campling 1963, Ulyatt et al. 1967). Thus, rumen capacity can limit intake before energy requirements are met. For diets not bulk limiting chemostatic regulation of intake occurs Conrad et al. 1964, Montgomery and Baumgardt 1965, Baumgardt 1970). For deer, bulk limitation appears to occur above a digestible energy concentration of 2.7 kcal/g DM (approximately 50% digestible dry matter) (Ammann et al. 1973). Therefore, gut capacity or rumen volume has important implications for animals consuming native forages. The source of material which contributes most to rumen fill is the hydrated fibrous fraction of the diet. Cell solubles contribute little to total volume (Mertens 1973:19). Thus, digestion kinetics of the fibrous fraction of forages is an important mechanism influencing voluntary intake. Van Soest (1966) proposed a mechanism of cell wall digestion which suggests digestion of fiber cells is accomplished by microbes "eating holes" in the cells leaving the incompletely digested cell walls to occupy volume and limit intake. Alternately, if, microbial digestion, rumination, and mastication completely shatter partially digested cell walls, the consequent decrease in their volume would theoretically allow increased voluntary intake. This alternate hypothesis is more plausible for wild ruminants consuming highly lignified diets, while the former hypothesis relates better to ruminants consuming diets high in cellulose. While rumen fill may be constant, some limited evidence suggests it can be altered (Fell et al. 1964, Tulloh 1966) •. Forage density can alter dry matter intake of diets at given levels of digestibility. High density feeds occupy less rumina 1 volume per unit weight. Tulloh (1966) reported volume of digesta contents were affected by season and reproductive state of the animal. Red deer were observed to increase intake of heather between January and April by 70%. However, no changes in. digestibility were observed with only small increases in rumen marker retention time (Miln et al. 1978). These investigators hypothesized that compensatory enlargement of the digestive tract had occurred. These observations suggest that rumen volume and related kinetics of digestion may also be influenced by seasonal requirements for nutrients. Seasonal cycles in voluntary intake and body weight of wild ruminants are well documented (reviewed by Moen 1973). Changes in metabolic rates and endocrine function are associated with these annual physiological alterations. Food intake has been shown to decrease in whitetailed deer (French et al. 1956, Ozoga and Verme 1970, McEwan 1975) and elk (Miln et al. 1978, Westra and Hudson 1981) during late fall, then increase to higher levels in spring and summer. Daily activity and energy expenditure have been reported to be less in winter than ' .. ,:. •.....- .. �120 any other season of the year (Holter et al. 1975, Simpson et al. 1978). Serum thyroxine (T4) and 3,5,3- triidothyroxine (T3) have also been shown to be related to lowered metabolic rate (Seal et al. 1972). These physiological traits have been suggested to be advantageous to wild ruminants consuming forages of low quality in winter. Little research has been done relating these physiological changes to seasonal rate of passage and apparent digestibilities of nutrients. Elk calves fed a pelleted concentrate ration reduced their intake from October to December and showed no increase by June (Westra and Hudson 1981). Apparent digestibilities of dry matter, energy and protein increased during winter while mean retention time decreased. Unexplicably, digestion of acid detergent fiber was depressed during the same time period. Other studies of red deer contradict these results. Voluntary intake of grass and browse diets increased 65-70% between January and April, however, digestibility and mean retention time of both diets did not decrease with seasonal increases in forage intake (Miln et al. 1978). It appears from these observations that many variables may influence voluntary forage intake by wild ruminants. Research on domestic ungulates suggests different effects of cold exposure on digestion. Dry matter digestibility in sheep was reduced by 0.18 percentage units per degree (C) drop in temperature (Chris~pherson 1976). This decrease was associated with an increase in reticulum motility and a decrease in mean retention time. Subsequent studies found a strong relationship between T3 and T4 levels and mean retention time of digesta and reticular motility (Westra and Christopherson 1976). These findings suggest that domestic livestock respond to cold exposure by decreasing rumen fermentation and increasing proportion of dietary nutrients which escaped microbial degradation and reach the hind gut unchanged. When this occurs, higher intake rates would occur at the expense of lowered fiber digestion. However, similar to the reason above, an increase in total supply of nutrients to the animal which results from increased intake could offset this inefficiency. Such compensation could be important for winter survival. In conclusion? relationships between rate of passage and nutrient intake and utilization are complex and poorly understood, especially for wild ruminants (Mertens and Ely 1979, Robles et al. 1980). The few extant studies of food passage rates for wild ruminants are characterized by small sample sizes and large variation between animals (Mautz and Petrides 1971, Milchunas et al. 1978). Such limitations preclude useful comparisons and predictions. More research is needed to explain the seemingly paradoxical occurrence of poorly digested foods in deer diets. In addition, further studies are needed to elucidate the effect of season and physiological condition on food passage 'rates and digestibility for large and small ruminants. Knowledge of these nutritional parameters is essential for evaluating ungulate diet quality, diet selection, and for resolute predictions of voluntary intake (Smith et al. 1969, Short et al. 1974, Milchunas et al. 1978:35). "" .. :', �121 Particulate and Solute Harkers A major problem confounding efforts to understand nutrient utilization and voluntary intake is lack of a reliable marker to calculate flows of digesta and related constituents. Before a substance qualifies as an effective nutritional marker, it should (1) be inert and produce no toxic physiological or psychological 'effects, (2) be neither absorbed or metabolized within the gastrointestinal tract (GI), (3) have minimal bulk, (4) mix and remain uniformly dLs t r'Lbut ed in the digesta, and react similarly to the passage of liquid and solid components of the digesta, (5) have no influence on GI functions, (6) have no effect on microflora, and (7) have chemical properties readily discernable throughout the GI tract, which al.l.owprecise quantitative measurements (Kotb and Luckey 1972). The types of particulate markers currently used in ruminant nutrition studies and the advantages and disadvantages of each are reviewed by Kotb and Luckey (1972), ~acRae (1974), Ellis et al. (1979), Faichney (1980). Recently, considerable interest has been directed toward use of "rare earth" elements (atomic numbers 57 through 71) for marking indigestible feed residues. At very low concentrations (below 10-12 M) rare earth elements exhibit radio-colloidial behavior, i.e., strong absorptive properties (Kyker 1962). Chemical and biological characteristics of these elements are described by Kyker (1962). Rare earth elements appear useful as particulate markers for rate of passage and digestibility studies. These markers have been reported indigestible and to bind strongly to particulate matter. Forages appear ~articularly well suited for use with rare earth markers since the element is apparently adsorbed onto the outside of the cuticle or lignified plant cell wall, which undergo relatively little digestion (Hartnell and Satter 1979a). Preliminary investigations have shown radioactive cerium (Huston and Ellis 1968), dysprosium (Ellis 1968, Young et al. 1978) samarium, cerium and lanthanum (Hartnell and Satter 1979a) to remain bound to feedstuff and digesta particles. However, some migration of the marker from one marked particle to another unmarked particle has been reported for highly soluble feeds; Hartnell and Satter (1979a) showed that approximately 10% of the rare earth marker migrated from particulate fraction originally labelled to other particles. Movement of marker from easily solubilized fractions may be either readsorbed onto other digesta solids or may form insoluble hydroxides which would move with the liquid or particulate phase of the ingesta (Kyker 1962, Ellis 1968). Hartnell and Satter (1979b) found marked hay particles contained 15 times the concentration of marker as marked grain particles after 24 hrs of in vitro incubation. It is unknown whe t he r forages high in cell solubles such as woody shrubs would show a similar response. These same forages, however, contain high proportions of fiber which would likely contribute to higher association constants and binding capacities (Teeter et al. 1979). The surface area available for readsorption would likely determine the ultimate fate of released marked particles. The utility of rare earth markers for measurements of rumen fill, retention time and ruminal turnover rates of ingesta has been �demonstrated (Hartnell and Satter 1979b). This study reported that for cattle there was no significant difference in total mean retention time estimated from rare earth marker or stained particle technique. Furthermore, grab sampling of feces compared to total fecal collections resulted in similar estimates of liquid and particulate turnover rates in the rumino-reticulum. In conclusion, application of inert rare earth elements for particulate markers in digestion studies offers the following advantages over other particulate markers (Ellis 1968): 1. Although there is considerable disagreement as to which method of marking forages is most appropriate, all methods are relatively simple. 2. They are non-radioactive 3. Multiple markers can be administered simultaneously for measuring intakes, rates of flow and rates of digestion of different components of a diet. 4. Total fecal collections may not be necessary to estimate rumen fill and turnover rates. As a result more animals can be used in digestion experiments. 5. They offer the potential of being used as external markers in pasture or range studies for qualitative and quantitative measurements of consumption. 6. Precision of estimates of mean retention time and digestion kinetics obtained with rare earth elements appear to be as good or better than for other available particulate markers. and relatively easy to analyze. In contrast, rare earth elements have unique problems common to all markers. These problems include: and share dilemmas 1. Lack of thorough in vitro and in vivo evaluation. More research needed to. determine if rare earth elements meet all established criteria for particulate markers. 2. The optimum method of marking has not been established. 3. More studies are needed to evaluate marker movement between solid and liquid phases. 4. Analysis is relatively feeds differing in chemical .- ,,', composition among particles expensive. Despite these short comings rare earth markers are currently method for studying a variety of digestive processes. Water soluble markers volume in the rumen. is are primarily These markers the best used to study water balance and follow the flow of solutes derived •..... ,.. ' �123 from either the diet or microbial metabolism. Several types of water soluble markers have been used in digestion studies with varying degrees of success. Polyethylene glycol (PEG) and chromium chelate of ethylenediamine tetracetic acid (Cr-EDTA) are the most common. PEG was found satisfactory in estimating approximate fill, rumen volume and weight of rumen contents of dairy cows (Sinka et al. 1970). In contrast, other studies showed PEG inconsistent in marking the liquid phase of rumen contents, and some association with the particulate phase of digesta was reported (Teeter 1981). Downs and MacDonald (1964) suggested use of Cr-complex as a substitute for PEG. Most studies have shown that 51Cr-EDTA is primarily recovered in the feces (85-91%) with generally less than 5% absorbed and excreted in the urine (Hogan 1964, Weston and Hogan 1967, Binnerts et al. 1968). This marker appears to be evenly distributed through almost all of the water of the digesta leaving the reticulum of sheep with no adsorption onto particulate material (Hogan 1964). Non-radioactive, chelating agents other than EDTA have been reported more stable for marking liquid phase of rumen contents (Ellis et al. 1979). Chromium-diethylenetrini-trilopentaacetic acid (Cr-DTPA) contains 2 additional election-pair donor atoms. This property contributes increased stability to any metal. Less absorption is likely to occur due to its larger molecular weight. Moreover, CrDPTA, unlike 5 1Cr-EDTA , does not cause problems associated with disposaL and handling of radioactive tracers in feces and urine. Unfortunately, few substances used as markers in nutrition studies completely satisfy all criteria. Conditions and objectives of the experiment will dictate the material which comes closest to meeting experimental need. In summary, although the objective of any range evaluation system is to assess the potential of various habitats to support and produce grazing ungulates, information necessary to make such predictions for wild ruminants is incomplete. Effective forage allocation and predictions of range carrying capacity must examine complex interactions among diet, animal and environment. Indexes of forage nutritive values are of limited utility without thorough understanding of animal digestive physiology and chemical and physical properties of forages and their utilization. Elucidation determinants of voluntary intake of grazed forages offers predictions extending beyond single, empirical observations and are essential for effective habitat and population management. B. OBJECTIVES The primary objective of this research program will be to quantify seasonal forage intake rates and nutrient utilization of mule deer and elk consuming for~ges of different nutritional composition and to use this information to improve the predictive capability of carrying capacity models. The general approach of this study will involve 2 interrelated phases of research. \~ile each phase will define specific ..•. - �124 objectives and hypotheses to be tested they will also provide a common framework of knowledge from which subsequent experiments can be designed In this way, a more thorough understanding of deer-elk digestive physiology can be gained and methods for estimating forage intake rates developed and evaluated. The two phases of this investigation are: , i Phase I--Comparative digestive physiology and nutrient utilization studies of deer and elk and evaluation of rare earth elements as particulate markers. Experiment 1: Comparative nutrient utilization turnover rates of deer and elk. and digesta Experiment 2: Effects of season-temperature on voluntary food intake and digestive functions of deer and elk. Phase II--Application of techniques developed in Phase I to estimate voluntary forage intake of deer and elk under range conditions. Specific objectives of Phase I; Experiment 1, are: 1. To examine patterns of food passage, nutrient utilization and digestion kinetics of mule deer and elk fed diets differing in chemical composition. 2. To examine inter-relationships between forage chemical constituents, rate of passage parameters and level of consumption for deer and elk. Specific objectives of Phase I; Experiment 2 are: 1. To determine combined effects of season and temperature on digestive function, intake and nutrient utilization by deer and elk. 2. To evaluate potential of rare earth markers rates and voluntary forage intake. Tentative objectives for estimating flow of Phase II are: 1. To estimate conditions. 2. To assess effects deer and elk. 3. To measure forage intake of deer and elk in pastures where food biomass is measured or manipulated prior to the experimental period (i.e., snow, domestic livestock grazing). 4. To begin to validate a simulation model of energy and nitrogen flow for deer and elk with empirical information obtained in the above studies . .' r.· ,,' ...~~:,,.. seasonal intake rates of deer and elk under range of changing forage quality on intake rates of �125 Detailed study plans designed to address written at the completion of Phase I. C. EXPECTED these objectives will be RESULTS AND BENEFITS Knowledge of deer and elk digestive physiology and nutrient utilization will provide resource managers with basic information necessary for identifying habitat requirements and diet selection processes of these species. Documentation of interrelationships between forage chemical constituents and level of consumption can be used by wildlife managers to predict individual animal condition and population performance. D. APPROACH Phase I: 1. Experiment l--Comparative turnover rates nutrient utilization Hypotheses H : Elk will digest test diets more completely a but digestible energy be greater for deer. intake per kilogram than mule deer, of body weight will H: o Digestive efficiency deer and elk. ~: Mule deer and elk will increase rate of passage as lignin concentration of the diet increases. H : Turnover rate of digesta will not be affected tration of the diet. H : Liquid turnover rates from the rumen will increase for both deer and elk as browse concentration in the diet increases. H : Liquid turnover rates will be similar both. deer and elk. o c o Rate of passage deer than elk. H : o 2. and digesta Digesta turnover and elk. and nutrient intake will be similar for from the rumen by lignin concen- for all test diets for for all test diets will be faster for mule rates of test diets will be similar for deer Rationale Elk have relatively large rumens with well developed physical barriers for slowing digesta. Protracted food retention allows elk to maximize rumen fermentation of fibrous diets (Hoffman 1973, Church and Hines 1978). In contrast, mule deer which have smaller rumens, more rapid turnover rates and higher energy requirements per metabolic body size, must increase rates of intake and rapidly ,'.~-..... .; .,,; '. �126 excrete ingesta at the expense of efficient rumen digestion (reviewed by Kay et al. 19.80). If ruminal fill is proportional to body weight, and energy demand is related to metabolic weight, then a smaller ruminant requires a faster rate of passage or must select more easily digested foods to meet energy requirements. Studies of domestic and wild ruminants suggest high concentrations of lignin cause plant cell walls to shatter during mastication and rumination. As a result, lignified forages may be broken down and passed out of the rumen faster than less lignified forages (Van Soest 1966, Smith 1968, Mertens 1973, Nagy et al. 1974, Milchunas et al. 1978). Thus, ruminants consuming highly lignified browse diets may depend on rapid turnover rates rather than rapid digestion. Rapid excretion would reduce the limitation of rumen fill and allow increased nutrient intake (Ammann et al. 1973). Whitetailed deer fawns consuming diets diluted with various levels of oak sawdust partially made up for decreased energy in feed by increasing dry matter intake. This increase in intake occurred as digestibility of the diet decreased to 2.3 kcal DE/g (Richens 1975). Reduction in rumen fill of animals consuming lignified diets is based on the theory that digestion of cellulose weakens cell wall structure contributing to ease of particle size reduction and passage and therefore reducing rumen volume (Mertens 1973:19, Milchunas et al. 1978). Since browse is characterized by higher cell solubles, an increase in this component in the diet should also result in higher liquid turnover rates. Furthermore, fermentation in the large intestine may compensate for reduced digestion in the rumen. 3: Methods a. Test Diets and In Vivo Digestibility Three diets will be fed to experimental 'deer and elk and will represent an array of chemical constituents encountered by wild mule deer and elk grazing in winter. At one extreme will be a 100% grass diet (Treatment 1) represented by smooth brome (Bromus inermus). This grass will be harvested when mature in late summer. At this time, chemical composition should be similar to that of grasses available to deer and elk in winter; cell wall constituents should range from 7080%, acid detergent lignin, 3-5%, crude protein, 4-6%, and dry matter digestibility 45-55%. Smooth brome was chosen because of its frequent occurrence in the winter diets of both mule deer and elk, ease of collecting large quantities and high palatability to both species (Baker and Hobbs 1981). A 100% browse diet (Treatment 3) will offer the opposite extreme in chemical composition. Blueberry (Vaccinum spp.) low growing shrub will be collected for this diet. This species is an important food item for deer and elk in summer (Regelin et al. 1974, Baker and Hobbs 1982). Chemical constituents of blueberry include low cell wa Ll, (40-50%), high lignin (15-20%), high crude protein (10-12%), and low digesti- �127 bility (25-30%). A third test diet (Treatment 2) will be composed of 50% Bromus inermus:50% Vaccinum spp. All forages will be air-dryed and coarse chopped to a uniform size (3 cm) in a hammer mill and stored in burlap bags until fed. Six adult elk and 6 adult mule deer will be used for comparative digestion studies. Mothershead et al. (1972) reported that 5 deer per treatment were sufficient to detect significant differences (~ < 0.05) in in vivo digestion coefficients. All animals to be used in these trials have been previously conditioned to digestion chambers, thus extensive training is not anticipated. Due to the availability of 6 digestion cages, 2 groups of animals, consisting of 3 elk and 3 deer each, will be randomly selected and fed experimental diets during November, January and March. Once assigned to a group, an animal will remain in that group throughout the experiment. A complete balance trial and rate of passage estimate will be made for each animal during the 3 sample periods. For each sample period, 2 animals of each species (1 from each group) will be fed 1 of 3 experimental diets. Thus, by the end of the experiment, all animals will have been offered each ration (Table 1). Each experimental period will be divided into 4 parts (Table 2). Group II animals will enter the d Lge s t Lcn .. cages __ as Group ~I enters the postrial period. Each experimental period will require 31 days to complete. Pretrial: Purposes of pretrial periods are (1) adjustment of animals and rumen microflora to specific test diets (Bryant and Burkey 1953, Church and Petersen 1960), (2) voiding of previous diet, and (3) determination of intake for balance trial. Adaptation of microflora within the rumen has been shown to occur during the first 4 days (Potter and Dehority 1973). During the first 5 days of the pretrial period the test ration will be fed ad libitum until daily intakes can be predicted. Daily rations of test diet will then be adjusted until no orts remain after each l2-hour feeding. Trial: Buildup and pretrial will last 14 days. Length of digestion time has been evaluated by several investigators. Davis et al. (1958) and King et al. (1960) suggest 6 days while Staples and Dinusson (1951) recommend 7 days. Mothershead et al. (1972) advised 10 days, but little statistical difference was noted for 7-day collections. Seven day collections will be used in this study. Feces and urine will be collected daily, subsampled and frozen. Urine will be collected in polyethylene jugs and kept at a pH of approximately 3 by addition of a 10% solution of sulphuric acid. Feed, orts and feces will be freeze-dried, then �128 Table 1. Tentative schedule for experimental and recovery periods. Experimental and recovery period Species Group .. Group II I Diet Recovery Period I October 1-31, 1981 Experimental Period I November 1-30, 1981 Elk A B C Deer A B C D E F 100% grass 50% grass:50% browse 100% br-owse D E F 100% grass 50% grass:50% browse 100% browse D E F 50% grass:50% browse 100% browse 100% grass D E F 50% grass:50% browse 100% browse 100'%grass D E F 100% browse 100% grass 50% grass:50% browse D E F 100% browse 100% grass 50% grass:50% browse Recovery Period II December 1-31, 1981 Experimental Period II January 1-31, 1982 Elk A B C Deer A B C Recovery Period III February 1-28, 1982 Experimental Period III March 1-31, 1982 Elk A B C Deer A B C Recovery Period IV April 1-30, 1982 End Tria1s--Hay 1, 1982 �129 Table 2. Feeding schedule for deer-elk in vivo digestion trials. Percent of diet Test Native grass % ration % Day Locationa .Recovery- 1-21 IP 100 a Buildup 22 IP 75 25 23 IP 75 25 24 IP 50 50 25 IP 50 50 26 IP 25 75 27 IP 25 75 28 IP 0 100 29 IP 0 100 30 IP 0 100 31 IP a 100 32 IP 0 100 33 IP a 100 34 IP 0 100 35 DC 0 100 36 DC 100 37 DC a a 38 DC 0 100 39 DC 0 100 40 DC 0 100 41 . DC a 100 42 DC 0 100 43 DC 100 44 DC a a 45 IP 25 75 46 IP 50 50 47 IP 75 25 48 IP 100 49-60 HP 100 a a Period Pretrial Trial Postrial Intake Group Ad libitum Group II 100 Adjust intake to 90% of ad libitum 100 Ad libitum a DC = digestion cage, IP = isolation pen, HP = holding pen. Group I �1:30 processed through a Wiley mill with a l-mm screen and analyzed for dry matter (DM), organic matter (OM), cell wall constituents (CWC) , acid detergent fiber (ADF) , lignin (LIG), nitrogen (N), and gross energy (GE). Postrial: This period will allow animals to slowly adjust to recovery ration. Between experimental periods, animals will be placed· in isolation pens and fed high quality chopped meadow hay ad libitum for 21 days. Recovery periods are designed to (1) prevent excessive weight loss of animals. (2) allow a baseline intake measurement (concomitant observation) to compare intake of test diets, (3) to partition season and temperature effects on intake and digestion parameters, and (4) to minimize carryover effects from the previous experimental diet. Retention time and ruminal turnover rates will be estimated for each animal and each test diet using 3 stable rare earth elements. YHerbium (Yb) will be used to mark all test diets. This will allow comparisons among all diets using the same marker. Cerium (Ce) will be used to independently mark the grass portion of each meal and Samarium (Sm) the Vaccinium spp. part of each meal. These additional elements will be used to assess possible associate effects of animals consuming diets containing different proportions of brmvse and grass. Chromium diethylenetrianinepentaacetate (Cr-DPTA) will trace the liquid phase of rumen ingesta. These markers were chosen due to their suitability for neutron activation analysis. They are suitable because their decay products do not yield gamma photons of similar energies and consequently can be resolutely separated by counting gamma emissions of different. energy. Cr-DPTA chelate has been reported to have higher stability constants than EDTA, thus, it would be less likely to be replaced by hydrogen in the lower gut (Ellis et al. 1979). Methods for marking meals and dosing animals will follow procedures of Ellis et al. (unpublished data). This method requites the soaking of a portion of a meal in a rare earth marker solution for 24 hrs, rinsing and air. drying prior to feeding. The amount of marker to be administered is calculated according to the following formula: dose (~g/lOO kg weight) = day MAL x F/100 kg body weight x 1/K2- x DX (1/.5) where MAL F k2 D ... ".' minimal analytical level for the marker, dry matter fill for segment body weight), approximate turnover expressed in days, of interest expected ~ (g/lOO kg for the marker days post dose of last collection . .. (Ug/g DM), �131 This calculation assumes (1) a rumen dry matter fill of 2% of body weight, (2) a daily turnover of 1/K2-day, and (3) a serial diluting effect of 0.5. Fecal collections will be made 2 hrs following dosing for the collections will be made every days. The entire fecal sample and stored for later analysis. feed and feces will be measured analysis (Young et ale 1975). just prior to dosing and every first 24 hours. Subsequent 6 hours for the following 6 will be collected, weighed Concentration of marker in using neutron activation Calculation and measurements of ingesta turnover rates and total mean retention times in the gastro-intestinal tract (GIT) will be calculated according to method of Grovum and Williams (1973). Fecal excretion of markers will be used to fit a 2 compartmental model described by the following equation (Brandt and Thacker 1958): y where: s and A constant rations of marker rate constant describing the reticulo-rumen, in feces dry matter, marker kinetics in rate constant pertaining primarily to digeRta kinetics in the caecum and proximal colon, t the period after injection of marker, TT transit time of mar.ker through the omasum, abomasum, and the small and large intestines. Previ~us studies with sheep suggest that these digestion parameters are biologically significant (Grovum and Williams 1973). The advantage of this model is that ruminal turnover rates of ingesta can be calculated based on fecal excretion of markers. 4. Analysis A randomized 3 x 3 Latin square (cross-over) design will be used such that each treatment is applied to the same animals in different periods (Cox 1958). Thus, 2 temporal replicates of a Latin square using 2 sets of animals will be as follows: .".J. �132 Treatment Treatment 2 I I r-l C1l .,-1 II III I 2 3 I Elka Elkb Elkc Deera Deerb Deer c 2 Elkd Elk e Elkf Deerd Deer e Deerf I EI~ Elk c Elk a Deerb Deer c Deer a 2 Elkf Elkd Elk e Deerf Deerd Deer e I Elk Elk EI~ Deer c Deer a Deerb 2 Elk e Elkd Deer e Deerf Deerd ,.. E-4 3 c a Elkf The mathematical model used for this experiment is: Y II klja where + Y klja + j A a + interactions as below + [. klja measured varible, common effect in all observations (true.mean of the population), treatment effect (3 rations) - fixed effect, month effect (3 experimental periods) - fixed effect, period effect (2 trials in each experimental period) fixed effect, A f a klja animal effect (6 animals per treatment per species) random effect, error term. -:. ".:/' .... ,:~',. �133 The analysis of variance table for this experiment Source of variation df Total 17 Between animals is: EMS 5 I [ F-test Time Animals (4) F-test (f" 4 <:r e: 2 + + (f <r A 2 <r e:2 + 6 cr 2 cr e:2 6 Treatment x Time 2 (f" e:2 2 cr e:2 + x Time + + F-test where Treatment x month (Time 1) 2· cr e:2 + Treatment x month (Time 2) 2 a- e:2 + 2 = treatment, 2 = month, T M time, (["7)2 cr A <r e:2 2 + 9 A (1'"3(['"2 Treatment Month ~Month [ 2 30"'" 12 Within animals [ e: 2 1 random variation true error. in animals, (f T 2 2 M 3(f'T 2 O 3<rMO 2 <r™ (!""™2 2 (["02 �134 Phase I: 1. Experiment 2--Effects of season-temperature on voluntary intake and digestive functions of deer and elk Hypothesis H : Voluntary food intake (VFI) for deer and elk is inversely related to mean retention time and apparent digestibility I of nutrients. .a H : Voluntary food intake is independent time and/or apparent digestibility. o 2. food of mean retention Rationale Seasonal cycles in voluntary intake are well documented for wild ruminants (reviewed by Moen 1973). For these ruminants, VFI appears consistently Lowe r in wi.nt er . From studies of dames tic ruminants, intake depends to. a great extent upon mean retentian time af digesta in the rumen (Ulyatt 1970, Oltjen et al. 1971, Tharnton and Minsan 1972,1973). Furthermore, retentian time in the rumen has been shown to. be inversely carrelated with digestible arganic matter intake (Tharntan and Minsan 1972, 1973). Thus, extent af digestion would likely be more complete at restricted levels af intake and associated higher mean retentian times. Far wild ruminants these relatianships have nat been clearly described. Preliminary investigations suggest digestive mechanisms af wild ungulates may not fallow traditional concepts described for domestic species and that other variables, particularly behaviaral effects may influence seasanal forage intake (Miln et al. 1978, Westra and Hudsan 1981). 3. Methads General pracedures far this experiment will follaw the methods described under Phase I, Experiment 1 with the exceptian that anly 1 forage, a low quality meadow hay, will be fed. Digestion trials will be canducted in October, December, February, April and June. Dry matter intake will be manitored daily beginning in September even when animals are nat in the digestian cages. Digestian parameters far all camponents af the test ratian will be calculated. Parameters estimating rate af passage of digesta will be determined using particulate and liquid phase markers. 4. Analysis Specific experimental design and statistical treatment be determined fallowing evaluation of Experiment 1. '- .. '~' ; '"""" .•... :' ... _ of data will �135 Schedule June-September 1981 Train deer and elk for digestion studies October 1981 to March (Experiment 1) 1982 April 1981 to September E. 1982 Digestion experiments elk consuming forages chemical composition Analyze digestion for deer and of different data October 1982 to June 1983 (Experiment 2) Digestion experiments to evaluate the effects of season-temperature on VFI and digestion kinetics of deer and elk July 1983-December Analyze 1983 digestion data July 1984 Completion date--Phase I including 1 or- 2 major publications January Begin Phase II 1985 LOCATION The study will be conducted at the Colorado Division of Wildlife Foothills Research Facility, Colorado State University Foothills Campus, Fort Collins, Colorado. F. RELATED FEDERAL PROJECTS W-144-Rl . ,-" �136 LITERATURE CITED Ammann, A. P., R. L. Cowan, C. L. Mothershead, and B. R. Baumgardt. 1973. Dry matter and energy intake in relation to digestibility in whitetailed deer. J. 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Range Manage. 26(2):106-ll3~ Kyker, G. C. 1962. "Rare earths". Pages 499-528 in C. L. Comar and Felix Bronner, eds. Mineral metabolism. Academic Press, New York. MacRae, J. C. 1974. The use of intestinal markers to measure digestive function in ruminants. Proc. Nutr. Soc. 23:147-154. Maloiy, G. M. 0., R. N. B. Kay, and E. D. Goodall. 1968. Studies on the physiology of digestion and metabolism of the red deer. Symp. Zool. Soc. Lond. 21:101-108. Mautz, W. W. 1978. Nutrition and carrying capacity. Pages 321-348 in J. L. Schmidt and D. L. Gilbert, eds. Big Game of North America. Stackpole, Harrisburg, Pa. 494pp. --- , and G. A. Petrides. deer. ~ .".- 1971. Food passage rate in the white-tailed J. Wildl. Manage. 35:723-731. �140 McEwan, E. H. 1975. The adaptive significance in cervid compared with the other ungulate Zhurnal 54:1221-1232. of the growth patterns species. Zoologichesky Mertens, D. R. 1973. Application of theoretical mathematical models to cell wall digestion of forage intake in ruminants. Ph.D. Thesis. Univ. of Washington. Seattle. l76pp. I --- , and L. O. Ely. 1979. A dynamic model of fiber digestion and passage in the ruminant for evaluating forage quality. J. Anim. Sci. 49:1085-1095. Milchunas, D. G., M. I. Dyer, O. C. Wallmo, and D. E. Johnson. 1978. In vivo/in vitro relationships of Colorado mule deer forages. Colorado Division of Wildlife Special Report No. 43. 43pp. Milne, J. A., J. C. MacRae, A. M. Spence, and S. Wilson. 1978. A comparison of the voluntary intake and digestion of a range of forages at different times of the year by the sheep and the red deer (Cervus elaphus). Br. J. Nutr. 40:347-357. Moen, A. 1913. Wildlife ecology: W. H. Freeman. 458pp. an analytical approach. San Francisco. Montgomery, M. J., and B. R. Baumgardt. 1965. Regulation of food intake in ruminants. 2. Rations varying in energy concentration and physical form. J. Dairy Sci. 48:1623. Mothershead, C. L., R. L. Cowan, and A. P. 'Ammann. 1972. Variations determinations of digestive capacity of the white-tailed deer. J. Wildl. Manage. 36:1052-1060. in Mould, E. D. 1980. Aspects of elk (Cervus canadensis nelsoni) nutrition and associated analytical procedures. Ph.D. Thesis. Washington State Univ. Pullman, Wa. 72pp. --- , and C. T. Robbins. to white-tailed deer. 1982. Digestive capabilities in elk compared J. Wildl. Manage. (In Press). Nagy, J. G., D. C. Baker, J. A. Bailey, D. S. deCalesta, D. E. Reeder, and G. G. Schoonveld. 1974. Middle Park cooperative deer study. Physiology and prevention of deer starvation. Colo. Div. Wildl. Fed. Aid Proj. W-38-R-28 Final Rep. 554pp. Oltjen, R. R., T. S. Rumsey, and P. A. Putman. 1971. All-forage for finishing beef cattle. J. Anim. Sci. 32:327. Ozoga, J. J., and L. J. Verm. 1970. Winter feeding patterns white-tailed deer. J. Wildl. Manage. 34:431-439. diets of penned Pearson, H. A. 1970. Digestibility trials: In vitro techniques. Pages 85-92 in Range and wildlife habitat evaluation, a Research Symposium. USDA Forest Service Misc. Publ. 1147. 220pp. �1~1 Potter, E. L., and B. A. Dehority. 1973. Effects of die.tary changes on rumen inoculation and upon subsequent daily digestibility in the ovine. J. Anim. Sci. 37:1408-1413. Regelin, W. L., O. C. Wallmo, J. G. Nagy, and D. R. Dietz. 1974. Effect of logging on forage values for deer in Colorado. J. Forestry 72:282-285 Richens, V. B. (ed.). 1975. Nutritional requirements and metabolism of white-tailed deer. Pages 65-66 in Report of the Coop. Wildl. Res. Units Prog. 1973-74. FWS USDA. Washington, D.C. Robbins, C. T., P. J. Van Soest, W. W. Mautz, and A. N. Moen. 1975. Feed analysis and digestion with reference to white-tailed deer. J. Wildl. Manage. 39:67-79. --- , Y. Cohen, and B. B. Davitt. 1979. J. Wildl. Manage. 43(2):445-453. Energy expenditure by elk calves. Robles, A. Y., R. L. Belyeau, F. A. Mautz, and M. F. Weiss. 1980. Effects of particle size upon digestible cell wall and rate of in vitro digestion of alfalfa and orchardgrass forages. J. Anim. Sci. 51:384-390. Ruggiero, L. F., and J. B. Whelan. 1976. A comparison of in vitro and in vivo feed digestibility by white-tailed deer. J. Range Manage. 29:82-83. Seal, U. S., L. J. Verm, J. J. Ozoga, and A. W. Erickson. 1972. Nutritional effects on thyroid activity and blood of white-tailed deer. J. Wildl. Manage. 36:1041-1052. Short, H. L. 1963. Rumen fermentations and energy relationships in white-tailed deer. J. Wildl. Manage. 27:184-195. --- , D. E. Medin, and A. E. Anderson. teristics of mule deer. 1965. Ruminoreticular characJ. Mammal. 46:196-199. , R. M. Blair, and C. A. Segelquist. --- digestibility by small ruminants. J. 1974. Fiber composition and Wildl. Manage. 38:197-209. Simpson, A. M., A. J. F. Webster, J. S. Smith, and C. A. Simpson. 1978. The efficiency of utilization of dietary energy for growth in sheep (Ovis ovis) and red deer (Cervus elaphus). Compo Biochem. Physiol. 59:95-~ Sinka, K. N., F. A. Mautz, H. D. Johnson, and L. Hahn. 1970. Use of polyethylene glycol in estimating volume and weight or rumen contents, and digestion in cattle. J. Anim. Sci. 30:467. Smith, L. W. 1968. The influence of particle size and lignification upon the rate of digestion and passage of uniformly labeled carbon-14 plant cell walls in sheep. Ph.D. Thesis. Univ. Maryland, College ·Park. l32pp. �142 _____ , D. R. Waldo, L. A. Moore, E. C. Leffel, and P. J. Van Soest. 1967. Passage of plant cell wall constituents in the sheep. J. Dairy Sci. 50:990. ----- , H. K. Goering, and C. H. Gordon. treatments upon digestibility Waste Manage., Cornell Univ. 1969. Influence of chemical of ruminant feces. Proc. Conf. Animal 88pp. Staples, G. E., and W. Dinusson. 1951. A comparison of the relative accuracy between seven-day and ten-day collection periods in digestion trials. J. Anim. Sci. 10:244-250. Teeter, R. G. 1981. Indigestible markers: in ruminant nutrition. Ph.D. Thesis. Stillwater. ---- , F. N. Owens, and G. W. Horn. Abstracts, American 1979. Ytterbium Soc. Anim. Sci. 4l2pp. Thornton, R. F., and D. J. Minson. tary intake and mean apparent J. Agr. Res. 23:871. ____ methodology and application Oklahoma State Univ., as a ruminal marker. 1972. The relationship between volunretention time in the rumen. Abstr. , and 1973. The relationship between apparent retention time in the rumen, voluntary intake, and apparent digestibility of legume and grass diets in sheep. Aust. J. Agr. Res. 24:889. Tilley, J. M. A., and R. A. Terry. 1963. A two-stage technique for the in vitro digestion of forage crops. J. Brit. Grassl. Soc. 18:104-111. t'Mannetje, L. 1978. Measurement of grassland vegetation and animal production. Commonwealth Agric. Bur. Bulletin No. 52, Hurley, England. 260pp. Troelson, J. E., and J. B. Campbell. 1968. Voluntary consumption of forage by sheep and its relation to the size and shape of particles in the digestive tract. Anim. Prod. 10:289. Tulloh, N. M. 1966. Physical studies of the alimentary tract of grazing cattle. IV. Dimensions of the tract in lactating and non-lactating cows. N.Z. J. Agric. Res. 9:999. Ulyatt, M. J. 1970. Factors contributing to differences in the quality of short-rotation ryegrass, perennial ryegrass and white clover. Proc. 11th Int. Grassl. Congr. 709pp. ---- , K. L. Blaxter, and I. McDonald. 1967. The relations between the apparent digestibility of roughages in the rumen and lower gut of the sheep, the volume of fluid in the rumen and voluntary intake. Anim. Prod. 9:463. Urness, P. J. 1981. Desert and chaparral habitats. Part 1. Food habits and nutrition. Pages 347-385 in O. C. Wallmo, ed. Mule and blacktailed deer of North America. Univ. Neb. Press, Lincoln. �143 ___ , D. D. Smith, and R. K. Watkins. 1977. Comparison of in vivo and in vitro dry matter digestibility of mule deer forages.~ Range Manage. 30(2):119-121. Van Soest, P. J. 1964. chemical procedures Symposium on nutrition and forage pastures: new for evaluating forages. J. Anim. Sci. 23:838-844. 1966. Forage intake in relation to chemical composition and digestibility: some new concepts. Pag~ 24 in Proc. Southern Pasture Crop Improvement Conf. 1967. Development of a comprehensive system of feed analysis its application to forages. J. Anim. Sci. 26:119-128. and Waldo, D. R. 1969. Factors influencing the voluntary intake of forages. Proc. Natl. Conf. Forage Qual. Eval. Utile P. E-l. \-Jallmo,O. C., and W. L. Regelin. 1981. Rocky Mountain and intermountain habitats. Part 1. Food habits and nutrition. Pages 387-421 in O. C. Wallmo, ed. Mule and black-tailed deer of North America. Univ. Neb. Press, Lincoln. --- , L. H. Carpenter, W. L. Regelin, R. B. Gill, and D. L. Baker. Evaluation of deer habitat on a nutritional 30:122-127. basis-. 1977. J. Range Manage. Weston, R. H. 1966. Factors limiting the intake of feed py sheep. 1. The significance of palatability, the capacity of the alimentary tract to handle digesta and the supply of gluconeogenic substrate. Austr. J. Agric. Res. 17:939. , and J. P. Hogan. --- roughages by sheep. 1967. The digestion of chopped and ground I. The movement of digesta through the stomach. Aust. J. Agric. Res. 18:789. Westra, R., and R. J. Christopherson. 1976. Effects of cold on digestibility, retention time of digesta, reticulum motility, and thyroid hormones in sheep. Can. J. Anim. Sci. 56:699-708. , and R. J. Husdon. 1981. Digestive function of wapiti calves. --- J. Hild1. Manage. 45: 148-155 . Young, B. A. 1981. Cold stress as it affects animal production. Sci. 52:154-163. J. Anim. Young, M. C., F. E. Haskin, M. E. Wacks, B. Theurer, and P. R. Ogden. 1975. Neutron activation analysis of dysprosium (a potential inert marker) in hay and feces. J. Anim. Sci. 41:178-184. -----: , B. Theurer, P. R. Ogden, G. W. Nelson, and W. H. Hale. 1978. Dysprosium as an indicator in cattle digestion trials. J. Anim. Sci. 43:1270-1279. �144 APPENDIX B PROGRAM NARRATIVE State: Project I. Colorado Title: Project No. Work Plan Title: Work Plan No. Job Title: Evaluation of Factors Influencing Elk and Mule Deer Nutritional Status and Population Performance In vivo/in vitro relationships deer-elk forages A. of NEED Intensive management of ungulate grazing systems is dependent upon a thorough understanding of the relationships between wild ruminants and their food resources. Productivity of these animals is not only a function of availability and utilization of food plants but also consumption, digestion and conversion of these foods into a form that can be efficiently assimilated into body tissue. Thorough documentation of forage nutrient availability and utilization is not well defined for wild ruminants. This information is an important ingredient for proper allocation of seasonal food supp Li.e s and more thorough ecological understanding of plant-animal interactions. Rate of Passage and Digestion Productivity of grazing ruminants is largely a function of the amount of forage voluntarily consumed and the efficiency of its digestion and metabolism. While voluntary forage intake (VFI) probably influences animal performance to the greatest extent, this variable has received little attention from wildlife scientists. While seasonal voluntary intake estimates of wild ruminants are invaluable to- resource managers knowledge of the factors controlling intake offers predictions extending beyond single observations and are essential for long-term range management decisions. Realistic predictions of intake and animal productivity are dependent upon defining in vivo and in vitro relationships of wild herbivore ,)Lets. From studies of domestic ruminants it has been suggested that voluntary intake is primarily a function of rumen fill and turnover time when consuming a bulk-limiting diet (Campling and Balch 1961, Weston 1966, Ulyatt et 'al. 1967. Forage turnover time is dependent on rapidity of clearance of forage from the digestive tract which allows additional \ intake. Forage can be rem~ved in two w~ys: _(1) through digestion and J absorption (2) through excretion. Excretion or passage of undigested matters appears to be controlled by the reticulo-omasal oriface which filters large forage particles (Troelson and Campbell 1968). Thus, rate of oJ· ••• �145 particle breakdown to an optimal size for passage is of major importance in reduction of rumen fill and ultimately forage intake. The rate of breakdown is largely influenced by means of rumination, mastication and digestion (Van Soest 1965a). Measurements of passage are usually made by giving the animal a pulse dose of an indigestible, recoverable marker. Collections of feces are then made over a period of days (Kotb and Luckey 1972). A limited number of feeding experiments with wild ruminants suggest that animals consuming browse diets high in lignin and low in digestibility may depend significantly upon rapid excretion to increase energy intake (Nagy et al. 1974 and Milchunas et al.1978). The second pathway of forage disappearance from the digestive tract is through digestion and absorption. Rate of digestion is defined by Van Soest (1982) as the quantity of feed digested per unit time and is primarily a function of the physical composition of the diet. Soluable constituents are generally digested rapidly, while structural substrates are fermented more slowly. The importance of rate of digestion on voluntary intake has been emphasized by several investigators (Van Soest 1965, Short et al. 1974, Milchunas et al. 1978). Total digestibility and rate of digestion have been shown to be closely related when making comparisons within forage classes (grass, forbs, shrubs) but for between class comparisons this relationship is less predictable. Different plant parts may have the same digestibility but very different rates of digestion (Short et al. 1974). Because intake of forages having rapid digestion rates is greater than for forages with slower rates, measurements of rates of digestion could provide an important assessment of diet quality (Holechek et al. 1982). Rate of digestion has been shown to be related to nitrogen content and fiber composition of forages. As nitrogen becomes limiting, microbial growth is depressed, resulting in decreased fermentation. This leads ~o an increase in fill effect since removal by digestion is decreased (Egan and Moir 1965). Physical characteristics of the cell wall can also influence rate of digestion. Generally increased lignin content decreases rate of digestion by acting as a physical barrier between cellulose and bacteria (Kamstra et al. 1955). Diet composition of wild ruminants in general and deer in particular have been characterized by forages low in digestibility and high in lignin (reviewed by Wallmo and Regelin 1981, Hobbs et al. 1982). A major component of deer diets contributing to this nutrient quality is highly lignified shrub material. Further examination of winter shrub species suggest that cell soluables constitute the major source of available nutrients of these plants (Hobbs et al. 1981). Since cell soluables are rapidly fermented and converted to volatile fatty acids for energy, it could be hypothesized that for small ruminants, with relatively higher �146 energy requirements, per unit body weight, a winter diet comprised primarily of shrub species would be nutritionally advantageous. This mechanism, together with rapid and efficient excretion of lignified forage particles as suggested by Van Soest (1966), Mertens (1973) and Milchunas et al. (1978:28) could allow both digestible and undigestible fractions of browse plants to be removed more rapidly from the rumen. Rapid turnover resulting from the combined influences of these mechanisms would result in less efficient digestion of fiber but total energy intake could be increased through reduced rumen fill. Few studies have described the kinetics of rate of digestion and passage of browse diets (Milchunas et al. 1978). Furthermore, description of the rate and extent of cell wall digestion of browse diets has not been reported. The information is essential for evaluating ungulate diet quality, diet selection and for predictions of voluntary intake. Rumen Fermentation and Volatile Fatty Acid Production For ruminants, volatile fatty acids (VFA's) are the most important product of rumen fermentation. These acids are absorbed from the rumen and provide the major source of energy to the animal. The relative composition of the individual VFA's present is modified by the quality and quantity of the consumed forage. More than 50 percent of the digestible dry matter intake is ultimately fermented in the rumen and converted to VFA's. This energy accounts for 50-80 percent of the ruminants overall energy requirements (Gray et al. 1976, Annison and Armstrong 1970). Generally, more VFA is produced in the rumen from forages containing high levels of soluable carbohydrate and protein than forages high in fiber and insoluable components (Hogan and Weston 1969). The fermentability of a particular substrate over time is a function of the physical characteristics of the substrate·(fiber composition of the cell wall); the surface and availability of hydrolyzable bonds; and the intrinsic properties of the cell wall carbohydrates (Kirk and Moore 1972). High fiber diets usually result in increased proportion of acetic acid while easily fermentable plants yield a greater relative proportion of propionic acid (Hungate 1966). Increased acetic acid has been shown to be associated with high roughage diets of black wildebeast Van Hoven and Boomker (1981) and buffalo Van Hoven (1980) in Africa. For mule deer, molar percentages of acetic acid were shown to increase in winter and early spring while food plants were dormant and decrease in summer when range forage was succulent (Short et al. 1966). As the ratio of acetate to propionate increases, a greater proportion of food energy is lost as methane (Demeyer and Wolin 1960, Demeyer and Giesecke 1973). Decreased methane production on a given diet has been shown to be related to an increase in digestibility of the ration (Church 1969). Given this relationship, an overall relation between rate of gas production and rate of dry matter disappearance would be likely, since the formation of acetic acid requires the production of carbon dioxide. The relation between fermentation rate at any given time during substrate digestion should the~depend on the composition of the substrate, which in turn, would be reflected in the composition of the gas. �147 Use of VFA concentrations of production rates as a sole parameter of forage quality or available energy is limited. Rumen concentrations at any given time are dependent not only on rate of production but also rate of absorption, rate of passage, saliva dilution, and incorporation of VFA's into rumen microbial bodies (Annison and Lewis 1959). However, when VFA measurements are used in conjunction with other in vitro laboratory techniques, they may provide a more thorough means of assessing habitat quality and animal productivity. B. OBJECTIVES The objectives are to: 1. Examine in vivo/in vitro relationships of deer and elk consuming native forage diets of different chemical composition. 2. Measure rate of passage and rate of digestion and elk fed the above forages. 3. Describe the relationship between total gas production matter disappearance for these forages • 4. Describe the relationship between production rate of hydrogen, methane and carbon dioxide gas and cell wall fermentation of test diets. 5. Measure total and individual VFA production ments to cell wall· fermentation .. 6. C. of this investigation Estimate produced EXPECTED kinetics for deer and dry and' relate these measure- the amount of metabolizable energy available during fermentation of cell wall substrates. from VFA's RESULTS Knowledge of deer and elk digestive physiology, nutrient utilization, and nutritional characteristics of wild ungulate diets will provide resource managers with basic information necessary for understanding habitat requirements and diet selection processes of these species. Furthermore, documentation of the mechanisms influencing voluntary forage intake can be used to improve the predictive capability of carrying capacity models and assessment of forage resources. D. APPROACH 1. Hypotheses - IN VIVO experiment H : Elk will digest test diets more efficiently a ~: than mule deer but digestible energy intake per kilogram body weight raised to the 0.75 power will be greater for deer. Mule deer and elk will increase rate of passage from the rumen as browse concentration of the diet increases. �148 H: e Rate of digestion of dry matter, rate of gas production and rate of VFA production is directly related to the concentration of browse (cell solubles) in the diet. Total dry matter d i ge s t ed; total gas production and total VFA produced is inversely related to the concentration of browse in the diet (these parameters will increase as cell wa~l digestibility increases). 2. Rationale Large ruminants, such as elk have relatively large rumens with well developed physical barriers for slowing digesta. Protracted food retention allows elk to maximize rumen fermentation of fibrous diets (Hoffman 1973, Church and Hines 1978). In contrast, smaller ruminants, such as mule deer, which have smaller rumens, with more rapid turnover rates and higher energy requirements per metabolic body size, must increase rates of intake and rapidly excrete ingesta at the expense of efficient digestion (reviewed by Kay et al. 1980). These differences in digestive characteristics would suggest different patterns of diet selection. Deer in winter consume relatively larger proportions of woody shrub material (reviewed by Wallmo and Regelin 1981, Hobbs et al. 1982) of low digestibility. These plants have been shown to contain large amounts of indigestibile cell wall substrate and relatively large proportions of cell solubles. Efficient utilization of browse plants by deer would require rapid digestion of available cell solubles and rapid excretion of undigestible cell wall material. Rapid digestion would result in a rapid initial rate of fermentation and production of VFA's. However, the highly lignified cell wall would prevent extensive degradation of this substrate, this reducing total fermentation compared to the less lignified cell wall of grasses or forbs. Rapid excretion of lignified residues could be accomplished through efficient particle size breakdown during rumination and mastication (Van Soest 1966, Mertens 1973, Milchunas et al. 1978). These two processes would theoretically result in a combined reduction in rumen fill, thus allowing increased energy intake. 3. Methods a. Methods pertinent to in vivo measurements of rate of passage, voluntary intake and digestibility have been previously described (see program narrative). b. Methods pertaining follows: to objectives 2, 3, 4, 5, and 6 are as (1) Rate of Digestion - the three test diets previously fed to experimental deer and elk during complete balance trials, plus 2 additional composited diets will be used to examine rate of digestion kinetics (see program narrative for more thorough description of diets). At one extreme will be a �149 100% grass diet, while the opposite extreme will be a 100% browse diet. The remaining diets will be made from combinations of these forages. Thus, the 5 diets tested will be 100% grass, 25% grass:75% browse, 50% grass:50% browse, 75% grass:25% browse and 100% browse. These diets will represent an array of chemical constituents encountered by wild mule deer and elk grazing in winter. The in vitro procedure to determine rate of cell wall digestion will follow the methods described by Goering and Van Soest (1970). This procedure determines the in vitro true digestibility of plant cell wall by removing bacterial residues with neutral detergent solution. Rate of cellulose and hemicellulose disappearance will be determined using sequential analysis without sodium sulfite in the neutral detergent boiling (Van Soest and Robertson 1980). Fermentation times will be 0, 3, 6, 12, 18, 24, 36, 48, 72 and 96 hours. Disappearance of fiber constituents for each diet will be plotted against each fermentation time. (2) Rate of Gas Production - gas production (C02, H2, CH3) will be measured continuously for each test diet. Total volume of gas produced and dry matter disappearance will be determined at each of the given fermentation times. Determination of dry matter disappearance will follow procedures of Mellenberger et al. 1970. Gas chromatographic analysis will be used to determine proportions of individual gases produced. (3) Rate of VFA Production - rate of total VFA production and molar percentages of acetic, propionic and butyric acids will be determined for each test diet. Volatile fatty acid measurements will parallel in vitro rate of fiber digestion analysis. Samples will be taken from in vitro flask after given fermentation times and prior to fiber measurements. All VFA analysis will be accomplished using a gas chromatograph. Inoculum source and preparation are the most critical aspects of rate experiments, especially where samples are incubated for short times (Mertens 1973). Lack of adaptation of microbes to the substrates of the in vitro system can produce a delay in digestion rate. Inoculum source appears to be an important varible influencing both rate and extent of digestion (Van Soest 1982). In vitro. cell wall digestion of tropical forages using rumen fluid from animals fed temparate forage was depressed relative to values where donar animals were fed tropical forages (Grant et a1. 1974). These types of interactions are likely more important for free-ranging ruminants where diverse substrates are eaten. Inocula from deer on browse were equal or better able to digest maple cell wall than cattle on grass hay, but deer inocula was less able to digest grass cell wall (Robbins et a1. 1975). Inocula �150 from goats fed a mixture of 65 percent shrub and 35 percent grass~alfalfa mix digested browse dominated diets greater (P < 0.005) than inocula from goats fed alfalfa alone (Sidahmed et al. 1981). Part of this depression in digestibility of browses may be due to secondary plant compounds in shrub species. Thus, microbes taken from animals on shrub diets are likely more adapted to these substrates and depression or lag in rate of digestion would be minimized. Based on these findings, 2 to 3 deer will be fed a diet consisting of 50 percent browse and 50 percent grass for 2 weeks prior to rumen fluid collection. Animals will be sacrificed and the entire rumen contents removed for inoculation of test diets. Inocula preparation will involve placing whole rumen contents in a waring blender and blending for 2 minutes. The blended mass will then be squeezed through 8 layers of cheescloth, mixing with nutrient-buffer solution and pipetting into in.vitro tubes (Mertens 1973). The purpose of this procedure is to dislodge microorganisms that attach to forage fiber and would otherwise be discarded when rumen fluid is strained (Van Soestll pers. corom.) This procedure significantly (P< 0.05) increased digestibility of range forages of deer when compared to strained or layered only techniques of inocula preparation (Milchunas and Baker 1982). 4. Analysis Linear regression using least squares criteria will be used to estimate rate constants for each test diet. An important characteristic of cell wall digestion is the presence of an ultimately indigestible residue that when mathematically treated as a separate entity allows first order kinetics to be applied to the digestible fraction (Waldo et al. 1972). Thus, the amount of neutral detergent residue as a percentage of original sample dry matter will be determined for each fermentation time. The 96-hour residue will be used as the estimate of the indigestible fraction and subtracted from the residue obtained at each of the other fermentation times to provide potentially digestible cell wall. Potentially digestible cell wall data will be analyzed using semilogarithmic plots of residue versus time and by regression using logarithmic transformation (Mertens 1973). The coefficient of regression of log natural of potentially digestible cell wall upon incubation time equals the rate constant for digestion (Smith et al. 1972, Mertens 1973, Robles et al. 1980). �151 Schedule Analysis Period July I-August 1, 1982 July la-August August August IS-August October ]) E. Prepare laborabory for in vitro experiments, a.cquire ~additiona,l equipment, calibrate gas chromatograph condition experimental deer to test diets. 9, 1982 9-August September Activity 13, 1982 Conduct in vitro experiments DOW lab. 30, 1982 I-September Chemical residue, 30, 1982 I-December 1, 1982 P.J. Van Soest. Animal Analyze in analysis of in vitro VFA and gas ana Lys i.s ; data. Prepare manuscript. Science Department, Cornell Univ., Ithaca, N.Y. LOCATION The in vitro study will be conducted at the Colorado Division of Wildlife Research Center. Gas chromatographic analysis will be conducted at the Colorado State University, Foothills Campus, Metabolic Laboratory, Fort Collins, CO. F. RELATED FEDERAL PROJECTS None LITERATURE CITED Annison, E. F., and D. Lewis. 1959. Metabolism in the rumen. Methnen and Co. Ltd., London. Wiley and Sons Inc., New York. l84pp. Annison, E. F., and D. G. Armstrong. 1970. Volatile fatty acid metabolism and energy supply, p 422-437. In: A. T. Phillipson (ed.). Physiology of digestion and metabolism in the ruminant. Orill Press Limited. Newcastle, England. 636pp. Campling, R. C., and C. C. Balch. 1961. Factors affecting the voluntary intake of foods by cows. I. Preliminary observations on the effect on the voluntary intake of hay, of changes in the amount of the reticulo-ruminal contents. Brit. J. Nutr. 15:523. �1,5.2 Church, D. C. 1969. Vol. 1. Oregon. Digestive physiology and nutrition of ruminants. Oregon State University. Church, D. C., and W. H. Hines. 1978.' 'Rumino-reticular characteristics of elk. J. Wildl. Manage. 42:654-659. Egan, A. R., and R. J. Moir. 1965. Nutr~tional status and intake regulation in sheep. I. Effects of duodenaliy infused single doses of casein, urea, and propionate upon voluntary intake of a low protein roughage by sheep. Austr. J. Agric. Res. 16:437-449. Goering, H. K., and P. J. Van Soest. 1970. Forage fiber analysis. Agricultural Research Service. U. S. Dep. Agr. Handbook. No. 379 20pp. Gray, F. V., A. Weller, A. F. Pilgram and G. B. Jones. 1967. Rates of production of volatile fatty acids produced in the rumen of sheep. Aust. J. Agric. Res. 18:625-638. Hobbs, N. T., D. L. Baker, J. E. Ellis, and D. M. Swift. 1981. Composition and quality of winter elk diets in Colorado. J. Wildl. Manage. 45:156-171. Hobbs, N. T., D. L. Baker, and R. B. Gill. 1982. Comparative nutritional ecology of montane ungulates during winter. J. Wildl. Manage. (In press) Hoffman, R. R. 1973. The ruminant stomach. Stomach structure and feeding habits of East African game ruminants. E. Afr. Monograph in BioI. 2:38-45. Holechek, J. L., M. Vavra, and R. D. Pieper. 1982. Methods for determ1n1ng the nutritiv.e quality of range ruminant diets: A review. J. Animal Sci. 54:363-376. Hogan ; J. P. and R. H. Weston. 1969. The digestion of pasture plants by sheep. III. The digestion of forage oats varying in maturity and in the content of protein and soluable carbohydrate. Austr. J. Agric. R~s. 20:347-363. Hungate, R. E. 1966. New York. 533pp. The rumen and its microbes. Academic Press, Kamstra, L. D., A. L. Maxon, and O. G. Bentley. 1955. Effect of lignification in plants on digestion of plant cellulose in vitro. J. Anim. Sci. 14:1238. Kay, R. N. B., W. V. Englehardt, and R. G. White. 1980. The digestive physiology of wild ruminants. Pages 743-761 in Y. Ruckbusch and P. Thivend. eds. Digestive physiology and metabolism in ruminants .AVI, Westport, Conn. �153 Kirk, T. K. and W. E. Moore. (1972). Removing lignin from wood with white-rot fungi and digestibility of resulting wood. Wood Fiber 4:72-79. Kotb, A. R., and T. D. Lukey. and Rev. 42:28-40. 1972. Markers in nutrition. Nutr. Abstr. Mellenberger, R. W., L. D. Satter, M. A. Millett and A. J. Baker. 1970. An in vitro technique for estimating digestibility of treated and untreated wood. J. Anim. Sci. 30:1005-1011. Mertens, D. R. 1973. Application of theoretical mathematical models to cell wall digestion and forage intake in ruminants. Ph. D. Thesis. Cornell Univ., Ithaca. l87pp. Milchunas, D. G., M. 1. Dyer, O. C. Wallmo, and D. E. Johnson. 1978. In vivo/in vitro relationships of Colorado mule deer forages. Colorado-Uivision of Wildlife Special Report No. 43. 43pp. , and D. L. Baker. 1982. In vitro digestion--Sources of within-and between trial variability. J. Range Manage. 35:199-204. ----- Moen, A. 1973. Wildlife ecology. W. H. Freeman and Co., San Francisco. Nagy, J. G., D. L. Baker, J. A. Bailey, D. S. deCalesta, D. E. Reeder, and G. G. Schoonveld. 1974. Middle Park cooperative deer study. Physiology and prevention of deer starvation. Colo. Div. Wild1. Fed. Aid Proj. W-38-R-28. Final Rep. 554pp. Robbins, C. T., P. J. Van Soest, W. W. Mautz, and A. N. Moen. 1975. Feed analysis and digestion with reference to white-tailed deer. J. Wildl. Manage. 39;67-79. Robles, A. Y., R. L. Belyea, F. A. Martz, and M. F. Weiss. 1980. Effect of particle size upon digestible cell wall and late of in vitro digestion of alfalfa and orchard grass forages. J. Anim. Sci. 51:783-790. Short, H. L. 1963. Rumer{ fermentations and energy relationships in white-tailed deer. J. Wildl. Manage. 27:184-195. ;D. E. Medin, and A. E. Anderson. 1966. Seasonal variations in volatile fatty acids in the rumen of mule deer. J. Wildl. Manage 30:466-470. _'J_· 9 R. M. Blair, and C. A. Segelquist. 1974. Fiber composition and forage digestibility by small ruminants. J. Wildl. Manage. 38:197. Sidahmed, A. E., J. G. Morris, L. J. Koong, and S. R. Radosevich. 1981. Contribution of mixtures of three chaparral shrubs to the protein and energy requirements of Spanish goats. J. Anim. Sci. 53:1391-1400. �154 Smith, L. W., H. K. Goering, and C. H. Gordon. 1972. Relationships of forage composition with rates of cell wall digestion and indigestibility of cell walls. J. Dairy Sci. 55:1140. Troelson, J. E. and J. B. Campbell. 1968. Voluntary consumption of forage by sheep and its relation to the size and shape.iof particles.dn the digestive tract. Animal. Prod. 10:289. Ulyatt, M. J., K. L. Blaxter, and I. McDonald. 1967. The relations between the apparent digestibility of roughages in the rumen and lower gut of the sheep, the volume of fluid in the rumen and voluntary intake. Anim. Prod. 9:463. Van Hoven, W. 1980. Rumen fermentation buffalo (Syncerus caffer, Sparrman, Koedoe, 23:45-55. and methane production in African 1779) in Kruger National Park. ____ "-'_ '" and E. A. Boomker. 1981. Feed utilization and digestion in the black wildebeest (Connochaetes ~, Zimmerman, 1780) in the Golden Gate Highlands National Park. S. AFr. J. WIldl. Res. 11:35-40. Van Soest, P. J., and J. A. Robertson. 1980. Systems of analysis for evaluating fibrous feeds. Pages 49-60 in W. J. Pigden, C. C. Balch, and M. Graham, eds. Standardization of analytical methodology for feeds. Int. Develop. Res. Center and Int. Union Nutr. Sci. Ottawa, Onto 1966. Forage intake in relation to chemical composition and digestibility: some new concepts. p24 in Proc. Southern Pasture Forage Crop Improvement Conf. Van Soest, P. J. 1965a. Symposium on factors influencing the voluntary intake of herbage by ruminants: voluntary intake in relation to chemical composition and digestibility. J. Anim. Sci. 24:838-843. 1982. Nutritional Inc., Covallis. 374pp. ecology of the ruminant. 0 and B Books, Wallmo, O. C., and W. L. Regelin. 1981. Rocky Mountain and intermountain habitats. ~art 1. Food habits and nutrition. Pages 387-421 in O. C. Wallmo, ed. Mule and black-tailed deer of North America. Univ. Neb. Press, Lincoln. Weston, R. H. 1966. The significance tract to handle Austr. J. Agric. Factors limiting the intake of feed by sheep. I. of palatability, the capacity of the alimentary digesta and the supply of gluconeogenic substrate. Res. 17:939. Wolin, M. J. 1960. A theoretical Sci. 43:1452-1459. Prepared by ~.I?~ D. L. Baker rumen fermentation balance. J. Dairy � https://cpw.cvlcollections.org/files/original/ce4c943dcf75af32f50f2c3928291370.pdf 72475e5b9d3a2b5a92401d57ab45f934 PDF Text Text Colorado Division of Wildlife Wildlife Research Report July 1983 155 JOB PROGRESS State of REPORT Colorado Project No. 45-01-503-15050 Big Game Investigations Work Plan No. Multispecies Job No. Animal and Pen Support Big Game Research Period Covered: Author: Personnel: - Noncervids Investigations Facilities for 7/1/82-6/30/83 P. H. Nei 1 P. H. Neil ABSTRACT Modifications to the facilities and wind damage repairs were completed to facilitate several research programs. Training of mule deer and antelope for use in digestion cages continued during the early portions of year. A study concerning the viability of bindweed seeds after passage through the pronghorn digestive system was undertaken. Details and results are reported in Appendix 1. Sample analysis and data summary were completed for the aspen bark palatability study. Results indicate that these pellets are at least palatable, comprise a low qual ity diet, and that the main value to the animal is derived from cell soluable material. The total captive big game research herd presently consists of 22 mule deer, 5 elk, 9 Rocky Mountain goats, 7 b Iqho rn sheep, 11 pronghorn antelope, and one domestic cow. ��I'J/ ANIMAL AND PEN SUPPORT FACILITIES FOR BIG GAME RESEARCH Paul H. Ne i1 P. N. OBJECTIVES To provide and maintain populations of captive big game animals and pen facilities to support big game research programs. SEGMENT OBJECTIVES 1. Continue to develop facilities at the CSU Foothills Campus and to maintain facilities and animals at the Wildlife Research Center and CSU Foothills Campus. 2. Coordinate rearing, training, and ,research activities with captive wild and tame big game animals for all big game cervid and noncervid research projects. 3. Integrate big game animal and physical plant support facilities, resources, and fiscal resources into a single budget. human METHODS AND MATERIALS Routine neonate rearing procedures were used to hand rear and train 17 mule deer fawns for future nutrition studies and supplement the big game research herd. Nine adult mule deer were transported via horse trai~er to the Kremmling facility in November, 1982, in support of the mule deer thermal cover study project 45-01-502-15050, WP2, Jl, Dave Freddy, Principal Investigator. Materials and equipment were obtained to facilitate winter and early spring maintenance and neonate rearing at the Foothills and Fort Collins facilities. Repair of the five remaining wind damaged isolation pens was completed. Ten barrel type bear traps were constructed using 55 gallon barrels and del ivered to Crawford, Colorado, in support of Black Bear Project 45-01-503-15050, WP5, Jt , Tom Beck, Principal Investigator. A digestion cage study was undertaken in cooperation with the Game Damage Section, CDOW concerning the viability of bindweed seeds after being subjected to the digestive system of pronghorn antelope. Two pronghorn were placed in digestion cages for 6 days and initially fed 2,000 bindweed seeds each and maintained on ad lib pelleted ration. Fecal material was collected daily and air dried. Recovery of seeds from fecal material for germination trials was then completed. �158 Data summary from the Aspen Bark Palatability was completed and results are contained in this report. RESULTS AND DISCUSSION Neonate rearing was considered successful this year with seventeen of twenty mule deer fawns, 3 bighorn sheep, and one Rocky Mountain goat being hand-reared and trained for digestion cage, energetics, and other nutritional studies. All mule deer fawns were weaned, ear tagged, and male fawns castrated in April, 1983. Eight of the nine adult mule deer used in the Thermal Cover Study at the Kremmling facility were returned to the Foothills facility in April, 1983. One female died of chronic wasting disease at Kremmling. Results of the pronghorn study concerning the viability of bindweed seeds after being subjected to the digestive system of two pronghorn are contained in a Colorado State University Special Report (Appendix A). Sample analysis and data summary were completed for the Aspen Bark Palatability Study conducted during the winter of 1982-83. Summary tables are contained in Appendix B. It is apparent that this is a low quality diet (Table l). Some variation in intake of the aspen pellets does exist and appears to be related to variation in body size of the deer (Table 2). In-vitro digestibility overestimates in-vivo digestibility of the aspen pellets by 10 units or approximately 33 percent (Table 3). Dry matter digestibility appears very similar among deer. Fiber constituent digestibility was relatively low and greater variation among deer exists. It is apparent that the aspen bark pellets comprise a low quality diet and that the main value to the animals is derived from cell soluable material. Twelve mule .deer and two elk died during this fiscal year. The total research herd presently consists of 22 mule deer, 5 elk, 9 Rocky Mountain goats, 7 bighorn sheep, 11 pronghorn antelope, and one domestic research cow. Other activities that occurred during the year included the following: Tour of facilities - September, 1982 - CSU students Lecture - March, 1983 - "Immobilization Equipment and Techniques" Colorado Division of Wildlife DWM Trainees Tour of facilities - April, 1983 - CSU Wildlife Nutrition Class Tour of facilities - April, 1983 - CSU Wildlife Management Short Course Lecture - May, 1983 - "Capture Techniques: Mechanical and Chemical Colorado Association of Animal Control Officers Prepared by __G:>auf.H Paul H. eil ~'..f Wildlife Technician III II - �APPENDIX A 159 Loreen Anne Ryan Department of Fisheries and Wildlife Biology Colorado State University Fort Collins, CO 80523 • VIABILITY OF BINDWEED SEEDS AFTER PASSAGE THROUGH THE PRONGHORN DIGESTIVE SYSTEM LOREEN ANNE RYAN, Department of Fisheries and Wildlife Biology, Colorado State University, Fort Collins, CO 80523 There is concern by landowners that pronghorn (Antilocapra americana) may be a major factor in the spread of bindweed (Convolvulus arvinsis) from infested croplands to their ra~gelands or croplands. Food habit studies in Colorado (Hoover et ale 1959) and California (Ferrel and Leach 1950) show that pronghorn eat bindweed in minute quantities. However, bindweed was not found as a food item in Oklahoma (Buechner 1950), Oregon (Mason 1952), Montana (Cole and Wilkens 1956), Saskatchewan (Dirschl 1963), New Mexico (Russel 1964), Wyoming (Severson et ale 1968), or Texas (Hailey 1979). -.If soil moisture is adequate, bindweed will remain green in wheat fields through the summer, flowering and producing seed until the first frost. According to Eugene Heikes (Extension Weed Specialist, Colorado State University), bindweed seed remains in the capsule until it is shattered by the plant drying or from some disturbance. Pronghorn may eat bindweed seed when they are grazing in stubble fields in the late summer and early fall. Ferrel and Leach (1952) found bindweed in the rumen of pronghorn with a 3% frequency of occurrence in California in the fall. Bindweed seeds are extremely durable. In soil, bindweed seeds can remain viable for more than 30 years (Phillips 1961). The seed is very �160 hard and germination percentages at anyone time are usually quite low unless the seed coat is weakened (scarified) mechanically or with acid. A study in Nebraska (Harmon and Keirn 1934) showed that viable bindweed seed passes through the digestive system of calves, horses, sheep and hogs. A thousand bindweed seeds were fed to various animals and the seeds recovered from the droppings for the next 4 days. unscarified tracts. controls. Germination of seed was increased by passage through the animals' digestive Acid-scarified seeds also had a higher·germination rate than The recovery period (4 days) in this study was probably a little short considering that in an 'Idaho study (Leherer and Tisdale 1956) viable seeds of other plant species remained in the digestive tracts of sheep for as long as 9 days. Viable seed of other plant spe- cies have been shown to pass through the digestive system of sheep and rabbits (Leherer and Tisdale 1956) and sheep and deer (Heady 1954). Since pronghorn would be expected to be much like sheep or deer, it is ~~ probable that bindweed seeds injested by pronghorn would be viable after passage through the digestive system. This study addresses the question of whether or not passage through the pronghorn digestive system effects the viability of bindw d s ds. METHODS Two tame pronghorn (a yearling buck - Speedy, and a 2-year-old doe - Tinker) were trained over a period of 10 days to stay comfortably wire-mesh bottomed digestive cages. wire-mesh flooring caught the feces. Collecting in trays underneath the Each pronghorn was fed by hand about 100g of grain mixture feed, maple syrup and 1000 bindweed seeds in the morning and afternoon on the first day of the trial. Feces were �161 collected from the collection trays twice daily (morning and afternoon) for 6 days, beginning with the afternoon of the seed feeding. Each day's feces were put in individual paper bags and marked with the animal's name and date. The pellets were air dried for several days. The seeds were separated from the pellets by carefully crushing each individual pellet with a mortar and pestle and searching the material for seeds. Samples of seeds from each pronghorn and a control sample of undigested seeds were germinated for 10 days. square plastic-covered dishes. environment germinator. the temperature day. Seeds were placed on moist blotters in The samples were put in a controlled The seeds were germinated in total darkness with kept at 300C for 8 hours a day and 200C for 16 hours a Four separate samples were germinated and cumulative numbers of seeds from each dish were recorded each day for 7 days, beginning the third day of germination. The seeds used were obtained from a commercial seed company (Valley Seed Service, ,Fresno, CA). Source or treatment of the seeds are unknown, however, alfalfa (Medicago sativa) seeds and pods were found among the bindweed seeds. It may, therefore, be assumed that the bindweed seeds were collected in conjunction with a commercial alfalfa seed operation. It is not known what chemical treatment of the fields, if any, was used. It is also unknown what phenological when harvested. stage the bindweed plants were in These treatments may affect seed coat hardness and, therefore, germination. As a check on the viability of the California seed supply a sample of naturally grown and ripened bindweed seeds from a vacant lot in Fort Collins, Colorado, were collected. The Colorado plants were not treated chemically and the seeds were naturally ripened on the vine and removed �162 from the pod by hand. The seeds were harvested in April after remaining on the vine and in the pod over winter. RESULTS AND DISCUSSION Ninety-nine and 107 bindweed seeds were recovered in the feces of Speedy and Tinker, respectively. A total of 40.8 percent of the seeds from Speedy germinated and 30.0 percent of the seeds from Tinker germinated. Chi-square analysis of germination ~uccess between the two pronghorn showed no significance the germination (P > 0.05, X2 = 1.43). Comparison of success of seeds recovered from the pronghorn feces to the control seeds showed for Speedy and X2 = a significant difference (P <"0.05, X2 = 6.813 13.500 for Tinker). These results indicate that there is a lower percentage of germinat(rtt.b1e I) ing seeds after passage through the pronghorn digestive syst~~. This differs from the Nebraska study (Harmon and Keirn 1934) which showed a higher germination percentage after passage through digestive systems of .. calves, horses, sheep and hogs. It seems unlikely that the pronghorn digestive system inhibits germination of bindweed seeds. Approximately percent of the 2000 seeds fed to each pronghorn were recovered from the feces. Therefore, nearly 95 percent of the seeds were digested. It is possible, then, that the seeds that were recovered were harder coated seeds that allowed their survival of the digestive process. These seeds may take longer to germinate or have a lower germination potential. The seeds used for this study were from California (1981) and were assumed to be recovered from a commercial alfalfa seed operation. Germination of the California-grown seeds were compared with seeds 5 �naturally grown and ripened in an abandoned lot in Fort Collins, Colorado. Chi-square analysis showed a highly significant difference (P < 0.05 X2 = 133.47). in germination Only 9 of 200 Colorado seeds germinated while 133 of 200 California seeds germinated after 10 days. It is possible the treatment of the California seeds, either chemical or mechanical, provided some degree of scarification of this hard-coated seed which resulted in higher germination. CONCLUSIONS Percent germination of California bindweed seeds was lower after passing through the pronghorn digestive system. Since there is such a difference in germination between Colorado and California seeds, this . study should be repeated using Colorado seeds. ACKNOWLEDGMENTS I would like to thank the following persons for their support and cooperation in this study: Tom Pojar, Paul Neil, and Andre' Duvall of the Colorado Division of Wildlife; Dr. Wm. Laycock and Dr. A. Wilson of the U.S.D.A., Agriculture Research Station; and Dr. D. Bowden of the Statistics Department, C.S.U. LITERATURE CITED Buechner, H. K. 1950. Range ecology of the pronghorn on the Wichita Mountains Wildlife Refuge. Cole, G. F. and B. T. Wilkins. Trans. N. Amer. Wildl. Conf. 15:627-644. 1956. The pronghorn antelope, its range use and food habits in central Montana with special reference to wheat. Montana Fish and Game Dept. Tech. Bull. 2:1-39. �164 Dirschl, H. J. 1963. Food habits of the pronghorn in Saskatchewan. J. Wildl. Manage. 27:81-92. Ferrel, C. M. and H. R. Leach. antelope of California. 1950. Food habits of the pronghorn Calif. Fish and Game 36(1):21-26. Ferrel, C. M. and H. R. Leach. 1952. The pronghorn antelope of California with special reference to food habits. Calif. Fish and Game 38(3):285-293. Hailey, T. L. Texas. 1979. A handbook for pronghorn antelope management in Texas Parks and Wi1dl. Dept. F. A. Rep. Series No. 20. Harmon, ~. W. and F. D. Keim. 1934. 50pp. The percentage and viability of weed seeds recovered in the feces of farm animals and their longevity when buried in manure. Heady, M. F. 1954. and deer. J. Amer. Soc. Agron. 26:762-767. Viable seed recovered from fecal pellets of sheep J. Range Manage. 7:259-261. Hoover, R. L., G. E. Till and S. Ogilvie. 1959. The antelope of Colorado: A research and management study. _Colo. Dept. of Game and Fish Tech. Bull. N&. 4. llpp. Leherer, W. P., Jr. and E. W. Tisdale. 1956. Effect of sheep and rabbit digestion on the viability of some range plant seeds. J. Range Manage. 9:118-122. Mason, E. 1952. Food habits and measurements of Hart Mountain antelope. J. Wildl. Manage. 16:387-389. Phillips, W. M. 1961. Field bindweed and its control. Agric. Leaflet No. 496. Russell, T. P. 1964. 7pp. Antelope of New Mexico. Fish Bull. 12:1-103. U.S. Dept. New Mexico Dept. Game and �165 Severson, K., M. May and W. Hepworth. 1968. capacities and forage competition sheep in Wyoming's Red Desert. Monograph Prepared by carrying between antelope and domestic Univ. Wyoming Agr. Exp. Sta. Sci. 10:1-51. \~avuUf, '1J.=' ~ Paul H. Nei 1 Food preferences, Q_ �166 Table 1. Comparison of bindweed seed germination between seeds passed through the pronghorn digestive system and control seeds. Sample Cumulative Number of Seeds Germinated by Daya 3 4 5 6 7 8 9 8 18 23 27 30 38 40 Control _ CAb 33 54 63 67 67 67 67 Tinker 13 22 24 26 28 30 30 Control _ CAb 33 53 61 63 65 66 66 Colorado #"1c 3 4 4 4 4 4 4 Colorado #2c 1 3 5 5 5 5 5 Speedy aGermination test begun 25 April 1983 which was day 1. No seeds germinated on days 1 and 2 . . bSeeds obtained from a commerci a1 seed supplier from California. cSeeds natural )y grown and ripened in a vacant lot in Fort Collins, Colorado. �167 APPENDIX B Table 1. Average Chemical ComQosition of Ration Aspen Bark Pellets (x of 3 samples) x Chemical Nutrient Ash OM GE (Cal/g) CP Nitrogen Cell Soluables NDF ADF ADL .. Hemicellulose Cellulose IVDDM LIG/ADF Ratio composition (%) 22.00 73.60 4264.00 5.64 .90 43.51 56.50 42.60 23.20 14.10 19.30 31.00 54.70 Table 2. Aspen Bark Pellets Intake Summary Species Deer Deer Deer Deer Animal No. 144 187 225 227 Average Body \~t (kg) Average dry matter intake g/day g/day/wt (kg) 75.00 68.00 33.75 1432 1429 594 19.0 21.0 17.6 59.0 60.4 42.0 44.00 1422 32.3 33.6 55.00 9.80 1219 208 22.5 3.4 61.0 8.5 g/day/wt (kg·75) (X) Hean SE �168 APPENDIX B (continued) Table 3. Appearent mean digestibilities Dry matter Organic matter DE Crude protein Cell sol. CWC Hemicell ulose ADF Cellulose 31 21 26 25 -24 44 2 -4 3 -27 21 227 31 20 46 25 -.5 35 9 12 8 14 -23 144 31 21 26 25 -21 84 4 14 -1 13 31 20.6 32.6 25 -15 54.3 5 7.3 4 -4 19 0 0.3 5.7 Q 5.7 13 1.8 4.9 1.8 10.4 2.6 Speci es No. IVDDM Deer 187 Deer Deer Mean SE (%) of aspen bark pellets consumed by mule deer ADL �Table 3. Appearent mean digestibilities (%) of a~pen bark pellets consumed by mule deer Dry matter Organic matter DE Crude protein Cell sol. CHC Hemicellulose ADF Cellulose ADL Species No. IVDDM Deer 187 31 21 26 25 -24 44 2 -4 3 •.27 21 Deer 227 31 20 46 25 -.5 35 9 12 8 14 -23 Deer 144 31 21 26 25 -21 84 4 14 1 -1 13 31 20.6 32.6 25 -15 54.3 5 7.3 4 -4 19 o 0.3 5.7 o 5.7 13 1.8 4.9 1.8 10.4 2.6 Mean SE '" I..D ��171 Colorado Division of Wildlife Wildlife Research Report July 1983 JOB PROGRESS REPORT State of Colorado 45-01-503-15050 Project No. Big Game Investigations Work Plan No. Multispecies Job No. Author: Investigations Noncervid Research Administration 3 PerIod Covered: - Noncervids 7/1/82-6/30/83 R. B. Gill Personnel: L. H. Carpenter, L. E. Lovett, N. McEwen ABSTRACT Improvements were made in the computerized budget accounting system which increased output accuracy and improved reporting punctual ity. Document storage and filing system was revised and a file descriptor was prepared to improve document storage and retrieval. Annual Work Plans were redesigned to be compatible with Department of Personnel IS new FAPAS performance evaluation system. Draft Program Plans were prepared to describe research objectives for the period 1983-84 through 1988-89. The Wildlife Research planning process was diagrammed to illustrate the process and the interrelationships among planning and monitoring documents. Efforts were increased to expand and improve internal communications with other non-research organizational units in DOW: Results were positive. Research productivity reached its highest point in the history of big game research in Colorado. Results of the research audit are reviewed and critiqued. This Job Progress Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the Director. ��173 NONCERVID RESEARCH ADMINISTRATION R. B. Gi 11 P. N. OBJECTIVE Supervise and administer research activities within the scope of Federal Aid Project 5503X (45-01-503-15050) Big Game Investigations - Noncervids. SEGMENT OBJECTIVES Supervise and administer all studies in Project W144R-2 (45-01-503-15050). ACKNOWLEDGEMENTS Collectively the members of the staff of Big Game Investigations Noncervids, deserve most of the credit for attaining the several research objectives of the project. Even though injury, sickness, personal and professional traumas, and abnormally adverse weather conditions presented formidable barriers, all of the research objectives were met within time and budget constraints. The unstinting dedication, commitment and effort each member of the Big Game Investigations - Noncervids staff gave to achieve those objectives was inspirational and immeasurably gratifying to me. This acknowledgement is a woefully inadequate way of saying thank you. INTRODUCTION In the broadest sense, the goal of the Big Game Investigations - Noncervids Project is to provide better service and better research products each year to the several research clients of the Colorado Division of Wildlife. Effective and efficient administration is one of the key tools necessary to meet that goal. Administrative services such as personnel supervision, accounting, purchasing, typing, file storage and retrieval, photocopying, and editorial services consume a significant proportion of the resources allocated to research programs. Accounting for the expenditures of those resources should be equally rigorous to accounting requirements demanded of research activities. The purpose of this report is to provide an account of how administrative resources were expended and of the products that resul ted. RESULTS AND DISCUSSION I established several specific objectives for the Big Game Investigations Noncervids Section for FY 82-83. Among these were: I. Improve administrative efficiency Investigations projects. and efficacy within the 2 Big Game - �174 2. Prepare revised and updated drafts of the 5-Year Operations the Big Game Investigations - Noncervids Section. Plans for 3. Develop improved systems and processes to communicate more effectively with research clients of the Colorado Division of Wildlife. 4. Increase the productivity of the Big Game Investigations Section through better supervision. 5. Develop and begin implementing of the Big Game Investigations - Noncervids an in-service training program for staff - Noncervids Project. 6. Develop a more relevant reward system to reward above-standard outstanding performance Noncervid project. and among the staff of the Big Game Investigations ADMINISTRATIVE - IMPROVEMENTS One problem the staff of the 2 Big Game Investigations Projects (Cervids and Noncervids) has faced was to develop an accounting system which provided timely and accurate accounts of the budget status of each research study upon demand. Each staff member of the Big Game Investigations projects has been delegated the responsibility and authority to manage the fiscal resources of his own research program. But the periodic budget status reports issued from the Denver office are not sufficiently detailed nor sufficiently timely to provide the level of service that is needed. This is especially true during the last! of the fiscal year when expenditures have to be monitored closely to prevent overspending. In 1978 we developed a computer program and a data storage and retrieval system that would provide a complete and accurate record of expenditures of each Big Game I.nvestigations staff member. Our goal was to routinely provide quarterly statements to each scientist detailtng his budget status for the first 3 quarters of the fiscal year. During the fourth quarter of the fiscal year, we attempted to provide monthly budget status reports. We also wanted the system to be updated every 2 weeks so that anyone could ask for a budget status report at any time and it would be accurate to within 2 weeks of the reporting date. Initially, coding error rates and program Ilglitchesll were unacceptably frequent. This year welve revised coding and recording systems and redesigned the file storage and retrieval so that error rates are acceptably infrequent. The result is that budget status reports were issued on time and were accurate. During FY 82-81 the entire document storage and retrieval system was redesigned and was purged of all unnecessary material. A typewritten file descriptor was prepared to facilitate document location by anyone unfamiliar with the system. During FY 82-83, qual ity control was improved within the Big Game Investigations internal peer review system. Transit time of manuscripts through the system has been r·educed and the qual ity of the reviews has been enhanced. �175 Year end purchases have always been hectic and somewhat chaotic because of changing and increasing regulations from State Purchasing. State rules and regulations require rapid and efficient reporting so purchasing activities can continue to the end of the fiscal year. In FY 82-83 we had problems wherein duplicate receiving reports were issued for 3 purchases. None of these incidents led to actual dual payments, but they could have. We changed purchasing protocol of the Big Game Investigations projects so that only the Big Game Investigations Secretary prepares receiving reports for the purchaser's signature. This should eliminate the problem of dual receiving reports in the future. The Department of Personnel implemented a new employee performance evaluation system in January 1983. This system is called FAPAS (Factor/Anchored Performance Appraisal System). I participated in 2 FAPAS workshops to develop Factors and Anchors for the Research and Wildlife Job Families. also critiqued the general FAPAS system and suggested ways to improve the system (Appendix A). During June and July 1983 I restructured all of the Annual Work Plans of the staff of the Big Game Investigations - Noncervids project to accommodate the new FAPAS format and reviewed these with each employee on' my staff. RESEARCH OPERATIONS PLANS In 1971 the Colorado Division of Wildlife formed a Planning Section to develop a comprehensive planning system for the then Colorado Division of Game, Fish, and Parks. This planning system was supposed to produce a hierachy of plans which al igned and integrated the annual work assignment of each CDOW employee with the overall CDOW goals and objectives defined in d statewide resource management plan called a Statewide Strategic Plan. Operation management plans were to be the next hierarchial level and these plans were supposed to define specific species goals for each organizational unit or program;and "plans to accomplish these goals" (Appendix B). The final planning hierachy was to be the Annual Work Plan of each employee assigning specific tasks to accomplish the objectives of the operation management plans and the statewide resource management plan. The first resource management plan was published in 1974 and was entitled "The Strategy of Today, for Wildlife Tomorrow" (Colo. Div. Wildl., 1974). However, no Division-wide operation management plan system or formats were established during the 5-year planning horizon encompassed by the 1974 Strategic Plan. In December 1977 the Colorado Division of Wildlife updated the first Strategic Plan and published the revision as "Today's Strategy ... Tomorrows Wi ld life" (Colo. Div. Wi ld l,, 1977). That plan declared ..... "The Iinks between the Strategic Plan and the way the Division of Wildlife spends money and resources are the strategies ..... When an organizational unit manager develops his annual operating plan to determine funding needs and work schedules, those activities which contribute most to the top priority strategies will be given first consideration for funding. Thus by designing projects which contribute most to top priority strategies, managers can become better competitors for limited funds as they improve the Division's progress toward its most important objectives." �176 In order to enhance the contribution of Big Game Research toward the accomplishment of Strategic Plan goals and objectives, the Big Game Research Section prepared a draft long-range research plan in 1977 which closely linked big game research proposals with top priority Strategic Plan strategies. That plan was subjected to thorough internal review and was approved for implementation in June 1978 (Gill, 1978). This plan was the first comprehensive research program plan. In 1981 work was begun to update the Strategic Plan for the third planning period, 1983-1988. The third edition of "Todays Strategy ... Tomorrow's Wi ld life" (Colo. Div. Wi Idl., 1983) was publ ished in January 1983. Each organization unit was instructed to prepare draft operations plans linking the objectives of each organizational unit to highest priority strategies listed in the new Strategic Plan. The deadline assigned to the Big Game Investigations - Noncervids Section was February 1, 1983. We completed draft plans by December 1, 1983 (Appendix C). However, before internal review of these plans was completed, the format was changed completely. The new format called for a 2 component operations planning" system - program plans and'implementation plans. Program plans relate the objectives of each organizational unit to Strategic Plan strategies, and implementation plans describe how products of those objectives will be implemented in DOWis species management programs. The Big Game Investigations - Noncervids Section submitted draft program plans for review by July 8, 1983 (Appendix D). Finally, as part of the requirements of the research audit, the 2 Big Game Investigations prepared a diagramatic representation of the research planning process. This diagram summarizes the relationships among hierachial plans and links the planning process to the FAPAS performance monitoring and evaluation system (Fig. 1). RESEARCH COMMUNICATIONS In FY 82-83 the Big Game Investigations - Noncervids Section made a concerted effort to improve our communications with operations staff. The primary audiences were Regional Wildlife Biologists, Denver Program Staff, Denver Wildlife Services staff and District Wildlife Managers. Communications goals were established for each Wildlife Researcher as part of his 1982-83 Annual Work Plan. In each case, minimum communications goals were met or exceeded. Results were a clearer understanding of what we are doing, why we are doing it, and what it will contribute to DOWis species management programs. Support for the Big Game Investigations - Noncervids program is increasing among Regional personnel. Goals for 1983-84 will be to increase awareness and support for these programs at the Director's Staff and Wildlife Commission leve ls, In 1983-84 we will produce a draft research information plan that identifies communications objectives, audiences, messages, and media. �177 WILDLIFE RESEARCH SECTION PLANNING PROCESS STRATEGIC PLAN Policy document 1. Identifies programs Establ ishes statewide and regionwide objectives 3. Defines strategies to meet objectives 4. Planning period: 5 & 15 yrs 2. STRATEGIES PROGRAM PLAN Action Program Outline Defines strategies into research problems 2. Identi fies tasks (studies) 3. Estimates resource requirements (funds. personnel. time) 4. Planning period: 5 yrs 1. TASKS (STUDIES) IMPLEMENTATION PLAN~ Research Strategy Document l. Establishes need (synthesizing Lit. Rev. ) 2. States hypotheses 3. Describes hypothesis testing strategies 4. Establishes schedule 5. Identifies research client and product targeted for each 6. Length of each study RESEARCH STRATEGI ES ANNUAL WORK PLAN Responsibility-Resource Allocation Document 1. Assigns work (products) Di,stributes fiscal resources 3. EstabHshes work standards (quantity-quality) 4. Planning period: 1 yr 2. PRODUCTS & SCHEDULES FAPAS FIG. 1 - Diagramatic summary of the Wildlife Research Planning Process and planning documentation. Performance Monitoring Document 1. Monitors resource use vs. production 2. Adjusts performance as required 3. Rewards performance (-I-' or .) 4. Planning period: 1 yr �178 RESEARCH PRODUCTIVITY Research productivity is not easy to measure because the primary research product is knowledge. The most reI iable index to the quality and quantity of knowledge production is numbers of scientific articles published in peer reviewed scientific publications. Productivity of the 2 Big Game Investigations Sections has increased dramatically since 1973 even while the size of the Big Game Research staff was declining (Fig. 2). During 1982-83,at least 15 peer reviewed technical manuscripts authored or co-authored by staff of the Big Game Investigations - Noncervids Section were published or accepted for publication. In addition, at least 10 papers were prepared and presented at professional society meetings and workshops. In summary, 1982-83 was the most productive year, in terms of scientific output, of the 2 Big Game Investigations Sections in the entire history of big game research in Colorado. RESEARCH Although individual staff enrolled in a 'no formal in-service This remains a high IN-SERVICE TRAINING members of the Big Game Investigations - Noncervids variety of continuing education programs in FY 82-83, training plan for the entire Section was prepared. priority goal for FY 83-84. RESEARCH PERFORMANCE AWARD SYSTEM In FY 82-83 I attempted to obtain approval for developing an incentive award system for Wildlife Researchers that would reward above-standard and outstanding work performance with increased opportunity for out-of-state travel to professional meetings and for research work trips to research labs and research stations of other states and other countries. I believe such a reward system would have 2 very positive benefits. First, it would provide a nonmumerative incentive for extraordinary work performance which should provide positive feedback into increased productivity. Second, it would provide a mechanism for person-to-person exchanges of research ideas with workers involved in similar research assignments. This should increase research competence, decrease duplication of effort, and incr ase work quality. Unfortunately, key members of the Director's staff remain unconvinced of the merits of this idea, and it still has not progressed beyond the discussion stages. RESEARCH LEADER PRODUCTIVITY During FY 82-83, I set a goal for myself to publish at least 2 technical articles in peer-reviewed journals. I did not achieve that goal. I did succeed in getting I manuscript accepted for publication in JWM (Gill et al., 1983a~ one paper published in'the Transactions of the North American Wildlife and Natural Resources Conference (Gill et aI, 1983b) and presented a paper to the Colorado Chapter of TWS on research philosophy which is being prepared for submission to the Wildlife Bulletin for publication consideration (Gill, 1983). �179 PERIOD PER 00 I n 25 Em D Papers at prof. mtgs. and symposia DOW scientific publications Peer reviewed articles 20 15 10 5 o 65 66 67 68 69 70 71 72 73 74 75 76 77 '78 79 80 81 82 YEAR Period I (1965-1973) Period II (1974-1982) Average Peer Reviewed publications/year 2.44 5.00 Average DOW Scientific publications/year 0.78 1.11 Average Papers at pr-ofessional meetings and symposia/year 0.78 1.22 Average Scientists productivity ( s of 3 above -:-9) 4.00 8.11 Average Productivity/scientist 2.65 5.71 Average Productivity/scientist/year 0.29 0.63 FIG. 2 - Productivity of the Big Game Research Staff, 1965-1982. �180 RESEARCH AUD IT Following the November, 1982 Wildlife Commission Meeting, it was announced that the Division of Wildlife was going to audit the real estate, planning and budgeting, and research processes. Initially, some members of the Wildlife Commission wanted to audit the entire Division of Wildlife to ascertain how well goals and objectives outlined in the Strategic Plan were being met. After considerable debate the Wildlife Commission compromised on an audit of the real estate, planning and budgeting, and research processes. Why these processes were singled out sti 11 remains unclear, but one reason given for the research audit was that key legislators and the Wildlife Commission wanted to know why the Colorado Division of Wildlife spends so much money on research. On November 29, 1982, Director Grieb outlined Audit as follows: the Divisionis the objectives philosophy of the Research 1. Define explicitly research. and policy regarding 2. Provide an acceptable research proposals. 3. Develop a Research Operations Plan (where we are vs. where do we want to be) and a control system (so we know when we get there) that supports the Strategic Plan. 4. Conduct an audit of the total Divisionis align research to Division objectives. system that allows adequate analysis of alternative research activity to better Since November 29, 1982, research audit business has commanded approximately 70% of my time and energies. Since that time and those energies were supported with Federal Aid funds, itls appropriate to give a brief accounting of how they were spent and what was accomplished with them. A research audit committee was formed and was comprised of Acting Assistant Director Ed Prenzlow, Chairman; Spencer Smith, retired Director of USFWS; Fred Samson, Leader Colorado Cooperative Wildlife Research Unit, Jim Lipscomb, CDOW Game Program Chief; Ed Kochman, CDOW Fish Program Chief, John Torres, CDOW Nongame Program Chief; Dick Hopper, Wildlife Research Chief; and Don Horak, Fisheries Research Chief. The first task of the research audit committee was to address objective 1 formulation of research philosophy. The deadline for drafts of the research phi losophy statements was January 31, 1983. I was assigned to prepare the Wildlife Research Sectionls draft. That draft was submitted to the audit committee members in January, 1983 (Appendix E). A final decision was supposed to be made regarding the final approved statement of research philosophY and policy in DOW by April 1, 1983. Despite the fact that this was the number one objective of the research audit and despite the fact that more than 60 typewritten pages of research audit transactions were devoted to this topic, no decision has yet been made regarding CDOWls philosophy and policy of research. In addition, this topic was discussed only once by the entire research audit committee. No drafts of a new �181 research policy resulted from that discussion. Commission adopted a policy on research in 1975 the research audit there were no discussions of extant policy which might have provided a basis Finally, the Wildlife (Appendix F). Throughout the inadequacies of this for constructive revisions. Objectives 2 and 3 of the research audit seem to reflect a lack of awareness on behalf of the research audit committee regarding current CDOW standard operating procedures. In August, 1982, Director Grieb implemented a research project selection process and a research implementation process via Administrative 1-1 and 1-2 (Appendix G). These were reviewed by the General Staff and the Director's Staff before they were implemented, so they should have reflected input from the entire staff of CDOW. The research audit committee seemed to be unaware of their existence. Objective 3 required the Research Sections to have draft Operations Plans prepared by February 10, 1983, for submission to the audit committee. Apparently the research audit chairman was unaware that the Wildlife Research Section had draft Operation Plans prepared before the research audit was commissionec;i. Objective 4 was never really addressed by the research audit committee. The only research projects which were audited were those of the Research Sections, not lithe total divisionis research activity." Regional research studies such as the interstate pronghorn distribution study of the Northwest Region, the Fruitland Mesa deer study of the Southwest Region, the Hugo pronghorn population study of the Southeast Region, the prairie chicken status study of the Northeast Region, etc., etc., were not audited. On May 18, 1983, the research audit recommendations were released to CDOW employees and to the general public via the Nongame Advisory Council. There were 15 recommendations contained in this report. None of them dealt directly with the 4 original research audit objec.tives. Two of those recommendations call~d for a 27% reduction in research staffing and funding and a reallocation of significant portion of the remainin~ research effort away from biological research towards operations research. Yet, part V of the research audit recommendations (which was included with the research audit recommendations) indicated that 24% of the studies reviewed by the research audit committee should be expanded with additional money and manpower. In addition 40% were recommended to continue with existing levels of resource allocations. Another 27% were recommended to continue at current funding and staffing levels, but were recommended for further reevaluation and slight redirection. Only 9% of the current studies audited by the research audit committee were recommended to be discontinued, and at least half of those will terminate on their own by July 1, 1983~ The research audit recommendations never define any problems that the recommendations are designed to correct. They do not list any alternative recommendations. They do not indicate what will be done with the people who are taken off research assignments. So there is no way to evaluate this staffing reduction recommendation in terms of what will be lost from the research effort vs. what will be gained by reallocating these people to other activities. These deficiencies in the research audit process and in the recommendations are glaring in light of the fact that the research audit has cost the Colorado Division of Wildlife an estimated $500,000 - �182 $750,000 in salaries, reduced productivity, per diem, etc., and still has not concluded. Much of this cost has accrued to Pittmann-Robertson and Dinge11~Johnson cost centers. LITERATURE CITED Colorado Division of Wildlife. 1974. The strategy of today, for wildlife tomorrow. Colo. Div. Wi1d1., Denver, CO. 103pp. Colorado Division of Wildlife. 1977. Today's strategy ... tomorrow's wildlife. Colo. Div. Wildl., Denver, CO. 96pp. Colorado Division of Wildlife. 1983. Today's strategy ... tomorrow's wildlife. Colo. Div. Wildl., Denver, CO. 96pp. Gill, R. G. 1978. Big game research program Ft. Coil ins, CO. 72pp. 1978-1988. Colo. Div. Wi1dl., Gill, R: B., L. H. Carpenter, R. M.' Bartmann, D. L. Baker, and G. G. G. G. Schoonveld. 1983a. Fecal analysis to estimate deer diets. J. Wildl. Manage. 47(4): (in press). Gill, R. B., L. H. Carpenter, and D. C. Bowden. 1983b. Monitoring large animal populations; The Colorado experience. N. Amer. Wildl. Natur. Res. Conf., Trans. 48: (in press). Prepared by -=-' _'~\-=-~__:"::=,,;;:-~:_:SJ:__.-=:"'<"=:.~=- R. Bruce Gi 11 Wildlife Research Leader _ �, r ., ~TATE Richard 183 APPENDIX A OF COLORADO O. Lamm. Governor DEPARTMENT OF NATURAL RESOURCES DIVISION OF 'NILDLiFE Jack R. Grieb. Director 6060 Broadway Denver. Colorado 80216 (825-1192) MEMO TO: R.M. Hopper FRQ.\l: R.B. Gill SUBJECT: Performance Appraisal Development Panel \ (\ ~.~. Dick: Following is my report on my act i \,1 t ies and thoughts regarding the PERFORH:\i\JCE APPR.!\ISALDEVELOP(-'rENT PA\TEL. First, by way of background, there were 6 people on the panel and 1 representative from the Department of Personnel. Four of the panelists we re in the Researcher series and 2 of us in the Wildlife Researcher series. To the best of my knowledge, I was the only panelist who supervised other certified. personnel. The impetuses for changes in the State's personnel appraisal system we re provided by Senate Bill 308 and general dissatisfaction with the previous appraisal systems, TR-l and TR-2. The new system - called FAPAS for Factor/Anchored Personnel Appraisal System - is intended to increase the objectivity of the appraisal process and to provide structured direction to the supervisor in making his performance appraisals. It wiLl do this by: a. grouping employees into "families" of similar jobs. For example - all personnel classified into the Researcher series or the Wildlife Researcher series ",ould be combined into the "Research Family" and would have identical appraisal systems; b. establishing more specific factors within the appraisal system than existed with TR-I and TR-2 (the old appraisal systems). The old systems (e.g. TR-I) rated employees in the following categories: Quanti ty of h'ork, Quality of Work, Taking Action Independently, Relationship with People, Work Habits, Effectiveness of Supervision, Equal Employment Opporttmity/Affirmative Action Performance and Other Pertinent Factors. FAPAS would replace those categories with the following: Program Administration; Problem Analysis; Research Skills; Occupational Competence; \\'rittenand Oral Communications; Human Resource ~!anagement/Sllpervision; Planning, Organizing, and Coordinating; Relationships wi th People; and Organizational Commitment and Adapt ab i li tv ; c. establishing "anchors" or guideline criteria wh i.ch indicate to the emnloyee and sunervisor what level of performance within each factor is ~~'"P~ctedat e~ch of :+ performance levels: Outstanding, Above Standard, St31ldard, and Below Standard. An example- of the "31lchors" for the four DEPARTMENT OF NATURAL RESOURCES. James smun, Vice Chairman Micnael 0 Monte Pascoe. t:xeCU(ive Director Richard Divettnss. Higbee. Memoer 0 Secretary Sam Cauciu. 0 0 WILDLIFE COMMISSION. Jean K. Tool. Member Member 0 0 Donald Fernancez. James C. Kennedy. WI!bur Redden. Memoer Memoer Chairman �184 rating categories under the Program Admi.nistrat i.on Factor might be: 4 - OUTSTAj\lDING(Met requirements for Above Standard, plus): Achieved the highest feasible levels of service in all major assignment areas given available resources. Achieved high level of accomplishment despite conditions of adverse circumstances; Made comprehensive program improvements; Conceived, developed and/or implemented a new program of significant impact on 01\lIl initiative; Developed and implemented a comprehensive cost-savings system that will have significant value; Significantly improved productivity on a fixed, allocated budget. 3 ABOVt STfu\~l\RD(Met requirements for Standard, plus): Achieved substantial improvement in quality; Maintained adequate productivity given conditions of adversity (i.e., reduced resources); Sought opportunities to affect program or project improvements \vithin and outside of assigned areas of responsibilities; Developed and implemented an i~novative procedure to reduce costs; Improved productivity on a fixed, allocated budget. 2 - STANDARD Schedules CL.'1d deadlines were usually met; Amonnt accomplished was adequate given available resources; Quality was acceptable; Range of services provided was adequate given available resources; Prepared an accurate, nnderstandable, and well-documented budget request; Allocated resources in conformance \flth needs and priorities; Made consistent use of appropriate cost-control procedures (e.g., monitored lnajor expenditures; detected and eliminated any excessive costs or expenditure): Assured adequate conservation of resources (e.g., incurred in unnecessary daInage to equipment or facilities); Took required steps to stay within budget, reducing costs as needed . .:> - 8ELOW STA\JDARD Often provided inadequate serVIce gIven available resources 1Il any major assignment area; Often failed to maintain acceptable quality; Created service backlog or had to cut services as a result of poor management; �18S .)- BELOW STAi'IDARD(continued) Seriously under- or over-budgeted requests; Prepareu inadequate documentation to support critical budget requests; Experienced significant problems due to poor financial planning or monitoring. d. the Factors are separated into 2 categories: Results Factors and Behavior Factors. Results Factors are those factors that result in accomplislunents or products while behavior factors are the activities or skills ·employed to effect results factors. Our job as participants on the Performance Appraisal Development Panel was (according to the letter we received inviting us to participate) to "help to develop and define the elements of your work and that of your coworkers that should be evaluated." Actually, there was only time sufficient to "fine-tune" the Factors which the Department of Personnel had already prepared for the Research Family from the job descr ipti.onsand job analyses of the,Researcher and \Vildlife Researcher Series. I encountered problems with both the FAPAS system and the Performance Appraisal Development Panel Process. The problems I have with the FAPAS system are: 1. One primary impetus for the new FAPAS system ivas that employee performance appraisals under the old system were frequently poor. They ivere subjective and some supervisors (the poor ones) did not communicate their expectations to their employees very well. In short, poor supervisors abused the old system. It was reasoned that when (if?) the state went to incentive awards for performance, that the proportion of poor ~mployee performance evaluations would have to be drastically reduced to make the incentive system work. Department of Personnel believed the way to do this was to implement a more detailed, specific appraisal system. It seems to me, however, that the new system will only penalize the good supervisor without impacting the performance of the poor supervisor. The old system afforded the supervisor considerable flexibility to individualize each employee's work assignment and appraisal. Good supervisors used the TR-l and TR-2 systems effectively because they had considerable latitude to fit the system to the individual. It was true that poor supervisors did not use the old system effectively. FAPAS will considerablY constrain the flexibility of the good supervisor because it wi ll force him to fit the individual to the system. FAPAS will not improve the performance of the poor supervisor because poor supervisors can still fail to use FAPAS effectively, and there arc no additional provisions in the FAPAS program to negatively reward the poor supervisor that do not exist with TR-l and TR-2. In my opinion, the Department of Personnel started at the wrong end to improve employee wor k assignments and performance appraisals. \lieneed to attack the problem by rest ructur ing the Civil Service system so that it 1S easier to discipline the poor performers - cmp lovee s and superv i sors . We also need to thoroughly retrain supervisors so they think in terms of assigning emp oyees work identified in terms of specific products related to agency goals and objectives and then rating each employee's �186 performance in relation to how well the employee met those production goals. Finally, supervisors need to be trained to include employees in all the decision processes that affect their work and/or their work envirorunent. Until this big job is undertaken, I think FAPAS will be just another band-aid solution. 2. The FAPAS system separates factors into results factors and behavior factors. Then the supervisor is supposed to rate an employee's performance in these two factor categories more or less independently. SO,311 employee could be rated above standard in all the results factors and ve t be rated below standard in a behavior factor- -such as Relationships wi th People. If you think about the relationship between results factors and behavior factors, however, this possibility should not even exist. Behavior factors are activities or skills used to produce job-relevant accomplishments or products. Therefore, if a person is below standard in a Behavior Factor, it should be reflected in Below Standard accomplishments or p rodnct s . I think the FAPAS sYstem should be restructured so that Behavior Factors become a subset of Results Factors (see the outline in Appendix A where I've 'tried to diagram my view of the relationships between Behavior Factors and Results Factors). The problems I had with the Performance Appraisal Deve lopment Panel were : 1. Only 2 days were allowed to address a very complex assignment with longterm and short-term impacts. The panelists were selected from a diversity of agencies and were all professionally trained employees. Given this diversity, it takes from 1-2 days just for the participants to sort out dominance and leadership roles before they can even begin to function as a creative team. Before this working relationship is established by the participants among themselves (it can't be imposed by an outside authority), the individual participants tend to react to the problem at hand via narrow self-interest responses. They respond by asking what the new system mean's to me, not what it means or can be constructed to mean for the long-term improvement in state employee performance appraisals. 2. Insufficient time was allowed to explore the philosophies of the old vs. the new system and why or if a new system is even necessary. In fact, because unrealistic deadlines had been imposed for the completion of the entire PAPAS review and revision process (December 15 was the deadline for incorporating panel input from all Families into the new system) only cosmetic revisions were possible. Substantive revisions would have taken too much time to accomplish within the 2day deadline and, therefore, were not even attempted. 3. As I mentioned in the introduction, I was the only supervisor on the panel. In addition, only Pat Dav i es and I were involved in traditional scientific research (i.e. hypothesis testing and experimentation). The other 4. panelists were classified tmder the Researcher series hut, in my opinion, would more appropr iat.eLy be called ana lys t s , If you read the job descriptions for the Researcher seri.es , it was intended to he a scientific research series, but several agencies hove classified employees as Researchers who really do little or no �187 scientific investigative work. When I tried to reorganize the Factors into a system that more logically reflected the scientific method (Appendix B), 4 of the panelists rejected that approach because they said it did not reflect their job assignments. There were no panelists who represented those considerable numbers of Researchers who do scientific investigative work. Therefore, the Factor prompters or "bullets" and the Factor anchors were generalized so they could fit both an analyst function and a scientific research flmction; but, in my opinion, they are not arrayed in any logical order. The end result will be that the good supervisor will have to wade through the entropy and bring some kind of logical order back into it, while the poor supervisor won't even try. 4. Since I was the only supervisor on the panel, I was the only one who had a real perspective of employee performance appraisal systems. I could think of how the system might be applied to me as an employee, and I also could think about how I might apply it to my employees Since the other 5 panelists tended to think only in terms of how it would be applied to them, it seemed to me that their cosmetic revisions of the system tended to be'defensive and employee -oriented vs. supervisor-oriented. What was needed \vas a panel composition that Hould have given a more holistic orientation to the cosmetic revisions. I'm not very optimistic that we panelists or the Department of Personnel have spent our time and resources most effectively to build a better system for assigning work and appraising employee performance. My "gut reaction" is that we have added additional shackles to the good supervisors and have not added any additional incentives to improve the performance of the poor supervisors, SO,on balance, I think we have built a more impoverished system than we had. In summary I would suggest the following to improve the system and the process of work assignment and employee performance appraisal: 1. Department of Personnel needs to devote at least as much time training supervisors in the art of assigning work in ways that identify products or accomplishments first and then outlines ways (behaviors) to achieve accomplishments or production. A large part of the reason the previous systems for appraising employee performance failed, I think, was because supervisors failed to be very specific in work ass i gnments and failed to identify products and/or accomplishments within those assignments. 2. 111e Results Factors need to be restructured for the Research family so they reflect products corrnnonto the Researcher and Wildlife Researcher series. Those products, in my opinion, are: :l. a succinct and snecific definition of the problem. Usually in research this is a statement of the research objectives or hypotheses; b. the rese:J.rchplan. This plan may he either formal (written) or informal, but all good research proceeds from some kind of plan ( ven case hi -tory or library research); �188 c. the research action program. By this I mean doing the research according to plan. It is not a tangible product, but it certainly is a necessary accomplishment from Hhich products Hill £1oH; d. the application of the research. The end product of all research should be a mow ledge or information package. iA/henthe research is funded by a client, the terminus of that research should be syntheses of that research to the larger body of existing mowledge on the subject and recommendations of what this information might mean or how it might be used by the client to meet the client's objectives and goals. 3. The Behavior Factors need to be structured so that they become subsets of the Results Factors. The structure of the appraisal system needs to show how behaviors are job relevant or how they contribute toward accomplishments and/or products. Behaviors of employees that are not relevant to job accomplishments or products are none of the agency's business. 4. Department of Personnel needs to Hork with state agency directors to develop supervisor training programs that are aimed at providing consistency among supervisors in the ways they assign work and appraise performance. S. Departffi8ntof Personnel needs to get to work to develop alternatives or improvemellts to the current Civil Service system to make it easier and quicker to discipline recalcitrant employees and supervisors. Currently there is a great deal of cynicism among the employees I mow. They are cynical because they believe that no matter how good the personnel appraisal systems are, the critical elements'are the employees. If they want to perform, they find ways to do it in spite of poor systems. Likewise, if an employee does not want to perform, he can find ways to do it in spite of good systems. 6. I would prefer to see general performance appraisal systems rather than specific ones because the general systems allow me sufficient flexibility to fit the system to the individual. Specific systems force the supervisor to develop depersonalized systems. We are so concerned with being fair and equitable that we forget the major function of any appraisal or testing system is to discriminate among employee performance. The key to that discrimination process is to develop objective measures of employee production and/or accomplishment. The bottom line, however, is accomplishment or production. The innovative supervisor is one who can motivate each employee individually to meet his/her potential. To do this i_tis absolutely essential to retain flexibility in our work assignment/performance appraisal systems. 11 cc: Lcs Canges �189 Relationships between Results Factors and Behavior Factors BERWlORS = ACTIVITIES RESULTS = OR APPLICATION OF SKILLS TO MEET SPECIFIC PRODUCTS OR ACCOi-IPLISHMEi\lJS SPECIFIED OBJECTIVES BY OBJECTIVES 1HE REL\TIONSHIP BETIVEEN BEHAVIORS AND RESULTS IN THE ORGANIZATIONAL SETTING IS THAT BEHAVIORS ARE USED TO EFFECT RESULTS. RESULTS MAY ALSO BE NESTED SO THAT ONE RESULT MAY FACILITATE AN01HER (E.G. RESULTS 1 MAY BE NECESSARY BEFORE RESULT 2 C~~ BE PRODUCED - IN THIS CASE, RESULT 1 COULD FUNCTION AS A BEHAVIOR IN EFFECTING RESULT 2). BEHi\VIOR 1 BEH!\VIOR 2 BEHAVIOR 3 BEHAVIOR 4 ~ \ ) <: Blli!\VIOR 2· \ BEHWIOR 5 RESULT 2 \\ BEHAVIOR 6 tl· BEH~VIOR 7 BEH;\VIOR 1 _ --------_j BEHt\VIOR 3 RESULT 3 ~ BEI-liWIOR 5 BEHAVIOR 7 BEHt\VIOR 2 l3EHAVIOR 4 \'vi BEHAVIOR 6 II --------..;\ Jj V// ~ __ -------_,) RESULT 4 ~ v \ J �190 PROGRA\! ADivlI:HSTRATION RESULTS FACTORS BEH~VIOR FACTORS PROBlliv! .'\i'iALYSIS RESULTS FACTORS BEHAVIOR FACTORS PLANNE\G, ORG\NIZING, RESULTS FACTORS AND COORDINATING �191 BEHAVIOR FACTORS RESE<\ROI ACTIO:.J PROGRt\M RESULTS FACTORS BEHA.VIOR FACTORS COl'-~'lUNICAnONS RESULTS FACTORS BEHWlOR fACTORS �192 PERSO?'J:"IJ:L~l~'\AGDlE\'T :\i\JD INTERPERSONAL RELATIONSHIPS RESULTS FACTORS BE{~VIOR FACTORS �193 Restructuring of Results and Behavior Factors according to a logical approach relevant to Scientific Research. RESULTS FACTORS (ACHI~ffi~TS/PRODUCTS) A. PROBLEM IDENTIFICATION AND DEFINITION B. RESEARCH PLAN C. RESE<\RCH PLAN IMPLH-IENTATION (ACTION PROGRAM) D. RESE<\RCH APPLICATION BEHAVIOR FACTORS (ACTIVITIES/SKILLS) A. RESEARCH SKILLS (COr-.IPETENCE) B. COvJt-1UNICATIONSKILLS C. PERSONNEL MA.t\JAGEMENT SKILLS D. ORGANIZATIONAL SKILLS E. INTERPERSONAL RELATIONSHIPS F. RESEARCU ADMINISTRATION SKILLS (HUlv1ANRESOURCE !v!AJ\JAGH1EJ\!,SKILLS) �194 RESULTS FACTOR (PRODUCTS OR ACCOMPLISI-IMEJWS) A. PROBLBl IDENTIFICATION AND DEFINITION Identifying, interpreting and defining the problem in terms that outline the contribution research can make towards the problem solution and in terms that give direction to the research process. Includes: Identifying problems accurately, hypothesizing the major problem elements and hypothesizing the interrelationships of problem elements. Identifying problem elements and problem element relationships where additional research is needed. Defining the researchable problem elements and problem element relationships in ways that give direction to the research (e.g. research hypotheses, objectives, goals) -. . Identifying \~hat is already known about the research problem in relation to its solution. Identifying what needs to be learned to solve the problem. Establishing priorities among research problem elements so that the most critical elements (i.e. those that are likely to contribute most to the problem solution) are addressed first. B. RESEARCH. PLAN Developing a research strategy that maxlmlzes the probability that the research will solve the problem or test the hypotheses. Includes: Developing unambiguous hypotheses, objectives or goals and establishing the relevance of these hypotheses, objectives, or goals to the research problem. Designing the research strategies or experiments that will be employed to meet the objectives, attain the goals or test the hypotheses. Selecting the data analysis procedures that will be used to objectively qualify the research results. Defining tolerable accuracy and precision limits. Establi shing research schedules and deadlines. Estimating requirements of agency fiscal and human resources to conduct the research. �195 B. RESE~CH PLJ'u\J (continued) Reviewing the research plan with qualified peers to improve the research strategy and enhance the probability the research will meet the planned objectives or test the predictions of the hypotheses. Adjusting the research strategy as new developments require. C. RESEARCH PLA.i\J Il\1PLEMENTATION(ACTION PROGRAM) Applying the agency's resources according to the specifications of the research plan to effectively and efficiently meet the research objectives or test the research hypotheses. Includes: Gathering data that are relevant to the research problem. UtiliZing appropriate sampling techniques to gather unbiased data. Doclmenting data clearly and in ways that are organized to maximize efficiency and accuracy of data processing. Analyzing data using appropriate methods and procedures as outlined in the research plan. Interpreting data logically so the interpretations are compatible with existing theory or suggest new theory or modifications of existing theory. Synthesizing the research results into the larger knowledge pool. Modifying existing techniques or developing new' techniques to accomplish the research and/or agency objectives and goals. D. RESEARCH APPLICATION Participating in the process of applying research results to the agency's problems. Includes: [dentifying potential applications of research results to agency operations. Developing alternative strategies for implementation research results into agency operations. Publicizing research results so they arc available to the general research commun.ity and research clientele. Serving as an expert consultant to apply information or knowledge in subject matter specialties to agency problem solving activities. �196 D. RESE\RCII APPLICATION (continued) Training other agency personnel in specialized methodology or in applications of information and/or knowledge to accomplish agency objectives and attain agency goals. �• 197 I BEHAVIOR FACTOR (ACTIVITIES OR SKILLS) A. RESEARCH SKILLS (COMPETENCE) Improving professional/technical Includes: competence related to one's area of expertise. Continuing educational growth to maintain current knowledge and skills and acquire additional expertise within the constraints of agency policy. Maintaining a working-level awareness of the status of the knowledge and state of the art in one's area of expertise. Utilizing state of the art techniques and equipment relevant to one's area of expertise to accomplish research objectives. Recognizing the nature and extent one's limitations to know when to calIon expertise of specialists to facilitate the attainment of the research objectives. Developing creativity and analytical skills. B. Co.\r-1UNICATIO~ SKILLS Communicating research results and/or proposals effectively and accurately to enhance agency problem solving and to advance knowledge. Includes: Defining the communications message clearly and concisely . . Structuring the message to audience. Choosing the media appropriate to the message and the audience. Conceptualizing and developing abstract ideas clearly and precisely so the message is accurately and effectively conveyed. Publishing research findings in professional journals appropriate to the field. Presenting papers at professional meetings appropriate to the field. Constructively criticizing research proposals and/or results of others in the field upon request. Testifying before judicial, legislative, and/or executive hearings or proceedings. �198 c. PERSONNEL !'-!AJ\JAG&lENfSKILLS (HUMAN RESOURCE MANAG&IENI' SKILLS) Improving employee productivity, competence and morale. Includes: Increasing employee efficiency, responsiveness and performance level. Selecting and retaining productive personnel. Establishing training strategies, providing opportunities or conducting training to enhance professional/technical competence of subordinate employees. Implementing corrective strategies to improve the performance of below-standard employees and when necessary, initiating disciplinary actions against recalcitrant substandard performers. Defining and communicating clearly the ,vork assignments and responsibilities for which employees will be held accountable. Improving morale by identifying and correcting factors which contribute to poor morale. Matching work assignments to employee interests and abilities as closely as agency objectives, goals and policies permit. Developing and implementing procedures and systems to involve employees in decision processes which affect their work assignments and work environments. Developing and implementing systems to evaluate employee performanse objectively and fairly. Effecting affirmative action goals and programs of the agency. Monitoring employee performance in relationship to agency goals and objectives. Developing and implementing work safety programs. D. ORGfu\JIZATIONAL SKILLS Structuring one's own work and the work of subordinates in ways that maximi ze productivity and efficiency in the accomplishment of agency goals and objectives. Includes: Organizing and scheduling actIVItIes according to priorItIes so that time is allocated proportional to the ilnportance and difficulty of the task. Developing systems and procedures that maximize efficiency without compromising quality standards. �199 D. ORG~~I~~TIONAL SKILLS (continued) Coordinating with other individuals and groups to establish and accomplish objectives. Developing contingency plans to cope with anticipated potential problems. Monitoring progress towards objectives. Adjusting priorities, goals, plans, schedules as contingencies dictate. E. INTERPERSONAL RELATIONSHIPS Interacting \vith others in ways that facilitate the achievement of agency objectives. Includes: Developing and maintalnmg constructive working relationships with publics that affect research objectives. Leading by example to encourage excellence in subordinates and/or co-workers. Enhancing the work environment to increase work quality and quantity. Confronting interpersonal conflicts in a timely and effective way to maintain a harmonious and productive work environment. ~~intaining a receptivity for and tolerance of ideas, attitudes and individuality of subordinates and co-workers, and a sensitivity to their personal problems. Using skill, tact, and sophistication in interviews with subjects, clients, experts and others to gather information. Nai.ntairring a positive professional image. F. RESEARCII AD~fINISTRATION SKILLS Employing agency resources, personnel, and service functions effectively to accomplish agency goals and objectives. Coordinating the planning, implementation, monitoring and adjusting functions of the research process to accomplish agency goals and objectives. ~~aging allocated agency resources to accomplish agency goals and objectives to effect the greatest return to the agency from resources invested. Defining and delineating the boundaries of your decision space so you know when to involve higher level administrators in your research administration assignment. �200 APPENDIX B Program Description of the Planning System for the Division of Gal1\e,Fish and Parks 1971 February 1971 (First Edition) �B-2 201 -- CONTENTS -- Introduction Need for a Planning System Justification for a Planning System Who Team Approach of the Planning System What Goals of the Planning System How Program Des<;:riptionof the Planriing System 10 Orientation of the Planning Team 2. Orientation of the Division 3•. Mea suremen t 4. Test 5. Documentation When Schedule and Parameters of the Planning System Appendices �202 -- WHY -- Introduction Planning is an act we all do. That is a matter of fact. However, there are differences in our ability to plan even for routine events. Why is this so? It has been determined that knowledge and experience helps, but that judgement plays the most important role in good planning. But for a planner to render good judgement, his ways must be analytical and systematic. But to be analytical, one should be inquisitive; to be systematic, one should deal in systems. What is a system?· The world is a system and everything in it, its sub-system. Every system is also a sub-system of another system or systems. The purpose of a system is to satisfy. the requirements of a larger system that it services. Therefore, anything or anyone that attempts to survive within itself will eventually' destroy itself or be destroyed. Why are these statements important? This is the basis for understanding systems or the analysis of systems, an important discipline of planning. Consider the following example? If the Division of Game, Fish and Parks determines its annual needs on the -.basis of how many people it employs, how much equipment it has and how much space it has, it will neither effectively meet demands for increases in service nor adjust efficiently to decreases in demands for service. Thus * if this Division does not recognize it is a sub-system of larger systems * if this Division does not plan for the future on the basis of the needs of the larger systems * if this Division does not study the planning of the larger systems, and * if this Division does not try to influence the planning of the larger systems, we will 1. 20 operate inefficiently and ineffectively be victims of our own destiny The questions then are: How do we plan to operate efficiently and effectively? How do we plan for the future? �203 THE LARGER RENEW( ABLE ESOURCE PRESSURE, ~====~V PLANNING SYSTEM I., DIVISION ADMISTRA. , TION I �204 Need for a Planning System Increasing public pressure for hunting, fishing and outdoor recreation opportunities has and will continue to exert in the forseeable future tremendous demands on" the fish and wildlife resources of Colorado. Fish and wildlife are renewable resources, provided that natural environments can be preserved and managed. Therefore, this Division will implement a goal-oriented plan- ning system so policy decisions generated by the Governor, the General Assembly, the Executive Director (Department of Natural Resources), and the Commission and administration of this Division, will satisfy and best serve the . demands" of outdoor enthusiasts and the general public consistent with our supply of natural resources. �205 Justification for a Planning System The state policy shall be to encourage, by every appropriate means, the full development of the state's natural resources to the benefit of all of the citizens of Colorado, and shall include, but not be limited to, creation of a resource management plan to integrate the state's efforts to implement and encourage full utilization of each of the natural resources consistent with realistic conservation principles. The governor, through the executive director of the Department of Natural Resources, shall develop and direct the resource management plan and shall,be responsible for negotiations with the federal government in all resource and conservation matters (C.R.S. 1963, section 3-15-3). It is hereby declared to be the policy of the state of Colorado that the fish and wildlife and their environment, and the natural, scenic, scientific, and outdoor recreation areas of this state are to be protected, preserved, enhanced, and managed, for the use, benefit, and enj9yment of the people of this state and visitors to this state. It is further declared to be the policy of this state that there shall be provided a comprehensive program of outdoor recreation in order to offer the greatest possible variety of outdoor recreation opportunity to the people of this state and its visitors, and that to carry out such a program and policy there shall be a continuous operation of planning, _acquisition, and development of outdoor recreation lands, waters, and facilities (C.R.S. 1963, section 62-1-2). �206 -- WHO -- Team Approach of the Planning System A team approach to develop a planning system is being taken to gain representation and coordination of our three missions: 1) land and water management, 2) wildlife management and 3) fish management. I Larger Systems I .' ,~ Division of Game, Fish and Parks , ., . '. Operational Management Plans Fish Planning Specialist ( P+annin~ or Coordinator Land & Water Planning Specialist Planning System , ; .•.. or Game Planning Specialist Resource Management Plan - External Planning Efforts t Comprehensive State & Federal Outdoor Plannfng Planning �207 -- WHAT -- Goals of the Planning System Develop a planning system that maximizes the effectiveness of the Divison of Game, Fish and Parks in fulfilling its mission. The objective of this goal is the preparation of a resource management plan that satisfies the legislative charge of the Reorganization act of 1968 (C.R.S. 1963, Section 3-15-3). Fit pianning system into the real day to day world of the Division. tives Objec- of this goal wi Ll, be operational management plans for the various fish and wildlife species, and the Division's programs and management areas (see Apendix A). �208 GOALS Develop Planning System Fit To Organization OBJECTIVES Resource Management Plan Operational Management Plans PERFORMANCE CRITERIA Units of Environment Preserved Number of Activity Days Number of Fish & Wildlife Utilized �209 -- HOW -- Basically, the planning system will influence multi-level decision making. Schematically and in a very simplified form, the framework appears as: Planning Team' -. Planning System r-. Decision Making Alternative . Alternative ~ ,n ,., ~ , . . 4J. ••• r First-Level Goals Alternative Ultimate A ....• . -Objec t Lves The arrows within this schematic represent functions of the planning system, whereas the boxes establish the system's structure. For example, the arrow between jPlanning System/ and /Decision Makers/ represents a number of functions or processes, including: How the planning system fits into the organization. How it communicates within the organization. How it monitors within the organization. �210 • Program Description of the Planning System Program 1 Orientation of the Planning Team-~Since the decision was made to convert "biologists" into "planners" rather than convert "planners" into "biologists", a considerable amount of time and cost will be necessary for proper orientation to the technologies of professional planning. Subprograms: a. Methods--determine how the planning system will be designed and determine which tools will make it operate more efficiently and effectively. * P.P.B.--Programs, Plans and Budgets * P.E.R.T.--Performance, Evaluation and * Network Analysis--Priority Allocation * Cost/Benefit Analysis * Cost/Efficiency Analysis Review Techniques (Critical Path Method) b. Coordination--determine who and ±f other planning systems (Land Use Commission, State Planning Office, Colorado Comprehensive Outdoor Recreation Plan, Feds., et al.) are 'compatible for mutual efficiency, effectiveness 'and results. c. Problem identification--this will be to gain an overall perspective which wil1 emphasize which route planning system should follow. Program 2 Or Len t a'tLon of the Division--a continual relationship between the planning system and the decision makers. Subprograms: a. Justification--why the planning system should exist and its proper role. b. Organization--establish the planning teams responsibility and authority (i.e., where it belongs in the organization). c. Communication--how planning system will be incorporated into the Division. d. 1. Persuasion (selling the planning system to those concerned). 2. Mutual Education (the planning system will guide decision makers and will instruct the Division as to how the planning system of the decision makers operates). 3. Politics (identify and if necessary alter the power structure of the Division). Implement planning system into the Division. �211 r----- I De c Ls Lon ~~ I •.. TP" Makers Team , Division I I . I I I Planning -;. Goals ---I ...• "'IIj I I L I I Alternatives .d '"<I I I I J �212 Program 3 Measurement--identify, classify, predict and monitor the structures and functions of the Division. A planning effort needs to be precise, accurate and general. If estimates are imprecise, -t.hen together all es t Lmat es may be intolerably vague. If estimates are, inaccurate, then the plan will tell managers the wrong things. If estimates are not general, the plan will only work in specific context and time. Subprograms: a. Identify decision makers--those people Who produce change in an organization. b. Identify the customers-1. 2. 3. 4. Who are they? What do they want? When do they want it? How do they want it? c. Determine appropriate alternatives--Planners need to encourage radical' viewpoints because when planning appears to follow tried and true procedures, then planning is in danger of becoming useless. The research and development section will be relied on to assist in creation of new alternatives. d. Identify the first stage goals--the purpose of setting first stage goals (missions) is to put the short-range objectives into proper perspective (Le. determine which are the most important objectives). First stage goals, when well written, should constitute policy of this Division. Therefore, first-stage goals need to be stated quite specifically or else their relationship to objectives becomes lost and their role as ~ntegrating function becomes meaningless. e. Identify ultimate objectives--objectives must be stated in very specific terms. f. Measuring the effectiveness of each first-stage goal for the ultimate objectives--this activity allows for understanding of the environment of the Division. The environment being the systems that exist outside of the Division which can be influenced but not controlled. Prediction is a necessary tool in attempting to understand the environmento g. Select optimal alternative--this should be done only after all of the preceding subprograms have been evaluated. �213 Program 4 Test--verify plans. Subprograms: a. Modelling and simulation--a method of analyzing the behavior of any system by identifying its structure and function. It is a technique which will be very useful in analyzing the effects of programs on fish and wildlife species. b. Reviewing----.--allowing the plans to be scrutinized by the practical approach of experienced Division personnel using their intuition, leadership and brilliance. c. Controlling the plan--once a plan is imple.mented, there must be feedback of information about the operation of the plan so changes can be made if needed. Prqgram 5 Documentation--Initially, as a product of the planning system, there will be three types of documents published. Each type of document will be written for a specific purpose. Subprograms: Resource Management Plan a. , b. Digest--this concise document will be written in a popular and illustrative style for the sole purpose of introducing the Resource Management Plan to the general public. It should be well advertised with unlimited distributiqn. Compendium--this document will include all categories of the Division's responsibilities (i.e. species, groups of species and programs). It will consist of several volumes which will outline the problems of the Division All background data will and the broad plans to rectify those problemso be presented in these volumeso The purpose of the compendium will be twofold: (1) disseminate additional information pertaining to the digest, and (2) a documented policy standard of the Division. Distribution should be limited to: (a) libraries, (b) other resource management agencies, and (c) Division employees. Members of the general public can obtain these documents, but only upon request (purchase). ?Jeration Management Plans a. There will be operational management plans written for each major species, group of species, program and management property. Documents will be formulated on a iter&tive basis consistent with budgets, change in policy or change in legislation. Each plan will be concerned with the current operational program and the immediate future. Specific objectives and the plans to accomplish these objectives will be included and related to budget allocations. �214 -- WHEN -- Schedule and Parameters of the Planning System At this time it is not possible to give an objective, universally valid estimate of the amount of activity required for each of the above mentioned subprograms. (e.g. How much persuasion and education will be needed to attain necessary understanding and positive acceptance of the planning system?). Also, only relative estimates can be offered on the cost and technology available to accomplish each subprogram. However, when time permits, each subprogram will be described in detail and assigned performance measures so decisions can be made as to there relevance. II Time Sub:erograms Cost.!1 Methods Coordination Problem Identification .IustLf Lca t Lon Organization Persuasion Mutual Education Politics Implementation Decision Makers Customers Alternatives Goal~ Objectives Effectiveness Optimal Alternatives Modelling & Simulation Anti-planning Control Digest Compendium Operational Plans H M H L Continual Periodically H M H M M L L Continual February July .!/ H Technology-- 1971 H M f1 M M H Continual M M M M L L M M M M L M L L M M H H M H M 1974 Continual Continual H M M M 1973 December M Continual Continual Continual Periodically Periodically Continual Continual Continual Continual M 1972 Continual means High, M means Moderate, L means Low. Periodically Periodically Continual �215 Definitions Missions--Statements related to purpose: they may contain who, why, when, how and what. Emphasis should be on what. Goals--something toward which an effort is directed. Theoretically goals should never be achieved, although every effort is made (e.g. improve efficiency, improve effectiveness). Objectives--something toward which an effort is directed, but it is feasible to attain (e.g. ~ efficient, most effective). Performance Measure (Criteria) -- measurable levels of success which are set for objectives. �216 Mission of the Division of Game, Fish and Parks Listed are four alternatives that may aid in establishing the first-level goal (mission) of this Division. When this decision is made, lower-level goals can be set for the various programs, management properties, species, and so on. Each subsequent goal will be cons i.st ent; with the Division's primary mission. Alternative 1 The mission of the Game, Fish and Parks Division is to provide the maximum possible hunting, fishing and parks related activities for the people of Colorado and for the visitors to this state. . Definitions Outputs Preserve and produce the fish and wildlife species that will provide maximum sustained human benefits Animals produced Animals utilized man-days use Assure survival of all fish and wildlife species in a natural state Units of habitat Preserve and enhance outdoor parks and recreation areas in Colorado for the purpose of providing a variety of human recreation activities Man-days use Alternative 2 To preserve all species of fish and wildlife in Colorado through preservation of their environment, and to manage the resulting animal populations and their habitat to produce the greatest public benefit within the prevailing natural and social limitations. Definitions and Outputs Preserve (animal) - To maintain a species at its greatest possible density and distribution with emphasis upon increasing the density and distribution of rare and endangered species. Fish and Wildlife - As defined in 62-1-3 of Colorado Revised Statutes. Environment - The eco:logical elements in an area of adequate si.zeto sustain a stable animal population. Elements and size vary according to animal species considered. �217 Manage - To manipulate the appropriate human and ecological elements to affect a predetermined result. ;-- .. Habitat - The current condition of an animal's environment. Public - All people, resident and. nonresident, but with emphasis on Colorado citizens. Benefit - Any human activity which contributes to an individual's health, happiness and/or knowledgeo Natural - Of other than human origino Social - Of human origin. Alternative 3 Provide maximum public benefits consistent with demand, by preserving, managing and restoring Colorado's diverse land and water environment, and by maintaining continually abundant fish and wildlife resources through proper conservation practices. Definitions Outputs Public Benefits Activity Days Environments Surface Units Fish and Wildlife Resources Animals utilized per Activity Days and Surface Units . \ Alternative 4 The special mission of the Game, Fish and Parks Division is to preserve, enhance, and manage fish and wildlife and their environment, and all recreation areas for the greatest public benefit. Definitions Outputs To cummunicate information essential to an optional level of public understanding of the benefits obtained. To increase density of rare and endangered species. To remove surplus fish and wildlife from all populations through fishing and hunting. Optimize supply and demand for the benefit of both fish and wildlife and the public. Provide maximum recreation in all parks within Colorado. Numbers of fish and wildlife produced for consumptive and non-consumptive utilization. �218 Problem Identification Please list, in order of decreasing importance three (3) problems which are of major importance to this Division. This will aid in giving the planning system proper perspective and orientation. �'') ( '-" / '. s.IR&ITGIC PLAtl-MAJOR PROGRA~': SIRATEG I C PLANPRQGJlllliS.: SIRlUEG J C PLAfl- QBJEillID: SPORT GAt1E NONCERVID RESEARCH 1. BIGHORN SHEEP 2. PRONGHORN ANTELOPE 3. MOUNTAIN GOAT 4. BLACK BEAR 5. MOUNTAIN LION ."." I'T1 1. To 2. 3. 4. 5. QR~IZATION8L UNIIW!1PONEtU OPERATI Drl PLANS » PROVIDE INCREASES OF 50% IN BIGHORN SHEEP POPULATIONS, 84% IN HARVEST, 29% IN HUNTERS, AND 39% IN RECREATION DAYS BY 1988 COMPARED TO 1980 BASE-YEAR LEVELS. To PROVIDE FOR INCREASES OF 5% IN PRONGHORN ANTELOPE POPULATIONS, 20% IN HARVEST, 18% IN HUNTER NUMBERS, AND 17% IN RECREATION DAYS AS COMPARED TO 1980 BASE-YEAR LEVELS. To MAINTAIN MOUNTAIN GOAT POPULATIONS AT ABOUT CURRENT LEVELS WHILE PROVIDING FOR INCREASES Of 28% IN HARVEST, 40% IN HUNTER NUMBERS AND 49% IN RECREATION DAYS BY 1988 COMPARED TO 1980 BASE-YEAR LEVELS. To MAINTAIN ABOUT THE SAME LEVELS OF BLACK BEAR POPULATIONS, HARVEST, HUNTERS, AND RECREATION DAYS THROUGH 1988 COMPARED TO 1980 BASE-YEAR LEVELS. MAINTAIN ABOUT THE SAME MOUNTAIN LION POPULATION LEVEL WHILE PROVIDING AN INCREASE OF 34~ IN HARVEST, 30% IN HUNTER NUMBERS, AND 27% IN RECREATION DAYS BY 1988 COMPARED TO 1980 BASE-YEAR LEVELS. WILDLIFE 1. 2. 3. 4. 5. 6. RESEARCH - BIG GAME HONCERVID BIGHORN SHEEP RESEARCH PRONGHORN ANTELOPE RESEARCH r10UNTAIN GOAT RESEARCH BLACK BEAR RESEARCH PUMA RESEARCH RESEARCH ~UPPORT Z o >< n SECTION _. N \.0 �, "\ \ , c...._ , ') N N o .,',' OPERATION PLAN COMPONENT - BIGHORN SHEEP RESEARCH 1. PROBLEM - THE NUMBER ONE PRIORITY STRATEGY FOR THE BIGHORN SHEEP PROGRAM IS TO ~INCREASE CARRYING CAPACITIES OF BIGHORN SHEEP HABITATS ON PUBLIC LANDS THROUGH CONTROLLED BURNING, CHAINING, FERTILIZING, TIMBER MANAGEMENT, AND SIMILAR METHODS." INCREASING THE CARRYING CAPACITY OF BIGHORN RANGES ULTIMATELY DEPENDS ON CORRECTING SPECIFIC DEFICIENCIES IN HABITATS. HOWEVER, BEFORE THOSE INADEQUACIES CAN BE REMEDIED, THEY MUST BE IDENTIFIED. MANY OF THE PROBLEMS WHICH AFFECT SURVIVAL AND REPRODUCTION OF BIGHORN SHEEP POPULATIONS CAN BE EXPLAINED BY POOR NUTRITION; LOW RESISTANCE TO DISEASE AND PARASITISM, REPRODUCTIVE FAILURE OF FEMALES, AND HIGH r~ORTALlTY OF LAMBS CAN P,Ll RESULT FRor~ r~ALNUTRITION. DESPITE THE APPEAL OF THIS EXPLANATION FOR POOR PERFORMANCE OF BIGHORN POPULATIONS, THERE IS NO COMPELLING EVIDENCE TO SUPPORT OR REfUTE IT. CONSEQUENTLY, IT REMAINS UNCLEAR WHETHER IMPROVING FORAGE SUPPLIES iN BIGHORN HABITAT, A PRINCIPLE OBJECTIVE OF MOST HABITAT MANAGEMENT STRATEGIES, WILL OFFER BENEFITS COMMENSURATE WITH ITS COST. THE ABILITY TO IMPROVE BIGHORN RANGE BY MANIPULATING VEGETATION WILL BE SUBSTANTIALLY ENHANCED BY DETERMINING WHETHER NUTRITION LIMITS BIGHORN POPULATIONS. IN THE ABSENCE OF THAT DEiERMINATION. HABITAT MANAGEMENT FOR BIGHORN MAY BE MISDIRECTED AND INEFFECT! VE. 2, SI8IUS - CURRENT KNOWLEDGE ABOUT BIGHORN SHEEP HABITAT REQUIREMENTS AND CARRYING CAPACITY OF IMPORTANT HABITATS IS LIMITED TO UNTESTED HYPOTHESES. HYPOTHESES ARE CONFLICTING. ONE SCHOOL OF THOUGHT HYPOTHESIZES THAT BIGHORN SHEEP POPULATIONS ARE LIMITED BY EPISODIC MORTALITY DUE TO RESPONSES TO STRESS. A SECOND SCHOOL HYPOTHESIZES THAT BIGHORN SHEEP POPULATIONS ARE LIMITED BY CHRONIC MORTALITY (PRIMARILY OF LAMBS) INDUCED BY PARASITIC LUNGWORM INFECTIONS. A THIRD SCHOOL OF THOUGHT HYPOTHESIZES THAT HABITAT DEFICIENCIES LIMIT BIGHORN SHEEP POPULATIONS, BUT THE CRITICAL FACTOR OF HABITAT IS COVER. A FOURTH SCHOOL HYPOTHESIZES THAT BIGHORN SHEEP POPULATIONS ARE FUNDAMENTALLY LIMITED BY INADEQUATE FOOD RESOURCES AND THE OTHER APPARENT LIMITATIONS ARE SYMPTOMS OF NUTRITIONAL DEPRIVATION, �r " 3. __ I ''] \ RESEARCH OBJECTIVE - TEST THE HYPOTHESIS --'-THROUGH NUTRITIONAL DEFICIENCIES. THAT HABITAT LIMITS BIGHORN SHEEP POPULATIONS 'MEDIATED PRIMARILY 4. nlPLEr1ENTATIorl - EVALUATING THE IMPORTANCE OF NUTRITION IN BIGHORN POPULATION DYNAMICS WILL INVOLVE FOUR PHASES OF RESEARCH. FIRST, NUTRITIONAL REQUIREMENTS FOR MAINTENANCE AND REPRODUCTION WILL BE ESTIMATED. SECOND, THE CAPABILITY OF BIGHORN TO ASSIMILATE AND METABOLIZE NUTRIENTS FROM TYPICAL FORAGES WILL BE ASSESSED. THIRD, THIS INFORMATION WILL BE SYNTHESIZED INTO A PLAN FOR FORAGE ALLOCATION FOR BIGHORN SHEEP AND RECOMMENDATIONS FOR HABITAT MANIPULATION TO IMPROVE AND ENLARGE BIGHORN RANGES. THESE FOUR PHASES WILL BE IMPLEMENTED IN THE FOLLOWING STEPS: A. DETERMINE ENERGY REQUIREMENTS OF BIGHORN FOR BASAL METABOLISM, ACTIVITY, AND THERMOREGULATION. USE THESE ESTIMATES TO PREDICT SEASONAL MAINTENANCE REQUIREMENTS. B. DETERMINE MAINTENANCE PROTEIN REQUIREMENTS FOR BIGHORN SHEEP CONSUMING FORAGE DIETS. C. DESCRiBE THE PHYSIOLOGICAL ABILITY OF BIGHORN SHEEP TO DIGEST AND METABOLIZE ENERGY AND PROTEIN FROM TYPICAL GRASS AND BROWSE DIETS. D. DETERMINE EFFECT OF PLANE OF NUTRITION ON REPRODUCTIVE PERFORMANCE IN BIGHORN SHEEP EWES. ESTIMATE ENERGY AND PROTEIN REQUIREMENTS OF EWES NECESSARY FOR PRODUCTION OF VIABLE LAMBS. E. DETERMINE ENERGY AND PROTEIN INTAKE OF FREE-RANGING BIGHORN SHEEP ON WINTER AND SUMMER HABITATS. F. SIMULATE CARRYING CAPACITY OF BIGHORN SHEEP WINTER AND SUMMER HABITATS. 5. BENEFITS - THIS RESEARCH WILL FACILITATE SELECTION OF APPROPRIATE HABITAT MANIPULATIONS INCREASES IN BIGHORN SHEEP POPULATION SIZE TO ACCOMMODATE INCREASES IN HUNTER HARVESTS, AND RECREATION DAYS. TO ACHIEVE DESIRED HUNTER NUMBERS, N N �.. ~ N' N N \ >. : )' DIVISION FIVE FORM NO. PLAN STRATEGY 1. Detennine energy requirements for bighorn sheep for basal metabo l i sm, act iv i ty and thermoregula t ion to predict seasonal maintenance requirements. OI'EMTION crminc mai ntcnance protein requirements for bighorn sheep consuming forage diets. Dct 3. Describe the physiological ability of bighorn sheep to digest and metabolize energy and protein from typical grass and browse diets. L Determine ef feet of plane of nut r it ion on reproductive perfor· mance in bighorn sheep ewes, WILDLIFE OPERATIONS PLAN (1982-87) ACTIVITIES /I.N:\LYSIS PREPARED BY R.B. Gill, N.T. Hobbs 2. YEAR OF TITLE cosr CENfER SS03X Noncervid Research Bighorn Sheep Research ACTIVITY PROGRAM PRODUCT ANALYSIS Bighorn Sheep DEADLINE 1. Rear and train experimental animals. 3 ewes per year for 3 years 2. Prepare a detailed study plan. Approved Study Plan 1 1 1 31 3. Conduct research. Annual Budget Request Annual Report Publication 1 May Annually 31 July Annually 31 July 1984 SS03X Noncervid Research 1. Approved Study Plan 31 July 1984 2. Conduct research. Annual Budget Request Annua l Report Publication 1 May Annually 31 July Annua ll.y 31 July 1985 5503X Noncervid Research 1. Prepare a detailed study plan. Approved Study Plan 31 July 1984 2. Conduct research. Annual Budget Request Annua I Report Publication 1 May Annually 31 July Annually 31 July 1985 S503X Noncervid Research 1. Prepare a detailed study plan. Approved Study Plan 31 July 1985 2. Conduct research. Annual Budget Request Annual Report Publication 1 May Annually 31 July Annually 31 July 1987 Prepare a detailed study plan. October 1983 October 1984 October 1985 July 1983 �' .....•. ~ " v TITLE Bighom OPERATION Sheep Research p[j\,\j STRATl:GY 5. Determine energy and intake of free-ranging bighorn sheep 011 winter and summer habitats. 6. Simulate carrying capac i t y of bighorn sheep winter and summe r hab i tat.s. (continued) AcrrVITY COST CENTER SS03X Noncervid Research SS03X NOllcervid Research PRODUcr ANALYSIS DEADLINE 1. Prepare a detailed study plan. Approved Study Plan 31 July 1985 2. Conduct Research Annual Budget Request Annual "Report Publication 1 May Annually 31 July Annually 31 July 1987 1. Conduct simulations. 2. Estimate carrying capacity. Annual Budget Request Annual Report Publication 1 May Annually 31 July Annually 31 July 1987 3. Implement results. Implementation Meetings with Regional and Staff Personnel 1 August 31 March 1987-88 \ N N W �fL'1 N N ~ OPERATI ON PLAN CO~lPONENT - PRONGHORN AiITELOPE RESEARCH I, PROBLEt1A, B. 2, THE NUMBER ONE STRATEGY FOR THE PRONGHORN ANTELOPE PROGRAM IS TO "PROVIDE INFORMATION ON ANTELOPE HABITAT NEEDS FOR USE IN LAND MANAGEMENT DECISIONS AND SEEK MITIGATION FOR ANTELOPE LOSSES WHEN iMPACTS ARE UNAVOIDABLE," PRONGHORN ANTELOPE RESIDE ON PRIVATELY OWNED LANDS MORE SO THAN ANY OTHER BIG GAME SPECIES. THEREFORE, THE KEY TO INCREASING PRONGHORN POPULATIONS BY 5% IN THE NEXT 5 YEARS WILL DEPEND, IN PART, ON ENHANCING PRODUCTIVITY OF PRIVATE LANDS FOR PRONGHORNS, DOMESTIC L~VESTOCK PRODUCTION CONSTITUTES A MAJOR USE OF THESE PRIVATE LANDS. RESEARCH IS REQUIRED WHICH WILL EVALUATE LIVESTOCK GRAZING PRACTICES THAT HAVE POTENTIAL TO INCREASE LIVESTOCK YIELDS AND TO ENHANCE PRONGHORN HABITATS AT THE SAME TIME, THE NUMBER TWO PRIORITY STRATEGY FOR THE PRONGHORN PROGRAM IS TO "IMPRBVE OUR ABILITY TO MATCH HARVEST REGULATiONS WITH AVAILABLE ANTELOPE POPULATIONS THROUGH BETTER COUNTS AND BETTER ANALYSIS OF COUNT DATA," INCREASED DEMANDS FOR FISCAL RESOURCES WITHIN DOW NECESSITATE IMPROVING THE EFFICIENCY AND EFFECTIVENESS OF EACH ACTIVITY. PRONGHORN CENSUS METHODS CURRENTLY IN USE ARE INTENSIVE AND EXPENSIVE, THE GREATEST PROMISE FOR INCREASING COST-EFFECTIVENESS OF PRONGHORN CENSUSING LIES IN COMBINING STATISTICAL SAMPLING TECHNIQUES AND POPULATION SIMULATION MODELING TECHNIQUES TO IMPROVE CENSUS ACCURACY; PRECISION) AND EFFICIENCY, STATUSA, THE COLORADO STATE UNIVERSITY AGRICULTURAL EXPERIMENT STATION IS PLANNING A SERIES OF STUDIES TO EVALUATE THE POTENTIAL OF VARIOUS GRAZING MANAGEMENT SYSTEMS TO LIVESTOCK PRODUCTION WITHIN SHORTGRASS PRAIRIE PLANT COMMUNITIES, THE OPPORTUNITY EXISTS TO DESIGN CONCURRENT EVALUATIONS OF THE EFFECTS OF THESE GRAZING EXPERIMENTS ON PRONGHORN HABITATS, CONSIDERABLE SAVINGS IN COSTS WOULD ACCRUE AS A RESULT COMBINING DOW's PRONGHORN RESEARCH PROGRAM WITH THAT OF THE CSU AGRICULTURAL EXPERIMENT STATION, B, RECENT TESTS WHICH HAVE APPLIED SAMPLING METHODS TO PRONGHORN CENSUS INDICATE THAT CENSUS EFFICIENCY CAN BE INCREASED AND CENSUS COSTS CAN BE DECREASED BY EMPLOYING SAMPLING STRATEGIES INSTEAD OF ATTEMPTING TOTAL ENUMERATIONS OF PRONGHORN POPULATIONS, HOWEVER, CENSUS ACCURACY IS STILL A MAJOR QUESTION. ESTIMATES OF PRONGHORN POPULATIONS GENERATED FROM TOTAL ENUMERATIONS, QUADRAT CENSUSES, STRIP CENSUSES, AND �, \ I, ''\ r- POPULATION SIMULATIONS DO NOT CONSISTENTLY ESTIMATES IS NOT KNOWN. 3, 4, COINCIDE. WHICH, RESEARCH OBJl~ A, EVALUATE THE POTENTIAL OF LIVESTOCK GRAZING SYSTEMS TO ENHANCE B, EVALUATE THE ACCURACY OF PRONGHORN CENSUS TECHNIQUES, IF ANY, OF THE METHODS PRONGHORN PROVIDES ACCURATE HABITATS, l11PLEMEflTLillOllA, STUDIES WILL BE DESIGNED TO EVALUATE THE EFFECTS OF LIVESTOCK GRAZING 'ON PRONGHORN HABITAT QUALITY. THE FOLLOWING RESEARCH PHASES WILL BE IMPLEMENTED: 1, TEST' THE HYPOTHESIS THAT PRONGHORN CAN AND DO MAXIMIZE PROTEIN AND ENERGY INTAKE THROUGH FORAGE SELECTION STRATEGIES, 2, DETERMINE THE EFFECTS OF LIVESToCK GRAZING SYSTEMS ON THE AVAILABILITY OF NITROGEN AND EN~RGY TO PRONGHORN IN SHORTGRASS PRAIRIE COMMUNITIES. 3. MODEL THE IMPACTS OF CHANGES IN AVAILABLE NITROGEN AND ENERGY THAT ARE INDUCED BY LIVESTOCK GRAZING SYSTEMS ON PRONGHORN NUTRITIONAL ECOLOGY. B. ACCURACY OF PRONGHORN CENSUS SYSTEMS WILL BE ASSESSED BY COMBINING MARK-RECAPTURE METHODS AND AERIAL SAMPLING METHODS. A KNOWN POPULATION OF MARKED ANIMALS WILL SERVE AS THE STANDARD BY WHICH TO COMPARE AERIAL SAMPLE COUNT PROJECTIONS. 5, BENEElI£ - A SYSTEM TO MORE ACCURATELY CENSUS PRONGHORN WILL RESULT, THIS WILL ALLEVIATE ONE OF THE MAIN WEAKNESSES OF THE PRONGHORN POPULATION MODELS AND FACILITATE MORE REFINED MANAGEMENT OF THE PRONGHORN RESOURCE, ASSESSMENT OF LIVESTOCK GRAZING PRACTICES ON PRONGHORN HABITAT WILL ENABLE THE DIVISION TO MAKE SOUND RECOMMENDATION IN LAND MANAGEMENT DECISIONS, IN ADDITION, INFORMATION WILL BE GAINED ON THE HABITAT FEATURES THAT ARE IMPORTANT FOR MITIGATION PURPOSES, N N V1 �( I "I '\ I ".' N' N 0' DIVISION FIVE FORM NO. PREPARED YEAR OF WILDLIFE OPERATIONS PLAN (1982-87) ACTIVITIES ANl\LYSIS BY R.B. Gill, T.~1. I'ojar Ol'ERATIO\,JPU\N STRATEGY TITLE COST CENTER PROGm1 Pronghorn Antelope Research ACTIVITY PRODUCT ANALYSIS Pronghorn Antelope DEADLINE l!.. 1. Test the hypothesis that pronghorn can and do maximize protein and energy intake through forage selection strategies. 2. 3, S503X NOllcervid Research animals. 1 October 1983 1. Rear and train experimental 2. Prepare a detailed study plan. Approved Study Plan 31 July 1983 3. Conduct research. Annual Budget Request Annual Report Publication 1 May Annually 31 July Annually 31 July 1985 Determine the effects of 5503X livestock grazing sysNOllcervid tems on the availability Research of nitrogen and energy to pronghorn in shortgrass prairie communities. 1. Prepare a detailed study plan. 2. Conduct research. Annual Budget Request Annual Report 1 May Annually 31 July Annually ~jodcI impacts of changes 5S03X Nonceririd in available nitrogen Research and energy that are induced by livestock grazing systems on pronghorn nutritional ecology. 1. Develop a nutritionally driven pronghorn population dynamics model. Publication :51 July 1987 2. Conduct pronghom population simulations using nutritional inputs based upon short-term evaluations of livestock grazing systems. Simulations 1. Prepare a detailed study plan. Approved Study Plan 31 July 1983 2. Conduct research. Annual Budget Report Annual Report Publication 1 Hay Annually 31 July Annually 31 July 1987 6-8 pronghorns • Approved Study Plan 31 July 1985 Spring-Summer 1987 b. Assess the accuracy of experimental pronghorn census methods 5503X NOllcervid Research �I j .. OPERATION PLAN COMPONENT - MOUiHAIN I GOAT RESEARCH 1. PROBLEM - THE NUMBER TWO PRIORITY STRATEGY FOR THE MOUNTAIN GOAT PROGRAM IS TO "DETERMINE THE DEGREE OF INTERSPECIFIC COMPETITION BETWEEN MOUNTAIN GOATS AND BIGHORN SHEEP THROUGH RESEARCH," MOUNTAIN GOAT POPULATIONS APPEAR TO BE CAPABLE OF VIGOROUSLY COLONIZING NEW HABITATS AND THEREBY EXTENDING THEIR DISTRIBUTION AS POPULATIONS EXPAND. BIGHORN SHEEP ARE NOT VIGOROUS PIONEERS OF NEW HABITATS. INSTEAD BIGHORN SH~EP DISTRIBUTION IS BEING EXTENDED THROUGH MANAGEMENT TRANSLOCATION PROGRAMS. BOTH PROCESSES - NATURAL EXTENSIONS OF MOUNTAIN GOAT DISTRIBUTION AND ARTIFICIAL EXTENSIONS OF BIGHORN SHEEP RANGES - INCREA?E THE POSSIBILITY OF COMPETITIVE INTERACTIONS WHICH COULD IMPEDE THE ACHIEVEMENT OF STRATEGIC PLAN OBJECTIVES. MOUNTAIN GOATS AND BIGHORN SHEEP APPEAR TO BE SEPARATED ECOLOGICALLY BY THEIR HABITAT CHOICES. THIS SEPARATION MAY DIMINISH~ HCWEVER~ AS POPULATIONS OF MOUNTAIN GOATS EXPAND AND DISPERSE. GIVEN THE LIMITED AMOUNT OF HABITAT AVAILABLE TO THESE BIG GAME SPECIES; INCREASES IN THEIR NUMBERS WILL LIKELY RESULT IN WIDESPREAD SYMPATRY. IT REMAINS UNCERTAIN WHETHER SUCH SHARED RANGE USE WILL DELITERIOUSLY AFFECT ONE OR BOTH POPULATIONS. PREDUCTING THE OUTCOME OF THIS INTERACTION DEPENDS ON ASSESSING THE CAPABILITIES OF EACH SPECIES TO SURVIVE AND REPRODUCE WHEN RESOURCES ARE IN RE~TIVELY SHORT SUPPLY. THIS~ IN TURN~ REQUIRES KNOWLEDGE OF COMPARATIVE HABITAT SELECTION PREFERENCES AND OF THE NUTRITIONAL ECOLOGY OF BOTH SPECIES. 2, STATUS - ~10UNTAIN GOATS (ilREAMNOS AMERICANUS) ARE NOT INDIGENOUS TO COLORADO. THEY WERE FIRST INTRODUCED INTO THE STATE IN L948 (DENNEY L977). SEVERAL ADDITIONAL RELEASES HAVE BEEN MADE SINCE THEN~ INCLUDING A 1961 RELEASE IN THE MOUNT EVANS AREA. THIS AREA WAS WITHIN THE RANGE OF AN INDIGENOUS BIGHORN SHEEP (0Yl£ CANADENSIS) POPULATION, THE PROBLEM IS THAT AS THESE UNGULATES OCCUpy THIS RANGE AND THEIR POPULATIONS GROW IT IS SPECULATED THAT AT SOME THRESHOLD DENSITY OF ONE OR BOTH SPECIES~ "COMPETITION" MAY OCCUR RESULTING IN A COMPETITIVE ADVANTAGE FOR NOUNTAIN GOATS. IT IS PLAUSIBLE THAT THIS HAS ALREADY OCCURRED~ AND THAT THE SYMPATRIC MOUNTAIN GOAT AND BIGHORN SHEEP POPULATIONS ARE ALREADY RESPONDING TO COMPETITIVE INTERACTIONS. 3, ~JECTIVE AND SPACE. - INVESTIGATE THE DEGREE TO WHICH MOUNTAIN 1'1 GOATS COMPETE WITH BIGHORN SHEEP FOR NUTRIENTS \ N N -...J �-'1 N N 00 ( 4. ~PLEMENIAIJDN - THE STUDY OF COMPETITION FOR SPACE VIA HABITAT SELECTION IS A LOGICAL FIRST STEP IN EXAMINING THE PROBLEM OF POTENTIAL COMPETITION. AN UNDERSTANDING OF POTENTIAL COMPETITION WILL REQUIRE THE FOLLOWING ACTIVITIES: A. TESTING THE FOLLOWING HYPOTHESES: (1) MOUNTAIN GOATS USE ALPINE HABITATS DISPROPORTIONATELY TO THEIR AVAILABILITY. (2) MOUNTAIN GOATS AND BIGHORN SHEEP USE THE SAME ALPINE HABITATS. (3) BIGHORN SHEEP USE ALPINE HABITATS DISPROPORTIONATELY TO THEIR AVAILABILITY. (4) MOUNTAIN GOATS AND BIGHORN SHEEP EXHIBIT THE SAME ACTIVITY PATTERNS IN ALPINE HABITATS. B. TESTING THE RESPONSES OF BIGHORN SHEEP HABITAT USE AND ACTIVITIES TO MOUNTAIN POPULATION REDUCTIONS. C. COMPARE ACTUAL MOUNTAIN GOAT DISPERSAL RATES TO MODEL PREDICTIONS. A SECOND AREA,OF RESEARCH INVOLVES THE COMPETITION FOR NUTRIENTS, THIS EVALUATION WILL FOCUS ON THE TWO SPECIES TO USE SCARCE FORAGE RESOURCES BY EXAMINING THEIR ENERGY REQUIREMENTS AND DIGESTIVE CAPABILITIES. ACTIVITIES WILL INCLUDE MEASUREMENT OF ENERGY REQUIREMENTS BY MOUNTAIN GOATS AND BIGHORN SHEEP DURING WINTER AND DETERMINATION OF DIGESTIVE EFFICIENCY OF THESE SPECIES CONSUMING LOW QUALITY GRASS AND BROWSE DIETS. IF GOATS CAN USE LOWER QUALITY FORAGE) AND HAVE LOWER ENERGY REQUIREMENTS FOR FORAGE, THEN THEY WILL POSSESS A SIGNIFICANT COMPETITIVE ADVANTAGE OVER BIGHORN SHEEP WHENEVER THESE SPECIES SHARE FOOD RESOURCES. 5. Bf]lEEITS SHEEP. THiS'RESEARCH WILL FACILITATE PREDICTION OF EFFECTS OF COMPETITION BETWEEN MOUNTAIN GOATS AND BIGHORN �''i " \ I DIVISION FIVE FORI-!NO. PREPARED BY ACTIVITIES YEAR OF WILDLIFE OPERATIONS PLAN (1981-87) ANALYSIS R.B. Gill, D.F. Reed, N.T. Hobbs TITLE Mountain Goat Research PROGRAM Mountain Goats OPERATION PLAN STRATEGY Test the hypothesis that mountain goats use alpine habitats disproportionately to their availabili ty, COST CENfER 5503X Noncervid Research 1. Conduct research. Annua l Budget Request Annua l Report Publication 1 May Annually 31 July Annually 31 July 1985 z. Test the hypothesis that mountain goats and bighom sheep use the same alpine habitats. 5503X Noncervid Research 1. Conduct research. Annual Budget Request Annual Report Publication 1 May Annually 31 July Annually 31 July 1985 3. Test the hypothesis that bighom sheep use alpine habitats disproportionate Iy to their ava i lab i Iity. 5503X Noncervid Research 1. Conduct research. Annual Budget Request Annual Report Publication 1 May Annually 31 July Annually 31 Jllly 1985 5503X . NOllcervid Research 1. Conduct research. Annual Budget Request Annua l Report Publication 1 May Annually 31 July Annually 31 July 1985 Test the responses of big- 5503X horn sheep habitat use Noncervid Research and activities to mountain goat population reductions. 1. Prepare a detailed study plan. Approved Study Plan Annual Budget Request Annual Report Publication 31 July 1985 1 May Annually 31 July Annually 31 July 1987 Compare actual mountain goat dispersal rates to model predictions. 1. 2. Approved Study Pl<1n Annual Budget Request Annu:1l Report Puhlication 31 July 1985 1 ~Iay Annually 31 July Annua lly 31 Jul.y 1987 1. 4. Test the hypothesis that mountain goats and bighom sheep use the same alpine habitats. 5. 6. 55U3X Noncervid Research ACTIVITI 2. Conduct research. Prepare a detailed study plan. Conduct research. PRODUCT ANALYSIS DEADLINE N N 1..0 �I , I 'i N" W o OPERATION PLAN COHPONErJT - BLACK BEAR RESEARCH I. PROBLEM - THE NUMBER ONE PRIORITY STRATEGY FOR THE BLACK BEAR PROGRAM IS TO ~DETERMINE BEAR LIFE HISTORY AND POPULATION CHARACTERISTICS THROUGH 'RESEARCH," THE STRATEGIC PLAN OBJECTIVE FOR BLACK BEAR POPULATIONS IN COLORADO CALLS FOR MAINTAINING 1980 BASE POPULATION LEVELS BUT AT THE SAME TIME MAINTAINING 1980 HARVEST LEVELS, HUNTER NUMBERS, AND RECREATION DAYS THROUGH 1988, THESE OBJECTIVES MAY BE INCOMPATIBLE, PRELIMINARY DATA FROM THE BLACK BEAR STUDY AREA SUGGEST THAT EVEN WITH NO LEGAL HUNTING ALLOWED, ILLEGAL HARVEST MAY BE SUFFlCIENT TO PREVENT POPULATION INCREASES, IN OTHER WORDS, THERE MAY BE NO HARVESTA£LE SURPLUSES OF BLACK BEAR IN THAT AREA. IF IT IS TYPICAL, THEN REDUCTIONS IN HARVESTS, HUNTERS, AND RECREATION DAYS MAY BE REQUIRED TO MAINTAIN BLACK BEAR POPULATIONS. AN ADDITIONAL PROBLEM IS THAT BECAUSE THE RATE OF INCREASE IS SO SLOW IN BLACK BEAR POPULATIONS THAT THE EFFECTS OF OVEREXPLOITATION WILL NOT BE APPARENT UNTIL SEVERAL YEARS AFTER THE FACT BECAUSE OF TIME LAG EFFECTS. 2. SJlU~- INVENTORY OF EXISTING RESOURCES IS A CRITICAL COMPONENT OF MANY RESOURCE MANAGEMENT PROGRAMS, UNFORTUNATELY, NO PRACTICAL METHOD EXISTS TO ACCURATELY AND PRECISELY ESTIMATE BEARS NUMBERS AND DEMOGRAPHY OVER A LARGE AREA, THUS MANAGERS ARE PRESENTED WITH THE OPTIONS OF GUESSING OR UTILIZING DATA FROM A SELECTED POPULATION WITHIN A MAJOR HUNTING AREA TO ANALYZE HUNTING REGULATIONS AND SUBSEQUENT DATA, A COMPARISON OF DATA FROM THE SCIENTIFIC LITERATURE AND COLORADO HARVEST STATISTICS STRONGLY INDICATES THAT SEVERAL IMPORTANT DAU's ARE BEING OVER8uNTED. COMPLICATING ANY ANALYSIS IS THE UNKNOWN BUT SUSPECTED HIGH LEVELS OF UNLICENSES AND/OR ILLEGAL KILLS. By STUDYING A BLACK BEAR POPULATION PROTECTED FROM LEGAL HUNTING WE SHOULD BE ABLE TO OBTAIN DATA ON BASIC NATALITY PARAMETERS (AGE OF FIRST BREEDING, AND DISPERSAL PATTERNS,) OBTAINING SUCH DATA FROM A POPULATION WITHIN A MAJOR BLACK BEAR REGION WILL FACILITATE THE ANALYSIS OF BLACK BEAR HARVEST DATA CURRENTLY AVAILABLE TO WILDLIFE MANAGERS. PRESENTLY SPORT HUNTING OF BLACK BEARS RANKS THIRD BEHIND MULE DEER AND ELK IN MAN-DAYS OF RECREATION AMONG BIG GAME OF COLORADO. INTEREST IN HUNTING BLACK BEAR DURING THE SPRING SEASON HAS DRAMATICALLY INCREASED DURING THE LAST 7 YEARS. THE COMBINATION OF RAPIDLY GROWING HUMAN POPULATION, INCREASING RESOURCE DEVELOPMENT, INCREASING DEMAND FOR BLACK BEAR HUNTING AND BEAR HARVEST DATA SUGGESTING OVERHUNTING STROflGLY DICTATE THE NEED FOR A STRONGER BIOLOGICAL DATA BASE UPON WHICH TO ESTABLISH BLACK BEAR MANAGEMENT PROGRAMS. �'1 " '1 3, RESEARCH OBJECTIVE - DESCRIBE POPULATION OF A SINGLE-SPECIES POPULATION MODEL, 4. l11PLErlENTAIlJJJi- RESEARCH IS NEEDED TO OBTAIN THE BASIC DATA PREREQUISITE TO ANY MEANINGFUL POPULATION MANAGEMENT PROGRAM, AT PRESENT MANAGERS DO NOT HAVE POPULATION DEMOGRAPHY DATA UPON WHICH THEY CAN EVALUATE BLACK BEAR HARVEST DATA, MANAGERS FIND THEMSELVES IN A DILEMMA OF BEING ABLE TO ESTIMATE HUNTER HARVEST. NOT KNOWING LEVELS OF UNLICENSED KILL AND NOT EVEN KNOWING THE POTENTIAL ANNUAL HARVEST SUSTAINABLE, THEREFORE. RESEARCH EFFORTS WILL SEEK TO OBTAIN THE FOLLOWING INFORMATION: A, FREQUENCY DISTRIBUTIONS FOR AGE OF FIRST LITTERING; PERIODS BETWEEN'LITTERS, AND LITTER SIZE, B, AGE-SPECIFIC CHRONIC MORTALITY RATES FOR AC 1.2. AND 3 BEARS, C, r1AN-RELATED MORTALITY RATES FOR BE'ARS AC 4 AND OVER, D, HABITAT PREFERENCE FOR FEMALE BLACK BEARS WITHIN THE OAKBRUSH-ASPEN-SPRUCE COMMUNITIES TYPICAL OF THE WEST CENTRAL PLAUTEAUS AND UNCOMPAHGRE DAU's, E, DENNING ECOLOGY OF BLACK BEARS IN WEST-CENTRAL COLORADO. DYNAMICS OF A SELECTED BLACK BEAR POPULATION TO ALLOW FOR DEVELOPMENT BASED ON THE INFORMATION OBTAINED. THE RESEARCH PERSONNEL WILL DEVELOP A SINGLE-SPECIES MODEL FOR COLORADO BLACK BEAR POPULATIONS AND INSTRUCT MANAGEMENT PERSONNEL IN THE USE AND POTENTIAL MiSUSE OF THE SYSTEM. INFORMATION ON HABITAT UTILIZATION AND DENNING ECOLOGY WILL BE PUBLISHED AND GIVEN TO APPROPRIATE MANAGEMENT PERSONNEL BOTH WITHIN THE DOW AND PUBLIC LAND MANAGEMENT AGENCIES. 5. BENEFITS - THE RESEARCH WILL PROVIDE BASIC INFORMATION ON POPULATION DEMOGRAPHY. SEASONAL MOVEMENTS, AND HABITAT USE OF A POPULATION IN THE IMPORTANT WESTERN PLAUTEAUS DAU. IMPACT OF HUNTING SEASON REGULATIONS WILL BE ANALYZABLE FROM A THEORETICAL RATHER THAN EMPIRICAL, POST-FACTO BASIS. IV W �l ' \ ~ ~ N ~ N DIVISION FIVE rOHM NO. PREPN<ED YEAR OF WILDLIFE OPERJI.TIONS PLAN (1982-1987) ACTIVITIES ANALYSIS BY R.B. Gill, r.D.I. Beck OPI:RATION FLA.'! STR1\TEGY TITLE COST CENTER ACTIVITY PROGRAM Black Bear Research PRODUCr ANALYSIS Black Bear DEADLINE Estimate natality rates of a selected population of black bear. SS03X Noncervid Research l. Conduct research. Amlual Budget Request Annual Report Publication 1 May Annually 31 July Annually 31 July 1987 ~. Estimate mortality rates 5503 Noncervid Research 1. Conduct research. Annual Budget Request Annual Report Publication 1 May Annually 31 July Annually 1 July 1987 Describe h3bitat prefcrcnccs for female black bears from a selected population of black bears. 5503X Noncervid Research 1. Conduct research. Annual Budget Request Annual Report Publication 1 ~lay Annually 31 July Annually 1 July 1984 Describe denning ecology of black bears from a selected black bear population. SS03X NOllcervid Research l. Annual Budget Request Annual Report Publication 1 Nay Annually 31 July Annually 31 July 1987 l. of a selected population of black bear. I. Conduct research. �y ") ~ OPERATION PLAN CO~1PONENT - t10UNTAIN LION RESEARCH 1. PROBLEM - THE NUMBER ONE PRIORITY STRATEGY FOR THE MOUNTAIN LION PROGRAM IS TO "OBTAIN BETTER INFORMATION ON MOUNTAIN LION POPULATION CHARACTERISTICS) HABITAT AND DEGREE OF PREDATION ON DEER AND OTHER WILDLIFE THROUGH RESEARCH." THE 1988 STRATEGIC PLAN OBJECTIVE IS TO MAINTAIN MOUNTAIN LION POPULATIONS AT 1980 BASE LEVELS WHiLE INCREASING HARVESTS BY 34%) HUNTER NUMBERS BY 30% AND RECREATION DAYS BY 27%. THESE OBJECTIVES ARE BASED UPON 2 UNSUBSTANTIATED ASSUMPTIONS: 1) THAT MOUNTAIN LION POPULATIONS CURRENTLY ARE UNDERHARVESTED AND CAN SUSTAIN INCREASED HUNTING PRESSURE WITHOUT DECREASING 1980 POPULATIO~ LEVELSj AND 2) THAT HUNTING MORTALITY REPLACES NATURAL MORTALITY IN HUNTED POPULATIONS OF MOUNTAIN LIONS. ACTUALLY) VERY LITTLE IS KNOWN ABOUT THE EFFECTS OF HUNTING ON MOUNTAIN LION POPULATIONS, THUS THE NUMBER ONE RESEARCH PRiORITY IS TO EXAMINE THE POPULATION DYNAI1ICS OF A MOUNTAIN LION POPULATION PROTECTED FROM AND THEN SUBJECTED TO HUNTING. 2. STATUS - DURING THE LAST LO YEARS) RESEARCH ON MOUNTAIN LIONS INCREASED DRAMATICALLY. NEARLY EVERY WESTERN STATE INITIATED INVESTIGATIONS OF MOUNTAIN LIONS PRIMARILY TO OBTAIN ESTIMATES OF MINIMUM DENSITIES OR POPULATION SIZE, THESE STUDIES GENERALL¥ WERE SHORT-TERM AND NARROW IN THEIR SCOPE, A MAJOR IMPETUS FOR THIS INCREASE IN MOUNTAIN LION RESEARCH WAS THE PERCEPTION OF A WESTERN-WIDE DECREASE IN MULE DEER NUMBERS AND THE SUSPICION THAT MOUNTAIN LIONS WERE CONTRIBUTING TO THAT DECLINE, ANOTHER MOTIVATION FOR ACCELERATED MOUNTAIN LION RESEARCH WAS THE CONCERN OF ANTI-HUNTING GROUPS THAT MOUNTAIN LION POPULATIONS WERE ENDANGERED BY SPORT HUNTING, IN ADDITION) THE DOMESTIC LIVESTOCK INDUSTRY WAS PRESSING FOR MORE STRINGENT CONTROL OF MOUNTAIN LION POPULATIONS, ALL 3 FACTORS COMBINED TO STIMULATE ADDITIONAL RESEARCH ON MOUNTAIN LIONS, BUT. ONLY ONE STUDY FROM IDAHO INVESTIGATED LONG-TERM CHANGES IN MOUNTAIN LION POPULATION DYNAMICS IN AN UNHUr~TED SITUATION, THAT STUDY SUGGESTED SEVERAL HYPOTHESES RELATING SOCIAL DYNAMICS OF MOUNTAIN LION POPULATIONS TO THE REGULATION OF THOSE POPULATIONS, RECENT SHORT-TERM STUDIES IN OTHER STATES HAVE INDICATED THAT RESULTS FROM THE IDAHO STUDY MAY NOT BE GENERALLY APPLICABLE TO MOUNTAIN LIONS IN OTHER ECOSYSTEMS. 3, RESEARCH OBJECTIVE - DESCRIBE THE DYNAMICS A SINGLE-SPECIES POPULATION MODEL, N OF A SELECTED MOUNTAIN LION POPULATION TO ALLOW FOR DEVELOPMENT OF \.AI \.AI �v / 'y i I I 4. .l.l1I:.LHlEfHATIQN - A LONG-TERM STUDY (CA 10 YRS) IS NEEDED TO OBTAIN A DATA BASE WHICH WILL ENABLE WILDLIFE MANAGERS TO MANAGE MOUNTAIN LIONS MORE INTENSIVELY. PRESENT- LEVEL OF KNOWLEDGE IS INSUFFICIENT TO ESTIMATE THE EFFECTS OF INCREASING HARVESTS BY UP TO 34% ON MOUNTAIN ~ION POPULATIONS, To OBTAIN THAT INFORMATION RESEARCH WILL ADDRESS THE FOLLOWING POPULATION CHARACTERISTICS: A, DENSITY OF RESIDENT MOUNTAIN LIONS IN A SELECTED MOUNTAIN LION POPULATION, B, SIZE OF HOME RANGES OF MOUNTAIN LIONS IN A SELECTED MOUNTAIN LION POPULATION, C. NATALITY AND POST-NATAL MORTALITY OF MOUNTAIN LIONS IN A SELECTED MOUNTAIN LION POPULATION, D, DISPERSAL CHARACTERISTICS OF YOUNG MOUNTAIN LIONS FROM A SELECTED MOUNTAIN LION POPULATION, E. MOUNTAIN LION - DEER POPULATION INTERACTIONS WITHIN THE RANGES OF A SELECTED MOUNTAIN LION POPULATION. 5, BE~LI&- DATA THAT RESULT FROM THIS STUDY WILL ALLOW POPULATION SIMULATIONS TO BE CONDUCTED WHICH ESTIMATE THE POTENTIAL OF A MOUNTAIN LION POPULATION TO SUSTAIN SPORT HUNTING. MORE REALISTIC LONG-TERM MANAGEMENT OBJECTIVES WILL RESULT, ,,,, W J:- �, I ;; DIVISION FIVE FORt-!NO. PREPNU:D BY ACfIVITIES '/ ') YEAR OF WILDLIFE OPERATIONS PLAN (1982-87) ANALYSIS R.B. Gill, A.E. r\nderson OPLRATIt1f l'DlJ-N-STRATEGY Estimate density of resident mountain lion in a selected mowltain lion population. TITLE COST CENTER 5S03X Noncerv i d Research Mountain PROGIWI Lion Research PRODUCT ANALYSIS ACTIVITY Mountain Lion DEADLINE 1. Prepare a detailed study plan. 2. Conduct research. Approved Study Plan AJIDllalBudget Request Annual Report Publication 31 1 31 31 January 1983 May Annually July Annually July 1987 Approved Study Plan Annual Budget Request Annual Report Publication 31 1 31 31 January 1983 May Annually July Annually July 1987 Approved 31 January 1983 Estimate home range sizes of mountain lions in a selected mountain lion population. SS03X Noncervid Research 1. 2. Prepare a detailed study plan. Conduct research. Estimate natality and pos t+nat al mortality of mOlmtain lions in a selected mount ain lion popUlation. 5S03X NOllcervid 1. Prepare a detailed study plan. Descrihe dispersal c11aracterist ics of young mountain lions from a selected mountain lion population. SS03X Noncervid Describe mounta in lion deer population intcract ions wi tili n the ranges of a sclectetl mountain l i on popul a t ion. SSI)3X NOllcervid Research 2. Conduct research. Study Plan Annual Budget Request Annual Report Publication 1 May Annually 31 July Annually 31 July 1987 Approved Study Plan 31 January 1983 2. Conduct research. Annual Budget Request Annual Report Publication 1 Nay Annually 31 July Annually 31 July 1987 1. Approved Study Plan 31 January 1~J83 Annual Budget Request J\nnual Report Publication 1 ~f:l y Aruma 11y 31 July J\nnually 31 July 1~87 1. Prepare a detailed study plan. Prepare a detailed study plan. 2. Conduct research. N w V'1 �I :;; ) ) N VJ 0' OPERATIONS PLAN COMPONENT - RESEARCH SUPPORT 1, eRQBill1A, B, C, D, PRODUCTIVITY IN RESEARCH UNITS REQUIRES A WELL-DEFINED SENSE OF MISSION, A CREATIVE ENVIRONMENT, AND CLEAR OBJECTIVES, MISSION STATEMENTS SHOULD INCLUDE DEFINITIONS OF THE RESEARCH UNIT!S PHILOSOPHY OR REASON FOR EXISTENCE AND ITS LONG-TERM (5-10 YEAR) GOALS, CREATIVE ENVIRONMENTS PROVIDE SECURITY, CHALLENGE. AND RECOGNITION, SECURITY IS PRODUCED THROUGH STABLE FUNDING. UNREGIMENTED ATMOSPHERE. AND THE DELEGATION OF AUTHORITY AND RESPONSIBILITY TO THE LOWEST POSSIBLE LEVEL, CHALLENGE IS PROVIDED BY SUBJECTING RESEARCH PEOPLE TO A DIVERSITY OF IDEAS RELATING TO THEIR ASSIGNMENT, RESEARCHERS HAVE TO BE ENCOURAGED AND TAUGHT TO SEEK CRITICISM, RECOGNITION IS PROVIDED BY REWARDING RESEARCH PERSONNEL IN WAYS THAT ARE RELEVANT TO THEIR INTERESTS, REWARDS INCLUDE REMUNERATION, PEER RECOGNITION. INCREASED AUTONOM~AND PROMOTION, CLEAR OBJECTIVES FOR RESEARCH RESULT WHEN RESEARCHERS PLAY A KEY ROLE IN THE PLANNING PROCESS FROM STRATEGIC PLAN TO STUDY PLAN AND WHEN SUPERVISORS PROVIDE CONSTRUCTIVE FEEDBACK TO THEIR EMPLOYEES, THIS RESULTS IN CLEAR STEP-DOWN PLANNING SO THAT EACH RESEARCH STUDY FUNCTIONALLY RELATES TO HIGHEST LEV~L DOW GOALS AND OBJECTIVES, THE CHALLENGE TO RESEARCH LEADERS IS TO PROVIDE A SENSE OF MISSION. A CREATIVE ENVIRONMENT. AND CLEAR OBJECTIVES WITHIN AN ORGANIZATIONAL ATMOSPHERE THAT VALUES CENTRALIZATION OF AUTHORITY BUT DECENTRALIZATION OF RESPONSIBILiTY AND THEREBY CREATES IMPEDIMENTS TO INDIVIDUAL CREATIVITY AND INITIATIVE, AN INTEGRATED PLANNING SYSTEM IS NECESSARY TO LINK INDIVIDUAL STUDIES LATERALLY AND VERTICALLY UP THROUGH THE ORGANIZATIONAL STRUCTURE SO THAT THE ENTIRE RESEARCH PROGRAM EVENTUALLY TIES CLEARLY AND LOGICALLY TO THE DIVISION'S HIGHEST PRIORITY GOALS. OBJECTIVES. AND STRATEGIES, PROFESSIONAL GROWTH IS CLOSELY LINKED TO PRODUCTIVITY OF RESEARCH PERSONNEL, ECOLOGY IS A RAPIDLY GROWING SCIENCE, CONTINUING PROFESSIONAL GROWTH OF RESEARCHERS MUST BE PROGRAMMED SO RESEARCHERS CAN CONTINUE TO BE CURRENT WIT~ ADVANCES IN THE SCIENCE OF ECOLOGY, INDIVIDIUALIZED IN-SERVICE TRAINING PROGRAMS WILL RESULT IN CONTINUING PROFESSIONAL GROWTH OF RESEARCHERS AND INCREASED PRODUCTIVITY, THE END PRODUCT OF ALL RESEARCH IS KNOWLEDGE, THE UTILITY OF THAT KNOWLEDGE DEPENDS LARGELY UPON HOW EFFECTIVELY THE KNOWLEDGE IS TRANSFERRED FROM THE KNOWLEDGE GENERATOR (THE RESEARCHER) TO THE KNOWLEDGE APPLICATOR (THE MANAGER), EFFECTIVE KNOWLEDGE TRANSMISSION REQUIRES AN EFFECTIVE INFORMATION AND �, _,) F. ') EDUCATION PROGRAM TO PACKAGE AND TRANSMIT THAT KNOWLEDGE TO THE APPROPRIATE CLIENT AT TIMES AND IN WAYS THAT ARE RELEVANT TO HIS NEEDS. THE LAG TIME BETWEEN KNOWLEDGE GENERATION IN DOW AND KNOWLEDGE APPLICATION IS TOO LONG (5-10 YEARS). KNOWLEDGE TRANSFERS CAN BE MADE MORE EFFICIENT WITH MORE EFFECTIVE INFORMATION AND EDUCATION PROGRAMS. THE WILDLIFE RESEARCH SECTION FUNCTIONS AS A LOOSELY INTEGRATED GROUP OF SEMI-AUTONOMOUS FUNCTIONAL UNITS EACH SUPERVISED BY A RESEARCH LEADER. EACH RESEARCH LEADER IS EXPECTED TO MANAGE A BLOCK OF FISCAL RESOURCES TO PRODUCE KNOWLEDGE ACCORDING TO PREVIOUSLY DEFINED GOALS AND OBJECTIVES. FOR RESEARCH THE BEST ADMINISTRATION IS THAT WHICH IS DECENTRALIZED TO THE LOWEST EXTENT POSSIBLE. THE PROBLEM WITH RESEARCH ADMINISTRATION IS TO DECENTRALIZE RESPONSIBILITY AND AUTHORITY TO THE LOWEST POSSIBLE ORGANIZATIONAL UNIT THAT CAN EFFECT THOSE DECISIONS WITHIN A STATE GOVERNMENT ENVIRONMENT THAT ATTEMPTS TO DECENTRALIZE RESPONSIBILITY BUT MAINTAINS A HIGHLY CENTRALIZED CONCENTRATION OF AUTHORITY. KNOWLEDGE OF THE RELATIONSHIPS BETWEEN THE DYNAMICS OF BIG GAME RUMINANT POPULATIONS AND HABITAT CHARACTERISTICS (IE CARRYING CAPACITY) HAS BEEN ADVANCED TO ITS CURRENT STATUS PRIMARILY THROUGH INDUCTIVE PROCESSES. ANIMALS AND POPULATIONS HAVE BEEN OBSERVED TO VARY APPARENTLY IN RESPONSE TO CHANGES IN HABITAT CHARACTERISTICS. BUT THESE RELATIONSHIPS ARE ONLY HYPOTHETICAL. IN ORDER TO TEST THESE HYPOTHESES THEY HAVE BEEN FORMALIZED IN10 PREDICTIVE MODELS OF CARRYING CAPACITY AND FORAGING STRATEGIES. FURTHER ADVANCES TOWARDS AN UNDERSTANDING OF THE CARRYING CAPACITY CONCEPT WILL REQUIRE EXPERIMENTAL AND EMPIRICAL TESTS OF THE MODEL PREDICTIONS (DEDUCTIVE PROCESSES). THE MOST PRODUCTIVE TOOL FOR TESTING OF THE MODEL PREDICTIONS HAS BEEN THE USE OF DOMESTICATED WILD RUMINANTS. THESE ANIMALS ARE SUFFICIENTLY TRACTABLE THAT THEY CAN BE SUBJECTED TO CLOSELY CONTROLLED EXPERIMENTAL ROUTINES AND STILL RETAIN NORMAL BEHAVIORAL AND PHYSIOLOGICAL FUNCTION. IT IS COSTLY IN TIME AND FISCAL RESOURCES TO DOMESTICATE NEW ANIMALS EACH TIME AN EXPERIMENT IS REQUIRED. IT IS MORE PRACTICAL TO MAINTAIN EXPERIMENTAL ANIMALS AND EXPERIMENTAL FACILITIES AS A MATTER OF ROUTINE. 2. SI8IUSA. SINCE 1975 THE BIG GAME RESEARCH SECTION HAS INCREASINGLY PROVIDED A SENSE OF MISSION TO BIG GAME RESEARCHERS BY ASSIGNING WORK THAT WAS CLEARLY IDENTIFIED \~ITH TOP DOW GOALS, OBJECTIVES, AND STRATEGIES. WE'VE ALSO INTRODUCED MANDATORY PEER REVIEW OF STUDY PLANS AND MANUSCRIPTS AND HAVE INTRODUCED A SEMINAR N W -....J �_. N \N 00 B, C, D. E. F. SERIES TO EXPOSE THE RESEARCH TO INTERESTED INDIVIDUALS FROM CONCEPTION TO COMPLETION. IN ADDITION) FUNDING HAS BEEN REASONABLY STABILIZED AND RESEARCHERS HAVE PLAYED A MAJOR ROLE IN SELECTING THEIR RESEARCH ASSIGNMENTS. THIS HAS CREATED A HEALTHY TENSiON BETWEEN SECURITY AND CHALLENGE IN THE RESEARCH ENVIRONMENT WHICH WAS INTENDED TO FOSTER PRODUCTIVITY. PRODUCTIVITY HAS INCREASED STEADILY FROM 1975 TO THE PRESENT. BEGINNING IN 1973 THE BIG GAME RESEARCH SECTION HAS WORKED CLOSELY WITH THE PLANNING SECTION TO CONSTRUCT AN INTEGRATED PLANNING SYSTEM. THIS SYSTEM IS NOW FUNCTIONAL. IT BEGINS WITH THE STRATEGIC PLAN WHICH IDENTIFIES HIGHEST PRIORITY GOALS) OBJECTIVES) AND STRATEGIES. NEXT) RESEARCH'S ROLE IN ATTAINMENT OF THOSE GOALS AND OBJECTIVES IS DEFINED IN OPERATIONS PLANS. SPECIFIG STUDIES ARE TIED INTO OPERATIONS PLANS VIA DETAILED PROGRAM NARRATIVES OR STUDY PLANS. ANNUAL BUDGET REQUESTS ARE DETAILED IN SEGMENT NARRATIVES. ANNUAL WORK PLANS ARE DETAILED IN THE PERFORMANCE EVALUATIONS WHICH ARE ALSO USED TO MONITOR PROGRESS TOWARDS GOALS AND OBJECTIVES. CONTINUING PROFESSIONAL GROWTH IN THE PAST HAS BEEN LEFT PRIMARILY TO THE INDIVIDUAL RESEARCHER. SOME HAVE VIRGOROUSLY PURSUED CONTINUING EDUCATION WHILE OTHERS HAVE AVOIDED IT LIKE THE PLAGUE. THIS PROCESS NEEDS TO BE INCREASINGLY FORMALIZED AND PROGRAMMED SO EVERYONE HAS THE OPPORTUNITY AND THE MOTIVATION TO KEEP CURRENT WITH DEVELOPMENTS IN THEIR FIELD. IN THE PAST) THE RESEARCH SECTION DID NOT HAVE AN AGGRESSIVE) PROGRESSIVE INFORMATION AND EDUCATION PROGRAM. WITH THE ASSIGNMENT OF GEOFF TISCHBEIN AS RESEARCH I&E SPECIALIST) THE POTENTIAL FOR IMPROVED I&E EFFECTIVENESS INCREASED TREMENDOUSLY. Now THAT POTENTIAL HAS TO BE REALIZED. STEPS HAVE BEEN TAKEN TO IMPROVE THE ACCOUNTING OF BIG GAME RESEARCH EXPENDITURES THROUGH IMPLEMENTATION OF A COMPUTERIZED BUDGET TRACKING PROCESS. THIS SYSTEM CAN ACCOUNT FOR EXPENDITURES AT THE JOB LEVEL AND IDENTIFIES EACH INDIVIDUAL TRANSACTION BY EACH RESEARCHER. As SUCH) IT OFFERS FAR GREATER RESOLUTION THAN THE CENTRALIZED ACCOUNTING SYSTEM IN THE DENVER OFFICE. WE HAVE ALSO BEGUN TO DEFINE ROLES AMONG RESEARCH STAFF OFFICERS. RESEARCH LEADERS ARE DEVOTING LESS OF THEIR TIME TO BUSINESS MANAGEMENT ACTIVITIES AND MORE TO RESEARCH LEADERSHIP ACTIVITIES. SIMULTANEOUSLY THE BIG GAME RESEARCH SECRETARY IS BEING GROOMED FOR INCREASING RESPONSIBILITY FOR THE BUSINESS MANAGEMENT FUNCTIONS OF THE RESEARCH ASSIGNMENT. A BIG GAME RESEARCH SUPPORT FACILITY HAS BEEN CONSTRUCTED AT THE CSU FOOTHILLS CAMPUS. A RESEARCH ANIMAL POPULATION HAS BEEN DEVELOPED AND CONTAINS MULE DEER, ROCKY MOUNTAIN ELK) BIGHORN SHEEP, PRONGHORN ANTELOPE, AND ROCKY MOUNTAIN GOATS. �'IJ j ~ 3. RESEARCH SUPPORT OBJECTIVES A. EXPAND PEER REVIEW TO INCLUDE KEY DOW WILDLIFE MANAGEMENT PERSONNEL. B. EFFECT CLEARER DEFINITIONS OF THE ROLES OF THE WILDLIFE RESEARCHER) WILDLIFE BIOLOGIST, AND DISTRICT WILDLIFE MANAGER AND DEFINE THE RELATIONSHIPS AMONG THOSE ROLES. C. EXPAND PEER REVIEW OF BIG GAME NONCERVID RESEARCH STUDY PLANS TO INCLUDE MORE EXTRADIVISIONAL REVIEWERS. D. DEVELOP FORMAL CONTINUING EDUCATION PLANS FOR EACH BIG GAME NONCERVID RESEARCH PERSONNEL INCLUDING THE DECENTRALIZATION OF AUTHORITY AND RESPONSIBILITY FOR OUT-OF-STATE TRAVEL TO THE RESEARCH LEADER LEVEL. E. DEVELOP A BIG GAME NONCERVID RESEARCH I&E PLAN WHICH IN DESCENDING ORDER OF PRIORITY WILL: INCREASE THE AWARENESS OF DOW PERSONNEL OF THE BIG GAME NONCERVID RESEARCH PROGRAM AND WHY WE ARE DOING WHAT WE ARE DOING; INFORM THE LAY PUBLIC OF HOW BIG GAME NONCERVID SPECIES FUNCTION IN THEIR RESPECTIVE ENVIRONMENTS AND WHAT ATTRIBUTES OF THOSE ENVIRONMENTS ARE MOST CRITICAL TO EACH SPECIES' SURVIVAL; PROVIDE INPUT OF OUR RESULTS INTO THE EDUCATION SYSTEMS, INFORM THE GENERAL WILDLIFE PROFESSION OF OUR RESEARCH RESULTS. F. EXPAND THE ROLE OF THE BIG GAME RESEARCH SECRETARY TO THAT OF A FULLY FUNCTIONAL ADMINISTRATIVE ASSISTANT AND RECLASSIFY THE POSITION FROM SECRETARY TO ADMINISTRATIVE ASSISTANT. G. IMPLEMENT TRAINING PROGRAMS TO TRAIN ALL BIG GAME RESEARCH ANIMALS TO BE USEFUL TO A WIDER VARIETY OF EXPERIMENTS. H. EXPAND THE USE OF THE RESEARCH FACILITIES AND ANIMALS TO EFFECT GREATER USE OF THESE ANIMALS AND FACILITIES BY CSU STAFF. I. DEVELOP A BETTER MAINTENANCE RATION (OR PERHAPS RATIONS) FOR BIG GAME RESEARCH ANIMALS. 4. l~PLEMENTATION - ALL OF THESE RESEARCH MANAGEMENT OBJECTIVES WILL REQUIRE CLOSE LIAISON AMONG THE 2 BIG GAME RESEARCH LEADERS, THE TERRESTRIAL WILDLIFE RESEARCH CHIEF, THE RESEARCH I&E SPECIALIST, AND THE ASSISTANT DIRECTOR OF STAFF. THE FIRST STEP IN IMPLEMENTATION WILL REQUIRE AGREEMENT ON THE OBJECTIVES OF THIS PLAN AND THE STRATEGIES THAT WILL BE USED TO IMPLEMENT THEM. 5. BE~ ACHIEVED - THE BENEFITS TO DOW WILL BE INCREASED BY INCREASED EFFICIENCY OF PRODUCTION. RESEARCH PRODUCTIVITY AT LESS RELATIVE COST. THIS WILL BE N W 1..0 �240 Divisional Correspondence Only APPENDIX D STA TE OF COLORADO DIVISION DEPARTMENT OF WilDLIFE OF NATURAL RESOURCES DATE: TO: All Noncervid Research Personnel FROM: R. Bruce Gill SUBJECT: Program PIans July 12, 1983 ~ This letter is being sent to you along with a plea for your understanding and indulgence. You will recall at the beginning of this fiscal year I requested each of you to prepare an Operations Plan outlining the research you intended to conduct over the next 5-year period and relating that research to priority Strategic Plan objectives and strategies. Despite several changes in format and delays in finalizing the new Strategic Plan, we completed that assignment before the February 1,1983, deadline established by Dick Hopper. The original formats and deadline were established by Dick Norman, Chief of Planning. However, sometime between February 1 and June 1,1983, Dick Norman was relieved of responsibility for developing Operations Plans. Jim Lipscomb was given that assignment instead. Jim redefined the concept of Operations Plans. Operations Plans are now the composite of 2 new plans--Program Plans and Implementation Plans. Unfortunately, by the time I learned of this change in concept and format, there was not time to ask for your input into the Program Plans. I recently wrote all of the Program Plans for all of the species included in the Noncervid Research Program. Copies of these Program Plans are included for your review and comment. You will notice that these Program Plans contain very little detail or justification. Detail and justification are to be included in our research implementation plans which will be our Program Narratives_and the species management plans. I tried to include everything we had listed in our Operations Plans in the new Program Plans. So, if you donlt see something in the Program Plans you think ought to be there, call me. I donlt know if changes can still be accommodated in drafts of these plans, but if you wish to change the content, let me know and 1111 do what I can. . 11 DOW·A-F-S �I. Il i~ ~ .'1 ij .,'" Pr oq rant I,:: -._--- r, f = r: ;;'tr'1I",.' ..... _.-. -_-------_ __ -_ .. __ . .. _-_ ~: 4-1 4-1-1 Ii: _ _-_._---_ .. _--_-_-_--_"_----_._--_._- .. .. Progress Reports LmpLomon ta ri.on 11'-(> I !, Bighorn SheeQ_ . .... __. ie ·U: ;ii', 1983-88 Progrnm Plan --------.r.-----. .._~~~I_~~LY.._~'_~.L(.'~~t_.i~~_~~ .. Increase carrying capacities of bighorn sheep habitats on public lands through controlled burning, chaining, fertilizing, timber management, and similar methods. . Test the hypothesis that bighorn sheep populations are limited primarily by nutritional deficiencies in critical habitats. a. Prepare a detailed study plan to investigate relationships between nutritional plane and parasitic resistance in bighorn sheep. b. Conduct pi lot experiments to determine the most effective method to challenge bighorn sheep with Protostrongylus sp. larvae. c. Conduct experiments testing the relationships between nutritional plane and parasite resistance. I'l.)ll .1. Product I Responsibility I .~ UtiLC' n },i P, I! V I, ! I .! Program Narrative Noncervid Research Work Plan 2, Job 4 Study Plan Research Annual Report Final Report Research Annual Report Final Report Publications Research 7-1-84 q i! \ I, f ~ ~ ~ hI i' 7-1-84 7-1-84 7-1 Annuall 7-1-86 7-1-87 ;! • ~. ~: r , :'1 i 'I •~ a I 1 ) ~ , N .::- �~ .1983-88 Program Program . Bighorn Plan Sheep 'N .: Reference Number 4-2 4-2-3 Strategy Objective Improve the distribution of bighorns by trapping and transplanting animals within the state and exchanging for desert bighorn sheep from other states. Investigate methods of dispersing sheep into suitable adjacent sites thereby increasing current populations. a. In cooperation with Regional Wildlife Biologists and Big Game Supervisor, develop plans to alter forest habitat structure via burning, logging, and firewood cutting to evaluate effectiveness of these techniques to expand bighorn sheep distribution. b. Assist Regional Wildlife Biologists in analysis and interpretation of results. c. Disseminate results of the investigations. Implementation Plan Bighorn Sheep Management Product P1aniHabitat Management portion of the plan Management Experiment Plans Management Experiment Evaluations Publications, in-service training program Progress Reports Responsibility Dates RegionS-Research Ongoing Regions-Research Ongoing Regions-Research Ongoing :~ �1903-88 Program Plan Program Bighorn Sheep R",[erence Number 4-4 4-4-1 Strategy Objective Provide information on bighorn sheep habitat needs for use in land management decisions and seek mitigation for bighorn losses when impacts are unavoidable. Develop specific quantified habitat -.requirements and habitat management guidelines for species. a. Determine the status of knowledge concerning bighorn sheep habitat requirements and management guidelines b. Develop a model of habitat and bighorn sheep population interactions. c. Test bighorn sheep habitat model. Implementation Plan Program Narrative Noncervid Research Work Plan 2, Job 3 Program Narrative Noncervid Research Work Plan 2, Job 3 Program Narrative Noncervid Research Work Plan 2, Job 3 Product Progress Reports Responsibility Dates Special Report Status of Knowl edge Paper Bighorn Sheep Habitat Model Research 1-1-84 Research 1-1-84 Annual Report Final Report 2nd generation Bighorn Sheep Habitat Model Research 7-1 Annual y 7-1-88 1-1-88 N ,J::VJ �~ 1983-88 Program Plan Program Bighorn Sheep N Rt;(ercncp .. Nurnbo r --_._--_. 4-5 4-5-1 Strategy Objective - Treat bighorn sheep to control disease where neces sarv , Assist the NE Region Wildlife Biologist staff to develop and implement plans to evaluate the efficacy of chemotherapeutic drug treatment of bighorn sheep to control lungworm parasitism. Implementation Plan Bighorn Sheep Management Product Plan Treat and transplant portion of the plan-Evaluation of drug treatment efficacy Progress Reports Responsibility NE Region-Research 4:..j::- Dates 7-1-84 I I I �t -..,\ . ;1': ',"1 :: 190J-00 Program Plan Pr og ram .,Il,n.teJJ1P.e- __·__ ............ --. ----_._-_ ... __ ._-_ .. _ .. _-_ __ ..._._-------_. __ ._._-----. .. f'·!·':ll· .... ;"1'11'(' r 6-1 6-1-1 b. c. 6-1-3 ._._. S_tE.:l_t"g~I_j.:~!:.!:.':.':. .. ._. Provide information on antelope habitat needs for use in land management decisions and seek mitigation for antelope losses when impacts are unavoidable. Develop specific quantified habitat requirements and habitat management guidelines for species. a. 6-1-2 I.. I 1Il.l'.lcmentil t. ion ..._.. I Determine the status of knowledge concerning antelope habitat requirements and management guidelines. Develop a model of habitat and antelope population interactions. Test antelope habitat model. Investigate the impacts of antelope winter wheat production. grazing on Investigate the potential of livestock grazing systems (deferred grazing, rest-rotation grazing, etc.) to enhance both livestock and antelope productivity. -- ______ I'}an Species habitat requirements and management guidelines Program Narrative Noncervid Research Work Plan 1, Job 4 Program Narrative Noncervid Research Work Plan 1, Job 4 Program Narrative Noncervid Research Work Plan 1, Job 4 Product I Pro'lress Reports Responsibility Pronghorn Antel pe Research habitat require ments and guidelines Special Report Research Status of knowledge paper Antelope Research Habitat Model I I Dates .' I I :1' I 7-1-84 I 1 r ! .. i I :f r I. .it I 1-1-84 ',. I 1 I: :L F Program Narrative Noncervid Research Work Plan 3, Job 4 Antelope Management Program Narrative Work Plan 3, Job 5 Plan Antelope Management Plan Annual Report Final Report 2nd generation Antelope Habitat Model Annual Report Final Report Damage Control portion of the plan Annual Report Final Report Habitat Management portion of the plan Research I 7-1-Annualll 7-1-88 1-1-88 !. :" .1 T Research Wi1dl i t e S",; Research '''-I I 7-1-Annualli 7-1-86 Resea rch Game ProgramResearch I 1-1-87 " :' ; 7-1-87 and fnnually Therea ter 7-1-92 1-1-93 N -I='" U1 �\ : \ i I. j 1903-00 P~ogr~m Plan Program ..- " ... ---..- .----------------.'•. f f"' r {'llf~ lllrlh-l 6-2 6-2-1 Antelope' I Impl emerrt e t i on _____ . . ~::.:.'tc:L'/_~L(.'cl.i.v~:, . • ~ L~ Ii,l· n IN ..- ..- .... ------------ (' __ ~ . - -, I Progress Reports Responsibility Product Dates', .~ '"., ~ ... ~ .~ , ~ ;' Improve our ability to match harvest regulations with available antelope populations through better counts and better analysis of count data, Develop improved antelope census systems. I' ~ )! ,\ ~ f Annual Report Program Narrative Noncervid Research Work Plan 3, Job 5 Antelope Management I Research I Final Report Plan Inventory pnrtion of plan I 7-I-Annua 11 Game ProgramRegions-Research 7-1-87 ~l ,~ 7-1-88 "J ~ t ~ f ~ r, j I;. 1 :} .~ 'I' ; �I, ~ L:i i:: u 19B]-U(J Pr oq r am Program l:: Pl,," r:·' J1uuntain Goat 1.,',' -J f," ._.~ - - o ____ • _____________ •••• _. ____ . _________ ••• ___ ._, ••• _______ •• ________ • ____ .': f rr·':tr:n ;'1'":11 or' r 10-2 10-2-1 10-2-2 __ ,__",________ Integrate findings into bighorn sheep and mountain goat management plans. • ________ Implementation Plan ~,:':.:.1_!_cgy __2_L:.~~r~t ivc ___________ Determine the degree of interspecific competition/ between mountain goats and bighorn sheep through research. Continue research on interspecific,competition. . ______________ .: • Product Progress Reports Respansibility I Dates i ;1 }~ I I I I I [1 Program Narrative Noncervid Research Work Plan 4, Job 1 Annual Report Mountain Goat and Bighorn Sheep Management Plans Population status portion of the plans Research 7-1-Annua11 , J. Final Report Region-Game Program-Research Director's Staff ,:~ 7-1-86 I f t I 7-1-88 ~ l It ',: i! , ~ I ! ~ i ',:,,~ . . t , r ;; " ~ • { { I , ! N ~ -...J �r "Ii ~\ ': r"I~f " 1.9fl3-00 Program Pr oq r am .- . frl 111:':11-,(" - .__ --.- ... --.----------------------_._._---, 12-1 12-1-1 ._ .. . _________ ~~ 1~.i~_t~:~r\(_ __~~I.2.-L("'!.~t.:_~_y_~? _.. _..__.________ Black Bear .__ .. ___ Imp].e",ent.:ttion P] an '. ___ -------, . Product Progress Reports Re s pons i b i I i ty - _- ' long-term research on black bears. " :'1\ t' '_':- , .s:00 Dates '1 '!1, Determine bear life history and population characteristics through research. Continue ~, t"l --, "Ilt"" r ~i':::~; I'lan ] Program Narrative Noncervid Research Work Plan 5, Job 1 Annual Report, Research 7-1-Annuall: Final Report Research 7-1-87 ,i: '. . j li ~r::,.t 11 j , .. 'to; ! ~ I I :r I J I i �Lii!' , }} Ii PI'\.\ ]~)n]-no lr oq ram Plan 1"'1"')<11',-,", _ t·' 'II'" t.· _!ng_~__El_~_a._L __ . r. J 'I H I': r r- t r::l("'~ ~!'1,.,1 ' • 12-3 12-3-2 12- 3- 3 12 - 3-4 Str,ltc'JY ._ Obj"cl. i-vc ..__ ._ _ _-----, Regulate hunting to increase harvest of bears causing depredation problems and adult males while protecting young and females where neces sa Q'_. Define what data will be maintaineJ by DAU and establish criteria to set bear harvest quotas by DAU. Develop a population bear DAU. simulation model for each Develop bear harvest regulations to target adult male bears while protecting juveniles and females. _- .. ImpLcmcnt~tion Plan I Product Progress Reports Rcsponsihility Dates !~, !J i;1 ri ! Bear Management Bear Management Bear Management Plan Plan Plan Section on population status and criteria for setting harvest quotas Population simulation model Bear harvest regulations Game ProgramResearch 7-1-84 ii't I: ". :/' ;'i1~.; ' ','i Research 7-1-88 . r, \, :'1 Game ProgramResearch 2-1-84 '; ·H" Ii Ii '/:' T''r; f ri " I; ; ~; " !:, ;_ " N ,J::- so �, :! \1 1983-08 Pr oq r arn Program :1 :;.1 L1'; ~'_I_ :: " PIon -,I ~-- 1: - _1U_ack___!!_~ ~, .' ~. !, lIi :' . - -.---- ----------_ ... 1·~1 frr( __ ._---_------_ _-_._---.. Imp l.ornen t a t i.on 11:-1""' rI"cd '-r 12-4 12-4-1 . . ~~te'l.LQlj_'~:_IJvc. . Provide information on bear habitat needs for use in land management decisions and to mitigate for , bear losses when impacts are unavoidable.· Develop specific, quantified habitat requirements and habitat management guidel ines .for species. a. Detennine the status of knowledge concerning black bear habitat requirements and management guidelines. b. Develop a model of habitat and black bear population interactions. .___ P}~ _ Product Test black bear habitat model. o __ t.os na ~ f I c. N V1 Progress Reports Responsihility ~ ~ ; 1- I -!l c.. Program Narrative Noncervid Research Work Plan 5, Job 2 Program Narrative Noncervid Research Work Plan 5, Job 2 Program Narrative Noncervid Research Work Plan 5, Job 2 Special Report Status of knowledge paper Black Bear I Habitat Model Annual Report Final Report 2nd generation Black Bear Habitat Model Research ~ I, Research ;i -f a -i'1 'r ~ ,t :-f .~. Research 7-1-AnnUal~) jf ! ~13 ~ 1-1- 2. ~ 1- ; 1. 1 I I I I I! �I il : \ ~ 1983-88 Pr oq r am - - ._---.---_ --_-----_._-------------- .. __ ... __._....... _____ 1'1ull __l1Q_y_o_tilin .L iQD_ f,;NiJ :;~f; ,~~f ~; ;.~ , ,i .~ . ~ - __ . .-·f (' r('nf:'r :11"Jj··('r Program ".",. ~I::_'-t:_:c_2X._.!:~~t('Ct i~ ___._____________ Implementati.on Plan Product Progress Reports Responsibility ill Dates ', i i 13-1 Obtain better information on mountain lion characteristics, habitat, and degree of predation on deer and other wildlife through research. 13-1-1 Conduct research on mountain lion ~opulation dynamics, habitat needs, and inventory techniques. ,., f Program Narrative Noncervid Research Work Plan 6, Job 1 Annual Report Mountain Plan Population Simulation Model Lion Management Research 7-1-Annuall 7-1-87 Final Report Resea rch 1-1-88 1:' I ~ 1 ~! 1,':; :" i f J -i .: ~ ,! .. ~ ~~ i~ I N IJ1 �252 APPENDIX E TOWARDS A PHILOSOPHY OF RESEARCH AND RESEARCH MANAGEMENT IN WILDLIFE MANAGINENT INTRODUCTION There is considerable confusion in the literature regarding meanings of the terms "philosophy," "science," and "research." Before we can define the research process or try to articulate policy regarding that process we should agree on what we mean by the terms "philosophy," "science," and "research." So in the interests of clarity, here is what we mean in this paper when we use those terms. Philosophy originally was defined as "a love of wisdom or knowledge." Mtmson (1981) has defined the purpose of science as the quest to broaden our understanding of the world and the things that are in it. So if philosophy is the love of knowledge, then science continues to redefine that which philosophy loves. How does science do this or what is the process by which science works? DEFINITION Goldstein and Goldstein (1978:67) in their book HOW WE KNOW define science as follows: "We choose acterized 1. It is found 2. to define science very broadly - as an activity charby three features: a search for ~~derstanding, for a sense of having a satisfying explanation of some aspect of reality. The understanding is achieved by means of statements of general laws or principles--laws applicable to the widest possible variety of phenomena. 3. The laws or principles can be tested experimentally." If we accept this as a definition of science, then are science and research synonymous? We believe so. The terra research comes from an historic theory of knowledge. Plato theorized that prior to birth each person knows all things. But ... "In being born we forget; but we may recover our memory and our knowledge though only partially: only if we seek truth again shall we recognize it. All knowledge is therefore re-cognition recalling or remembering the essence or true nature that once we knew." (Popper 1962:10). In this sense, research--the process of searching again--is simply an approach to the search for knowledge which is science. This attempt to define the philosophy of research in wildlife management will address 3 questions: 1) What is the research process? 2) What is the ideal role of research in wi.Ldl i fe management? and 3) How should research be managed in an organizational setting'? These thoughts are presented as ideas. not dogmas, as a basis for discussion and ultimately decisions about the research process as applied by researchers in a wildlife management; agency. In the process of developing �253 these thoughts we've consulted over 100 books and publications on the subject to learn about the subject. But as Karl Popper (1969:29) once remarked in his book CONFRONTATIONS AND RERITATIONS ... "It might be well for all of us to remember that while differing widely in the various bits we know, in our infinite ignorance we are all equal." TIlE RESEARCH PROCESS Medawar (1969:2) in his book INDUCTION AND INTUITION IN SCIENCE said of the research process ... '~~eall know in rough outline what lawyers do, or clergymen, physicians, accountants, and civil servants; we have a vague idea of the codes of practice they must abide if they are to succeed in their professional duties, and if we were to learn more about them we should be edified, no doubt, but not surpris~d. But what are scientists like as professional men, and how do they set about to enlarge our understanding of the world around us? There seems to be no one answer." In this statement, Medawar encapsulates 2 persistent and pernicious problems that have long infected wi Idl i fe management. First, as Medawar says, it is difficult to define specifically what it is that scientists do, so it is difficult to distinguish the scientists from the nonscientists, particularly since Aldo Leopold exorted a11 of us to become ever more "scientific" in our practice of wildlife management. This ambivalence about what it is that scientists do in the field of wildlife management is reflected in the titles we give to people in wildlife mariagement agencies. We have numerous wildlife managers and wildlife biologists but scant few wildlife scientists and wildlife researchers. The 2nd problem with science in lVildlife management, stems from the 1st. Because we are ambivalent about what it is that wildlife scientists do, we are equally ambivalent about what wildlife science can produce. We tend to think of science and problem-solving as synonymous. As a result, we think the application of science should automatically result in solutions to our problems and when it does not, we fault our science. The late O. C. Wallmo used to say "Problems are artifacts of men's minds." By that he meant that ecology recognizes no problems, only processes and change. Problems occur only when those processes do not result in changes that are desirable to society. That is why people try to control processes to effect desired changes. Barriers to control are called problems. If we had complete knowledge about the systems we seek to control, there would be no ba~ers and thus no problems. But we rarely confront barriers to our efforts ~ control which can be resolved solely by application of knowledge. Most frequently we resolve problems with some knowledge or fact, some intuition or guesswork, some chance or luck, and considerable uncertainty. "Munson (1981:195) acknowledged this when he said ... "It should be no surprise that contemporary medicine despite its commitment to science, continu 5 to employ practical success or �254 ( control as its basic criterion for evaluating rules, procedures, and causal claims. Medicine is an eminently practical enterprise that must attempt to meet immediate and urgent demands. It cannot afford to wait for the acquisition of the appropriate scientific knowledge but must do the best it can in the face of ignorance and llilcertainty. We do not look to medicine to tell us what the world is like. Rather, we count on medicine to act against disease and suffering. "Science, by contrast, is a leisured pursuit. It may be prodded by external demands to solve practical problems, but its internal standard of success continues to be truth. It can afford to wait and work until this is met. For medicine, the constant external demand to promote health is identical with medicine's internal aim, and as a result, practical success is both the external and internal standard. i~at we expect of medicine is what it expects of itself." Science has a role to play in problem solving, but that role is to generate more and more. knowledge about problem components so that decisions are made with less and less uncertainty. But we should not expect science to make the uncer.tainty disappear from the deci.sion-maker+s problem-solving process. It should also be pointed out that researchers can playa more comprehensive role in the problem-solving process than simply by generating new knowledge pertinent to those problems. Researchers - like managers - have intuition, guesses, and opinions that should be sought. But when they offer these intuitions, guesses, and opinions, they are no longeI' functioning as scientists. Instead, they function as educated players in the problem-solving theater. As researchers, they only generate knowledge via the scientific process. So, in the interest of clearing up the waters and at the risk of muddying them even further. 111e are going to take a stab at defining what science ought to be in wildlife management and what it ought to produce. Becht (1974) in a paper called SYSTEMS THEORY, THE KEY TO HOLISM AND REDUCTIONISM, outlined an iterative or repeating process wh ich he called the empirical cycle that we think characterizes the research process. OBSERVING ~ ~ EMPIRICAL CYCLE CHECKING \8 .~ PREDICTING GUESSING ~j) ...... r Research usually doesn't start with random observations. Generally there are reasons for wan t ing to know something. So, in a sense> all research is goal oriented and all research starts with hypotheses (Popper 1()7.2). The primary objective of the observation phase of research is to detect any patterns or relationships among the observations. These observations of p3.tterns then lead to guesses or hypotheses about the nature of cause of these relationships. Then, if the �_- ._ .. 255 . hypothesis is stated in an unrunbiguous way, predictions can be deduced from the h}~othesis such that if the hypothesis is true, the predictions must also be true, and vice versa. Finally, checks or experiments are devised to see if the predictions can be verified. The process is repetitive because the experiments are themselves observations from which additional hypotheses result. Let's illustrate this process with some real wildlife examples. In the 1950's C. W. Severinghaus and co-workers observed regional differences in body size, weights, and reproductive performance among white-tailed deer in New York. They hypothesized that these differences were caused by regional differences in carrying capacity of winter habitats. They predicted that if deer populations could be drastically reduced in the impoverished regions, deer size, weight, and productivity would increase because per capita deer food would increase. They proposed a management experiment to test their hypothesis. They advocated eithersex deer hunting to increase deer harvests. This, they reasoned, would increase the per capita amount of food available to the remaining deer which would result in larger and more productive deer. Severinghaus and his colleagues, however, went 1 step further. They tried to sell their hypothesis as a management prescription. Unfortunately the story ended there because the politics of the proposed hunt .precluded its implementation. . Severinghaus could be faulted as a problem-solver because his solution did not solve "the problem." It didn' t solve it because it wasn't politically practical. Severinghaus could also be faulted as a scientist because he didn't test his hypothesis before he sold it as fact. But science did not fail management in this example. The failure resulted from a faulty application of research to a problem. In the 1960's, Lou Verme of ~lichigan set up eA~eriments with penned deer to test 1 of the predictions of Severinghaus, i. e. that nutrition is directly correlated to changes in deer productivity. Venne found that does which were offered adquate, nutritious diets produced greater numbers of fawns than does which were fed restricted diets. These effects were particularly pronounced among younger does. Verme also ·noticed, lIDexpectedly, that malnourished does had a tendency to produce more male offspring that! female offspring which 'led him to an additional hypothesis concerning maternal nutrition and sex of the progeny. This hypothesis led to further predictions, which led to further experiments, and so on. Ultimately, Verme and others linked the nutritional hypotheses into a more general hypothesis about white-tailed deer and habitats. This hypothesis postulated that optimum deer habitat is provided by secondary successional stages of northern mixed hardwood-conifer forests. The Michigan Department of Natural Resources adopted this hypothesis as the major strateb~ to meet a goal of 1 million deer in Michigan by 1980. Verme's successes can be cont rast.ed to Severinghaus' failures as an example of how research ought to be used in the management or problem-solving process. Verme used science to test his hypothesis and then used the results of his experiments to formulate an even broader and more general hypothesis. He cOImnlIDicated both the hypotheses and the results of his experiments to his peers. But the wildlife managers--not Verme the researcher--took his results and transformed them into a management prescription. Venne kept most of his attention. on further experiments and hypotheses which would generate even more knowledge about deerhabitat interactions for the management problems and prescriptions of the future. �256 This abbreviated and highly abstracted example highlights several important aspects of the research process. First, the Severinghaus experience illustrates that researcll rarely solves problems. Biological research can only solve problems that are strictly biological in nature. This is because the only product of the research process is knowledge about the system that is being studied. Only rarely do we ever have enough information at hand to resolve multidimensional problems on the basi s of fact alone. Research --Li.ke law enforcement --is only I of several tools the wildlife manager needs to solve her problems. Research should be evaluated on the basis of its contributions to knowledge, not on how we1l it solves management problems. Cal.dwelI (1966:526) in his paper PROBLEMS OF APPLIED ECOLOGY talked to this issue when he said.... "To argue that applied ecology is not very useful because it does not move the Corps of Engineers or the Bethlehem Steel Corporation is somewhat like arguing that insulin is of no particular value to a diabetic who will not take it. 111e arguments do not disprove the efficacy of ecology or of insulin. But they do indicate that the effectiveness of applied science depends upon the circumstances of application as well as upon the validity of the science." l The 2nd point illustrated. by the example is that research is most productive when it is allowed to revolve throtlgh several repetitions of the empirical cycle. This ailows the researcher(s) the opportunity to link small hypotheses into more general and rrore useful (in an explanatory sense) larger hypotheses. Several examples come to mind where unirrterrupted repetitions of the empirical cycle have resulted in significant contributions to wildlife ecology. Among these are the long-term studies of Keith and. others regarding snowshoe hare ecology; the long-term studies by Chitty and associates on population regulation in small mammal populations; the work by Caughley concerning the feedback regulation mechanisms between ungulate populations and plant popUlations; the long-term studies of Mech and others on the sociobiology of wolves. None of these studies would have fulfilled their expectations without continuity and longevity. Finally, a 3rd lesson in the white-tailed deer examp.Ie is that all too frequently wildlife science never progresses beyond the hypothesis stage. Romesburg in his paper GAINING RELL\BLE KNOWLEDGE THROUGH WILDLIFE SCIENCE (1981:294-295) spoke to this issue when he stated ... "Because wildlife science hardly uses the H-D (hypotheticodeductive) method, it is stuck with no way of testing the many research hypotheses generated by retToduction. Herein lies the main cause of urrreli.abIe knowl edge . The research hypotheses either are forgotten, or they gain credence and status of laws through rhetoric~ taste, authority" and verbal repetition." Romesburg's message is that Our science is poor because ses but we've seldom tested prematurely because as Gill ( ~ we have been guilty of poor science in wildlife ecology. itls incomplete. We've generated scores of hypothethem. Instead, we accept hypotheses as fact (1976: 10=.,-104) suggested In an earlier paper. .. "We have been too eager to believe, too quick to appJy. and too busy to think." �257 One final word about the research process concerns what it is not. Research is all too often regarded as a data gathering process. This is the so-called "background" research. This unfortunate misconception arises because several data gathering activities have been labeled research. Thus we have histori~al research, legal research, etc., all of which are basically data gathering processes. But if we accept research as synonymous to science, then research is much more than a data gathering process. Saxon (1983:14-15) in a paper in the AMERICAN JOURNAL OF PHYSICS, spoke to this issue when he said ... "First, it (curriculum) should be designed to help students understand the nature of physical laws, what they are and what they are not; what they can tell us about the physical world and what they cannot; how they are arrived at; and in what sense they are true. "Second, it should provide some grounding in the laws of probability and chance, and thus some understanding that in a world as complex as ours both statistical fluctuations and the accidental coincidence of unrelated events happen all the-time. ~hny events are unexplainable, are simply happenstance, much as we would like to believe otherwise. "Third, it should convey the important idea that science is not a collection of isolated facts but a highly unified and consistent view of the world. In many ways it can be described as a web--a conceptual, empirical, theoretical, and historical web, nearly but not quite seamless. We need to give our students some sense of why we believe the world and everything in it is made up of small particles; why we believe in our modern grand theories, such as relativity, when past grand theories have been proven wrong. Newton's theory, after all, has been superseded by Einstein's; the chemists' proof that matter is immutable, itself a correction of'the earlier views of the alchemists, has been superseded by the discovery through nuclear chemistry that it is indeed possible to transmute base metals to gold. Pastuer's proof that spontaneous generation was impossible has been replaced by the views of modem biologists, who are convinced that life itself most likely began spontaneously." . "The reason, of course, is that science has a foundation of large general laws that link together various observations about the physical world and which provide a framework within which various potentialities, facts, and theories can be evaluated. And when a generalization that has long been believed proves no longer true, it is usually because the physical domain in which that generalization was valid has turned out to be a limited one, more limited than actually exists in nature. The discovery of radioactivity demanded radical revisions in our view of nature because it arose entirely beyond the boundar i cs of what was then known. It should be possible--I have no doubt t}~t it is possible-to convey to students both the power and the limits of general scientific laws and why we can, in the light of.both, draw reliable conclusions from those laws." �258 ( Most of the sting of these crItIcIsms we aim at reasearchers, not at managers. But we think a large part of our problem in wildlife ecology stems from the fact that we haven It clearly defined and distinguished the roles of the wi.Idl.i.feresearcher from those of the wildlife biologist and the wildlife manager. TIlE RESEARCH ROLE Wildlife administrators have argued forcefully that role relations between the wildlife researcher, the wildlife biologist, and the wildlife manager should be ambiguous and vague. They base their argument on the observation that in agencies where there were organizational distinctions among these functions, effective cOITllTrtmications were impeded because of rivalries for status and prestige. We believe effective communications have been impeded precisely because role relationships have remained ambiguous and this has led to overlap of responsibility and competition for authority. Romesburg (1981:311-312) at the conclusion of his paper said ... l "I regard medical science and wildlife science as fields with equal· potentials for achieving reliable knowledge. I think, however, that medicine has come closer to its potential, whereas wildlife science has lagged. I thip..kmedicine owes its success to the strict attention it pays to scientific method. Scores of books on the philosophy of clinical experiments have been published, yet I know of few comparable books in the natural resource sciences. Medical science obviously cares for and is committed to the quest for reliable knowledge . It is a good role model. !I If Romesburg is correct, perhaps we can also find in the medical profession model role relationships analogous to those of the wildlife researcher, the wildlife biologist, and the wildlife manager. According to a paper by Munson (1981) entitled WHY MEDICINE CANNOT BE A SCIENCE, even though medicine uses science to make lnedicine more scientific, the 2 disciplines are fundamentally different. He claims that the "basic internal aim of science is the acquisition of knowledge and understanding of the world and the things that are in it." The basic aim of medicine, on the other hand, is "to promote the health of people through the prevention or treatment of disease." Munson (1981:194) contrasts these disciplines in the following way ... "In seeking knowledge, science can be described as a quest for truth about the wor Id , In seeking to promote health, medicine can be described as a quest for control over factors affecting health. Knowledge or under stand.ing of biological processes is important to medicine because it leads to control. But where information is lacking, medicine will seek control in other ways. In particular, it will rely on empirical rules that are validated (at least partially) by practical success." We apologize for quoting Munson at length, but we think the relationship of science to medicine is closely analogous to the relationship of science to wildlife management. �259 Wildlife science should provide knowledge about how animals relate to their environments and wildlife management should use that knowledge to control factors regulating animal populattons in ways that serve the interests of the public owners of the wildlife resource. To pursue the medical analogy a bit further, in medicine 3 distinct but integrated roles have evolved to promote health: the role of the medical researcher, the role of the research. physician, and the role of the practicing physician. The medical researcher generates knowledge a.bout hOI";living organisms function and what factors regulate or a.lter those functions. The research physician has a dual role. She seeks to extend knowledge about the ftmctions of the human organism but also she experimentally tests treatments to control or prevent disease. The practicing physician seeks to promote health. In so doing she uses every weapon in her arsenal including facts, hypotheses, guesses, and intuition. Of the 3 roles, that of the practicing physician is certainly the most risky because human lives are at stake as well as the physician's livelihood. We believe these 3 roles are analogous to the ideal roles of the wildlife researcher, the wildlife biologist, and the wildlife manager. If we are correct, then the ideal wildlife researcher would formulate and test hypotheses concerning plant-animal interactions, animal-animal 'ineractions, population regulation mechanisms, harvesting strategies, etc. In other words, the ideal wildlife researcher'S goal would be to expand our knowledge about animals and their relationships to their environments. The ideal wildlife biologist would take that information from the wildlife researcher and think of ways that it might be used to manipulate animal populations and their environments in ways that are potentially useful to societal goals. The ideal wildlife biologist also would design and conduct experiments to test management hypotheses. Finally, the ideal wildlife manager would apply the knowledge gained from the ecological experiments of the w i Ld.li.feresearcher and the management experiments of the wi Idli fe biologist on 'a broad scale to achieve desired animal population and environmental objectives. Like the practicing physician, we think the ideal wildlife manager's job would be the most risky because she will not be able to manage populations and environments of wild animals on a totally scientific or factual'basis. ~IDSt of the time he will have to manage with a pinch of fact, a slice of intuition, a cupful of experience, a shovelful of luck, and a warehouse full of pants seats. She will learn,most effectively through the frequent experience of failure. His procedural tools lrill be those of the practicing physician: examination, diagnosis, prescription, and monitoring. EXAMINATION ~ ~ DIAGNOSIS MONITORING oj) ~ r PRESCRIPTION ~--..•-"",J' Because He have tested so few hypotheses in wild.life ecology, we have mostly conjecture and little fact in our medical bags. The wise manager will not prescribe large sweeping wildlife management therapies. Instead she will prescribe her therapies as limited management applications. He Hill prescribe them in ,.•. ays �260 ( that will pennit objective and quantitative evaluation. Then she will carefully monitor results of these experiments to see if broader-scale application is warranted. Lindblom (1959:86) in his paper THE SCIEl\JCEOF "MUDDLING THROUGIf' described this process as follows: "Making policy is at best a very rough process. Neither social scientists, nor politicians, nor public administrators yet know enough about the social world to avoid repeated error in predicting the consequences of policy moves. A wise policymaker consequently expects that his policies will achieve only part of what he hopes and at the same time wi l.L produce unanticipated consequences he would have preferred to avoid. If he proceeds through a succession of incremental changes, he avoids serious lasting mistakes in several ways. l "In the first place, past sequences of policy steps have given him knowledge about the probable consequences of further similar steps. Second, he need not atten~t big jumps toward his goals that would require predictions beyond his or anyone else's knowledge, because he never expects his policy to be a final resolution of a problem. His decision is only one step, one that if successful can quickly be followed by another. Third, he is in effect able to test his previous predictions as he moves on to each further step. Lastly, he often can remedy a past error fairly quickly--more quickly than if policy proceeded through more distinct steps widely spaced in time." (See also Bailey' 5 [1982J paper IMPLICATIONS OF ''MUDDLING THROUGH" TO WILDLIFE MANAGEMENT.) In our view, the Hildlife manager functions as the developmental researcher of the wildlife profession. Both the wildlife researcher and the wildlife biologist should function as consulting physicians to help the wildlife manager design and evaluate his management applications. Now those of you who have been reading carefully and critically may have noticed that we've just done some semantic magic. We started out by telling you that the method of research was the empirical cycle which is characterized by the repetitive or iterative processes of: _----.A (!'-> OBSERVING CHECKING t:=-:~J ./7 ~ PREDICTING GUESSING <:"-:::'/ Next we told you (without actually telling you) that the ideal wildlife biologist was actually the applied researcher of wildlife ecology. Finally we told you that the method of the wildlife manager \Vas characterized by the iterative processes of: �261 f . M:)NITORING if> ~ EXAMINATION ~ DIAG'JOSIS PRESCRIPTION ~!) 'v-- And we said the wildlife manager was the developmental researcher of the wi Idlife profession. Except for the words chosen to describe it~ the method of the wildlife manager is indistinguishable from the method of the wildlife researcher. This is as it should be because this method has evolved as an effective way of establishing truth or fact, and truth is not the sole domain of scientists. It belongs to us all. The primary differences beuveen sicentist, biologist, and manager are their purposes or functions within society, or more specifically, within organizations. Perhaps the best way to illustrate this is through another analogy. The function of a wildlife researcher is analogous to a generator--she generates knowledge. The function of the wildlife biologist is analogous to a trans former- -11e transf~rms knowledge into a potentially useful form. And the function of the' wildlife manager is analogous to the user--she plugs into this useful knowledge and puts it to work to complete worthwhile tasks. (Churchman 1977) Earlier in this discussion, we mentioned in passing that we believed a large measure of the dissension which exists among members within wildlife agencies results from failures to clearly define roles and role relationships within those agencies. We said that because definition of roles and role relationships is 1 of tilecornerstones of organizational theory. ~Vhen organizations work, they work because the diverse talents of individuals have been directed in ways that those talents compliment each other rather than compete with each other. It's like the music played by allorchestra. Orchestras are composed of musicians who play instruments which make different sounds. But orchestras only play music when each musician knows what sounds to make and when to make them. Musical scores are written so that each musician knows his role. Conductors constantly adjust the tempo, intensity, and transition of those sounds to give the performance harmony. Without the conductor or the score you have only disharmonious noise. You have cacophony--not symphony. Francis (1977: 21-22) in his book PRINCIPLES OF R&D ]'\!ANAGE1ENT, spoke to the need for role definitions in organizations when he sai~ "An organization serves to bind the various individuals in the endeavor and integrate their separate activities along a single direction to achieve common goals. Organization accomplishes this first by mobilizing diverse resources within the structure. More than that, however, it is a mechanism for countering those competi t ive forces whi.ch would tL'1C' e rmi.ne human collaboration. It is designed to minimize or resolve conflicts and neutralize the effect of individual behavior which deviates from group standards and it attempts to do this, rroreover, without stifling individual creativity. �262 Finally, organization introduces stability into intragroup relationships by reducing uncertainty regarding the nature of group structure and the individual roles within it." ( All of which brings us to the final question posed in our introduction-"How should wildlife research be managed in an ideal setting?" RESEARCH MANAGEMENT In his book, TIlE MANAGBvlEl\1T OF RESEARCH AND DEVELOPl'vtENT, Walters (1965) identified 6 key elements in the rese-ardl management process: Planning, Organization, Leading, Performing, Administering, ruldEvaluation. Books have been written about each of these subjects so our treatment of them is extremely superficial. We will try to highlight some of the lessons learned. from industry that we think are particularly pertinent to wildlife management agencies. PLANNING l Walters indicates that the foremost consideration ln the research planning process is the selection of projects. Effective project selection depends upon welldefined and relatively stable agency goals and objectives" Research is most effective when it is directed towards obstacles that will have to be sunnounted to achieve the top priority goals of the agency. In other words, the most productive research is directed. towards the agency's most pressing problems. Most government agencies today are committed to the philosophy of management by objectives. Consequently, the agency's goals and objectives are usually written and communicated to each employee. But, as Aldous Huxley (1969) remarked in his book ENDS AND .tvlEANS ... "With regard to the goal, there is ....a very good general agreement. Not so with regard to the roads that Lead to that goal. Here unanimity and certainty give place to utter confusion, and to the clash of contradictory opinions~ dogmatically held and acted upon with the violence of fanaticism." It almost sOlmds like Aldous Huxley once worked for the Colorado Division of Wildlife. The dilemma that faces wildlife agency administrators is that wildlife managers, on the one hand, are the people with their feet to the fire. They are responsible for the grass roots management of the wildlife resources. They are acutely aware of the problems they face in that management responsiblity better than anyone. QUite naturally, they think that they should connnand the lead role in the selection of research problems. Researchers, on the other hand, are the agency's experts within their subject matter fields. They know better than anyone what we already know and have good ideas about what remains to be learned and which among; those things needs to be learned 1st. (_ Perhaps a satisfactory way out of this dilermna can be found in the planning process. Wildlife managers should playa central role in the definition of the �263 agency's wildlife management problems. These problems will be reflected in the agency's statements of goals and objectives and the priorities the agency attaches to each of them. Wildlife researchers should play a central role in determining \vhat new information needs are likely to contribute most towards achieving those goals and objectives. Neither group should work in a vacuum. Pelz and Andrews -(1976) suggest in their book SCIENTISTS IN ORGANIZATONS that researchers are most productive in a climate characterized by both security and challenge. Security is provided by a working enviror~ent that gives the researcher considerable freedom to pursue her own ideas, encourages evaluation of the work by peers inside and outside of the organization, and provides the researcher with ready access to everyone who influences research assignment decisions. Challenge is provided by critical peer review of the researcher's hypotheses, research strategies, and results. Researchers should be challenged by biologists and managers to defend the logic of their biological problem analyses. Biologists and managers should be challenged by researchers to defend the logic of their management problem analyses. Both should be given the security to use their individual strategies to make their integrated but separate contributions to the solutions of the organization's problems. One final caveat, neither wildlife managers nor wildlife researchers can be productive in organizations where the agency's direction changes each time the political winds change. ORGAl'JIZATION Walters (1965), in his book, presents organization charts for some of the most competitive and innovative corporations in the nation. All of those organizations have at least one thing in common. Research is plugged into a decision process at the very apex of those organizations. Most have vice presidents whose only assignment is to manage research and development functions. This is because all of those organizations realize that research is absolutely essential to maintain the organization's competitive edge. If research isn't plugged into decisionmaking at the very top, it won't be keyed to the organization's most important goals and objectives. Walters (1965:113-114) stressed this point by saylng ... "An indication of the place. status or importance of research and development within a company is the level at which the chief R&D administrator is located in the organization or on the organization chart. If he reports to the president or chief executive officer, he is usually considered more important to the organization than if he reports to the chief officer of engineering or another operating or staff department. In a separate study of 52 organizations the author found a majority having a chief administrator of research and development reporting to the president, chief executive officer or executive vice president." WI Idf,L One final remark on the chain of command in the ideal ~ is highlighted by an additional quote from Walters (1965:109) . l}1allc.<¥\'1U.~t- agency 0 • �264 "The consensus on the chain of cOJ11I1land appears to be that the number of administrative levels through which a scientist or engineer reports to the top should be as short as possible. The recent trend of decentralization of responsibilities in research and development tends to decrease the chain of command by placing responsibilities for decisions nearest to where they are made. Since they are usually made by the individual scientist, and. since creativity usually or often rests with the individual scientist, this accentuates decentralization and the need for it in research and development." LEADING The consensus of the sources we consulted was that the leaders of the research sections should be former researchers. The reason is most of these sources believed it was more effective to train a scientist to manage and administer than it was to train a nonscientist to understand science and the scientific method. Leadership of researchers can best be summarized by what we call the 4 gets of research management. l 1) get good people 2) get them directed towards meaningful objectives 3) get them adequate resources to do the job 4) get the hell out of their way. PERFORMING Walters sums up the key to productive performance in research and development sections with a single word---decentralization. He defines decentralization as the delegation of authority, responsibility, and accoLmtability to the lo\vest level of management at which the decision can be made and the function performed. Frequently we try to evaluate research on the basis of how well or how effectively the research has been used in the management process. This is a mistake for 2 reasons. First, as we pointed out previously, effective application depends on a willing applicant as well as effective science. But probably, more importantly, we regard the research clientele too narrowly. We look only to the wildlife manager to see what impact research has had on society. In reality, research serves a diverse clientele. Within the Division of Wildlife, for example, researchers not only talk to District Wildlife Managers and Regional Wildlife Biologists but to environmentalists, wi.Id li.fetechnicians, infonnation and education specialists, administrators and commissioners as well. Any time knowledge is required to assist the problem-solving process, researchers can contribute. l There is a diverse extra-agency clientele as \-:e11. Politicians, educators, other agencies? and wildlife citizen advocacy groups all require and use the knowledge generated from research in their respective social roles. So perhaps a broader evaluation of this clientele and the effectiveness of research in meeting their demands is in order before we judge the efficacy of the research. �265 mUNISTERING CorporatIons which use research and development most competitively and most effectively are those which not only decentralize authority, responsibility and accountability of research decisions to the lowest possible management level, but decentralize administrative functions as well. Administration includes the following activities: personnel administration, budgeting and accounting, research support facilities, training, ~nd public relations. In the highly successful companies,the research and development sections have their o\vn staff officers to handle these administrative func ions. The result is administrative officers who view their roles as service roles rather than watchdog roles. COORDINATION AND EVALUATION Industry and most federal government agencies now routinely monitor performance of all organizational w1its and the organization as a whole through periodic performance audits. Function of the audit is to determine how well the organization and its organizational units are functioning by comparing their performance records against their goals and.objectives. The audit also recommends adjustments to improve performance. The audits are most frequently conducted by -in-house teams comprised of senior research and operations managers. In summary, we can characterize our image of the ideal wildlife management in relation to how it understands, programs, and manages research to maximum advantage. Such an organization would have the following characteristics: 1. It would have a clear understanding of what the research process does and what ,it cannot do. It would use research to generate knowledge, but would realize that research is only 1 of several tools used in problem-solving. 2. It would clearly distinguish the roles of the wildlife researcher, the wildlife biologist, and the wildlife manager and equally clearly define the expected relationships among those roles so they would pull together rather than pull apart. 3. It would direct its research towards top priority goals of the agency but with the realization that the research of today is is aimed towards the problems of tomorrow. It also woul.d realize that research is most productive when there is continuity and longevity to the research assignments so the empirical cycle can revolve through several turns. 4. I'tiwou.ld plug its research executives into the apex of the organizational structure to insure that research was directed towards problems of the highest priority. 5. It would recruit researchers for research administration positions and then train them to be good administrators. 6. It would decentralize authority, responsibility, and account.ahili ty for research decisions to the Iowes t possible level. �266 ./ ( 7. It would decentralize administrative possible organizational level. officers to the lowest 8. It would periodically audit activities of the total organization and its organizational units--including research--to maintain productivity and progress toward agency goals. 9. It would COrrmD_tadequate, stable and long-terrn funding to the research effort. ruE STATUS OF REALITY Up to this point we've tried to create a concept of how the ideal wildlife management agency would view science, how it would define the role of science in relation to the organization's philosophy. goals, objectives, and structure, and finally how it would manage (illa business sense) researchers and the research process to maximize its utility to the organization. We used the analogy of the medical profession to illustrate our ideas. We did not use the example of corporate industry where most. of the research about the sociology of research has been concentrated. We did not use industry as a role model because lye believe that industry represents a special case where research has been used effectively to solve industry's "problems." We do not think the same conditions apply within wildlife management. First, industry has been able to use research effectively because it has a simple, easily monitored index to success - profit. Second, the systems to which industries have applied research are also simple. They are technological systems designed towards the manufacture of mechanical or chemical products. These systems are relatively easy to control because they are realtively simple. Since they can be controlled, they can be successfully directed towards useful products with profitable frequency. The industrial analogy is unique to itself for a 3rd most important reason. Industry has been able to tap a considerable and refilling reservoir of basic knowledge at little expense to corporate resources. Universities have maintained traditions of strong institutional support for the so-called basic sciences of physics, mathematics, and chemistry. These disciplines have been uniquely endowed by the universities with stable academic and fiscal environments. Consequently, industry did not have to invest heavily in either science or scientists. Instead, industry predominantly could hire chemica19 mechanical, and electrical engineers. Engineering problems, to some extent at least, can respond to increased infusions of money with increased productivity. The medical profession has not been blessed with the same traditional institutional support from the universities that the universities have bestowed upon physics, mathematics, and chemistry. The medical profession had to build its own basic research institutions within and outside of the universities. ~lost of these institutions have been financed with government funding. Government is acutely aware of 'the successes research has brought to manufacturing. The Manhattan Proje~t and the Space Projects are just 2 major examples. Consequently, goveTllffient expects the medical profession to be able to solve the nation's health problems. All that is needed is to pour money on the problem and cancer, heart disease, multiple sclerosis would all be conquered in short order. �267 ..,.,._"" ~. They were not! They were ~ot because the systems were inconceivably complex and the basic research was too ~ager to develop and test predictive hypotlleses which would lead to control. Our expectations of science have been too great in the medical'arena. However, research institutes have been built and staffed. If they can be left alone long enough, the practical payoffs will come--but at their own pace.· More money and less patience will not spur productivity in the medical profession as it has in Indus try :(see Borek 1982 and Chargaff 1980 for expansions on this theme). ~ l~t then of wildlife management? In wildlife management we have not yet even begun to build the basic research'institutions. The universities have allowed their wildlife science to be sold:to the highest bidder. For the most part, university wildlife scientists have been funded by clients who expected products in short order. The universities :have responded by turning out predictable science. Chargaff (1980:378) spoke tp this best when he said of the medical profession... . "When I hold those stories of woe against the st.m1ptuously funded 'institutes,' 'centers of excellence,' 'clinical and ~iagnostic centers,' etc., + can see that the trend is all toward the creation of very large scientific conglomerates in which, under 'the leadership of men with managerial qualifications, .the predictable will be discovered in ton lots. II In wildlife management we've been able to discover the predictable in ton lots even without St.m1ptuouslyfunded institutes, centers of excellence, and clinical and diagnostic centers, but we haven't been able to discover much of the unpredictable; and we do not understand well the systems we manipulate. The only stable funding sources available to 'wildlife management and wildlife science have been the Pitmann-Robertson and the Dingell-Johnson funds. If wildlife management is going to grow into a more socially responsive profession, we are g6ing to have to do a much better job of unraveling the mysteries of the extremely complex systems we are trying to control. That can only corne about by better wildlife science aimed directly at problems of ecosystem function. ! -"'!'!~'t.;~· In the short-term (for the remainder of the 20th century), the best hope for better wildlife science lies within the federal and state wildlife management agencies. Those agencies have the fiscal and human resources for the job. But those agencies are going to have to exert leadership and responsibility beyond their narrow self-interest to make, significant progress. They will have to accept their responsiblity for developing and filling the reservoir of basic .research in wildlife management. Then we need to build the basic research institutions and fund them with P-~ and D-J moneis. Finally, we need to redefine the roles of the wildlife biologist in wildlife management so the wildlife biologists gradually identifies with and accepts the role of the applied researcher and the wildlife manager with the role of the wi.l.d li fe producer or information user. We think that progress in wildlife management can be made only when the wildlife researcher, the wildlife biologist, and the wildlife manager evolve, respectively, into bioscientists, bioengineers, and biodoctors. Only then can their individual sounds be harmonized into a s)~hony. �268 LITERATURE CITED l Bailey, J. A. 1982. Implications of "muddling through" for wildlife management. Wildl. Soc. Bull. 10(4) :363-369. Becht, G. 1974. . 24:569-579. Systems theory, the key to holism and reductionism. Borek, E. 1982. 313-315. Congress as a research director. BioScience Trends Biochem. Sci. 7(9): Caldwell, L. K. 1966. Problems of applied ecology: perceptions, institutions, methods, and operational tools. BioScience 16:524-527. Chargaff, E. 1980. In praise of smallness - how can we return to small science? Persp. BioI. Med , 23(3):370-385. Churchman, C. W. 1977. Towards a holistic approach. pp. 11-24. in R. A. Scr.ibner, and R. A. Chalk. Adapting science to social needs: mowledge, institutions, people into action. Amer. Assoc. for the Advancement of Science. Washington, D. C. 312 p. Gill, r <; R. B. 1976. Mule deer management myths and the mule deer population decline. pp. 99-106. in G. W. Workman, and J. B. Low. !'vfule deer decline in the West. A symposium. Utah State University College of Natural Resources and Utah Agricultural Experiment Station. Logan, Utah. 134 p. Goldstein, M., and I.F. Goldstein. 1978. How we know. An exploration of the scientific process. Plenum Press. New York, NY. 357 p. Hansen, C. S. 1978. Social costs of Michigan's deer habitat improvement program. Mich. Dept. Natur. Resour, Wildl. Div. Rep. No. 2808. 61 p. Lindblom, C. E. 19:79-88. 1959. The science of "muddling through." Publ. Admin. Rev. Medawar, P. B. 1969. Induction ru1d intuition in scientific thought. Philosophical Society. Philadelphia, PA. 62 p. Munson, R. 1981. Why medicine cannot be a science. Amer. J. !'vIed. Phil. 6:183-208. Pelz, D. C., H. Meyer, and S. W. Gellerman. 1975. Organizing the organization for better R&D. Amacom. New York, NY. 45 p. Popper, K. R. 1963. NY. 412 p. Conjectures and ·refutations. Basic Books. New York, Popper, K. R. 1972. Obiective knowledge. An evolutionary approach. University Press. Oxford. Engl and , 380 p. Romesburg, H. C; 1981. Wildlife science: J. wn,n. Manage. 45 (2):293--313. gaining reliable knowledge. Oxford �269 S~xon, D. S. 1983. The place of science and technology in the liberal arts ' curriculum: conference on science and technology education for civic and professional life - the undergraduate years. Amer. J. Physics 51(1); 12-15. Walters, J. E. 1965. Research management: Books. Washington, DC. principles and practice. Spartaq Williams, L. P. 1967. History of science. pp. 1-14 in B. A. Leerburger, Jr., (ed). Cowl.es encyclopedia of Science, industry andtechno1ogy. Cowles Education Corporation. New York, NY. 510 p. �270 APPENDIX F r: STATE OF COLORADO WILDLIFE COMMISSION July 10, 1975 POLICY NO. B-9 SUBJECT: RESEARCH The Wildlife Commission recognizes that proper management of wildlife resources is dependent upon continuously expanding factual knowledge related to terrestrial and aquatic wildlife species and their environments. Re s ea r ch to obtain this knowledge shall be subject to the following general guidelines: 1. Research shall be conducted for the purpose of developing necessary knowledge and effective techniques and procedures for use by the Division in the proper management of the wildlife resources of this state. The Division shall consider the Colorado Cooperative Wildlife Research Unit and the Colorado Cooperative Fisheries Re s ea r ch Unit to be integral parts of the Division's research establishment. Activities and services performed by these Units shall be closely coordinated with the research program and responsive to the management objectives of the Division, and utilized to the greatest extent possible. 2. Research needs, objectives and responsibilities shall be determined by the Director and his staff. Research projects shall be correlated with Strategic Plan goals and objectives with guidance from the Planning Section. Projects shall be subje ct to annual review and modification. 3. Research findings must be currently translated and disseminated by the research staff to the Commis sion and to the various units of the Division in a form and manner to insure effective u nde r standi.n g and application of research information. 4. It shall be the responsibility of all personnel of the Division to seek the latest research findings applicable to operational programs and to utilize such information in all decisions affecting management of the resources. 5. Wherever and whenever possible, research ac tivi ti e s of the Division shall be coordinated with those of other agencies, organizations and institutions engaged in corollary research. It shall be the responsibility of the research staff to use every pos sible means to eliminate duplication of research effort and to take advantage of com.bined efforts and talents to provide the most knowledge for the least cost. / 6. Implementation of research findings shall be in augu r at ed within the parameters here defined by de cis ion s of the Director and his staff. �271 APPENDIX G STATEOF COLORADO DEPARTMENT OF NATURAL RESOURCES DIVISIONOF WILDLIFE Administrative ;' Directive .; .: .• " .~.;::: :'~-~.:~-'.~ .. .:.~.;':':.q~' ., 'J' August 3~ 1982 ", No. I-I SUBJECT~.RESEARCH PLANNING ANDPROJECT .... '..: i.JMPLEMENTATION ... '.... I. i:,'~.::~.'.;:~.:.' ..'::,'.,; 1'~'~:!":'·"'-' 'r-' ' ,·;or ..~.}.:.'.•',~.·.. .~;' '.... ·" . I .: _) ". ~4..~~~} ... ?~·/i.~:,.' -,'.~. "'.. . . ::-i . This directive sets forth" guidelines for the development of research plans and assigns responsibilities for their preparation and implementation. Flexi= ...." bility is provided to incorporate attention to short-term problems as they arise. .». "_':,. ,.,.:1: r,» ~ " . ".J', I I. .; PLANNING PROCESSANDFORMAT A. Research plan development will involve three steps~ 1. ........ '::": Preparation of Five-year ,Operations Plans. . These plans address Division needs as outlined in the statewide Strategic Plan, are updated in time with Strategic Plan revisions and '.'involve budget considerations. . 1 2. Preparation of a detailed Project Narrative or basic plan for the execution of each job or study. These plans cover in detail the studies outlined in the above Five-year Operations Plans. . .:. ' ," '~. "'~:;,' 3. Preparation of Annual Work or Segment Plans. B. All planning steps will be developed within the framework of the Division's program planning process and. planning system format. Operationsp as well as Staff personnel at all levels, are esponsible for identification of research needs.' C. Each program category following: '. in the '·f,' -, 1. five=year plans ,'. -- ",' will .include .::~:,.:,:./ .. the .. ':.'.:';:' ::"\"_"_' A brief review of past Division management'" and efforts and a discussion of future needs for both. : research , .,: '.' 2. 3. D. : A plan for meeting those · and procedures or a list · projects. future needs, including objectives and brief description of proposed . ,- A list (or discussion) of related, ongoing research being done by the Cooperative WildE fe and Fisheries Units and by other ....agencies and institutions. Separate but integrated plans will game and nongame research. be develo ed for Division fishp E. ' Plans will be developed in terms of program, subprogram and species categories defined by recogni.zable environmental units and/or complexes. i �272 " . 'Administrative Page 2 Directive '.~41~;I~~> No. I-I ,,(~,It,:' .III. PLAN PREPARATION AND REVIEW '''; A. '. Research planning will be the responsibility of r~~earch personnel .and Research Section Chiefs in coordination with. the .. Program Sec.' . tions. These persons will integrate the proposed 'plan into the '. research and DIvision program planning structureo. ~;;:,. ~~~'::.:.<,~::~',. .. "'., .... • > , "B. :' ... ~ : c' D. "":' ': .' • '-. •••. • • "" .";~:':'~~; ~. (~~~~?;~: i' . E. B.' ." • : ':.-_, . ........ '.;,.:'_ ','\-, ~ ) --_,-,,-,,_,_-, ...•+ ........ :ij: . . -",. '-.y •• ~.'''' I', " ..~. • ..;~ ,.'" . ,~~~ -. .... .~i~~( ::.:.,,:, .C The final approved five-year research Operations Plans will be incorporated into the Division's comprehensi ve plans by the . Research and Planning Sections and disseminated as a part of the Division's statewide plan documentation. ,::_i' .' PLANIMPLEMENTATION A. .-" Revised plans then will be prepared by the Research Sections, and following review by appropriate Program personnel and approval by the Director's Staff, submitted to the Chief of Planning for incor.' poration into the Division'S comprehensive planning process. This process will include authorization for new research starts at the general staff meeting for annual resource allocation. Preparation and revisions of research Five-year Operations Plans will be '. synchronized in time with management Five-year Operations Plans revtsfons. It is intended that research and management Five-year Operations Plans be developed to complement each other.' . . ".,' IV. • The respective research staffs will consult with the,.Regional Man. agers and appropriate program management staffs prior"to prepara'. tion of the first drafts of the Five-year Operations Planso Together, these groups will agree on the general direction research should take to meet the needs outlined in the Strategic Plano .I~\?:\..:.~J~::ff~:~:~: ::. .:' C. The research staffs will prepare draft 'editions of the Five~year Operations Plans and submit them to Section Chiefs. for crt tical review and response, with sufficient lead time' to integrate -research plans with other- Oivision planning processes.' Sections included will be: Planning, Nongame Programy Fish Program, Game Programy Ecological Services, Wildli fe Servi ces, Information' and . Education, Law Enforcement and Regions. .""~~t . .. ' " Research Chiefs, in collaboration with the Research leaders, will . assign research personnel to specific studies according to their individual expertise and interestsp insofar as these are compatible with the priorities of research plans. Assignment of research personnel to non-research tasks will be done only after the assignments have been reviewed with the appropriate Research Section Chief to determine the impacts of those non-research assignments upon ongoing research activities. Research personnel will design specific studies, sary project documentation 9 conduct the research results. prepare the necesand report on the '~, :- !.~j; ..•.,.. �273 ,.... Administrative Directive Page :3 / c. ."_,', ." . No. I-I .:: . :" "Research, personnel will be responsible for presenting and inter, preting the results of their research to other Division personnel " _in an appropriate and timely manner. Final reports will normally be submitted within 12 months following termination of each study. . " .:_ ",. _' . ...-: _';; ...... ::.. :,' ~ ••. : t'" ;,' ,,';.' : ,:' " . .. -: !,' ' ::;. ~~; ' .'. .", " .. " . ' i . "- . " :, ;. .- . ', ... " . , "/' ·"'l '. " _::J':-" ,~,." ' .'. �274 ST~TEOF COLORADO DEPARTMENT OF NATURAL RESOURCES DIVISIONOF WILDLIFE .... Administrative 'Directive No. 1-2 August 5 ~ 198 2 ,', " . 9JBJECT: It-A...EMENTATION OF RESEARCH RESULTS . . 1. "(:.' PURPOSE . '. .. .:; ~ " ,,':', ,.""., . . . .' Research and management activities need to be integrated to design improved This directive provides guidelines operations based upon research results. for development. of a research-management information and implementation network. . ,. ":.' .' . . II. . RESPoNsIBILITIES .',.' A. .<" .... 8. - .. Research 1. At the conclusion of each research study, research personnel will submit the results of their research for publication, normally within 12_ months of the termination of the study. These publications will be given to management personnel. Research personnel will serve as advisors to Operations and Staff personnel to interpret results of research and discuss implications to management. 2. Research personnel will aid in the design of new managemeqt activities and modification of existing activities. Research personnel will transmit to Operations and Staff personnel new and/or improved techniques applicable to management. 3... Research resul t s, when applicable, will be implemented as the result of administrative action subsequent to fiscal and manpower approval. Where implementation of research findings cannot be accommodated through ongoing or "matntenance" fiscal and manpower resources, budgetary project request forms will be submitted not less than 16 months prior to the anticipated implementation date. Implementation of research results will be clari fied in the annual Resource Allocation Plan where either reallocation of existing money and manpower or new monies and/or manpower are involved. 4. Research personnel will be available to conduct Division of Wildli fe training on subjects within their areas of exper=' tise. Such training will be coordinated by the Division's Trainf~ Coordinator. Plans to incorporate research findings into management activiies will be developed at these training schools. Staff 1. The Director and his staff agement activities at the leveL will implement research into manAssistant Director for Operations �,', 275 / Administrative Page 2 2. Directive 'j~;:-' . '.~.j~~~/ .~}:;:/~l .'r~)' 1-2 b, The PlanningSection will assist the Regionsin developing strategies for incorporating research results planning and resource allocation proce~s. ", 3. ~n~o.~he fiscal .ittiJt~' :)~t '. i, ,~~ Staff Section Chiefs = Game, Fish, Nongame, Law Enforcement ~ ~~,;':I Information Services':'"' will recommendstrategie~.);:f.pr ..!ncorpor- ';~~~'~'~':':: ating research results into manaqenent , . ',:' ;'.~';:"::;'-.'. };'f.:;--'~;:::: "i~'; c. ~era;~;i~~l:;ict;tse;~i~:; 2. .' : :ilgr~:;el~~na~§~i~l~~~d~~; for implementation of research by Operations -. ;;,;:~,~~~~:,::r:.~ . <12~9~ .1 .•.· . , ..:;.,~,:;,.:..,.:.:::,; ;}-,;;p Operations personnel at all levels will be responsible for ,·!4~:.!.!'~~. implementation of research results as contained in the annual Resource Allocation Plan. \,41 ,II: ',:;,/1 • :Xi,~;. .'i}kt,~:H:'~ le·:, :. ~'~r>~ , iJt..:' I,' " :1~ ~{~~I ~'::0,~ .~,:. , . ':: ', . " .", ... , /,', . ~,' .'. '..;,.. " ". < r , ��277 Colorado Division of Wildlife Wildlife Research Report July 1983 JOB PROGRESS REPORT State of Colorado Project No. 45-01-503-15050 Multispecies Work Plan No. 4 Job No. Period Covered: Author: Big Game Investigations - Noncervids Investigations Big Game Forage Selection Dynamics 7/1/82-6/30/83 R. B. Gi 11 Personnel: R. B. Gill ABSTRACT Two hundred and thirteen references were located during the segment. Approximately 1/3 have been read. No progress was made towards developing a draft Program Narrative because research audit activities commanded the majority of my time. This Job Progress Report represents a prel iminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the Director. ��279 BIG GAME FORAGE SELECTION DYNAMICS R. Bruce Gi 11 P. N. OBJECTIVE To develop an approved Program Narrative forage selection by big game ruminants. to investigate factors influencing SEGMENT OBJECTIVES 1. Review and synthesize the literature tion theory and data. 2. Prepare a Program Narrative selection theory. relative to big game forage selec- to test predictions of big game forage RESULTS AND DISCUSSION Two hundred thirteen references were located and approximately 1/3 of them were read during the FY 82-83 segment. No comprehensive theoretical synthesis was attempted because not all of the reference material has been read yet. No Program Narrative was begun because most of my time was dominated by research audit activities. But some preliminary thoughts can be explored in this report. Traditionally, wildlife biologists have regarded animal forages from a taxonomic perspective. Investigators of wild animal food habits and forage preferences most often have described forage preferences as priority lists of taxonomic items occurring in diet samples of various kinds (Kufeld 1973). The taxonomic approach to animal food choices has influenced wildlife habitat evaluation procedures and competition and forage allocation concepts. We assumed that since we Homo sapiens recognized plants and animals according to our hierarchial taxonomic categorizations that other animals and plants perceived one another in the same fashion. Recent evidence from several studies raised serious questions about the utility of the taxonomic paradigm to understand animal forage choices. Bell (1969) reported that 4 species of Serengeti wild ungulates appeared to be selecting for different plant parts within plant species. Wildebeest seemed to select for grass leafs, zebra selected for grass stems, and topi were intermediate. Selection patterns seemed to be related to chemical composition (protein) of forage items, morphology of the digestive systems of the grazer, and the nutrient requirements of the grazer. Bellis (1969) observations have been confirmed by several other investigators (Jarman 1974, Arman et al. 1975, Hoppe 1977, Hoppe et al. 1977, Kreulen and Hoppe 1979, Drozdz 1979, Kay et a 1. 1980, Hobbs e t a 1. 1982, Owen-Sm ith and Nevel 1ie 1982, Hobbs et al. 1983). But all of this evidence is observational. It supports the hypothesis that wild ungulates select diets along nutrient gradients rather than along �280 taxonomic gradients. Janzen (1979; 338-339) talked to this problem when he said "In short, there seem to be two productive directions to take in working out the preferences displayed by animals. One is to develop realistic artificial diets and then tip in solitary and combined secondary compounds and nutrients to define the limits of tolerances and pick out those compounds that 1hvoke exceptionally strong reactions. The other is to focus on a few key species of plants, work out their defense repertoires in detail, and then focus on the specialist herbivores that get around these defenses and the general ists that are deterred by them." The most productive research agenda probably would include both approaches. Recently, several researchers have incorporated the nutrient gradient diet selection hypothesis to evaluate wild ungulate habitats (Sinclair 1975, Bobek 1977, Mentis and Duke 1976, Mentis 1977 and 1978, Wallmo et al. 1977, Hobbs et al. 1982). However, all of these systems regard plants as a relatively nonresponsive component of the plant-animal grazing system. It has become abundantly clear in recent years that both plants and animals are highly interactive in this system. Plants react to herbivory and herbivores respond to the plant reactions (Oates et al. 1977, Abaturov 1979, Chapman and Blaney 1979, McNaughton 1979 and 1983; Bryant and Kuropat 1980; Waterman et al. 1980; Bryant 1981; Batzl i 1983, Bryant et a l, 1983). The research proposed under Work Plan 1, Job 4 will be designed to test experimentally and empirically some of these emergent hypotheses concerning the nutrient gradient of forage selection and plant responses to herbivory. Preliminary thoughts are to grow selected grass, forb, and shrub species in a greenhouse environment. Plants within each forage class will be treated with supplemental nitrogen to enhance plant protein and potential digestibility. Forage selection of treated vs. control plants by at least 1 wild ruminant species [currently pronghorn because they most closely fit Hoffman's (1973) classification of concentrate selector or selective feederJ. Additionally, my intent is to monitor selected secondary plant compounds in grazed vs. ungrazed plants. These thoughts are no more than "brain-storm" ideas at the present time. As I do more reading and involve more people in discussions of these ideas these tentative plans may change. This research is neither trivial nor academic. Much of the current thinking regarding habitat evaluation on a nutritional basis depends upon an accurate and detailed understanding of forage selection dynamics. Those dynamics are unpredictable when forage selection dynamics are interpreted along taxonomic gradients. They become potentially more predictable if, as hypothesized by Hobbs et al. (1982), wi ld ruminants select forages along nutrient gradients. Given enough knowledge about forage selection mechanisms, it might be possible to predict forage yields from phytomass yields from chemical analyses of phyomass samples (Jung and Fahey, Jr. 1983, Moir and Ebersohn 1983). This possibly has important implications to big game ruminant habitat evaluation, forage allocation, and game damage assessment procedures. LITERATURE CITED Abaturov, B. D. 1979. Peculiarities of tropic interrelationships involving plant-animal interactions in pasture ecosystems. Agro~Ecosystems 5:317-327. �281 Batzl i, G. O. factors. 1983. Responses of arctic rodent populations Oikos 40:396-406. to nutritional Bell, R. H. V. 1969. The use of the herb layer by grazing ungulates in the Serengeti. Pages 111-124 in A. Watson (ed.). Animal populations in Blackwell Scientific Publ., Oxford, relation to their food resources. England. Bobek, B. 1977. Summer food as a factor limiting roe deer population Nature 268:47-49. size. Bryant, J. P. 1981. Phytochemical deference of snowshoe hare browsing by adventitious shoots four Alaskan trees. Science 213:889-890. Bryant, J. R., and P. J. Kuropat. 1980. Selection of winter forage by subarctic browsing vertebrates: the role of plant chemistry. Ann. Rev. Ecol. Syst. 11 :261-285. Bryant, J. P., F. S. Chapin, III, and D. R. Klein. 1983. Carbon/nutrient balance of boreal plants in relation to vertebrate herbivory. Oikos 40:357-368. Chapman, R. F., and W. M. 'Blaney. 1979. How animals perceive secondary compounds. Pages 161-198 in G. A. Rosenthal and D. H. Janzen (eds.). Herbivores: their interaction with secondary plant metabolites. Academic Press. New York. 718pp. Drozdz, A. 1979. Seasonal intake and digestibility roe-deer. Acta Theriologica 24(13):137-170. of natural foods by Hobbs, N. T., D. L. Baker, and R. B. Gill. 1983. Comparative nutritional ecology of montane ungulates during winter. J. Wildl. Manage. 47(1): 1-16. Hobbs, N. T., D. L. Baker, J. E. Ellis, D. M. Swift, and R. A. Green. Energy- and nitrogen-based estimates of elk winter-range carrying capacity. J. Wildl. Manage. 46(1) :12-21. 1982. Hoffman, R. R. 1973. The ruminant stomach. East Afr. Monogr. BioI. East African Lit. Bureau. Nairobi, Kenya. 354pp. 2. Hoppe, P. P., S. A. Quortrup, and M. H. Woodford. 1977. Rumen fermentation and food selection in East African Zebu cattle, wildebeest, Coke's hartebeest, and topi. J. Zoo 1. , London. 181:1-9. Jarman, P. J. 1974. The social organization of antelope their ecology. Behaviour 48(3-4) :215-267. in relation to Janzen, D. H. 1979. New horizons in the biology of plant defenses. Pages 331-350 in G. A. Rosenthal and D. H. Janzen (eds.). Herbivores. Their interactTOns with secondary plant metabolites. Academic Press, New York. 718pp. �282 Jung, H. J. G., and G. C. Fahey, .Jr . 1983. Interactions among phenolic monomers and in vitro fermentation. J. Dairy Sci. 66:1255-1263. Kay, R. N. B., W. V. Engelhardt, and R. G. White. 1980. The digestive physiology of wild ruminants. Pages 743-761 in Y. Ruckebusch and P. Jhivend (eds.). Digestive physiology and metabolism in ruminants M.T.P. Press Limited, Lancaster, England. 854pp. Kreuten, D. A., and P. P. Hoppe. 1979. Diurnal trends and relationship to forage quality of ruminanal volatile fatty acid concentration, pH and Afr. J. Ecol. 17: osmolarity in wildebeest on dry range in Tanzania. 53-63. Kufeld, R. C. 1973. Foods eaten by the Rocky Mountain elk. Manage. 26(2):106-113. J. Range McNaughton, S. J. 1979. Grazing as an optimization process: grassungulate relationships in the Serengeti. Amer. Natur. 113(5):191-203. McNaughton, S. J. 1983. Compensatpry herbivory. Oikas 40:329-336. plant growth as a response to Mentis, M. T. 1977. Stocking rates and carrying capacities for ungulates from African rangelands. S. Afr. J. Wildl. Res. 7(2):89-98. Mentis, M. T. 1978. Economically optimal species - mixes and stocking rates. Internat1. Rangeland Congr., Proc. 1:146-149. Mentis, M. T., and R. R. Duke. for large wild herbivores. 1976. Carrying capacities of natural veld S. Afr. J. Wi ldl. 6:65-74. Moi r , K. W., and J. P. Ebersohn. 1983. Evaluation of pasture for beef cattle from measurements of cell wall in separated leaf and stem fractions. J. Agr. Sci., Comb. 100:5"13-518. Oates, J. F., T. Swain, and J. Zantovska. 1977. Secondary compounds and food selection by colobus monkeys. Biochem. System Ecol. 5:317-321. Owen-Smith, N., and P. Novellie. Amer. Natur. 119(2):151-178. 1982. What should a clever ungulate eat? Sinclair, A. R. E. 1975. The resource limitation of trophic levels in tropical grassland ecosystems. J. Anim. Ecol. 44(2):497-520. Wallmo, O. C., L. H. Carpenter, W. L. Regelin, R. B. Gill, and D. L. Baker. 1977. Evaluation of deer habitat on a nutritional ba~ls. J. Range Manage. 30(2):122-127. Waterman, P. G., C. N. Mbi, D. B. McKey, and .J.. S. Gartlan. 1980. African rainforest vegetation and rumen microbes: phenolic compounds and nutrients as correlates of digestibility. Oecologia 47:22-33. ~ Prepared by \' C', \ -, \\ .\~_'--\.,~(:' ~I._Q_l -=-R-.-B~r_';'_L_"lce Gill .------.----.-.----.--Wildlife Research Leader � https://cpw.cvlcollections.org/files/original/073a99ff42140d03d78c81bfcacf4311.pdf 96a4e38cf0757497e1989c1345f97e0f PDF Text Text Colorado Division of Wildlife Wildl ife Research Report July 1983 283 JOB FINAL REPORT State of Colorado Project No. 45-01-503-15050 Work Plan No. 2 ------------------- Job. No. Personnel: - Noncervids Bighor~ Sheep Investigations Prescribed Burning of Bighorn Sheep and Mule Deer Winter Range Period Covered: Authors: Big Game Investigations 7/1/80-6/30/83 N. T. Hobbs and R. A. Spowart N. T. Hobbs and R. A. Spowart ABSTRACT Prescribed burns were carried out in mountain shrub and grassland communities to test hypotheses about efficacy of fire as a method of improving habitat for mule deer (Odocoileus hemionus) and mountain sheep (Ovis canadensis). Prescribed burning elevated the concentration of protein and in vitro digestible organic matter (IVDOM) in winter dieti of mountain sheep and mule deer. We observed no effect of burning on ungulate nutrition during spring. Effects of burning on diet crude protein persisted for 2 years in both communities. Treatment effects on diet IVDOM lasted for 2 years in mountain shrub, but were absent during the second year in grassland. Simulation modeling of energy and nitrogen balance revealed that thesedifferences in the nutritional quality of ungulate diets caused substantial enhancement in overwinter nutritional status. Simulated mule deer and mountain sheep maintained or gained weight throughout winter on burned mountain shrub and grassland plots, but lost weight on controls. Enhanced nutritional status appeared to be related to increased carrying capacities re su ltInq from burning in both communities. Increases in the amounts of high quality forages on burned plots allowed burned communities to carry more animals at high levels of nutrition, although controls could carry more animals over all diet quality levels. Burning increased diet overlap between mule deer and mountain sheep~ This' increase persisted for 2 years following treatment with no reduction in magnitude. Increased diet overlap on burned plots resulted from deer switching from browse dominated diets on controls to diets dominated by grass similar to those chosen by sheep on burns. Nitrogen mineral ization rate was increased 1 year after the burn in both communities. This increase persisted for 1 year in the grassland and for 2 years in the shrub community. Ambient levels of N02- + N03- and NH4+ ��285 PRESCRIBED BURNING OF BIGHORN SHEEP AND MULE DEER WINTER RANGES N. T. Hobbs and R. A. Spowart INTRODUCTI QN An important objective for managing populations of mule deer and mountain sheep in Colorado is to increase the carrying capacity of their ranges through habitat manipulations including timber management, mechanical treatment, and prescribed burning (Colorado Division of Wildlife 1983). For several reasons burning is a particularly appealing technique; it is relatively cheap (Duvall 1970, Kufeld 1983); it is widely used by federal land management agencies (Scotter 1980), and mule deer and mountain sheep appear to be attracted to burned areas. Despite these unambiguous practical and economic advantages of burn treatments, it remains uncertain wnether burning offers any direct biological benefits to these animals. Although there is evidence that the nutritional quality of forages improves following fire, these improvements often are small and transitory (reviewed by Bendel 1 1974). Such changes in forage qual ity may have little impact on selective feeders like mule deer and mountain sheep (Hobbs et al. 1983). Consequently, we are left with the question, "Does prescribed burning improve habitat for mule deer and mountain sheep in a demonstrable, meaningful way?" Here, we report experiments designed to answer that question and to provide information on how burning can be done most effectively to benefit mountain sheep and mule deer. P. N. OBJECTIVES Objectives of our studies included the following: 1. Quantify the effects of burning mountain shrub and grassland communities on nutritional status of mule deer and bighorn sheep during winter. 2. Determine the change in nutritional carrying capacity of mountain shrub and grassland winter range for mule deer and bighorn sheep which is brought about by burning. 3. Examine the effect of fire on food niche relations and ecological separation of mule deer and bighorn sheep. 4. Explain changes in responses of forage resources, both quantity and quality, in terms of processes in the nitrogen cycle. DESCRIPTION OF AREA We conducted our experiments in a south-facing valley in the Front Range Mountains at 2300 m in elevation, 2.5 km NE of Rustic, Colorado. The area offers winter and spring range for populations of mule deer, elk, and mounta in sheep;"· *The accepted common name for bighorn sheep was changed to mountain sheep after the objectives of this study were written. �286 We worked in 2 upper-montane (Marr 1967) plant communities--a grassland dominated by bluebunch wheatgrass (Agropyron spicatum), needle and thread grass (Stipa comata), and Kentucky bluegrass (Poa pratensis), and a mountain shrub community containing predominantly big sagebrush (Artemi:sia tridentata) and wax current (Ribes cereum). Important understory species in the mountain shrub community included mountain muhly (Hulenbergia montana), sulfur flower (Eriogonum umbellatum), bluebunch wheatgrass and needle and thread grass. Soils in these communities are formed· of decomposed granite and mica-schists and are classified as Dystric Eutrochrepts. j \ METHODS AND MATERIALS Experimental Design and Fire Measurements We chose 3 pairs of plots in each plant community, the members of each pair being similar in slope, aspect, vegetative cover, and soil moisture. Plots were 0.3 ha in size in grassland and 1.0 ha in mountain shrub. Pretreatment fuel loads were estimated according to Brown et al. (1982)(Table 1). Burn treatments were assigned randomly to one member of each pair. Treatment plots were burned according to prescription {Table 2) on 29 and 30 September 1979. Flame height and rate of spread were estimated visually by fire observation teams. Energy release was estimated by water can analogs (Beaufait 1966) . Table 1. Pre-burn fuel loading (kg/ha) contributed ground and surface fuel components Fuel category Herbaceous Grassland Mountain Shrub vegetation Live 53 42 Dead 124 77 Letter 119 200 Downed wood 305 700 Shrub 739 2661 -- 1340 3680 Total by �287 Table 2. Prescribed and achieved conditions mountain shrub and grassland communities.a for experimental Achieved Prescription Prescribed Time of day range 1000 - 1400 Air temperature Relative humidity Mountain shrub 1700 - 1730 1000 1200 1200 - 1700 c 17 - 21 C 15 - 19 13 - 35% 29 - 32% Wind velocity 5 - 17 kph Wind direction S - SE range Grassland c 7 - 26 burns in 30 - 31% 6 - 14 kph 8 kph S - SE E - NE aData from Omi and Laven, in preparation. FIRE BEHAVIOR Prescribed burns were more intense and more homogeneous in distribution in the mountain shrub community than in grassland (Table 3). This difference resulted from greater fuel loads in mountain shrub and the more level topography in grassland. Two grassland plots burned incompletely because they were in afternoon shadows at the time of ignition. Table 3. Fire behavior characteristics communities. Fire type Plant community in mountain shrub and grassland Rate of spread (m/sec) X Flame length (m) SE X SE a Energy release kcal X SE Mountain shrub back fire 0.01 0.001 1.3 0.5 32.0 8.8 Grassland head fire 0.03 0.01 0.5 0.2 7.8 3. 1 .flank fire 0.05 0.01 c c aFireline intensity (I) may b~ computed directly from flame length(F) by the formula: I = 5.76F2. 17· bDeri~ed from water-can analogs. cNot measured. Following treatment the perimeter of each plot was fenced with 2.5-m high panels of nylon netting strung between steel posts 5 m apart. We fenced plots to facilitate controlling experimental animals and to prevent confounding effects of grazing by native ungulates. Methods specific to each experiment will be covered in sections describing that experiment under Results and Discussion. b �288 RESULTS AND DISCUSSION Experiment I: Fire Effects on Nutrition of Mountain Sheep and Mule Deer During Winter and Spring Literature Review and Rationale It is widely believed that fire benefits ungulates by enhancing their nutrition. Although intuitively appealing. this belief is largely unfounded; evidence on the effects of fire on ungulate nutrition is highly equivocal. Some workers have shown that the nutritional quality of forage improves following fire (Dewitt and Derby 1955. Swank 1956. Lay 1957. Smith and Young 1959. Leege 1969. Hallisey and Wood 1976. Springer 1977. Wilms et al. 1981b) but others found no improvement (Leege 1969. Dills 1970. Christensen 1977. Merrill et al. 1980. Meneeley and Schemnitz 1981. Rowland 1981), or demonstrated a detrimental effect of fire on forage quality (Swank 1956, Koelling and Kucera 1965, Leege and Hickey 1971, Young and Bailey 1975, Springer 1977). This lack of consensus is compounded by ambiguities in interpreting the nutritional significance of fire-induced changes in forage. Inferences on the nutritional quality of animal diets drawn from observations on the quality of forages are unreliable whenever animals feed selectively. However, measures of fire effects on diet quality are rare (Taber 1953, Rao et al. 1973, Rowland 1981). Here we test the hypothesis that prescribed burning improves the nutritional quality of diets of mule deer and mountain sheep grazing in montane plant communities during winter and spring. Methods We began our observations 13 months after treatment. During November. January, March, and May of 1980-81 and 1981-82 we conducted grazing trials to examine fire effects on nutritional quality of diets of tame mule deer and mountain sheep. Our deer included 1 castrated male and 3 females. All were approximately 6 months old during the first trial. We observed 2 female and 2 castrated mountain sheep who were 3-4 years old. Animals were reared and trained according to Neil et al. (1979) and Hobbs and Baker (1979). Prior to each trial animals were transported from pens at Fort Collins, Colorado, to the study area and were allowed to graze and wander freely on burn and control plots for 2 days prior to data collection. For 6 days following this acclimation period animals were observed daily on a single burn-control replicate. At the beginning of an observation period, a pair of animals (2 sheep or 2 deer) was released on a burn or a control plot. After 30 minutes of observation, we switched the animals to the other plot in the replicate for another 30 minutes of grazing, and then moved them back and forth again for 2 additional 15-minute observation intervals. Half of our group of animals was released initially on control plots, the other half on burns. By balancing where animals started feeding, we prevented effects of satiation from biasing burn-control comparisons. �289 A single observer counted the number of bites of each forage eaten by each animal. We differentiated our counts among plant species and plant parts and between green and dead material. Simultaneously, another observer handplucked forage samples for nutritional analysis. These samples mimicked the material consumed by experimental animals~ At the conclusion of each trial, we collected additional samples for estimation of bite weight. These procedures were described in detail by Hobb~ (1979), Hobbs et al. (1979), and Baker and Hobbs (1982). We defined principal forages as plant species and plant parts contributing 2% or more of bites eaten by mule deer or mountain sheep on any plot. Samples of principal forages were analyzed for crude protein, dry matter, and ash according to A.O.A.C. (1965). In vitro digestible organic matter (IVDOM) was determined by the method of Tilley and Terry (1963) as modified by Pearson (1970). Organic matter digestibility coefficients were calculated based on ashed residues as recommended by Alexander and McGowan (1966). Rumen fluid was obtained from a fistulated Holstein cow fed native grass hay for in vitro determinations of winter samples, and fed 3rd-cutting alfalfa hay for determinations of spring samples. Diet of the donor cow was changed to reflect seasonal shifts in protein content of diets or mule deer and mountain sheep. There is conflicting evidence on the importance of inoculum donor to prediction of IVDOM (Troelsen and Hanel 1966, Ward 1971, Robbins et al. 1975, Palmer et al. 1976, Mould 1980:67, Palmer and Cowan 1979, Zeeman and Coetsee 1984 Pedersen and Welch 1982). However, we decided that for the purpose of compa~ing digestibility of diets among treatments, the control gained by using a single, reliable innoculum source outweighed the potential value of collecting fluid from wild sheep and deer (Clark and Mott 1960, Scales et al. 1974, Milchunas and Baker 1982). A single in vitro run was made for each month1s forage samples. Differences among runs were tested with a one-way analysis of variance on 3 laboratory standards included in each run. Because we found no differences (p > 0.42) among in vitro runs, data were not adjusted to compensate for among-run variation (Clark and Mott 1960). We calculated diet digestibility and crude protein content on an organic matter basis as the sum of products of forage quality values of principal forages times their normalized proportions in ungulate diets (Hobbs et al. 1979:48-49). Calculations were performed on an organic matter rather than a drymatter basis to prevent bias from soil contamination (Alexander and McGowan 1966). We were concerned about this problem because much more bare ground was exposed on burned plots than on controls. This difference offered a potential confoundment because analyses of forage samples collected from burned plots were more likely to be influenced by adhering particles of soil. Differences in diet quality values were analyzed with a factorial analysis of variance for a randomized complete block design with repeated measures. We replicated by plots and animals and repeated over months. Blocks, animals, months, and years were considered random effects. Differences among individual means were established with Bonferroni t statistics (Miller 1966) at a simultaneous confidence level of P < 0.10. We examined treatment effects on forage quality with paired t-tests. �290 Results With few exceptions1 effects of prescribed burning on nutritional quality of forages of mu 1e deer and mounta in sheep tended to be sma 11 (Tab 1e 4). Th is tendency was particularly apparent for forage digestibility which was enhanced only in grasses from burned grassland plots, only during the first winter. Forage protein was somewhat more s~nsitive to treatment. We observed increases (p < 0.10) in grass p rote i n 'in both plant communities, and increases in protein content of forbs during both years. These difference s va lthouqh significant, were usually less. than a few percentage points. j a Table 4. Crude prote1n (% of organic matter) and IVDOM content of forages of mule deer and mountain sheep in burned and unburned mountain shrub and grassland communities during 2 years following prescribed burning. P 1ant Seasonc Community _Year Mountain shrub 1980-81 winter spring 1981-82 winter spring Grassland 1980-81 winter spr ing 1981-82 winter _l " spri ng a IVDO Mb Burn Control X Forage grass forbs shrubs grass forbsb shrubs grass forbs shrubs grass forbs shrubs grass forbs shrubs grass forbsd shrubs grass forbs shrubs grass forbsd shrubs -X Crude rot e l b ru e voprotein Burn Control 54 41 27 76 66 52 41 25 78 66 13.1 8.9 6. 1 26.3 29.6 ,;', 47 37. 23 74 53 44 37 22 74 61 8:5 9. 1 8.8 20.2 -19.8 ,,;', 43 44 27 77 67 7.3 7.4 6.4 16.2 18.0 "1: 46 39 23 70 61 7.9 8.5 4.8 16.1 19.9 48 46 28 77 68 50 39 25 70 58 -;', it~ "l: "k -}: ,,;', * 'I: ;': 10.1 7.4 5.8 27.5 24.4 5.9 8.0 5.7 21.4 19.8 6.3 6.6 5.7 18.5 19.4 8. 1 7.5 6.3 18.6 20.5 Includes forage spec~es ~o~tr~buting >2% of bites on a burn and control replicate during any month. Samples were collected during grazing trials to mimic material chosen by experimental animals. bMeans separated by * are significantly different at P < 0.05. cWinter = November, January, March; Spring = May. dNo shrub material in diet. �291 In marked contrast to our observations on the quality of individual forages, we found that prescribed burning substantially increased the nutritional quality of winter diets of mule deer and mountain sheep in both plant communities. During spring, we observed no consistent nutritional improvements attributable to burning. . Mountain sheep diets chosen on burned plots in the mountain shrub community during winter contained higher levels of crude protein than their diets selected from controls during both years (treatment P < 0.02, Fig. 1). The size of these differences remained constant during November-March of both winters (treatment x month P > n.12). During May, the effect of treatment was reversed, but this difference was not significant (p = 0.10). Improvements in protein content of mountain sheep diets persisted for 2 years with no reduction in magnitude (year x treatment ~ = 0.26). Similar to mountain sheep, mule deer chose diets higher in protein from burned mountain shrub plots than from controls during both years (treatment P < 0.02, Fig.l). During Year 1, this improvement in dietary protein level increased with advancing season during November-March, but disappeared in May (treatment x month p = 0.007) •. Effects of treatment also depended on month the following year (treatment x month P = 0.02). Averaged over months, the magnitude of fire effects on deer diet protein in mountain shrub declined sharply during Year 2 (treatment x year ~ = 0.02)~ Fire increased the level of IVDOM in diets of mountain sheep in the mountain shrub community (treatment P < 0.03, Fig.2). Size of treatment effects was constant during the first winter (treatment x month.P = 0.10) and remained constant throughout winter and spring of Year 2 (treatment x month P = 0.29). The increase in sheep diet digestibility resulting from burnin~ mountain shrub persisted for 2 years (treatment x year ~ = 0.77). We observed a substantial improvement in digestibility of deer diets attributable to burning mountain shrub communities (treatment P < 0.02, Fig. 2). However, the size of improvements in diet digestibility were sharply diminished during the second year (treatment x year P = 0.005). :\,we also observed va rl.a t lon in fire effects on deer diet IVDOM among months; although size of treatment effects remained constant during winter of Year 1 (treatment x month.~ = 0.35), those effects decreased drastically during May. During Year 2, the effect of burning on deer diet IVDOM was greater during November and March than during January and May (treatment x month ~ = 0.0001). Mountain sheep diets selected from burned grassland plots contained more crude protein than their diets chosen from controls (treatment P < 0.03). During both years the magnitude of fire effects on diet protein-depended on_ month (treatment x month P < 0.01). Although the reversal of treatment effects we observed durin~ May contributed a large component of this interaction, effects of fire also varied in size during the winter months. In contrast to this monthly variation, the effect of burning did not change with year (treatment x year P = 0.26). �292 \ ,. \ \ ,I r f C' ., ! l 25 MOUNTAIN SHEEP c z B w 6 '0:: (L 15 /B, / W 0 / ::J o:: <, / / ,......B// 5 25 * * * NOV JAN MAR MULE DEER ••.... w (L 15 / ,/ ,/ / / B"" "" - B-- MAY - c NOV JAN MAR MAY C B ",,"" B" ,/ B........ ./' ./' --B~ c~c-""""-""'c (.) , * / B " ,/ 0:: " * /. 0 . _ ..J ..' * B ::J \ ," !. ,.-, """" --B" c_____ c c____ W ~ 0 / c/ Z 6 0:: / ",B/ -c (.) ~ 0 ./' B__ c- B" I. ~ c 5 * * * NOV JAN MAR c * * MAY NOV C JAN MAR MAY Fig. 1. Crude protein content (% of organic matter) of diets of mountain sheep and mule deer in burned and unburned mountain shrub communities. Monthly means for burned plots are shown by B, for unburned plots by C. Pairs of monthly means with ~'.below them are different at P < 0.10. Data for 1 year after burning are on the left, 2 years post-burn are on the right. �293 75 ~ 50 o o > eft. 25 MOUNTAIN SHEEP •..•.•.•....•. "'" I~ ...... 8_...... A ..••.•..••.• / S S" CCC * * * NOV JAN MAR MAY * * * * NOV JAN MAR MAY //s;/// 75 ~ 50 o o > eft. 25 /8 C '/ C C S----S// /C--C C C * * * NOV JAN MAR * * MAY NOV JAN MAR MAY Fig. 2. In vitro digestible organic matter (IVDOM) content of diets of mountain sheep and mule deer in burned and unburned mountain shrub communities. Monthly means for burned plots are shown by S, for unburned plots by C. Pairs of monthly means with ,',below them are different at P < o. lO~ Data for 1 year after burning are on the left, 2 years postburn are on the right. �294 Mule deer chose diets higher in crude protein from burned grassland plots than from controls during Year 1 (treatment.P:= 0.007, Fig. 3). However, this improvement declined dramatically the second-year (treatment x year P = 0.03) such that treatment effects only approached significance (p := 0.058)-during Year 2. The magnitude of effects of burning on deer diet protein levels depended on the month diets were observed (treatment x month ~ < 0.03). Prescribed burning elevated IVDOM levels of diets of mountain sheep in grassland during November and January of Year 1 (treatment.P = 0.0003, Fig. 4). During the remainder of that year and during Year 2, there was no difference (p > 0.10) in IVDOM content of sheep diets selected from burned plots compared to diets from controls. Mule deer chose diets containing higher levels of IVDOM from burned grassland plots than from control~ during November-March of Year 1 (treatment p=0.004, Fig. 4). During May we observed no effect of treatment (p > 0.10). Similar to mountain sheep, mule deer chose diets consistently, but-not significantly higher in IVDOM on burned grassland plots compared with controls during Year 2. Discussion Our observation that effects of fire on forage quality tend to be small agrees with results of other worker:s. Improvement in forage protein and digestibility resulting from burning frequently average less than few percentage points (reviewed by Bendel 1 1974:101). Given our findings on forage quality, it may seem paradoxical that ungulates were able to choose diets of substantially higher qual ity on burned plots compared with controls. This result can be explained, though, by changes in diet selection. The nutritional quality of herbivore's diet is determined by the quality of individual forages it eats, as well as the way that animal chooses among forages of differing quality (Ellis et al. 1976). Quality of forages was changed little by fire; however, patterns of diet selection shifted markedly. In particular, mule deer and mountain sheep ate more green grass, primarily leaves of Kentucky bluegrass (Poa pratensis), b1uebunch wheatgrass (Agropyron spicatum), and cheatgrass (BrOiilUstectorum), on burned plots than controls during winter (Spowart and Hobbs, unpublished data). Much of the variation in ungulate diet quality was explained by the percentage of green grass in ungulate diets (Table 5). This was the case because green grass was much more nutritious than other forages during winter. For example, its crude protein content often exceeded 25% when other forage averaged 3-5%. As a result, the amount of green grass an ungulate could find and eat during winter profoundly affected the qual ity of its diet. The importance of green grass as an influent on ungulate diet quality has been emphasized by others (Hamilton et al. 1973, Field 1976, .Lang1ands and Sanson 1976, Langlands and Holmes 1978, Jarman and Sinclair 1979, Shinn 1980). �295 25 MOUNTAIN SHEEP c z w b 0: a.. 15 w o ::::> 0: u ?f? 5 25 * * NOV JAN * MAR MAY * NOV JAN MAR * * * NOV JAN MAR MAY MULE DEER z w b 0: a.. 15 w o ::::> 0: u ?f? 5 * NOV * JAN MAR MAY MAY Fig. 3. Crude protein content (% of organic matter) of diets of mountain sheep and mule deer in burned and unburned grassland communities. Monthly means for burned plots are shown by B, for unburned plots by C. Pairs of monthly means with ,',below them are different at P < 0.10. Data for 1 year after burning are on the left, 2 years post-burn are on the right. �296 75 ~ 0 50 0 > ~ 0 /8 8----8_ C//// /-/ MOUNTAIN SHEEP . c 9 /,../ __9::---C. 9 9-C----- . C ~C C 25 75 * * NOV JAN MAR ~ 0 50 9/ ./ ,.. NOV C 9 ____ 9-- C MAY MAR JAN C " ~C 0 > MAY MULE DEER 9-_ ./ --9/ ./ ~ 0 /8- c C C -- fC /9-;/9 / 9/ . C 25 * * * NOV JAN MAR MAY NOV JAN MAR MAY Fig. 4. In vitro digestible organic matter (IVDOM) content of diets of mountain sheep and mule deer in burned and unburned grassland communities. Monthly means for burned plots are shown by B, for unburned plots by C. Pairs of monthly means with * below them are different at P < 0.10. Data for 1 year after burning are on the left, 2 years post-burn are on the right. �297 . (R2)a. In nutrltlona .. 1 quality of winter Table 5. P ercentage 0 f· variation diets of mule deer and mountain sheep explained by the proportion of green grass in their diets. Mule deer Crude protein Mountain sheep Crude protein IVDOM Plant Community Year IVDOM Mountain shrub 1980-81 1981-82 85 37 84 46 65 29 72 63 Grassland 1980-81 1981-82 56 67 65 52 a NS 23 57 32 aN = 72 for each regression. P < 0.05 unless marked NS. All regressions are significant at Green grass on burned plots was nutritionally similar to green grass on controls. However, animals appeared to be able to find it much more readily on burned plots, apparently because it was more available. We observed more green grass on burns than controls and attribute this difference to 2 causes. First, green grass on controls tended to be obscured by down litter and standing dead herbage. Old growth of grasses can obstruct feeding of mule deer on basal tillers (Willms and McLean 1978, Willms et al. 1980, Willms et al. 1981a). Second, we suspect that soil conditions on burned plots favored grass growth. Soils of burned areas tend to be warmer than soils from unburned ones (Weaver and Rowland 1952, Kucera and Ehrenreich 1962, Scotter 1963, Old 1969, Lloyd 1972, Peet et al. 1975). This tendency results from enhanced absorption of sunlight by blackened soil surfaces unshaded by plants and litter (Woodmansee and Wallach 1981). Such warming could stimulate the growth of cool season grasses we observed on burned plots throughout winter. Our finding that the improvement in quality of diets caused by burning exceeded the improvement in forage quality which resulted from fire offers an important caution in interpreting studies of fire effects. We warn that inferences on benefits of fire for ungulates based on studies of forages alone may severely underestimate the improvements in ungulate nutrition which result .from burning. This finding may explain the seemingly anomalous observation that preference of ungulates for burned areas persists long after differences in forage quality are detectable (Lay 1957, Davis 1977, Asherin 1974, Peek et al. 1979, Oefinger and Seifes 1978). Prescribed burning improved the quality of ungulate diets during winter but provided no consistent nutritional benefits during spring. During May the nutritional quality of diets from control plots almost always exceeded the quality of diets from burns, although this difference was not significant. We surmise that the nutritional advantage attributable to burning during winter was lost in May because of green-up of high quality forage. Ubiquitous growth of green forage allowed animals to select nutritious diets wherever they fed. However, reversal of treatment effects during spring may be related to an effect of fire which is nutritionally beneficial to ungulates. With few exceptions forages were higher in quality on controls than burns during May �298 (Table 2). We suggest that spring forages were nutritionally superior on controls because they were phen~logic~lly younger. Similar to Kucera and Ehrenreich (1962), we observed t~at green-up occurred 1-2 weeks earlier on burned plots than controls, usually in late April or early May. Such patterns are plausible because of enhanced warming of burned soils we described earlier. Differences between burn and control plots in initiation of plant growth may benefit ungulates by offering 2 temporally distinct flushes of nutritious plant tissue, early on the burn, later on the control. In this way, burning may prolong the period when .young forage is available to ungulates and may, in effect, shorten the nutritional deprivation of winter. Effects of prescribed burning on ungulate nutrition were more per~istent in the mountain shrub community than in grassland. Because burns were cooler and more heterogeneous in grassland then in mountain shrub, it is difficult to infer whether the relative transience of fire effects in grassland was attributable to characteristics of the community itself, or to effects of a less intense burn resulting from differences in time of ignition. Despite this confoundment, we suspect that fire effects were more short-lived in grassland because burns were less complete there. The patchy nature of burns in grassland plots allowed conditions to return to pre-burn status much more rapidly than in mountain shrub. Specifically, standing crop burned grassland plots equalled biomass on the control by the end of the second growing season following fire in contrast to mountain shrub plots where the standing herbage remained only 40% of the control value during Year 2 (Hobbs and Spowart, unpublished data). Many of the effects of fire on diet qual ity we observed depended on the presence of green grass on burned plots. The increased crop of dead herbage in grassland during Year 2 appeared to reduce the availability of this important forage. Experiment II: Simulation Modeling of Fire Effects on Energy and Nitrogen Balance of Mule Deer and Mountain Sheep Literature Review and Rationale Few studies have shown that the changes in plant communities following fire improve the condition of animals which use those communities. Although domestic livestock have shown greater weight gain feeding on burned pastures and rangelands compared with unburned areas (Lemon 1946, Duvall and Whitaker 1964, Anderson et al. 1970, Smith and Owensby 1973, McGinity et al. 1983), improvements in the nutritional status of wild animals following burning scarcely have been studied. With few exceptions (Springer 1977, Rawland 1981) the only evidence showing that fire enhances the condition of wildlife comes from studies of blacktailed deer (Odocoileus hemionus columbianus) in the California chaparral (Taber 1953, 1956, Taber and Dasmann 1957). Measuring changes in animal condition directly is extremely difficult and expensive. Consequently, we chose to estimate those changes indirectly using computer simulation. �299 Methods We used a generalized model of energy and nitrogen balance for ruminants (Fig. 5, Swift 1983) to simulate changes in nutritional status of mule deer and mountain sheep following fire. Flow of energy, nitrogen, and microbial biomass are simulated in 3 separate submodels. Material moved varies with submodel; in the energy submodel, units are kilocalories; in the nitrogen and rumen microbe submodels, units are grams of nitrogen and microbial protein, respectively. The model is generalized in the sense that it can simulate functioning of a variety of ruminant species by altering parameters which describe an animal's digestive physiology and meta601ism. The model requires 47 input parameters, 15 of which are species specific (Table 6). Principal driving variables are diet digestibility values and nitrogen concentrations over time, and daily minimum and maximum air temperatures. Weather data were taken from National Oceanic and Atmospheric Administration records (1980, 1981, 1982) for Red Feather Lakes, Colorado, approximately 11 km north of our study area. The ~odel is constructed of 33 difference equations operating at 1 day time steps simulating a single animal. The model predicts energy and nitrogen balance in response to changes in diet digestibility and nitrogen content and ambient temperature. Rates of forage intake, digestion and metabolism of energy and nitrogen, partitioning of energy and nitrogen within the body and losses of those materials from the body are predicted. Energy and nitrogen balance predictions allow estimation of animal weight changes as well as changes in body composition. All input parameters, variables, and constituent equations were described by Swift (1983). Driving variables for the model, diet digestibility and protein content, were determined in Experiment 1. We ran 16 simulations to examine the potential effects of fire-induced differences in diet quality on animal nutritional status. All model input parameters except diet digestibility and nitrogen concentration were the same for simulations of each species feeding in burned or unburned plant communities. Mean values for IVDOM and nitrogen concentration of mule deer or mountain sheep diets eaten during each grazing trial were input at trial dates. Ambient temperature input ranged from· -200 F to 75° F and -250 F to 670 F for 1980-1981 and 1981-1982 runs, respectively. Simulations were initiated on 16 November and terminated on 25 May. Based on the age and size of our experimental animals, we assumed that simulated deer weighed 32 kg and were 210 days old, and sheep weighed 80 kg and were 1278 days old at the beginning of Year 1. Second year simulations were initiated with 50 kg, 575 day old mule deer or a 90 kg, 1643 day old mountain sheep. Hypothetical deer had fat reserves of 10% of their body weight; sheep, 15%. Results Burning improved the nutritional status of simulated mule deer and mountain sheep in both plant communities during both years. In mountain shrub during �300 Rumen Microbial Protein X21 Fig. 5. Generalized model of energy and nitrogen balance of nutritional status of mule deer and mountain sheep. used in sumulation �301 Table 6. Species specific parameters used in simulation nitrogen balance of mule deer and mountain sheep. Parameter Mule Deer 28.8 (Year 1) 45.0 (Year 2) Init ia 1 fat (kg) (Yea r 1) 3.2 (Yea r 2) 5.0 Lower critical temperature (oC) o Cca Thermal conductance 1. 7 e Passage rate of digestible forage 0.65 f Passage rate of indigestible forage 1.0-1.3 Fat threshold for intake regulation 1.0~ Metabolic fecal N (g/kg drymatter intake) 7.6 h Endogenous urinary N (g/wt·75/d) .115 Maximum life span (days) .1800.0 Initial lean body (kg) of energy and .Mounta in Sheep 68.0 76.5 12.0 13.5 -20b d 1. 4 e . 0.45 f 0.7-0.9 1. 5: 7.31 •102 i 2190.0 aWeiner. (1977) b Chappel and Hudson.(1980) CWesley d (1971) Chappel and Hudson (1978) eModel tuning fMautz and Petrides gTorbit (1971) (1981) hRobbins et al._(1974) i Calculated from Hebert.(1973) Year 1,simulated deer grazing on controls were predicted to lose 100%of their fat reserved and 37% of their lean body during the winter months (Fig. 6). On burned plots in mountain shrub, simulated deer gained weight, primarily lean body, throughout winter. Thus, control plots offered diets deficient in both energy and nitrogen whereas burn plots provided levels of dietary energy and nitrogen sufficient for growth. During spring, simulated deer rapidly regained the weight lost during winter on control plots. However, this regrowth was not adequate to equal the steady overwinter growth by simulated deer on burn plots. By mid-May, simulated deer on burns weighed more than 10 kg more than simulated deer on controls. During the following year, the nutritional benefits of burning mountain shrub for simulated deer were substantially smaller (Fig. 7). Although simulated deer lost weight on controls and gained weight on burns during winter, there was no difference in lean body weight of simulated deer grazing on burns or controls by late May. We observed similar effects of burning on energy and nitrogen balance of simulated mul~ deer in grassland (Figs. 8,9). On controls, simulated deer lost weight throughout both winters, but they gained weight on burns. The rapid regrowth of simulated deer on control plots during March through May showed that burning has no effect on spring nutrition. . �\.N . 0 N· 60, SHRUB CONTROL 60 56 56 52 52 ~ G 40 ~ I- 36 TOTAL 'I LEAN BODY I ~ 32 ~ 28 w , 'I ~ >- g .- 44 44 - J TOTAL 48 48 G BURN 1 SHRUB .-' 40 I- 36 I ~ 32 ~ 28 w ••.... ••.... ••.... - ...•.. _ ...•.. ...•.. ...•.. ------ __ LEAN BODY ,.....""" >- g 24 24 CD CD 16 16 12 1 12 :j ...•... 8 4--1_._ .•.......•..••. o NOV Fig. 6. unburned DEC '-. JAN FEB MAR APR ._FAT MAY 0 NOV Simulated changes in body composition and total weight mountain shrub communities 1 year after treatment . .... .",.,..._. DEC ' __ ._'_' __ JAN of simulated FEB _._._._.'_'_' ' MAR mule deer ~-. . APR FAT " . MAY in burned and .-,,:,,--- �76 76 SHRUB CONTROL 72 72 68 68 SHRUB BURN __ --TOTAL TOTAL 64 64 60 -:x: - :x: •.... 52 / •.... 52 / I 48 ~ 48 w 3: 44 / / LEAN BODY .: ,// /,/ >0 o 40 ./ (D (D 16 16 12 12 8 -l 8 ., 4 _---/ I / w 3: 44 >0 o 40 /_-------- B 56 / B 56 ~ 60 / LEAN BODY -_. . -.-.-.-.-.-.-.-.-.~./. / FAT ./ . O~--r---~----~----~--~-----r--~ NOV DEC JAN FEB MAR APR MAY Fig. 7. unburned ."....-._._.- FAT ,/ ,/ .--.-~-.-.O~--r---~----~----r---~----~--~ NOV DEC JAN FEB MAR APR MAY 4 -l -._ Simulated changes in body composition and total weight mountain shrub communities 2 years after treatment. .. ..,.,. ./ of simulated mule deer in burned and \J.) o I..N �w o +:- 60 GRASSLAND 60 CONTROL 56 56 52 52 48 48 TOTAL 8 40 / 36 w ~ I- 36 / I ~ 8 40 'I LEAN BODY ...., ~ ~ ...., I ~ / 32 ~ ~ 28 ~ / ,;- 28 LEAN BODY CD CD 16 16 12 12 8 8 / 4 _._.,;' . ...-. ...•••....•........•..... 0 ."..- ,; ------ _---- >0 o 24 ~'/ >0 o 24 32 W ./ BURN TOTAL 44 44 I- GRASSLAND NOV DEC JAN FEB MAR· APR . MAY FAT 4 _._._._._._._._._._._._ -1_._'- OJ_--~--~r---~-----r----~----~--~ NOV DEC _--_--_- Fig. 8. unburned Simulated grassland FAT changes in body composition and total weight communities I year after treatment. JAN .... _-_ FEB .. of simulated ----_. __ .. MAR _-_ .... APR MAY -_-_ .. _-_-------_.-- __ ._. mule deer in burned and �76 76 GRASSLAND BURN 72 _TOTAL 68 64 64 8' 56 w 3: 44 >- g - _/ ---_/ 60 LEAN B ODY ",--- ~ ~ 52 :r: ~ 48 w 3: 44 >- g 40 BODY I / •....... ",// ••••LEAN / 8' 56 / ~ •.•..... ~ 52 :r: ~ 48 _------- /' CONTROL TOTAL 68 60 GRASSLAND 72 / / --__ -,--------'-- 40 -: " / / / '" II) II) 16 16 12 /._._._._._ FAT 12 .,- . ...._ ._ 8 4 '-. .•..•• /' --._. __ .......• .-.' ./. O~--r---~----,-----r---~----~----, NOV DEC JAN FEB MAR APR' MAY Fig. 9. unburned Simulated grassland 8 4 ._ - ./ ,,/ ._._._. __ • __ • __ ._ FAT ./ .*""""o O~~r----r----r----.----,---~----~ JAN FEB MAR' APR . MAY NOV DEC changes in total body composition and total weight communities 2 years after treatment. of simulated mule deer in burned and I.N o \J1 �306 Nutritional benefits of burning were smaller for simulated mountain sheep than for mule deer. In mountain shrub during Year 1, simulated sheep grazing on controls during winter were predicted to lose 63% of their fat reserves (Fig. 10). On burned plots, they increased their fat stores by 43% during the same period. During spring, simulated sheep regained some of their fat reserves lost on controls and rapidly gained lean body weight. However, these gains did not equal the overwinter and spring weight gains of simulated sheep grazing on burns. By the end of May, simulated sheep on burns weighed 10 kg more than simulated sheep on controls. We observed similar differences in the nutritional status of simulated mountain sheep grazing on burns compared with controls in grassland during Year 1 (Fig. 11). During Year 2, the nutritional lated sheep were not as great plots, losses of fat reserves in lean body, less. However, over 6 kg more than simulated benefits of burning mountain shrub for simuas we observed previously (Fit. 12). On burn were greater than the year before and gains in late May, simulated sheep on burns weighed sheep on controls. On grassland controls during Year 2 simulated sheep initially lost a much smaller percentage of their fat reserves during the second winter (Fig. 13). However, during spring, weight gain of simulated sheep on controls was fast enough to surpass the steady weight gain of simulated sheep on burns. By late May simulated sheep on controls weighed nearly 9 kg more than simulated sheep on burns. Discussion We demonstrated in Experiment 1 that fire caused statistically significant increases in the nutritional quality of diets of mule ,deer and mountain sheep feeding in grassland and mountain shrub communities during 2 years following treatment. The question remained, however, whether these differences were physiologically significant. To address that question, we simulated energy and nitrogen balance of mule deer and mountain ~heep obtaining diets similar to those measured on burned and control plots. Simulations suggest that burning can substantially enhance the overwinter condition of those ungulates, particularly during one year following treatment. Fire-induced changes in mountain shrub and grassland communities presented overwinter weight loss and, in some instances, allowed substantial growth by wintering ungulates. Our results indicate that mule deer benefited from burning to a greater extent than mountain sheep. This result is reasonable because burning elevated the digestibility and crude protein content of deer diets more than sheep diets (see Experiment 1). Moreover, the simulated effects of fire on animal nutritional status decreased in second year simulations in response to the decline we observed in deer and sheep diet quality, and changed monthly as the quality of animal diets varied within years. For example, deer and sheep grazing control grassland plots in January 1981 obtained unusually high dietary protein which was reflected in simulations of body weight dynamics. Between March and May of both years, there were large increments in dietary nitrogen and energy on control plots. Simulated animals dramatically increased their weight on control plots at this time. �116 116 SHRUB CONTROL 112 _-------- 104 104 TOTAL 100 100 96 96 r" 92 ~ - I / I a 72 0 ...•.. ...•.. 64 ",,"-_...-"'" _- -- I / _/ 80 I / / / / / I b 0 72 CJ) 68 I / / 64 20 20 /._._.-.-.-.-_._.,.....,.- 16 16 12 ~ . I 3: 76 I >- 84 LEAN BODY / ijj / 3: 76 - I- I 80 68 ~ I 84 _------------ 92 -(!) 88 I 88 ijj CJ) LEAN BODY I I- ~ TOTAL 108 108 B SHRUB BURN 112 12 ._._._ .-.-._._ 8 /' 8 4 4 O~I--~----~----~----~----~----~----~ i DEC i JAN i -._._._o/ /FAT .-....--.--..._._.". NOV FAT ,/ FEB i MAR i APR i MAY i O~~----~---,----~---.----r---~ JAN FEB MAR APR MAY NOV DEC Fig. 10. Simulated changes in body composition and total weight unburned mountain shrub communities 1 year after treatment. of simulated mountain-sheep in burned and w o -.....J �v~ 116 116 GRASSLAND CONTROL TOTAL 108 108 104 104 100 100 96 96 92 88 - 84 ::.:: I / 80 ~ 76 W / >- o 72 0 CO 68 -_ <, •..•. / -> _-_ / / -./ - 84 w I ~ 76 >o 72 0 CO 68 / 64 64 20 20 16 '. 12 8 ....•... .•.... " 4 '- /., ._._._._./ ~./ 12 / Fig. 11. grassland DEC JAN FEB MAR / I I / / / // ........ _._.-::.-.-.- FAT .".....-.-.-.-.-.~./ 8 ./ 4 o~~----,---~----~--~----~--~ NOV / / 16 /fAT .•.... / § 80 I <, 88 I- BODY I <3 ::.:: / / /_----------LEAN 92 / LEAN BODY I- G TOTAL 112 112 <3 . ex> GRASSLAND BURN APR MAY Simulated changes in body composition communities 1 year after treatment. of simulated O~--r---~----~----r---~----'----. JAN FEB MAR APR MAY NOV DEC mountain sheep in burned and unburned �116 116 SHRUB CONTROL SHRUB BURN 112 .112 -----TOTAL .> TOTAL 108 108 104 104 100 100 I 96 / ~ 84 I- / --............. is 0 72 CD 68 ---_ / / / ///"""" ~ 80 jjj 3: 76 I ------..", / ::.:: / ~ 80 ijj 3: 76 / -(!) 88 / ~ ~ 84 I- / 92 I I S 88 LEAN BODY / I I 92 /_-------- 96 LEAN BODY is o / /// / 72 CD 68 64 64 20 20 • 16 16 ..•.. 12 .,<, _._._.-.-.-._..••.•.. 8 ._ -.•.•... ...,... .• FAT .."". / ,._. 12 _._._.-- ..".,-._._.- ·-·-FA . / T ..•...... ..".,.. 8 4 4 n~I__~~ NOV i ~ _ __ LJt:1,; i ~ JAN i ,FEB i -r MAR i -r APR i O~-,~--r--~--r--~--.-~ ~ MAY i NOV DEC JAN FEB MAR APR MAY \oN Fig. 12. Simulated changes in body composition and total weight unburned mountain shrub communities 2 years after treatment. of simulated mountain sheep in burned and o I.D �126 GRASSLAND w o CONTROL TOTAL 122 GRASSLAND BURN 116, __ 118 112 114 108 110 104 106 100 102 96 ------ TOTAL ,.......-------- LEAN BODY 1 98 G I 94 86 I w ~ I ~ 78 0 !Xl74 -__ ..... 70 __ / 80 / Cl o 1/ / 72 64 20 /-FAT 26 .•.•••._._._._.-._._._ 16 ./ .... "..-", 22 14 / rn 68 _/ .-..----- 30 18 / ./ / 3: 76 >- _ ... / / W / >Cl ", I- / 3: 82 88 -- 84 I I- ~ G / ~ •..•. 90 / 92 1- LEAN BODY . --._._. __ ....,... _.-._ ..•.••...•.••. DEC JAN _."" .. .""" 8 4 O~--'-----r---~-----r----'-----r-~~ 12~~~~_,--~_r~~,_~~,_~_,~~~ NOV ..._.._ 12 FAT FEB MAR ••• APR M •• • __ MAY "_ ._ ._ • NOV DEC JAN FEB MAR APR MAY _ Fig. 13. Simulated changes in body composition and total weight and unburned grassland communities 2 years after treatment. of simulated mountain sheep in burned �311 Although the absolute values of model predictions of energy and nitrogen balance may be biased, we believe the magnitude of the differences between treatments and controls offers ~ reliable indicator of fire effects on animal condition. This is because biases in the model should operate similarly on simulations of animals on burns and controls. Thus, simulations are useful for comparing effects of divergent dietary regimes on nutritional status, even if the predicted values for each treatment are biased. The patterns of weight gain and loss of simulated deer and sheep in unburned plant communities correspond well with measured weight flux of these ungulates (Wood 1962). Exceptions to this correspondence include the predicted gain in weight by mule deer consuming high quality diets on burned plots; deer ofte~ lose some weight during winter, even when consuming nutritious diets because of hormonal effects on metabolism (McEwan 1975). This phenomenon is not represented in the model. In addition, patterns of catabolism of lean body and fat do not occur as predicted. The model predicts fat reserves will be totally catabolized before any lean body is lost. That is, the model assumes that maintenance of lean body has a higher priority than maintenance of fat. Torbit (1981) found no such clear cut priority exists; mule deer catabolized both lean body and fat from the outset of negative energy balance. We conclude that prescribed burning can substantially enhance the physical condition of mule deer and mount~in sheep in mountain shrub and grassland communities. The differences in patterns of weight loss we observed would likely result in differences in animal reproduction (Verm~ 1963, 1965, 1969, 1979) and survival (deCalesta 1977).;' Experiment III: Fire Effects on Nutritional Carrying Capacity of Mountain Shrub and Grassland Habitats Literature Review and Rationale Many have observed that large herbivores are attracted to plant communities which emerge following fire (Lay 1967, McCulloch 1969, Kruse 1972, Miller and Watson 1973, Davis 1977, Roppe and Hein 1978, Singer 1979, Riggs and Peek 1980, Willms et al. 1980). This attraction is often explained by nutritional benefits offered by herbage from burned areas. Enhanced nutrition, in turn, is believed to stimulate growth of herbivore populations feeding in burned habitat (Leopold 1950, Taber 1952, Taber and Dasmann 1957, Biswell 1961, Miller and Watson 1973, Vierreck 1973, Salwasser et al. 1978, Scotter 1980). However, the nutritional explanation for the effects of fire on the habitat preferences of individuals and the numerical responses of populations appears at odds with previous reports of fire-induced changes in herbage. Standing crops of plants may be reduced by fire (Oaubenmire 1968, Harniss and Murray 1973, Miller and Watson 1973, Nimir and Payne 1978, Young and Evans 1978, Uresk et al. 1980), and the nutritional quality of herbage frequently changes little (reviewed by �312 Bendel 1 1974) or may even decline following burning (Swank 1956, Koelling and Kucera 1965, Le~ge and Hickey 1971, Young and Bailey 1975, Springer 1977). Thus fire may diminish the absolute abundance of potential food for herbivores. We propose that the effects of fire on herbage can be reconciled with its effects on herbivores by focusing on the relative abundance of food. In a given standing crop of herbage there is a fraction of the total plant biomass which is nutritionally adequate to meet 'animal requirements for maintenance and production. White (1978) argued that this "good food" fraction exists amid a much larger portion of non-nutritious herbage and that, as a result, herbivores confront a relative rather than an absolute food shortage. That is, there is a deficiency of herbage with adequate concentrations of nutrients, particularly nitrogen, rather than a shortage of food in general. We test the hypothesis that a fundamental effect of fire on plant communities is to reduce a relative shortage of food for ruminant herbivores by increasing the fraction of herbage standing crops which contains high concentrations of nitrogen and digestible organic matter. We propose that fire increases the nutritional carrying capacity of grassland and mountain shrub habitats for mule deer and mountain sheep by increasing the proportion of "good food." Methods We estimated the aboveground biomass of a large array of herbage categories (Table 7) at the end of the growing season 1 and 2 years after treatment. The members of this array were chosen to maximize variation in nutritional quality among categories, and to include important forages consumed by mule deer and mountain sheep. Biomass'was estimated by clipping 30 1/4 m2 plots in each burn and control replicate, separating the contents into the appropriate categories, drying them at 1000 C for 48 hours and weighing them to the ne.are s t 0.1 gm. Ten separate samples of each category were also collected from each replicate, composited, and stored at -100 C. These samples were dried at 400 C for 48 hrs and analyzed for drymatter, ash, and nitrogen content according to A.O.A.C. (1965). In vitro digestible organic matter (lVDO~)was determined according to procedures of Tilley and Terry (1963) as modified by Pearson (1970) using innoculum from a fistulated cow fed alfalfa hay. All residues were ashed for calculation of IVDOM (Alexander and McGowan 1966). Distributions of IVDOM and nitrogen within herbage standing crops were described by partitioning the biomass of herbage categories (Table 7) into distinct groups of nitrogen and IVDOM concentration. The herbage biomass in each group was expressed as a total amount (g/m2), and summed and divided by the biomass in all categories and expressed as a percentage of the total biomass (for example, Fig. 1). �313 Table 7. Categories content, and IVDOM. Taxanomic group of herbage analyzed Species categories for aboveground separated biomass, nitrogen Plant parts separated Grasses and sedges All species composited green leaves, dead leaves, culms, inflorescences Shrubs Ribes cereum Purshia tridentata Artemesia tridentata Rubus deliciosus Miscellaneous shrubs leaves, fruits, current growth, old growth Forbs Agoseris glauca Antennaria parvifolia Chrysopsis villosa whole plant except where noted leaves, stems, inflorsences leaves, inflorescences Eriogonum u~bellatum Galium boreale Geranium fremont ii Astragalus spp. Oxytropis spp. Lupinus greenii Thermopsis divaricarpa Potentilla spp. Solidago nuna Miscellaneous forbs leaves, stems, fruits Treatment effects on herbage biomass and nitrogen and IVDOM content were analyzed with a factorial analysis of variance for a randomized complete block design. Replicates and years were considered to be random effects. Differences among individual means were established with Tukey's Q simultaneou~ comparisons at ~ = 0.10. Results With few exceptions, effects of fire on the nutritional quality of herbage tended to be small (Table 8). The exceptions to this tendency were leaves of grasses from burns which generally contained more nitrogen and IVDOM than grass leaves from controls. In the mountain shrub community, leaves of shrubs and some forbs showed some nutritional enhancement resulting from burning. In grassland, no forages other than grasses were improved by fire. Fire caused pronounced shifts in the distribution of nitrogen and IVDOM within plant communities. During both years, control plots in the mountain shrub community were dominated by plant tissue containing dilute concentrations of nitrogen and IVDOM (Fig. 14, 15, 16, 17); more than 75% of the aboveground biomass contained less than 50% IVDOM and less than 1.5% nitrogen. In general, plant tissue containing high concentrations of nutrients on unburned mountain shrub plots was exponentially rare relative to tissue with dilute nutrient concentrations. In contrast, the standing crop in burned mountain shrub contained a preponderance of plant tissue with relatively high concentrations of nitrogen and IVDOM. Although the �Table 8. Mean a concentration (gm!gm om) of nitrogen and in vitro digestible organic matter in herbage from burned an~unbLirned mountain shrub and grassland communities at the end of the growing season. ~1eans separated by~'~are sig-nlfrcant-ly-cHfferentatP-< 0.05. Mountain shrub' % IVDOM Herbage category Grasses green leaves dead leaves culms infloresences Shrubs old growth current growth leaves inf10resences Forbs legumes non-legumes Year Burn 1980 1981 1980 1981 1980 1981 1980 1981 62 '62 57 40 44 32 43 41 1980 1981 1980 1981 1980 1981 1980 1981 10 12 26 27 47 52 _b 40 1980 1981 1980 1981 69 62 55 54 Control -;', it: °l: it: il: ";~ i': ;': blnsufficient material for sample. Control Burn- 52 56 50 47 40 37 40 38 2.5 1.8 2.0 0.8 1.0 0.8 1.5 1.1 11 11 27 27 38 49 34 38 0.5 0.5 0.8 0.7 2.3 1.7 ' b 1.7 60 59 47 52 2.5 2.3 2.1 1.7 aAveraged over reps and species categories. Grassland % IVDOM % N it: it: it: it: it: l i : "k. Burn,. 1.6 1.4 1.4 0.8 0.7 1.1 L2 1.1 68 64 54 44 31 48 39 46 0.5 0.4 0.9 0.8 1.7 1.6 1.5 1.4 13 12 29 26 40 36 _b -b 2.3 2.6 1.5 1.3 65 62 52 48 i': it: ;': "'k % N Control Burn 59 51 47 48 33 49 41 50 2.2 1.7 1.3 0.9 0.5 0.6 1.2 1.2 12 11 25 28 37 39 31 42 0.5 0.4 0.9 0.8 2.1 1.9 1.1 - b 69 69 49 47 1.9 1.8 1.4 1.5 Control 7: 1.5 1.4 1.1 1.1 0.3 0.8 1.3 1.0 0.5 0.5 0.7 0.9 1.8 1.7 1.2 1.6 ;': 2.8 2.4 1.3 1.4 - w -l:" �315 50 BURN ; 20 40 15 . 30 T J_ 1 T 1 10 20 ::r: I'T1 o, 0 5 10 cr (.) (.!) z 0 z 0 ~ 50 I T 1 1 T I 1 1 T I .L . :::0 rn l> G) ITl 1 0 0 :::0 -< s:: cd: l> -i -i CONTROL u, 0 :::0 . ~ 40 0 I'T1 120 G) ....•. s::I\) 30 80 20 10 0 T ~ T 1 40 T 1 - o ~. >0-10 >10-20 >20-30 >30-40 >40-50 >50-60 >60-70 >70-80 >80-90 % IVDOM Fig. 14. Distribution of in vitro digestible organic matter (IVDOM) in burned and unburned mountain shrub communities 1 year following treatment. �316 / 50 BURN f- 60 40 .. 30 f- 40 T 2() a... 0 10 U Z 0 Z I 0 I 20 1 1 0:: (!) T 1 T ~ ::0 CD 1> G> 1"'1 T 1 o I 0 ::0 -< <t I~ 50 CONTROL LL. 0 ~ ~ - 140 ~ ::0 ~ 40 T G> .L r- 120 T 30 <, ~ N 1 - 80 20 T J.. 40 10 T ... 0 I >0-10 I I ; >10-20 >20-30 T ... >30-40 I >40-50 o >50-60 >60-70 >70-80 >80-90 % IVDOM Fig. 15. Distribution of in vitro digestible organic matter (IVDOM) in burned and unburned mountain shrub communities 2 years following treatment. �317 50 BURN 20 , '. 40 15 30 ; I r 10 T -r 20 a... 0 a:: T I <!) z 5 0 i:! 50 1 T 10 (..) 1 1 I . I 1 I I 5. 1 T 1 ::x: :::0 fT1 1 II . tll » Gl fT1 T 0 0 z :::0 -< CONTROL (/) 120 u, 0 3: » -i -i fT1 40 :::!! :::0 0 l 30 80 1 " 40 1 1 0 >0.0-0.3 >0.3-0.6 >0.6-0.9 ...•.. 3: 20 10 Gl >0.9.-1.2 T L 1 T T r >1.2-1.5 >1.5-1.8 ,% >1.8-2.1 1 >2.1-2.4 >2.4-2.7 I >2.7-3.0 >3.0-3.3 >3.3-3.9 NITROGEN Fig. 16. Distribution of nitrogen in burned and unburned communities 1 year following treatment. mountain shrub o �318 1 50 BURN 1 60 40 .. 30 40 T 20 , T 1 a, 0 a:: 10 T u (!) Z \ T T 1 I 0 T 1 o 0 I/) s:: 50 !=i CONTROL u, -I 140 fT1 0 ~ ::u -< z « l- 0 ::u 40 ~ G'l r 30 120 <, s:: N T 80 T 20 1 _I_ 40 1 10 o 0 :>0.0-0.3 :>0.3-0.6 :>0.6-0.9 :>0.9-1.2 :>1.2-1.5 :>1.5-1.8 :>1.8-2.1 >2.1-2.4 :>2.4-2.7 1'/0 :>2.7-3.0 :>3.0-3.~ NITROGEN Fig. 17. Distribution of nitrogen in burned and unburned mountain communities 2 years following treatment. shrub �319 proportions of high quality herbage were greater in burned mountain shrub, the total amounts of herbage on offer were substantially greater on control plots in that community. During Year 2, distributions of IVDOM and nitrogen became more similar to controls than they had been the previous year because burns tended to contain a smaller fraction of herbage in the higher IVDOM and nitrogen categories, but more herbage biomass overall. Fire effects on nutrient distributions in grassland were similar to the influences we observed in mountain shrub. Durinq Year 1, more than half of the standing crop in burned grassland exceeded 50% IVDOM and 1:5% nitrogen? only 1/4 of the standing crop in unburned grassland contained IVDOM concentrations greater than 50%, only 1/5 of that biomass exceeded 1.5% nitrogen (Fig~ 18, 19). During Year 2, the distributions of IVDOM and nitrogen on burned grassland plots resembled distributions on controls more closely (Figs. 20,21). However, there remained a tendency for burns to contain a large portion of herbage in the higher categories. These differences in nutrient distribution, caused sharp divergence in supplies of food between burn and control plots. Assuming that 50% of the standing crop in any category can be eaten by deer or sheep and allowing a drymatter intake of 1.5 kg/day for an adult deer (Alldredge et a1. 1974) and 2.0 kg for adult sheep (Hebert 1973), we calculated nutritional carrying capacities on burn and control plots (Table 9). In general, fire reduced the capability of mountain shrub and grassland habitats to support animals on a low nutritional plane, but dramatically increased the carrying capacity of those habitats for animals obtaining high quality diets. If all animals were to consume the highest quality diets (>50% IVDOM, >1.5% N), burned grassland and mountain shrub could consistently support more deer than controls. At intermediate levels of nitrogen (>50% IVDOM, 71.5% N) carrying capacities of burns also exceeded controls except in grassland during Year 2. Controls could carry more deer and sheep only if their diets contained relatively low quality herbage (>40% IVDOM, >0.9% N). Thus it is plausible that burned plots, despite their lower total biomass, could support more animals in productive condition because of the high levels of diet quality necessary to meet the costs of production, lactation, and growth (reviewed by Robbins 1983:207-226). This demonstrates that although burning may diminish the absolute abundance of food for herbivores, it substantially increases food supplies of relatively high quality. Discussion Fire caused 2 changes in mountain shrub and grassland communities which accounted for much of the difference we observed in nutrient distributions. Fire increased the amount of herbaceous biomass relative to the amount of woody biomass in both plant communities. Because herbs tended to contain higher levels of nitrogen and IVDOM than woody growth, standing crops in burned areas were dominated by higher quality herbage; standing crops in unburned areas contained relatively more low quality plant tissue. Moreover, fire increased the nitrogen and IVDOM content of grass leaves, the dominant herbaceous component of both communities, and this change accentuated the difference between burns and controls in the amount of herbage containing the highest levels of IVDOM (>50%) and nitrogen (>2.2%). �Table 9. Densities (animal days/ha)a of mule deer or mountain sheep which could be supported levels of diet qual ity in burned and unburned grassland and mountain shrub communities. IVDOM > 50% Burn Control at different N 0 N bO% Burn Control > 2.1% Burn Control Year ~O% Burn Control 1980 1981 136 363 166 283 94 273 90 200 39 76 10 13 153 356 256 466 133 256 ' 123 67 60 33 43 20 1980 1981 253 363 296, 433 253 190 156 276 123 183 20 98 250 226 376 350 136 193 93 80 60 33 13 12 1980 1981 102 272 124 212 70 204 67 150 29 57 8 10 115 267 192 349 100 192 92 50 45 25 32 15 1980 1981 190 272 222 324 190 142 117 207 92 183 15 74 188 169 282 262 102 145 70 60 45 24 10 9 > > 0.9% Burn Control > 1.5% Burn Control > Mule deer Mountain shrub Grass land Mountain sheee Mountain shrub Grass land aDensity calculated as (1/2 herbage biomass which exceeds diet quality level) f (drymatter w intake 1b) �321 50 BURN T ! 40 T 1 40 30 30 ; i 20 20 a. 10 10 0 a::: z rn T 0 0 C/) -< s: 50 ~ J> CONTROL u, 0 0 0 :::0 0 z ~ CD J> (j) u (!) ::r: :::0 IT1 T -I -I 80 ",:::0 - 40 (j) ...•.. 60 30 I\) + 20 s: T T --r T ~ '_l 40 1 i 20 10 T .L 0 :>0-10 >10-20 :>20-30 r I ..I " T :>30-40 >40-50 % >50-60 >60-70 0 >70-80 >80-90 IVDOM Fig. 18. Distributions of in vitro digestible organic matter (IVDOM) in burned and unburned grassland communities 2 years following treatment �322 ( i 50 BURN 40 40 .. 30 T 30 1 T 1 ; ; 20 20 I ::u llJ o, 0 10 a: o (!) ~ 0 ~ fTI T f T I I T I 1 10 1 I T 1 . l> G'l fTI 0 0 0 ::u -< 3: z ~ 50 ~ CON TROL en 100 ~ ::u u, 0 ~ 40 80 0 G'l <, s::: 30 20 I\) 1 1 60 T . r T ~40 T J. 10 I I o ~ zo .. T i , >0.0-0.3 >0.3-0.6 >0.6-0.9 >0.9-1.2 >1.2-1.5 >1.5-1.8· >1.8-2.1 >2.1-2.4 >2.4-2.7 >2.7-3.0 >3.0-3.3 !"/o NITROGEN Fig. 19. Distribution of nitrogen in burned and unburned communities 1 year following treatment. grassland 10 �323 50 BURN L 50 40 40 30 30 20 20 :c IT! a.. 0 a:: :::0 OJ 10 T o (!) z 0 z 10 1> T --'O_ 0 J G> 1 IT! _T 0 :::0 -< ~ ~ CJj 50 1> -i CONTROL IJ... 100 ~ 0 tft. :::0 40 80 i 0 ~ 30 I\) 60 T 1 20 10 40 T ~ T T 1 1 0 G> <, .___ >0-10 T >10-20 >20-30 >30-40 20 .' >40-50 0 >50-60 >60-70. >70-80 >80-90 % IVDOM Fig. 20. Distributions of in vitro digestible organic matter (IVDOM) in burned and unburned grassland communities 2 years following treatment. �324 50 BURN T J 40 50 1 1 -. 40 30 30 : 20 \ Q.. 0 0::: o 10 (!) T Z 0 0 Z <t I- 50 C/) I T 1 r T I 1 I 20 :r: ITI ::u III 10 l> G) ITI 0 ::u -< s: CONTROL 100 ~ LL -I 0 ~ 0 0 ITI 40 80 G) <, 30 60 20 40 10 I T r T 1 1 0 ::u 1 :'~ 20 1 >0.0-0.3 >0.3-0.6 >0.6-0.9 >0.9-1.2 >1.2-1.5 >1.5'-1.8 >1.8~.2.1>2.1-2.4 .>2.4-2.7 >2.7-3.0 >3.0-3.3 % NITROGEN Fig. 21. Distribution of nitrogen in burned and unburned grassland communities 2 years following treatment. 0 s: I\) �325 During two growing seasons following treatment, fire reduced the total amount of aboveground herbage in mountain shrub and grassland communities and increased the amount of herbage containing high concentrations of nitrogen and IVDOM. These changes offer important implications for animal diet selection and for the carrying capacities of mountain shrub and grassland habitats. The shifts in nutrient distributions we observed should facilitate selection of high quality diets by individual animals. On burned plots. food items containing high concentrations of nitrogel1 and IVDOM were relatively unobstructed by less nutritious biomass. For example, in burned mountain shrub during Year 1, a foraging herbivore would have to search through 24 gm of plant tissue to find a single gram of herbage which exceeded 60% IVDOM; in unburned areas it would.have to find that gram amid 276 gm of less nutritious herbage. These differences appeared to influence the quality of winter diets of ungulates; mule deer and mountain sheep chose diets with substantially higher levels of nitrogen and IVDOM from burned plots compared to controls (see Experiment 1). The differences in nutrient distributions we observed are consistent with theory on plant succession and ecosystem development. During early stages of secondary succession following a disturbance like fire, the rate of primary production within communities exceeds the rate of respiration (Odum 1969). As the community matures, production and respiration approach equality. It follows that early succession communities are dominated by productive plant material and later seres contain more biomass devoted to maintenance. Growing plant tissue contains higher concentrations of nitrogen and digestible organic matter than tissue which is not growing. Thus, changes in community production and respiration should be reflected in the distribution of nutrients within those communities. Recently disturbed communities, like the ones we observed, should be dominated by productive, and therefore nutritious plant tissue. We see that trend by observing that burns, compared with controls, contained a larger fraction of nutritious herbage during both years. Moreover, that fraction of herbage declined as burned communities matured; during Year 2 proportionally less of the total biomass on burned plots contained the high concentrations of nitrogen and IVDOM that we observed the previous year. We conclude that fire reduces a relative shortage of food for herbivores. This reduction results from shifts in the way nutrients are distributed within herbage standing crops; fire increases the fraction of aboveground herbage which contains nutrients at high concentration. Experiment IV: Fire Effects on Ecological Separation Between Mule Deer and Mountain Sheep Literature Review and Rationale Differences between sympatric species in their food habits and habitat choices reduce competition for shared resources (reviewed by Schoener 1982). These mechanisms of ecological separation may depend on the successional �326 state of the plant communities those species share. In climax mountain shrub and grassland communities, mule deer and mountain sheep are separated ecologically by differences in their diets (Hobbs et al. 1983). Moreover, mountain sheep appear to be best adapted to use steep, open, rocky hillsides (Geist 1971); mule deer prefer more gentle terrain with more extensive shrub and tree cover (Wallmo 1981). However, both species are attracted to plant communities which emerge following fire (Lay 1957, McCulloch 1969, Roppe and Hein 1978, Riggs et al. 1980). Consequently, fire may reduce the extent of ecological separation along the habitat dimension of niches of mule deer and mountain sheep. If fire also increases diet overlap, then competition between these species will be likely. Here, we test the hypothesis that fire increases diet overlap between mule deer and mountain sheep in mountain shrub and grassland communities. Methods We estimated botanical composition of diets of mule deer and mountain sheep grazing on burned and unburned plots according to procedures described in Experiment 1. The contribution of each principal forage to the diet was calculated as: Wi bi N_ L Wi bl i=i where: Wi mean weight of 25 hand-plucked bi = number of bites of principal N bites of principal forage i, forage i, and = total number of principal forages (those diet items contributinq >2% of observed bites on any burn or control plot). Percentages of principal forages in mountain sheep and mule deer diets in burned and unburned ~lant communities are given in Appendix A. The degree of overlap in diets of mule deer and mountain sheep was calculated using the proportional similarity index of Schoener (1968): .. N Ro = 1 - 1/2 z a=l j >, I a - P. I· Ja where Pia and Pja are the proportions of the total diet of animal species i and animal species j taken from food category a. The overlap coefficient R. varies from 0, when diets are completely distinct, to 1 when diets are identical. Differences in diet overlap indices were analyzed with a 4 x 2 x 2 factorial analysis of variance for a randomized complete block design with repeated measures. We replicated by plots and repeated over months. Blocks, months, and years were considered random effects. Significant differences among individual months by treatment means were examined using Tukey's w-procedure, �327 also called the honestly significant difference simultaneous confidence level of P < 0.10. (hsd) procedure, at a Results With few exceptions, we found that prescribed burning increased diet overlap between mule deer and mountain sheep in both plant communities (mountain shrub treatment P = 0.051, grassland tre~tm~nt P = 0.105, Fig. 22). Averaged over months, the-size of treatment effects remained constant for 2 years after burning (mountain shrub treatment x year P = 0.771 grassland treatment x year P = 0.696). However, within years we observed substantial monthly variation in fire effects on diet overlap. In general treatment elevated diet overlap during winter months; however, effects of treatment were inconsistent during May. There were exceptions to this trend; during March of Year 1, diet overlap was higher on controls than burns. We attribute this reversal of treatment effects to the influence of snowcover during that month. Although sheep pawed through the snow to eat grasses, snowcover appeared to prevent deer from consuming grass on burns. Shifts in the percentage of grass and browse in diets of mule deer and mountain sheep accounted for much of the increase in diet overlap on burned plots. Averaged across years and months, deer consumed more grass on burned mountain shrub (treatment P = 0.0002, Table 10) and burned grassland plots (treatment P = 0.048, Table-II) than on controls. Sheep also ate more grass on burned plots in mountain shrub than they did on unburned plots (treatment P = 0.066, Table 10). Their diets were consistently high in grass content on both burn and control grassland plots (Table 11). a Table 10. Mean percentage grass in deer and mountain sheep winter and spring diets during 1980-82 in burned and control mountain shrub habitat in Colorado. Year and month Burn Mule deer Control Mountain sheep Burn Control 1980-81 Nov Jan Mar May 63 86 82 82 20 11 -7 64 90 93 99 91 65 47 41 75 44 36 87 96 10 21 15 87 67 79 84 94 54 76 66 83 1981-82 Nov Jan Mar May aN 4. �328 a Table 11. Mean percentage grass in mule deer and mountain sheep winter and spring diets during 1980-82 in burned and control grassland habitat in Colorado. Year and month Burn Mule deer Control Mountain sheep Burn Control 1980-81 Nov Jan ~r May 48 77 70 71 25 37 23 66 69 81 57 91 61 52 67 87 1981-82 Nov Jan Mar May 25 30 81 81 12 41 69 85 73 65 68 87 53 83 84 96 aN = 4. The increased contribution of grass to ungulate diets on burned plots was relatively greater for mule deer than mountain sheep. As a consequence of these shifts, diets of deer and sheep contained similar portions of grass of the same species and plant par~on burns. On controls, sheep tended to eat more grass and less browse than deer. We observed similar results when we analyzed burning effects ~n the percentage of green grass in ungulate diets. Mule deer (treatment P < 0.006, Tables 12, Ii and mountain sheep (treatment P < 0.059, Tables-l~, 13) consumed more green grass on burns than controls Tn both plant communities. Increases of green grass in deer diets were more d~amatic than in sheep diets. Burning resulted in a decrease in the percentage of browse in ungulate diets. Averaged across years and months, deer consumed less browse on burned mountain shrub (treatment P = 0.027, Table 14) and grassland (p = 0.008, Table 15) than on controls-and sheep consumed less browse on burned than unburned mountain shrub plots (treatment P = O:OO~ Table 14). Again, burning effects on diet botanical composition-were more dramatic for deer than sheep, especially during the winter months. This relatively greater decrease in percentage of browse in deer diets than in sheep diets contributed to greater diet overlap on burns. We observed no effect of burning on forb content of ungulate diets in either plant community ( ~ > 0.05). The percentage of green grass in mule deer diets was directly correlated with diet overlap between deer and mountain sheep (Fig. 23); the percentage of browse in deer was inversely correlated with diet overlap (Fig. 24). Thus, the increase in diet overlap we observed resulted largely from deer switching from browse dominated diets on controls (which were dissimilar to sheep diets) to diets with more green grass on burns (which were similar to sheep diets). �329 a Table 12. Mean percentage green grass in mule deer and mountain sheep winter and spring diets during 1980-82 in burned and control ~ountain shrub habitat in Colorado. Year and month Burn Mule deer Control Mountain 1!.Burn sheep Control 1980-81 Nov Jan Mar May 52 84 82 82 20 11 7 54 23 54 26 91 14 19 6 75 44 16 87 96 10 0 14 87 44 12 63 94 31 4 26 83 1981-82 Nov Jan Mar May Table 13. Meana percentage green grass in mule deer and mountain sheep winter and spring diets during 1980-82 in burned and control grassland habitat in Colorado. Year and month Burn Mule deer Control Mountain Burn sheep Control 1980-81 Nov Jan Mar May 36 77 63 71 23 37 6 66 5 68 8 91 16 29 2 87 25 5 81 81 12 52 19 49 87 23 1981-82 Nov Jan Mar May aN 0 69 85 0 26 96 4. a Table 14. Mean percentage browse in mule deer and mountain sheep winter and spring diets during 1980-82 in burned and control mountain shrub habitat in Colorado. Year and month Burn Mule deer Control Mountain Burn sheep Control 1980-81 Nov Jan Mar May 26 6 16 7 53 53 72 3 0 0 0 2 9 7 22 0 �330 Table 14. (continued) 1981-82 Nov Jan Mar May 11 26 2 0 0 0 0 0 56 46 22 0 0 3 0 0 aN = 4. a Table 15. Mean percentage browse in mule deer and mountain sheep winter and spring diets during 1980-82 in burned and control grassland habitat in Colorado. Year and month Burn Mule deer Control Mountain Burn sheep Control 1980-81 Nov Jan Mar May 22 1 24 6 42 41 63 4 3 0 10 1 10 2 14 0 1981-82 Nov Jan Mar May 26 16 2 0 44 39 21 0 0 0 0 0 4 0 3 0 aN 4. Table 16. Meana percentage forbs in mule deer and mountain spring diets during 1980-82 in burned and control grassland Colorado. Year and month Burn Mule deer Control sheep winter and habitat in Mountain Burn sheep Control 1980-81 Nov Jan Mar May 29 21 6 23 32 23 14 30 27 19 34 8 29 47 19 13 1981-82 Nov Jan Mar May 49 54 16 19 44 25 10 15 27 35 24 13 43 16 14 4 aN .~ 4 . �331 a Table 17. Mean. percentage spring diets during 198d 82 Colorado. forbs in mule deer and mountain sheep winter and in burned and control mountain shrub habitat in p Year and month Burn Mule deer Control Burn sheep Control 27 36 21 35 10 7 1 6 26 46 37 17 34 33 63 13 33 20 16 6 46 21 34 17 Mountain 1980-81 12 8 2 Nov Jan Mar May 11 1981-82 45 37 Nov Jan Mar May 11 4 aN 4. Discussion Increased diet overlap does not necessarily indicate increased competition between sympatric species. However, when the similarity of diets is great and the quantity of resources shared by those species is small, then the likelihood of competitive interactions becomes substantial. This is the situation we observed following prescribed burning. Diets of mule deer and mountain sheep were much more similar in burned than unburned mountain shrub and grassland communities. The absolute amount of food for these herbivores decreased following fire (see Experiment III). Consequently, we infer that one possible detrimental effect of burning is an increased potential for competition between mule deer and mountain sheep. Experiment Literature V: Fire Effects on Nitrogen Cycling Review and Rationale The influence of fire on the nitrogen economy of ecosystems has been widely studied. Fire acts as a potent mineralizing agent, causing the rapid transformation of organic nitrogen to inorganic forms (Christensen 1973, St. John and Rundel 1976, Sharrow and Wright 1977, Dunn et al. 1979, Jurgensen et al. 1981). Fire results in alterations of the abiotic environment, which, in turn, lead to changes in biotic processes (reviewed by Ahlgren and Ahlgren 1965, Viro 1974, Wells et al. 1979, Raison 1979, Woodmansee and Wallach 1981). Elevated soil temperatures that result when the plant canopy is removed and the release of cations in ash improve conditions for microbial growth in the soil environment (Jorgensen and Hodges 1970, Renbuss et al. 1973, Christensen and Muller 1975, Tiwari and Bharat 1977, Raison and McGarity 1980). It follows that nitrogen transformations mediated by microbial populations should be similarly altered by burning. �332 GRASSLAND GRASSLAND 1980 - 81 1981 - 82 .8 C .7 B- B f\ .6 w \ z Q. \ .3 c---c \ C-C 0 I- \ B ...J ...... B / B ...... \ / -c a: w > / \ I I .4 / / \ I 0 / c \ I .5 / \ / / x / / \ ~./ ..2 \ w , C, 'B 0 .1 '" '" NOV MOUNTAIN MAR JAN SHRUB '" MAY NOV '" MAR JAN MOUNTAIN 1980 - 81 SHRUB MAY 1981 - 82 c .8 B .7 I " / I "- / x o ~ \ I \ \ Q. -c ...J a: w > 'B I B w " I / -> o IW c o * NOV JAN MAR MAY .* '" NOV ._JAN '" MAR '" MAY Fig. 22. Diet overlap between mule deer and mountain sheep in burned and unburned mountain shrub and grassland communities. Means (n = 3) for burned plots are given by B, for control by C. Pairs of means with ,':below them are different at P = 0.10. Data for 1 year following treatment are shown on left; 2 years after treatment are on the right. �DIET OVERLAP-REGRESSION I 1.0 I .8 x ~ x I '" x XxX X X X .6 Xx x X X X XX x x X ~ X X )f( X X X X ~ X x Cl X ' ___....--x X ___....-- x X x .4 x ~ '% X x 2fx .... x "X • 0.0 x xx )0( X L_'_.. ~ x X x ._ X_ ~ N x x x x x x x X X ___0_ __ ~ ~ .. __ X X._ ~ ~ ~,,_J ~ ~ ....•. AVE%GGRSS Fig. 23. Relationship between diet overlap green grass in diets of mule deer. index (DINDX) for mule deer and mountain sheep and the % w w w �" \.N \.N J:- DIET OVERLAP-REGRESSION 1.0 .9 ... x f. x .0 .7 .6 ~ z H •u x· " fA~ x ." " 'P.' _. .5~ x X x t.:a .4 lV.lI\ .3 he x x ~ x x 0.0 x x XX XX x .2 ~ .1 J. x x " * x x 'Sc x x x x I x x x x x x L m x Q ...• ro cu t.J ('l) Q- v m ttl L__ t.'i) ~ nVE%BROl~SE Fig. 24. Relationship between diet overlap browse in diets of mule deer. L (;J "" x I co CX) index DINDX) for mule deer and mountain m en m...• sheep and the % of �335 Despite the scrutiny that ecosystem effects of firehave received, few studies have examined the influence of fire on biotic transformations of nitrogen. Dunn et al. (1979) reported that nitrogen mineralization was stimulated following fire, but Schimel (1982) found that repeated burning reduced potentially mineral izable nitrogen. Although Jorgensen and Wells (1971), Youngberg and Wollum (1976) and Grove et al. (1980) observed that fire increased rates of nitrogen fixation in forest ecosystems, no work to date has examined these effects in mountain grassland or shrub communities. Prescribed burning is widely used in these communities to improve range conditions for wild and domestic animals; however the ecosystem' effects of such application are poorly understood. Here, we report experiments on the influence of fire on nitrogen mineralization, nitrogenase activity, and soil inorganic nitrogen in mountain shrub and grassland communities durinq 2 years following burning. Methods We estimated soil concentrations of N0 - + NO - and NH4+ and the rates of 2 nitrogen mineralization and fixation on burned and unburned plots during 2 growing seasons following treatment. Mineralization was estimated by the buried bag technique of Eno (1969). On 4 June 1980 and 1981, 20 soi 1 cores were taken to a depth of 5 cm at random locations on each plot. Each core was split longitudinally and sieved through a 2-mm mesh screen to remove large roots and rocks. Half of each core was placed in a polyethylene bag and replaced in the ground. The remaining portion was immediately taken to the laboratory for analysis. Within 10 hours of taking samples, we extracted a 3-g subsample of each core with 2 M KCl with 5 ppm phenyl-mercuric acetate added to inhibit microbial activity. Moisture content of a separate 5-g aliquot was determined according to A.O.A.C. (1965). Ammonium content was determined by a_colori~etric ammonium reaction with salicylate nitroprusside at pH 13; NO~ + N02 was determined by a cadmium reduction using a continuously coppefized reduction column. Six weeks later we retrieved the buried polybags and treated those samples the same as described for the initial samples. Net min~raliz~tion _ was estimated as the increment in total inorganic nitrogen (NH4 ' N03 ' N02 ) between 2 sample dates. Potential heterot~ophic nitrogenase activity was estimated with the acetylene reduction procedure of Hardy et al. (1973), as described by Hersman and Klein (1979). Thirty 5-cm soil cores were taken from random locations on each plot on 10 June 1980, and 12 June 1981. For each plot, cores were composited into 3 samples of 10 cores each, placed in plastic bags and immediately cooled at approximately 15 C in a portable ice chest. Upon returning it to the laboratory, each composite was sieved through a 2-mm screen and thoroughly mixed. Twenty subsamples were selected from different locations within each composite and were mixed to form a 10-g sample for analysis. Samples were stored in plastic bags at 10 C. Prior to analysis, samples were wetted to field capacity. Responses were analyzed with an analysis of variance for a split plot design with plant community type forming the whole plot, and burn vs. control contributinq the split plot. Differences among individual means were established �336 with Tukey's Q simultaneous comparisons were considered random effects. at p;:: 0.10. Years and replicates Results We observed that fire resulted in significant (p < 0.10) increases in NH4+ and N02- + N03" contents of grassland and mountain shrub soils 10 months after the burn (Figs. 24, 25). This effect, however, disappeared in the second year (year x treatment interaction P ;::0.05); soil mineral N values were significantly higher on burns during Year 1, bur returned to control levels during Year 2. We attribute this return to uptake of N by the recovering plant community; above-ground plant biomass on burned plots was almost 3 times greater durin~ Year 2 sompared with Year 1 (N. T. Hobbs, unpubl. data). Reduced levels of NH4 and N03 in June of the second year may also have been due to increased microbial uptake. Surface soils were probably temporarily sterilized. The winter following the burn was exceptionally severe; microbial populations had little opportunity to recover prior to the first sample. Effects of fire on soil mineral N also depended on plant community (community x treatment interaction f;:: 0.12). The larger increases in NH4+ in mountain shrub relative to grassland likely resulted from the more even nature of the burn. The patchy distribution of burned areas within grasslands resulted in some treatment plots being similar to controls while in the mountain shrub practically all the area was burned. Net mineralization of soil N was significantly greater (p ;::0.10) on burned grassland plots during Year 1, and on burned mountain shrub plots during both years (Fig. 26). Dunn et al. (1979) and Sharrow and Wright (1977) also found that rates of nitrogen mineralization were increased by burning. However, although a single fire may stimulate mineralization, the cumulative result of several fires may inhibit it. Schimel (1982) and Biederbeck et a1. (1980) suggested protracted annual fi re regimes led to decreased mineralization as a result of reductions in pools of readily decomposable C and N and consequent decrements in microbial biomass. The magnitude of fire effects on net mineralization depended on year (year x treatment interaction.P ;::0.11) and vegetation type (community x treatment interaction P = 0.005)~ treatment effects were greater in mountain shrub than grassland and declined in magnitude during Year 2. Main effects and interactions can be explained by changes in the soil environment. Soils in burned grasslands are typically 3-16 C warmer than similar unburned sites (Weaver and Rowland 1952, Kucera and Ehrenreich 1962, Scotter 1963, Old 1969, Lloyd 1972, Peet et al. 1975). This increase results primarily from absorption of solar radiation on blackened soils unshaded by litter and plants (Woodmansee and Wallach 1981). Nitrogen mineralization rate increases with increasing temperature (Stanford et al. 1973, Campbell et al. 1981). The effect of increased soil temperatures on N mineralization may have resulted in the elevated mineralization rates we observed. Immobilization of N into remaining litter may also have been a factor. Both of these explanations are consistent with the temporal and spatial interactions in treatment effects. Plant biomass on the burned plots of grassland was equal to the control by June of the second year; the difference in soil temperatures �25 20 ~ BURN D CONTROL '"' 0' <, 0' _...::J.. 15 'N o z + c 10 11'0 o a Z 5 o I V/((4 1980 I (////4 1981 GRASSLAND I [/((0 1980 I ve((a I 1981 MOUNTAIN SHRUB Fig. 24. Soil concentrations of N03- + N02- in mountain shrub and grassland communities during the growing season, one and two years after prescribed burning. Different letters indicate differences in means at p = 0.10. w W -.....J �25 . w w 00 ·e 20-1 r///)/f I ~ ~ BURN D CONTROL ~ 15 Ol - :t. +v J: Z 10-1 I 5 o .(] . ~ I V///A I I rm/J I 1980 b u..-- V///IJ 1981 GRASSLAND b ~ I V///A 1980. I V///I] I 1981 MOUNTAIN· SHRUB Fig. 25. Soil concentration~ of NH4~ in mountain shrub and grassland communities ,during the growing season, one and two years after prescribed burning. Different letters indicate differences in means at P ~ 0.10. �~ -z 25 ~ 20 0' ::l a o - a CD ::> ~ BURN D CONTROL U z - ~ W 15 W b ?: <D z 10 o W c N _J <{ 5 0:: W Z ::IE o z I V///Il 1980 I W//4 I 1981 GRASSLAND Fig. 26. Net mineralization rate in mountain one and two years after treatment. Different V(((O 1980 I r////4 I 1981 MOUNTAIN SHRUB shrub and grassland communities during the growing season, letters indicate differences in mean values at P = 0.10. w W 1..0 �340 and the consequent divergence in N mineralization, disappeared by Year 2 in that community. In contrast, plant biomass in the mountain shrub community had not recovered to control levels by Year 2, and large areas of bare soil remained. As a result, we surmise that e)evated soil temperatures would have persisted into Year 2. A similar pattern existed with respect to the immobilization potential of remaining litter. The mountain shrub community was burned intensely enough to remove all litter, while in the grassland some litter remained. Thus, the prolongation of increased N mineralization in the mountain shrub community is attributable to the greater intensity of the burn which occurred there, the slower recovery of the plant canopy, and the complete removal of the litter layer. The effect of fire on N mineralization in grassland was more brief because the fire was more heterogeneous and less intense. The recovery of plant canopy was more rapid, and some litter remained. Potential nitrogenase activity was lower on burned plots than controls during Year 1; however, this effect was significant only in mountain shrub (p = 0.10) (Fig.27). Magnitude of treatment effects depended on year (year x treatment interaction P = 0.07) but not on plant community (community x treatment interacti08 P = 0.74). The reduction in nitrogenase activity may have resulted from sterilizing effects of fire. However, the inhibition of nitrogenase activ---i~yon burned plots during Year 1 and the return to control levels the fpllowing year appeare~ to b~ relat~d to the concentration of mineral N in soils. Mineral N (NH4 ,N03 + N02 ) was increased in soils of burned plots during Year 1, but not in Year 2. The reduction in potential nitrogen fixation was greatest in the mountain shrub community and the increase in soil inorganic N was also greatest there. Soil mineral N levels accounted for 52% of the variation in potential nitrogen fixation (Fig. 28); free-living and symbiotic nitrogen fixers decrease fixation when mineral N is elevated (Oghoghorie and Pate 1971, Manhart and Wong 1979, Wong 1980). Our observation of a curvilinear relationship between nitrogenase activity and soil mineral N suggests that fixation responds in a threshold manner with respect to mineral N and that N fixation is reduced drastically at soil mineral N concentrations >10-20 ~g/g dry soil. These findings are at odds with reports that fire increases rates of N fixation (Jorqensen and Wells 1971, Youngberg and Wollum 1976, Grove et al. 1980). However, we cannot exclude the possibility that N fixation may become significant after 2 or more years, or that inorganic N would not have been elevated had the burn been followed by a more normal winter. A better understanding of the controls on N-fixing organisms would be required to better predict post-fire responses. Discussion Fire caused increased rates of N mineralization for 1 year in a mountain grassland, and for 2 years in a mountain shrub community. The rapid recovery of the vegetation in the grassland caused rapid convergence of burned to unburned conditions. The shrub community burned more evenly and more severely and required longer to return to control conditions. We attribute most of the increase in N mineralization to increased soil temperatures. Nitrogenase activity was depressed by fire 1 year after the burn in the mountain shrub community. This may have resulted from partial sterilization �25 • ,_ I ~ 01 1 °1 2 ~ BURN D CONTROL ._>- > I0 « w en 15 10 « z w o 0 a:: 5J a a "h I I a ~ i I a ~ab I- Z o I '(((//], 1980 '«(Cd 1981 GRASSLAND I V««I' 1980 MOUNTAIN '({Cf?! , 1981 SHRUB Fig. 27. Nitrogenase activity (acetylene reduction) in mountain shrub and grassland communities during the growing season, one and two years after treatment. Different letters indicate differences in means at p = 0.10. \.N ,.!:- �'_ Fig. 28. Relationship between potential nitrogenase activity (acetylene mountain shrub (open circles) and grassland communities (closed circles) tion is given by y = e2.66 - 0.53x. P = .00005, R2 = .52. reduction) and soil mineral during the growing season. N in Equa- �343 of the soil. In addition, soil inorganic N levels were elevated in burn relative to control plots and this probably contributed to the depression nitrogenase activity. in The effect of fire was to increase the rate of N mineralization, and presumably, the availability of N to vegetation. Nitrogen losses undoubtedly occurred from the vegetation and these may not have been rapidly compensated for by N-fixation. Consequently, we recommend that prescribed burns for range improvement in montane communities not be repeated frequently on the same sites until fire effects on N budgets in these communities are better understood. RECOMMENDATIONS We conclude that prescribed burning can improve grassland and mountain shrub habitats for mule deer and mountain sheep by increasing the capability of those habitats to carry larger populations of animals at a high plane of nutrition. The potential for using fire to benefit deer and sheep can be maximized by specific burning practices. Based on our studies, and our synthesis of the results of other workers, we recommend that the following points be considered in planning prescribed burns: Objectives of Burn Treatments Based on this study and others (Riggs et al. 1980, Risenhoover and Bailey 1980, Risenhoover 1981), it appears that burning can be used to achieve 3 primary objectives for habitat improvement. First, burning can improve the nutritional status of individual animals. If such improvement translates into enhanced reproductive performance of females, then burning will result in increased productivity of populations. Second, burning can be used to improve the structure of habitats, particularly for mountain sheep. Converting forest and brushland to open grasslands increases habitat visibil ity, which is believed to be an important feature of quality mountain sheep range. Finally, because burned areas attract deer and sheep, burning may be useful in increasing dispersal into formerly unoccupied range. Deciding which of these objectives, nutritional improvement, structural improvement, or . increased dispersal, is the primary management goal will influence when, whe re, and how to bu rn. . Site Loca t ion Appropriate location of sites to be burned will depend strongly on the objectives for habitat improvement. If nutritional enhancement is the primary consideration, then we recommend burning south-facing slopes on winter range in areas without persistent snowcover and which are traditionally used by the target population. Such practices will facilitate production of green grass, which is largely responsible for the nutritional benefits we observed, and will insure that animals "find" the burned area. To achieve structural enhancement of habitat for mountain sheep we suggest burning visibility obstructing vegetation in areas adjacent to openings currently used by sheep. Indentification of seasonal ranges and migration routes will greatly facilitate �344 choosing the optimum sites for these burns. Si.m!1arly? we recommend burning sites near areas of traditional sheep concentration to enhance dispersal and reduce density of sedentary populations. It may be necessary to locate burns to minimize competition among sympatric ungulates. For example, if burns are being planned to benefit mountain iheep, competition with deer or elk could reduce those benefits. Thus, where other ungulates share ranges with sheep, we recommend burning in steep, rocky terrafri of the type lfk~ly to be used exclusively by mountain sheep. In any case, burns should always be located in proximity to escape cover appropriate for the target species. Size of Burns Total area of burns is critical to successful habitat improvement; they must be large enough to offer meaningful benefits to a significant portion of the target population. Moreover, because the biomass of forage is sharply reduced in mountain shrub and grassland communities following fire and because of the attractiveness of burned areas to ungulates, concentration of animals and damage to the emerging plant community from overgrazing is probabl~ whenever burned areas ~re small relative to unburned habitat. We are uncertain what size burned areas should be to maximize benefits to mule deer and mountain sheep and prevent these deleterious effects. However, as a first approximation, we suggest that burns encompass at least 10% of the currently used range. For mountain sheep, this 10% should be distributed over relatively large patches, no less than 15 ha, to create the open habitat they appear to favor. For mule deer, smaller patches are desirable (1-3 ha) to allow for deer preference for shrub and timber cover adjacent to open areas. Timinq of Burns and Fire PrescriPtions Our overriding recommendation for planning burns for habitat improvement is to consult with fire ecology experts on the specifics of all burns. They will .be invaluable in translating management objectives for animals into appropriate prescriptions for fires. Because of the tremendous variation among sites, our suggestions cannot provide the detai 1 necessary to develop site-specific fire prescriptions. However, we can offer some general principles influencing choice of season of burn ing. Spring burns tend to be cool; this characteristic has advantages and disadvantages. Because of the wet conditions typical of spring in Colorado, it may be very difficult to carry extensive fires during that season. Despite this drawback, spring burns may prevent damage to fine-leaved perennial grasses which can result when fires are hot. Spring burns allow more rapid recovery of shrubs, which mayor may not be beneficial depending on the species the burn is intended to benefit. In general, early spring burns are detrimental to cool season species (which are often important winter forages) and enhance production of warm season ones. The primary advantage of spring burning on winter range is that the plant community has a full growing season to recover prior to winter. �345 Frequency of Burning There is insufficient information to recommend a burning schedule which will maximize benefits to deer and sheep without incurring long-term environmental degredation. However, our findings on fire effects on N fixation caution against overly frequent burning on the same site. It is possible that burning too often could deplete the nitrogen supplies of burned areas. Consequently, we recommend that burning schedules be conservative and that burning new areas be emphasized in preference to reburning previously treated areas. Future Research and Monitoring This study provides evidence that prescribed burning can improve the nutritional status of mountain sheep and mule deer. It remains uncertain, however, whether these improvements will lead to tangible changes in the performance of their populations. The next question which needs answer is "Does prescribed burning of ungulate habitat cause measurable increases in the productivity of ungulate populations?" That is, we need to know if the ultimate objectives of habitat improvement, more animals, is realized by our management practice. Thus, we recommend that monitoring be directed at evaluating the success of burning with particular emphasis on assessing effects of burning on recruitment. For bighorn sheep, it will also be important t6 know if burning can be used to increase dispersal from high density, sedentary populations. Inappropriate concentration of sheep on small, preferred patches of habitat can be related to many of the problems they are believed to confront, inc~uding inadequate nutrition, susceptibility to parasitism and, disease, and endocrine abnormalities related to chronic stress. Thus, mo~itoring fire effects on mountain sheep should determine whether burning habitat adjacent to traditional ranges can reduce the density of those populations by increasing dispersal. �346 LITERATURE CITED A. O. A. C. 1965. Official methods of analysis, Agric. Chern., Washington, D.C. 957pp. 10th ed. Assoc. Official Ahlgren, I. F., and C. E. Ahlgren. 1965. 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Spowart �356 APPEND,X A a in diets of 4 Mean percentage (g/100 g drymatter intake) of major forages mountain sheep grazing in burned and unburned mountain shrub communities in Colorado during November, January, March and ~ay 1980-81 and 1981-82. B, -. Jan Cd ,: B C 43 8 Nov Year Forage .t::I GRAM INOIDS Agropyron spp. (dead leaves) 1 2 77 11 Agropyron spp. (green tillers) 1 2 2 Agropyron spp. (inflorescences) 1 2 Bromus tecto rum (green leaves) 1 2 Bromus tectorum (inflorescences) 1 2 Carex spp. 1 2 Festuca ovina 1 2 Hesperoch1oa king ii 1 2 Poa spp. -rcfead leaves) 1 2 Poa spp. \green leaves) 1 2 Poa spp. -rrnf1orescences) 1 2 Sitanion hystrix Dead leaves) 1 2 Sitanion hystrix (green tillers) 1 2 30 March B 9 47 6 1 3 63 C 24 11 May B C 76 23 67 15 5 5 10 5 4 1 2 2 15 5 5 6 8 48 11 18 12 15 14 19 4 46 3 25 5 10 35 10 14 3 12 61 7 33 9 19 6 37 4 4 25 11 42 11 22 10 39 2 2 Total Green Graminoids 1 2 14 45 10 40 56 10 19 3 21 68 7 33 15 24 18 42 Total Graminoids 1 2 86 82 64 71 94 70 41 81 97 83 45 63 95 94 88 82 �357 APPENDIX A (cent inued) I',' Nov Forage Year Narch Jan .d "d B C B C B 2 5 11 May C B C FORBS Antennaria Arenaria Aster parvi fo1 ia fend1eri 1aevis Aster porter i 1 2 5 1 3 15 1 2 1 2 2 1 2 Cerastium arvense 1 2 Chrysopsis spp. 1 2 Cirsium 12 2 4 2 3 41 23 32 2 spp. 2 Co11insia parvif10ra Erigeron spp. Ga1ium borea1e Lupinus greeni 1 2 1 2 2 2 1 1 2 1 2 1anceo1ata Mertensia 1 3 1 2 coronop ifo 1ia Oenothera 1 2 Potenti11a spp. 5 7 1 2 Sisymbrium a1tissimum 1 2 Taraxacum Total officina1e Forbs 1 2 1 2 5 17 26 8 3 12 43 12 1 12 34 15 3 8 2 �358 APPENDIX A (continued) Nov Forage b Year BROWSE Arctostaphylos c .d B C B May March Jan d C B C B C 1 uva-ursi 2 Artemisia frigida 1 2 Artemisia tridentata (inflorescences) 1 2 Erigonum umbellatum (1eaves) 1 2 Physocarpus (leaves) monogynus 1 2 Physocarpus (stems) monogynus 1 2 Pinus ponderosa 1 2 Populus tremuloides 1 2 Purshia tridentata 1 2 Rosa acicularis 1 2 1 2 Total Browse 5 2 8 21 3 3 4 3 1 22 5 3 17 2 4 1 7 9 8 4 2 1 11 2 12 2 19 1 21 1 5 4 22 5 3 17 aMajor forages include those which contributed 2% or more of total bites eaten by mountain sheep on any replicate plot. bForages were differentiated green and dead material CYear d 1 = 1980-81 Yea r 2 - 1981-82 , B = Burn C = by plant species and plant part and between Control �359 APPENDIX B Mean percentage (g/100 g drymatter intake) of major forages in diets of 4 mountain sheep grazing in burned and unbu~~ed grassland communities in Colorado during November, January, March; and May 1980-81 and 1981-82.a Nov Forage cd· Year B b May ~1arch Jan d' B C B C B 48 8 50 3 40 41 80 7 23 54 79 18 30 29 12 52 11 60 11 60 91 89 C C I GRAMINOIDS Agropyron spp. (dead leaves) 2 Agropyron spp. (green ti llers) 1 2 Agropyron spp. (inflorescences) 2 Bromus ciliatus (green leaves) 1 2 Festuca idahoensis 1 2 Hesperoch1oa kingii 1 66 6 2 3 2 1 1 1 2 Juncus spp. 1 2 Poa spp. Wead leaves) 2 Poa spp. Tgreen leaves) 1 2 Stipa comata 1 2 Total Green Graminoids Total Graminoids 13 10 1 4 54 1 4 2 54 1 71 2 60 23 28 65 20 38 3 4 15 23 68 22 38 64 41 76 61 56 87 15 23 4 2 52 22 6 2 52 22 13 52 57 55 55 73 91 82 �360 �361 B (cent inued) APPENDIX Nov Forage b Year c B d C B May March Jan d B C B C C BROWSE Artemisia tridentata (inflorescences) 1 2 Erigonum 1 2 1 8 Populus tremuloides (1eaves 1 2 3 Potentilla 1 2 umbel1atum fruiticosa Ribes cereum 1 2 Rosa acicularis 1 2 Rubus deliciosus (stems) 1 2 Total Browse 1 2 5 6 2 5 9 5 1 15 6 1 10 3 2 2 8 3 11 14 6 2 5 3 3 3 2 13 15 12 9 1 10 aM ajar . forages include those which contributed 2% or more of total bites eaten by mountain sheep on any repl icate plot. b c d Forages were differentiated green and dead material. Year 1 = 1980-81 B = Burn by plant species and plant part and between Year 2 - 1981-82 C = Control �362 APPENDIX C Mean percentage (g/IOO g drymatter intake) of major forages in diets of 4 mule deer grazing in burned and unburned mountain shrub communities in Colorado during November, January, March, and May 1980-81 and 1981-82.a Nov Forage b Year GRAMINOIDS Agropyron spp. c B d I 2 5 Agropyron spp. (green t i ller s) I 2 II Bromus ci Iiatus I 2 Bromus tectorum (green leaves) I 2 Bromus tecto rum (inflorescences) 2 Carex spp. Jan Ce C B May C 4 17 B C 43 9 34 7 25 33 6 5 21 7 24 II I 3 18 4 3 2 12 I I 2 Hesperochloa B March kingii 4 I 5 2 Ko le r ia cr istata 3 3 2 I I 2 Poa spp. ldead leaves} 1 2 Poa spp. \green leaves) 1 2 Sitanion hystrix (green tillers) 1 2 Total Green Graminoids Total Gramoinoids FORBS Agoser is glauca Antennaria Arenaria parvifolta fendleri 41 28 20 6 16 15 70 10 9 1 2 52 45 20 9 88 14 12 1 2 62 45 20 9 88 38 12 21 1 2 70 10 2 1 5 10 21 21 6 62 37 85 16 11 5 37 34 85 16 I1 5 85 64 2 2 1 2 2 29 1 2 2 2 5 7 5 �C (continued) APPENDIX Nov c B,r:i Year b Forage FORBS (cont inued) Aster laevis Aster porteri Cerastium Chenopodium Chrysopsis Cirsium freemonti spp. spp. 1 2 Equisetum 1 2 Ga 1 ium bo rea 1e 1 2 Oenothera 1 2 Penstemon vi rens 1 2 Potenti11a spp. 1 2 Senecio spp. 1 2 integerrimus 1 2 Sisymbrium Taraxacum Tragopogon altissimum officinale dubius 2 7 6 Forbs 6 2 B C 2 13 9 5 15 14 2 6 11 4 2 4 4 10 5 1 2 3 8 12 10 2 5 1 2 1 2 Total C 1 2 1 2 Selagine11a B 2 1 2 Co 11 ins ia parvi flora co ronop ifo 1 ia C B May 2 1 2 arvense March 1 2 1 2 arvense Jan Cd 1 2 18 10 37 23 16 5 37 22 30 2 11 14 32 2 3 25 0 �364 APPENDIX C (continued) Nov Forage b Year c bd Jan c e b March c b May b c c BROWSE Ame1anchier (leaves) a 1n ifo 1ia Ame1anchier (stems) a 1n ifo 1ia 1 2 1 2 Arctostaphylos uva-ursi 1 2 Artemi s ia frigida 1 5 2 2 Eriogonum umbe11atum (inflorescences) 2 Eriogonum umbe11atum (leaves) 2 9 Physocarpos (leaves) monogynus 1 2 6 Physocarpus (stems) monogynus 1 3 1 1 2 11 8 2 6 11 30 3 4 2 Populus tremu10ides (leaves) 1 2 5 56 Populus tremuloides (stems 1 2 3 Potentil1a fruiticosa (inflorescences) 1 2 Prunus virginianus (stems 2 Purshia tridentata (stems) 2 Ribes cereum (leaves) 2 3 24 15 10 11 16 5 1 5 10 11 0 18 11 9 1 2 4 7 7 1 Ribes cereum (stems) Rosa acicu1aris 1 .2 1 2 18 Rubus deliciosus (leaves) 1 2 2 2 8 7 Rubus deliciosus (stems) 1 5 30 2 Total 2 Browse 1 2 4 5 30 18 - 2 2 2 6 3 3 23 19 18 5 2 2 14 6 56 76 7 26 69 44 15 4 66 47 �365 APPENDIX C (continued) aMajor forages include those which contributed 2% or more of total bites eaten by mountain sheep on any replicate plot. bForages were differentiated by plant species and plant part and between green and dead material. cYear 1 = 1980~81 d B = Burn Year 2 = 1981-82 C = Control �366 APPENDIX D Mean percentage (g/100 g drymatter intake) of major forages in diets of 4 mule deer grazing in ~urned and unburned grassland com~unities in Colorado during November, January, March, and May 1980-81 and 1981-82.a Nov Forage b Year c B d Jan Cd B May March B C B C C GRAMI NOI DS spp. Agropyron .1 2 4 6 Agropyron spp. (green tillers) 1 2 Bromus tecto rum (green leaves) 1 2 16 Carex spp. 1 2 2 Hesperochloa kingii 1 2 2 8 2 27 36 10 9 2 10 Poa spp. ""l'dead leaves} 1 2 Poa spp. Tgreen 1eaves} 1 2 31 32 Total Green Graminoids 1 2 Total Gruminoids 1 2 2 2 5 15 26 55 79 4 77 27 69 27 78 76 5 26 81 81 4 77 55 71 72 88 76 33 26 32 86 81 20 77 57 79 72 89 6 14 4 28 28 20 10 76 5 31 32 20 10 43 32 21 10 FORBS Agoseris Antennaria glauca parvifolia 1 2 1 2 Aster flaevis 1 2 Aster porter i 1 2 Cerastium Chrysopsis arvense spp. Cirsium spp. 1 2 1 2 1 2 5 5 8 5 4 4 17 5 2 21 20 12 2 3 27 2 2 4 2 2 4 9 5 ,1 1 �APPENDIX Forage D (cont inued) Nov c Bd Ct:i Year b Jan March B C B May C B C FORBS (continued) Erigeron spp. 1 2 Galium boreale 1 2 Geranium 1 2 freemontii Oenothera coronop ifo 1ia Penstemon virens Potentilla spp. Selaginel1a Senecio spp. 1 2 1 2 Total Forbs BROWSE Artemisia frigida 2 2 1 2 officinale dubius 1 7 1 2 1 2 Tragopogon 6 1 2 integerrimus Taraxacum 3 2 4 1 2 27 1 2 34 35 1 2 Erigonum umbellatum (1eaves) 1 2 Physocarpus (leaves) monogynus 2 Physocarpus (stems) monogynus 11 2 6 4 1 17 16 21 6 3 7 3 5 4 16 15 8 19 29 33 22 48 27 28 2 1 1 7 5 6 3 4 11 8 7 8 1 2 2 5 2 1 2 2 4 1 2 9 2 Populus tremuloides {1eaves 1 2 Potentilla fruiticosa (inflorescences) 2 28 44 2 15 1 4 Potentilla fruiticosa (stems and leaves) 1 7 2 10 Prunus virginianus (leaves) 2 17 5 1 2 6 31 2 12 3 3 2 �APPENDIX D (~ontinued) Nov Forage b Year c Bd BROWSE (continued) Prunus virginianus {stems 2 Ribes cereum (Ieaves) 2 Ribes cereum (stems) 2 Cd B B C I 2 3 7 C 11 18 5 2 9 5 2 Rubus deliciosus (leaves) 1 2 6 2 2 Rubus deliciosus (stems) 1 12 5 2 2 1 2 24 30 Total Browse B 3 I 2 Rosa acicularis C 17 1 1 May March Jan 5 2 50 2 48 55 19 40 11 1 27 63 15 7 aMajor forages include those which contributed 2% or more of total bites eaten by mountain sheep on any replicate plot. b c d Forages were differentiated by plant species and plant part and between green and dead material. Yea r 1 = 1980-81 B = Burn Year 2 - 1931-82 C = Control � https://cpw.cvlcollections.org/files/original/17bcc0fc18d44d185d7d16cd091c3c2c.pdf 31a238af4e59468889ea23e4e412bda8 PDF Text Text JOB PROGRESS REPORT STATE OF PROJECT Colorado -------------------------NO. FW-26-R-1 ----------------------- WORK PLAN NO. JOB TITLE: Research JOB NO. Evaluation of Law Enforcement Period Covered: Author: Law Enforcement Activites 1 July 1982 through 30 June 1983 John F. Smeltzer Personnel: Clait E. Braun, Dave Croonquist, Richard M. Hopper, Donald L. Horak, Bob Leasure, Kris Moser, John F. Smeltzer, Area Wildlife Managers, District Wildlife Managers, Colorado Division of Wildlife. ABSTRACT An evaluation of the daily activities of District Wildlife Managers (DWM's) and Area Wildlife Managers (AWM's) of the Colorado Division of Wildlife (CDOW) was conducted from 1 July 1982 through 30 June 1983 as the initial phase in a newly created and ongoing wildlife law enforcement research program. Daily activity reports (DAR's) were collected from 12 of the 16 CDOW administrative areas. Quantifiable data, including days and hours worked per month; days off per month; and days leave taken per month were totaled for each DWM. A total of 514 months of daily activity reports from 32 DWM's was examined. A subsample of 133 months of activity-coded DAR's from 10 of the 32 DWM's was analyzed by activity-type in addition to the more general analysis. Four activities: wildlife inventory, law enforcement, public relations, and administrative and clerical services accounted for as much as 88% of the total time worked by DWM's during a given month. Mean values for DWM activity time were: 201.5 hours worked per month, 23.48 days worked per month, 5.6 days off per month, 1.36 days leave per month. AWM's spend the majority of their time in budget preparations, meetings, distribution of equipment and supplies, and report preparation. There is no line authority between the AWM's and the Chief of Law Enforcement. The Regional Law Enforcement Coordinators (RLEC's) are responsible for planning, implementation, monitoring, and coordination of all Regional law enforcement activities. The RLEC positions may be extremely important to the standardization and coordination of law enforcement activities statewide as well as regionally. A review of the wildlife law enforcement research literature demonstrated 3 basic facts: significant amounts of descriptive and philosophical wildlife law enforcement literature are available; wildlife forensic research is an accepted research area with much potential for growth and problem-specific �2 application; non-forensic wildlife law enforcement research is slowly evolving as a science and has virtually unlimited growth potential if it develops as an integrated discipline. Recommendations include: continued use of the present daily activity reporting system until a practical and functional computer-based system can be developed, the reasons for the system must be clearly defined; development of quantifiable wildlife law enforcement objectives; identification of specific areas in which need to evaluate levels of efficiency and effectiveness; development of an interactive computer-based information retrieval system available in all Regions to include information on hunting/fishing license revocations, general license information, hunter safety card information, and special files for recording and analysis of other law enforcement information considered worthy enough to collect; development of statewide, regional, and district wildlife law enforcement priorities ranked with district fisheries and wildlife management priorities; development of comprehensive district profiles. we �3 EVALUATION OF LAW ENFORCEMENT ACTIVITIES John F. Smeltzer Enforcement of wildlife laws is an integral part of successful wildlife management programs. Increased demands on wildlife and fisheries resources from increasing numbers of recreationists (hunters, fishers, animal watchers, etc.) and decreasing amounts of quality habitat make it imperative that fishery and wildlife populations be properly managed for the benefit of all people. With increasing numbers of people interested in fish and wildlife, more leisure time, and sophisticated equipment, wildlife managers find it difficult to efficiently enforce wildlife regulations over large areas. Wildlife law enforcement research is relatively new although substantial research has been done on wildlife forensics particularly the identification of cause and time of death of a variety of animals, and species identification. Additional research on wildlife law enforcement problems is needed to measure and improve the efficiency and effectiveness of wildlife managers and to provide tools for use in the field. However, before wildlife law enforcement research needs can be clearly identified, it is necessary to evaluate present wildlife law enforcement activities so that research efforts can be properly directed. P.N. OBJECTIVES 1. Examine and evaluate available wildlife managers in Colorado. data on law enforcement 2. Prepare a study plan on an appropriate research topic. wildlife activities of law enforcement S.N. OBJECTIVES 1. Examine available data on law enforcement Area Wildlife Managers. activities 2. Evaluate available data on law enforcement Area Wildlife Managers. 3. Interview selected personnel of the Colorado Division of Wildlife concerning law enforcement research needs. activities of District and of District and 4. Review available literature on wildlife law enforcement problems and past research and management efforts. 5. Prepare a study plan on an appropriate approved wildlife law enforcement research topic. 6. Compile data, analyze results, and prepare progress report(s). �4 METHODS Daily Activity Reports Daily activity reports (DAR's) were collected from 12 of the 16 Division of Wildlife administrative areas. All 4 Regions were represented. The sample consisted of 514 months of activities by 32 District Wildlife Managers (DWM's, 30% of total) from January 1979 through June 1982. Total months evaluated for any individual DWM ranged from 4 to 28. Quantitative data, including days and hours worked per month, days leave. taken per month, and days off per month were totaled for each DWM. The information was used to calculate ranges and means by month for the following: days worked, hours worked, days leave taken, and days off. Weighted grand means were calculated for each parameter for both annual and monthly totals. Coded Daily Activity Reports Coded daily activity reports were received from 10 D\.JM's: Their daily activities had been coded using a series of 18 activity classifications developed for the Division of Wildlife "EARS" program (Employee Activity Reporting System, Appendix A). This reporting system is no longer in general use. Work time per activity was recorded to the nearest 0.5 hour. A total of 133 months was evaluated. This sUbsample represented approximately 25% of the total months examined. Quantitative data were totaled by coded activity for each DWM. Calculations were made to determine mean hours and mean percent time spent on a coded activity by each DWM during a l-year period, January 1981 through December 1981 (106 months of coded work) and a 1.5-year period, January 1981 through June 1982 (133 months of coded work). Weighted monthly and annual means were calculated by activity-type for both periods. Annual leave hours (Code 78) were excluded in 2 series of calculations (Tables 1, 2). The percentages shown reflect time worked per activity divided by total hours actually worked (x 100). Code 78 data (annual leave) were included in Table 3. These data represent time spent on an activity divided by the total hours the DWM was paid salary by the State (x 100). The category "uncoded" was included (Tables 1, 2, 3) to show an approximate coding error rate expressed as a percent of total hours worked. All hours included in this category had not been coded or were obviously miscoded. Field Observations Twenty-nine days were spent on field observations of the law enforcement effort. Personal contact was made with 32 DWM's, 6 Area Wildlife Managers (AWM's) and 4 Regional Law Enforcement Coordinators (RLEC's) in addition to the Chief of Law Enforcement and his 2 assistants. Subjective interviews were conducted with these personnel regarding wildlife law enforcement activities in Colorado and research needs. �5 Literature Review and Current Research Available wildlife law enforcement research information and wildlife law enforcement articles of a descriptive or philosophical nature, published and unpublished, were collected and reviewed. A short, informal questionnaire was sent to each of the 50 states, 4 Canadian provinces, and Puerto Rico. The questionnaire was intended to identify current or recent law enforcement research/evaluation efforts and perceived wildlife law enforcement related problems. Study Plans Two study plans were drafted, a 3rd outlined, and a 4th proposed. The 1st, Illegal Purchase of Resident Licenses by Non Residents, is in its 2nd revision. The 2nd, Preparation of an Annotated Bibliography for Wildlife Law Enforcement Personnel is being revised. A draft-outline has been completed on the development of a 3rd study plan tentatively entitled: Public Perceptions of the Law Enforcement Effort and the Role of Wildlife Law Enforcement in Colorado. The 4th proposed study plan would undertake the development of measurable law enforcement objectives. Initial recommendations for improving the efficiency and effectiveness of the wildl ife law enforcement effort of the Division of Wildlife are suggested. RESULTS AND DISCUSSION Coded Data District Wildlife Managers of the Colorado Division of Wildlife operate under the "multi-purpose" concept. Their job is, by description, multifaceted; a composite of many activities (Appendix B). Law Enforcement is only one component of this multifaceted position. Any analysis of the law enforcement activities of the DWM must be done in relationship to other activities. Reported time relationships between these activities are illustrated in Tables 1, 2, and 3. Four activities: Code 10, wildlife inventory; Code 18, public relations; Code 38, law enforcement; and Code 70, administrative and clerical services accounted for as much as 88% of the total time worked during a given month (~ ~ 77.5%, range 63.0-88.0%, Tables 1, 2, 3). Percent time spent on wildlife inventory (Code 10) was highest in January ( ~ 17.5%) and lowest in February (~ 7.0%). This was probably attributable to early winter survey counts of big game and waterfowl. Percent time spent on public relations (Code 18) remained relatively stable throughout the period but showed a slight increase during both May (early fishing) and October (big game season) reflecting increased public contact by DWM's. Law enforcement (Code 38) represented only 25% of the worktime in February but peaked at �Table 1. Percent from 1 region).a time spent on an activity by month. (Based on reported coded data from Dally Activity Reports of 10 District Wildlife Managers (J'\ Code 10 Month 14 18 - 20 26 30 34 38 b 42- --46 50 54 17.750.1914.021.202.634.150.5527.380.030.134.051.78 Jan 58 62 66 70 74 Uncoded 0.26 2.163.140.1318.981.46 feb 7.00 2.28 13.02 2.34 0.42 3.38 1.84 24.83 1.10 0.63 5.49 3.76 0.89 5.68 0.08 18.16 8.67 0.44 Mar 10.10 1.30 17.67 0.72 0.69 3.97 0.50 27.68 3.62 1.26 4.80 1.68 1.43 2.80 0.08 14.83 5.83 0.34 Apr 12.184.5415.871.240.223.950.1829.240.330.375.85 May 11.58 0.66 20.54 0.74 0.39 3.37 0.04 33.11 0.66 1.47 5.30 1.05 1.16 1.32 0.00 13.98 4.59 0.64 Jun 11.00 0.96 17.63 0.05 1.42 4.53 0.14 32.31 0.00 1.63 7.88 0.96 0.91 0.29 0.05 15.90 5.19 0.24 Jul 12.94 0.50 19.05 0.00 0.00 2.68 0.00 34.27 0.20 r.07 7.38 0.13 1.34 0.80 0.27 16.53 2.62 0.20 Aug 11.980.8117.140.270.074.761.0029.351.010.946.410.13 1.14 1.270.4015.335.84 2.15 Sep 11.760.0017.601.801.113.560.5240.531.561.243.980.00 2.97 0.00 0.0012.100.83 0.45 Oct 11.66 0.21 20.24 0.16 0.52 1.66 0.26 45.31 0.23 0.00 4.69 0.00 2.07 0.31 0.52 10.81 0.31 1.04 Nov 12.18 1.19 18.40 0.17 0.00 2.71 0.07 42.44 0.07 0.20 3.81 1.59 1.26 0.60 0.20 14.13 0.93 0.07 Dec 13.34 0.69 19.35 0.07 0.55 2.80 0.00 38.91 0.14 0.73 3.90 0.35 2.14 0.76 0.00 15.34 0.73 0.21 Weighted annual mean 12.03 1.24 17.26 0.75 0.76 3.43 0.46 32.67 0.79 0.79 5.28 1.26 1.49 1.88 0.33 15.59 3.48 0.50 apercentages bSee Appendix derived from 133 months of coded activities A for description of activity. 1.650.952.301.1017.322.38 from January 1981 through June 1982. All leave 0.33 hours excluded. �Table 2. Percent from 1 region).a time spent on an activity by month. {Based on reported coded data from Daily Activity Reports of 10 District Wildl ife Managers Codeb 10 14 26 30 34 50 54 58 62 66 70 74 Jan 16.66 0.26 14.96 1.94 1.94 1.89 0.84 29.31 0.00 0.10 4.61 2.88 3.04 0.31 0.00 18.96 1.99 0.31 Feb 6.85 2.72 13.03 2.55 0.65 2.46 2.26 27.90 1.10 0.65 6.30 5.69 1.36 1.26 0.00 18.17 6.63 0.55 Mar 10.20 0.05 20.60 0.76 0.98 3.96 0.71 31.59 1.47 1.79 6.62 1.68 1.66 0.95 0.11 14.33 2.28 0.27 Apr 11.65 4.11 16.18 1.46 0.31 3.85 0.26 28.64 0.47 0.52 7.34 1.72 1.35 0.73 1.30 16.62 3.02 0.47 May 9.87 0.95 20.49 0.22 0.56 4.21 0.06 33.31 0.00 0.62 6.78 0.90 1.46 0.28 0.00 13.18 6.42 0.59 Jun 9.62 1.40 18.48 0.07 0.88 2.92 0.28 33.26 0.00 0.49 10.82 0.56 1.07 0.42 0.00 15.60 3.65 0.21 Jul 12.94 0.50 19.05 0.00 0.00 2.68 0.00 34.27 0.20 1.07 7.38 0.13 1.34 0.80 0.27 16.53 2.62 0.20 Aug 11.98 0.81 17.14 0.27 0.07 4.76 1.00 29.35 1.01 0.94 6.41 0.13 1.14 1.27 0.40 15.33 5.84 2.15 Sep 11.76 0.00 17.60 1.80 1.11 3.56 0.52 40.53 1.56 1.24 3.98 0.00 2.97 0.00 0.00 12.10 0.83 0.45 Oct 11.66 0.21 20.24 0.16 0.52 1.66 0.26 45.31 0.23 0.00 4.69 0.00 2.07 0.31 0.52 10.81 0.31 1.04 Nov 12.18 1.19 18.40 0.17 0.00 2.71 0.07 42.44 0.07 0.20 3.81 1.59 1.26 0.60 0.20 14.13 0.93 0.07 Dec 13.34 0.69 19.35 0.07 0.55 2.80 0.00 38.91 0.14 0.73 3.90 0.35 2.14 0.76 0.00 15.34 0.73 0.21 Weighted annual mean 11.61 1.09 17.98 0.81 0.65 3.11 0.52 34.46 0.51 0.68 6.03 1.34 1.79 0.63 0.25 15.08 2.92 0.54 Month apercentages bSee Appendix derived ~ -18~----22 from 106 months of coded activities A for description ~-42-~q6 from January 1981 through December Uncoded 1981. All leave hours excluded. of activity. -...J �Table 3. Percent time on an activity by month. from 1 region).a {Based on reported coded data from Daily Activity Reports of 10 District Wildlife Managers 00 Codeb 10 111 Jan 17.19· 0.19 Feb 6.86 Mar 42-- 46 50 511 58 62 0.13 3.93 1.73 2.09 3.05 1.07 0.62 5.37 3.68 0.87 26.63 3.48 1.21 4.62 1.61 0.18 28.66 0.32 0.36 5.73 3.32 0.04 32.60 0.65 1.45 1.17 2.68 0.12 26.67 0.00 0.00 0.00 2.42 0.00 30.95 15.41 0.24 0.06 4.28 0.91 0.00 15.17 1.55 0.95 3.07 11.66 0.21 20.24 0.16 0.52 Nov 12.18 1.19 18.40 0.17 Dec 12.32 0.64 17.87 0.06 Weighted annual mean 11.361.1716.290.710.72 22 26 30 13.58 1.16 2.54 4.02 0.53 26.52 0.03 2.23 12.75 2.29 0.41 3.31 1.80 24.32 9.72 1.25 17.00 0.70 0.66 3.81 0.48 Apr 11.94 4.44 15.56 1.22 0.22 3.87 May 11.40 0.65 20.23 0.72 0.38 Jun 9.08 0.79 14.56 0.04 Jul 11.69 0.45 17.20 Aug 10.76 0.72 Sep 10.13 Oct Month 18 70 74 78 Uncoded 0.13 18.39 1.41 3.14 0.25 5.56 0.08 17.79 8.49 2.07 0.43 1.38 2.70 0.07 14.26 5.61 3.81 0.33 1.61 0.93 2.26 1.08 16.97 2.33 2.01 0.32 5.22 1.03 1.14 1.30 0.00 13.77 4.52 1.53 0.63 1.35 6.50 0.79 0.75 0.24 0.04 13.13 4.28 17.45 0.20 0.18 0.97 6.67 0.12 1.21 0.73 0.24 14.93 2.36 9.96 0.18 26.38 0.90 0.84 5.76 0.12 1.03 1.15 0.36 13.78 5.25 10.13 1. 93 0.45 34.92 1.34 1.07 3.43 0.00 2.56 0.00 0.00 10.43 0.72 13.83 0.39 1.66 0.26 45.31 0.23 0.00 4.69 0.00 2.07 0.31 0.52 10.81 0.31 0.00 1.04 0.00 2.71 0.07 42.44 0.07 0.20 3.81 1.59 1.26 0.60 0.20 14.13 0.93 0.00 0.07 0.51 2.58 0.00 35.93 0.13 0.67 3.61 0.32 1.98 0.70 0.00 14.17 0.67 7.66 0.19 31\ 38 -66-- 0.47 3.240.4430.840.750.744.991.191.411.770.3014.723.295.60 apercentages derived from 133 months of coded activities from January 1982 through June 1982. bSee Appendix A for description of activity. AI I leave hours included. �9 45% in October during the big game seasons. Administrative and clerical services (Code 70, "paperwork") required 19% of the total worktime in January but only 11% in October. These numbers are virtually meaningless when considered alone in the absence of objective statewide standards for each activity. The information they provide relates to the relative distribution of worktime as described by this subset of 10 DWM's from 1 Region when those DWM's were restricted to 18 coded activities and definitions. They cannot be used to project specific statewide values with any degree of confidence. The percentages may suggest the relative ranked priority of an activity during a month as described by the DWM's. These percentages would not be the best indicator of total workload during that month. Workload must be considered on a combination of characteristics and not just hours worked. Workload Beattie (1976~) described the workload of wildlife law enforcement officers: "The term workload, as it appl ies to wi ldl ife law enforcement sections and their employees, will vary with the role of the officer. The workload of an officer represents the sum of all activities he (she) performs or is expected to perform. Actual (based on objectives) and perceived workload may be viewed differently by individual officers and the wildlife law enforcement agency. Some officers may, in actuality, have a lighter workload (based on agency criteria) than other officers but may perceive themselves as having a heavier workload. The important point is that agencies must establish and articulate meaningful criteria for determining individual officer workload and wildlife agency organizational workload." Wildlife officer workload would include the hours spent by an individual officer in the performance of his/her duties. Hours worked or percent time spent per activity would be one indication of workload. However, hours worked or percent time does not address the quality of those hours or the efficient and effective use of the time coded into a given activity category. District Wildlife Managers throughout the State consistently reported work hours in excess of those required of State employees by personnel regulations. It is the feeling of most field personnel that extra hours must be expended to properly do the job required of them. This assumption mayor may not be true. McCormick (1970) documented the contact rate between wildlife agents and the public prior to and following the enactment of a mandatory 40-hour work week for all wildlife officers. McCormick's results suggested that, with improved planning of efforts and better definition of priorities, field contacts per agent actually increased after the 40-hour week was enacted. A 40-hour work week may not be practical in Colorado under the present multi-purpose concept. A clear definition of priorities for the DWM position would be necessary along with average completion times by activity and average number of activities per day before a 40-hour week could be considered. What McCormick's work does suggest is that with an increased planning effort efficiency may be increased. �10 Major Work Period October was the peak work period for law enforcement activity and for total hours worked during the 12-month period January-December 1981. Approximately 45% of all hours worked 'during October were assigned to law enforcement. Mean DWM time expenditure during October was 275 hours and 27 days worked statewide. This is approximately 107 hours more than required by state personnel regulations (DWM's are exempt). On an annual basis, based on the information provided, the "average" DWM worked 417 hours in excess of those required (Table 4). This would suggest that statewide, DWM's reported approximately 47,000 "excess" or "overtime" hours during the period January-December 1981. The qual ity of those hours cannot be determined. Table 5 lists calculated values for individual DWM's. These values include: months evaluated, mean days worked per month, mean days off per month, mean hours worked per month, range in hours worked per month, and mean leave days (of all types) taken per month. A partial profile of an "average" DWM would read something like this: Every month he/she worked an average of 23.48 days; took off 5.6 days; worked approximately 201.5 hours, of which 34 hours could normally be considered "overtime"; and used 1.36 days leave. Hours worked per month could range from 45 (excluding leave hours) to 420. During 1981 he/she issued approximately 40 Penalty Assessments (PAis) and Court Citations combined. Figure 1 graphically illustrates the average hours worked per month on each activity during a 1-year period, January-December 1981. Wildlife inventory, publ ic relations, law enforcement, and administrative and clerical services are the predominant activities as described by this subset of 10 DWM's. Division equipment and facilities maintenance (Code 50) is the only additional activity that on average required over 10 hours per month to complete. These figures represent an accurate picture of relative hourly work loads as identified by the individual DWM's. If these are to accurately represent the true distribution of DWM activities during January-December 1981 several assumptions must be made: 1. All activities have been defined in a similar manner by every DWM. 2. Time reported for each activity accurately represents the actual time spent on that activity. The quality standard between DWM's is the same. 3. All activities actually undertaken are represented in the daily activity report. If these assumptions are not true, then the validity of the results can be questioned. If these assumptions cannot be satisfied, the use of this technique to gather information must be questioned. Definition Problems Poor definitions can be a severe constraint to categorization. The problem being that "what is" to one person "is not" to another. Careful and thoughtful definitions must be a part of any attempts to code and categorize activities. The system must be thoroughly explained through training so that al 1 �Table 4 . Weighted mean days and hours of reporteG work per montha by 32 DWM's during the period January 1979 - June 1982. (514 months of DAR's evaluated.) Month Days/ month Hours/ month Mean hours/ actual days worked Required work days/month Required hours/ month Mean overtime hours/month Jan 24.86 200.14 8.05 21 i68 32. 14 Feb 20.68 172.29 8.33 18 144 28.29 Mar 24.63 196.02 7.96 22 176 20.02 Apr 23.38 200.52 8.58 22 176 24.52 May 24.72 208.48 8.43 21 168 40.48 Jun 21.96 187.78 8.55 22 176 11 .78 Jul 24.05 210.99 8.77 21 168 42.99 Aug 22.32 192.30 8.62 21 168 24.30 Sep 21 .73 199.20 9.17 . 21 168 Oct 26.95 274.70 10.19 21 168 45.54 Nov 24.08 213.54 8.87 21 168 53.54 Dec 21.56 176.82 8.20 21 168 8.83 ,'" aAnnually calculated mean hours/month = 201.52. Annually calculated 106.7 mean days worked/month = 23.48. �12 Table 5. Calculated values derived from daily activity reports (DAR's) of 32 District Wildlife Managers of the Colorado Division of Wildlife from January 1979 through June 1982. DWM 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 Months evaluated Mean days worked/mo. Mean days off/mo. Mean hours worked/mo. Range hours v:orked/mo. Mean days leave/mo. (a11 t vo e s ) 18 18 13 18 24.50 5.10 212.44 136-319 0.86 23.00 22.08 21.56 6.50 6.70 7.66 198.66 207.85 184.11 153-232 45- 261 56-254 0.94 13 4 12 6 18 18 18 18 24.15 22.00 21. 75 22.33 21.94 21.22 25.30 5.77 8.00 6.67 6.17 6.78 235.88 175.00 192.83 168.17 196-305 144-193 114-264 23.72 26.50 21.54 22.54 24.00 24.92 5.72 3.83 7.15 6.00 23.79 20.42 23.26 23.87 21.45 23.73 22.67 25.92 21.08 24.08 24.96 5.21 7.08 5.83 5.52 6.82 4.64 172.83 207.39 205.52 168.27 182.18 6.50 2.67 7.25 6.08 4.18 25.68 23.17 24.54 3.96 6.33 4.08 23.69 23.48 12 13 24 24 24 24 12 23 23 11 11 12 12 12 12 28 25 12 13 13 Mean (weighted) 7.39 3.94 5.38 4.08 173.90 167.19 187.50 202.80 260.67 172.62 199.00 226.73 242.08 191.42 101-221 108-228 107-208 95-252 150-248 192-328 76-224 143-313 129-420 152-351 95-262 103-259 87-311 114-278 1.54 1.22 0.46 0.00 2.08 1.33 1.67 2.28 1.60 1.06 0.00 1.77 1.91 0.96 1.44 1.29 2.83 1.35 1.04 89-207 116-224 2.09 2.00 204.96 225.67 192.96 188.75 198.04 142-293 162-328 115-240 140-238 71-241 1.25 1.83 2.17 0.25 1.32 5.38 241.52 216.67 176.38 164.08 114-296 134-306 114-329 ')1-226 0.76 0.92 1.85 1.46 5.60 201.52 118-271 1.36 �13 ACTIVITY 10: ACTIVITY 141 LOLl FE INVENTORY 14: REGULATION RECOMMENDATIONS 10 30 8 6 20 4 10 x= 21.63 SE = 5.29 2 o D ACTIVITY 18: PUBLIC FMA RELATIONS MJJ o A ACTIVITY 22: HUHTER ACTIVITY 30: GAME DAMAGE fl D SAFETY 50 4 40 ACTIVITY 26: RESEARCH 10 4 R 6 2 4 o o F M A M A SON Fig. 1. Mean hours spend per month on a defined activity by 10 District Wildlife Managers, N.E. Region, January-December 1981, (note variable scale for hours). D �14 ACTIVITY 34: DISTRIBUTE WILDLIFE ACTIVITY 38: LAW ENFORCEMENT 110 4 = 0.97 x 100 3 90 80 2 70 60 50 X 40 o 64.18 SE = 16.46 J F ACTIVITY M 42: A M HABITAT J J A S o N D DEVELOPMENT ACTIVITY 46: PUBLIC FACILITIES 3 2 o SE = J ACTIVITY 50: DIVISION FACILITIES F ACTIVITY II A II J J 51.: ENVIRONMENTAL o A o N PROTECTION 19 10 x 16 R SE = = 2.49 2.56 6 13 4 10 7 0 J Fig. 1. (continued) F M A M J A 0 N D �15 ACTIVITY 58: RESOURCE ACTIVITY RECONNAISSANCE 62: PLANNING/BUDGETING 5 x = 1. 18 4 0.65 2 3 2 x SE = = 3.33 1.29 o 0 F ACTIVITY 66: J M A M A N 0 F 0 REAL ESTATE ACTIVITY M 70: A M J A SON D AOMINISTRATIVE/CLERICAL 3 x = 2~.09 35 SE = 3.55 2 30 25 20 o J FHA ACTIVITY M J A S () N o D ACTIVITY IN-SERVI CE 74: 78: N D LEAVE 40 15 x = 3.85 30 12 9 20 6 10 3 0 0 F Fig. 1. H A H J (continued) J A 0 N 0 H A M A S 0 il 0 �16 participants will respond in similar fashion to the question '~hat category should I place this activity in?" and all participants must report all activities correctly. The coded information examined in this study must be treated carefully with the understanding that it represents the best information available concerning daily activities of DWM's, but that it may contain serious reporting errors based on differential judgment regarding definitions of different activities. Area Wildlife Managers The Colorado Division of Wildlife (CDOW) has 20 Area Wildlife Managers (AWM's or Area Supervisors), with 1 assigned to each of the 16 CDOW Administrative Areas and 4 serving as Regional Law Enforcement Coordinators (RLEC's). One RLEC is assigned to each Region. Like the DHM, the AWM position is multi-faceted (Appendix C). Generally speaking, AWM~s are selected following an examination from the DWM ranks, although in theory it is possible to move into the AWM position after 4 years of experience at the DWM level or above. This experience must have included enforcement arid administrative responsibilities. AWM's are directly responsible to the Assistant Regional Manager in their Region. The majority of the AWM's time (exclusive of those AWM's serving as RLEC's) is spent in budget preparation, attendance at meetings, distribution of equipment and supplies, report preparation, and public contacts. AWM's typically work with DWM's as a supervisory field enforcement officer during peak law enforcement periods and on special enforcement problems. There is no line authority from the Chief of Law Enforcement or his assistants to any field law enforcement position, including the AWM and DWM as well as other CDOW commissioned officers. This lack of line authority can complicate the implementation of new law enforcement procedures. Likewise, it has hindered the standardization of law enforcement practices on a statewide basis. With the creation of the RLEC position some of these difficulties may be surmounted. The 4 RLEC's are responsible for planning, implementation, monitoring, and coordination of all Regional law enforcement activities. It is the RLEC's position to conduct or supervise special regional investigations; plan, conduct, evaluate, and coordinate regional law enforcement training; conduct public education and information programs; and to serve as regional officer for license suspension hearings. The RLEC position may prove to be critically important to the standardization and coordination of law enforcement activities statewide as well as regionally. CONCLUSIONS Coded Daily Activity Reports Coded daily activities can be effectiveness of a program or activity are weighted against can be evaluated. This would valuable in evaluating the efficiency and an individual. When total hours expended per agency objectives, the direction of the efforts not be a total measure of effectiveness, only �17 an indicator. Before a complicated questions must be answered: reporting system is constructed 1. What do we want to measure the efficiency 2. How would these efforts and effectiveness difficult of? impact our total program? 3. Can we make such a system work? 4. How will the information obtained be used? 5. How are the results to be communicated to the individuals who provided the information? 6. Who will be responsible for implementing, monitoring, and evaluating these efforts? Identification of the answer(s) to question #1 is (are) extremely important. Levels of efficiency and effectiveness can be measured only if we can clearly define what we are to measure and then are given a standard to use in comparison. Lacking a standard, one must first be developed. This must be done before any modifications are made to a program if we expect to measure the results of those modifications. Unless we do, we are not truly concerned with those impacts. Standards can be developed in 2 ways: known outputs such as miles driven, contacts made and recorded, programs given, articles written, reports completed, and arrests made can be used to set a "standard". The 2nd way involves establishing "desired" levels for difficult to quantify areas, such as tolerable levels of violation for specific laws. These standards would be developed based on informed discussions; establishing and implementing methods to measure current levels; and finally adjusting the standard as more information becomes available or as collection techniques improve. Objective evaluations are not possible without standards for comparison. The impact of any evaluation on the overall program or portions thereof must be considered. Just because we have the ability to evaluate a specific activity or function does not mean that we should evaluate it. Internal and external resistance to any evaluation program should be considered. A comprehensive reporting system will only be as good as the data received and the support system developed to evaluate it. It will be necessary to clearly define all activities or functions that will be monitored and evaluated. It is important to remember that just because we can measure or report something doesn't automatically mean that we should do so. To encourage accurate reporting we should allow flexibility in some activity definitions and demand specific definitions in other activity areas. Above all, we should use the information obtained. Use of the information obtained is vital to the continuation of the system. If we don't use it, we probably shouldn't collect it. Use of the information to positively impact the overall program will encourage accurate and consistent reporting of data. �18 Any information collected, analyzed, and reported should be made available to those individuals who spent time in the collection and analysis of the data. This is true of all data collection functions. Failure to complete the communication process by not supplying feedback to those who originally supply the data will result in rapid deterioration of the reporting system. Responsibility for the implementation, monitoring, and evaluation of any data collection efforts must be clearly defined. Failure to do so will result in another system in which lack of accountability causes the system to fai 1. Current Daily Activity Reports The current daily activity reporting form (DARls uncoded) provides the information necessary to evaluate days and hours worked per month, days leave taken per month, days off per month, and gives a brief narrative description of the daily activities of the DWM or AWM. In its present form and as it is typically used, the DAR does not provide a convenient way to list daily activities by category or to quickly analyze them. +t is, however, an adequate method to provide the information currently used by the CDOW to monitor the work of its' personnel. With the addition of clearly defined, coded activities, it could be used as a coding form and could provide additional information on specific daily activities of all personnel. A support system would be needed to collect and analyze any information gathered. Training would be required to promote consistent reporting of activities. Any changes in the system must be evaluated and directed to impact the overall Division functions. Standardized Reporting Lack of standardized reporting of law enforcement information is due largely to the decentralization of law enforcement authority. Regional 1awenforcement programs have evolved based in theory, on the needs of a particular Region. Historically, the Division has apparently seen no need to standardize its reporting of law enforcement information or has found the task to be too complicated. The creation of the RLEC position should aid in the standardization of statewide law enforcement efforts. Standardized reporting and a system to collect and analyze these data must be in place before substantive efforts can be made and sustained to evaluate present levels of enforcement efficiency and effectiveness. Concurrent with these efforts must be the development of measurable wildlife law enforcement objectives. Beattie et a1. (1977a) stated: "A clear and precise formulation of wildlife law enforcement objectives must precede an enforcement research program; they must be operationally defined so progress can be measured.11 Simply stated, we must know where we are, where we want to go, and how we will know when we get there. Furthermore, we must be able to articulate these objectives to others. �19 Recommendations 1. Continued use of the present Daily Activity Reporting system is recommended with reservation. Administrative discussion and decisions must occur that specifically address the need for and the benefits of a more comprehensive activity reporting system. This system should be based on coded activity data which would be computer filed, analyzed, and used to impact the overall Division function. The system must be practical, functional, and have specific objectives. 2. A system similar to that described in #1 above is necessary before substantive sustained evaluations of the statewide efficiency and effectiveness of field law enforcement/management personnel can occur. Evaluation efforts initiated prior to the implementation of the described system will be superficial because they can only look at site-specific information and not at statewide problems. 3. Quantifiable wildlife law enforcement objectives must be considered and an initial attempt made to develop an array of objectives specifically designed to address the current and future need of wildlife law enforcement efforts. These objectives must be operationally defined so progress can be measured. 4. Standardized reporting forms should be developed and their use required, after testing, in all Regions. Space should be provided on the forms so that information of Regional interest only can be recorded, if needed. There should be statewtde discussions regarding the nature and importance of this information, how it is to be recorded, how it should be tracked, and how it should be evaluated. The RLEC's should playa major role in the development of a standardized reporting system. These efforts should be coordinated through the Regions and promoted by the Denver Wildlife Law Enforcement Staff. 5. An interactive computer information retrieval system should be available in each Region to permit entry, analysis, and print-out of information that has previously been determined valuable enough to collect. The system should include current data on hunting/fishing license revocations, prior violation records, license information, hunter safety card information, and special files to record data used in overall analysis of the law enforcement effort. 6. Statewide, regional, and district wildlife law enforcement priorities should be developed ranked with district ment priorities. These ranked priorities Managers clearly identify their wildlife ment, and fisheries management roles and activities. wildl ife and fisheries managewould help District Wildlife law enforcement, wildlife managewould aid the scheduling of �20 7. District profiles should be developed through a joint effort of CDOW personnel to provide information specific to a district. This information would be used as an aid to trainees or transfers into a new district and as a reference manual for established DWM's. It should be periodically updated. It should include information on specific management practices used in that district; law enforcement activities, including historic information on contact rates, arrests, particular law enforcement problems; influential individuals and publics; human population levels, socio-economic structure and distribution; major geographical and environmental features, miles of road by surface type, acres of water, miles of river; major fisheries and wildlife concerns, both game and non-game; past or present research efforts and any written and/or published materials resulting from those efforts including approved study plans and job progress or final reports. Overview of the Literature Wildlife law enforcement research does not have a well developed literature base., Much of the information that is available is fragmented, poorly reported, incomplete, or the results are based on unfounded assumptions and poor research design. A significant portion of all published wildlife law enforcement information is philosophical or descriptive in nature but does generally contribute to the overall understanding of problems surrounding wildlife law enforcement efforts. Wildlife forensic research and much of the administrative-type research conducted at Virginia Polytechnic Institute and State University (VPI & SU), under the guidance of R. H. Giles, Jr. are notable exceptions to t~is descriptive or philosophical approach. In both cases, diligent attempts are being made, or have been made to maintain the standards of the scientific method. Wildlife law enforcement needs well designed research to help it evolve as a science. Forensic research must be carefully conducted so that results and applications can withstand rigorous testing in the Courts. Work by Brohn and Korschgen (1950) helped establish the use of the Precipitin test as a tool of wildlife law enforcement following earlier work by Gay (1908) and Clark (1914). This test is useful in the identification of many animal species using only meat or blood samples. Oates et al. (1974) and Glover and Korschgen (1980) have recently experimented with this particular technique. Johnson et al. (1980) used potassium levels in the vitreous humor of eyes from mule deer (Odocoileus hemionus) to calculate time of death. This work has recently been replicated in Missouri (R. Glover, pers. commun.) and Illinois (Woolf and Roseberry, no date). Much additional forensic research has been conducted and interested readers are referred to the publication by Wilson (1977) for additional information. �21 Virginia Polytechnic Institute Studies Significant wildlife law enforcement research was undertaken by R. Giles and associates at VPI & SU during the 1970's as part of a joint law enforcement research effort. Funding came from a variety of sources including: the Wildlife Management Institute, National Wildlife Federation, National Rifle Association, American Petroleum Institute, Georgia State Game and Fish Division, South Carolina Wildlife and Marine Resources Agency, Tennessee Wildlife Resources Agency, and the Virginia Commission of Game and Inland Fisheries (Giles 1976b). These studies were some of the first attempts at an objective examination-of the wildlife law enforcement activity as an integral part of the overall job of wildlife and fisheries management. Giles has been a primary motivating force behind wildlife law enforcement research since the 1960's when he served as advisor to J. R. Vilkitis at the University of Idaho. Vilkitis' research on the illegal take of big game and characteristics of big game violators in Idaho (Vilkitis 1968) was the first attempt to quantify poaching losses of big game. At VPI & SU, Giles continued law enforcement research projects into the late 1970's (Giles 1971; 1976~, ~; 1978; Giles et al. 1971). Other examples of the work and extent of the projects supervised by Giles at VPI & SU include: Kaminsky (1974) examined the problem of white-tailed deer (Odocoileus virgianus) spotlighting in Virginia. His analysis helped to demonstrate that research could be conducted in wildlife law enforcement. Ritter (1975) developed measurable law enforcement objectives and performance criteria for state wildl ife agencies. Ritter's work~co~ld serve as the foundation in the development of comprehensive evaluation programs by other state wildlife agencies. Beattie (1975; 1976a, b; 1978a, b; 1979; 1981a, b) and associates (Beattie et al. 1977a, b, c; 1978a, b, c; 1980) has been-aprolific writer and an advocate of welT designed-law enforcement research projects. C. Cowles, a contemporary of Beattie's at VPI & SU, and also a student of Giles was instrumental in the development of optimum deployment techniques, based on computer models, for wildl ife law enforcement agents (Cowles 1977; Cowles et al. 1977, 1978, 1979). The collective efforts at VPI & SU demonstrate the diversity possible for research in wildlife law enforcement. This work was by no means definitive, but should serve as the foundation for more comprehensive, integrated wildlife law enforcement research. Integration will be a key word in successful wildlife law enforcement research. The problems facing wildlife law enforcement today are not strictly biological. The disciplines of Biology, Chemistry, Ecology, Sociology, Psychology, Politics, Economics, Computer Science, Statistics, Engineering, and Police Science must be fully integrated to help solve these problems. �22 Other Research Efforts Other individuals and agencies have had active roles in the evolution of wildlife law enforcement research. W. B. Morse, now with the Wildlife Management Institute, gathered wildlife law enforcement information for many years on a regional and later on a national basis. His reports to the Western Association of State Fish and Wildlife Agencies and other published works have stimulated thoughts on needed law enforcement research (Morse 1957, 1958, 1963, 1968, 1972, 1973, 1976, 1980, 1982). The law enforcement divisions of Michigan, California, and New Mexico have each conducted research projects to measure the effectiveness of their law enforcement efforts (Hussain 1977; Purol 1982a,b); Purol and Fournier 1979; Purol and Zambetis 1982; McCormick 1968, 1970; Mikel 1981a,b; Pursley 1977a,b; Vaught 1975; Vaught and Turner 1975). These projects have met with varying success but have been successfully used to impact their law enforcement operations. Field oriented wildlife law enforcement research efforts have been reported from most states (Smeltzer 1983, Appendix D; Beattie and Giles 1979). These projects have generally been problem specific. The tendency to pursue only problems of immediate concern may be a weakness of many present wildlife law enforcement research projects. Wildlife law enforcement research appears to be slowly evolving into an accepted scientific discipline. The process has been and will remain slow. Researchers and other individuals involved in law enforcement can aid this process of evolution by lending support and constructive criticism to the overall process. LITERATURE CITED Beattie, K. H. 1975. Anti-poaching campaigns: a tool of wildlife law enforcement? Proc. Southeast Assoc. Fish and Wildl. Agencies 29: 728-743. tors. 1976a. A descriptive assessment of Mississippi game law cooperaM.S. Thesis, Miss. State Univ., Starkville. 96pp. 1976b. A review of crime load, workload, and manpower standards in wildlife law enforcement. VPI & SU, Southeast. Reg. Wildl. Law Enforcement Res. Proj. 53pp. 1978a. A comparison of hunting satisfaction of Virginia Wildlife and Colorado Outdoors hunter - subscribers. Proc. Southeast Assoc. Fish and Wildl. Agencies 32:738-744. 1978b. An initial bibliography of wildlife VPI & SU,-Southeast. Reg. Wildl. Law Enforcement law enforcement. Res. Proj. 21pp. 1979. A social psychological investigation Virginia sportsmen towards laws and regulations. VPI & SU, Blacksburg. 220pp. of attitudes of Ph.D. Diss., �23 1981a. The influence of game laws and regulations on hunting satisfaction. Wildl. Soc. Bull. 9:229-231. 1981b. Warnings versus citations Wi ldl. Soc. Bull. 9:323-325. in wildlife , and R. H. Giles, Jr. 1979. A survey of ---research needs and current research. Wildl. , ---w~ildlife Agencies , ---~P-roc. , and C. J. Cowles. law enforcement data. 31:698-708. law enforcement. wildlife law enforcement Soc. Bull. 7:185-188. 1977a. An analysis of nationwide Proc~ Southeast Assoc. Fish and Wildl. , and 1977b. Fines in wildlife law enforcement. Southeast Assoc. Fish and-Wildl. Agencies 31:690-697. enforcement. and 1977c. Lack of research in wildlife Wildl. Soc. Bull.-5:170-174. law 1978a. Quasi-experiments, multiple indica----;--- , and tors, and enforcement effectiveness. Proc. West. Assoc. Fish and Wildl. Agencies 58:76-91. , and 1978b. Relative importance of enforcement obje-c-t~i-v-e-s-and seriousness of violations in relation to objectives. Proc. Southeast Assoc. Fish and Wildl. Agencies 32:808-815. , and 1980. Estimating illegal kill of deer. -'7"::"--=7" Pages 65-71 in R. L. Hine and S. Hehls, eds. White-tailed deer population management in the north central states. Proc. 1979 Symp., North Cent. Sect., The Wildl. Soc. , T. A. Pierson, and H. L. Gilliam. 1978c. A readership preference ------s-urvey of Virginia Wildlife subscribers. proc. Southeast Assoc. Fish and Wildl. AgencIes 32:732-737. Brohn, A., and L. J. Korschgen. 1950. Precipitin test -- A useful tool in game-law enforcement. Trans. North Am. Wildl. Conf. 15:467-478. Clark, F. C. 1914. Forensic value of the precipitin test in the enforcement of game laws in California. Univ. Calif. Pub], Pathol. 2:131-138. Cowles, C. J. 1977. Optimum deployment of wildlife law enforcement agents: a problem analysis. VPI & SU, Southeast. Reg. Wildl. Law Enforcement Res. Proj. 31pp. , R. H. Giles, Jr., and K. H. Beattie. 1977. Dyanmic deployment of ------..,wildlife law enforcement manpower -- A decision aid. Proc. Southeast Assoc. Fish and Wildl. Agencies 31:679-689. �24 , K. H. Beattie, and R. H. Giles, Jr. 1978. A survey of methods of ------r-ecording reports of fish and wildlife violations. Fisheries 3:8-11. , ------~ pliance , and estimators. 1979. Limitations of wildlife Wildl. Soc. Bull. 7:188-191. law com- Gay, F. P. 1908. A contribution to the forensic value of the musculoprecipitin test. J. Med. Res. 19:219-224. Giles, R. H., Jr. 1971. Wildlife law enforcement and research needs. Pages 131-133 in R. D. Teague, ed. Manual of wildlife conservation. The Wildl. Soc~ Washington, D.C. 1976a. Alpha-man: A theory of wildlife law violation. Southeast~ Reg. Wildl. Law Enforcement Res. Proj. 12pp. VPI & SU, 1976b. The Southeastern wildlife law enforcement research project: -progress and perspectives. Proc. Southeast Assoc. Fish and Wildl. Agencies 30:680-684. 1978. Wildlife Wildlife management. law enforcement. Pages 343-377 in R. H. Giles, W. H. Freeman and Co., San Francisco, Calif. , M. Kaminsky, and J. ------research -- The context McLaughlin. 1971. Wildlife law enforcement and the needs. Proc. Southeast Assoc. Fish and Game Comm. 25:677-687. Glover, R. L., and L. J. Korschgen. 1980. Evaluation of two methods of preparing antisera for precipitin tests. Wildl. Soc. Bull. 8:118-122. Hussain, N. G. 1977. An evaluation of the Michigan wildlife law enforcement effort. Proc. West. Assoc. State Game and Fish Comm. 57:35-66. Johnson, B. C., L. Maguire, and D. Anderson. 1980. Determining time of death in mule deer by using potassium levels in the vitreous humor. Wildl. Soc. Bull. 8:249-252. Kaminsky, M. A. 1974. Analysis of the spatial and temporal occurrence of deer spotlighting violations in Virginia. M.S. Thesis, VPI & SU, Blacksburg. 170pp. McCormick, J. B. 1968. A procedure for evaluating the effectiveness of wildlife law enforcement. Proc. West. Assoc. State Game and Fish Comm. 48:626-639 . . 1970. An evaluation of wildlife law enforcement -----W~est. Assoc. State Game and Fish Comm. 50:517-540. effort. Proc. Mikel, H. C. 1981~. Application of deterrents to reduce fishing viola~ tions. Proc. West. Assoc. State Fish and Wildl. Comm. 62:246-249. 1981b. Illegal harvest of warmwater game fish. Game and fish Fed. Aid Proj. F-51-R. 14pp. New Mexico Dep. Morse, W. B. 1957. The conservation officer, a personnel challenge. Proc. West. Assoc. State Game and Fish Comm. 37:63-69. �25 1958. Western wildlife law violators. Game and Fish Comm. 38:292-295. Proc. West. Assoc. State 1963. The new look in wildlife law enforcement. Assoc. State Game and Fish Comm. 43:287-294. Proc. West. 1968. Wildlife law enforcement - 1968. Game and Fish Comm. 48:683-694. Proc. West. Assoc. State 1972. Wildlife law enforcement - 1972. Game and Fish Comm. 52:118-137. Proc. West. Assoc. State 1973. Law enforcement - one-third of the triangle. Bull. 1:39-44. Wildl. Soc. 1976. Wildlife law enforcement - 1976. Game and Fish Comm. 56:127-145. Proc. West. Assoc. State 1980. Wildlife law enforcement - 1980. Fish and Wildlife Comm. 60:162-180. Proc. West. Assoc. State 1982. Sidearms policy and assaults on wildlife law enforcement officers. Ann. Meeting, Midwest Fish and Game Law Enf. Off. Assoc. Steamboat Springs, Colo. 6pp. Oates, D. W., C. W. Brown, and D. L. Weigel. fication of selected birds and mammals. Comm., Fed. Aid Proj. W-38-R. 91pp. 1974. Blood and tissue identiPart I. Nebr. Game and Parks Purol, D. A. 1982a. Field estimating the legality of harvested deer. Mich. Dep. Nat. Resour., Law Enf. Div. Rep. 4. 43pp. 1982b. Recreational trespass enforcement in southern Michigan. Mich. Dep~ Nat. Resour., Law Enf. Div. Rep. 2. 43pp. , and T. J. Fournier. ------as a tool of enforcement deer hunting seasons. , and ----~~ license 1979. A review of the arrest: contact ratio using prosecution data from the 1977 Michigan Mich. Dep. Nat. Resour;, Law Enf. Div. 1. 25pp. M. M. Zambetis. 1982. Enforcement of Michigan1s hunting revocations. Mich. Dep. Nat. Resour., Law Enf. Div. Rep. 3. 19pp. Pursley, D. 1977a. Illegal big game harvest during closed season. Mexico Dep. Game and Fish, Santa Fe. 5pp. New 1977b. Reducing illegal harvest of trout in streams. New Mexico Dep. Game and Fish, Santa Fe, Fed. Aid Proj. F-22-R-18. 27pp. �26 Ritter, A. F. 1975. Objectives and performance criteria for state wildlife law enforcement agencies. M.S. Thesis. VPI & SU, Blacksburg. 198pp. Vi1kitis, J. R. 1968. Characteristics of big game violators and extent of their activity in Idaho. M.S. Thesis. Univ. of Idaho, Moscow. 202pp. Vaught, J. R. 1975. Wildlife law enforcement research in New Mexico. Proc. West. Assoc. State Game and Fish Comm. 55:152-162. , ---and and F. L. Turner. 1975. Wildlife officer district evaluation wildlife officer district law enforcement survey. New Mexico Dep. Game and Fish, Fed. Aid Proj. FW-15-R. 37pp. Wi Ison, M. (ed.). enforcement. Div. 171pp. 1977. Bibliography Alberta Recreation, of forensic science in wildlife law Parks and Wildl., Fish and Wildl. Woolf, A., and J. L. Roseberry. No date. Forensic science studies: application of techniques to estimate the post-mortem interval of white-tai led deer in III inois. Final rep., Ill. Dep. Conse rv. , Div. Law Enf. 10pp. Prepared by �27 APPENDIX ACTIVITY Activity Code 10 DEFINITIONS Activity AND CODES Name Wildlife Inventory Wildlife Regulation Recommendation Public Relations Hunter Safety Research Game Damage and Animal Control Propagating and Distributing Wildlife Law Enforcement Habitat Development and Maintenance Publ ic Facilities Development and Maintenance Division Equipment and Facil ities Maintenance Environmental Protection Resource Reconnaissance Planning and Budgeting Real Estate Administration and Clerical Services In-Service Training Leave 14 18 22 26 30 42 46 50 54 58 62 66 70 74 7 10. A Wildl ife Inventory Any work intended to provide information about the number, composition or distribution of wildlife, wildlife habitat, or publ ic use of wild1 i fe. 14. Wildlife Regulation Recommendation Any part of the process of translating inventory recommendations for season regulations. 18. information into Publ ic Relations Any work done to inform, educate or assist the publ ic with the intent of increasing publ ic understanding or use of a wildlife resource. 22. Hunter Safety Any work performed with the intention of increasing fied hunter safety students. 26. Research Any work necessary project. 30. the number of certi- to accomplish the objectives of an approved research Game Damage and Animal Control Any work intended to control animals which are a nuisance or a danger to people or to properties and any investigation or processing of wildl ife damage claims. �28 Appendix A (continued) 34. Propagating and Distributing Wildlife Any work intended to supplement or introduce wildl ife by releasing captive animals into the wild. This category includes the capturing and caring for animals which are to be released for this purpose. 38. Law Enforcement Any work action. 42. Habitat Any work 46. intended to detect violations Development and bring the violators to legal and Maintenance intended to enhance Publ ic Facil ities Development the abil ity of habitat to support wildlife. and Maintenance Any work intended to result in the construction or upkeep of facil ities designed to provide comfort or convenience for wildlife users. 50. Division Equipment and Facilities Maintenance Any work intended to result in the construction or upkeep of facil ities or equipment used by Division personnel in their daily activities. 54. Environmental Protection Any work intended to minimize the adverse effects of land or water development on wildlife or their habitats. 58. Resource Reconnaissance Any work intended to famil iarize an individual with his geographic of responsibil ity. 62. Planning and Budgeting Any work intended to influence the final form of a Division Plan, Operation Plan or budget document. 66. area Strategic Real Estate Any work intended to result in the transfer of some degree of surface control of land to the Division. 70. Administration and Clerical Services Any work where the primary intention is to increase the efficiency or effectiveness of other Division personnel. Work in this category will result in some form of written or oral communication. Task force assignments are included in this category. 74. In-Service Tra-ining Any formally scheduled Division employees. effort to improve the knowledge or skills of �29 Appendix A (continued) 78. Leave Any regularly scheduled working day when an employee is not on duty, except for holidays, weekends and compensatory time, should be recorded in this category. This includes all paid leave (annual, sick, funeral, etc.). Hours should be figured at the rate of 8 hours per working day. �30 APPENDIX DISTRICT WILDLIFE (Statutory Title: B MANAGER Wildl ife Conservation Officer) NATURE OF WORK This is a multiple range class with a broad range of duties and responsibilities from the entry trainee through the fully operating journeyman level District Wildl ife Manager. This position requires the exercise of independent judgment, professional skills and abilities and technical expertise while engaged in wildlife management, including law enforcement, wildlife inventories, population analysis, habitat maintenance and improvement, environments impact assessments, information and education, and a variety of additional wildlife-oriented responsibilities. RANGE A This is the beginning professional trainee level with formalized on-the-job classroom and field training under continuous supervision and control. The employee becomes knowledgeable of the Division's operation, appl ication of the State's laws, rules, and regulations, and techniques of wildlife resource management. RANGE B This range is a continuation of the developmental concept. This range requires less supervision and the individual is assigned a district or area of responsibility and is given more latitude to gain confidence and experience necessary to perform at the journeyman level. Although assignments given at this level are broader and more in-depth, the employee is still in a developmental stage. RANGE C This range identifies the fully functioning journeyman, full scope of assignment, usually non-supervisory level. A person of this level has full responsibil ity for all wildlife and related management practices and enforcement of wildl ife laws in an assigned district. This level receives general supervision from an area supervisor but the district programs and how well they are managed and how effective they are is essentially the employee responsibil ity. SOME EXAMPLES OF WORK Initiates and conducts census and interprets other data. Initiates and performs complex wildlife wildlife research studies. management studies; assists in complex Initiates and assists in development of game management areas related to share crop, grazing leases and contract farming arrangements, in addition to controlling and developing state-owned lands. �Appendix 31 B (continued) DISTRICT WILDLIFE Page 2. MANAGER (Con't) Develops district objectives and plans to achieve state goals and prepares budget estimates to achieve set goals. Develops technical wildlife material and writes articles for departmental and public use. Attends and participates in professional conferences and meetings and reviews literature to improve knowledge in the field of wildlife management. Performs liaison, coordinative dictions and other entitites. and contact work with various agencies, juris- Works with radio, television and press media, sportsmen, public schools, recreational ists and rancher organizations and other groups in developing and promoting understanding and support of department policies and wildlife management objectives. Develops and supervises hunter safety programs within an assigned district. Conducts aerial and ground census trends, trappings, tagging, and banding studies; introduces and transplants game, non-game, fur bearers, birds and fish for experimental or management purposes and writes summary and conclusions. Conducts age-growth, length-weight, migration, distribution, food and life history studies; measure depth and surface areas of water, water flow, and the basic chemical composition of water. Conducts ecological surveys; investigates and assists in determining feasibil ity of planting of warm and cold water fish in streams, lakes and rivers in the state. Participates in experiments with various feeds and rations of feeds, either in a laboratory or in the field. Responsible for initiating and planning of exclosures ranges to determine util ization of forage. and enclosures on game Performs field reconnaissance and planning for wildl ife habitat improvements; gathers information for mapping seasonal ranges and routes of migration of various wildlife. Assists in the planning, program. preparation and planting of the annual fish planting Assists in checking stations operation of various game species. for tabulation of wildlife harvest Investigates wildlife damage claims, plans and initiates actions to alleviate the damage and makes recommendations on possible restitution; coordinates distribution on preventative materials within an assigned district. Investigates non-damage wildlife harassment and initiates actions on how such harassment can be alleviated. �32 Appendix B (continued) DISTRICT WILDLIFE Page 3. MANAGER (Con't) Coordinates with staff specialists district programs and projects. Participates Conservation professional and other conservation officers in the training of and supervIsion and evaluation Officer A's. Supervises subordinate professional employees. Acts for and represents the Area Wildlife Supervisor in of Wildlife and non- upon assignment. Initiates and conducts investigation of violations involving Division of Wildlife and Division of Parks and Outdoor Recreation laws and regulations, including federal statutes. Prosecutes said violator in courts of law. Initiate recommendation for state statute sion" regulation improvement. improvement and "Wildlife Commis- Responsible for the implementation and coordination within a given county or counties of the land and water use law. (Examples, HB 1041 and SB 97). Assists or initiates acquisition district. of wildlife habitat within an assigned Initiates and performs environmental analysis, participates in writing completion of environmental impact statements and Environmental Assessment Reports, i.e., highway projects, water development, coal and oil shale, land reclamation, etc. Must review, assign and investigate all land use on private lands, responding in the preparation of technically written reports and effectively express orally, the interpretation of these technical reports to County Planning and Zoning Commissioners, County Commissioners, and Developers. Selects by rivers, tributaries and natural lakes to determine scientifically, minimum stream flows and lake levels to legally avoid dewatering by other users and to protect the natural environment and aquatic ecology. Generates publ ic support and is contributing witness to various water boards and water courts. Performs related duties as assigned or required. KNOWLEDGES, SKILLS AND ABILITIES Knowledge of state, federal, wildlife and outdoor modern law enforcement procedures and techniques. Knowledge of theories, principles of land development practices. and techniques Knowledge of stream and lake biological procedures. Knowledge surveying of aquatic ecology and invertebrate recreational laws and of wildlife management and and sampling methods and zoology. �Appendix B (continued) DISTRICT Page 4. WILDLIFE 33 MANAGER Knowledge of mibrobiology, to wildl ife management. (Con't) and organic and inorganic Knowledge species. of the habits and ecology of wildlife Knowledge of related federal, Skill of firearms, outdoorsmanship in the use of laboratory Skill and ability and happenings. including state, and local cooperative Knowledge of the relationships of wildlife resource conservation concepts. Knowledge chemistry in relating conservation and all outdoor as they relate game and non-game programs. to total natural recreation functions. and field equipment. in-depth observations of wildlife incidents Abil ity to observe and classify wildlife from aircraft, fixed-wing helicopters and also from the ground under varying conditions. Abil ity to complete complex wildlife management and biology assignments. Abil ity to analyze data and apply relevant wildlife the solution of problems. Abil ity to express oneself clearly and concisely, Abil ity to establish and maintain effective courts, district attorneys, law enforcement Abil ity to enforce wildlife rights of others. biologic principles to both orally and in writing. working relationships with officials, and the publ ic. laws firmly and tactfully, Abil ity to make independent decisions determined course of action. and and act quickly with respect for the and decisively on the Ability to create public awareness of and involvement in wildlife commission's programs and objectives and to provide leadership to the publ ic in this activity. Abil ity to work under stress and strain for prolonged Ability to util ize and maintain MINIMUM PREPARATION Education and extended periods. issued equipment. FOR WORK and Experience Graduation from an accredited college or university with a Bachelor's degree in Wildl ife Biology, Biology or related field. �)If Appendix B (continued) DISTRICT \~ILDLIFE MANAGER Page 5. Necessary (Con't) Special Requirement Valid Colorado Driver's License at time of appointment. NOTE: Employees are not adjusted to the "B" range until they have successfully completed a one-year training program and to the "c" range until they have completed one year at the "B" range with satisfactory performance or above, unless otherwise approved by the Division of Wildlife Director. �35 APPENDIX AREA WILDLIFE C SUPERVISOR NATURE OF WORK This is professional supervisory and administrative life management and lands program. work in the state wild- Under general direction of an assistant regional manager, plans, organizes, supervises, coordinates, and participates in wildlife management and enforcement activities, environmental impact programs and budget preparation for an assigned regional geographic area, or performs a wide variety of staff coordination functions in a regional office such as regional law enforcement coordination or similar staff positions within a wildlife region. Area wildlife supervisors assigned to an area within a wi1d1 ife region must supervise three or more District Wildlife Managers and may also supervise Wildlife Technicians. Staff coordinator positions in a region must have regionwide responsibility for planning, implementing, evaluating, and coordinating a major wi1d1 ife activity such as law enforcement or a comparable wildlife activity. SOME EXAMPLES OF WORK Plans, organizes, supervises, coordinates and participates in a wide variety of wi1d1 ife management and enforcement activities, within an assigned geographic area. Conso1 idates and evaluates findings and reports of subordinate personnel, and prepares management and enforcement and other recommendations for the area supervised. Compiles a variety of reports on wildlife population and game land and environment conditions; develops program changes and management plans for present and future needs. Plans and inspects construction and maintenance projects, wi1d1 ife improvement projects, and usage of state game lands and private lands under state agreement or contract. Reviews reports and performs investigations in espec1a11y complex or unusual situations or 1itigation resulting from arrests, appeals, or hunting accidents. Promotes pub1 ic relations by preparing new releases, giving sl ide lectures, appearing on radio and television, preparing exhibits, and developing information for District Wildlife Managers. Estab1 ishes and maintains working relationships with federal and local governmental agencies and other state agencies and provides assistance to other divisional programs on request. �36 Appendix C (continued) AREA WILDLIFE Page 2. Coordinates SUPERVISOR (Con't) and supervises the area's hunter safety programs and schools. Consolidates area reports and recommendations and assists the regional manager in making evaluations, correlations and regional consol idations for consideration and action at the division and higher level. Coordinates the area's wildlife management and enforcement activities, publ ic use of division properties, and area environmental issues and projects. Assists with budget preparation, performs business management prepares a variety of reports and correspondence. duties and Coordinates the assignment of regional staff personnel, both professional and support, within and between areas and on special assignments. May serve as a regional staff coordinator for specialized programs such as regional law enforcement coordinator: Plans, implements, monitors, and coordinates regional law enforcement activities; conducts and/or supervises special investigations of major and complex cases of suspected violation of wildl ife statutes; plans, conducts, evaluates, and coordinates law enforcement training programs for regional staff; conducts public education and information programs; serves as regional hearings officer for license suspension hearings; performs related staff coordination activities. Performs related work as assigned or required. KNOWLEDGES, SKILLS AND ABILITIES Thorough knowledge of state wildlife laws and modern law enforcement procedures; the impact on the environment of natural resources development projects and the incidence of and problems inherent in publ ic use of division properties. Thorough knowledge control principles of wildlife management, and practices. land development and environmental Considerable knowledge of federal, state and local cooperative related to wildlife management. Considerable knowledge of the practices relations program. and objectives programs of an effective Knowledge of modern administrative practices and procedures, including budgeting, supervision, management, planning and organization. Skill in communicating effectively, both orally and in writing. publ ic �Appendix AREA WILDLIFE Page 3. Ability changes 37 C (continued) SUPERVISOR (Con It) to prepare a variety of reports and to keep personnel and additions to procedures and programs. informed of Skill and abil ity in establishing and maintaining effective working relationships with federal, state, and local officials, fellow employees and the publ ic. MINIMUM Education PREPARATION FOR WORK and Experience Graduation from an accredited college or university with an appropriate Bachelorls degree and four years of experience as or at the level of a District Wildlife Manager or above. The required experience may be in wildlife management or wildlife research and must have included enforcement and administrative responsibilities. Enforcement being defined as enforcing appl icable wildlife statutes. Administrative responsibilities must have included developing and implementing work plans, preparing budget requests, providing input for operating and policy decisions. �38 APPENDIX 1982-83 SUMMARY OF D STATE/PROVINCIAL RESEARCH/EVALUATION IN WILDLIFE LAW EFFORTS ENFORCEMENT by john Colorado F. Smel tzer Division of \-'ildlife Law Enforc~ment Research Wi ldl ife Research Center 317 West P~ospect Road Fort Coll ins, Colorado 80526 March 1983 �Appendix D (continued) 39 TABLE PART I OF CONTENTS State/Provincial Summary of Research/ Evaluation Efforts - a narrative summary PART II ..... Responses to survey question: What are the most serious problems facing wildlife law enforcement today? • . . . . . . . . . . . . 7 PART III .... Responses to survey question: What are the needs for wildlife law enforcement research? 9 PART IV Names and addresses of contact persons throughout the U.S. and Canada who are most familiar with their attempts to evaluate or research wildlife law enforcement problems ...•........ 10 �40 Append ix D (cont inued) Part l I STATE/PROVINCIAL SUMMARY OF RESEARCH/EVALUATION EFFORTS The following is a narrative summary of the information provided in response to the Colorado Wildlife Law Enforcement Research/Evaluation Survey. is not intended as a comprehensive summary of all but simply as a source of information to responding agencies. It is based only on the information supplied. Alabama: a.) Special Task Force development b.) Investigating possibilities of covert operations Alaska: a.) Computerization Arizona: a.) 10-year research plan being developed which will include wildl ife law enforcement research b.) 1 person assigned to coordinate Wildlife L.E. research plan Arkansas: Alberta: No research/evaluation data, violations/contact efforts reported a.) Hunter attitudes toward wildlife laws and wildlife officers in Alberta b.) Factors associated with wildlife law violations in Alberta c.) The use of aircraft in wildl ife law enforcement d.) Compendium of Alberta Fish and Wildlife District Work Analysis Status 1980-81 e.) 1982 -- Helicopter patrol program Nov. 1-30, 1982 f.) A selected annotated bibl iography of literature on wildlife enforcement g.) Permanent personnel are assigned to Wildl ife L.E. research/evaluation Cal ifornia: Colorado: of violation It No report a.)Computerized monthly and annual violation, court case, dismissed, and fines report b.) Wildlife Law Enforcement Plan -- 1981-84. Developed and being implemented. c.) Has wi ldl ife law enforcement researcher d.) Draft study plans for the following: Illegal license purchase evaluation -- Development of measurable enforcement objectives -- Public attitude survey e.) Developing a liS-yr. Operations Plan forWildlife Law Enforcement Res ee r ch" f.) Using select college information. students to quantify historic law enforcement �Appendix 41 D (con t lnued ) Connecticut: No research/evaluation activities reported Delaware: No research/evaluation activities reported Florida: a.) b.) c.) d.) e.) f.) g.) h.) i.) j.) k.) 1.) m.) A practical field method for blood and tissue identification Intermediate non-lethal weapons for Florida wildlife officers Decriminalization and recodification of Florida's wildlife code A three-year plan to curtail the commercialization of certain rare Florida wildlife What is enough wildlife law enforcement The scope of the wildlife trade in the United States Development of a statewide citizen crime patrol reporting project (Wildlife alert) and impact New approaches toward wildlife crime patrol General orders manual for wildl ife law enforcement Wildlife officer productivity computer printouts Spec iali zed so 1ut ions to a number of wi 1dl ife 1aw enforcement prob 1ems Undercover investigations program Accountabil ity-wildl ife enforcement goal for the 1980s Georg ia: No report Hawai i: No research/evaluation activities reported. Do have in-house review measures which were not elaborated on. Idaho: a.) b.) c.) d.) 111 ino is: a.) b.) c.) d.) e.) f.) g.) h.) i.) Indiana: Vilkitis poaching study Evaluation of purchase of resident licenses by non-residents Computerized violation, license,. and suspension files Currently adding one person to staff to be responsible for wildlife law enforcement research projects Time of death studies - deer Blood identification test kits for.deer Sonar research Lead and copper test kits Identification of illegal furs using scanning electron microscope (recommend discontinue) Long distance hearing devices Signal transmitters on illegal devices and commercial wildlife Muscle tissue/parasite determination Post-Mortem vitreous humor potassium levels a.) Developed an extensive Standard Operating b.) Coded violations c.) Coded activities by category Procedures Manual �42 Appendix D (~ontinued) Iowa: a.) Evaluation of wildlife enforcement officer productivity b.) Use of aircraft to detect spotl ighters Kansas: a.) Evaluate effectiveness of uniformed vs. non-uniformed personnel b.) Evaluate effectiveness of aircraft in aiding detection of spotlighters c.) Computerized boating registration files d.) Computerized activity analysis Kentucky: No report Louisiana: No research/evaluation Maine: a.) b.) c.) d.) Manitoba: Rec~ntly Maryland: No research/evaluation Massachusetts: efforts reported Computerized officer daily activity reports Computerized prosecution reports Computerized complaint file Have looked at illegal purchase of resident licenses by non-residents replicated Vilkitis poaching study activities -- no result released yet reported No report Michigan: a.) An evaluation of the Michigan wildlife effort b.) Work on compliance estimation c.) Estimation of poaching levels law enforcement Minnesota: No report Mississippi: a.) Officer performance and operational cost analysis b.) Computerized citation files -- coded violation forms by: species, county, violation �Appendix D (continued) Hissouri: a.) b.) c.) d.) e.) Montana: No report Nebraska: No research activities reported, doing substantial forensic work Nevada: No research/evaluation activities reported New Jersey: No research/evaluation activities reported New Mexico: a.) b.) c.) d.) e.) f.) g.) h.) New York: 43 Has wildlife law enforcement researcher Deer poaching and poacher study Lead detection analysis Meats and blood identification Computerization of arrest reports and agents monthly activity reports f.) Potassium levels in vJtreous humor for time of death analysis - deer g.) Developed operation game thief program similar to New Mexico h.) Covert operations however, Dave Oates has been Illegal harvest evaluation of warmwater game fish Illegal harvest of big game during closed season Reducing illegal harvest of trout in streams Wildl ife officer district evaluation andwildl ife law enforcement Wildlife Law Enforcement research in New Mexico Implication of illegal harvest on deer management Remote sensing equipment and radio telemetry testing Evaluatjon of illegal purchase of resident licenses by non-residents a.) Computerization of violation information b.) Development of activity codes c.) Establishing objectives North Carol ina: No research/evaluation efforts North Dakota: No research/evaluation efforts reported Ohio: No research/evaluation activities Oklahoma: No report Ontario: reported reported a.) Development of regional and district law enforcement plans b.) Development of a uniform officer reporting system and standard incident reporting system c.) Development of data base information on contacts, warnings, charges, time analysis, miles driven, etc. d.) Draft proposa I -- IIImprovi ng Enforcement Eff ic iency" -- by Outdoor Recreation Group. 1981-03-06. e.) Report on the "Role and Responsibil ities of the Conservation Officer" surv �44 Append ix D (cont inued ) Oregon: No report Pennsylvania: No research/evaluation activities reported Rhode Island: No report Quebec: a.} Identificationofwi1d bird blood, feather and bone b.} Identification of deer and moose meat c.} Computer ana 1ys is of wi 1d 1ife 1aw enforcement effort -- poss ib 1e report to be given to mid-west L.E. meeting, Springfield, I11 ino is in June 1983 Saskatchewan: Canadian Wildlife Saskatchewan: Tourism and Renewable a.) Computerized b.} Computerized analysis South Carolina: Service - NO.research activities reported Resources prosecution report work management report -- time/activity No current research activities report. Have been involved with research activities conducted through Virginia Polytechnic Institute. South Dakota: a.} Considering electronic surveillance devices for back road use b.} Documented violations to estab1 ish projected/actual losses, and periods of highest illegal activity c.} Use of news media to influence public attitudes towards poaching d.} Covert operations Tennessee: a.) Study completed on "High visibility versus low visibility vehicles in wi 1d1 ife law enforcement" b.) Participated in VPI studies for 3 years c.} No additional research activities currently planned Texas: a.) Development of Statewide Manning Standards and Assignments Program. Deployment based on population density, type and area of patrol (land and water), local hunting/fishing activity b.) Development of violation trend charts and tables c.} Evaluation of reorganization in 1975 d.} Development of field contact survey program (FY-78) e.} Development of "Wi1d1 ife Law Enforcement Division Operational P1an" FY 1983. This is a 58-page plan including: work load summaries, 1982 accomplishments and 1983 objectives for numerous wi1d1 ife law enforcement activities. Utah: a.} b.) c.} d.} e.) Vermont K-9 use in wildlife law enforcement Effectiveness of two-man vs. one-man patrol Effectiveness of various patrol techniques Effectiveness of preventative law enforcement Time of death studies No report methods �Appendix 45 D (continued) Virginia: Virginia: a.} Have been involved with research activities b.) Some limited covert operations at VPI Virginia Polytechnic Institute and State University (V.P.I. S.U.) Studies -- supervised by Dr. Robert H. Giles, Jr. a.} Objectives and performance criteria for state wi1d1 ife law enforcement agencies b.} Analysis of the spatial and temporal occurrence of deer spotlighting violations in Virginia c.} Optimum deployment of wi1d1 ife law enforcement agents: a problem analysis d.) A review and appraisal of crime load, workload, and manpower standards in wildlife law enforcement e.) An initial bibliography of wi1d1 ife law enforcement f.) Estimating illegal kill of deer g.) The influence of game laws and regulations on hunting sat isfact ion h.) Warnings vs. citations in wildlife law enforcement i.) A survey of wi1d1 ife law enforcement research needs and current research j.) Anti-poaching compaigns -- a tool of wildlife law enforcement? k.) Quasi - experiments, multiple indicators, and enforcement effectiveness 1.) Fines in wildlife law enforcement m.) An analysis of nationwide wi1d1 ife law enforcement data n.} Relative importance of enforcement objectives and seriousness of violations in relation to objectives 0.) Wi1d1 ife law enforcement research - the context and the needs p.) Alpha - man: A theory of wi1d1 ife law violation q.) Wildlife law enforcement r.) Limitations of wildlife law enforcement compliance estimators s.} Dynamic deployment of wildlife law enforcement manpower a decision aid Washington: West Virginia: a.} Regional workload assessment program b.) Development of data base information a.) No research activities reported b.) Do have standardized complaint forms Wisconsin: No report Wyoming: a. ) Computerized b. ) c. ) d. ) e. } f. ) arrest records by violations reported by citizens, observed by officers, and by arrests from reports or observations. Developed case file referral system to keep track of current investigations and time spent per case Have a stop-poaching program District violation summaries (1978) A study of user attitudes and violations Has personnel assigned to wildlife law enforcement research/ eva 1uat ion �46 Appendix PART II D (~ontinued) a What are the most Table 1. Survey responses to the question : serious problems facing wildlife law enforcement today? Response 1. 2. 3. 4. 5. 6.' 7. 8. 9. # of responses Funding problems - lack of funds Poaching, commercial poaching, marketing Lack of personnel Public attitudes More people - increasing number of users Inabil ity to evaluate enforcement effort Program administration problems Civil suits - lawsuits against officers Spotlighting aTwenty-six Other responses each) . \ (26) States/Provinces suggested 12 11 9 5 4 3 2 2 2 responded these additional to this question. problems. (Mentioned only once Habitat loss Inadequate detection rate Need to educate publ ic Low salaries Lack of equipment Native hunting and fishing privileges Trespass problems Increasing wildlife law enforcement responsibilities Some examples of complete responses are: IIWe are seeing more lawsuits against our officers his/her duties.1I IIWe need to educate the hunting and fishing public and respons lb ll ltv ;!' in the performance of in ethical conduct IIHow to do more, with less!1I IIOur biggest problem is poor publ ic attitude and the inabil ity to prove the worth of law enforcement activities in the eyes of the public and the l eq i s la tur e ;" IIWe are finding more sophisticated II attitude II lack of criteria law violators.1I of the publ ic -- the silent majority.1I for officer evaluation.1I IICommercialization of wildl ife resources, is our most serious problem." fish, game and non-game �47 Appendix D (continued) "lack of money and personnel to handle increasingly sophisticated violators." "More people, that's our problem." "Program administration, including budgeting problems, collective bargaining issues, compo time accrual, enforcement coverage, and productivity." "We are seeing increased enforcement drugs, theft, recreational vehicles involvement in peripheral to name a few." "Trespassing on posted lands ... resulting relationships." II ••• a serious misunderstanding for enforcing those laws ;!' areas; in bad landowner/hunter of the laws and the agency responsible �50 Append ix D (con t lnued] KENTUCKY: Steve Yontz, Director Division of Law Enforcement Dept. of Fish & Wildl. Res. #1 Game Farm Road Frankfort, Kentucky 40601 (502) 564-3400 MINNESOTA: Fredean Hammer, Director Division of Enforcement Dept. of Natural Resources 300 Centennial Building 658 Cedar Street St. Paul, Minnesota 55155 LOUISIANA: Susan K. Brittain Staff Development Specialist Enforcement Division Louisiana Dept. of Wildlife & Fisheries P. O. Box 15570 Baton Rouge, Louisiana 70895 (504) 568- 5667 MISSISSIPPI: Philip J. Strong, Chief Law Enforcement Mississippi Dept. of Wildl ife Cons. Southport Mall, P. O. Box 451 Jackson, Mississippi 39205 (601) 961-5300 MAINE: John F. Marsh, Chief Warden Dept. Inland Fisheries & Wi ldl. 284 State Street State House Station 41 Augusta, Maine. 04333 (207) 289-2766 Detective Richard Hennessey Augusta Regional Headquarters Dept. Inland Fisheries & Wi ldl. 8 Federal Street Augusta, Maine 04330 (207) 289-2175 MANITOBA: MARYLAND: MASSACHUSETTS: S. A. "Bud" Mcivor, Chief Field Services & Enforcement Manitoba Dept. of Natural Res. 1495 St. James Street Winnipeg, Manitoba CANADA R3H OW9 (.204) 786-9132 Ann Williams (Computer (601) 961-5313 John Given (Computer (601) 961-5331 Program) MISSOURI: Ron L. Glover Protection Research Specialist Missouri Dept. of Conservation Fish & Wildlife Research Center 1110 College Avenue Columbia, Missouri 65201 (314) 449-3761 MONTANA: Erwin J. Kent, Administrator La,,!Enforcemen t Dept. of Fish, Wildlife & Parks 11120 East Sixth Helena, Montana 59601 (406) 449-2452 NEBRASKA: Donald C. Schaepler, Chief Law Enforcement Division Nebraska Game & Parks Commission 2200 North 33rd Street P. O. Box 30370 Lincol~, Nebraska 68503 (402) 464-0641 Jack T. Taylor, Deputy Supt. Dept. of Natural Resources Natural Resources Pol ice Tawes State Office Building Annapol is, Maryland 21401 (301) 269-2248 James Jesseau Supervisor of Enforcement Dept. of Fisheries, Wi Idl ife & Recreational Vehicles 100 Cambridge Street Boston, Massachusetts 02202 (617) 727-3900 Program) Dave Oates, Forensic Research Nebraska Game & Parks Commission 2200 North 33rd Street P. O. Box 30370 Lincoln, Nebraska 68503 (204) 464-0641 NEVADA: Darrel D. Harold, Acting Chief Division of Law Enforcement Nevada Department of Wildlife 1100 Val ley Road, P. O. Box 10678 Reno, Nevada 89520 (702) 784-6214 �Appendix NEW HM'lPSHIRE: Major Mason S. Butterfield Chief of Law EnforCement New Hampshire Fish & Game Dept. Box 2003, 34 Bridge Street Concord, New Hampshire 03301 (603) 271-3421 ~JEW Hudson G. Amory District Conservation Officer Bureau of Law Enforcement CN 400 Trenton, New Jersey 08625 (609) 292-2965 JERSEY: 51 D (~ontinued) NEH MEXICO: Harry Mikels Wildlife Law Enforcement Researcher H.M. Game i Fish Department Vi llagra Bui lding Santa Fe, New Mexico 87503 (505) 827-2550 NE~I YORK: George Firth, Asst. Director Division of Law Enforcement H.Y. State Dept. of Environmental 50 Wol f Road Albany, New York 12333 (518) 457-5680 NORTH CAROLINA: Gene H. Abernethy, Chief Division of Enforcement N.C. Wildlife Resources Commission Archdale Building 512 H. Salisbury Street Raleigh, North Carol ina 27611 (919) 733-7191 NORTII DAKOTA: Harold II. Spitzer, Chief Lal~ En f orccmen t D ivis ion State Game & Fish Department 2121 Lovett Avenue Bismarck, tlorth Dakota 58505 (701) 22/1-2180 OHIO: Richard Francis Game Protection Manager Ohio Department of Nat. Resources Division of Wildl ife Fountain Square Columbus, Ohio 43224 (614) 466-5854 OKLAHOMA: Kenneth Van lIoozer, Chief Law En forcemen t Dept. of Wildl ife Conservation 1801 Horth Lincoln P.O. Box 531165 Oklahoma City, Oklahoma 73152 (405) 521-3719 ONTARIO: W. Da Ie Ga rt Iey Provincial Enforcement Spec. Room 2342, Whitney Block Queen's Park Toronto, Ontario CANADA M7A lW3 (416) 965-5661 OREGON: L. R. Hyder Oregon State Police Fish & Game Enforcement Salem, Oregon 97301 PENNSYLVANIA: Edward W. Manhart, Chief Law Enforcement Division Pennsylvania Fish Commission P.O. Box 1673 Harrisburg, Pennsylvania 17120 (717) 787-2350 G. D. Kirkpatrick, Chief Division of Law Enforcement Pennsylvania Game Commission P. O. Box 1567 Harrisburg, Pennsylva~ia 17120 (717) 787-5743 Cons. ' QUEBEC: Jos-A. Saint-Pierre Canadian Wildl ife- Service P. O. Box 10100 Tour Champlain, 4th Floor Ste-Foy, Quebec CANADA GIV 4H5 (418) 694- 3914 RHODE ISLAND: Stephen P. Fongere, Chief Division of Law Enforcement Dept. of Environmental Mgt. 83 Park Street ~~ovidence, Rhode Island 02903 (401) 277-2284 SASKATCHEWAN: Garry Bogdon Enforcement Coordinator Western & Northern Region Canadian Wildl ife Service 115 Perimeter Road Saskatoon, Saskatchewan CANADA S7N ox4 (306) 665-4087 Barry F. Tether, Director Field Services Saska t chewan Tour ism & Renewable Resources 3211 Albert Street Regina, Saskatchewan CANADA S4S 5W6 (306) 565-2323 �52 Appendix SOUTH CAROl! NA: SOUTH DAKOTA: TENNESSEE: D (continued) Pat Ryan, Director Division of law Enforcement & Boating South Carol ina Wildlife & Marine Resources Department Rembert C. Dennis Building P. O. Box 167 Columbia, South Carol ina 29202 (803) 758-00~2 Ronald P. Catlin law Enforcement Staff Specialist S.D. Game, Fish & Parks Sigurd Anderson Building ~~5 East Capitol Pierre, South Dakota 57501 (605) 773-3381 A. J. Gulley, Jr. Tennessee Wildlife Resources Agency 216 E. Penfield Street Crossville, Tennessee 38555 (800) 262-6704 TEXAS: Chester l. Burdett law Enforcement Director Texas Parks & Wildl ife Department ~200 Smith School Road Austin, Texas 787~4 (~15) ~79-~8C8, ext. ~845 UTAH: Bruce Johnson Enforcement Specialist Utah Division of Wildl ife Respurces )596 West North Te~ple Salt Lake City, Utah 84116 (801) 533-9333, ext. 217 VERr10NT: Roger Whitcomb Chief Ga~e Warden Fish & Game Department Montpel ier, Vermont 05602 (802) 823-3371 VIRGINIA: Col. John H. McLaughlin, Chief law Enforcement Division Virginia Game & Inland Fisheries Box 11104 Richmond, Virginia 23230 (804) 257-1000 Dr. Robert H. Giles, Professor Wildl ife Management V.P.I. s S.U. Blacksburg, Virginia 24061 (703) 961-5910 WASH INGTOtl: Mike Shockman Wildlife Enforcement Department of Game 600 H. Capitol Way 01 ymp ia, Ilashing ton (206) 753-5740 98504 WEST VIRGINIA: Nelson B. Shaw Law Enforcement Division Department of Natural Resources 1800 Washington Street East Charleston, West Virginia 25305 (304) 348-2783 . I~ISCONS IN: Donald L. Beghin Bureau of Law Enforcement Department of Natural Resources Box 7921 Madison, Wisconsin 53707 (608) 266-1115 IIY0r'lI NG: Tom Moore, Research aiologist Game & Fish Research lab Biological Science Building Box 3312 University Station Laramie, ~/yoming 82701 (307) 766-6313 Steve Smith Law Enforcement Section Wyoming Game and Fish Department Cheyenne, Wyoming 82002 � https://cpw.cvlcollections.org/files/original/d7779e49ba1dd746977d11c6afc0176b.pdf f164311209bfaeb125e1188932578148 PDF Text Text Colorado Division of Wildlife Wildl ife Research Report October 1983 JOB PROGRESS REPORT State of Colorado ------~~~~~------------Migratory w-88-R-28 Project No. Work Plan No. Job No. Job Title: Waterfowl Period Covered: Personne I: Bird Investigations Production Surveys April 4, 1982 through June 19, 1982 M. Bauman, G. Byrne, J. Creasy, J. Corey, J. Dennis, J. Ellenberger, J. Frothingham, H. Funk, J. Gray, J. Hicks, S. Hinshaw, J. Kauffeld, D. Kenvin, G. Lorentzson, D. Masden, M. Nail, T. Rause, J. Ringelman, W. Russell, G. Saville, S. Steinert, M. Szymczak, and R. Weldon. ABSTRACT Water conditions were good for waterfowl production throughout the state except for the San Luis Val ley where surface water continues to decl ine. Snow pack was good with a slow runoff. There was no major flooding reported in the state. The total number of breeding pairs of ducks declined 42% from the longterm average. The Yampa Valley was the only area surveyed which showed an increase in the number of breeding pairs. Breeding pairs of Canada geese showed an increase in all trend areas except in northcentral Colorado which indicated a reduction of 21% in gosl ing production from the long-term average, and a reduction of 37.7% from 1981. ��3 WATERFOWL PRODUCTION SURVEYS Gerald Lorentzson P. N. OBJECTIVES 1. To estimate the number of duck breeding pairs, by species, on selected major waterfowl nesting areas in Colorado. 2. To estimate the number of goose breeding pairs, and in some cases, obtain production data on selected goose nesting areas in Colorado. 3. Compile data and submit reports to appropriate state personnel and the Fish and Wildlife Service for use in monitoring status of the various species and establishing hunting season recommendations. SEGMENT OBJECTIVES 1. To estimate the number of duck breeding pairs, by species, in the San Luis, Cache la Poudre, South Platte and Yampa Valleys, and in North Park and Brown's Park, using procedures presented in the Program Narrative. 2. To estimate the number of goose breeding pairs in the San Luis Valley, on the Yampa, Little Snake and Green rivers in northwest Colorado and in northcentral and westcentral Colorado using procedures presented in the Program Narrative. 3. Evaluate the present boundaries of the area sampled for duck breeding pairs in the San Luis Valley. 4. Alter the duck breeding pair survey in the Cache la Poudre and South Platte areas to compensate for sample sections which can no longer be counted because of changes in land use patterns. METHODS AND MATERIALS The 1982 duck breeding pair surveys were conducted during the period of May 11 through June 17. Surveys in North Park, the Cache la Poudre and South Platte Valleys were conducted exclusively from the air. Brown's Park and the Yampa River Valley were made on the ground. Aerial counts in the San Luis Val ley and North Park were adjusted for visibility by air ground comparison studies. Pairs estimates for the Monte Vista National Wildlife Refuge in the San Luis Valley were obtained from nesting transects. All survey methods remained the same as in previous years. �4 Canada Goose surveys were conducted within the April 4th through June 19th period. Estimates of the Colorado, White River, Yampa and Little Snake River populations were obtained by direct counts from a fixed wing airplane. Aerial counts were not done on the North Fork of the Gunnison River this year. Brown's Park continues to be a ground count conducted by the Brown's Park NWR personnel. All flying was done with Cessna 185 aircraft. Two observers were used when flying transects, while one observer was used for sampl ing sections. RESULTS AND DISCUSSION Water conditions for duck production were good in most sections of the State. Surface water continues to decline in the San Luis Valley due to the installation of more irrigation systems and a conflict with landowners over water in the San Luis Lakes area. Reservoirs in the northeast portions of the state were not at capacity at survey time but were completely filled soon after. Snowpack was good throughout the mountainous regions and a cool spring resulted in a slow runoff with little or no flooding. The total duck breeding pair population in Colorado decreased by 62.8% from 1983 and 42.5% from the long-term average (Table 1). Part of the decrease is due to a correction in the visibil ity ratio in 1982 which was not done in 1981. Even though the total numbers of ducks decl ined, the species composition was very close to the long-term average (Table 2), and none of the species showed as spectacular a change as in 1981. Totals by species and area are shown in Table 3. Table 1. Summary of Colorado's in selected areas, 1982. Area San Luis Valley North Park South Platte Valley Cache 1a Poud re Va 11ey Yampa Valley Brown's Park Totals duck breeding pair population Total estimated breeding pairs Long-term average 1981 1982 estimates Percent change Long-term average 1981 12,975 5,545 5,905 5,800 3,587 1,265 26,338 40,599 14,838 8,377 2,572 1,476 26,865 17,989 7,715 4,452 2,605 1,123 -50.8 -86.7 -60.2 -30.8 +28.3 -14.2 -51. 7 -69.2 -23.5 +30.2 +26.4 +12.6 35,077 94,199 60,749 -62.8 -42.3 �5 Table 2. lation. Species composition of Colorado's 1982 duck breeding pair popu- Number of breeding Ea irs 1954-81 average 1981 Percent species composition 1954-81 average 1982 1981 Species 1982 Mallard 15,029 18,186 26,372 42.8 19.3 48.0 Blue winged and cinnamon teal 5,781 16,272 6,343 16.5 17.3 11.2 Gadwa 11 2,957 12,204 5,980 8.4 13.0 10.7 Pintail 2,093 3,556 3,566 6.0 3.8 6.4 Shoveler 2,033 4,152 3,637 5.8 4.4 6.5 2,147 3,572 3,302 6. 1 3.8 5.9 2, 194 16,243 3,255 6.2 17.2 5.5 447 3,923 1,194 1.3 4.2 2. 1 2,192 15,909 2, 155 6.2 17.0 3.5 204 182 35,077 94,199 Green-winged teal Redhead American wigeon Other divers Mergansers TOTALS Table 3. .6 55,804 Estimated population of duck breeding pa irs in Colorado, San Luis Valley North Park Poudre River Ma lla rd 4,310 2,239 2,715 Blue-winged & cinnamon teal 3,340 1982. Yampa Vall ey Brown's Park Total 3,712 1,816 237 15,029 1,014 374 796 257 5,781 S.Platte River Gadwall 904 880 395 346 234 198 2,957 Pintail 670 533 458 232 96 104 2,093 Shoveler 924 373 333 183 96 124 2,033 125 104 206 52 2,147 208 441 138 156 202 27 396 289 Green-winged teal Redhead 1,660 1,167 240 Wi geon Other divers 1,280 Mergansers TOTALS 12,975 5,545 5,800 2, 194 62 447 227 2, 192 22 178 4 204 5,905 3,587 1,265 35,077 �6 The estimated show a sl ight adults (Table personnel and number of Canada geese on the Yampa and Little Snake Rivers increase in the breeding pairs but a decline in the total 4). The Brown's Park count continues to be done by the NWR shows a slight increase over 1981 (Table 5). Table 4. Number of Canada geese observed on aerial surveys Routt counties. Nesting pa irs Nonnesting pairs in Moffat and Total adults 1982 1981 1982 1981 1982 1981 12 23 1 18 11 7 15 4 27 6 10 12 2 11 11 26 50 3 70 41 65 59 46 12 16 5 25 22 80 38 65 18 126 94 341 12 26 21 8 22 24 30 16 38 29 46 46 Yampa River Steamboat Sp r ings-C ra ig Craig-Juniper Springs Juniper Canyon Juniper Springs-Cross Mt. Lily Park TOTALS 190 Little Snake River Cross Mt .-Powderwash Br. Powderwash-Baggs TOTALS Table 5. NWR. Number of Canada geese observed on ground counts Nesting pairs Area Brown's Park 1982 76 1981 Total adults 55 70 125 142 48 190 in Brown's Park Estimated no. of gosl ings1 1982- 19~1 1982 1981 305 277 294 245 Breeding pairs in west central Colorado Canada goose areas increased about The total numbers of non-breeding birds declined about 36% from 1981 (Tables 6 & 7). The Gunnison River was not counted in 1982 which would account for some of the difference. 6%. �7 Table 6. West central Colorado Canada goose breeding Area Singles pair survey, 1982. Pai rs Groups White River Meeker-Rio Blanco Lake Rio Blanco Lake-Rangely Rangely-Utah Line Subtotal 13 46 1 13 24 2 39 60 4 13 16 14 7 2 1 0 6 9 45 36 36 7 0 4 1 6 b3 144 1 1 10 2 12 17 51 0 68 Colorado River Newcastle-Silt Silt-Rifle Rifle-Parachute Parachute-DeBeque DeBeque-Pal isade Palisade-Grand Ave. Bridge Grand Ave. Bridge-Fruita Fruita-Horsethief Canyon Horsethief Canyon-Utah Line Subtotal 14 52 23 23 5 3 15 0 37 172 Roaring Fork River El Jebel-Carbondale Carbondale-Glenwood Subtotal Gunnison Springs 2 44 0 44 River Hotchk iss-De 1 ta Delta-Mesa Co. Line Mesa Co. Line-Whitewat~t Whitewater-Grand Junction Subtotal GRAND TOTAL NOT C 0 U N TED I N 1 982 104 216 284 �8 Table 7. Comparison of the number of Canada goose singles and pairs observed west central Colorado, spring 1978-82. in aerial surveys in Number observed Area Wh ite River Roaring Colorado Fork River 1981 39 39 2 Pairs 1980 1979 1978 1982 1981 1979 1978 30 46 31 60 74 50 26 2 6 3 2 12 7 7 0 56 31 47 41 30 133 105 74 77 35 7 9 6 4 4 11 14 11 12 4 Not done 0 2 0 0 Not done 4 5 " 6 3 0 4 10 8 5 5 104 87 94 94 71 214 138 152 70 39 River Glenwood Springs5th Street Bridge Bridge-Utah Gunnison 1982 Singles 1980 Line River Hotchkiss-Delta Delta-Grand Junction TOTALS 216 0 The estimated number of nesting pairs of Canada geese in the San Luis Valley was up 171% in 1982. The total number of geese observed is up 80% from 1981 (Table 8). This represents the largest number of geese observed since the fixed wing counts were instituted in 1976. T2ble 8. Results of San Luis Valley Canada goose breeding Year Nesting Pairs pair survey. Projected total number Non-nesting Pairs Grouped Birds 1975 Helicopter 59 59 110 1976 Hel icopter Fixed wing 92 77 64 30 1977 Hel icopter Fixed wing 100 100 101 91 130 97 99 96 58 141 226 154 90 159 244 376 No counts 1979 Fixed wi ng 1980 Fixed wi ng ~ Fixed w i n9 1982 Fixed wing 169 �9 The northcentral Colorado Canada goose production survey was done on June 17th and 19th. The results are presented in Table 9. The total number of birds decreased about 11% from 1981. The number of adults decreased by 60 birds, a decrease of 1.4% (Table 10). The large decrease was in the number of goslings produced. This decrease is shown in all areas except the area around Boulder. The total number of goslings decreased 37.7% from 1981 and 20.9% from the long-term average (Table 11). �10 Table 9. Production area Fort Collins Results of northcentral Water area Peterson Ponds Herring Pond Maxwell Pond College Lake Dean Acres Andrijeski Marsh Claymore Lake Sterling Gravel Pits Fort Collins Gravel Pit Lindenmeier Lake Grey Lakes Novak Reservoir Winick Ponds Flatiron Gravel Pits Andersons Pond Parkwood Lake Kitchel Lake Timnath Res. Romily Gravel Pit Fossil Creek Shuelke Res. Horseshoe Lake Equalizer Lake Wolaver Ponds Subtotal Loveland Boulder Colorado goose census, June 17-19, 1982. Flatiron Res. Boedecker Res. Ma r ino Res. Flatiron Gravel Pits Kauffman Gravel Pits Big Thompson River McNeil Res. Welch Lake Res. #12 Subtotal Total no. gosl ings 0 0 11 30 6 0 15 85 0 0 30 13 0 4 Total # adults & yearlings 12 51 8 281 26 12 51 19 311 32 o o 91 217 106 302 o o 72 108 10 72 138 23 o o 17 21 22 96 59 129 54 37 14 8 4 92 51 8 Total birds 31 25 23 0 0 0 0 98 299 1,204 1,503 4 12 100 16 147 47 0 0 0 0 36 45 0 132 Ish Lake 6 Crystal Lake 5 Terry Lake 15 Faivre Ponds 10 Sawhill Pond & Walden Pond 86 Valmont Res. 47 Boulder Valley Farm 5 King Pond 0 Eddy Pond 0 Angus Ranch Pond 2 Subtotal 176 29 14 o 19 o o o 19 o o o o 25 14 25 14 o o 60 113 62 386 96 158 62 ---=-518 13 12 19 17 96 49 212 81 39 126 142 2 189 7 o o 6 6 4 599 2 423 �11 Table 9. (continued) Production area Larimer Denver GRAND TOTAL Water area Total no. ,goslings Total # adults & yearl ings Total birds 77 Terry Lake Ervin Pond Res. #4 Deines Res. Launer Pond Wood Pond Douglas Lake Stewart Pond Poudre #1 Dry Creek N. Poudre #3 N. Poudre #2 N. Poudre #5 N. Poudre #6 Bureau of Standards #2 Bureau of Standards #1 Reservoir #8 Elder Lake #8 Annex Country Club Pond Long Pond Van Sants Takes Pond Cobb Lake Management Area Cobb Lake Hinkley Res. Dale Pond Watson Lake Curtis Lake Beghtol Lake Subtota I 26 0 0 0 51 o o 28 32 60 0 0 13 o o 37 37 35 48 9 6 15 0 21 0 6 0 0 7 0 27 27 15 36 Ketring Centennial Columbine Country Club Cooley Sand Gravel Bowles Lake King's Pond Tule Lakes Grant Ponds Marston Res. Pinehurst Country Club Clarefield Res. Ward Res. Kendrick Lake Fed. Center Pond Sloans Lake Stanley Lake Denver City Park Colo. Blvd. @ Quincy Blackmer Res. Subtotal 27 o o 15 15 o o 8 14 16 16 o o 13 85 4 6 85 39 37 42 o o 79 0 0 14 0 0 0 0 27 10 11 Not done o o 41 o 219 219 o o o 9 19 183 194 o 9 o o o o 852 6 14 43 o 1,038 94 76 121 30 44 82 87 96 206 206 147 92 o 147 20 8 72 35 3 50 53 38 72 110 55 55 43 o 380 409 o 55 55 o 24 24 24 24 407 1,561 4,272 5,219 29 o 154 1, �12 Table 10. Number of adult Canada geese observed production trend areas, 1982. Area Wellington Fort Coll ins Loveland Boulder Denver Total 1982 in northcentral Colorado No. of adults average 1981 1970-1981 Percent change from 1981 1970-1981 852 1,204 386 423 1,407 864 1,005 458 476 1,529 734 726 266 530 1,233 - 1.4 +19.8 -15.8 -11 .1 - 8.0 +13.8 +39.7 +31 .1 -20.2 +12.4 4,272 4,332 3,489 - 1.4 +18.3 Table 11. Number of Canada goose goslings produced in northcentral Colorado production trend areas, 1982. Area Wellington Fort Coll ins Loveland Boulder Denver Total No. of goslings average 1982 1981 1970-1981 Percent change from 1981 1970-1981 186 299 132 176 154 308 424 212 175 401 263 318 116 206 294 -39.6 -29.5 -37.7 -61.6 -29.3 - 6.0 +13.8 -14.6 -47.7 947 1,520 1,197 -37.7 -20.9 �13 Colorado Division of Wildlife Wildl ife Research Report October 1983 JOB PROGRESS State of Colorado Project No. w-88-R-28 Work Plan No. REPORT Migratory Job No. Ecological Job Title: Bird Investigations 14 Studies of the Fl ightless Period of Ducks in Colorado Period Covered: Personne I: 1 Apri I 1982 - 31 March 1983 J. Corey, S. Porter, S. Steinert, G. Tischbein, J. Wagner, J. Ringelman and ~. Szymczak. Colorado Division of Wildlife. ABSTRACT Thirty-seven wetlands in North Park were selected to study the flightless period of adult ducks. Most wetlands were photographed from the air and cover maps prepared. A buoy grid system 1,000 feet apart was placed on Walden Reservoir to aid in census and observation of marked and/or unmarked molting ducks. Active invertebrate fauna were sampled twice during the summer period at Walden, MacFarlane and L. John Annex. Counts of fl ightless ducks were conducted on all wetlands at least twice during the summer period. Some flightless birds were observed on most areas, but only Walden, MacFarlane and Sneed Reservoir and Lake John Annex and Boettcher Lake contained substantial numbers. Primarily because of the lack of flightless adult mallards, only gadwall were studied intensively in 1982. Visual markers were placed on 12 adult male and 7 adult female gadwall and radio-transmitters were attached to 1 adult male and 3 adult females. The locations of the marked birds were plotted periodically during their flightless period. Time budget data was collected on adult male gadwall for 1565 minutes and 1064 minutes during the pre-flightless and fl ightless periods, respectiv~ly. Morphological measurements were recorded for over 300 adult males and over 180 adult males. A significant weight loss was observed in both sexes from the mid- to later stage of molt. Markers had no unusual effect on fl ightless gadwall in terms of aberrant weight loss or gain carcass analysis of 24 adult males collected at specific molt stages indicated no significant variation in weight, percent fat or percent protein in relation to stage of molt, but sample sizes were considered too small to detect differences. ��15 ECOLOGICAL STUDIES OF THE FLIGHTLESS OF DUCKS IN COLORADO PERIOD Michael R. Szymczak James K. Ringelman P. N. OBJECTIVES 1. Document the species and sex composition and seasonal ducks molting on selected wetlands in North Park. 2. Identify the physical by molting ducks. 3. Investigate spatial and temporal differences molting ducks. 4. Determine the behavioral molting ducks. 5. Investigate differences in duration of the fl ightless period of selected species of ducks in relation to sex, body condition, habitat qual ity, and time of molt. 6. Determine the survival rate of molting ducks in relation condition, habitat quality, and time of molt. 7. Identify and quantify and biological characteristics abundance of wetlands in use of wetlands of used by time budgets and net energy balance of duck molting wetlands to sex, body in Colorado. SEGMENT OBJECTIVES 1. Document the species and sex composition and seasonal abundance ducks molting on selected wetlands in North Park. 2. Identify the physical and biological by molting ducks. 3. Investigate spatial and temporal differences molting ducks. 4. Determine the behavioral ing ducks. 5. Investigate differences in duration of the flightless period of selected species of ducks in relation to sex, body condition, habitat quality, and time of molt. characteristics of wetlands in use of wetlands of used by time budgets and net energy balance of molt- �16 INTRODUCTION This report covers the first year of a study of ducks during the flightless period. As with most studies, the first year of field work was hampered by unexpected occurrences. The major problem was the lack of molting adult mallards for study. From 1965 through 1969 over 1,700 flightless adult mallards were trapped on Walden and MacFarlane reservoirs and Lake John Annex in North Park. From 1971-80, numerous flightless mallards were captured in bait trap even though the trapping period was beyond the peak period of molt for that species. In 1982, a considerable number of molting mallards were found only on Boettcher Lake. Therefore, the emphasis of species data collection during the 1982 ~eason was directed toward gadwall. METHODS Study Wetland Characteristics and Waterfowl Use Utilizing USGS 1:24,000 topographic maps and larger scale maps of recent origin of the Arapaho National Wildlife Refuge, all wetlands that were estimated to be at least 5 acres in size were assigned identification numbers. The wetland classification (Cowardin et al. 1979) was obtained by examining draft maps provided by the U.S. Fish and Wildlife Service, National Wetlands Inventory. Forty of the 60 wetlands identified were selected for study. Five areas known to support fl ightless adult ducks and 5 wetlands subject to heavy fishing pressure were selected. These 10 selected areas were all classified as members of the Lacustrine (lake) system. The remaining wetlands were stratified according to wetland classification and study areas selected at random according to the percentage composition of the various wetland types. Subsequent field investigations revealed 3 of the 40 wetlands were dry in 1982 and those were dropped from study. Study wetlands were photographed from the air on 5 August, 1982 under contract, by the Colorado State Forest Service. Three wetlands were inadvertently not photographed. Wetland cover maps (Appendix A) were prepared by that same agency designating area and number of patches of each aquatic vegetation type, upland (islands), and open water and perimeter. The data were entered and stored in a computer file in a format enabling further analysis. The size and location of the study wetlands are presented in Table 1. In addition the composition of vegetative maps were examined on sample wetlands number 2, 3, 10, 11, 18, 19, 21 and 22. �17 Table 1. Number Some characteristics Name of study wetlands UTM grid locationa l. John 1 2 L. John Annex Walden Res. MacFa r 1 ane Res. Damfino Res. Pole Mountain Res. S. Delaney Butte L. E. Delaney Butte L. N. Delaney Butte L. Boettcher L. Sneed Res. Tricks Pd. #1 Ca r 1 strom Res. Tricks Pd. #2 Holzingers Home Pd. Antelope Pd. Case #2 Res. Case #3 Res. 76 Pd. Elk Pd. Goose Pd. Marsh Pd. Eagle Pd. S. School Section Pd. \4illford Pd. Spring Creek Pd. S. Allard Pd. Geisses Pd. Hebron Clayton Res. Addison Res. Rich Pd. HUdspeth Wattenburg Pd. Richard Pd. E. Boettcher 3 4 5 6 7 8 9 10 11 12 13 14 16b 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31b 32b 33 34 35 36 37 40 in North Park. 4515 4515 4509 4489 4486 4490 4506 4507 4508 4520 4524 4522 4521 4523 4509 4507 4503 4502 4502 4503 4501 4502 4501 4500 4501 4502 4501 4595 4490 4490 4484 4485 4486 499 4513 4516 4523 x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 375 376 388 391 381 374 376 377 376 372 371 389 3~9 389 395 392 390 387 385 388 387 388 388 391 393 394 395 391 386 385 388 386 380 379 378 376 372 aGrid location near the center of the wetland. b Not photographed. Area (ha) 201.99 72.40 1904.65 112.80 8.95 49. 16 57.47 25.92 44.55 49.37 68. 11 3.03 37.46 5.32 15.97 7.49 6.91 5.42 5.81 8.74 4.27 3.91 2.06 1. 73 3.45 10.26 3.32 3.32 2.58 2.31 4.90 4.28 5.71 2.64 �18 Because of the absence of flightless adults in early July and the workload in late July through September, counts were conducted on most wetlands only twice during the study period. Counts on Walden Res. were made more frequently. Early season counts began on 28 July 1982 and were completed on 10 August 1982 while late season counts began on 26 August 1982 and were completed on 9 September 1982. Counts were conducted primarily during early morning or late evening hours. All adult ducks observed on the wetland were recorded to species, sex, if possible, and as to whether they were fl ightless or not. Starting and ending time of the count and temperature and wind data were also recorded. Data were stored in a computer file in a format which would allow analysis. Some summary statistics were computed. Characteristics of Molting Wetlands and Duck Habitat Selection Walden Reservoir was selected for intense study of flightless adults since it was known to be a major molting area. To aid primarily in locating the position on the reservoir of birds observed, and census, a buoy grid system was establ ished on the reservoir (Figure 1). Base 1ines were established along the south and west sides of the reservoir and marked at 1,000 foot intervals. The anchored buoys were placed at the intercept of lines ~rpendicular to the base lines at each interval (Appendix A). Placement required radio communication between 2 observers with transits and 2 people in the boat. The orange buoys were sponge-type floats 6 in diameter and 14" long anchored by polypropylene rope to cement blocks. Undulating terrain along base lines, buoy placement error, wind action and water level fluctuation resulted in buoys which were not floating at exact 1000 ft. intervals. Some adjustments were made throughout the study period. Fence posts were used as a substitute for buoys in shallow water and as shoreline markers along buoy lines. Each square of the grid was assigned an alpha-numeric designation with the buoy in the southeast corner of that grid having the same designation. Water level fluctuations were monitored through measurement of the corner stake at G7. 11 Active invertebrate fauna were sampled at Walden and MacFarlane reservoirs and L. John Annex using activity traps as described by Whitman (1974) modified to enable suspension of 3 years about 12 inches below the water level. At Walden, traps were open for approximately 24 hours, attached to buoys Bl, E2, FO, G2, G4, G6, Hl, J4 and J7 on 14 July 1982 and 8 September 1982. Traps were also set at B2 during July sampling but at Fl in September because the Bl buoy was missjng. Trap sites were selected to represent different depths and locations on Walden. At MacFarlane and Lake John Annex traps were set at 5 locations for a 48-hour period on 20 July 1982. On both areas traps were selected to provide good coverage of the Reservoir. Water depth and notes on aquatic vegetation were recorded at each trap site. Invertebrates collected were frozen in water for future analysis. �19 Samples of aquatic vegetation were collected at Walden for analysis of chemical composition. Species collected were Potamogeton filiformis, Potamogeton richa~dsonii, Polygonum amphibium, Ranunculus circinatus, Ceratophyllum demersum, .Myriophyllum exalbescens and filamentous algae. A portion of each sample was dried and a second portion frozen for storage prior to analysis. Data on habitat utilization on Walden were collected by recording (1) location of flocks of flightless waterfowl during periodic counts of ducks on Walden, (2) location and activity of gadwall observed during time budget data collecting bouts during the pre-flightless and flightless periods, (3) location and activity of individual gadwall marked with back tags (Furrer 1979), and (4) location of radio-marked gadwall. All data, except those collected during time-budget bouts were recorded on maps similar to Figure 1. Visual markers and radio-transmitters were appl ied to molting gadwall captured with a long-handled dip-net from an airthrust boat. Both types of markers were attached to the ducks using a back-pack harness (Dwyer 1972). Numbers were stitched onto back-tag visual markers to allow individual identification. Markers were applied periodically throughout the gadwall's flightless period beginning on 3 August, 1982 with the last markers applied on 2 September, 1982. Tags were placed on 12 adult males and 7 adult females. Radio transmitters were attached to adult male (3 August 1982) and 3 adult females (2 September 1982). Instrumented bird locations were determined through triangulations utilizing a vehicle-mounted precision direction finding antenna array, vertically mounted and spaced 1 wave-length apart. Observations and radio-locating of marked birds began on 10 August 1982 and were conducted periodically throughout the flightless and post-flightless period until 23 September 1982. Observations and radiolocations for each individual overtime were pooled and crude home range estimates computed. Behavior Time Budgets and Energy Balance Time budget data for adult gadwall were collected during a pre-fl ightless (just prior to remige molt) and flightless period. Birds observed during the flightless period-were recorded as early or late in the molt sequence depending on observed feather length. When collecting data through scan sampling, molt stages of the birds observed varied and the stage of molt was classified as "unknown". Time budget bouts using the "focal bird" technique involved a maximum of 90 minutes of continuous observation of one bird recording behavior data, as listed in Appendix A, every 20 seconds. Focal bird time budget data were limited almost exclusively to adult male observations in 1982. "Scan" time budget bouts, recording the instantaneous activity of each individual as a group of adult gadwall was scanned, were also done. Since sex and stage of molt, in most instances, cannot be determined during scan sample, gadwall observed during most collection bouts were classified as fl ightless, but of unknown sex and stage of molt. �20 Time budget data were recorded on micro-cassettes and transcribed to computer coding forms along with other pertinent information (Appendix B). A summary of time budget observations are presented in Tables 2 and 3. Data were entered into a computer file for analysis. Table 2. A summary of focal bird time budget observations and flightless adult male gadwall in North Park, 1983. Time interval 0530-0830 0831-1130 1131-1430 1431-1730 1731-2030 Pre-fl ight less 393 264 251 392 265 Minutes of observation Fl i9htless Late stage Early stage 58 47 89 37 of prefl ightless Unk. stage 95 319 177 91 65 86 Table 3. A summary of scan sample time budget observations of prefl ightless and fl ightless adult gadwall in North Park, 1982. Time interval 0530-0830 0831-1130 1131-1430 1431-1730 1731-2030 Prefl ight less Minutes of observation Fl i9htless Early stage Late 708 181 Unk. stage 308 371 274 130 577 Adult gadwall trapped in conjunction with a banding operation were weighed and measured. Data collected were designed to classify stage of molt and be used in the development of a condition index for molting birds. The categories presented in Appendix C are self-explanatory except that (1) primary feather lengths were measured from the outer edge of the sheath to the tip, (2) U-B length was the distance from the uropygial gland to the tip of the nail of the bill, (3) wing-P length is the distance from the wrist joint to the distal end of the manus, (4) bill length was from the gap to the tip of the nail of the bill and (5) girth was the circumference of the body at the base of the wing measured with a seamstress tape. The tape had a loop-type buckle stitched on each end and was cinched around the girth of the duck to a spring-scale tension of 1000 grams. If a bird was subsequently recaptured only primary lengths, weight and girth measurements were recorded. �21 The adult gadwall measured were classified as to stage of development of the 10th primary. The number of birds measured by stage of development is presented in Table 4, including those birds captured more than once. Data were entered on a computer file for analysis. Table 4. Number of adult gadwall measured in North Park, 1982. by stage of wing feather molt Sex Primary #1 status (mm) Males Pre-molta-20 21-50 51-90 91-130 131-170+ aprimary 48 64 84 108 53 Females 88 47 39 24 4 feathers still in place. Between 3 August and 24 August 1982, 24 adult males were collected for carcass analysis; 5 each in the first 4 molt development stages and 4 in the last stage. The birds were measured, according to Appendix B, plucked and frozen in plactic bags for storage. The carcasses were later thawed, the contents of the G. I. tract, bill and feet removed, and the following tissues disected and weighed: pectoral muscle (1), leg (1), gizzard, liver, small intestine and large intestine. The whole carcass minus feet and bill was then ground to a uniform consistency through a Hobart 4852 commercial grinder. A 15-20 gram sample of the homogenate was dried at 1000 C for 48 hours and lipids extracted using a Soxhlet apparatus with petroleum ether distillate for an 8-hour period. An additional sample was ashed in a muffle furnac~ at 5500 for 12 hours. Extensive attempts to collect only 4 birds being collected; (39 mm) and 1 flightless male from the dead bird within 1-2 gadwall for food habits studies resulted in 2 pre-flightless females, 1 flightless female (136 mm). Esophageal contents were extracted minutes after collection. All observations (time budgets, duck counts) were conducted using a Questar telescope with some limited use of a 20 x 60 variable Bausch and Lomb spotting scope. Temperatures were monitored throughout the 1982 study period by a thermograph placed in the old Walden Fish Hatchery on Case Flats in the Arapaho National Wildlife Refuge. �22 RESULTS AND DISCUSSION Analysis of data to date has been limited primarily to morphological measurements of captured flightless birds and carcass analysis. Duck counts on sample wetlands Counts of flightless ducks on sample ponds were considerably more difficult than anticipated. During both count periods it was sometimes difficult to determine whether an adult bird was flightless or not and during the later period it was also difficult to differentiate between a fl ightless adult and a Class III duckling, particularly when viewing at long range. Adult diving ducks were more difficult to classify to status than dabblers. Therefore counts do not represent the exact number of flightless ducks on each wetland but only those whose status could be discerned. Flightless adults were observed on a variety of wetlands (Table 5). Generally the smaller wetlands contained only a few flightless birds. Wetlands with a comparatively large number of flightless adults were Walden, MacFarlane, Pole Mountain and Sneed reservoirs, Lake John Annex and Boettcher Lake. Walden Reservoir is not included in Table 5 and MarFarlane Reservoir was not included in the second count period. Table 5. Sample wetlands on which flightless adult ducks were observed, by species, during the 2 regular count periods in North Park, 1982. Inclusive dates 28 July - 10 August Wetland nos. Species Mallard 2,4,6,10,11,12,17,19,21,22,28,29,30,40 Gadwall 2,4,6,10,11,19,21,22,23,24,26,33,34 American Wigeon 2,4,6,10,11,21,22,28,31,34,35,37,40 Pintail teal Lesser scaup 10,11,12,19,30,31,37,40 1,6, 10, 11 ,27,28 Redhead Ring-necked 2,10,",19,22 2,5,6,10,11,18,21,22,23,24 2,6,10,18,21,23 2,4,5,6,10,11,13,19,20,29,35,40 Blue-winged and/or Cinnamon teal 2,5,10,12,13,14,19,20,21,22,30,34,40 Green-winged 26 August - 9 September Wetland nos. 2,10,11,12,19,21,37 duck 2,6,10,11,28,110 18,21,22,24,35 20,21,22,23 6,10,11,18,19,20,21,32,33,37 10,11,18,22,35,37 10,11,37 Canvasback 2,10 10 Ruddy duck 2,10 10,37 Northern shoveler 35 �23 Morphological Measurements of Gadwall Of the 5 morphological body measurements recorded for captured molting gadwall, length, wing and bill were considered to be stable measurements that would not fluctuate with body condition. Weight obviously changes and girth has been indicated as a dependent variable in estimation of fat stores of mallards (Ringe1man and Szymczak, unpub1. data). Data presented in Table 6 indicate slight variability within sex in the gadwall population according to the former 3 measurements. Girth also showed 1 ittle variation, although data have yet to be analyzed in relation to fat stores. Weight did vary and preliminary implications are discussed in another section. Walden vs. MacFarlane MacFarlane Reservoir was chained completely during the summer of 1981 and refilled in the Spring of 1982. One evaluation of the effect of drainage on the molting population was to compare morphological measurements of gadwall captured at MacFarlane with those captured at Walden Res. Very few gadwall used MacFarlane during their fl ightless period in 1982. Only 10 birds of each sex were available for analysis. A grass analysis of all morphological measurements, disregarding stage of molt, indicated a significant difference (p = .039) in weight of males. Unexpectedly, the MacFarlane birds were heavier (x = 835 gr vs. 794 gr). Bills were also significantly larger for MacFarlane males (£ = .005) indicating that structurally, MacFarlane males may be larger than males in the Walden population. Females showed no significant difference in weight or girth in spite of being significantly longer (UB length) at MacFarlane (p = .004). Marked vs. Unmarked Birds Attachment of radio-packages and back-tag resulted in no obvious unusual behavior of marked birds during the flightless period. Very late in the molt, after the marked birds had reacquired the ability to fly, some with back tags expressed a reluctance to fly. Fifteen marked birds were recaptured during the molt period after initially attaching the markers. No abrasive wear was noted on the birds from the harness. No significant differences were noted in total weight loss and percent of weight loss between recaptured marked and unmarked birds on both a seasonal and daily basis. Use of Energy and Energy Reserves Weights of gadwall captured were examined in relation to stage of molt to get an indication of weight dynamics of the birds as a group during the flightless period. Means by sex were calculated and plotted by stage as indicated in Figure 1. The graphic presentation indicates that the weight dynamics of both sexes are quite similar. No significant differences were found when comparing total and rate of weight change by sex. A significant decline in weight was noted for both sexes between the 3rd and 5th stage (~~ .05) and females weights also increased graphically but the means were not significantly different. �24 Specific information on weight dynamics of individual birds was obtained by comparing weights of birds captured more than once during the molt cycle, excluding those birds captured in the pre-molt stage. Calculations indicated a mean weight loss of 7 gr/day for males and 6 gr/day for females. The rate of loss was significant for both sexes (p < 0.001). No attempt was made at this time to assign recaptures to specific molt stages. These two tests indicate that weight loss does occur in gadwall at least at some period during remige molt indicating that birds rely in part on endogenous reserves during the period. Carcass weight, percent fat and percent protein values for individual birds collected are presented in Table 7. These data are summarized by stage of primary molt in Table 8. Note that these data are grouped according to primary growth stages indicated in Figure 1. Analysis of variance indicated no significant difference between molt stages in weight, percent fat or percent protein. Most likely sample sizes were too small to detect any differences, such as those that occurred in weight for the larger data set (Figure 1). LITERATURE CITED Cowardin, L. M., V. Carter, F. C. Golet and E. T. LaRoe. 1979. Classification of wetlands and deepwater habitats of the United States. U.S. Dep. Interior, Fish and Wildl. Serv., Office of Biol. Servo FWS/OBS79/31. 103pp. Dwyer, T. J. 1972. 43:282-284. An adjustable radio-package for ducks. Bird Banding Furrer, R. K. 1979. Experiences with a new back-tag for open-nesting passerines. J. Wildl. Manage. 43(1);245-249. Whitman, W. R. 1974. marsh management. The response of macroinvertebrates to experimental Ph.D. Thesis, Univ. Maine, Orono. 114pp. �Table 6. Summary of morphological measurements in North Park. (mm) of gadwall captured during the molt period Males Females No. Y - S.D. 427.0 - 429.7 187 394.8 12.1 104.5 - 105.3 188 97.7 4. 1 97. 1 - 98.3 51.9 188 47.8 2. 1 47.5 - 48.1 9.3 227.2 - 229.0 207 215.7 9.7 214.4 - 217.0 62.2 788.8 - 801.1 213 702.0 64.4 693.3 - 710.7 Measurement No. X - S.D. Length 314 428.3 12.3 \n ng 324 104.9 3.5 Bill 323 51.7 1.9 Girth 390 228.1 We ight 394 795.0 95% C. I. 51.5 - 95% C. I. 393.0 - 396.5 N V1 �26 Table 7. according Date-tO Carcass composition of adult male gadwall to development stage. Number 08-03-82-1 Length of (mm) #10 Primary Prefl ightless Plucked Carcass weight 836 08-03-82-2 II 08-03-82-3 08-10-82-4 II 08-24-82-2 II 08-10-82-3 08-10-82-6 08-12-82-1 08-18-82-6 08-19-82-4 08-18-82-1 08-10-82-5 08-18-82-2 08-10-82-1 27 30 70 70 74 719 713 667 08-10-82-2 08-12-82-2 08-18-82-4 75 110 110 08-18-82-3 08-19-82-1 08-18-82-5 08-24-82-1 08-24-82-3 08-19-82-3 08-19-82-2 a II collected in North Park % Fat 32.7 14.6 14. 1 703 774 787 28.9 % Protein 57.1 72.9 75.4 59.9 61.9 749 762 26.9 18.5 778 801 712 822 30.5 25.3 18.3 38.7 53.5 758 33.7 14.8 "'Ob 57.7 72.9 88.5b 17.3 70.3 704 31.1 736 681 19.0 26.7 111 730 25.1 59.9 70.0 64.5 64.2 111 789 723 800 22.9 65.1 17.3 17.2 ll18 757 779 28.3 24.3 69.9 70.5 62.7 64.2 161 787 37.5 54.6 31 33 37 69 113 136 144 Results of fat analysis dependent on fat analysis. are questionable; % protein calculation 70.3 60.0 63.7 69.6 �Table 8. Weight, Pr ima ry #10 status (mm) fat and protein dynamics Number of birds of adult male gadwall Plucked we igh t (gr) X - S.D. collected in North Park, 1983.a % Protein % Fat X - S. D. - X S.D. Pre-fl ightless 5 769.8 49.0 23.4 8.6 65.4 8.2 0-30 2 770.0 11.3 24.5 8.5 65.2 7.3 31-60 3 778.3 58.4 27.4 10.4 62.3 8. 1 61-90 4 712.0 37.7 24.2 9.6 65.2 7.5 91-120 5 731.8 38.6 22.2 3.9 66.7 2.9 120 4 780.8 18.0 26.8 8.5 63.0 6.5 > a'ncludes all males except 08-18-82-2 from Table 7. N --...J �N co 56 825 68 800 1~ -;;;- MALES 775 I :0:: 117 <t rr: .::. 750 I- := '-" . I w.J 725 44 J: >- I CI ,',Sarnp1e \ 56/ 5 ize 0 '" 700 -I /' 5 \ FEMALES 40 " 675~ "27 650 1 pre-rno1t 2 0-30 mm (pri 10) 3 31-60 MOL Figure 1. Relationship of body weight T 4 61-90 S TAG 5 91-120 6 >120 E of gadwall to flightless stage in North Park, 1983. �29 WALDEN RESERVOIR (3) - 95 % A photo scole - 1'10000 mop scele-I'9500 B S-I (18) = 229.10 he S -2 (34) = 242.54 he S-5 (5) = 20.71 he S-6 (I) = 1405 he S-7 (2) = 1.21 ho E-I (I) = 3.42 he E-2 (9) = 5.35 he E- 3 (36) = 110.09 he U( 15) = 45.41 he 0(6) = 1219.66 he c TOTAL AREA = 1904.65 he PERIMETER = 16898.32 m D ,. F J K o Appendix 1 A. 2 3 4 5 6 8 Cover map of Walden Reservoir indicating grid system used in buoy placement. A buoy or marker stake was placed where the lines intercept. �30 COLill/IN (S ) 1-3 4 5 6-9 10-15 16-17 18-19 20-23 24 25+ DATA CODE AOU number- species AOU l=fernale; 29'!'.2.1e; O=urJ.:::O':m 1=adul,t; 2=1...7.2ture; 3=un.\mo':m wet.Land numoer code month-day-year (see oe Iow ) sex age location date life stage skip st ar-ting t irae sarrp Le type benavior-a'l data sleeping swirrntng .. walking co<.U'ort movement 102.i'i..'1g on land/ice loafing in watersur-race feeding bill-dip feeding head-dip feeding tippir:.~ field feedi..'1g feedlot 1'eeting intraspecific aggression i..'1terspecific aggression courtship alert flyi..Y1g 24-hour military time l=1'oc21 a'1i:.al; 2=scan 01 02 03 04 05 06 07 08 09 10 11 12 13 14 15 16 17 .. 18 89 vocalize . out of sight end of data 99 24-hour mi1itarj time temperature o? wi.nd speed 1...'1 mph endir!g time avg , terr:perature avg. wi.nd speed 'lime ~"ite!"'v'2.1s: Life stages: 0530-0830 0830-1130 1130-1430 1430-1730 1730-2030 10 breeding (gener-al ) pre-nesting nesting brood-rearing post-breeding (general) pre-flightless early fli~'1tless late fliv.htless flightless - stage unknown post-molt fall mtgratcry '..jinterin~ (~er.eY'al) early wintering mid wintering late Wintering spring migratory 11 12 13 20 21 22 23 24 25 30 1:0 41 42 43 50 Appendix B. Codes for collection of behavioral data. �WATERFOWL Species Number Sex/Age Primary 1 CONDITION Lengths 5 10 U-B INDEX DATA Lengths Wing-P Bill Girth Weight Comments I J , i , ---_.- _-- ---- ... w Appendix C. Recording form for measurement of adult gadwall captured in North Park, 1982. ��Colorado Division of Wildlife Wildlife Research Report October 1983 33 JOB PROGRESS State of Colorado Project No. w-88-R-28 Migratory Work Plan No. Bird Investigations Job No. Development Job Title: and Use of a Physiological for Monitoring Period Covered: Personne I: REPORT Wintering 15 Condition Mallard Nutrient Index Reserves 1 April 1982 - 31 March 1983 G. Berlin, J. F. Corey, H. D. Funk, G. M. Lorentzson, M. K. Pinkham, I. ~. Rl nqe lrnan , s. F. Steinert, M. L. Stevens, M. R. Szymczak, C. A. Weinland, Colorado Division of Wildlife; M. Schoenfeld, Colorado State University; R. Field, U. S. Fish and Wildlife Service. ABSTRACT During early-, mid-, and late-winter 1982-83, 1,183 mallards were weighed and measured at four sites in eastern Colorado. These data, approximately balanced over time and among age/sex classes, indicate that regional and temporal differences in body weight and hence physiological condition exist within Colorado .. Birds captured at Bonny Reservoir appeared to be in better condition in 1983 than in 1982. Ninety-six mallards were collected at Bonny Reservoir, plucked, ground to a uniform homogenate, and samples submitted for whole carcass analysis of body I ipid and ash. When the analyses are completed, the physiological condition index will be developed and applied to live bird weight/measurement data to precisely determine. the nutritional status of wintering populations. ��35 DEVELOPMENT AND USE OF A PHYSIOLOGICAL CONDITION INDEX FOR MONITORING WINTERING MALLARD NUTRIENT RESERVES James K. Ringelman Michael R. Szymczak P. N. OBJECTIVES 1. Develop a physiological condition index to accurately reserve levels of wintering mallards. 2. Determine if differences exist in the physiological mallards wintering in several areas of northeastern 3. Document temporal changes in mallard major Colorado wintering area. physiological assess nutrient condition Colorado. condition of on a 4. Relate spatial and temporal differences such differences cereal grain. exist, to weather in mallard condition, if and the availabil ity of waste SEGMENT OBJECTIVES 1. Develop a physiological condition index to accurately reserve levels of wintering mallards. 2. Determine if differences exist in the physiological mallards wintering in several areas of northeastern 3. Document temporal changes in mallard major Colorado wintering area. physiological assess nutrient condition Colorado. condition of on a 4. Relate spatial and temporal differences such differences cereal grain. exist, to weather in mallard condition, if and the availabil ity of waste METHODS AND MATERIALS Salt Plains bait traps were used to capture mallards during the winter of 1982-83. Two wintering populations, one at Bonny Reservoir and the other in the vicinity of Kodak Ponds/Woods Lake, were sampled during early-, mid-, and late-winter. Two additional populations at Segelke Slough and Valmont Reservoir were sampled during mid-winter only. Trapped birds were weighed to the nearest 10 grams and the following measurements taken: total body length, wing length, bill length, and girth. When necessary, trapped birds were held 12-24 hours until bait corn was digested, thus minimizing the effect of food weight on body weight measurements. �36 Initial objectives dictated that a minimum of 20 mallards of each age/sex class be measured during each time period. Preli~inary analyses of within class variation in December measurements suggested that a minimum of 40 birds in each class should be measured to be 95% certain of detecting a 10% difference in body weight. Thus, following the December trapping effort, the initial quota of 20 was increased to 40 birds per age/sex class to allow for increased precision and statistical power. During each of the three trapping periods, 32 mallards (8 of each age/sex class) were collected at Bonny Reservoir. After being weighed and measured in the standard manner, birds wer~ euthanized, plucked, labeled, doublewrapped in airtight plastic bags, and frozen. In the laboratory, birds were thawed and after removal of the right wing at the humeral-ulner articulation, ground whole in a Hobart 4852 commercial meat grinder to a uniform consistency. The lipid content of the right wing, used in another cooperative study, will be included in the whole body lipid estimate prior to data analysis. Soxhlet extraction apparatus using petroleum ether distillate were used for lipid extractions performed for 8 hours/sample. Percent moisture was determined by air-drying a 15-20 gram sample at 1000 C for 48 hours. Ash was determined by ashing in a muffle furnace at 5500 C for 12 hours. Nonfat, lean, dry weight, assumed to be nearly entirely protein, was determined by subtraction. Simple and mUltiple correlation analyses will be performed to derive the combination of weight and measurements that best predict actual whole body fat and protein levels. RESULTS AND DISCUSSION A combined total of 1,183 mallards were weighed and measured during winter 1982-83 (Table 1). Quotas of each age/sex class were met or exceeded in all but four cases. The early trapping period included the first week in December, mid-winter trapping ran the period 11-21 January, and the latewinter period lasted 24 February - 3 March. Body weight and measurement data were coded and entered on computer. PreI iminary summary statistics have been computed, but detailed statistical analyses await completion of whole carcass lipid extraction data. Body weights of all age/sex classes combined appeared to vary by region and over time (Fig. 1), although these means have not yet been subjected to statistical tests of significance. The average weight of first-year females (979 g), the lightest age/sex class, was 15% less than the weight of adult males (1151 g), the heaviest class. During mid-winter, variations in body weights of females among different areas appeared to vary more than male body weights (Table 2). �37 Compared to body weights taken in an identical manner during mid-winter 1982, Bonny Reservoir mallards were heavier during 1983. Adult female birds in particular were appreciably heavier in 1983 (Table 3). Future Work The reasons behind temporal and spatial differences in body weights will be investigated. Under the hypothesis that weather and/or food availabil ity regulates body nutrient reserves, weather records and grain crop abundance data for each study site will be examined and correlated with body weight and fat reserves. The condition index, to be developed from the 96 collected birds, will be appl ied to weight/measurement data obtained at the four sites to enable use of actual fat reserve levels in evaluating condition. Detailed statistical analyses will be performed to determine if and where real differences exist in nutrient reserves and physiological condition. Prepa red by \~.r-~,%~ <::tamesK. Ringel man''''-.,..;j Wildlife Researcher B �Table 1. w Numbers of mallards captured and measured by trapping location and date. Traeping Locations-Number Bonny Reservoir 00 Captured Kodak-Woods Lake Valmont Reservoir Segelke Slough Age/Sex Class 12/2/8212/7/82 1/15/831/21/83 2/27/833/3/83 12/2/8212/4/82 1/11/831/15/83 2/24/833/2/83 1/17/831/21/83 1/22/831/26/83 Totals Adult females 33 46 40 20 29 39 42 40 289 Adult males 30 57 40 20 40 40 40 40 307 First-year females 33 40 43 20 40 40 36 38 290 First-year males 37 40 40 20 40 LIO LIO 40 297 133 183 163 80 149 159 158 158 Totals �Table 2. Mallard body weights during mid-winter, 1983, at four eastern Body Weight Age/Sex Adult Class female Adult male First-year female First-year male Bonny Reservoir Kodak/woods Lake Colorado locations. All weights in grams. by Location Valmont Reservoir Segelke Slough 1143 985 997 1045 1164 1135 1181 1166 967 986 991 1012 1083 1105 1105 1147 W 1..0 �40 Table 3. Comparison of 1982 and 1983 mid-winter weights of mallards at Bonny Reservoir. All weights in grams. captured % difference Age/Sex Class Adult female Adult male First-year female First-year male 1982 1983 1982 to 1983 959 1143 19.2 1090 1164 6.8 902 967 7.2 1029 1083 5.2 �41 ? 1120' / -- / / en ·E <, tU '0) s: 0) / <, / <, / ... <, <, 10'80' <, <, CD ;: "0 / <, "e' / >0 OJ 10'40' oKodak/Wood's Lake .Sonny Res. .6Segelke's Slough .•.Valmont Res. Dec. Ja!1. Feb. Figure 1. Body weights (grams) of mallards at four eastern Colorado locations during early-, mid-, and late-winter. Weights represent pooled values for all age/sex classes combined. ��43 Colorado Division of Wildlife Wildl ife Research Report October 1983 JOB PROGRESS REPORT State of Colorado --------------------------- 2 Work Plan No. Population Period Covered: in Southeastern Bird Investigations 9 Job No. Monitor Banding of the Shortgrass Job Title: Personne 1: Migratory w-88-R-28 Project No. Prairie Canada Goose Colorado January and February 1983 Jennifer Slater ABSTRACT There were no geese banded in Southeastern Colorado postseason A snowstorm forced the wintering birds out of the area towards of the season. Some birds returned towards the end of January, would not respond to bait. Efforts will be made to band short prairie geese in 1984. in 1983. the end but they grass ��45 Colorado Division of Wildlife Wildlife Research Report October 1983 JOB PROGRESS REPORT State of Colorado ----------~~------------Migratory vJ-88-R-28 Project No. Work Plan No . 2 Distributional Job Title: Bird Investigations 10 Job No. Characteristics of Some Populations of Canada Geese Inhabiting Colorado Period Covered: Personne 1: 1 May 1982 through 28 February 1983 M. Bauman, L. Budde, G. Bryne, J. Corey, D. Crawford, M. Creamer, L. Crooks, T. Davis, K. Dillinger, J. Ellenberger, M. Etl, H. Funk, J. Frothingham, M. Gardner, J. Gray, J. Gumber, W. Haggerty, R. Kahn, J. Leslie, §: Lorentzson, S. Porter, F. Pusateri, C. Reichert, J. Ringelman, S. Steinert, M. Szymczak, J. Wagner, K. Wagner, P. Will, Colorado Division of Wildlife; G. Patten and staff, Arapaho National Wildlife Refuge; and C. Cesar, Bureau of Land Management. ABSTRACT The number of large Canada geese banded on production/brood rearing areas exceeded quotas in West central Colorado (309), North Park (246), Northeast Colorado (199) and South Park (148). There were no birds banded on production areas in northwest Colorado or on wintering birds in westcentral Colorado. Trapping efforts post-season birds being bahded. in northeast Colorado resulted in 63 large ��47 DISTRIBUTIONAL CHARACTERISTICS OF SOME POPULATIONS OF CANADA GEESE INHABITING COLORADO Gerald Lorentzson P. N. OBJECTIVES 1. To document the wintering range and harvest distribution of Canada geese nesting in (1) northwest Colorado, (2) west central Colorado, (3) North Park, (4) northeast Colorado and (5) South Park. 2. To ascertain the breeding range of Canada geese wintering central and northeast Colorado. 3. To contribute data to the various Central and Pacific Flyway management plans for specific populations of Canada geese. in west SEGMENT OBJECTIVES la. Trap and band at least 150 Canada geese on production central Colorado. areas in west lb. Trap and band at least 150 Canada geese on production areas in northwest Colorado. 1c. Trap and band at least 100 Canada geese on production areas in North Park. 1d. Trap and band at least 150 Canada geese on production east Colorado. 1e. Trap and band at least 75 Canada geese on production areas in South Park. 2. Trap and band 250 Canada geese on wintering Colorado. 3. Trap and band at least 250 Canada geese on wintering areas in northeast Colorado and take the measurements mentioned in the Program Narrat ive. 4. Submit banding schedules and recovery reports to the Bird Banding Laboratory. 5. Prepare progress report. areas in north- areas in west central �48 METHODS AND MATERIALS Canada geese were trapped on production/brood rearing areas in west-central Colorado, northeast Colorado, North Park and South Park during their flightless period using standard drive trapping methods. All captured birds were banded and released. Winter trapping in west-central and northeast Colorado was attempted using cannon nets. Only large Canada geese were banded in northeast Colorado. We did not take physical measurements of banded geese this year. RESULTS AND DISCUSSION Summer Populations Banding quotas were exceeded in 4 of the 5 areas in which summer banding was proposed (Table 1). We were unable to locate concentrations of brood rearing geese in northwestern Colorado. We suspect that these areas exist but so far all concentration of molting geese have proven to be nonbreeding subadult and adult birds. Table 1. The number of Canada geese banded on production/brood areas in Colorado, summer, 1982. Number banded Unknown LF Area/location AM AF LM West-central Colorado Colorado River White River (Meeker) 27 106 _l 27 2 __!l 30 29 119 30 2 5 46 50 E 7 3 10 11 83 87 9 6 4 4 Subtotal North Park McFarlane Res. Case Flats Home Pond Walden Lake Subtotal Northeast Colorado Johnson Pond Haxton Lagoon Guenzi Property Jumbo Annex Subtotal South Park Antero Reservoir Eleven Mile Reservo ir Subtotal rearing Total 120 11 -131 280 29 309 31 12 5 11 -16 44 100 10 31 105 246 36 19 15 9 79 89 37 48 25 17 10 5 21 38 18 19 7 82 199 5 21 26 3 17 20 28 53 81 20 57 77 56 148 204 5 �49 Comparisons of the numbers banded on rearing areas in Colorado are shown in Table 2. Table 2. Summary of geese banded on production/brood rearing areas in Colorado, summers of 1981 and 1982. Number banded West-central Colorado North Park Northeast Colorado South Park 19 1 LM LF AM AF LM 19 2 LF 29 30 29 119 131 309 65 363 83 87 31 44 246 82 93 232 17 21 82 79 199 _2_ 61 _2_ 220 26 20 81 .i: 204 202 223 228 844 156 157 313 313 958 AM AF 3 2 11 13 114 115 69 29 28 45 191 Total Unknown Total Winter PopUlations Efforts to band Canada geese in the winter have been frustrating and relatively unsuccessful. The absence of snow and abundance of feed have made it difficult to attract geese to bait sites. In 1983 there were no geese banded in west-central Colorado and only 63 banded in northeast Colorado (Table 3). Table 3. Canada geese banded in northeastern 1983. Colorado, winter, Number banded AHYF IF AF AHYM Area/location AM 1M Northeast Colorado Johnsons Pond Red Lion 13 2 22 7 2 1 7 1 3 2 3 0 Total 20 3 25 9 5 Prepa red by: 'Cera Id M. Lorentzson ~; �\.r1 o Table 2. Summary of geese banded on production/brood rearing areas in Colorado, summers of 1981 and 1982. Number banded 19 1 West-central Colorado North Park Northeast Colorado South Park --- -- - --- - - ~-- -- 19 2 AM AF LM LF 3 2 11 13 114 115 69 29 28 45 191 AM AF LM LF 29 30 29 119 131 65 363 83 87 31 44 82 93 232 17 21 82 79 57 61 57 220 - -26 -20 -81 77 202 223 228 844 - 156 157 313 313 Total Unknown Total 309 1 246 199 1 204 - 958 �51 Colorado Division of Wildl ife Wildlife Research Report October 1983 JOB PROGRESS REPORT State of Colorado ---------------------------- Project No. Work Plan No. 3 Job Title: Job No. Bird Investigations 8 Monitor Banding of Eastern Colorado Mallard Populations Period Covered: Personnel: Migratory w-88-R-28 30 April 1982 - 31 March 1983 M. Babler, G. Berlin, L. Budde, L. Childers, J. Corey, D. Crawford, M. Creamer, L. Crooks, T. Davis, J. Dennis, K. Dill inger, M. Ette, M. Gardner, R. Kahn, G. Lorentzson, T. Lynch, F. Marcoux, R. Moore, R. Oehlkers, C~ Pabst, F. Pusateri, F. Rinell i, J. Ringelman, M. Szymczak, S. Smith, H. Spear, Colorado Division of Wildlife. ABSTRACT A total of 2,903 mallards were banded postseason in seven locations of eastern Colorado in Segment 28. Some work was accompl ished on an update of analysis of banding data but publication results have been delayed. ��53 MONITOR BANDING OF EASTERN COLORADO WINTERING MALLARD POPULATIONS Gerald M. Lorentzson P. N. OBJECTIVES 1. To establish monitor banding of wintering mallard populations eastern Colorado as an annual management function. in 2. To continually document, through monitor banding and analysis of recovery data, the annual and long-term status of eastern Colorado wintering mallards to provide a basis for annual hunting season recommendations. SEGMENT OBJECTIVES 1. Band a minimum of 4,000 mallards during the post-season period including a minimum of 500 birds in each of the following general areas of the South Platte Valley and Arkansas Valley: (1) Denver-Greeley, (2) Fort Collins-Loveland-Windsor, (3) Greeley-Fort Morgan, (4) Fort Morgan-Sterling, (5) Sterl ing-Julesburg, (6) Bonny Reservoir, (7) Manzanola-Lamar, and (8) Two Buttes Reservoir area. Divide the banded samples in each area equally among the four age and sex classes. 2. Submit banding schedules and recapture data to Bird Banding Laboratory. 3. Conduct an updated analysis of band recovery data, including the following major determinations for important population segments of mallards wintering in eastern Colorado: (1) distribution of harvest, (2) recovery rates, and (3) survival rates. 4. Prepare progress report and publish pertinent findings. METHODS AND MATERIALS Procedures and equipment remained the same as in previous segments. The research section banded the ducks on Bonny Reservoir and the rest of the banding was done by the Southeast and Northeast Regional personnel. RESULTS AND DISCUSSION Only 2,903 mallards were banded postseason during January and February 1983. The areas, Two Buttes and Manzanola-Lamar, only contributed 5 birds to the sample because of inclement weather and the loss of wintering birds on the Arkansas River. Most areas in the Northeastern section were ice-free and birds were not congregated on the usual warm water sloughs. Adult female mallard quotas were difficult to attain, but the four age and sex classes were well represented in the banded sample. All banding schedules and recovery reports were prepared and sent to the Bird Banding Laboratory. �54 Table 1. Mallards banded postseason banding areas, January, 1983. by age and sex in 7 eastern Colorado Number of ducks banded Age and sex Banding area Bonny Reservoir Sterl ing-Julesburg Fort Morgan-Sterl ing Greeley-Fort Morgan Denver-Greeley Fort Collins-Loveland-Windsor Br istol (Arkansas Valley) Totals Updated Analysis AM SM AF SF Total 156 125 128 125 171 128 _3 836 148 125 125 129 136 138 --2 803 93 89 137 70 85 136 138 139 110 62 107 98 610 654 535 478 500 386 499 500 5 2,903 of Band Recovery Data A previous segment report indicated the progress made through Segment 24 in regard to the updated analysis of banding data from eastern Colorado wintering mallard populations (Hopper 1979). This same report discussed the steps followed in the analysis, as well as the quantity of data used. Most of the analysis was finalized during Segment 25, and considerable progress was made in Segments 26, 27 and 28. However, preparation of a final report by way of a publication will be completed during the next segment under Work Plan 6, Job 1 and submitted for publication in an appropriate technical series. LITERATURE CITED Hopper, R. M. 1979. Monitor banding of eastern Colorado wintering mallard populations. Colo. Div. of Wildl. Fed. Aid Game Res. Rept., Oct. Pp. 49-56. Prepared by: �55 Colorado Division of Wildlife Wildlife Research Report October 1983 JOB PROGRESS State of REPORT Colorado --------------------------Migratory ~J-88-R-28 Project No. 3 Work Plan No. 10 Job No. Some Population Job Title: Bird Investigations Characteristics of Adult Gadwall Molting in North Park Period Covered: Personnel: 03 August 1982 to 22 September 1982 J. Corey, J. Ringelman, S. Steinert, G. Tisthbein, and M. Szymczak, Colorado Division of Wildlife. J. Wagner, ABSTRACT Trapping of gadwall (Anas strepera) in North Park resulted in the fol lowing number of birds being banded: adult males = 332; adult females = 191; immature males = 9; immature females = 7; local males = 28; local females = 21. ��57 SOME POPULATION CHARACTERISTICS OF ADULT GADWALL MOLTING IN NORTH PARK Michael R. Szymczak P. N. OBJECTIVES 1. To document the distribution of harvest of gadwall young or molting adults in North Park, Colorado. 2. To estimate recovery and survival North Park, Colorado. banded as fl ightless rates of adult gadwall molting in SEGMENT OBJECTIVES 1. Trap and band 400 adult males, 350 adult females and, if available, up to 100 local gadwall in North Park on molting and brood rearing areas in North Park. 2. Submit banding schedules and recapture reports to the U.S. Fish and Wildlife Service's Bird Banding Laboratory. File return information at the Colorado Division of Wildlife Research Center. 3. Prepare progress report. METHODS AND MATERIALS All birds were captured from an air thrust boat at night using a hand-held 12-volt landing light and a long-handled net. Nearly all fl ightless adults were captured, weighed and measured prior to being banded and released on conjunction with a study of the flightless period in ducks (Szymczak and Ringelman 1983, in press). Thirteen birds were fitted with back-tags or radio transmitters. The band numbers of birds captured that had been previously banded were recorded. Banding schedules and recapture reports were prepared and submitted to the U.S. Fish and Wildlife Service's Bird Banding Laboratory. Information on returning birds recaptured in the same 10-minute grid of banding were filed at the Colorado Division of Wildlife Research Center. RESULTS As in 1981, trapping efforts fell short of producing the number of gadwall needed to meet banding quotas. Walden Reservoir was the major concentration area for molting adults and atypically high water allowed molting birds to disperse throughout the reservoir during night-time operations. MacFarlane Reservoir, which was dry in 1981, contained an average amount of water in 1982 but did not attract molting adult gadwall. �58 Table 1. Number of gadwall banded in North Park, August through September, 1982. Age and sex Location a Walden Reservoir Pole Mountain Reservoir MacFarlane Reservoir Totals AM AF 1M IF LM LF Total 320 6 6 175 10 6 0 9 0 0 7 0 22 6 0 17 4 0 536 42 12 332 191 9 7 28 21 590 aAn additional 6 adult males and 7 adult females were banded, but also were radio-or-backtagged marked. Prepared �Colorado Division of Wfldlife W1ldlife Research Report October 1983 59 JOB PROGRESS REPORT State of Colorado --------~~~------------ Project No. Migratory W-88-R-28/5506X Work Plan No. Job No. 3 Banding of Preseason Waterfowl Job Title: Bird Investigations 11 Populations in the San Luis Valle Period Covered: Personnel: 1 July to 15 November 1982 M. Nail and staff, Monte Vista and Alamosa National Wildl ife Refuges; J. Corey, G. Lorentzson, Colorado Division of Wildlife. ABSTRACT Totals of 1,449 mallards and 785 other ducks were banded in the San Luis Valley during the summer of 1982. Water conditions were better this year in most of the Valley. Quotas were met of all the birds except for adult female mallards. ��61 BANDING OF PRESEASON WATERFOWL POPULATIONS IN THE SAN LUIS VALLEY G. M. Lorentzson P. N. OBJECTIVES The objective of this job is to continually document, through monitor banding and analysis of recovery data, the status of the San Luis Valley preseason mallard population. SEGMENT OBJECTIVES 1. Trap and band at least 300 mallards of each sex and age group, adult males, adult females, immature males and immature females. 2. Band all other waterfowl lard trapping. 3. Submit banding schedules and recapture reports to the U.S. Fish and Wildl ife Services Bird Banding Laboratory, file resulting recovery cards at the Fort Coll ins Research Center. species captured during the period of mal- 4. Prepare Progress Report. METHODS AND MATERIALS Ducks were trapped in the San Luis Valley from August 9 through September 17, 1982. Mallards were the target species but all other ducks captured were banded. Salt Plains types of duck traps were used in the banding program for capturing the birds. Barley was used for bait. Ducks were banded and recorded according to age and sex. All banding reports were sent to the U.S. Fish and Wildl ife Service's Bird Banding Laboratory. RESULTS AND DISCUSSION A total of 2,234 ducks were banded in the San Luis Valley of Colorado during the summer months of 1982. The number and composition of each species are presented in Table 1. The quota of 300 mallards of each age and sex was attained, except for adult females. There appeared to be more water available in 1982 than in 1981 due to an abnormally wet fall. We were not able to trap at San Luis and Head Lakes this year as both of them were dry. Water still seems to be the limiting factor for duck production in the San Luis Valley as more irrigation systems are installed. �62 Table 1. Number of ducks banded, by species, during the pre-season period, 1982.1 in the San Luis Valley Age and sex IF LM Species AM 1M AF Ma 11a rd 416 404 184 422 Blue winged and/or cinnamon teal 17 151 13 Pintail 29 128 52 Redheads LF Total 11 12 1,449 117 2 0 300 20 80 3 0 260 29 26 18 0 0 125 6 25 3 13 10 66 Gadwa 11s 0 15 9 0 0 32 Lesser Scaup 0 0 0 2 25 22 2,234 Green-winged teal Totals 16 0 520 753 247 667 11ncludes ducks banded by personnel of the Alamosa and Monte Vista National Wildlife Refuge. (j ~~ Prepared by: /~ " n /1 {' '//1. tfl' �63 Colorado Division of Wildlife Wildlife Research Report October 1983 JOB PROGRESS State of REPORT Colorado ------~--~--------------- Project No. Work Plan No. 6 ------~--------------- Job Title: Migratory Period Covered: Personne 1: Migratory W-88-R-28 01 Bird Investigations Job No. Bird Publications April 1982 - 31 March 1983 Paul Curtis, Clait Braun, Howard Funk, Jim Ringelman Mike Szymczak. and ABSTRACT Work was continued and near completion on a publication regarding the high mountain duck study. Publications accomplished or submitted under this job during the segment were as follows: Curtis, P. D. Colorado. 1981. An albinistic band-tailed Wes t. Birds 12: 185. Ringelman, J. K. 1983. Solving the puzzle. Outdoors 16 June 1983). Prepared by: ~cuVr-Y;~_h (HOWard D. Funk Wildlife Research Leader pigeon (Submitted in Evergreen, to Colorado ��65 Colorado Division of Wildlife Wi Id life Resea rch Repo rt October 1983 JOB PROGRE~S REPORT State of Colorado --------~~~------------- Project No. Migratory Bird Investigations w-88-R-28 8 Work Plan No. Computerized Job Title: Job No. System for Storing and Retrieval of Banding, Recovery and Recapture Per iod Covered: Personne I: .1 Information 30 April 1982 - 31 March 1983 J. Corey, H. Funk, J. Ringelman, M. Szymczak, Colorado Division of Wildlife; M. Schoenfeld, Colorado State University. ABSTRACT A system was designed to file and access banding, recovery, and recapture data. Over 260,000 bandings and 35,000 recoveries were read from computer tapes, sorted, reformatted, and re-read onto a series of five computer tapes. These data can now be easily accessed. Additionally, about 6,500 "r'etur n" type recaptures have been coded and submitted for keypunching. Banding data are kept current by annual updates facilitated by three computer programs. These programs reformat a short-hand data file of bandings, check for errors, and produce a banding schedule for submission to the Bird Banding Laboratory. ��67 COMPUTERIZED SYSTEM FOR STORING AND RETRIEVAL OF BANDING, RECOVERY, AND RECAPTURE INFORMATION James K. Ringelman Michael R. Szymczak P. N. OBJECTIVE Major objectives of this job are to establish a computer system for banding and recovery data, then to enter all necessary data from past and future banding programs into the system for accurate and efficient record keeping, analysis, and reporting programs. SEGMENT OBJECTIVES 1. Complete the conceptual design of the computerized system. 2. Develop software programs necessary to access, reformat, and update banding and recovery data files. 3. Complete coding and entry of banding tub and "re turn" recapture data. METHODS AND MATERIALS Banding and recovery tapes were obtained from the Bird Banding Laboratory, U.S. Fish and Wildlife Service. All data sets were written on 2400 foot computer tapes stored in the main computer center site at Colorado State University. The "Gold" CSU Cyber computer was used in program design and appl ication. RESULTS AND DISCUSSION Banding Tub Archives Over 260,000 banding records were read from computer tape, sorted by species, location, and date, and read onto three new banding record tapes in the desired format (Table 1). Data are now easily accessed for use in determining the origin of recaptured birds and for use in recovery matrix/ survival rate analysis programs. Banding Update Winter 1982-83 bandings of 4,300 ducks and geese were entered, reformatted, and saved on tape as part of the annual update of banding records. Three programs were developed for use in data input and output. Fortran program �68 BANDIN provides a shorthand means of entering banding data where repetitive fields are frequently encountered. In the shorthand format, changes in age, sex, or both age and sex can be entered without re-entering redundant data fields (Fig. 1, top). Entire lines are entered if changes in other fields (species, location, date, etc.) are needed. Program BANDIN then reformats these shorthand data and writes a complete banding data record (Fig. 1, bottom). To facilitate submission of banding data to the U.S. Fish and Wildlife Service Bird Banding Laboratory, program BSGEN generates a computer-produced banding schedule (Fig. 2). Duplicate data written to tape wi 11 be used for di~ect input into the master banding files at the Bird Banding Lab. Prior to submission of data, program CHECK reviews data files for errors in recording or entry of data. Recovery Archives Nearly 36,000 band recovery records have been read, sorted, and re-read onto two tapes in the desired format (Table 2). Over 24,000 mallard recoveries compose the bulk of recoveries from 1963-81. Annual updates of these tapes will be made from data obtained from the Bird Banding Laboratory. Coding of Return Data "Re tur ns!", or birds recaptured in the same 10-minute grid of banding, have been recoded from field recapture forms and submitted for keypunching. Once keypunched, these return data will be matched-merged with original banding tub data to complete the recovery record. The estimated 6,500 recaptures coded in this manner will be used to supplement data on hunterkilled, banded birds in updated analyses to determine annual survival rates more precisely. Future Work An extension of this work plan will enable coding of return data obtained from the U.S. Fish and Wildl ife Service Refuge Division of recaptures in the San Luis Valley. Additionally, computer programs will be written that (1) produce banding summaries suitable for annual progress reports, (2) search banding tub files for banding origin data on recaptures, and (3) reformat Colorado DOW banding data and writes these data onto tape for direct submission to the Bird Banding Laboratory. Prepared by ~~/}:_: -esK. nge Iman ~ R'i Wildl ife Researcher ,> B �Table 1. Contents of banding Fi 1e # 1 2 3 4 5 6 7 8 9 10 11 12 13 Tape 2. 1 2 Tape 3. 2 3 4 5 6 7 8 # Records File contents 1. Mallard Tape tub tapes 1, 2, and 3. bandings, 1967-79. 1967 1968 1969 1970 1971 1972 1973 1974 1975 1976 1977 1978 1979 11 ,656 11,417 10,080 8,751 9,377 9,984 9,753 10,815 12,622 11,653 10,921 12,815 10,956 1980 1981 (part) 10,945 5,473 Combined blue-winged teal and cinnamon teal bandings, 1967-81 Green-winged teal bandings, 1967-81 Gadwall bandings, 1967-81 Pintail bandings, 1967-81 Small Canada goose bandings, 1967-81 Large Canage goose bandings, 1967-81 Redhead bandings, 1967-81 Wigeon bandings, 1967-81 11,115 19,853 5,393 40,350 9,349 13,627 2,253 1,337 Mallard Mallard Mallard Mallard Mallard Mallard Mallard Mallard Mallard Mallard Mallard Mallard Mallard Mal lard bandings, bandings, bandings, bandings, bandings, bandings, bandings, bandings, bandings, bandings, bandings, bandings, bandings, bandings, 1980-81. Mallard Mallard bandings, bandings, Other duck and goose bandings. �70 Table 2. Fi 1e # Tape 4. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 Tape 5. 1 2 3 4 5 6 7 8 9 Contents of recovery tapes 4 and 5. Recovery year(s) Mallard recoveries, # Recoveries 1963-81. 3,224 3,041 1,657 1,769 1,523 1,689 1,488 1,146 1,541 1,356 1,457 1,547 1,398 1,166 1963-67 1967-69 1969-70 1970-71 1971-72 1972-73 1973-74 1974-75 1975-76 1976-77 1977-78 1978-79 1979- 80 1980-81 (in part ?) Other duck and goose recoveries. Blue-winged teal recoveries, 1963-1981 Green-winged teal recoveries, 1963-1981 Gadwall recoveries, 1963-1981 Pintail recoveries, 1963-1981 Small Canada Goose recoveries, 1963-1981 Large Canada Goose recoveries, 1963-1981 Blue-winged and cinnamon teal recoveries, Redhead recoveries, 1963-1981 Wigeon recoveries, 1963-1981 1963-1981 304 896 484 3,037 2,503 3,409 332 275 162 �71 Figure 1. Short-hand banding data file (top) and completed banding record (bottom) produced by computer program BANDIN (data files reduced in scale). -08001' Nl' T3770450i o 002 . 00003 00004 A SY 00005 AASY 00000'00007 NL 137704507 00008 00009 A SY~00010 AA5Y 1320300 ASY~-'1 4215 '012082 132J 300 ASY M 4215 012082 REPLAClS 1077-5b2~5 REPLAC~S 777-93756 888H 00013 ,nn4 00015 oco i e 5 F A 5Y At.SY A SY AASY'888H 00019 A 5Y 00020 AASY 00021 p.;L 137704521 1320 00022 S M 00023 00024 A SY 0002S AASY ooo z e A SY AAS Y 00027 00028 00029 00030'NL 137704530-1320 00031 A SY 00032 AASY 00033 00034 NL . 137704534 1370 00035 AASY 00036 00037 .. 00038' :;--"-F" 00039 00040 ilL 137704540 1320 i'I 00041 S 00042 00043 AASY 00044 A,Y 00045 AASY 00046 00047 NL 137704547 1320 00048 00;)49 00050 300 ASY M 4102 012082 300 A5Y F 4102 012082 300' ASY M 4102 012082 300 ASY F 3601 0120a2 300 ..SY F 2802 012062 88gg~~~BH8t~8rTrE8'~8f1H ~ tH~ 8r:~8:g~ 137704503 132.0 300 ASY M 4215 01-20-e2 gggg;~.HHg~§g~ iH:8 ~~8 AH ~ 4H§ 81:~g:g~ 00003 00006 -137704506 00007 137704507 00008 137704508 00009 137704509 00010 137704510 00011 137704511 00012 137704512 132.0 30,) A5Y 132.0 300 ASY 132.0 30C A5Y 132.0 300 SY 132.0 300 ASY 132'8 300 ASY 132. 3~O A5Y M M M M M M M 4215 4215 4215 4215 4215 4215 4215 01-20-62 01-20-~2 01-20-82 01-20-~2 01-20-62 01-20-82 01-20-02 REPLACES 1077-56295 gggi~ i~tt8~§l21~~:8 ~3g i~~~4~i§ 8r:~g:§~ 00015 137704515 132.0 30~ SY F 4215 01-2C-b2 88819·i~t;8=~i9 i~~:8~88 Ai~ ~ ~it~ 8t:~8:~~ 00018 137704518 132.0 300 ASf F 4215 81-20-e2 00019 137704519 00020 137704520 00021 137704521 '00022 137704522 00023 137704523 00024 1377C4524 00.)25 "137704525 00026'- 137704526 00027 137704527 132.0 300 SY F 4215 1-20-62 132.0 300 ASf F 4215 01-20-e2 132.0 300 ~SY M 4102 01-20-82 132.0 300 ASf ~ 4102 01-2C-82 132.0 300 A~Y H 4102 01-20-62 132.0 300 SY M 4102 01-20-E2 132.0 300 liSY M 4102 'H-20-82 132.0 300 SY M 4102 01-20-62 132.0 300 ASY M 4102 01-20-62 883~~ 11ttg4~~Sl~~:g~38 ~~~ ~ ~I82 81:28:~~ 300 ASY F 4102 01-20-E2 REPLACES '00030 137704530 132.0 00031 137704531 132.0 00032 1?7704532 132.0 00033 137704533 132.0 00034 137704534 137.0 00035 137704535 137.0 00036 137704536 137.0 OC037 _.137704537'137.'> 00033 - 1377C4538 137.0 00039 137704539 137.0 00040 137704540 132.0 00041' 137704541 132.0 0004~~ 137704542 132.0 00043 137704543 132.0 0044 137704544 132.0 0045 137704545 132.0 00046 137704546 132.0 g 300 300 3JO 300 300 300 300 300 300 300 300 300 300 300 30J 300 ASY F Sf F tSY F ~5Y M ISY M ASY M A5Y M ASY F ASY F ASY F ASY M A5Y M ~$Y M Sf M ASY M A5Y M 4102 4102 41~2 4102 4102 41~2 4102 4102 4102 3601 3601 3601 3601 360~ 360~ 3601 01-20-62 01-20-€2 01-20-l2 01-20-82 01-20-e2 01-20-02 01-20-E2 01-20-82 Ol-20-a2 01-20-f2 01-20-t2 01-20-62 01-2u-82 0~-20.a2 OL-20-22 01-20-b2 gg5=~ 137704549 l§t~g~~~~i~~:g~gg ~~ ~ ~~g~ gl:~g:g~ 132.0 30) SY F 28)2 01-20-62 OG049 00050' 137704550 132.0 300 SY F_2b02 Ol-2W-82 777-93756 �72 BAN HASTER MASTER D I ~ G S C H E. U l j.) E ..-~-- PER~IT NO' 06529 PERMITTEE' COluRADO --- ------- -. BAND NOS DIVISION OF WILDLIFE 1377-04~01 THRU 04550 SANDING lOCATIONS A WINDSQR, .ElD CO., CO D :lOULDER, BOULDER CO., CO B P~EwlTT RESeRVOIR, ~ASHINGTJN CD.,CO E C FORT COLLINS, LARIMER CO., CO F BAND NUMBER COMMO:~ NA:-Ic AOJ STATUS AGE SEX ~"GIJN lAT-LJNG LOC DATE 1377<----MO-DAY-YR 04501 MALhARD 13~.0 300 A?oY ~ 317 "v2~1045 ~ 01-2~-82 C2 03 04 05 06 07 ~EPLACES g~ --- -. --- 1077-56295 ,",- H 15 •• ".. SY ASY " f,SY SY ASY SY ASY .... " ~3 26 27 28 n 3J REPLACES .. .... .... " " .. AMERICAN .. wIGeON .. 137.0- .... 36 --------- ------ 37 38 39 40 41 "2 -It 3 "4 45 46 47- ~g ------ 04550 -~~9AI~.lA~E; .30 REPlAf.~ Figure 2. .. .... .." " .." .... - .. ".. - .... .... .... .... .... .. .... .. " .... 402-1032 H F SY ASY .. .. .... .. .. M ".. " ".. .. .... .... .... .... .. C M .. F -.. .. I' ft " .... .. 41)0-1051 .... .. F SY A~Y SY It .. .... It ".. " 777-93756 MAL hARD .. .... " .... .... .. F " H 23 .. It .... ..•• SY .. .... " " 16 17 ld 1" 20 33 34 35 SY ASY .... MALhA~O 10 11 12 ~~ INCLUSIVE .... 0 .... .... 1077-56295 777_Q<7~~ Computer-produced banding schedule of data in Figure 1. Schedule produced by program BSGEN (schedule reduced in scale). �73 Colorado Division of Wildlife Wfldlife Research Report October 1983 JOB FINAL REPORT State of Colorado --------------------------- Project No. Work Plan No. 9 ---------------------- Job Title: Migratory Period Covered: Personnel: Migratory W-88-R-28 Bird Investigations Job No. Bird Research Planning 01 April 1981 through 30 March 1983 Howard Funk, Gerald Lorentzson, Jim Ringelman and Mike Szymczak ABSTRACT Planning associated with this job was in association with new starts during the 2-year period and for the long-range period plus the Strategic Plan. Involved were an aborted long-range plan (10 yrs) due to alterations in Division policy, a Research Operations Plan which was altered by policy to a Program Plan (S-yr period) as part of the Strategic Plan (also a S-yr period). The Strategic Plan is in operation while the Program Plan is being reviewed for final write-up and inclusion with other Division plans, probably in 1983. ��75 MIGRATORY BIRD RESEARCH PLANNING Howard D. Funk P. N. OBJECTIVES To prepare long-range research plans for w-88-R for a period of 10 years. METHODS AND MATERIALS During the period covered, time was spent in reviewing past accomplishments of the Migratory Bird Section, examining needs for future research needs, review of literature, correspondence and personal contacts with other migratory bird researchers on ongoing projects and discussion of needs with them, and in personal contact with Division of Wildlife (DOW) sections and individuals in search for their feelings on needs for migratory game bird research and/or management results. Also considered in the way of research needs were results of Technical Committee and species management plan sub-committee results and recommendations. Section personnel then assembled the results, made recommendations on specific research jobs, prepared plans and submitted them through the DOW review process. RESULTS Initial plans and efforts to prepare long-range (10 year) plans were altered by DOW pol icy in the period covered to shorter range plans of 5 years. The thrust was to incorporate general research needs in the revised Strategic Plan (5 years) which was made operational during the period covered by this job (Today1s Strategy ... Tomorrow1s Wildlife, January, 1983, Third Edition, Colorado Division of Wildlife). Input into this effort was made by section personnel in the way of suggestions for research and management needs and by solicitation of the same items from other sections and individuals. Meetings were held in the various regions to obtain ideas and discuss alternatives. Upon submitt~ng these to Denver, these were incorporated with other section data, assembled, sent out for review, revised, and priorities set by species and strategy by Region. The Strategic Plan was approved by the Commission and is established as the general guide for activities through the 5-year period. Section personnel then participated in establishing an Operations Plan, which was to be a step down from the Strategic Plan. This included information and plans, by job, for the section including Job Title, Problem, Status, Objectives, Implementation and Benefits. Also included were Operations Plan Strategies, Activity, Product Analysis and Deadline in formation as well as cost estimates and personnel needs. These were completed, including recommendations for new jobs during the 5-year period, in early 1983. �76 Alterations in DOW plans excluded the Operations Plan as such with the substitute being the Program Plan as the step down from the Strategic Plan. Thus, Operations Plans were condensed, in some categories made more general, and incorporated into the Program Plan. Besides research and management needs and ongoing programs in the section, more general needs and activities were placed in the Program Plan, including such items as recommendations for habitat purchase, easements, and improvement as well as specific recommendations on programs conducted by other Sections and the Regions in the way of improved public relations, hunter education, special surveys, cooperative management efforts, etc. The draft Program Plan data were submitted for review by other sections and the Regions and additional effort was then expended to make format similar in all aspects, but also to incorporate additional specifics into the research portions; somewhat more specific than initially but not as specific as the Operations Plan. The Program Plan is under review, as of the end of Segment 28, and is to be finalized in 1983. Thus, the overall guide, as stated previously, is the Strategic Plan and the first step down is the Program Plan. Section personnel contributed to these plans with information on ongoing plans and recommendations for new jobs in research and other management related activities. The Implementation Plan, the next step down, in the case of the Research Sections, will be a continuation of the Program Narratives as has been the procedure in the past. () Submitted by: )b9c~x;e_-e£c¥~ Howard D. Funk Wildlife Research Leader �77 Colorado Division of Wildlife Wildl ife Research Report October 1983 JOB PROGRESS REPORT State of Colorado Project No. W-88-R-28 Work Plan No. 10 job ----~~------------ Job Title: Cooperative Period Covered: Personnel: Migratory Management Bird Investigations No. Programs 01 April 1982 - 31 March 1983 john Corey, Howard Funk, Gerald Lorentzson, jim Ringelman, Steve Steinert, Mike Szymczak, Colorado Division of Wildl ife. ABSTRACT Work accompl ished during this segment consisted tive management programs with the four regions in the Central Flyway. Help was also given to Service consistent with the objectives of this of assistance in cooperaof Colorado and the states the U.S. Fish and Wildlife work plan. ��79 COOPERATIVE MANAGEMENT PROGRAMS Gerald M. Lorentzson P. N. OBJECTIVES The major sections matters, upon the objective of this job is to provide 1iaison with the various DOW in order to assist and work with them in migratory game bird particularly those relating to management responsibil ities placed various sections. SEGMENT OBJECTIVES The procedures for this job will be to provide liaison and expertise to the Regions and other DOW entities in assisting them with management activities for which they have been assigned main or partial responsibilities. This assistance will be given in the way of training and/or help in design of cooperative management programs such as monitor banding and various migratory game bird surveys. Assistance will be given in design of goose transplant programs and monitoring of results. Procedures for federal hunting season requirements and 1imited permit hunts will be explained and assistance given in setting up seasons and monitoring results. As necessary, assistance will be given in sampl ing schemes, hunter surveys and statistical analyses of results. Cooperative effort will be suppl ied in preparation of strategic plans for migratory game birds. Project personnel will take part in various technical committees, status and regulations meetings and management plan workshops both instate and out of state as necessary for cooperative research and/or management-oriented purposes. METHODS AND MATERIALS Due to the diverse nature of this program, various procedures and material were used to accomplish the objectives. The methods and materials used for trapping, banding, inventory studies, and the compilation of harvest data have been well documented in other segments of this program and will not be repeated in this report. RESULTS During this segment assistance was given to the Regions in the areas of breeding pair counts in the San Luis Valley and along the Snake and Yampa River in northwestern Colorado. Assistance was also given for the goose production surveys and the December and January inventories taken on waterfowl. Many contacts were handled regarding goose nesting structures and other day to day problems concerned with waterfowl. �80 Several training sessions and other informative regional personnel and other community-oriented meetings were held for organizations. We did another study for the Central Flyway Technical Pesticide Committee regarding the use of chlorinated hydrocarbons in agricultural regions of Colorado. Various species of ducks were collected and tested for the presence of chlorinated hydrocarbons or their residues, in the body fat of these ducks. Considerable work was accomplished on the Strategic Plan for the Division of Wildlife and initial work on the Operations Plan was begun. Work was also started on the goose transplant site inspection plan. Aid was given to the Fish and Wildl ife Service in collecting and speciating the duck wings for their waterfowl parts collection program. Help was given them setting up material for the Wing Bee and for actual assistance during the Wing Bee. Cooperative efforts with other states and the Central and Pacific Flyway included administration of Colorado1s dove call-count routes and submission of data, the Rocky Mountain Canada Goose Sub-committee, the Central Management Unit Technical Committee Meeting, the spring and summer meetings of the Central Flyway Waterfowl Technical Committee and Central Flyway Council meetings, and the Subcommittee on Nesting Studies in the Dakotas and eastern Montana as well as the Research Needs Sub-committee. Bands and banding materials were distributed to the regional personnel and banding records forwarded to the Bird Banding Laboratory. Mallards banded in west-central Colorado are reported in Table 1. Birds banded in other areas are reported in other work plans in this segment. Table 1. 1982. Mallards banded in the west-central Area De 1 ta Walker Wildl ife Area Mack (Hi Line Lake) Total portion of Colorado, AM SM AF SF Total 102 17 83 101 30 92 13 .is ...22. 105 23 109 400 83 317 202 201 160 237 800 ~. -4/L-c~_,<.'_____; Gerald M. Loreritzson h Senior..,Wildlife Biologist k:o-<iJr9IL,J;;_ I'fowa--rd D. Fun k Wildlife Research January Leader � https://cpw.cvlcollections.org/files/original/28ce06b82e7e16c234734822c9256404.pdf 8a49e04e6d89554132d8df49ae38ffa0 PDF Text Text Colorado Division of Wildlife Wildlife Research Report January 1984 1 JOB PROGRESS REPORT State of Colorado --~~~~~------- Project N-1-R-2 (SE-3-5) Work Plan 1 (II) Job 1 (I) Period Covered: Author: Nesting Performance of Peregrine Falcons iIi.Colorado 1 January - 31 December 1983 G. R. Craig Personnel: D. Berger and G. Craig, Colorado Division of Wildlife; J. Enderson, The Colorado College. ABSTRACT In 1983, the number of occupied peregrine falcon (Falco peregrinus anatum) breeding territories increased to 13. Released falcons were documented as members of pairs at 3 territories and an adult male released in 1976 was found at a recently discovered nesting site. Eggshell thickness measurements were obtained from 27 wild eggs removed during augmentation efforts and 8 nonviable eggs have been submitted for pesticide analysis. An annual adult survivorship of .078 was obtained using telephotographic documentation of returning breeding adults. This Job Progress Report represents a preliminary analysis and is subject to ch&Lge. For this reason~ information presented herein ¥iAY NOT BE PUBLISHED OR QUOTED without permission of the Author .. ��3 NESTING PERFORMANCE OF PEREGRINE FALCONS IN COLORADO Gerald R. Craig P. N. OBJECTIVE The objectives of this study are to annually monitor breeding numbers and reproduction of Colorado peregrine falcons to document further population declines as well as record the population's responses to recovery efforts. Additionally, health of the population will be monitored indirectly by analyzing pesticide residue levels in the falconsv eggs and principal prey. Information obtained from these investigations will be made available through annual reports to the Rocky Mountain/ Southwest Peregrine Falcon Recovery Team as well as cooperating agencies to aid in evaluation of recovery efforts. SEGMENT OBJECTIVE I. Annually monitor the number of breeding pairs of peregrines and reproduction in Colorado. 2. Annually monitor organochlorine pesticide levels in wild breeding peregrines. 3. Monitor recruitment of reintroduced peregrines into the wild breeding population of Colorado. 4. Compile data and submit reports to appropriate state and federal personnel and the Rocky Mountain/Southwest Peregrine Falcon Recovery Team for use in evaluating recovery efforts. (These objectives correspond to Jobs 1113.,1114., 212.9 221.~ .121.L" 321209 adn 333. of the approved American Peregrine Falcon Recovcrv r~I.RX'~ for the Rocky Mountain/Southwest Population). METHODS AND MATERIALS Methods and materials used in this study have been described prf..v:<:'p'LcJ) (Craig and Enderson 1981). RESULTS AND DISCUSSION Territory Occupancy The number of occupied breeding territories increased from 11 in 1982 to 13 in 1983 (Table 1). Pairs returned to 6 of the sites occupied in 1982. 2 sdtes frequented by lone adults in 1982 were vacant , 2 former'Ly �4 vacant sites were reoccupied, and 2 previously undocumented territor:ies were discovered. The 2 additional occupied breeding territories located in 1983 incrL~oLd the total territories recorded for Colorado to 36. It is probable that site 38 was occupied historically since the habitat is ideal for peregrinel" but until recently, the landowner prohibited access to the area. A caretaker encountered the male about 1 mile from the nest cliff, dying of injuries suffered from a powerline collision. The Division was contacted.snd subsequent investigations located the female on the nest cliff. The 2nd new territory (site 39) was reported late in the breeding season and although adults were observed, no young had been produced. It is possible that this pair relocated from a release site they were frequenting in 1982. Pairs returned to sites 8, 25, 30, 31, and 34 in 1983 and all successfully fledged young. Although both adults and their brood disappeared from site 9 in 19829 an adult male and yearling female reoccupied the territory in 1983. The female at site 34 successfully attracted another adult male when her previous mate failed to return. Territory 27, which had been frequented by a yearling male the previous year, was inhabitated by.a pair which successfully fledged young. A pair of yearling peregrines was observed at site 5 which was vacant in 1982. After a 7-year hiatus, site 3 was reoccupied by an adult male (released in 1981) and a yearling female. Sufficient numbers of peregrines were present in the wild to replace 1 pair lost at 1 site, a male at another, and to reoccupy 2 vacant territories. Unattached peregrines observed interacting with breeding pairs at sites 30,. 34, and 35 are also indicative of population recovery. Reproduction Reproductive success in 1983 was good (Table 1). In all, 7 pair~ were conf Lrmed to have laid eggs and all were manipulated to auglh<l< reproduction. It is possible that falcons at sites 38 and 39 may L,: ro, prod cod eggs but these sites were located too late in the breedau., season to confirm incubation behavior. In all, 27 eggs were removed from the 7 wild nests (3.86 eggs/clutch) for artificial incubati .IL Seve.ra.L eggs were dented or cracked and others exhibited exceas lve l'later lose, all indicative of poor shell condition. A to .4.. or IS young was successfully hatched from wild eggs and returned to t.he'\Jld. Undoi .•btedly, reproduction would have been less had the eggs been left: with the wild adults to be incubated naturally. In 1983, 27 captivehatched young were placed at the 7 wild sites (3.86 young/pair) and 21 :3uccesl:lfu ly fledged (3.00 young/pair). Golden eagle (Aquila chrysaetos) predation probably accounted for the death of 1 young at site 8. Five young (1 at site 25~ 2 at site 31, and 2 at site 35) disappeared from 3 other broods prior to fledging but the cause of mortality could not be determined. �Table 1. Occupancy and productivity of peregrine falcon eyries in Colorado~ 1972-83. Parameter 1972 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 Known territories ~2 22 22 221 25 29 31 32 34 34 36, 38 Occupied territories 11 12 9 ''7 8 12 11 12 13 11 11 13 Adult pairs 8 11 7 5 5 11 7 6 7 7 9 9 Immature pairsa 0 0 0 1 2 0 2 2 3 1 1 3 Lone adults 3 1 2 1 1 1 2 3 3 3 3 1 Successful pairs 0 1 5 2 4 6 5 4 5 6 6 7 No. ~ng. fledged No. yng. augmented 0/0 2/0 13/2 5/2 6/4 11/5 16/15 12/12 16/13 15/21 20/26 21/27 Young fledged/ adult pairb 0.00 0.18 1.57 0.83 1.20 1.00 2.29 2.00 2.29 2.14 ,2.22 2.33 Young fledged/ successful pair 0.00 2.00 2.60 2.50 1.50 1.83 3.20 3.00 3.20 2.50 3.33 3.00 Sites occupied~ %c 50 55 41 32 32 41 35 38 38 32 31 34 Site w/adult pairs, % 53 49 29 27 26 35 22 18 24 21 26 26 aAt least ! member' of the pair was in juvenile plumage. bl. 25 young fledgei/p&.ir is 'coris Lde'red normal reproduction. cFor a normally reprod.llci~,g pcpul.a t ton , 80-90% of the eyrie sites should normally be occupied in any particular yearc \J �6 ~ggshell Thickness Shell t.hd.ckneaa measurements obtained. from 19 of the 27 eggs taken from the wild in 1983 averaged 0.316 mm (with membrane). This average is approximately 12% thinner than pre-DDT era eggshells which is a 3% decrease in thickness from 1982. This is the first evidence of downward movement in an improving trend in shell thickness first observed in 1977. It is possible that the addition of shell thickness measurements from the 8 eggs sent to Patuxent for pesticide analysis may improve the average. Organochlorine Residues in Egg Contents Pesticide residue analysis was performed by the U.S. Fish and Wildlife at the Patuxent Wildlife Research Center on contents of 6 nonviable eggs collected in 1982. DDE residues averaged (arithmetically) 22.06 ppm (range 7.11-32.09) fresh wet weight basis in the 6 eggs. While this limited sample represents 4 sites (89 9. 34. and 35), there was an overall increase 1.nresidue levels from a sample of 8 eggs collected in 1981 (13.7 ppm arithmetic mean). At the same time, eggshell thickness increased from an average of 0.318 mm in 1981 to 0.323 in 1982. Due to the limited sample of eggs subjected to residue analysis, more credance should be given to the statistically larger sample of thickness measurements which continue to demonstrate an improving trend toward thicker eggshells. One sobering note from the 1982 pesticide analysis was that a known-age female hatched in 1979 accumulated critical levels of 32 ppm DDE in her eggs within 3 years. An additional 8 nonviable eggs collected in 1983 were submitted to Patuxent for pesticide residue analysis, but due to a backlog as well 'as reprioritization, work had not been undertaken at the time of this report. -, ~ecruitment of Released Peregrines into the Wild A banm. 10 released at a nearby hack site in 1980 returned. t.O I8 for the Znd yar t.omate with the wild female which was also pr. se .,,:; in 1982. The female was sufficiently aggressive to be caught at t.ht:: eyrIe in 1982 and was banded. In 1983. the plastic marker was no Longer af ILxcd to her leg" but the U. s. Fish and Wildlife Service band I'tnk . _.0.1 at tac.ned, A male released at a hack site adjacent to site 11 In 19(-;. l.ras paired lvith an unhanded (wild-p'roduced) yearling fema Le at: ch::('A. ; approximate.ly 48 kID. west of his release site. The most significant event was discovery of a banded male at site 38. Th:ts tiercel appears to have been breeding at that location for several years. In 19769 this male was fostered into a brood at site 7 about 29 km cast of site 38. It survived 7 years in the wild before dying after a collision with a power line. Word was also received that a falcon hacked at site 36 in 1982 was observed paired w.th a wild adult ma.le at an eyrie in New Mexico and subsequent.Ly �7 produced young. A male released from the same site in 1982 returned and fed with the young released there in 1983. A 3rd female falcon released from site 36 in 1982 was trapped at Matamoras, Tamaulipas, Mexico, south of Brownsville, Texas. Finally, a female released at site 11 in 1982 was observed interacting with young released at a hack site in Wyoming in June 1983. Photographic Documentation of Wild·Adults Photographs were obtained of 13 adults occupying territories in 1983 bringing to a total 43 useable photographs of individual adults since 1980. Overall, the data includes 28 instances where it could be ascertained that an adult returned in a subsequent year and there are 4 more instances where it is near-certain that determinations could be made. Of the 28 instances, 21 adults were alive the following year. The survivorship rate was 0.75 per year and if the 4 near-certain determinations are included, the survivorship is 0.78. In 1984, there is the potential of increasing the sample size from 32 to 46. LITERATURE CITED Craig, G. R., and J. H. Enderson. 1981. Nesting performance of peregrine falcons in Colorado. Job Prog. Rep., Colo. Div. Wildl., Wildl. Res. Rep., Jan., pp 13-23. Prepared by (] (l ~ Gerai~(R. Cra Wildlife Researcher C ��9 Colorado Division of Wildlife Wildlife Research Report January 1984 JOB PROGRESS REPORT State of Colorado N-I-R-2 Project (SE-3-5) Work Plan 1 (II) Job 2 (2) Period Covered: Author: Reintroduction and Augmentation of Peregrine Falcon Production 1 January - 31 December 1983 G. R. Craig Personnel: D. Berger, G. Craig, T. Fowler, B. Grebence, R. Meese, and M. Robert, Colorado Division of Wildlife; J. Enderson, Colorado College; J. Hogan and S. Petersburg, U.S. Department of Interior, National Park Service. ABSTRACT Fledging success of 7 wild breeding pairs of peregrine falcons (Falco peregrinus anatum) was increased to 3.00 young/pair through augmentation efforts in 1983. The 1983 hacking effort displayed remarkable success when 23 of 25 young reached independence from 6 release sites. Hacked young successfully bred at 1 site in New Mexico and 1 site in Colorado and an adult male released in 1981 paired with an immature female at an historic nesting cliff. Tnis Job Progress Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the Author. ��11 REINTRODUCTION AND AUGMENTATION OF PEREGRINE FALCON PRODUCTION Gerald R. Craig P. N. OBJECTIVE The objective of this program is to sustain the wild breeding peregrine falcon population in Colorado through augmentation of poor natural production and re-establishment of breeding pairs at vacant sites by release of captive-produced falcons. SEGMENT OBJECTIVES 1. Augment poor wild production by placement of captive-hatched wild young and captive-produced young into occupied wild nests. 2. Release captive-hatched wild young and captive-produced young to the wild through "hacking" at potential and abandoned wild nests. 3. Develop and implement release of adult falcons at potential or abandoned nest sites and at wild nests occupied by lone adults. 4. Monitor results of the efforts, compile data, and submit reports to appropriate state and federal agencies and the Rocky Mountain/ Southwest Peregrine Falcon Recovery Team. (These objectives correspond to Jobs 222., 3133., 321., 322., 331., 332., and 3211. in the approved American Peregrine Falcon Recovery Plan for the Rocky Mountain/Southwest Population). METHODS AND MATERIALS Methods and materials for this study have' been described pnevdou: <. '/ (Craig 1982). RESULTS AND DISCUSSION Augmentation Efforts The 7 wild breeding pairs of peregrines encountered in the 1983 season (sites 8, 25. 27. 30. 31, 34. and 35) were manipulated to increase produc t.Lv-tt.y . One other occupied territory (38) was discovered during in.cubation. but the male had been killed and the decision was made not to attempt augmentation. No sites were recycled since The Peregrine Fund anticipated that captive production was sufficient to provide young for the augmentation effort. In retrospect. it might have been advantageous to recycle several sites to bring them into synchron.y with captive production. Thus, young of the proper age (at least 18 �12 days of age) were not available and sites 31, 34, and 35 had to be delayed with broods of prairie falcon (Falco mexicanus) chicks for 13 days until the captive-hatched peregrines were the proper age for placement in the wild. A total of 27 eggs were removed from 7 breeding pairs (3.86 eggs/pair) for artificial incubation at The Peregrine Fund's Fort Collins facilities and 27 captive-hatched young were replaced in the 7 nests (3.86 young/ pair). The 7 pairs fledged 21 young for a success of 3.00 young/pair. Five nestlings were lost from 3 broods due to unknown causes. A 6th fledgling disappeared (site 9) at time of fledging, possibly due to golden eagle (Aquila chrysaetos) predation. Manipulation of wild eggs and broods obscured the nesting success the peregrines would have experienced had they been permitted to reproduce naturally. It is possible. however, to develop an estimate of natural reproduction based upon: (1) hatching success of eggs brought into the laboratory, and (2) nestling mortality experienced in the wild during the brood rearing phase. Thus, it can be assumed that in the wild, only 18 of the 27 eggs would have hatched and 6 nestlings would have died, yielding an expected natural fledging success of 12 young or 1.71 young/ successful pair. This estimated natural fledging success is optimistic since the laboratory treatment of eggs probably increased hatching success and nestling exposure to mortality was reduced because young were placed in the wild at 20-24 days of age. Hacking Efforts Hack sites were activated at 6 locations throughout the state in 1983. The releases experienced excellent success with 23 young reaching independence of 25 released. The 2 failures were caused by great horned owl (Bubo virginianus) predation at site 11. One young from site 8 was recovered about 48 km away with a broken wing about 1 month <.t~~t,,',." ind.epe.ndence. Evi.dp..nce of hacking success began to accumulate in 1983. One felMi ...: rcle.llsedat site 36 in 1981 bred with a wild male and successfully reared young at a wild site in northern New Mexico. A male released i.n 1981 at site 11 .pairedwith an unbanded female at site 3 (45 Ian lj_.8..:'un.:.:j The n.aLe released at site 8 in 1980 returned a 2nd year 9 p fred wi.th tb·· same wild adult female, and fledged 3 young. Another f emaLe nL(:·;~.s ' at. site 11 in 1982 was observed interacting with recently hs cb'··.~ V(~') ,. at a Wyoming site in 1983. It is also probable that at least ].membLi of the newly discovered pair which were observed late in the season at ;-;'tr '39 may have been released at site 36 about 32 km away. Adult Release Baaed upon the poor results encountered in 1981 and 1982, release of adult falcons was not implemented in 1983. , �13 LITERATURE CITED Craig, G. R. 1982. Reintroduction and augmentation of peregrine falcon production, Job Prog. Rep., Colo. Div. Wildl. Res. Rep., Jan., pp 53-57. Prepared by er C ��15 Colorado Division of Wildlife Wildlife Research Report January 1984 JOB PROGRESS REPORT State of Project Colorado N-1-R-2 Work Plan 1 (II) Job 3 (3) Period Covered: Author: Peregrine Falcon Captive Maintenance (SE-3-5) 1 January ~ 31 December 1983 G. R. Craig and W. Burnham Personnel: W. Burnham, D. Konkel, C. Sandfort, E. Levine, G. Eitemiller, The Peregrine Fund, Inc.; G. Craig, Colorado Division of Wildlife. ABSTRACT Due to funding restriction, this job was not activated in 1983 and probably will be discontinued in the future. This Job Progress Report represents a preliminary analysis and is subject to change. For this reason, information presented herein y~y NOT BE PUBLISHED OR QUOTED without permission of the Author. ��17 PEREGRINE FALCON CAPTIVE MAINTENANCE Gerald R. Craig and William Burnham P. N. OBJECTIVE The objective of this program is to maintain a colony of adult peregrine falcons (Falco peregrinus ana tum) which is genetically representative of the wild population, free of disease, and capable of reproductive activity. SEGMENT OBJECTIVES 1. Annually contract with The Peregrine Fund, Inc. to maintain 35 adult ana tum peregrine falcons in captivity. 2. Prepare an annual report of maintenance the contract. efforts associated with METHODS AND MATERIALS Methods and materials for this study have been described previously (Burnham and Craig 1981). RESULTS AND DISCUSSION The study was discontinued in 1983 due to funding restrictions lack of Section 6 monies as well as decreased state revenues. caused by LITERATURE CITED Burnham, W., and G. R. Craig. 1981. Peregrine falcon captive madrrtonance , Job Prog. Rep., Colo. Div. Wildl., Wildl. Res. Rep., Jan.» pp 34~37. Prepared by Gerald R. Craig ( Wildlife Researcher C ��19 Colorado Division of Wildlife Wildlife Research Report January 1984 JOB FINAL REPORT State of Colorado Project SE-3 Endangered Wildlife Investigations Work Plan III Job 3 Period Covered: Author: Peregrine Falcon Captive Maintenance 1 July 1979-31 December 1982 Gerald R. Craig ABSTRACT From 1979 through 1982, 35 adult peregrine falcons were maintained at the Fort Collins facilities of The Peregrine Fund, Inc. for captive propagation. Due to funding restrictions, the job was suspended in 1983 and subsequently terminated. ��21 PEREGRINE FALCON CAPTIVE MAINTENANCE Gerald R. Craig P. N. OBJECTIVE The objective of this program is to maintain a colony of adult peregrine falcons (Falco peregrinus ana tum) which is genetically representative of the wild population, free of disease and capable of reproductive activity. SEGMENT OBJECTIVES 1. Annually contract with The Peregrine Fund, Inc. to maintain 35 adult ana tum peregrine falcon in captivity. 2. Prepare an annual report of maintenance contract. efforts associated with the METHODS AND MATERIALS Methods and materials for this study have been described previously (Burnham and Craig 1981). RESULTS AND DISCUSSION Captive propagatipn was recognized as an intergral component of the peregrine falcon recovery effort in the approved Rocky Mountain/Southwest Peregrine Falcon Recovery Plan. The Recovery Team recommended that The Peregrine Fund's w_estern program be funded entirely by the Flah and Wildlife Service to avoid funding complications, avoid politics, and assure stable, long term support. Unfortunately, the Fish and \vildl:i..fe Service successfully procured only a portion of the support leavi5.1g The Peregrine Fund to make up the difference from the states, feder~l land management agencies, and the private sector. The majority of the contributions were associated with production of young for release, Jebndng little available to maintain the captive flock of breeding adults. 1'hi.8 job was designed as an interim measure to meet the funding shortfa.ll by contributing to maintenance of the breeding adults and from 1979 through 1982, 35 adult peregrine falcons of the anatum subspecies ue!:<G'. supported at The Peregrine Fund's Fort Collins facilities. During that period, the falcons remained in good condition and no disease problems or injuries were encountered. In 1983, Section 6 funding available to Colorado dwindled necessitating temporary suspension of the job. The fiscal problems have continued and the job has been terminated. It is not anticipated that total Fish and Wildlife Service support will be achieved, so the Peregrine Fund will have to rely upon private donations and fluctuating support from other agencies. �22 LITERATURE CITED Burnham, W., and G. R. Craig. 1981. Peregrine falcon captive maintenance" Job Progress Rep., Colo. Div. Wildl. Jan. Pp 34-37. Prepared by C.4. ~. Gerald R. Craig . Wildlife Researcher �23 Colorado Division of Wildlife Wildlife Research Report January 1984 JOB FINAL REPORT State of Colorado Project W-124-R Work Plan I Job 1 Period Covered: Author: Raptor Investigations Bald and Golden Eagle Nesting Studies 1 Mar 1979-28 Feb 1980 Gerald R. Craig ABSTRACT This job was transferred to the four regions for administration. Subsequent surveys will be reported by the regions under other project numbers. ��25 BALD AND GOLDEN EAGLE NESTING STUDIES Gerald R. Craig P. N. OBJECTIVES The objectives of this study are: (1) to estimate the breeding numbers and obtain production data of bald and golden eagles nesting in Colorado; (2) to identify important nesting areas and associated hunting habitats of bald and golden eagles in Colorado; and (3) to compile data and submit reports to associated state personnel and federal agencies for use in delineating and protecting eagle nest sites. SEGMENT OBJECTIVES 1a. Continue to locate and map nesting sites of golden and bald eagles throughout Colorado. Nest searches will be conducted with fixedwing aircraft and in some instances with helicopter. Additional field·work will be done from the ground by vehicles. Photographs of nest sites will be taken and pertinent information recorded about physical features of the habitat. lb. Nesting areas will be stratified by such features as climate, elevation and habitat type. Sample areas then will be delineated which are representative of important nesting areas. The sample areas throughout the state will be flown with a fixed-wing aircraft in late April and early May to ascertain the number of active sites. The same sites again will be checked from the air in June to determine the nesting success and productivity of the sample sites. The percent of active sites, percent of successful pairs, and total production will be extrapolated for each region of the state. 2a. When nests are visited in la, details will be recorded as to characteristics of the nesting habitat. Nesting eagles also 'c.; be observed from a distance to locate and map key hunting arl2oftS, 2b. Radio transmitters may be attached to several young bald eagl<es and a sample of young golden eagles to follow their movements after they fledge. Color markers mayor may not be used to mark the eagLes as well. 2c. A sample of sites will be selected and visited to determine the prey species present at the nest sites. This information will be valuable 5.nassessing the prey composition and availability to eagles. 3. Analyze all data obtained and prepare a report of the findings. �26 METHODS AND UATERIALS Methods and materials have been described previously (Craig 1979). RESULTS AND DISCUSSION Nesting surveys of bald and golden eagles have been transferred to each of the regional nongame programs for administration. Results will be reported by the regions under project numbers assigned to them. LITERATURE CITED Craig, G. R. 1979. Bald and golden eagle nesting studies. Wild. Res. Rep •. January: 81-95. Prepared by Wildlife Researcher C Colo. Div. �27 Colorado Division of Wildlife Wildlife Research Report January 1984 JOB FINAL REPORT State of Colorado Project W-124-R Work Plan I --~~----------- Job Bald and Golden Eagle Winter Population Surveys 2 Period Covered: Author: Raptor Investigations 1 Mar 1979-28 Feb 1980 Gerald R. Craig Personnel: Gerald Craig, Joe Frothingham,· 'and Wayne Russell, Colorado Division of Wildlife. ABSTRACT This job was transferred to three regions for administration. Subsequent progress will be reported under other project numbers assigned to the regions. ��29 BALD AND GOLDEN EAGLE WINTER POPULATION SURVEYS Gerald R. Craig P. N. OBJECTIVES The objective of this study is to obtain winter population trend information for bald and golden eagles on selected wintering areas in Colorado. The information will be used to estimate wintering populations of bald and golden eagles throughout the state. SEGMENT OBJECTIVES 1. Aerial Counts of Wintering Golden Eagles: Once annually an aerial flight will be made on census areas in the San Luis Valley, northeastern and northwestern portions of the state. These census areas were established in 1972 and the procedures will be essentially the same. Random transects will be flown throughout each study are~ with a Cessna 185 aircraft and all eagles observed within ~ mile of each side of the transects will be counted and classified as adult or juvenile. From the transects, an area estimate will be obtained as to eagles per 100 square miles. Identical census areas are'also flown in Wyoming, Idaho, Montana, North Dakota, New Mexico and Nevada by the U.S. Fish and Wildlife Service and cooperating agencies. Information forthcoming from these states are then collected and analyzed by the Fish and Wildlife Service to obtain population estimates for the West. Age ratio information which is obtained provides an indication of the previous breeding season's reproduction. 2. Aerial Counts of Wintering Bald Eagles: Since bald eagles tend to congregate primarily along river C;;H~'8ee and impoundments, aerial flights will be made along major river courses throughout the state and a direct count will be made of al bald eagles observed. The eagles will be classified as to a.ge in order to ohtain information about reproduction. The flights wll1 l.e made in January when the highest concentration of eagles are. ,(FlEa- ':~y present. Flights will be made along the South Platte River ~ Yampa River, White River and Rio Grande River. It is proposed to expand the censuses to the Colorado River and possibly the Gunnison~ Dolores and San Juan Rivers. 3. Compile data and prepare annual progress and final 'reports and submit them to.appropriate personnel and agencies. �30 METHODS AND MATERIALS Methods and materials have been discussed previously (Craig 9 1979). RESULTS AND DISCUSSION Aerial censuses have been transferred to the Southwest, Northwest, and Northeast regions for administration through their respective nongame programs. Subsequent progress accomplished through these surveys will be reported by the regions under project numbers assigned to them. LITERATURE CITED Craig, G. R. 1979. Bald and golden eagle winter population surveys. Colo. Div. Wildl. Res. Rep. Januar:87-9S. Prepared by c.a· c.~ Gerald R. Craig Wildlife Researcher C �31 Colorado Division of Wildlife Wildlife Research Report January 1984 JOB PROGRESS REPORT Colorado State of Project N-I-R~2 (W-124-R) Work Plan 2 (II) Job 1 (1) Period Covered: Author: Osprey Nesting Studies 1 January - 31 December 1983 G. R. Craig Personnel: E. Bowden, G. Claassen, G. Craig, G. Rosendale, S. Vana, and J. Wagner, Colorado Division of Wildlife. ABSTRACT Nest site occupancy by osprey (Pandion halieetus) in Colorado increased from 12 in 1982 to 14 in 1983. Eggs were laid at 8 sites but young were fledged at only 3 sites. Shell fragments were collected from 15 eggs and contents of 9 eggs were preserved for pesticide analysis. Observations from blinds continued, in an interrupted fashion, at 2 low disturbance and 2 high disturbance sites to determine effects of human activities upon reproduction. This Job Progress Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the Author. ��33 OSPREY NESTING STUDIES Gerald R. Craig P. N. OBJECTIVES The objectives of this study are: (1) locate previously unknown nesting ospreys and monitor productivity of all sites, (2) determine causes of reproductive failure and develop methods to restore normal reproduction, (3) determine nesting habitat requirements and implement protective measures to assure continued occupancy, and (4) compile data and prepare annual and final reports. SEGMENT OBJECTIVES lao Locate and map previously unknown osprey nesting sites throughout Colorado. lb. All. known nest sites will be visited annually to establish reproductive success. 2a. Observe nesting pairs from concealment to identify behavioral abnormalities, weather conditions, predation, or human disturbance which might impact reproduction. Sites in remote localities will be compared with those adjacent to public use areas to determine responses to human activities. 2b. All unhatched ~ggs and shell fragments encountered during nest visitations described in lb. will be collected and submitted for pesticide analysis. Shell fragments will be measured and compared with pre-DDT era eggs for possible thinning. 3. Habitat features such as hunting areas, topography, vege catLve type climate, and geology will be recorded for each nest site to establish habitat requirements. p 4. Analyze data and prepare a report of the findings. METHODS AND MATERIALS Methods and Materials for this study,have been described previously (Craig 1982). RESULTS AND DISCUSSION In 19839 14 sites were occupied, 2 of which were frequented by lone adults (Tables 1~ 2). Eggs were produced at 8 sites occupied by �w .p.. Table I.' Osprey reproduction in Colorado~ 1973-83. Parameter 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 Total Total nests 5 8 16 16 17 17 15 16 16 17 16 159 Occupied nests 4 6 6 8 13 6 8 13 11 12 14 101 Active nests 4 5 6 8 13 6 8 10 9 10 12 91 Successful nests 0 3 0 2 2 3 2 1 5 2 3 23 Young hatched 0 7 0 3 3 4 2 2 10 4 7 42 Young fledged 0 7 0 3 3 4 2 1 9 3 7 39 Young/occupied site 0.00 1.17 0.00 0.38 0.23 0.67 0.25 0.08 0.82 0.25 0.50 0.39 Young/active site 0.00 1.40 0.00 0.38 0.23 0.67 0.25 0.10 1.00 0.30 0.58 0.43 0.00 2.33 0.00 1.50 1.50 1.33 1.00 1.00 1.80 1.50 2.33 1.70 Young/successful pair �Table 2. Site Osprey reproduction 197.3 1974 1/3/3& 1/1/0 ?/1/0 JA-l .'::' JA-2 JA-3 JA-4 JA-S JA-6 JA-7 LA-1 LA-2 LA-3 GR-l GR-2 GR-3 GR-4 GR-S GR-6 GR-7 GR-8 GR-9 GR-10 GR-ll GR-12 GR-13 GR-14 GR-IS GR-16 GR-17 GR-18 GR-19 GR-20 GR-21 GR-22 GR-23 cGR-20 1976 1977 1978 1979 1980 1981 1982 1983 1/?/0 1/1/2 1/111+ TO 2+/0/0 1/1/0 1/?/1+ UOil UO AD AD UO OC UO OC TO 3/0/0 AD CC UO 2/0/0 2/0/0 3/0/0 AB TO TO AB 2/1/1 TO uo 1/? /2 7/1/0 1/1/0 , 1/0/0 2/0/0 2/0/0 ART 1/1/0 1973-83. 1975 1/1/2 1/1/0 lJO 1/0/0 1/0/0 uo ART ART 1/0/0 1/0/0 VO 1/1/1 1/0/0 OC 1/0/0 UO 1/0/0 UO ART vo 1/0/0 DC 711/0 111/0 TO AD UO UO 1/1/2 OC UO 1/0/0 1/1/0 1+/0/0 OC 1/0/0 TO 1/0/0 UO OC 1/1/0 TO 3/0/0 3/1/1 ? 11/2 OC 1/1/1 uo 3/0/0 3/0/0 1/0/0 TO AB 3/0/0 UO AB AD 3/0/0 UO 3/0/0 AB UO All . AD 0/0/0 '1/2/1 AB UO 1/1/0 TO 1/0/0 2+/0/0 2+/0/0 TO OC AB AB 3/0/0 pO UO AB 2+/0/0 1/0/0 0/0/0 2+/0/0 TD 3/2/2 AB 3/2/2 AB AB uo 3/0/0 AB 3/3/3 AB Uo TO 1/2/1 TO UO 0/0/0 UO aproduction bStatu9 by oite in Colorado, code: codes: 1/3/3 = ? eggs, 3 you~g hatched, uo AB 3/1/1 AD AB LA AD AB AD 2/0/0 UO LA 3/0/0 3/3/3 AB 3/0/0 AB AB 2/0/f!! UO AB 3/0/0 AD 3/0/0 3/3/3 TO TO TO AB AB TO 1/0/0c UO 1/0/fl,-2/0/0 2/0/0 AD TO 5/0/0 AB 4/0/0 AB AB 3/0/0 4/0/0 AB 3/1/1 AB 3/0/0 UO 3/2/1 AB UO TO TO UO OC UO 1/0/0 2/0/0 3 young fledged OC m occupied breeding territory. UO - unoccupied breeding territory. AB - abandoned. ART • artificial nest constructed. TD •• tree 0•• nest: blown down. th - lone ~~ul~ present. foile~ Gnd recycled Q~ GR-? ~ v �36 adult pairs. but only 3 pairs (GR-2, GR-13~ and JA-4) successfully produced young for an average of 0.30 young/active site. As in previous years, all nesting failures occurred.during the incubation phase. Of the nests that failed, 4 were due to infertile or dead eggs, 2 nests blew down during high winds, and 2 were abandoned for undetermined causes. Pairs of ospreys at 2 sites recycled after their 1st nest failed. The pair nesting at GR-22 failed when the nest blew down on 13 May, prior to completion of the clutch. The pair rebuilt the nest in the same tree and initiated incubation of 2 eggs on 28 May. The 2nd clutch was abandoned between 9 and 11 June. The pair nesting at GR-20 abandoned their eggs on 19 May after 11 days of incubation. Cause of abandonment could not be ascertained, but human disturbance was unlikely since it was in a remote locality. The pair relocated 2 km away at GR-7, which is considered to be a high disturbance location, and began incubating 2 eggs on 18 June. The renest attempt failed and 1 nearly full term dead egg was collected after 42 days of incubation. In 1983, 15 fractured or intact, nonviable eggs were collected and will be submitted to optical measurements for shell thickness. The 9 intact eggs were encountered in clutches at GR-1, GR-4, GR-5, GR-7, GR-I0t and JA~4 and will be submitted for chlorinated hydrocarbon insecticide analysis. The Colorado Epidemiological Pesticide Studies Center at Colorado State University analyzed samples from 7 eggs collected between 1978 and 1982 and reported DDE values (uncorrected) from 0.10 to 36.07 ppm. These samples will be corrected for water loss for comparison with values obtained from the 1983 samples. Four sites were selected in 1983 for intensive observation from blinds to document the impact of human activities upon reproduction (Table 3). Sites GR-l and GR-2 were chosen as representative of high disturbance nests and were observed throughout incubation (160 hrs) until it was determined from prolonged incubation that the eggs were nonviable. Cause for failure could not be attributed to human disturbance. Low disturbance nests were represented by GR-10 and GR-20, but due to premature failures, GR-22 was substituted for GR-10 and GR-2 was substit.uted for GR-20. GR-2 was subsequently observed through h•..• .:'1:1 (40 hrs) , but GR-22 failed after only 16 hours of observation. Observation was then switched to GR-7 which was considered a high disturbance location and 40 hours of observation were accrued until the eggs failed to hatch. In no instances could nesting failures be attributed to direct human disturbance. In anticipation of the final report, site specific information wa~ recorded for each nest, meteoroligca1 data and late ice conditions were co11ected~ and information on recreational use was obtained. LITERATURE CITED Craig, G. R. 1982. Osprey nesting investigations. Job Prog. Rep., Colo. Div. Wild1., Wi1d1. Res. Rep., Jan., pp 147-153. �Table 3. Intensive blind observation of nesting osprey~ 1983. Disturbance level Initiation of observation Termination of observation Total hours GR-l High 14 May 19 Jun 80 Observed throughout incubation; ended after collection of nonviable eggs. GR-4 High 15 May ·18 Jun 80 Observed throughout incubation; ended after collection of nonviable eggs. GR-I0 Low 03 May 27 May 34 Observation ended with nest failure on 27 May. GR-20 Low ,08 May 19 May 18 Observation ended with nest failure on !9 May. GR-22 Low 29 May 04 Jun 16 Observation ended with nest failure on 8 Jun. GR-2 Low 21 May );5 Jun 40 Observed through incubation to hatch. GR-7 High 21 Jul 24 Jul 40 Observed until nesting failure. Total hours 308 .Site Comments I.IJ .....• �38 Prepared by C i? ~ Gerald R.' Crrad.g Wildlife Researcher C �39 Colorado Division of Wildlife Wildlife Research Report January 1984 JOB FINAL REPORT State of Colorado Project W-124-R Work Plan Job III --~~~--------~-- Prairie Falcon Nesting Studies 1 Period Covered: Author: Raptor Investigations 15 March 1978-28 February 1980 Gerald R. Craig ABSTRACT The statewide nesting survey was terminated and nest site locations and other pertinent data were transferred to the regions. Data are being analyzed with the intent of examining productivity through a computer program of the Mayfield technique. A manuscript will be prepared and submitted for publication. ��41 PRAIRIE FALCON NESTING STUDIES Gerald R. Craig P. N. OBJECTIVE The objectives of this study are: 1. To document the breeding range and estimate the number of breeding pairs of prairie falcons in Colorado. 2. To obtain production data at selected nesting areas throughout the state and estimate total production of prairie falcons. 3. To delineate nesting habitat requirements of prairie falcons. 4. To document movements and mortality of prairie falcons throughout Colorado. SEGMENT OBJECTIVES 1. Locate and map all known nesting sites of prairie falcons throughout Colorado. From data obtained through approach 3, extrapolate the number of breeding pairs potentially occupying similar habitat types throughout the state. 2. Establish study areas in select habitats which represent important nesting areas. Study areas will be selected that represent shortgrass prairie, foo~hills and mountain nesting populations. Productivity will be determined and compar.ed between areas. 3. Physical and biological parameters of each nest site visited will be recorded on appropriate field forms and analyzed to establish those features which favor occupancy by prairie falcons. The f:teld information will be gathered in conjunction with approach 1. 4. When nests are visited to determine productivity, the young will be banded with Fish and Wildlife Service lock-on bands and their movements will subsequently be traced through reports filed with the Office of Migratory Bird Management. Should the occasion permit, transmitteJt8 will be placed on several breeding adults to determine extent of hunting ranges. Radio transmitters will also be placed upon young at fledging and their movements and activities will be monitored. 5. Compile data and prepare annual and final reports. �42 METHODS AND MATERIALS Methods and materials were described previously (Craig 1979). RESULTS AND DISCUSSION Field data recorded during previous years have been organized and summarized. Productivity information will be analyzed through a computer program which involves a modified Mayfield approach. A manuscript is in progress and will be submitted for publication. LITERATURE CITED Craig, G. R. 1979. Prairie falcon nesting studies. Colo. Df.v , Wildl. Jan. Pp 99-104. Prepared by G, ({. ~ , Gerald R. Craig~ Wildlife Researcher C Job Progress Rep., �Colorado Division of Wildlife Wildlife Research Report January 1984 JOB FINAL REPORT State of Colorado Project W-124-R Work Plan __ =I=I=I Job _ Prairie Falcon Breeding Population Characteristics Studies 2 P-eriod Covered: Authors: Raptor Investigations 15 April 1977-28 February 1980· Steve Platte ABSTRACT Two manuscripts and 2 articles were prepared as partial fulfillment of a Doctor of Philosophy degree. The first manuscript is titled "Population dynamics of. the prairie falcon in northeastern Colorado" and the second is titled "Individual vocal identify of alarm calls of the prairie falcon." In addition, an article titled "Successful breeding of innnature prairie falcons in northeastern Colorado" was published in Raptor Research 11:81-82 and the second article titled "Longevity of Herculite leg jess color markers on the prairie falcon (Falco mexicanus)" was printed in the Journal of Field Ornithology 51:281-282. These manuscripts and articles have been combined in a manuscript titled "Prairie falcon:" aspects of population dynamics, individual vocal identification, marking, and sexual maturity", a copy of which is on file in the library, Wildlife Research Center, Colorado Division -, of Wildlife, Fort Collins. Approved by: ��45 Colorado Division of Wildlife Wildlife Research Report January 1984 JOB FINAL REPORT State of Colorado Project W-124-R Work Plan IV Job 1 Period Covered: Authors: Raptor Investigations Ferruginous Hawk Nesting Studies 15 April 1977-28 February 1980 Gerald R. Craig and William C. Andersen ABSTRACT Field investigations were terminated and data are being analyzed. manuscript will be prepared and submitted for publication. A ��47 FERRUGINOUS HAWK NESTING STUDIES Gerald R. Craig and William C. Andersen P. N. OBJECTIVE The objectives of this study are: 1. To document the breeding numbers and productivity of ferruginous hawks in eastern Colorado. 2. To delineate habitat parameters and disturbances impacting breeding ferruginous hawks in eastern Colorado. 30 To evaluate potential techniques to enhance nesting pairs or encourage expansion of ferruginous hawk breeding populations. SEGMENT OBJECTIVES lao Locate~ map, and photograph all known nest sites of ferruginous hawks on the eastern Colorado plains. lb. Known nest sites will be observed from a distance to establish the presence of courting or incubating adults. Occupied sites will be revisted prior to fledging and the young will be counted, ~anded, and an attempt will be made to identify all prey items present. 2. Habitat features such as vegetative type, type of nesting structure, topogxaphy , soi-l type, climate, vicinity of human habitation, roads, and other disturbances will be recorded for those nests identified in la and lb. The physical and biological features will be evaluated to establish those parameters which favor successful nesting of ferruginous hawks. 3a. A sample of a predetermined number of nests will be stabilized w:1.th wire baskets in an attempt to enhance nesting success. Production of the manipulated nests later will be compared with unaltered sites to determine the effectiveness of the efforts. 3b. Between 1 and 2 dozen artificial nest structures will be placed itlsuitable habitats which are not occupied by breeding pairs. All breeding pairs adjacent to each treatment area will be located prior to placement of the nest structures and will be monitored after placement of the structures to be certain that the structures actually encourage pioneering by new pairs and do not cause relocation of adjacent pairs. 4. Analyze data and prepare annual progress reports and a final report. �48 METHODS AND MATERIALS Methods and materials have been described previously 1979). (Craig and Andersen RESULTS AND DISCUSSION Data collection was terminated during the previous segment and are currently being analyzed. The untimely death of William Andersen has complicated evaluation of the field notes and nest manipulation activities. A manuscript will be prepared and submitted for publication. LITERATURE CITED Craig, G. R., and W. C. Andersen. 1979. Ferruginous hawk nesting studies. Colo. Div. Wildl. Res. Rep. Jan. pp 111-120. Prepared by G.R.~ Gerald R. Craig ( Wildlife Researcher C �49 Colorado Division of Wildlife Wildlife Research Report January 1984 JOB FINAL REPORT State of Colorado Project W-124-R Raptor Investigations Work Plan IV --------------------- Population Surveys of Small Owls Job 2 Period Covered: Author: 15 March 1978-28 February 1982 Bruce Webb ABSTRACT A M.A. thesis entitled "Distribution and nesting requirements of montaine forest owls in Colorado" has been prepared, approved, and a copy is on file in the library, Wildlife Research Center, Colorado Division of Wildlife, Fort Collins. The abstract is given below: The breeding distributions of flammulated, northern pygmy, spotted, boreal, and saw-whet owls were investigated during 1978 and 1979. Flammulated owls were found in aspen-dominated stands throughout the south and central Colorado mountains. A total of five nests in live and dead aspen was studied. Pygmy owls were less commonly encountered than anticipated, with only one encounter during the two survey seasons. There are many more winter than summer records of this species, probably reflecting a downslope movement in winter:. Spotted owls were not found in any of their historical locations. Two male spotted owls were observed and heard calling in Navajo and Cliff canyons of Mesa Verde National Park. This species probably nests in the pinyon-juniper dominated cliff canycn habitat of western Colorado, but i~ extremely rare. Its nest has not been found in the state. There ~re many recent winter records of boreal owls in C.olorado~ and more spring records are occurring in the northern portion of the state. Populations of this species probably exist in isolated, high elevation stands of spruce-fir forest, particularly in northern Colorado. The sawwhet owl proved to be the most widely encountered of the five study species s' in terms of habitat and elevational range. Four nests were found 6f this most readily encountered species. Approved by: , :_,C .....: .,-'--/~'( ,_.,_d=' -,,-,~=' ',=' '0;;-' .• Gerald R. c1aig Wildlife Researcher C _ ��51 Colorado Division of Wildlife Wildlife Research Report January 1984 JOB PROGRESS REPORT Colorado State of Project N-1-R-2 (W-136-R) Population Surveys of Selected Bird and Mammal Species in Colorado Work Plan 3 (1) ~----~~-------- Job la Period Covered: Author: (1) 1 January - 31 December 1983 Gary C. Miller Personnel: R. Calderon, S. McCollough, G. C. Miller, Colorado Division of Wildlife; K. Webster, Colorado State University. ABSTRACT Population and h~bitat data for great blue herons (Ardea herodias) and double-crested cormorants (Phalacrocorax auritus) were collected to meet Program Narrative Objectives. Data are on file at the Nongame office, Wildlife Research Station and at the Cornell Laboratory of Ornithology, Colonial Bird Register. The sum of highest counts for each of 43 great blue heron colonies active between 1978 and 1983 totaled 1,918 breeding pairs or nest platforms. Most cottonwood trees (Populus spp.) encountered were small «25 mm diameter breast height), but the numbers of the next-largest size class (>25-75 rom) were among the fewest. Data analyses of habitat and other environmental variables are continuing. This Job Progress Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the Researcher. ��53 POPULATION SURVEYS OF SELECTED BIRD AND MAMMAL SPECIES IN COLORAiIO Gary C. Miller P. N. OBJECTIVES 1. For all great blue heron and double-crested cormorant nesting colonies in Colorado, both active and inactive, document the following: (1) colony size and past population trends, and (2) distances to different forms of human development and activity. 2. For all active great blue heron and double-crested cormorant nesting colonies in Colorado document the anticipated future human development and activity patterns near the sites, and the number and condition of the trees being used as nest sites. 3. Document the reactions of nesting great blue herons and doublecrested cormorants to different types and quantities of human activities at nesting colonies. 4. Determine the availability of future alternate nest trees for the active great blue heron and double-crested cormorant colonies in Colorado. SEGMENT OBJECTIVES 1. Review current literature on determinants of nesting by great blue herons and double-crested cormorant. 2a. Analyze data. 2b. Write Quarterly, Job Progress, and Job Final Reports. 30 Write manuscripts and submit to appropriate outlets. 4a. Answer information requests by CDOW personnel. 4b. Develop guidelines for maintenance of heron and cormorant co Iorrl.es in Colorado, and present to Regional personnel. ,. METHODS AND MATERIALS Methods and materials have been described previously (Miller and Vos 1982). By the end of the reporting period, all known colonies in Colorado had been visited. Protocols used in colony site measurements are in Appendices A-D. �54 DESCRIPTION OF STUDY AREA The study was conducted in the riparian and reservoir-riparian zones of the major drainages of Colorado and their tributaries. Population data in 1983 were collected primarily in northeastern Colorado. RESULTS AND DISCUSSION Data were tabulated for population trends of all known great blue heron and double-crested cormorant colonies in Colorado. The records are on file, colony-by-colony, at the Nongame office, Wildlife Research Station, F.ort Collins. Additio.nally, population information has been transmitted to the Colonial Bird Register, Laboratory of Ornithology, Cornell University and is stored in their computerized data banks. These data may be accessed by CDOW personnel. Data were added in 1983 for the Fossil Creek, Timnath, Arikaree, South Fork Republican, Empire, and Riverside colonies. Eighty colony sites were identified in Colorado; 43 were known to be active at some time between 1978 and 1983. No obvious relationships between human activities and status of colonies were detected; further data analysis will be needed. These data are tabulated and on file, colony-by-colony, at the Nongame office, Wildlife Research Station, Fort Collins. The sum of high counts of great blue heron nest platforms, or breeding pairs, for each colony during 1978-83 totaled 1,918. A simultaneous count of all colonies was not performed. A total of 846 sample plots was measured to characterize the nesting resource of great blue herons. Cottonwoods (populus spp.) occurred in 88% of all plots. Young cottonwoods «25 mm diameter breast height) occurred in 16% of the plots that contained this species. Saplings (:>25-75 mm diameter breast height) were encountered in 9% of the plotg (Fig. 1)0 A large proportton of the trees encountered in each of :, stand categories (active, inactive, unused or "alternate") were of .he <25 mm dhb category (Fig. 2). Fewest trees were in the sapling (>25---75 pole (>75-150), and large, over-mature (>750 mm) size classes. Nest trees, identified by the presence of platforms, had mean diameters at breast height for active colonies ranging from 297 to 1,110 mm 'toihile means for all stands ranged from 187 to 661 mm , For inactive stands, mean dbh of nest trees was 449-1,163 mm; means for all staujg were 216-638 mmo The significance of these statistics has yet ·to be ascertained. At the end of this reporting period, substantial data analysis has yet to be performed. A workshop for Southeast Regional personnel is planned for 01 March 1984. YlXt<1'l) > �50 ra ACTIVE COLONY P,LOTS (236)' lSI .INACTIVE n C/) W:::J 40 U....J Z:::J Wa.. 0:0 COLONY PLOT~ (119) 40 ALTERNATE PLOTS (816) ...•.•..•.••• •• a: a.. 30 :J. 28 UJ: U'I- o~ "_'C/) ZIWo U....J a: a. W a.z 20 16 12 Ir;·;·;·;·~ ·.... .~.~.~.~. I It=.:.:.:.~ ·.... .... ·.... .... 1°t-~1 0 > 750 DIAMETER Fig. 1. Percent occurrenee of co t tnnwood trees, BREAST HEIGHT (mm) by size class, in plots with cottonwoods 0 \JI \JI �69 10 IJt 0'\ rn:ACTIVE COLONY CSI· INACTIVE 60 o. STANDS (1,104) COLONY- STANDS (1,062) ALTERNATE STANDS (4,801) '5-0' 50 C/) w w a: ••••• 40 LL ••• ···... .... ·... 0 • ••••• z w o a: w 30 •• It :::::=: .:0:.:. ·0·.·.. 27 ::::::: a. 20 10 ::::::: ·.. ~:~:~:~ ·.. ::::::: .:.:.:. ....··... ..·0· ·... ····....,..'. ........ ·.. " • • 0 0 •••••• 0 C! • •• II • •• 1'•••• 0 ••• • o • •• 0 •• •• e ! ·0-25 10, ······ ...-... ··· ... 1<1.:.:.:.:.1: ····........ ·.... 9 t res .~ < 2. ••• ·23 ····· ..... ··.·.....•.... ···· .... 14· .:.:.:.: ···· ....I ···· .... :.:.:.:. ······ ...... · . ······ ...... 22 < < ' 7 2 < • .... .. ... .... ..... e 2 Size composition of co t tonerood trees number of cot tonwood stems sl?...mple,:t < in great .0 ••••• ftJ I eo. •••• •••••• 76-150 'DIAMETER Fig. ·· .. ::::::::: · ... ~~~~~~~~~ ·...· ... ·:.:.:.:.: · .. ·:·:·:·:·l!2J1 ···... · ..... ° ····.... · .. ··· ... ···. .... o • eo· ••••• 0 t G •• 3'76-750 151-375 11 '3 .••••••.•• .k . . < < < areas. >1 75(" BREAST HEIGHT (rnm) blue heron nesting G Numbers in pa rent heae e indicate �57 LITERATURE CITED Miller., G. C., and D. Vos. 1982. Population surveys of bird and mammal species in Colorado. Job Prog. Rep., Colo. Div. Wildl., Wildl. Res. Rep. Jan. Pp 73-107. Snedecor, G. W., and W. G. Cochran. 1967. Statistical Methods, 6th ed. The Iowa State Univ. Press, Ames. 593pp. Prepared by ~ C. JY1_~ Gary C. iller Wildlife Researcher C �58 APPENDIX A HERON STUDY PROTOCOLS PRIOR TO DEPARTURE 1. Locate heronry on topographic map 2. Determine landownership 3. Inform D.O.W. district manager of schedule. 4. Contact landowner prior to leaving Ft. Collins - explain purpose of study, a. get permission for access* b. get permission to take cores and mark trees 5. When arriving at site - check with landowner. 6. If landowner is not located at site, contact adjacent landowner. 7. After owner contact, return to heronry, begin sampling schedule • .* Make sure that if access to heronry requires access via land other than that of heronry "owner", all landowners are contacted and permission to cross property is granted. �59 APPENDIX B HERON STUDY PROTOCOLS On Site Techniquesll Enter on Sketch Map 1. After locating heronry, measure maximum length of heronry stand, noting beginning and ending of nest trees. Sketch Map 2. Measure width of stand from enough points along long axis to provide an estimate of total area of stand. Again, note beginning and ending of nest trees. Sketch Map 3. From steps 1 and 2, draw map of heronry (and stand) on "Sketch Map" sheet. Indicate magnetic North on map, and geographic center of stand. Sketch Map 4. Verify legal description. Locate first plot center (See "Plot Center Determination) and mark with stake. Enter stand number (0 if heronry stand) in column 10, plot number (0 if < 500 m2) in column 11. 5. Enter plot centers on sketch map. If entire ~ot does not fall within stand boundary, determine the actual area of plot that occurs within stand (See "Table of Partial Plot Areas") Field Data Sheet 6. Record plot size. Field Data Sheet 7. Take measur~ments as indicated on "Field Data Sheet" on all trees within plot boundary.J:./ Field Data Sheet 8. Secure numbered tags, at eye level, to all nest trees within plots and record under increment core number. �60 APPENDIX B cont. Field Data Sheet 9. After sampling 500 m2 take required measurements and secure tags on all nest trees within heronry that were not included in sample plots. Record these data on separate data sheet and note on sheet "Non-plot data." 10. Sample all adjacent stands of trees within 500 m of geographic center of colony stand. If no stand < 500 m sample "nearest stand". Sketch Map 11. Draw on sketch map (from step 3) and number each separate stand. Field Data Sheet 12. Sample all adjacent stands according to steps 4-7. Record stand number in col. 10, plot number in col. 11. 1/ An accurate count of active nests (or of indviduals) for great blue herons and other colonial nesting species within the heronry should be made when possible. 2/ For trees;~ 3 inches G6nnn) d.b.h., use circular plot = 100m2 "" raddus 5.64m, measure all trees; for trees < 3 inches d.b.h. 9 use cirCl!:'.cX' . 2 plot = 25m = radius 2.82m count all trees and record average condition for groups. Both plots use same origin. Record data for both plot sizes, regardless of presence or absence of trees less than 3 inches d.b.h. �61 APPENDIX C HERON STUDY PROTOCOLS FIELD DISTANCE MEASUREMENTS - Heron Study 1. After drawing sketch map locate geographic center of heronry on sketch map - designate as 0 2. Locate center of heronry. 3. Triangulate with compass on permanent landmarks (2), record azimuth readings on sketch map. 4. Locate landmarks used in Step 3 on topographic map, and designate geographic center of heronry on map as 0. Add new landmarks to map as needed. S. Measure required distancesl/ on topographic map, record on data sheet. (Do this in vehicle after field data collected). 6. Note any disturbance not accounted for on data form under comments. 1/ Distances defined: to nearest water: measured to nearest edge. ~ to nearest road; measured to nearest edge. to nearest habitation: measured to nearest house (rural). to hearest habitation: measured to nearest edge of development (ur.ban). to nearest recreation boating: to nearest edge of water where boating is allowed. fishing: to nearest edge of fishable waters. camp/picnic: hiking: to nearest permanent campground or picnic table. to nearest established trail. �62 APPENDIX D HERON STUDY PROTOCOLS PLOT CENTER DETERMINATIONS Enter stand and proceed to geographic center. From random number table!/ generate a random number between 0 and 360 as an azimuth reading. Follow this reading to stand perimeter. Select random number (0 to 9). This number indicates the number of meters into the stand along the random compass heading (-1800) the first _plot center is located. Succeeding plot centers will be located at 12-m intervals from the first plot center, along the random compass heading. If the opposite end of the-stand is reached prior to sampling 500 m2 proceed at 90% to the original directional been sampled. 11 (again at 12-m intervals) until 500 m2 have Record transect and plot centers on sketch map. Snedecor and Cochran (1967) �63 Colorado Division of Wildlife Wildlife Research Report January 1984 JOB FINAL REPORT State of Colorado Population Surveys of Selected Bird Proj ect No. _.:o.:N_-.::.1-_R::.:..,_-,;:;2 _ Work Plan No. Job No. 3 ~--------------lb. ~~------------------- Period Covered: Author: and Mammal Species in Colorado Response of breeding great blue herons to human disturbance in Northcentral Colorado 1 February 1981-30 June 1984 Diana K. Vos Personnel: C. E. Braun, A. T. Cringan, S. A. Flickinger, W. D. Graul, C. L. Mahoney, G. C. Miller, R. A. Ryder, and D. K. Vos. ABSTRACT Reactions of nesting great blue herons (Ardea herodias) to human disturbance were studied during the 1981 and 1982 breeding seasons at the Fossil Creek and Lonetree Reservoir heronries. All human activity within 100 m of the heronries was monitored and rates of human .intrusion were documented. Reactions of herons to human activity were grouped into 3 categories: minimal, local, and general responses. Rates of human intrusion were lowest (minimum = a intrusions/hr of observation) early in the breeding season, and peaked in June and July (maximum = 1.33 intrusions/hr of observation). Sixty-seven percent of all human intrusions caused minimal response. Local responses were elicited towards 27% of the human disturbances and only 6% resulted in a general res~nse. Herons were most disturbed by landrelated activity and least by boating activity. Heron response to human activity decreased as the breeding season progressed, with an increasing percentage of minimal responses (28.6 - 95.9%) being elicited each month. Heron response also varied between sites. Productivi-tyranged from 2.65 to 2.82 young/nesting attempt and 2.82 to 2.96 young/successful nest, and was sufficient to maintain a stable population. Recommendations to reduce human disturbance of breeding great blue herons are discussed. ��65 THESIS RESPONSE OF BREEDING GREAT BLUE HERONS TO HUMAN DISTURBANCE IN NORTHCENTRAL COLORADO Submitted by Diana Krammer Vos Department of Fishery and Wildlife Biology ~ In partial fulfillment of the requirements for the Degree of Master of Science Colorado State University Fort Collins. Colorado Spring. 1984 �66 COLORADO STATE UNIVERSITY Sprin g , 1984 WE HEREBY RECOMMEND THAT THE THESIS OUR SUPERVISION OF BREEDING BY DIANA KRAMMER VOS PREPARED ENTITLED UNDER RESPONSE GREAT BLUE HERONS TO HUMAN DISTURBANCE NORTHCENTRAL REQUIREMENTS COLORADO BE ACCEPTED FOR THE DEGREE OF Committee MASTER OF SCIENCE. on Graduate Depar-tment Head ii AS FULFILLING Work IN IN PART �67 ABSTRACT RESPONSE OF BREEDING GREAT BLUE HERONS TO HUMAN DISTURBANCE IN NORTHCENTRAL COLORADO Reactions of nesting human disturbance seasons - general o within intrusions in June and July tion) • Sixty-seven percent season (28.6 tte pt maintain turbance and of herons minimal, local, were lowest 1. 33 intrusions/hr towards in a general activity to and (minimum = in the breedin g season, were elicited by land-related All and of observa- caused minimal 27% of the human response. Herons and least by boating Heron response to human activity decreased as the breeding ~ progressed, with an increasing percentage of minimal l"(,;,:ponl3es - 95.9%) being between = heronries. Reactions of all human intrusions and only 6% resulted were most disturbed activity. early to 1982 breeding was monitored into 3 categories: (maximum Local responses disturbances Reservoir were documented. /hr of observation) herodias) 1981 and Rates of human intrusion peaked response. the and Lonetree were grouped responses. during (Ardea 100 m of the heronries of human intrusion human activity blue herons were studied at the Fossil Creek human activity rates great sites. elicited Productivity each month. ranged population. of breeding great also varted from 2.65 to 2. 82 young/m<~ting nd 2.82 to Z.96 young/successful a stable Heron response nest. Recommendations blue herons and wa to reduce .uffici O1t to human dis-· are discussed. Diana Krammer Vos Department of Fishery and Wildlife Biology Colorado State University, Fort Collins. Colorado Spring, 1984 �68 ACKNOWLEDGEMENTS Major funding Division special of Wildlife through thanks opportunity to Dr. major advisor, continual Drs. equipment. In addition, and unstinting Lastly. possible. enthrall for his enoral I thank Their me. the great grace I truly to Dr. R. A. Ryder, and understanding, assistance S. A. Flickinger during for their love, of all my endeavors, support and elegance hope my efforts never for supplying sugges- of the project. financial support, as well as my and unending blue herons I also positive criticism, the final stages faith in me. which made the study ceased will benefit my and and suggestions. for his constructive my parents encouragement Pete, patience the study, guidance I thank I extend and gave me the and C. L. Mahoney for their and C. E. Braun and invaluable husband, is also extended to provide throughout program. study. for his constant encouragement tions. this A. T. Cringan by the Colorado the nongame research appreciation willingness was provided W. D. Graul who initiated to pursue My sincere thank for this project to fascinate them. and �' ... ; 69 TABLE OF CONTENTS INTRODUCTION. .... • STUDY AREA 75 ... METHODS •• RESULTS. Nesting • 41 • • 85 .•• Phenology •• 85 AZ'rival~ Courtship. Egg-laying and Nest Construction Chronology 85 • • • 87 of Human Intrusions 88 ........ Fossil Creek Reservoir Lonetree Reservoir Overall Intrusion Heron Response 88 • • 93 Patterns. 95 to Human Intrusion Heron Response at Different 95 Sites. • .99 in Heron Response Experimental Nesting 85 and Incubation •• Nestling Development Changes 81 l(JO Intrusions 10.0 . Success •• DISCUSSION MANAGEMENTRECOMMENDATIONS LITERATURE CITED 6 • • • e 0 . 0 0 0 0 .. 0 .· .· . · 0 101 108 115 118 �70 LIST OF TABLES Table· 1 Phenology of great blue heron nesting at Fossil Creek and Lonetree reservoirs, 2 Colorado. 1980-82. .• 89 91 Reservoir. Colorado. 1980-82. Heron response by type of human intrusion at the heronries studied in 1980-82 5 86 Human intrusions observed per hour of observation " (Rra-FLonetree 4 •• Human intrusions observed per hour of observation (R) at Fossil Creek Reservoir, 3 Colorado. 1981-82. Heron response 97 . • • • to land-related and boat intrusions 99 at different heronzies , 1980-82 • .. • 6 Heron response to human intrusions at heronries 99 studied in 1980-82 f) 7 Average distances (m/month) at which herons were first disturbed by a person approachin g the heronry experimental intrusion for 3 daily time segments •• 0 105 �71 LIST OF FIGURES Figure 1 Page Location of Fossil Creek and Lonetree Larimer County. Lonetree 4 Human intrusions 77 Colorado Fossil Creek Reservoir 3 reservoirs. Reservoir heronry , Colorado. heronry. Colorado. 1981-82 1981-82 .• 78 79 - A * heronry. 5 observed 1980-82 .• Human intrusions Lonetree average reservoirs. caused distance herons Reservoir. 1 Average person distance caused Lonetree herons intrusions 102 reacted to a at the Fossil Creek heronries. intrusion. Colorado. for a person 1982" to be disturbed Colorado. 102; approach- • • • • • • • . • • • • at which experimental reservoirs. 96 • • . . intrusion Reservoir by herons at Fossil Creek at which herons approaching distance flights at which experimental Site by month interaction Average 1981-82. and the the 1981 season. to be disturbed ing the heronry 9 Colorado. Colbrado.1982 and Lonetree 8 94 ••••••• number of normal activity Average (R) at each per hour at Fossil Creek and per hour throughout 6 0 per hour 104 intrusions at Fossil Creek and 1982 106 �72 INTRODUCTION The great blue heron and intensively 1962). studied It occurs southern heron across Mexico north range and south 1962) • Although on the east to the northern great McAloney 1973; Edford (Cottrille and Cottrille extensively examined studied States (Graber et al. wariness of great North America, areas (Thompson 1977. Mathisen and Richards important factor that that distance in selection 1967; Kushlan few studies United (Vermeer sites colonies is detrimental 1978, Stephens from human activity of nesting ThE: of many breeding human disturbance ~S. responsible in the midwestern 1978) and Canada have blue hc' r;, great as a major factor and remoteness from urban (1943) suggested 1970, 1972; 1978. Kelsall and Simpson 1979). blue herons suggests (Pratt on breeding declines 1978, Thompson 1973. Markham and Brechtel biology this range. 1960. Mock 1976) have been has been implicated population within (Palmer 1979), and social behavior of human disturbance blue heron of its breeding ecology (Hedeen 1958, Meyerrieks throughout Human disturbance for great breeding - impacts part coast of South America 1976), feeding 1976, 1978; Willard 1977; Parris from Union 1983). (Am. Ornithol. have been recognized blue heron breeding Alaska on the west and Nova much of the southern Nine subspecies 1926, Palmer in North America (Bent to southern throughout most widely distributed, most of North America, Scotia and New Brunswick It winters is the largest, 1980). Miller was the most by herons. In Oregon, �73 Werschkul size, et al , (1976) found an indirect the number the distance Parker occupied that associated heronries within a colony was correlated with distance Great blue heron well quantified. occurring In the early colonies containing Rockwell 1909). herons Surveys conducted inactive great blue hero,p colonies the presence (Graul 1980). Three human recreational activity, and blue a state- of abandonment development and highway and 21 additional in 1980 (McGrath had from 1 to 40 nests trees, large great of 38 active heronries removal of nest that of colonies was in northeastern causes as by the Colorado Division of Wildlife nests. include occupying More recently, from 1 to 85 nests, Documented were listed with 23 colonies identified contained ~10 nests. not been 1909, Hersey Active heronries Inactive from roads, 1973 indicated colonies were identified concentration of They were described (Felger et al , 1979). 1978 and 1979 documented The greatest 1912). in Colorado, between and 2 inactive and 0.62 km The number blue herons in the northeast, in 1965 and in 1965 and 18 in 1973 (Ryder survey rate, operations. in Colorado have of 100 nests were still widespread wide waterbird (Sclater especially upwards trends 1900's great Colorado as a common breeder, colony to roads. population throughout fledging in Montana averaged developments. with proximity between with logging and 0.71 km from urban decreasing active within a colony, of human disturbance (1980) reported from roads nests of nests relationship however, 19,,_i). Colorado. 61%had <15 with 83% containing of colonies in Colorado of reservoirs construction and associated (Graul 1980). �7'4 The human population during the last decade in Colorado has increased (Weaver 1980) and large areas land have been converted to urban Thus. in nestin g areas human disturbance become a serious problem. colonies intensify. their nests exposure could result ment of a colony. herons respond (2) nest desertion. it is important blue herons pressures adult herons mortality 1978). could around to leave of young due to or (3) complete abandonto understand how great blue to human disturbance. of human disturbance and (2) quantify of this study at selected responses, were to: great of great human activity to learn if response (either the day or during sites. of great that causes of agricultural and Perri as recreational in (1) increased Thus. The objectives during use (Anderson especially Harrassment or predation. by over 30% (1) document current levels blue heron colonies in Colorado blue herons varies to different types with type of activity. the breeding season). of time or between �75 STUDY AREA The study the Front western Divide was conducted in northcentral Range of the central extension Plains, (Palmer Divide), on the east. and Crosby of the Great 1982). north of the Platte-Arkansas This area is within the Colorado Plains physiographic The region is drained Vrain rivers. These flow east-northeast. in the region. short grass rivers In addition, The original prairie (Salix interior), sargentii) areas -25 to -30 C. fluctuate Winter temperatures Summer temperatures often reaching 35-37 C (Hansen 19 to 25 C (Berry cm of which about average annual Heronries Larimer County. reservoirs was typical support sandbar and plains and occur of willow cotton- 75% falls between snowfall approximates with mean annual average average et al. 1974). at Fossil Creek and and Wasser 1980). The climate of the ifregion is continental tur-es of 9-10 C. irrigation maple (Acer negundo), (Scott River Big Thompson, of the region Riparian (Hansen in the Rocky Mountains numerous vegetation (Dix 1974). boxelder wood (Populus originate province by the South Platte and its major tr-ibutar'ies ; the Cache la Poudre, St. by Rocky Mountains on the west and the of the Great Piedmont section Colorado delineated 1978). 21 C with daytime highs Daily temperatures precipitation April and September. 122 cm (Berry where a wide diversity lows oi -2 C, reaching Mean annual Reservoir tempera" and Lonetree may is 30-35 The 1974). Reservoir of human activity in was known to �76 occur, were selected observations for observation were reviewed described by McGrath (1981). The Fossil Creek heronry shore cottonwoods reservoir and incorporated. of Fossil Creek following the contour and surrounding waterskiing occurred Land north human activity riders. Those near for nesting The Lonetree of Berthoud large grove north and south edges The eastern Reservoir horseback heronry The heronry many trees the several nesting Nesting pairs eastern herons abutted (Fig. 3) 0 used 3 km northwest The heronry occured The western site of the reservoir grove bordered a causeway the western 0 The 1980. The Herons 1960's after per s . commun.). site since a Lonetree site in the late (Co Cummings, W< ..::: In to the east. edge of McNeal Reservoir. were shot as agricultural separating a smaller reservoir site from the western have reoccupied had been formerly was approximately on the east shore site was on the south and 0 from McNeal Reservoir. moved to Motorboating people on foot, of the cottonwood border use was included 0.5 km apart of cottonwoods owned; The Other - about 2). for agriculture. in the NE! of S9, T4N, R69W. 2 sub colonies (Fig. row of side of the heronry. However, Reservoir was on in a U-shaped club. and motor vehicles. had died and fallen The site was used the heronry 50 years. were 14 km southeast was privately boating and Chatfield heronries of the shoreline shoreline of the heronry for at least Reservoir along the south motorcyclists, eastern Creek was approximately limited to members of a private land. In additron , some Collins in the SW! of S9. T6N, R68W. the northwest active 1). in 1980 by E. McGrath at Boulder Reservoir of Fort (Fig. �_c.----=- __ _. -=-- __~--- ~ LARIMER ~C£;O_ COUNTY N " FOSSIL CREEK t-:t RESERVOIR ~ COLORADO o 10 LONETREE RESERVOIRClt 20 BERTHOD KIL0METERS • .-..z Fig. 1. Location of Fos s l L Creek and Lonet r ee reservoirs, Larimer County, Colorado. ~ -...J -...J �78 .. I FOSSI L CREEK HERONRY "I N I / Cl / o / I I I I l \ FOSSIL CREEK \ RESERVOIR \ \ \ '--, .•..•.. <, <, -,\ \ \ \ ,/ / ./ --~~ ./ / / ...._ _.-_ II-X ~ HIGH SHORELI NE LOW SHORELINE FENCE TREES @> TRE ES WITH HERON NESTS Fig. 2. ----- ...•.• / / ./ o E3 so i Fossil Creek Reservoir heronry, Colorado,· 1981-82. 60 90 F*H METERS �LONETREE HERONRY N HIGH SHORELINE _-LOW SHOREU NE •.•.•...•..•.• CAUSEWAY BENCH c:;:;::>. TREES ~ TREES WITH HERON NESTS LONETREE RESERVOIR ---- ---,-=--- ___ --.// ..-_.",.--- _.....- _--- _...,.,.......- ...•.. ------ WELCH --- ...•... ......_ RESERVOIR -_ ---30 60 90 ,~ E METERS ./ . ......_ ....•... 0 / .•.....••. -....J \0 Fig. 3. Lonetree P,eservoir her onrv C,)lorado, 1981-82. �80 Land bordering privately the 2 heronries owned but considerable at Lonetree human traffic Reservoir originated public campsite on the west shore of the reservoir. farming activity occurred regularly was from the In addition. near the heronries. �81 METHODS Field work was conducted and 1982. Fossil Creek twice a week on a rotating on weekdays heronries throughout of the breeding season. research herons However, herons within any human activity to react was recorded. outside into 3 response categories: general r sponse the heronry. from their All observations scope from a distance herons by my presence. (termed of 100 m was selected that during beyond nests, of herons a human because the nesting that 100 m which obviously distance. caused heron to human intrusions wer'e (1) none or minimal r'esponee ( 2) local response in the area closest - temporary could be related With each human intrusion. grouped of nests was followed to human activity Reactions abandonment phenology 100 m of the heronries was recorded. flushed 0600-1200 or cycle. response no herons the either of 6 hours, A distance reacted of On each day of observation (McGrath 1981) indicated rarely Reservoir an equal number season so that observations was monitored. preliminary 1981 July and Lonetree to include Great blue heron nesting All human activity intrusion) schedule for periods the breeding to major stages Reservoir and weekends. were watched 1200-1800 hours. through 15 March and 21 July 1981, and 6 March and Between 20 July 1982, I visited visits from late February abandonment - temporary to the intrusion, of some nests were made using and (3) throughout a 15-4SX spotting of 250 m or more to prevent Comparable intrusion - disturbance rate of the and heron response �82 data gathered during a preliminary study were combined with data I obtained dence between of intrusion. the number site. by herons 25 active nests randomly short each hour obtain an estimate Because of additional occurred a series in 1982. intrusions. known to occur were: operated heronry. including (>300 m) by 4 to per hour. of controlled 1981) and 1981. spaced experimental included in 1980 in time intrusions a minimum number known to have The experimental in time and short in duration Intr-u- to min" }.i·~f: to the herons. The experimental person at was and long flights well below the number sions were widely spaced vities all flights, human intrusions The experiment Herons segment were multiplied flights of uncontrolled in 1980 (McGrath disturbance obtained the number on each day of ob ser+ segment a heronry tables. in 1981. 1981 was too low and irregularly analyses. was conducted beneath and the type the day. a IS-minute IS-minute of normal activity 1981) and for statistical were monitored The values the number during each hour was counted this to the ground were counted. activity Indeperr- X2 contingency using of observation. During away, (McGrath during at both heronries chosen. flights in heron 1981) sample sizes. in each category and month was tested of flights During to increase of responses To examine variation vation , in 1980 (McGrath intrusions at Fossil Creek consisted and Lonetree (1) a person approaching riding a motorcycle past near the heronry, and All 4 types of 4 types reservoirs. the heronry the heronry. These on foot. (3) a tractor (4) a motorized of experimental of human ac ti vi ry (2) a being boat passing human intrusions ;:dj-· the were conducted �83 at Fossil Creek approaching Reservoir. and the boat intrusions The person towards At Lonetree approaching the heronry. in the experiment. approached the heronry occurred During as I walked slowly of 12-16 kph at closer the motorcycle on a motorcycle motor was used. In the tractor could be done. intrusion For the boat intrusion. HP outboard only the person I wore similar clothin g each time to maintain consistency 16 k ph . Reservoir, a 3.7-m. V-bottom boat with a 9! The heronries a tractor I at a speed of approximately and 'closer distances intrusion. intrusion. were passed at a speed un til the herons was driven towards reacted. the heronry at a speed of <8 kph , The motorcycle. intrusions were conducted The person approaching time segments: day - between Each intrusion intrusions was repeated mental intrusion fledglings occurring. 0700 and 0900 hours. during 3 different 0700 and 0900 hours, and (3) afternoon at a specific (2) mid- - between 1600 time segment was con- the chance were conducted throughout of herons becornin g was measured disturbed were first season. The distance. disturbed to Each to the by each experi- with a Leitz Rangefinder. by an intrusion when present) on a monthly basis the breeding twice each month. 10 m, at which herons considered was conducted days to reduce examine heron response nearest intrusion experimental to the intrusions. Experimental intrusion in the morning between 1100 and 1300 hours, on different habituated and tractor (1) morning - between and 1800 hours. ducted motorized boat. Herons were when 2 or more adult herons flew from their nests If only 1 adult heron was present while the activity on a nest during (or was a human �84 intrusion enough (common later to be left unattended), from its nest heron in the breeding was measured. response or between the distance Data were grouped nesting each site was counted success, and productivity of nestling counts. they began climbing onto branches (1971). to examine whether. with type of disturbance, the impact of the current blue heron to represent at which that heron flew time of disturbance. sites. To assess on great varied were old season when nestlings fledging levels of human disturbance the number of active nests was estimated A count of young within nests success Young at that surrounding through just their prior nests at a series to when was used as recommended by Henny and Bethers time were approximately 45 days of age. �85 RESULTS Nesting Arrival. began to arrive each year herons Herons trees Reservoir nests. present at Lonetree to 3 March. reservoirs between Reservoir, Reservoir wary, at 12 returned to 24 and 28 February. among the higher position. at Fossil Creek were present In 1982, herons They were extremely in an alert and early March blue herons on 28 February; and Lonetree outstretched herons In 1981, 20 great prior Great blue herons sites in late February at this time were perched near necks 1). had arrived Fossil Creek and Nest Construction.- at the study (Table Fossil Creek than C()urtship, Phenology branches standing with their In late February took flight of the 1981, all when I was more 200 m away. Pair formation began March pairs continued at the heronries. first. within a few days after arrival. to form and more and more nests Nests r'ernairring from previous followed by construction of new nests. years Throughout were O~.Tupied were occupied Nests were occupied within dead as well as live trees. Egg-laying and Incubation Eggs were laid over a period started after both adults the first Peak egg-laying of several egg was laid. were generally was completed. c-- only 1 heron occurred days and incubation While the clutch probably was being seen on or near the nest. was on the nest in April. throughout laid, When the clutch most of the day. �00 0\ Table 1. Phenology of great blue heron nesting at Fossil Creek and Lonetree reservoirs. Colorado. 1981":'82. Year Site 1981 Fossil Creek Reservoir Lonetree Reservoir 1982 Fossil Creek Reservoir Lonetree Reservoir Mean interval (Days) aApproximately Peaka incubation Colony occupancy Initial incubation 28 Feb 27 Mar 25 Apr 3 Mar 30 Mar 23 Apr 26 Feb 24 Mar 27 Feb 27 Mar Initial Peak 80%of pairs at a particular Initial Last 19 May 30 Jun 24 Jul 30 Apr 18 May 29 Jan 22 22 Apr 24 Apr 19 May 23 24 Apr 28 Apr 22 May 26 Jun 2 May 32.5 stage. Jun , "_ 28.0 Fledging Hatching 58.7 Jul 20 Jul 22 Jul �87 Incubating herons more than 6 hours. arrival Nest relief occurred (Mock 1976). heron was brief. pairs for long periods, often 1 to 2 times a day. Upon of its mate, the heron on the nest rose and gave a stretch display other remained on nests It then stepped moved to the center taking <30 seconds, as 1 adult left the nest Nestling asynchronous seen at Fossil Creek on 3 May. nestlings. between throughout between in May. ages. Hatching Nestlings nestlings were observed was were first on 5 May 1981 and at Lonetree Their above the perimeter herons relief ceremony Reservoir on 27 April at Reservoir. The 2 and 7 days of age at the time. were brooded ing of young continued Brooding to hatch and on 28 April at Lonetree most of the day. seen bobbing The nest was no interaction in young of varied Reservoir Reservoir and there Eggs began In 1982, the first Fossil Creek of the nest. as the area until the next relief session. Development.resulting to the edge of the nest gray downy heads could only be of the nest for approximately could be distinguished during feeding. 1 week after Brood- hatching. from those still incubating eggs in that the former moved more frequently to accommodate the nestlings. In June, both parents nestlings foraged. were generally left unattended By approximately were climbing onto branches surrounding wings. I observed By 50 days of age. surrounding branches Peak fledging 60 days of age. in the nest 4S days of age. their nests nestlings the nestlings to exercise taking while short their hops from into the nest. occurred in July. Nestlings After fledging, young herons fledged continued as they reached to return to �88 their nests weeks. and remained in the vicinity of the heronry By the middle of July most nestlings Chronology Fossil Creek Reservoir.- 0.19 human intrusions Human intrusions graphers approaching During observation the heronry (:R = was lower unusual intrusion passing directly had fledged. March 1981 there of nature watchers as well as boats passing rate a hot air balloon approaching and "(R) at Fossil Creek than in March (Table 2). also increased during the heronry but not to the level observed April of both years consisted on foot as well as horseback <100 m above the heronry flying April 1982 During and 2 motorcycle intrusions. the heronry. by helicopters One above the heronry. 19810 Intrusions approaching the per hour of 0.04) than in 1981 (Table 2). in 1982 involved was higher the intrusion and photo- passing of human intrusions In April 1981 the rate of human intrusions Reservoir was a total of "(R) (Table 2). of observation primarily In March 1982 the rate heronry. 2 of Human Intrusions observed/hour consisted for about in of people riders. trucks Aerial intrusions were also ob<_;.';;, ved in 1982. In May 1981, the rate of intrusions the May rate remained human intrusion Most intrusions heronry June 0 (R) the same as in April. increased in June each year did so to gather In June, and in 1982 the rates from those measured of in Mayo 1981 and 1982 (86.0%)' were boats passing People on foot constituted 1981 and 1982. "(R) decreased only 12% of the intrusions Many of the people walking beneath wild asparagus. the in the heronry �A Table 2. Human intrusions Colorado. a 1980 -82. Type of intrusion Land People on foot 1980 1981 1982 b Motor vehicle 1980 1981 1982 Motorcycle 1980 1981 1982 Horseback 1981 1982 Water All boatsC 1980 1981 1982 Air Helicopter 1982 Hot-air balloon 1982 observed per hour of observation Mar '" R - 0.083 0.028 Apr N - 3 N 0.062 0.055 3 3 0.021 0.018 1 1 0.037 2 0.021 1 0.167 0.037 '" R 0.104 0.037 1 0,083 3 ,... Jul Jun May N 5 2 1 0.018 0.018 '" R (R) at Fossil Creek Reservoir. 0.074 4 8 0.400 0.062 5 3 2 0.037 2 '" R N 0.472 0.074 0.037 15 4 0.032 1 0.018 1 0.032 1 0.220 0.667 0.130 7 36 7 it N 0.067 1.333 0.083 1 48 3 2 00 1.0 �a Data from McGrath (1981). blncludes trucks. cars, and farm machinery. clnc1udes motorized boats. unmotorized boats, sailboats. and canoes. �Table 3. Human intrusions observed per hour of observation '" (R) at Lonetree Reservoir. Colorado, 1980a-82. Mar Type of intrusion '" R - . May Apr N - '" R - N - '" R - Jun N - '" R - Jul N - '" R - N - Land People on foot 1980 1981 1982 b 0.042 2 0.042 2 0.062 3 0.079 0.050 0.018 2 0.987 25 0.066 2 0.071 1 0.021 1 0.021 1 0.230 7 .0.071 1 0.093 5 0.083 3 3 1 Motor vehicle 1980 1982 Motorcycle 1980 0.040 1 0.158 4 Horseback 1980 1982 0.021 1 0.021 0.208 1 10 0.021 1 Water All boatsC 1980 1981 1982 0.021 1 0.671 0.050 0.148 17 3 8 Air Airplane 1982 \0 ••• �\0 N 3. Table Continued. Mar Type of intrusion Apr A R N - - Jun May A A R N - - Jul A A R N - N R - - - R N - - Total number of intrusions/month 1980, 1981 1982 NO OBSERVATIONS 0 5 49 6 9 1 15 9 0 7 2 0 3 Total hours of observation /month 1980 1981 1982 18.0 48.0 50.0 48.0 25.3 60.0 54.0 30.5 48.0 48.0 14.0 30.0 36.0 0 0.104 0.020 0.312 1.934 0.100 0.167 0.295 0 0.146 0.143 0 0.083 Total number of in trusions / hour/month (:R) 1980 1981 1982 a Data from McGrath bInc1udes trucks. crnc1udes motorized (1981). cars. and farm machinery. boats. unmotorized boats, sailboats. and canoes. �93 During July, decreased. nestlings No nature all human intrusions heronry. the rate were observed, 1981 and 1982 were boats of human intrusions the heronry passing and the per hour of observa- high levels in June and July (Fig. 4). (66.1%) of human intrusions. The on foot disturbance (20.9%) was a common as well. Lonetree Reservoir.- human intrusions = within the heronry or photographers the major source people approaching (R and activity each month reaching Boats constituted in trusion watchers in July Overall. tion increased fledged Observations were detected 0.10) were observed were made in March 1981 but no (Table 3). Only 5 human intrusions in March 1982. A During passing April 1981, only 1 human intrusion the heronry Reservoir. pecially Many other during (probably at a distance boats weekends, because (R = 0.02), of 80 m, was observed were seen on the reservoir but few came within the water receded a sailboat at Lonetree in 1981, es- 300 m of the heronry from below the heronry). In A April 1982, the rate of that year (Table sions were boats of human intrusion 3). passing 60 m above the heronry Sixty-six (R) reached percent the heronry. the highest of the April One airplane level 1982 intz-u- was seen. Hying as well. A During May 1981, the rate A highest level (R = of human intrusion 0.10) at Lonetree Reservoir. (R) reached its In 1982 the May int sion rate decreased observed in May 1980 and 1981. occurred over the last weekend in May (Memorial Day). 'U- from that in April but was similar to the rates Most intrusions in May each year A In June Reservoir and July, decreased the rates of human intrusion from those observed in previous (R) at Lonetree months (Table 3). �94 0.6 FOSSIL CREEK RESERVOIR .,_.,...•••.LONETREE R SERVOIR -.'- 0.5 .I:: U) '\ I \ I I 0.4 I 0 .-. \ I I I «I) ::> a: l- \ \ \ I 0.3 \ \ \ \ I I I I I :) :x: La. \ \ 0.2 \ \ \ I 0 w \ \ ~ ••••• <t // ex: 0.1 0.0 Fig. 4. ./ ./ /' "- """-="""""'""'-F----,.----...,...-----r---__.... MAR APR MAY JUN Human intrusions observed per hour (!) at each heronry, 1980-82. JUL �95 No intrusions were observed in June there were only 10 intrusions boats passing peaked boats (52.8%) passing on foot constituted Overall the (raining 1981 breeding P < 0.05) than The number kph , N = calm days and/or = season There 20). 5.18. on warm. (t = than reservoirs 0900 and while local responses Boats passing of 'the intrusions resulted caused = (N 20). >24 from the number on on weekends on weekdays. pattern during 1982 (Fig. morning 1.82. 1981 and and late afternoon of 5). at while most human in tr'uaione 1500 hours. to Human Intrusions during Only 6% of the intrusions the heronry days = the daily activity (66.9%) caused were elicited on 1981 and 1982 combined the number between in the early Most human intrusions 4). during and human intrusions between sunny (t (maximum win d velocity 1. 27. P > 0.05) reservoirs ~ < 0.001) was lower of human intrusions Heron Response (Table the heronry <10 C. N = 7) throughout on windy days and Lonetree (89.6%) occurred was of human intrusions of intrusions The number were most active Fossil Creek The number Reservoir was no relationship blue herons of intrusion at Fossil Creek and Lonetree (!_ = of human intrusion People approaching temperatures 6) was not different was higher Herons Patterns.- the number at Fossil Creek great the heronry. In 1982 of which 80% were The most common type of intrusions (N the rate 1981. 35%of the intrusions. Intrusion cooler days Overall. 4). 1980 and in June and July. the heronry. in May (Fig. or July only minimal responses 27%of the human intrusions resulted in a general minimal (92.1%) responses. in a local response. response. Only 8% All boat intrusions �96 25 /~ (I) •••• l: - C) _J 1.1.. I , \ \ \ 20 \ \ LL(/) oo wa:: UJ :t: \ 15 \ \ / . /-.. \ '\ \\ :J)o- 10 CD W C) <[ 5 ~ o (I) 18 0:: LaJ :> Z .•... I~ ::> 0:: 14 (/) t=- \ 12 zrt') en 10 :,:)11 8 - :t: ZI 0 6 I- 4 U 2 a:: w 0 f2 8 co 0 0 I 8 to 0 0 0 en 0 0 0 0 - ~• 0 0 0 0 0 0 0 0 0 0 1'-= 0 co 0 en 0 0 0 0 0 0 rt) 0 0 N ,... 0 0 tD 0 0 an I - 0 0 0 -• v- C\I 0 0 0·· 0 0 0 CD 0 tot) 0 V 10 I U) ,... TIME Fig. 5. Human intrusions reservoirs. activity Colorado. flights per hour at Fossil Cr~ek and Lonetree 1981-82. and the average by herons per hour throughout number of normal the 1981 season. �97 Table 4. Heron response by type of human intrusion at the heronries in 1980a-82. studied Heron Minimal Type of intrusion % resEonse Local N General % N % 68.3 54.5 8.3 77.9 56 6 1 7 8.5 27.3 66.7 11. 1 N = 114) Land (N People on foot Motor vehicleb Motorcycle Horseback 23.2 18.2 25.0 11. 1 Overall 19 2 3 1 61.4 21.9 7 3 8 1 16.7 = Water (N 203) All boatsC 92.1 187 7.9 16 100.0 75.0 1 3 25.0 1 = Air (N 6) Airplane Helicopter Hot-air balloon Overall Total (N = aData b 323) from McGrath Includes cIncludes canoes. trucks, motorized 100.0 66.7 16.7 16.7 66.9 26.9 6.2 1 (1981). cars, and farm machinery. boats, unmotorized boats, sailboats, and �?98 which elicited a local response were caused and canoes which were maneuvered No boat intrusions duration resulted in a general intrusions resulted (61. 4%), while minimal responses intrusions. Only 17 caused responses under response were elicited the response often Reservoir (Table 4). season. However. a hot-air resulted Overall. general the response by Iand+related a general between differed and land-related of response sites elicited at which it occurred. was (Table 5). (X2 = Motorcycle (66.7%) (Table 4). at Fossil Creek (X 2 = Reser- 252.5, Great blue herons and least by boating at Different to land-related for both boating elicited response as well. was dependent intrusions response intrusions on response. Heron Response Great blue heron tree. in minimal responses P < 0.001) upon the type of disturbance. disturbed a heronry A general responses balloon above the heronry voir in March 1982 caused General on 2 May 1981 when 4 people approached (66.7%) caused Aerial intrusions 22%of the land when people approached early in the breeding at Lonetree intrusions category with nests. throughout during the heronry and one of the people climbed a nest differed trees most often in local responses a general were often elicited especially caused directly of the study. Land-related foot. by slow-moving boats activity. Sites and boat intrusions The number of responses activity (X 2 = 9.16, 11.44, ~ < 0.05). by similar intrusions were most ~ < 0.05) Thus, was dependent per the type upon the site �Table response Heron 5. to land-related and boat intrusions at different Type General Local N - % Site % N - % N 19.6 16.3 7.1 10 8 1 Land (N = 114) Fossil Creek Reservoir Lonetree Reservoir Boulder Creeka 9.8 26.5 50.0 5 13 7 70.6 57.1 42.9 36 28 6 Boat (N = 203) Fossil Creek Reservoir Lonetree Reservoir Cha tfield Reservoira 96.7 83.9 88.4 117 47 23 3.3 16.1 11. 5 4 9 3 a Data from McGrath 6. Table Heron (1981). response to human Mar intrusions at heronries Apr studied in 1980a-82. Jun May Type of response % Minimal 28.6 4 64.7 33 51. 6 47 66.7 Local 42.9 6 21. 6 11 40.7 37 General 28.6 4 13.7 7 7.7 7 a Data from McGrath a 1980 -82. of response Minimal Intrusions heronries. N - % N - % -N % Jul % N 62 95.9 71 31.2 29 4.0 3 2.1 2 N - (1981), \0 \0 �100 Changes Great blue herons in the breeding disturbance. longer in Heron Response were most responsive season and flushed At that present. from their herons Attachment to the nest appeared in full foliage. it was sometimes possible any herons to abandon ing season progressed responses upon the types nests. was elicited (Table 6). each month. each month differed Overall. responded intrusions distances by the 4 types Fossil Creek Reservoir There intrusions, from land-related herons experiment. Uncontrolled lea ve their (~= nests. an increasing percent- as the breeding in type of response elicited dependent each month. Intrusions intrusions Distances 6). 0.36. ~ > 0.25) between of the uncontrolled heron response (Table 4). boat intrusions the types rarely of For the activity The experiment during at at which herons intrusions. to boating nests were first conducted (Fig. to fly from their were without ~ < 0.001) and were partly of experimental intrusions sions caused a heronry at which great blue herons unlike results uncontrolled and returned were also fewer local and which occurred varied did not differ 9 nests to human intrusions Experimental disturbed eggs had egg laying and incuba- to enter The changes (X2 = 63.67. of intrusions The average after when the young were older and the trees age of minimal responses general at the shghtest strengthened to be less willing to abandon Later, early until the cause was no During more readily. causing nests time they did not return been laid and once young were present. tion, to human intrusions differed 1 boat intz-u- each month of the caused herons to �101 Distances at which herons approached differed interaction, site by month. indicating that between the rate months was different The average approaching distances mid-day. intrusion varied occurred nests. not significant. response Although there between months voir of which at least produce any young the ground. Three were unsuccessful. Reservoir fledged while 4 nests renesting At Lonetree 2.74 ± the day was Success in 1981 at Fossil Creek from 107. were abandoned or late nesting Six pairs Reser- failed to and 1 nest attempts Mean (± SO) productivity fell to occur-reo observed and all at Fossil Creek attempt and 2.S5 nest. Reservoir. 27 nests in 1981 of which 25 were successful. duced. flushed ~ < 0.001) in heron in 1981 was 2.65 ± 0.98 young/nesting ± G.6S/successful first 9). was occupied 1 young The time at during 201.2, (~= time intervals the day did not influence time of disturbance (Fig. between to a person 7). at which herons Nesting A total of 115 nests responded (Table during was ia change An ~ < 0.005). changed in the 3 different and afternoon) 7). 8). at which herons O. 52. ~ > 0.25) the distance from their response as I (Fig. (F = 3.6, was also significant (Fig. nests ~ < 0.001) 10.6, (~= at each site which human disturbance (~ = sites flew from their at which heron the heronry (morning. first 1. 23/active nest were occupied Seventy-four and 2.96 ± at the eastern young 0.99/successful site were pronest. �•...• : o N 0' ngo 60 disturbed re'·1 ,off •. ' «4 It ,.-.-.-.] !<? 1;'-"-" ./? peile'n'cn! 1{4 I Average distance at which experimental intrusionl!i caused h~z:pn~ to, be at Fossil Creek Re~eirvoirp Colorado, 1982. Sample sizes are in parentheses. .,. , �180 (6) 11._. !65~ SOl I E -z 135 !6) ~ D LONETREE 'RESERVOIR (WESTERN SITE) I I FOSSIL CREEK RESERVOIR E:I LONETREE RESERVOIR ( EASTERN SITE) 0 - CJ) ::> 0:: I- Z 90 LL 0 7 W U Z 60 ~ (j) 0 o' K < <, , ( ..~ U i MAR V < <. i APR MAY I-' JUN JUL Fig. 7. _Averas.ediatances at which herons reacted to a person approaching intrusion at the Fosst] Creek and Lonetree Reservoir heronries. parentheses. Colorado. 1982. Sample sizes>are in o w �104 180 FOSSIL CREEK RESERVOIR 160 ----- \ \ ". _._.- \ , LONETREE RESERVOIR (WESTERN SITE) LONETREE RESERVOIR .( EASTERN SITE) \ \ 140 \ \ \ \\ \\ -z , \ '.\ E 120 \\ '\ \\ ..- 0 \\ (J) :::::> 0:: \\ 100 '~ I- .\ z '~ LL 0 80 w u z ~ en .," \\ " '\: ~, 60 0 \ ,. _.;-:.. \ ''-.-_._o" ..",. \ ,,"'" \v " . _- -_... ___ 40 20 OL---~--------r-------~------~------~ JUl MAR Fig, 8. APR MAY· JUN Site by month inter ction for a person approaching the heronry intrusion. �Average Table 7. the heronry distances experimental (rna/month) intrusion MD were first disturbed by a person approaching for 3 daily time segments. Mar Site at which herons Jul Jun May Apr MD A M MD A M MD A M MD A M MD A Fossil Creek Reservoir 140 125 130 40 60 55 25 40 35 65 45 55 70 70 60 Lonetree Reservoir (western site) 175 175 160 80 80 95 50 40 45 60 60 55 50 60 50 Lonetree Rservoir (eastern 165 130 175 90 75 80 50 50 60 55 60 60 70 60 50 site) aEach value based bM = morning on a sample of 2 except (0600-0900 hrs), MD = in July. mid-day (1100-1300 hrs), A = afternoon (1600-1800 hrs). I-' o VI �•.... o 0\ 150 -o ,E z 13 12 ~ - 10 a:: 9 z 7 I' (26) (f) ::> ( 13) ~ IJ.. . (26) o s w u 4 z ~ en o (25) (24) 3 i ! MAR Fig. 9. disturbed APR MAY JUN Average distance a~ which experimental intrusions caused. herons to be at Fossil Creek and Lonetree reservoirs, are in parentheses. Colorado, 1982.' Sample sbes JUL �107 In 1982. 94 nests Young were produced young/nesting attempt were abandoned; ing the nest nesting At Lonetree in 1982. in 92 nests. and 2.82 vertically. 0.84/successful supporting There Reservoir. All were successful 22 nests was 2.76 nest. and produced broke. leav- or late in producing young. at the eastern site a total of 62 young of young in 1981 and 1982 was more than adequate (nesting produced at both when compared the value of 1. 91 young/breeding pair by Henny to maintain a stable (1972) to be necessary ± 0.93 Two nests the nest wer e occupied The number Reservoir , were 2 renesting and both were unsuccessful (2.82 ± 1. 03 young/nest). heronries ± at Fossil Creek and productivity 1 when the branch hanging attempts were occupied attempt) calculated population. to �··108 DISCUSSION The rates of human intrusion seemed to be most influenced human intrusion varied each site occurred breeding that season by weather. between in March. of human intrusion. sites For example. 10. 7 cm while average low. turbance early had been sites. in the breeding to make the ground are extremely season (Cottrille when herons first arrived to abandon and less likely to abandon. tivity in subsequent flew from their each month. to human disturbance were (Natl . months. nests illustrating 1958. 1976). if chere at the breeding the heronries. to become habitu- even with increasing The average decreased during the change as the breeding to human dis- and Cottrille ated to the sites mental intrusions rates in There muddy sensitive in March may have allowed herons first at (~O.025 cm), precipitation Less disturbance which herons in the in May 1981 totaled for May is 7.4 ern, may have been more inclined of human disturbance early the observed 1970. Dennis 1971, Brandman more disturbance they at 1981). Admin. blue herons 1960, Pratt Meyerrieks the lowest rates of human intrusions Precipitation 8 days of which were wet enough great of of cooler temperatures the rate 16 days in May 1981 with measurable Because 4). studied peak rates also influenced precipitation Oceanic and Atmospheric Although (Fig. the result Precipitation May 1981 was comparatively at the heronries The low human disturbance was probably time of the year. observed season rates distance at the experi- in heron sensi- progressed (Fig. 9). �109 Although flushed intruders could come much closer from their less disturbed nests, at that in May when herons nestlings probably in their nests There because 4). Longley (1960), habituate tinually boating passing being past for newly hatched had considerable investment them. in heron least response disturbed was relatively in Florida. common near a heronry of great herons to sudden. such as people walking below nest the to boats as blue can such as fishermen as opposed of by Miller (1943). Apparently. activities to types by boating Habituation has also been reported (980). were was strongest may have been habituated and Nordstrom herons the herons attachment to abandon activity (960) activity that eggs or caring herons to common and expected disturbances trees con- unexpected or a motorcycle the heronry. No significant other Boating and herons to boating nest difference with herons by Meyerrieks herons adult was a significant studied reported Instead, and were reluctant (Table heronries time. did not indicate were incubating human disturbance activity this in May before types intrusions difference of experimental at Fossil Creek ing the heronry at speeds experimental intrusions 12-16 kph , Furthermore, boat only passed intrusions Reservoir the heronry tances until a response the heronry during was passed a response (Fig. boating 6). consisted of approximately the heronry study. intrusions, was found between activity boats at speeds at closer During boat intrusions, intrusion and closer most experimental at distances the of only once while in the experimental was only elicited pass- 45-55 kph while during most uncontrolled was elicited. boat Most uncontrolled of motorized was passed repeatedly and nearer disboat to the a �110 shore than most boats water. Experimental the few uncontrolled namely, could usually come safely because of shallow boat intrusions actually boat intrusions which did elicit a local response. slow-moving boats seemed to characterize which were maneuvered directly below the heronries. There were too few aerial intrusions the study that to determine fixed-wing (Nordstrom aircraft 1980). both helicopters do not generally drastically aircraft disturbed distances of 30 to 267m above the ground. the aircraft. habituated The herons south of the reservoir; day of observation (Table 5). at least One factor sity of disturbances, at Fossil Creek Reservoir however s lingering consisted rapidly most boat intrusions was an airport by flying at along the edge of the heronry, each season. at different this may have been the intenfrom nest >50 m, fishermen and duration. most boating of water skiers involved of 3.5 km were seen overhead For example, at distances of may have been to human intrusions in terms of distance at each heronry. the heronry Reservoir there influencing which occurred passing and egrets the 1982 breeding differently and He found no response 10 airplanes throughout Herons responded sites because of noise levels or the presence at Fossil Creek to aerial activity the effects (1978) examined the effects on herons increased colonial nesters colonies when flying at alti- In Ohio, Grubb to either agree on colonial wading birds noise levels from aircraft birds during Most observers (1979) tested increased from nesting 6) observed bother Kushlan and fixed-wing as low as 60 m, = impact on herons. In Florida. found that neither tudes their (N activity and speed boats At Lonetree Reservoir, in slow-moving boats, often for more than an hour. �111 Another response factor between at Lonetree sites was heronry Reservoir were not as readily Creek Reservoir There study the study. to leave nests to thermal stress. occurred highest. during The eastern grove of trees heronry and intruders as they were at the Fossil impacts that human disturbance during herons in heron which was more open. were numerous adult subjected was within a dense heronry differences structure. visible to the herons had on productivity causing which may have influenced For example. could have disturbance can cause eggs or young to be Most disturbances observed during the middle of the day when temperatures this were Bartholomew and Dawson (1954) found that even at moderate air temperatures (24.1-27.4 C). radiation significant elevations caused great blue herons. Blus et al. and 2 embryos in a nest a brief exposure to direct in body temperature (1980) reported subjected to excessive solar of nestling the death of 2 nestlings solar radiation during human disturbance. Adult herons flushing can cause breakage from the nest (Bjorklund 1969. Nordstrom be killed as a result et al , 1967). to leave the nest will no longer Nestling or disturbed. regurgitation herons possibly when suddenly to be accidentally 1980). dislodged Most dislodged to the nest. feed and defend 1974). If they will most likely die as them. are also known to regurgitate in loss of weight of food by disturbed nestlings can cause older nestlings as well (Teal 1965. Armistead resulting disturbed of a 30 m fall to the ground Human interference prematurely they are unable to return parents nests of eggs or nestlings (Jenni would probably from their nestlings food when frightened (Jenni was observed 1969). No in this study. �112 (1978) examined the impacts of human disturbance Potyraj and development of tricolored herons (~. egrets <.~.' thula} affected caerulea) , cattle tinuous egrets (Egretta ibis}, responsible and snowy activity. for the decreased could also disrupt little blue growth was significantly amounts of investigator was partly disturbance tricolor), {Bubulcus and found that nestling by increasing regurgitation herons on growth feeding Nestling growth. schedules, Con- affecting nestlin g growth. Predation temporarily during upon nestlings leave the nest. this study. by crows and aura) may also increase No predation however. ravens attacks (Corvus have been reported when adult herons on nestlings on nestling great blue herons spp , ) and turkey (Bent was observed vultures (Cathartes 1926, Temple 1969. Ives 1972, Krebs 1974. Kelsall and Simpson 1979). Bald eagles as nestling (Haliaeetus great 1952. Bayer blue herons heronry during caused loud squawking about preying was observed this study. adult herons noises. Presence once in 1981 at Fossil Creek of the eagle within the The eagle remained of great blue herons (Fischer An immature golden to circle the heronry 4 minutes and made no attacks similar reaction on adult as well has also been documented 1979. Kelsall and Simpson 1979). eagle (Aquila chrysaetos) Reservoir leucocephalus) frantically, perched on nestling making on a branch or adult for herons. A to a golden eagle was reported by Wilburn (1970). Great horned as predators fresh owls (Bubo virginianus) on great blue herons. remains of herons under a tree have also been implicated Cottril1e and Cottrille used by a great {1958} found horned owl's �113 newly fledged young. tion however. and many observers nesting This was only circumstantial (Miller 1943. Page 1970. Bjorklund English 1978. Warren 1979). horned owl nesting ignored landed herons flushed nests owls their 1976, a great in 1982 was completely At Lonetree when a great Reservoir horned owl within the heronry. Although there are potential ance on productivity. nesting of human interference observed. no effects of disturbance mixed-species heronry found no detrimental turbance aurttus) adverse success impacts of human disturb- was not affected by the levels Goering and Cherry frequency (93% cattle on reproductive egrets). in a colony of double-crested (1971) also found success DesGranges effect on reproductive success cormorants in a large and Reed (1981) with human dis(Phalacrocorax in Quebec. The lower rates of human disturbance at the heronries studied gators human disturbance studying human disturbance occurred early in the season (Hunt 1980. Ollason and Dunnett and during egg-laying Productivity However. nesting in colonial nesting success. birds to reproduction 1980). Black-crowned season Investi- have found when it 1972. Ellison and Cleary were most susceptible 1978. night-herons to disturbance just before (Tremlay and Ellison 1979). was adequate other early in the breeding may have influenced to be most detrimental (Nyd::icorax nycticorax) study. Reservoir. my observations. from their horned et al , 1967, Edford At Fossil Creek during of preda- seemingly accepting within 200 m of the heronry by the herons however. Cairnes have found great within heron colonies with the herons presence evidence factors to maintain the population besides human disturbance. during this including �114 food supply and weather were not measured. impacts in years Human disturbance when other addition, even though sufficient for population the number Werschkul nests of pairs et al. active higher activity colonies. could influence the number stability, occupying (1976) found in relation in undisturbed factors of young away from the point affect heronr+es , of disturbance In was may be limiting heronry. and occupancy of nests) detrimental per nest within a particular density. factors productivity. produced human disturbance to the number vs . disturbed These could have severe adversely nests nest productivity. (number of to be significantly A shifting was also noted of nesting in disturbed �115 MANAGEMENT Management of great into 3 categories: pretive blue heron nesting (1) restrictive or educational improve great RECOMMENDATIONS measures. or warning habitat. Impacts of human disturbance on great sites. buffer of this study, and 150 m in water is recommended. herons distances plus an additional reasons. First. be disturbed 1975). appeared beyond nests 50 m, prior Although a buffer These distances The additional to flushing from their Individual heron response and the past varied history herons for flight at distances Further, by human activities or stressed (Thompson et al. heronries nests, the caused the breeding were recorded. at which they flew. were since animals can be disturbed b eha vioral responses encompass activities adjust no exact measurements to be disturbed nesting season. for 2 may already of humans since by the time they fly. to be aware of human intruders have appeared around 50 m is suggested had to physiologically the distances can be reduced zone of 250 m on land at any time throughout by the presence they have already blue herons at which human recreational to abandon (2) inter- to maintain and zones free from human activity Based on results greatest can be divided measures, arid (3) measures blue heron breeding by establishing areas (Geist herons more than herons may not yet show no overt 1968). between of human activity 50 m while they actually should be examined independently significantly often sites. since Vegetative at a particular structure site may demand �116 lesser or greater ation. distances At some sites. size of a buffer next of 150 m, there zone. to' a heronry for buffer zones depending may be physical For example. there limtiations restricting may be a narrow where it would be difficult In such a situation. upon the situ- to create human activity the channel a buffer zone should be kept to a minimum by limiting the number of boats within the area and restricting boating breeding activity the middle of the day and early in the season. Buffer herons during zones should be maintained arrive at breeding been deserted for creating Buckley sites until early for the year. buffer (1976). from mid-February Restrictive August signing zones on land are reviewed before when sites have and fencing methods by Buckley and Buoys can be used to form boundaries around heron- ries in water. In addition educational signs with heronries if appropriate. what heronries and discuss some areas. or warning measures. should be placed at entrances and. should describe ing there. to restrictive boardwalks can be established. interpretive to recreational describe why human interference and depict lectures Signs species breed- is detrimental. or viewing outlooks with educational In addition. areas within sight of heronries. are. and or tours In exhibits can be given by area personnel. Besides ance. efforts be considered. for breeding. species protecting breeding great blue herons to maintain and improve existing Great blue herons The importance has been recognized from human disturb- breeding in Colorado require of riparian habitat as well (Carothers habitat should riparian habitat for many other and Johnson 1975. �117 Gaines 1977. Bull 1978). ductive and valuable 1977. Fitzgerald ecosystems wildlife habitats 1978. Schrupp Colorado is steadily of trees Riparian being lost wherever 1978). (Borden tree cutting Riparian This lack of regeneration ment and cattle grazing Nesting of great of cottonwoods is due primarily (Crouch the non-breeding deteriorate. artificial structures effective season if possible. artificial In addition 1982). nest structures additional Cottonwood regeneration around are improved. species heronries. tree cutting absent. to death of nest 1969. Wiese 1978). reinforced If nest and preserving breeding heron Artificial species methods for by Meier' (1981). existing habitat to blue herons have been discussed potential continue could be developed. 1980) and other To help by managers trees by great trees habitat. an should be made. should be promoted as recommended and out- lined by Beeson (1983) and. and shrub and land- in water manage- Design and implementation to maintaining to create of cottonwoods structures 1978. Sandilands (Wiese 1976. Hafner 1980). of extensive can contribute have been used successfully (Henny and Kurtz (Tubbs to changes could be structurally nesting 1979) and cutting 1979). blue herons a heronry , trees throughout has been virtually (Miller 1943. Kerns and Howe 1967, Vermeer preserve (Hubbar-d in public areas to the consequences and removal as regeneration attempt habitat 1978. Crouch should be prohibited owners should be educated nest they occur for firewood has become a major problem Unregulated during are among the most pro- when methods for successful plantings can be planted should be done. to provide better propagation Other vegetative tree buffers �118 LITERA TURE CITED Anderson, R. L .• and K. Perri. in the northern Coop. American Ext. Colorado Union. 6th ed . Lawrence. Armistead, Smith Island of rural lands Colo. State Univ . 26pp. 1983. Check-list Union, of North Allen Press American Inc , , 877pp. 1974. Lower Chesapeake to Barren G. A .• Jr., regulation Urbanization Range - 1978. Am. Ornithol. Kans. H. T. Bartholomew. Front Serv •• Fort Collins. Ornithologists' Birds. 1978. in young Island. heronries Maryland and W. R. Dawson. pelicans. herons of Maryland. Birdlife 30:9-27. 1954. Temperature and gulls. Ecology 35:466- 472. Bayer. R. D. 1979. Murrelet Beeson, T. 1983. Management riparian A. C. U.S. Berry, Sr •• associations 1926. Natl. J. W. 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Group. censuses Comment on the flight 3 colonial waterbird Mathisen. of helicopter Wildl. Manage. 1960. blue heron. Effects U.S. 21pp. Administration. Dep . Commerce, 1981. Climatological Natl . Oceanic and Atmos- �125 Nordstrom. W. R. Alberta 1980. Colonial waterbird Dep , Rec , and Parks, and Manage. C' Ollason , J. Sec. and G. M. Dunnet. D P. J. 1970. J. San Joaquin Palmer. W-S4-R-2. R. S.. Parker. J. Montana: M.S. Parris. W. R. foraging State Field Orni th , River ecology growth H. M. Md. 1972. herons Ryder. habitat (Ardea 1. Missoula. of great Calif. Dep , Wildt. Invest. American birds. Yale 576pp. herodias ) in northwestern of human disturbance. 82pp. blue heron Lake Erie. (Ardea M.S. herodias) Thesis, Ohio Ll Opp. The effects of human interference M.S. Thesis, Towson on heron State Univ .• 117pp. biology California. Nesting and movements Group. study. use and. the effects success in the San Francisco Waterbird 51 : 39- 54. of North herons in southwestern R. A .• W. D. Graul, bution in the fulmar s Rep .• Spec. Vol. Aspects 1970. Breeding in central Branch Conn. and development. Baltimore, egrets blue Urriv ; , Columbus. 1978. rookery Handbook Urriv , Montana. 1979. Poty raj , J. J. Pratt. Great nesting Thesis. 1962. New Haven. 1980. Nest failures 9pp. Editor. Univ . Press. Div , , Resour , Assessment 1980. Fish and Game. Wildl. Manage. Proj. program. 125pp. the eff ec ts of ob servers. Page. Parks protection of great Condor of common egrets Bay region. of Ciconiiformes herons and common 72:407-416. and G. C. Miller. 1979:49-58. blue and Condor 1979. in Colorado. great blue 74:447-453. Status. distri- Proc , Colonial �126 Sandtlarids , A. P. herons. Schrupp. 1980. Blue Jay D. L. habitat The wildlife values as related Chapter. nesting structures to other habitats and stream habitat of lowland river in Colorado. in Colorado: blue 1912. A history and Co .• London. U.K. of the birds Pages and stream 42-51 in a symposium. The Wildl. Soc. and Colo. Audubon Sclater , W. L. for great 38:187-188. 1978. Lowland river Scott. Artificial Colo. Counc . Greeley. of Colorado. Witherby 576pp. To G •• and C. H. Wasser. 1980. plan ts for wildlife biologists. Checklist of North American The Wild1- Soc , , Washin gton , D. C • 58pp. Stephens. H. A. Kansas Teal, J. 1980. Acad , Sci. M. 1965. Wilson Bull. Temple. S. A. herons. Thompson. Group. Thompson, Nesting success 1969. and of egrets and herons A case of turkey Wilson Bul1. Trans. in Georgia. 1977. within vulture piracy on great blue 81: 94. Declines the floodplain in populations of the upper Group. 1978. Declines egrets in five midwestern of colonial waterbirds Mississippi River . 1977:26-37. in the population states. of great blue herons and Proc , Colonial Waterbird 1978: 114-127. R. D•• C. V. Grant. 1968. in Kansas. 83: 161-186. Proc . Colonial Waterbird great blue heron 77:257-263. D. H. nesting The great Cardiac grouping. responses Am. J. E. W. Pearson, of starlings Physiol. and G. W. Corner. to sound: 214:41-44. effects of lighting �127 Tremblay, J., and L. N. Ellison. on breeding Tubbs. A. A. Pages black-crowned 1980. U.S. K. Nat. 1969. Auk communities of western 96:364-369. of the Great forests Dep . Agr'ic , , For. Great blue heron Great blue heron in the prairie N. M. Idaho. provinces. 1979. M.S. G. D., State Plains. and grasslands Sere Gen. colonies in Alberta. Can. Field-Nat. Ecology of great Thesis, Editor. Univ. 1980. Tech. Can. Field- blue herons Idaho, Moscow. Geography Fort on the great cormorant colonies 87:427-432. on Silver Creek, 63pp. of Colorado. Collins. 2nd ed . Colo. 186pp. D. F .• E. McMahon, and M. Lertschuh , of human activities blue heron 1976. Some effects in Oregon. Wilson 88: 660-662. 0 Wiese. J. and double-crested Univ .• Dep . Economics. WerschkuL H. Auk 1976. Courtship and pair formation in the great egret. 93: 709-724. 1978. Pages Heron nest-site 27-34 in A. Sprunt Wading birds. J. Calif. W. Natl. 1970. selection IV, J. Audubon C. Ogden. Soc. Lincoln great and its ecological Res. blue heron D. E. of herons Proj. 1977. W-54-R-2. The feeding in southeastern effects. and S. Winckler. Rep. Dep . Fish and Game. Wildl. Manage. WildI. Invest. Willard, herons. of human disturbance 83:237-242. 1973. Wilburn. Effects INT-86. Vermeer. Bull bird 419-433 in Management Rep. Weaver. night Riparian for non game birds. Warren. 1979. eds , 7. rookery Branch study. Rep.. 1970. Spec. 20pp. ecology and behavior New Jersey. Condor of five species 79: 462-477. �128 Prepared by Diana K. Vos Graduate Research Assistant Approved by C1ait E. Braun Wildlife Research Leader �129- Colorado Division of Wildlife Wildlife Research Report January 1984 JOB PROGRESS REPORT State of Colorado Project N-I-R-2 Work Plan 4 -'-------- Job Development of a Preservation Program For Insular Populations of Prairie Grouse .~~------------------------------ 1 Period Covered: Author: 1 January '.-31 December' 1983 G. C. Miller Personnel: C. E. Braun, R. Calderon, G. C. Miller, T. J. Schoenberg, J. Sedgwick, Colorado Division of Wildlife; K. Webster, Colorado State University. ABSTRACT Greater prairie-chicken (Tympanuchus cupido pinnatus) lek densities in the Arikaree Study Area increased by 45% from 1982 to 1983, but lek extinction rates ranged from 29 to 43% annually. In 2 years of intensive study, 49 prairie-chickens have been captured, and 26 have been intensively monitored with radiotelemetry techniques. Fourteen nest sites exhibited a mean of 1.80 (+ 0.86) shrubs/m2, 10.86 (+ 7.58) tall and mid-grasses/m2, and 24.6% (+ 12.2) bare ground. No nests of radio-marked or unmarked birds known to have been initiated in 1983 were successful. Male prairiechickens were released on the Tamarack Prairie in July and November 1983. Continuation of lek surveys, habitat restoration activities, and transplants of greater prairie-chickens to the Tamarack Prairie is recommended. This Job Progress Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the Researcher. ��131 DEVELOPMENT OF A PRESERVATION PROGRAM FOR INSULAR POPULATIONS OF PRAIRIE GROUSE G. C. Miller P. N. OBJECTIVES 1. By 1986~ ascertain for 1 species of prairie grouse in Colorado the combinat::!.on(s) of habitat island size, condition, inter-island distance~ and number (by sex and age-class) of grouse needed to establish and maintain populations with a persistence probability of P > 005 and extinction time of T > 100 years. SEGMENT OBJECTIVES lao Review current literature on theories of population ecology and the population dynamics of prairie grouse and similar species. lb. Continue data collection on birth and death rates. and intrinsic growth rates for greater prairie-chicken study populations, using radiotelemetry (20 bir.ds), banding. and line-transect sampling during nesting, brood-rearing, brood dispersal. and winter flocking activities. 2a. Review current literature on ecology of small populations and insular populations. '2bo Continue data collection on relationships between habitat quantity, quality. arrangement, and study population levels and distribution, using lek surveys, radiotelemetry. line transect sampling, and vegetation data collection from occupied and unoccupied habitats. Data collection will be compatible with multivariate analysis techniques. 381. Develop a technique to transplant greater prairie-chickens prairie relicts unoccupied by prairie-chickens. to 3b. Monitor behavior of experimentally transplanted prairie-chickens with respect to colonization and persistence probabilities (mortality, natality, emigration, immigration). 4. Publish research results in the form of scientific papers, popular articles. aridjob progress and/or final reports. 5. Assist the management branch of the Colorado Division of Wildlife in applying research r.esults to the greater prairie-chicken restoration efforts on the South Platte Wildlife Management Area (Tamarack Prairie). �132 METHODS Annual protocols for this project are provided in Appendix Au Surveys to find leks in the intensive study area were conducted as described pr.-eviously (Miller 1982) and are described in more detail elsewhere in this report. Surveys for leks in other portions of Yuma, Washington, Phi-llips, and Logan counties were conducted by Northeast Region, CDOW personnel from 1981 to 1983. Locations are given in unpublished reports by B. F. Van Sant (1981, 1982, 1983), in the files of the Northeast Region, CDOW. Prairie-chicken distribution was delineated using lek presence as the primary criterion. Areas were classified as unoccupied by prairie-chickens if leks were not present and if winter flocks or other evidence of prairiechicken use was n.ot found. Nearest-Iek distances (NLD) were calculated a.nd 2 (NLDmax) was used to classify leks as belonging to the same or different populations. With this criterion, there were 3 populations of leks i.n the study area--differentiated as "red", "yellow", and "blue". Data from this investigation were .compared to the findings of Swope (1953), Evans (1964) ~ and unpublished surveys from CDOW files (Miller 1983). Population ind1.ces were computed from previous studies and compared to the lek surveys of 1981-83. Beginning in December 1981 and continuing into this reporting periodp techniques of trapping and radiomarking greater prairie-chickens were evaluated. The techniques included night-lighting, walk-in funnel traps, and cannon nets over bait or on leks. All birds captured were weighed9 inspected for external characteristics of sex and age, and banded with a uminum leg bands. Birds not fitted with radio transmitters were banded with colored leg bands. Radio transmitters used (model SPCB-1250-3X) were solar-powered units manufactured by Wildlife Materials, Inc. (Carbondale, Ill) and weighed approximately 16 g without the attachment apparatus. Additonal details are given elsewhere in this report. LocatiollS of radio-marked greater prairie-chickens were ascertained from directional readings of radio signals at predetermined "locator points" which were marked on the ground with vinyl flagging and recorded on 7.5--minute USGS topographic maps. Directional readings, taken with hand-held compasses (liquid-filled Suunto or Silva models), were recorded from a minimum of 3 locator points and plotted on topographic maps (Appendix B). The resultant polygon (usually a triangle) was considered usable for analysis if all signals used to generate the polygon were recorded within a I-hour period. Vegetation measurements relating to habitat use followed procedures in Appendix C. Other vegetation mea.surements were collected between. 1981 and 1983 as described by Miller (1981, 1982). Ancillary data regarding other vertebrates of the study area were collected to the extent such aCl:ivities did not measurably impact the primary study objectives. DESCRIPTION OF AREA General study area descriptions have been provided previous y (l'tl.ller 1981). The intensive study area for greater prairie-chickens, selected �133 in January 1982~ was in Yuma County in northeastern Colorado (Fig. 1). The study area encompasses that portion of Yuma County lying north of the Arikaree River~ bounded on the west by State Highway 59 and on the north by County Road 26. The eastern boundary is irregular, delineated by County roads AA, Z, 20, Y, 23, and X. The st~dy area is 650 km2 in area and has as its potential natural vegetation Kuchler's (1964) sandsage-b1uestem (Artemisia-Andropogon) prairie type. It is referred to as the Arikaree Study Area. Soils are sands of the Valent series and sandy 10ams of the Vona and Dailey series. Mean growing season precipitation is 36 cm annually. Some work was performed at the South Platte Wildlife Management Area in Logan County. Approximately 1,500 ha of the area, lying south of Interstate Highway 76, is referred to as the Tamarack Prairie and was described by Miller (1980). RESULTS AND DISCUSSION Survey and Census Several techniques have been used to inventory greater prairie-chickens in Colorado in the past. These techniques varied among years and areas, depending primarily upon fiscal and human resources available, and the objectives of the inventory. Inventories included extensive voluntary "listening routes" conducted by interested private citizens, intensive 100% sur~ws of 2-mi2 (5.2-km2) areas selected with a stratified random sampling scheme, extensive 100% surveys of large units of potential or suspected prairie-chicken habitat (E. F. Van Santo unpubl. rep. Colo. Div. Wi1dl. 1983) and systematic surveys of smaller areas (Miller 1983). Data collected during the current study may serve to develop a standardized technique for use in monitoring greater prairie-chicken population trends. Data have been assembled that reveal human resource requirements for desired intensities of survey. Time of Survey.--Greater prairie-chickens have been found on leks each month except August. Although low-intensity vocalizations occur in autumn and early spring, it is not until approximately the last 10 days of March (1982, 1983) that the "booming" sounds can be discerned at I-mile (1.6-km) and greater distances. Additionally, males seem to congregate on a small proportion of leks until the latter part of March; the numbers of males at these leks then declines rapidly and additional leks become occupied at about the same time. Numbers of males 011 leks stabilize by early April. Patterns of hen attendance have varied between years, but peaks of attendance occurred in mid-April in 1982 and 1983 (Fig. 2) • Vocalizations are most constant from approximately 0.5 hour before sunrise to 1~ sometimes more than 2 hours after sunrise from the latter part of March to mid-May (Fig. 3). Periods of quiet ranging from 3 to 105 minutes �134 . GREATER PRAIRIE-CHICKEN DISTRIBUTION ARIKAREE STUDY AREA, 1 11 GREATER PRAIRIE-CHICKEN DISTRIBUTION ARIKAREE STUDY AREA, 1812 VERNON HI!!ARTSTRONO Hl!AftTSTRONO LOCATION IIA' o e A Aftft = •• • ~. a: ROUTE e 0 I!! U. S. • • • e 38 lit • U. S. ROUT!! 0 0 31 JOEl I I ! 1£ ; i' aD':' ~------~----=-----------------I '! t I ; I I ~ OCCUPII!D !, i' ~£----------------------------= YUMA COUNTY • I Q COLORADO KILOllnus , I '! I ; " LOCATIOII • YUMA COUNTY t 3 4 5 IIU& I ~ I IIILOMITUS COLORADO OCCU •••• D U!II LOCATIOII LI!II LOCAnOIl o UIIOCCU,oeD GREATER PRAIRIE-CHICKEN DISTRIBUTION ARIKAREE STUDY AREA, 1983 VERNON HEART STRONG o _--, o • o 4» • ~~ •• ~I\I",,,"EE U. S. ROUTE 38 JOES I " • 0 , 2 a • 0 MILlS L1.!!.U-'~_~I!.~ o YUMA COUNTY COLORADO _ OCCUPI D LEK LOCATIOII UNOCCUPllD LEA LOCATION Fig. 1. Greater prairie-chicken intensive study area and lek locations, 1981-83. �100 90 ~ W 80 Z 70 CIJ 60 o:t: ~ 50 ....J 0 W ....J ~. W I' I ' U. 40 .1Z W :\ I I 30 W , , I ~ o II: , I ,, \ \, 20 a, ~ 10 . '\ \,. _A ~-'""""""' FEB Fig. 2. MAR APR p--o ," .."d ,: • MAY Pattern of greater prairie-chicken hen attendance on leks, and 1983 (0-- -<l). 1982 (--'i) have been observed at other times during the mornings. Raptors passing over a lek may cause the birds to remain quiet or to flush from the lek. Usually~ booming will resume soon after a raptor passes, but it mayor may not take place at the usual lek location. Resource Reguirements.-- Surveying for leks of greater prairiechickens is time-consuming. Mornings unsuitable for survey due to rain, impassable roads~ and high winds are frequent, having ranged from 36 (1981) to 57% (1983). If the objective is to search previously unsurveyed areas for leks and also locate and count birds on all leks, approximately 13 mi2 (33.7 km2) may be surveyed each morning (from data by B. F. Van Sant, unpubl. rep., G6lo. Ddv , Wild!. 1983). Surveys of areas for presence and location of leks only, in areas familiar to the observer may cover 23 ~ 30 mi2 (59.6 - 77.7 km2) per suitable morning at a minimum (this study). Detection Distances and Lek Densities.-- In the Arikaree Study Area, the greatest distance at which males have been heard booming was 2.3 mi (3.7 km). Maximum detection distances measured perpendicularly to routesof-travel were 1.9 ± 0.3 mi (3.1 ± 0.5 km) on mornings with winds �136 SUNRISE ·0.0 -0.1 LL rJ) ·0.2 o t5 ·0.3 C) i= ·0.4 ~ ~ ·O.S Z .0.6 ~ -;J_ ·0.7 C) 0 .O.S Wo CO> .0.9 ·1.0 ·1.1'-'1 S----'-=----l.'::-S---'-----:'l'::-S ----:--~1:':S--:30 MAR a MAY JUN DATE 2.4 2.3 2.2 2.1 2.0 CJ) Z 1.9 0 1.8 i= 1.7 0:( 1.6 N 1.5 _I 1.4 0:( o 1.3 0 1.2 1.1 > 1.0 u, 0.9 0 0.8 0 0.7 Z 0.6 W 0.5 0,4 0.3 0.2 0.1 0.0 SUNRISE b APR 15 15 MAR APR 15 MAY DATE 15 30 JUN Figo 3. Beginning (a) and ending (b) of greater prairie-chicken morning vocalizations on leks with respect to sunrise, Yuma County, Colorado~ 1982-83. �137 < 3 mph (4.8 kph) (N=6) and 1.3 + 0.4 mi (2.1 + 0.6 km) with winds> 3-5 mph (4.8 - 8.0 kph)-(N:8). Dete~tion distance; were highly variable on mornings with winds> 5 mph (8 kph)~ and winds> 10 mph (16 kph) made conditions unsuitable for lek surveys. It may be possible to estimate lek densities from the percentage of listening stops at which booming is heard. Present data are inadequate to perform this task with high confidence (Fig. 4). Other variables would have to be tightly constrained. x o Z ~ 0 0 en <, x x S x x U) a, X 0 1-- X X x U) LL. 0 X 1-- Z X W o 0:: w a, X x x X X X X x 2 3 4 5 NUMBER OF LEKS DETECTED Fig. 4. Numbers of leks detected vs. percentage of listening stops where booming was heard, Arikaree Study Area, 1981-83. Listening stops were at 0.8-km intervals. �138 Recommendations for Surveys.-- Data collected to date do not indicate a need to stop and listen for prairie-chicken vocalizations at intervals less than 1 mi (1.6 km). To compensate for "quiet" periods and the influences of time-of-day upon booming, the practice of conducting routes in 2 directions should be continued. Beginning at mile 0, listen for booming for 3 minutes and proceed along the selected route stopping for 3 minutes at each 2-mi (3.2 km) interval to the end of the route. Retrace the route 1 mile (1.6 k~), stop, then proceed again at 2-mi (3.2 km) intervals to mile #1 of the route. A 12-mile (19.3-km) route, would theoretically require approximately 1.6 hours to accomplish, driving at 25 mph (40.2 kph). Surveys should begin no later than 0.5 hours before sunrise. Sunrise times (MST) are provided in Appendices D and E. The practice of actually locating the lek should continue with the location plotted on a USGS topographic map. Detection distances and angles can be used to compute lek densities (Burnham et a1. 1980, Gates 1980). Weather data should be recorded precisely. With adequate data a relationship might be developed relating weather, detection distances and lek densities 9 and percentage of stops with booming. Surveys should not be condu.cted when Winds exceed 10 mph (16 kph). A sample data collection form is provided (Appendix F). Management may desire to estimate the numbers of birds or numbers of males on leks although Carmon and Knopf (1981), Wells (1980), and R. J. Robel (pers. commu.n.) indicated this statistic was of questionable value. Such numbers vary greatly with time of year, time of day, weather patterns. and hen attendance patterns (Sisson 1976, this study). In sandsage-bluestem areas, I have been unable to gain reasonable sex ratio estimates without observing the lek for about 1-2 hours. A total~ accurate count of birds on a lek can be obtained only by flushing the birds from the lek, at which time ascertaining sex is difficult. Flushing birds from the lek carries the potential of an artificially high lek density estimate, since the flushed birds may establish 1 or more "temporary" leks, at distances up to 007 mi (1,.1 km) ,:cforthe remainder of the mornfng-v-Leks which _. might be eounted by an observer, If numbers are desired for leks, It may be desirable to use size (number) classes, estimated from a dicu.nce and without flushing the birds. One option to obtain numbers of males on leks is to count them on a sample of occupied leks in February before all leks are occupied and intensive "booming" begins. Surveys during the peak of activity, during the period of frequent "whooping" and flutter-jumping (which allows more efficient lek detection) could be directed solely toward estimating lek densities. Study Population Levels and Distribution In 1981, during the study area selection phase of the study, 23 active greater prairie-chicken leks were located during a 100% survey of the Arikaree Study Area (Table 1~ Appendix G). Another lek, in eastern Washington County, initially was considered for inclusion in the Lnt.enafve study area but was eliminated for logistical reasons. �139 Table I. Greater prairie-chicken lek turnover, Arikaree Study Area, Yuma County~ Colorado, 1981-83. 1981 1982 1983 Leks persisting from preceding year (%) No data Ii (48) 13 (65) Inactive leks, active in preceding year (%) No data 10 (43) 4 (20) Leks established or re-established No data Total active leks Net change from previous year (%) 9 14 23 20 29 No data -3 (-13) +9 (+45) In 1982. 20 active leks were located in a 100% survey of the Arikaree Study Area (Table 1, Appendix G). Eleven leks persisted and 10 sites were unoccupied fr m the previous year. Nine leks ~ere established for the first time during the study. The most notable non-occupancy of the previous years' lek sites was in the southeastern p()rtion of the:"blue" population. Reasons for lek abandonment were not obvious. Nearest lek distances were computed for the study populations, and NLDmax = 4.3 km (2.7 mL) (Appendix H). In 1983, 29 attive leks were located in a 100% survey of the Arikaree Study'Area (Table 1, Appendix G). Four sites active in 1982 were not occupied in 1983. Six lek si~es occupied in 1981 but unoccupied in 1982-,were re-occupied---in1983. Eight of 41 Lek sites active at S01ne time. from 1981 to 1983 persisted all 3 years. Mean NLD decreased from 1982 (3.0 km) to 1983 (2.0 km) but NLDmax increased from 4.~ km (1982) to 6.2 km (Appendix H). Trapping, Marking, and Radiotelemetry Studies In 1982-83, 49 greater prairie-chickens were captured. Two (1 male, 1 60-day-old chick, sex unknown) were captured by hand. The remaining 47 were captured with cannon nets on leks. All birds were banded with aluminum leg bands, weighed, and classifed by sex and age-class. Birds not fitted with radio transmitters were banded with colored, numbers, plastic leg bands coded for location and year of capture. Films of blood collected from 26 birds in 1983 were prepared. The sex of captured birds was ascertained from external characteristics. A total of 26 females~ 22 males, and 1 sex unknown (chick) has been captured. �140 Solar-powered radio transmitters, were I, J) and 8 males. A poncho apparatus transmitters placed on 4 females and 1 apparatus held the transmitters on all attached to 23 females (Appendices (Armstrup 1980) held the radi.o male in 1983. A backpack harness others (Fig. 5). Fig. 5. Backpack harnesg used to attach radio transmitters to greater prairie-chickens in Colorado, 1982-83. Numbers are measurements (mm) be~veen junctions in the harness apparatus for females. Transmitters were removed from 3 females (backpack harness) and 2 males (1 backpack, 1 poncho) for different reasons within 3 days of fitting. Thus~ 20 females and 6 males were intensively monitored for movement, habitat use, and fates. Radio-telemetry monitoring in the Arikaree Study Area ceased in July 1983 due to a combination of budget reductions and the beginning of transplant activities on the Tamarack Prairie. At that time 4 radio-marked females were alive~ One female's radio .was.known to have malfunctioned in 1982. The minimum longevity of the remaining 15 females varied between 1982 and 1983 (Table 2). Three radio-marked tn.":lles were experimentally transplanted to the Tamarack Prair:i.ein 1983; 2 males monitored in the Arikaree Study Area in 1982 were known to have been killed, and the fate of 1 male is unknown. In 1982, 343 different locations were obtained for radio-marked females (Appendix I). Characteristics of 6 nest sites and 25 brood-rearing sites �141 were collected (Miller 1983). Vegetation characteristics were measured for 90 location polygons. In 1983, 113 different locations were obtained for radio-marked females (Appendix J). In 1982-83~ 14 nests of radio-marked greater-prairie chickens were found. In addition, 3 nests of non-radio-marked hens were found and 2 radiomarked hens were killed during laying. Clutch size ranged from 5 to 16 eggs. Nesting success for 6 nests in 1982 was 33.3% (2 nests) at a minimum. A 3rd nest may have hatched successfully in 1982--a severe hail storm on the evening of the 25th day of incubation reduced the nest contents to shell fragmeIltsp but the hen's movements away from the nest were similar to those of a hen with a young brood. Intensive searches for the brood were conducted later. but no young were found. ,At least 1 hen (adult) renested after its nest was destroyed during incubation. Nesting success for 19 nests known to have been initiated in 1983, was zero. Distances between nests and the nearest active lek ranged from 0.2 to 4.7 km. A hen experimentally transplanted from the "yellow" to the "red" population accounted for the maximum distance. The mean distance between nests and the nearest active lek, experimental transplant excluded. was 1.3 + 0.7 km (range 0.2 - 2.4) (N=18). Characteristics of vegetation at nest-sites were measured (Table 3). Table 2. Survivorship of greater prairie-chicken females after attaching radio transmitters, 1982-83. Hen# 1982 Min. days alive SR 238 236 188 69 180 403 lOR llR 14Ra. 19R 21R Mean SD 249 90 Henll 27R 29R 39R 40R 41R 45R 46R 47R 102R 1983 Min. days alive 69 38 32 42 32 19 2S 06 21 32 18 �142 Table 3. Characteristics of greater prairie-chicken nest sites, Yuma County, Colorado, 1982-83 (N = 14). Mean SD Range Tall ~rasses illmL. ae, m Midgrasses 111m2 ne , m Tallgrass and midgrass combined 111m2 Ht , m o- 4.79 0.64 13 0.05 - 1.02 6.07 0.38 4.94 o - 18 0.24 0.05 - 1.04 lO.86 0.61 7.58 0.24 o - 25 0.29 - 1.04 1.80 0.45 0.86 0.14 1 - 4 0.26 - 0.76 24.:6 12.2 5.0 - 40.0 Shrub 111m2 Ht, m Bare ground, % Experimental Transplants On 19 April 1983, 1 AHY female greater prairie-chicken was caught by cannonnet at 0542 hours at the "yelLow" -S29 Lek , She was held all day in a dark box. At 1845 hours,after processing and fitting her with a radio transmitter, the female (#39R) was released approximately 13 km from the place of capture. The release took place in another portion of occupied range between "red" S36 and 35 leks in an area known to bc: used by females, and which subsequently was found to contain a high densLty of prairie-chicken nests. For 3 days following release, the female remained within 2 km of the release site and visited habitats such as lightly grazed pastures (2.5 .ee] AUM) and sublrrigated meadows used by resident birds. On day 4 postrelease, the weather changed from the rain and drizzle of the previous 2 days to sunny and calm. At 1000 hours on 23 April, I found the female beyond the range normally occupied by prairie chickens. She was on the south side of the Arikaree River in an area dominated by clay soils, short grasses p and yucca (Yucca _glauca)• I flushed her and she flew northwest toward the sandsage-bluestem prairie on the north side of the Arikaree. Radio contact was lost except for 1 occasion between 24 April and 02 May. Contact was re-established on 03 May--the female was aga:f.n in the area usually unoccupied by prairie-chickens. The remains of the fenmle were found at the nest on 25 May. but no egg remains were found. �143 She had been killed apparently by a canid. between 20 and 25 May. The nest of female 39R was farther from an existing lek (4.7 lan) than any other nest found. The distance from point-of-release was 8.5 km , the 2nd greatest movement of all hens monitored in 1982-83. Even though the hen was released into an area with resident birds, she displayed a tendency for erratic movements and a failure to use habitats used by resident birds" She did establish a nest and did not make the large movements reported for transplants into unoccupied range (Kruse 1973). On 01 July 1983 following several days of non-attendance at leks by prairie-chickens, a loop tape of booming was played on "red".S 36 lek and a taxidermy mount of a female was placed on the lek in front of a cannonnet. Three males flew to the lek and 2 were captured. The males were held in a darkened box all day. At 2015 hours, after processing and fitting the males with radio transmitters, the birds were released on the Tamarack Prairie. The birds ran and flew a short distance. One male, 49R, remained near the release site at least through 15 September. All locations except 2 were within 0.8 km of the release site. The bird moved up to 2.7 km from the release site between 15 and 26 July, but returned to the original release site. The second male, 48R, remained within 3 km of the release site for ppproximately 1 week post-release. By 15 July, the bird was off the Tamarack Prairie. approximately 9 km from the release site. At least through 08 September, the bird was inhabiting a I-section ungrazed pasture, 9.3 kIn from the release site. The summer release demonstrated that, for males at least, birds can be trapped and transplanted later than is normally done (Kruse 1973). The technique may tend to reduce the initial long movements observed with other prairie grouse transplant techniques. Prairie-chickens molt during summer and habitat cover values should be at or near their maxima at that time (Jo Toepfer, pers. commun.). Applying the summer transplant technique to hens and/or hens with broods would be time-consuming, but appears worthy of investigation given the high failure rates of autumn, winter, and spring transplants in other areas (Kruse 1973, Lawrence and Silvy 1981). Development of a transplant technique for prairie grouse may be one of the most pressing need for prairie grouse management in Colorado. Several Nongame Program objectives depend upon being able to move birds into unoccupied range with reasonable expectation they will remain in the vicinity. A population. of greater prairie-chickens needs to be established on the Tamarack Prairie to justify further habitat restoration activities and to begin ascertaining population parameters for the species. The Southeast Region, CDOW, is investigating potential sites for the reintroduction of the plains sharp-tailed grouse (Tympanuchus phasianellus jamesi) (C. Loeffler~ pers. commun.). Personnel of the Northeast Region, CDOW, have expressed �144 interest in attempting to establish prairie grouse in other areas of sandsage-bluestem prairie along the South Platte River (C. Leonard , pers. commun.). Other Activities and Ancillary Data Data collected on bird species occurrence in the Arikaree Study Area during 1981-83 were summarized in 1983 (Appendix K). Data exist, but have not been summarized for location and habitat of upland sandpipers (Bartramia longicauda) on the Arikaree Study Area, 1981-83. In 1982, several prairie~chi~kgn nests were destroyed--the eggs remained in the nest CUP9 virtually undisturbed in position, but the e;xposed egg surfaces had been removed. Another nest may have been destroyed by a severe hail storm •• To discover if nest losses in which eggs were not physically removed from the nest could be prevented, 4 clutches of eggs were remo"1ed from nests of incubating hens. The same number of plastic replicas were placed in each nest, in the same position as the live eggs had been. Colors and egg markings were matched to the extent possible. The live eggs were art~ficially incubated. In 2 instances, clutch replacement occurred during the period the hens were off the nest feeding, as part of their normal activities. In 1 instance, a hen vIas flushed from her nest by the investigator and the clutch was replaced. In these instances, the hens returned and continued normal incubating behavior. In 1 instance a hen was accidentally flushed from her nest, the clutch was replaced, and the hen was never known to return to the nest. She was found dead 2 days later. This hen may have been killed while off the nest during replacement. Alternatively, she may have been a renesting hen or been in the early stages of incubation either of which may have contributed to her failure to return to the nest. p All 3 manipulated nests to which females returned subsequently suffered coyote or mustelid destruction. The plastic eggs were gnawed, 1 was missing from the nest area, and all were physically removed from the nest cup. I replaced the eggs in the nest as soon as I detected destruction, but none of the hens was known to have returned to the nest. I was able to ascertain the dates ..o f incubation start and subsequent' nest destruction for 7 ~ests. The 3·manipulated nests survived the longest period of time, (7~12, 19-·22, and 17 days of a 25-·' day incubatidn period). Four non-manipulated nests were destroyed at 1 to 8 days of incubation. If the hens on the plastic replicas had been exhibiting incubating behavior at the time their artificially-incubated clutches had pipped, the pipping eggs would h rve been placed in the nest and the plastic repUcas removed. A contingency plan was to trap the hen on the nest and reunite the hen with live chicks in specially-designed brooder boxes, monitoring for hen acceptance of the chicks prior to release. A positive result would have carrted implications for efficient study of the brood-rearing phase of �145 prairie-chicken life history, and might have had utility in attempts to transplant hens with broods. The results of this exercise, however, were inconclusive. On 08 November 83 T. Davis captured an BY male greater prairie~chicken in Sterling, Colorado. I processed, banded, and placed a radio transmitter on the bird, and released it on the Tamarack Prairie 09 November 83. Subsequent searches failed to yield locations of this bird. In 1983, a review of the greater prairie-chicken research was performed during the CDOW's Research Audit in March and another review 6f the project took place in September. The latter review was performed by Leo Kirsch, Woodworth, N. D. (Appendix L). A prospectus was developed in November 1983 for a community study of sandsage-bluestem prairie (Appendix M). Recommendations 1. Lek survey activities should continue, both to serve management and research objectives. a. Samples of leks from populations exhibiting high and low lek turnover rates should be monitored annually. b. Lek routes should be as straight as possible--corners and curves may cause misrepresentation of total area surveyed. c. Routes should be conducted in 2 directions on a given morning, stopping at alternate listening posts for a minimum of 3 minutes each. d. Surveys should begin no later than 0.5 hours before sunrise and continue until at least 1 hour after sunrise or until winds exceed 16 kph , e. f. g. 2. Surveys should not be conducted if winds exceed 16 kph, winds < 4.8 kph are most preferable. Surveys may be conducted from late March until mid-May, but leks are most easily detected during April. Lek locations should be plotted on USGS topographic maps and counts or estimates of numbers of birds present may be made. Leks should not be flushed unless safeguards against counting "temporary" leks exist. Methods used to trap, radio-mark, and monitor prairie-chickens should undergo constant evaluation, modification, and improvement. a. Alternatives to cannon-netting should be evaluated with respect to ability to capture females and manpower requirements. b. Refinements to radio transmitter attachments should be sought which reduce weight and/or cumbersomeness and/or visibility of radio transmitters on hens while retaining current transmitter output. c. Monitoring of radio-marked hens from pre-nesting to the incubation phase should result in 2 precise locations/week--obtained by quietly circling the marked bird at 60-80 meters. Three or more locations/week may be less precise--ascribing the birds' Lccat.Lon �146 d. to a 1/8 section. Flushing the marked bird or driving near marked birds should be avoided. Monitoring of incubating radio-marked hens should be performed daily from a distance of 400 m or more. At least 1 all-day (sunrise to sunset) monitoring should be performed to ascertain the hen's foraging area-and daily activity pattern. In the latter stages of incubation~ the hen should be monitored twice/day. The hen should be circled only once during this period to ascertain nest location within roughly 0.3 - 0.5 ha area. The hen should not be flushed from the nest during this process, and the approach should be made in such a manner to minimize the possibility of observer-induced nest depredation. 3. Transplants of greater prairie-chickens to the Tamarack Prairie should continue. a. Male and female prairie-chickens should be captured on leks in April 1984. A sample should be radio-marked. b. The least labor-inten.sive technique and the one which can be evaluated soonest is to release the captured birds on the Tamarack Prairie the same day. c. An alternative technique~ or one which could serve as a contingency plan, is to release radio-marked birds at the place of initial capture. Hens which nest successfully could be recaptured with their broods and transplanted. Unsuccessful hens and males could be recaptured and transplanted in summer, at about the time of molt onset. This technique would require increased resources. d. Another alternative is to trap a large number of males (ca. 30). in June or early July and transplant them. The following spring, a number of these birds can be expected to have established leks on or near the Tamarack. Additional males and females trapped that spring wou l.dbe released at those already active leks (J. Toepfer, pers. commun.). 4. Habitat restoration activities on Tamarack Prairie should continue as if a breeding population of prairie-chickens was present. Activities such as seeding and burning may take several years to yield positive results. Windmills and the accompanying moist overflow areas should be kept operational permanently. Whether or not transplanted birds remain on/near the Tamarack in the first few days following their release will probably dictate the success or failure of a transplant effort. Birds failing to detect appropriate cues from the habitat can be expected to make erratic movements into unsuitable and insecure habitats? and suffer abnormal mortality rates. LITERATURE CITED Amstrup9 s. C. 44:214-217. 1980. A radio-collar for game birds. J. Wild!. Manage. �147 Burnham, K. P., D. R. Anderson, and J. L. Laake. 1980. Estimation of density from line transect sampling of biological populations" Wildl. Monogr. 72. 202pp. Cannon, R. W., and F. L. Knopf. 1981. Lek numbers as a trend index to prairie chicken populations. J. Wildl. Manage. 45:776-778. Evans, K. E. 1964. Habitat evaluation of the greater prairie chicken in Colorado. M.S. Thesis, Colo. State Uriiv., Fort Collins, 98pp. Gates, C. E. 1980. Linetran, a general computer program for analyzing line-transect data. J. Wildl. Manage. 44:658-661. Kruse, A. D. 1973. Prairie chicken restoration projects. Pages 40-46 in W. D. Svedarsky and T. Wolfe, eds. Proc. Conf. Prairie Chicken in Minnesota, Univ. Minnesota, Crookston. Kuchler, A. W. 1964. Potential natural vegetation of the conterminous United States. Am. Geogr. Soc. Spec. Publ. 36. 39pp. Lawrence, J. S,.; and N. J. Silvy. 1981. Movements and mortality of transplanted Attwater's prairie chickens. Proc. Prairie Grouse Tech. Counc , Conf. 14:7-8 • Miller, G. C. 1980. Development of a preservation program for three species of prairie grouse, Job Prog. Rep., Colo. Div. Wildl., Wildl. Res. Rep. Jan. Pp. 76-94. 1981. Development of a preservation program for three species of prairie grouse. 'Job Prog. Rep., Colo. Div. Wildl., Wildl. Res. Rep. Jan. Pp. 38-52. 1982. Development of a preservation program for insular populations of prairie grouse. Job Prog. Rep., Colo. Div. Wildl., Wildl. Res. Rep. Jan. Pp. 63-71. 1983. Development of a preservation program for insular populations of prairie grouse. Job Prog. Rep., Colo. Div. Wildl., Wildl. Res. Rep. Jan. Pp. 33-62. Sisson, L. 1976. The sharp-tailed grouse in Nebraska. Nebr. Game and Parks Comm., Lincoln. 88pp. Swope, H. M. 1953. Surveys to determine the population status of the prairie chicken. Job Completion Rep., Colo. Dep. Game and Fish, Proj. W-37-R-6. Pp. 77-80. Wells, R. 1980. Kansas prairie chicken annual report, 1979. Fish and Game Comm. Fed. Aid Proj. W-23-R-18, 36pp. Prepared by Go..~ ~. lYJ:..~ Gary t. Miller Wildlife Researcher C • Kans. �148 APPENDIX A Annual protocols: populations. habitat-quantity relationship to prairie-chicken Month Major Activities Jan P.I.a/_ Contact landowners or representatives for previous season grazing data (pasture-by-pasture cattle numbers, entryexit dates)~ public relations effort (explain findings, proposed activities, identify potential problems for up to 29 ownerships. Feb P.I. - Continue grazing data collection, begin field preparations~ hire temporaries, draft progress report. Mar P.I. - Draft progress reports,. orientation and training of temp'oraries. T.~7 - Preparation for trapping. Collect population and distribution data (lek surveys), equipment transport to Yuma County. Apr P.I. - Trap, process, mark prairie-chickens. Final draft progress reports, train temporaries for radiotelemetry work. Quality-control of data collection: Test transplant techniquescollect nesting data. T. - Collect population and distribution data (lek surveys). Monitor radio-marked birds. Collect habitat use data. P.I. - Quality-control of data collection. Test transplant techniques. Collect nesting data. T. - Collect population and distribution data (lek surveys). Monitor radio-marked birds. Collect habitat use data. Count cattle numbers and record entry dates (quality-control procedure) for selected pastures. Jun P.I. - Quality-control of data collection. Test transplant techniques. Collect nesting data. T. - Collect habitat use and brood data. Monitor radio-marked birds. Jul P.I. - Quality-control of data collection. Test transplant techniques. Collect nesting data. Year-end reports. T. - Same as June. Aug P.I. - Data storage, summary. Equipment replacement. Test transplant technique. T. - Equipment storage at Fort Collins. Equipment maintenance, repair. �149 APPENDIX A (cont.) Month Major Activities Sep P.I. - General project administration, data summary, and analysis. Writing. Periodic monitoring of transplants and radio-marked bird fates. T. - Vegetation measurements of selected areas. Oct P.I. - Maintenance level activities. Nov P.I. - Maintenance level activities. Dec P.I. - Project administration. Data analysis. Evaluation and revisions of techniques, protocols, budget, and human resource needs. ~I P.I. - Activities listed are those performed by the principal investigator. bl T ~ Activities listed may be performed by temporary employees or the principal investigator. These include the primary duties of temporary employees but may not. overall, be done primarily by temporaries. �150·· APPENDIX B PRAIRIE GROUSE INVESTIGATIONS Radiotelemetry-monitoring 1. Preparation A. Equipment 1. Kit 2. 3. 4. B. II. of Radio-marked Birds receiver, antenna, headphones data recording forms Locator Point map listing of frequencies, leg band numbers land ownership map Compass M2 frame and vegetation forms Long-handled net Review recent data forms for general locations of birds 1. Follow established schedule, guidelines in selecting "target" birds. Data Collection - always record data as they occur r: A. Know whose land you are one 1. Has permission been given (through principal investigator)? 2. Does owner prefer "concentrated" or "dispersed" prairie-driving? 3. Guard against vehicle-start fires. 4. Go only through established gates, leave gate as you found it, even if you intend to return soon (you may not). a) If gate found open, but cattle in pasture, note and contact landowner when possible. S. Minor fence repair should be done as a "good-neighbor" gesture. 6. Major fence repair, sick cattle, other problems-contac£ landowner same day. B. Obtain moderate-to-strong signal from radio-marked bird at an established Locator Point. I. If established Locator Point is inappropriate, establish new one, mark and label on topographic sheet. 2. Locator Points should be: on high ground sufficiently unique to guard against error in map location"no slip" on/near improved !Coad or 2-track 3. Record Locator Point, frequency, magnatic compass reading 00 form (attached). a) The most accurate readings are made with the antenna in a horizontal position and using the "null" for df.rec t Lon , b) Guard against bias - establish signal direction "blindli 4. Move to next Locator Point and repeat. �151 APPENDIX B (Cont) 5. C. Note: The best location polygons result when the Locators Points approximate 1200 intervals around the bird. If bird is flushed for any reason, take vegetation measurements with the "1 + 4" method at that time. Record polygon 10 Using the field forms and the Rotangle Protractor, draw the signal direction lines from the appropriate Locator Points (convert magnetic reading to Az). Number the resultant polygon on the data map and record in the appropriate bird's file. a) Polygons are numbered sequentially, preceded by the aluminum leg band number of the bird. Therefore, a hen carrying transmitters 150.920 with aluminum band number 8 will have all polygons labeled with an "8" prefix (8-1, 8-2, 8-3, etc.) 2. Be sure to initial the data collection forms upon completion of the procedures. �152 APPENDIX C Mar 82 GCM PRAIRIE GROUSE STUDY VEGETATION MEASUREMENTS 1. II. Preparation: A. Work straight 8 hour shift; B. Equipment: Appropriate field form compass tape measure, stakes M2 frame flagging veg. working map or_radio-location map Field Data Collection A. Locate polygon boundaries (or other sampling unit), flat. B. Reconnoitor sampling unit for sign of roosting, feeding, dusting, nesting. L If precise activity site found, use the "1 + 4" technique (M2 centered on site, then measure at 5-m distance in cardinal directions). In either case, proceed with the following: C. For sampling unit, record dominant slope, aspect (AzO), topography. D. Ascertain and flag ends of longest axis of unit. 1. E. Generate 3 random numbers compatible with length of long axis (i.e., if axis length in meters is a 2-digit number, generate doublets' if 3-digit number, generate triplets). Measure and mark these 3 points on axis, measuring from the southernmost extremity of the axis (i.e., the end pointing at >900, <2700 Az). Establish transects across entire unit, oriented NE-SW, intersecting long axis at flagged points. Take m2 readings at IO-m intervals along each transect, beginning at point 0 at the most southerly end of the transect. �153 APPENDIX D Sunrise Times for Arikaree Study Area, Yuma Co., Colorado - Mountain Standard Time. Day May Apr May Juri 01 02 03 04 05 06 07 08 09 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 0624 0622 0621 0619 0618 0616 0615 0613 0612 0610 0608 0607 0605 0604 0602 0601 0559 0557 0556 0554 0552 0551 0549 0548 0546 0544 0543 0541 0540 0538 0536 0535 0533 0532 0530 0528 0527 0525 0524 0522 0521 0519 0518 0516 0515 0513 0512 0510 0509 0507 0506 0504 0503 0502 0500 0459 0457 0456 0455 0454 0452 0451 0450 0449 0447 0446 0445 0444 0443 0442 0441 0440 0439 0438 0437 0436 0435 0434 0433 0432 0431 0431 0430 0429 0428 0428 0427 0427 0426 0425 0425 0424 0424 0424 0423 0423 0423 0422 0422 0422 0422 0422 0421 0421 0421 0421 0421 0421 0421 0421 0422 0422 0422 0422 0422 0423 0423 0423 0424 0424 0424 0425 �.154' APPENDIX E Sunrise Times for Wray, Yuma Co., Colorado - Mountain Standard Time Day 01 02 03 04 05 06 07 08 09 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 '" 31 •. Mar Apr May Jun 0622 0620 0619 0617 0616 0614 0613 0611 0610 0608 0606 0605 0603 0602 0600 0559 0557 0555 0554 0552 0550 0549 0547 0546 0544 0542 0541 0539 0538 0536 0534 0533 0531 0530 0528 0526 0525 0523 0522 0520 0519 0517 0516 0514 0513 0511 0510 0508 0507 0505 0504 0502 0501 0500 0458 0457 0455 0454 0453 0452 0450 0449 0448 0447 0445 0444 0443 0442 0441 0440 0439 0438 0437 0436 0435 0434 0433 0432 0431 0430 0429 0429 042"8 0427 0426 0426 0425 0425 0424 0423 0423 0422 0422 0422 0421 0421 0421 0420 0420 0420 0420 0420 0419 0419 0419 0419 0419 0419 0419 0419 0420 0420 0420 0420 0420 0421 0421 0421 0422 0422 0422 0423 �155 APPENDIX F LEK SURVEY FIELD FORM Data Processed: Date By ------_ CHECKLIST Pens Topo maps Compass Thermometer Wind gauge Psychrometer Binoculars ROUTE: DATE: --:-da. mo. yr. TIME: 'End start Elapsed Time INVESTIGATOR: MILE - Booming No Booming 0 2 0 4 0 6 0 8 0 10 0 12 0 11 0 9 0 7 0 5 0 3 0 1 SUNRISE Temp. _~ __ Wind Speed Direc. Rel. Humid. o- 0 ~~ _ _ 0 Direction (Mn) Location (mark on map) Size class!!./ ~I If complete, accurate count, circle number, otherwise classifyas: 0-5, 6-10, 11-20, >20. �156 APPENDIX G LEK LOCATIONS AND STATUS,a ARlKAREE STUDY AREA, 1981-83. Lek location SE NW NW NE SW SW SE NE SE NW SE NE NE sm NW NW SE SE NE SE NW SW NW NE SE NW SW NW SE NE SW NE SE NE NE NW 1 T3S R48W 3 T3S R46W 4.T4S R47W "5·T3S R45W 5 T4S R47W 6 T3S R46W 6 T4S R47W 7 T3S R47W 9 T3S R47W 11 T2S R46W 11 T3S R46W 11 T3S R47W 12 T3S R48W 13 T3S R47W 18/SW 17 T2S R45W 18 T3S R46W 19 T3S R45W 19 T3S R45W 19 T3S R47W 20 T3S R46W 20 T3S R47W 21 T2S R45W 21 T3S R47W 22 T2S R47W 23 T3S R46W 25 T3S R46W 26 T2S R47W 27 T3S R47W 28 T3S R47W 29 T2S R47W 29 T2S R47W 30 T2S R45W 30 T2S R46W 30 T3S R45W 33 T3S R47W 34 T2S R47W 34 T3S R47W 34 T3S R47W 35 T2S R46W 35 T2S R47W 35 T3S R47W 35 T3S R47W 36 T3S R47W 36 T3S R47W 1981 1982 1983 + + 1984 1985 nd + + nd nd + + + + + + + + + + + + + + + + nd nd + + + + + + + + + + + + + nd + + + + + nd + + + + nd + + + + + + + + + + + + nd + + + + + + + + + + + + + + + + + + a Lek locations classified as active (+), inactive (-), or no data collected (nd). �APPENDIX LEK LOCATIONS, 1982 Nearest Location 1*1 19, SE I, SE 29, SE 22, NE 28, SW 34, T3S, T3S, T2S, T2S, T3S, T3S·, NE II, T3S, SW 6, T3S, NE 36, T3S, NW 20, T3S, SE 35, T3S, unoccupied unoccupied unoccupied unoccupied unoccupied unoccupied unoccupied unoccupied unoccupied unoccupied unoccupied unoccupied unoccupied unoccupied unoccupied - DISTANCE lek TO NEAREST unoccupied . I,.T3S, 29, T2S, 22, T2S, 29, T2S, 34, T3S, 28, T3S, R48vl R4m R47H R47W R47W R47W 4.3 4.0 3.9 3.9 2.0 2.0 R47W R46W R47W R46W R47W SW NE SE NE NE 6, 11, 35, 36, 36, R46W R47W R471, R471, R47W 2.7 2.7 1.7 4.1 1.7 --- - Q Precise locations 2 leks in 1983. - - - and, therefore, -- - - .-unoccupied - ---- -- - - - SE 7, T3S, .R47W SE 20, T3S, R47W NE 21, T3S, R47W 1*1 NE NE NW NE NW SW SW 13, 35, 18, 29, 5, 4, 26, 30, .T3S, T3S, T3S, T2S, T48, T4S, T2S, T2S, R47W R47W R46W R47W R47W R47W R47W R46W ----- NW 27, NE 34, NE 35, Nt, 23, .T3S, T3S, T2S, T3S, R47W R47tV R46W R46W - nearest-lek 1982-83a D(km)· lek SE 7, T3S, R47vl NW 29, T2S, R47W SW 26., T2S, R47\-1 2.7 0;9 1.6 Nt, NE NE NE NE SE .NE SE 13, 11, 35, 18, 35. I, 21, 20, T3S, T3S, T3S, T3S, T3S, T3S, T3S, T3S, R47W R47W R47W R46.1 R47W R48t, R47W R47W 2.0 2.8 1.4 1.3 0.7 2.7 1.5 1.5 NE SE NW SE S~ NE SW SW II, 35, 20, 29, 33, 5, 30, 6, T3S, T3S, T3S, T2S, T3S, T4S, T2S, T3S, R47W R47W R46W R47W R47W R47W R46W R46W 2.0 0.7 1.3 0.9 2.1 1.2 .3.2 2.6 NE NW NW NE 34, 27, 23, 19, T3S, T3S, T3S, T35, R47W R47W R46W R45W 1.3 1.3 6.2 4.5 unoccupied ------- - STUDY AREA, 1983 Nearest Location SE SE SK SE SW NE -- LEK, ARlKAREE D(km) R47.1 R48W R47W R47H R47W R47W T3S, T3S, T3S, T3S, T3S, II distances I were not obtained for 4 leks in 1.98.2; I-' \J1 ""-I �RADIO-MARKED Nest D(km) from lek .'I-' V1 00 -- 1982 Percent days preeip. Date previous during pr ec Ip, ..incubation Date prevo preeip N radio locations Found dead predation 01 Dec NO 75 45.0 Last signal 16 Nov ND 64 03 Jun 77.8 Found dead, appar. coyote pr ed ,, 12 Oct NO 54 Nest. destr. 23 Jun 23 Jun 46.2 17 !! eggs N nest vtsits 5.5 11 2 Hatched 11 eggs, 14 Jun 14 Jun 40.7 26 May 2.3 11 2 Nest destr. poss. small mammal, 14 Jun 14 Jun 26 May 7.6 9 1 Nest destr. poss. small mammal, 03 Jun 11 Jun 3.4 unk. 0 Date Band (I(radio ~1H3) marked Incubation start whe r e 8R (150.920)a 24 Mar 19 May lOR (151. 261) 26 Mar llR (151.018) b 26 Mar -- APPENDIX I HEN GREATER PRAIRIE-CHICKENS SUMMARY SHEET trapped Nest fate (,date Hen fate (,date 14R (151.309)e 06 Apr 14-15 May 2.8 12 3 Hatched 9, 2 unhatched (near term), 1 unk, 09 Jun 03 Jun 37.0 Killed by vehicle 13 Jun 11 Jun 16R (151.060)d 08 Apr Not appUe. Not appUe. Not appUe. Not appUe. Not appUc. Not applie. Not appUe. Radio removed hen released 08 Apr Not applie. 19R (l51.359)e 10 Apr Assume 1.7 NO 0 14 May 0.0 Known alive La'st signal 06 Oct NO 67 1.3 NO 0 Assumed destr. 2-14 Jul 11 Jul 31.8 Not applie. Not Not applic. Not applie. Not appUc. Not applie. Not applic. Last signal 10 May Not appUe. 10 app l Ic , 9 May 1.5 11 1 Nest·destr. hail, '02 Jun 2 Jun 48.0 Still alive 18 May Assume .Assumed destr. 2122 May 23 Jun 20R (lSI. 060) f 21R (151.162) 10 Apr 14 Apr 55 a Last known alive on 16 Nov. b Off lek with booming male, 03 June; last known alive on 29 Sep. e Found carcass of 1 chick with hen in wheel-track. d Injured wing during trapping. e Off lelc with booming Qale, 04 June, Nesting, incubation assumed from non-movement. f Probable transoitter malfunction. Nest destruction assumed fr~ sudden, rapid movements. ��•.... 0- 0 Appendix J (cont'd) Nest D(km) from lek Incubation start where Band lI(radio HH3) Date marked 41R (151.109) 20 Apr 14-17 Hay 1.2 trapped N !!. nest eggs visits 13 0 Nest fate & date Destroyed, coyote or Date previous precip. Percent days precip. during incubation Date Hen fate & date prevo pr ec Ip , N rad io locations 18-19 Hay 66.6 Last signal 21 Hay 20 Hay 3 badger 18-19 Hay 42R (151.388) 27 Apr 22-26 Hay 1.0 5 2 Destroyed 2-3 Jun I, 2, 3 Jun 61.5 St Ll I aliv.e 29 Jun Not applic. 8 45R (150.859) 28 Apr Not applic. ND Not applic. 0 Hen killed prior to inc. partly-formed egg w/remains Not applic. Not applic. Killed by coyote during laying, 1618 Hay 15, 17, 18 Hay 4 46R (151.187) 28 Apr ND ND ND ND ND 20 Hay ND Killed, 22-23 May, appar. coyote pred. 20 ~lay 6 47R (151.001) 29 Apr Not applic. ND Not 0 applic . Hen killed Not app Ltc ; Not applic. Killed by coyote during laying, 4-11 May 4, 6 May 2 prior Not applic. Not applic. Killed 15-16 14, 16 Jun to inc., partly-formed egg w/remains 102R (150.888) 26 May ND ND ND 0 Not applic. Jun, appa r , coyote 9 �APPENDIX K Species List - Arikaree Study Area Heek of first si~~ Species a 2 Har 3 Am. White Pelican Great Blue Heron Little Blue Heron Snowy Egret White-faced Ibis Snow Goose Canada Goose Green-winged Teal Mallard Ra N. Pintail Blue-winged Teal Cinnamon Teal N. Shoveler Gadwall Am. Wigeon Lesser Scaup Turkey vulture R N. Harrier R Sharp-shinned Hawk N. Goshawk Swainson's Hawk Red-tailed Hawk R Ferruginous Hawk Rough-legged Hawk Am. Kestrel R Merlin Peregrine Falcon Prairie Falcon Ring-necked Pheasant R Greater Prairie-chicken WUd Turkey N. Bobwhite R Virginia Rail Am. Coot Sandhill Crane Killdeer R 4 5-1 2 83 82 81 A r 3 4 5-1 2 Ma 3 Conunents 4 5 83 83 81 83 82 83 83 82 81;82 81 83 82 83 83 83 83 83 83 81;82 81 81 82 81 83 83 81 83 82 81 81;82 83 82 83 83 81 81;82 83 82 83 83 83 81 ;82 82 83 82 R 83 83 83 82 81 81 81 81 8°2 83 82 83 81 83 83 83 82 81 I-' 0\ I-' �.•.... ; 0\ N species list (cont'd) 2 Species 2 Gr. Yellowlegs Lesser Yellowlegs Solitary Sandpiper Willet Spotted Sandpiper Upland Sandpiper Long-billed Curlew Least Sandpiper Stilt Sandpiper Long-billed Dowitcher Common Snipe Wilson's ?halarope Rock Dove R Hourning Dove E. Screech-Owl Great Horned Owl R Burrowing Owl Short-eared Owl Common Nighthawk Common Poorwill Belted Kingfisher Red-headed woodpecker DOwny Woodpecker R Hairy Woodpecker R N. Flicker-Both races EmEidonax sp. E. Phoebe Say's Phoebe I~.Kingbird E. Kingbird Horned Lark R Tree Swallow N. Rough-winged Swallow Barn Swallow Blue Jay R Black-billed Magpie R Am. Crow R Black-capped Chickadee Brown Creeper Rock Wren House I~ren Mar 3 4 5-1 2 A r 3 4 5-1 2· Ha 3 Comments 4 5 81 81;83 83 83 83 81 82 83 6/23/83 81 83 83 83 81 83 83 81 83 83 81 82 83 81 83 83 81;82 82 81 83 83 83 81 83 83 R 81 82 83 83 81 81 81;83 83 83 83· nest 4/10/81; 81 81 83 83 83 83 83 83 82 83 83 83 83 �species list (cont'd) 3 Species 2 Marsh ~lren Ruby-crowned Kinglet E. Bluebird Mountain Bluebird Swainson's Thrush Hermit Thrush Am. Robin R Gray Catbird N. Mockingbird Brown Thrasher N. Shrike Loggerhead Shrike European Starling R Bell's Vireo Orange-crowned Warbler Yellow Warbler Magnolia Warbler Yellow-rumped I~arbler Blackpoll Warbler Black and lfuite Warbler Am. Redstart MacGillivray's Warbler C. Yellowthroat Wilson' s I~arbler Mourning Warbler Yellow-breasted Chat Western Tanager Indigo Bunting Rufous-sided Towhee Cassin's Sparrow Am. Tree Sparrow Chipping Sparrow Clay-colored Sparrow Brewer's Sparrow Field Sparrow Vesper Sparrow Lark Sparrow Lark Bunting Grasshopper Sparrow Song Sparrow R Lincoln's Sparrow Swamp Sparrow Har 3 Ma A r 4 5-1 2 3 4 5-1 2 3 Comments 4 5 83 83 6/15/83 82 83 83 83 81;82 83 83 83 83 83 83 81 81 82;83 6/15/83 83 83 83 83 83 83 83 83 81 83 83 83 83 83 83 83 83 83 81 83 83 83 83 81 82;83 81 83 81 81 83 83 83 83 81 I- a v �species list (cont'd) 4 I-' Q'\ ..,.. Species 2 ~fuite-crowned Sparrow Dark-eyed Junco Lapland Longspur Chestnut-collared Longspur Red-Hinged Blackbird R H. Headowlark R Yello>1-headed Blackbird C. Grackle Brown+headed Cowbird Orchard Or"iole N. Oriole House Finch Am. Goldfinch R House Sparro>1 R -_ a Mar 3 4 5-1 A r 3 82 83 83 83 83 83 2 4 5-1 2 Ma 3 Comments 4 5 81· 81 83 81 82 nest 4/16/81; 83 81 81;82;83 81;83 83 83 83 81 83 83 81 R - Resident species ',. �165 APPENDIX L Woodworth, N.D. Sept. 28, 198) Dr. Clait E. Braun Wildlife Research Leader J17 West Prospect Road Fort Collins. Colorado Dear Clai t , It seems an age since bouncing about the Colorado prairies with you and Gary Miller. Attended the Prairie Grouse Technical Council Meeting at Emporia, Kansas and saw many of the pra~r~e grouse biologists currently active. It was rather sobering to find that Don Christensor. of the Missouri Conservadon Commission and myself were the only ones who had attended the last P.G.T.C. meeting held in Emporia 24 years ago. Was pleased to note that there are a number of success stories now. In 1959 there was one from the Hammerstroms in Wisconsin. I hope to see the day that Colorado has a success story to tell at the P.G.T.C. meeting and I am sure it will if management is applied to the grasslands. Now to the more difficult part of this letter. I have examined the Program Narrative Outline for project number W-145-R, ·work plan. 1. Job 1, titled Prairie Grouse Investigations. In my opinion this study costing approximatlHy $186,578 will result in little if any information useful to the Colorado Division of Wildlife in its effort to save the greater prairie chicken. In fact, I question if most of the major objectives can be attained. These include size and condition of habitat islands and the number of grouse by sex and age class needed to.establish and maintain populations. The use of radiotelemetry is an important technique for even partial success of this project. I fear that prairie grouse carrying radios harnessed on their backs or breasts are radioed grouse and may not function as other grouse do. This fear was Lnc r-ea.aed by the fact that 71% of the prairie ,chicken females studied in Colorado during 1983 were dead by September 1. A paper on the Attwater's prairie chicken presented at the P.G.T.C. meeting at Emporia reported that 67 percent of 1.5 females carryinv. radjo~ were dead by September. Previous to this T hno read W. Daniel Svedarsky's study of radioed prairie chickens in Minnesota. Dan reported that there was excess mortality among radioed hens. Examination of published literature on radioed sharp-tailed grouse reveals that they die at a rate of about 1 percent per day. Neither prairie chickens or sharp-tailed grouse would survive if true mortality rates approached this. In view of this it seems that putting radio packs on Colorado prairie chickens is not a wise thing to do. ',. ' 5"8<1'% �166 APPENDIX L cont'd. To prove that I'm not compl~tely negative, I do agree with Walter D. Graul's 'Analysis of State School Lands in Yuma County As Greater Prairie Chicken Habitat." Anything done to reduce grazing is a step in the right direction for prairie chicken management. Now I must renew the attack. Frankly speaki.ng, most of the current studies of prairie chickens in Yuma county have little potential for finding the needed answers. The most important part of the study is annual spring surveys of booming ground males. To learn something about habitat requirements you must have longterm land use and habitat manipulation control. Safe and acceptable nesting habitat is the bottle-neck which restricts both prairie chickens and sharp-tailed grouse throughout their range including Colorado. To prove this and ultimately save the chicken I suggest a new research project. You must let the prairie chicken tell you what kind of habitat he thrives in and to do this you must give him Choices. This can be accomplished on the school lands of Yuma county where prairie chickens still exist if complete land use control is obtained. I suggest that at least 6 sections (square 640 acre blocks) separated by at least 4 linear miles be purchased or leased for ten years. Manage two of these by grazing at or near the present rate, two by not grazing and two by prescribed burning in early or mid May of the year before the study begins. The burn must be of sufficient intensity to remove all or 90 percent of the sandsage. Prairie chicken response could be measured by simply counting males on all booming grounds within 1 or maybe 1.5 miles of each block each spring. If more intensive response studies are desired, search the study block in early May and late May with a cable-chain nest drag to measure nesting response. A similar search during the first two weeks of July would measure brood use. Nest counts will be minimal as prairie chickens when incubating are apt to ignore the cable-chain, however I believe they are suitable for comparing nest densities on the different treatments. I have used this type of response study on sharp-tailed grouse and the booming ground count .. method on prairie chickens North Dakota. (See attached paper)., in Height-density measurements should be made each year during the spring before new growth influences the physiognomy of the vegetati.on. Vegetative transects to measure species occurren\::c and canopy coverage should also be made. Gads! its 12:15 A.M. and have said enough for now. this is useful. Sincerely, f~ -/ . .•,...•....:.r Leo Kirsch Biologist Hope �167 APPENDIX L cont'd. STATE OF COLORADO RIchard D. Lam"" Governor DEPARTMENT OF NATURAL RESOURCES DIVISION OF WILDLIFE Jack R. Grieb, Dlroctor 6OSOBroadway . Denver, Colorado 80216 (297·1192) Leo Kirsch Woodworth, N.D. Wildlife Research Center 317 West Prospect Road Fort Collins, CO 80526 03 October 1983 58496 Dear Leo: Appreciate talking to you on 30 September and receiving your letter of 28 September 1983. I have reviewed your letter several times and have considered its contents at length. Basically I concur that our present prairie chicken research project needs redirection. While I am a firm believer in the use of radiotelemetry as a tool, my experience indicates that any object around the wings or a body of a grouse markedly reduces the survival of that particular bird. Radiotelemetry can help us learn many things but surv lva l rates i's,not 1 of them. The proposed research that you suggest is exciting and long term in nature. The basic problem is acquiring control over the land for a long enough period to measure results of the treatments. Most'indlviduals in the CDOW are negative about this approach as they do not believe we can get some control over state school lands for this purpose, I, as always, am optimistic and will be formulating an action plan to start working 'towards this goal. The first thing I will need is public support. It would also be nice to have Internal support. In any event, I plan to work hard to make the Tamarack renovation a success and in redirecting our present prairie chicken research effort. Progress can be made if the CDOW is serious. We will now see how serious they are. Appreciate your counsel and willingness winter. I will keep you posted. to help. Have a good fall and Sincerely, ~t.~ (p) Clait E. Braun Wildlife Research Leader Small Game and Nongame CEB/jeb enclosure xc: W. Graul R. Hopper ;;;-~~M l).rer.~ W. Snyder DEPARTMENT OF NATURAL RESOURCES, David H. Getches, Executive Director'WILDLIFE COMMISSION, Richard L Divelbiss, Chairman James C. Kennedy, Vice Chairman'Wilbur L Redden, Secretary· Donald A. Fernandez, Member'Michael K. Higbee, Member Timothy W. Schultz, Member'James T. Smith, Member'Jean K. Tool, Member �168 APPENDIX M PROSPECTUS Community Management Study: Sandsage-Bluestem Prairie Background Colorado's sandsage-bluestem (Artemisia-Andropogon) prairie once comprised 3.5% of the state's natural vegetation. Historical information is incomplete, but it appears this prairie originally was dominated by warm-season grasses such as bluestem, switchgrass (Panicum virgatum), prairie sand reed (Calamovilfa longifolia), and indiangras~ghastrum nutans). Sandsage (A. filifolia) may have been co-dominant or sub-dominant, depending on site characteristics (see Ramaley 1939). Much of the original extent of this prairie has disappeared. By 1967 48.5% of the state's sandsage-bluestem prairie had been converted to other uses (data from J. M. Klopatek--see Klopatek e,ta1. 1979). Work in 1979-80 in Colorado has shoW'rlthat, in some areas, over 45% of the remaining prairie is in a disclimax stage, dominated by sandsage and/or grasses such as blue grama (Bouteloua gracilis) and muhly (Muhlenbergia spp.) .(Miller 1981, Miller and Schrupp198T)'":The sandsage+bLuestem community contains a number of vertebrates that are essentially restricted to this vegetation type for reproduction. Greater and lesser prairie-chickens (Tympanuchus cupido pinnatus and T. pollidicinctus, respectively) are classified in Colorado as endangered and threatened species, respectively. Upland sandpiper (Bartramia longicauda) densities appear greatest in the sandsage-bluestem and long-billed curlews (Numenius amerLcanus) have been known to nes t in this type. Preliminary analysis of state distributional information indicates that at least 110 species of terrestrial vertebrates (18.1% of the state's total) in Colorado use this vegetation type for some part of the annual cycle. Need The Colorado Division of Wildlife (CDOW) is statutorily obligated to prevent the extinction of any native species in Colorado. In order to meet this obligation, it will be necessary to restore and maintain some sandsage-bluestem prairie in a natural state. In addition to benefitting such threatened and endangered species as the pra~rie-chickens and such stenotopic species as upland sandpipers, this restoration and maintenance will provide for other prairie-adapted species. The CDOW has as a goal the development and implementation of ecosystem management principles, as stated in the Strategic Plan (Colorado Division of Wil!1life 1983). Although of primary interest to the Nongame Program because of the threatened and endangered species, maintenance of sandsagebluestem has been identified, in the Strategic Plan, as important for Game Program ~bjectives also. The CDOW Haster Plan for Habitat Acquisition lists this type as the 5th highest priority in the state. �.169 APPENDIX M cont'd. The loss of a natural sandsage-bluestem prairie took place over many years. Techniques to restore this system from its present disclimax are unknown, and will take time to develop and implement. This prairie tYFe probably evolved with fire, but fire has been suppressed by man for many years through most of the area. Periodic grazing by large herbivores was also a part of the sandsage-bluestem prairie system, but it is doubtful if the grazing regimes employed at present even remotely resemble the natural situation. In order to restore and maintain a natural sandsage-bluestem community, in keeping with ecosystem management pr i.ncLp.Ie s (Graul and Niller, in ~.) it will be necessary to test and modify various management techniques and measure the impacts upon vertebrates. In order to meet CDOW goals and objectives, a management strategy for the sandsage bluestem community must be developed that allows the optimization of management for certain wildlife species within the constraint of not lOSing any species of that community. Objective ·Ascertain by 1990 whether prescribed burning is an appropriate tool for restoration and maintenance of sandsage-bluestem prairie. If so, develop and implement, in conjunction with operations personnel, appropriate methodologies for enhancing prairie grouse and other species to desired levels while maintaining acceptable levels of all native species of this prairie. Expected Benefits Wildlife managers \Jill have a tool to economically manage a large number of wildlife species and meet CDOW objectives. Quantifying costs and benefits of sandsage-bluestem prairie management ~ill be possible. Threatened and endangered species management programs for prairie grouse may benefit. Secondary benefits may accrue to the private sector, with the development of a possible range management technique to benefit ranchers. Avproach 1. Review literature dealing with prairie ecology limited to, effects of fire upon vertebrates. 2. Review prairie. data collected previously in Yuma County including, but no and on the Tamarack 3. Develop hypotheses to be tested. These hypotheses will probably relate to: a) comparision of vertebrate species richness on a grazed, ungrazed only, ungrazed and burned, ungrazed, burned Rnd seeded treatment plots b) m~asuring use or selection of various treatment plots by selected species including, loot not nece ssar i Ly limited to, prairie grouse and upland sandpipers. �170 APPENDIX M cont'd. 4. Test hypotheses a) select sampling units; treatment and controls NLT i ~~~1984. b) develop sampling design NLT ~.<.h._!984. c) take pre-treat~ent measurements NLT 30 April 1984 d) obtain treatment data, including fire characteristics agd costs. e) take post-treatment measurements 2 times/year for 5 years (1984-1988) f) compile, ana Lv ze , and report da t a armua lly ; ass is t in development of modifications as requested. 5. Publish resul ts NLT 30 June 1990. Personnel Assignments Gary C. Miller Vacant Principal Investigator Technician I-A or graduate student Location Field work will be accomplished on the Tamarack prairie portion of CDOW's South Platte W!-L". and on adjacent lands administered by the State Board of Land Commissioners (State School Lands). Related Projects This project accomp an i cs a project measuring vegetation (habitat) responses with the same treatments. A nongame research project (N-I-R-4-1 Prairie Grouse Investigations) will be conducted in the same location during this period). Literature Cited Colorado Division of Wildlife 1983. Today's strategy ...tomarrow's Wildlife. 3rd ed. Colorado Division of Wildlife, Denver. 96pp. Klopatek, J. ~., R. J. Olson, C. M. Emerson, and J. L. Joness. 1979. Land-use conflicts with natural vegetation in the United States. Environ. Cous erv . 6(3): 191-199. Miller, G. C. 198!. Development of a preservation program for three species of prairie grouse. Job Prog. Rep., Colo. Div. Wildl., Wildl. Res. Rep. Jan. pp. 38-52 . ., and D. L. Schrupp. 1981. Characteristics of greater prairie chicken range in Colorado. Proc. Prairie Grouse Technical Counc' Council Conf. 14. (abstract only). Ramaley, F. Monogr. 1939. Sand-hill vegetation 9(1). 51pp. of northeastern Colorado. Ecol. �171 Colorado Division of Wildlife Wildlife Research Report January 1984 JOB FINAL REPORT State of Colorado Project N-I-R-2 Work Plan 5 Job 1 Period Covered: Author: Mountain Plover Transplant-Experimental 1 January 1982 - 31 December 1983 G. C. Miller Personnel: B. Cordova, L. Fisher, W. D. Graul, G. C. Miller, J. Sedgwick, Colorado Division of Wildlife; M. Hames, Central Michigan University; B. McCaffery, Cornell University. ABSTRACT In 1982-83, 68 mountain plover (Charadrius morttanus) chicks were trapped in Colorado, transported to Kansas and released in an attempt to re-establish a population. No chicks died during capture or captive maintenance in Colorado. It is recommended that Colorado's efforts be terminated until the outcome of the transplants becomes clear. ��173 MOUNTAIN PLOVER TRANSPLANT-EXPERIMENTAL Gary C. Miller Historically, mountain plovers nested throughout western Kansas. Early records exist (Allen 1872, Goss 1891, Menke 1894), but records since that time have been scarce (Long 1940, Goodrich 1946, Rising and Kilgore 1964). Graul (1973) did not find evidence of recent nesting in western Kansas. Graul and Webster (1976) believed that the Pawnee National Grassland in Weld County, Colorado? represented the stronghold of the species' breeding range. Restoration of mountain plovers to Kansas was identified as a desirable objective for that state's nongame program (M. Schwilling, pers. commun.). A letter from B. Hanzlick, Director, Kansas Fish and Game Commission, to J. R. Grieb, Director, Colorado Division of Wildlife, dated 21 April 1982', requested 40 mountain plover/year over a 3-year period in a trade for greater prairie-chickens (Tympanuchus cupido) (Miller 1983). Thus, a possible restoration technique for mountain plover could be tested at little expense to Colorado. P. N. OBJECTIVES This project, funded through nongame checkoff funds, was initiated prior to implementation of Administrative Directive I-I, Research Planning and Project Implementation (3 Aug 1982). The documents that exist stipulate the general objective of providing mountain plovers to Kansas in return for greater prairie-chickens (Miller 1983). SEGMENT OBJECTIVES lao Review literature on mountain plover ecology and transplants of precocial avian species, especially charadriiforms. lb. Review literature on capture and captive maintenance of charadri::f.forms 2a. ca~ture young mountain plover from Pawnee National Grasslands at 18 days of age, prior to fledging. Band plovers as captured. 2b. Hold young plovers in captivity until groups of at least 8 are attained. Maintain records on developm~nt and care. 2c. Co-ordinate with M. Schwilling (Kansas Fish and Game) to receive plovers at Kansas-Colorado border. 2d. Transport plovers to border, transfer to Kansas Fish and Gam.e personnel (E. and J. Schulenberg). 0 �174 3a. Kansas Fish and Game personnel will release plovers in suitable brood habitat in groups of 3-5 immediately upon arrival at release site. 3b. Exchange information with Kansas personnel upon completion of transplant. 4. Monitor Pawnee grasslands in 1983 and 1984 for presence of banded, transplanted birds. 5. Publish results of maintenance of plovers and results of transplant. DESCRIPTION OF STUDY AREA Graul (1975) described the area from which mountain plovers were taken (Pawnee National Grassland, Weld County, Colo.). The potential release areas in Kansas were described by L. Fisher in a report submitted to the Kansas Fish and Game Commission on 8 June 1982. The site where the plovers were released was in Wallace County, 32 km northeast of Sharon Springs, Kansas. Details of the release were provided in a report by J. Ptacek, Kansas Fish and Game Commission, dated 4 August 1982 (Miller 1983). METHODS Reproductive activity was monitored on the Pawnee National Grasslands between April and June 1983 by the principal investigator, and nonDivision personnel M. Hames and B. McCaffery. Arrival, courtship, and nesting was monitored by Hames and McCaffery, and all surveyed for the presence of young and the presence of white leg bands on plovers (from 1982 transplants) during this period. Mountain plover chicks were captured by hand or with long-handled ;..""t:3" Only chicks that were within approximately 2 weeks of fledging wert taken from the area. Suitable chicks were those with feathering of the humeral tracts and at least some feathering of the capital tract, and with the longest primary feather partially exposed. Captured chicks were banded and held in captivity at the CDOW's Wildlife Research Station northeast of Fort Collins. Prior to placing the chicks in the holding facility, it was cleaned and disinfected, and a clean sand over cement substrate was provided. Captive plovers were maintained on a diet of 28% protein turkey starter and mealworms (Tenebrio molitor). RESULTS Late spring storms with snowfall and many hailstorms occurred during plover nesting causing nest losses and low nest success and/or asynchrony �175 of hatching. No plovers transplanted to Kansas in 1982 were encountered on the Pawnee National Grassland or at the release site in Kansas in 1983 (M. Schwilling, pers. commun.). Eighteen mountain plover chicks were captured, maintained, and delivered to Kansas personnel between 9 and 29 July 1983. Plover hatching occurred 2-3 weeks later in 1983 than in 1982. No report of 1983 activities was received from Kansas Fish and Game personnel. DISCUSSION AND RECOMMENDATIONS Through 1983, 68 mountain plover chicks were transplanted to Kansas. Until the outcome of the transplants becomes clear, I recommend ceasing capture and transplant of plovers. This strategy is acceptable to Kansas personnel (M. Schwilling, pers. commun.). Monitoring for presence of white banded plovers on Pawnee National Grassland in 1984 will occur at no cost to CDOW; B. McCaffery and M. Hames will monitor in the course of their studies in the area from which plovers were taken. Kansas personnel will monitor the release site again in 1984. LITERATURE CITED Allen, J. A. 1872. 6:263-275. Ornithological notes from the West. Goss, N. S. 1891. History of the birds of Kansas. Topeka, Kans. 692pp. Goodrich, A. L. 1946. Topeka. 340pp. Graul. W. D. system. 1975. 87:6-31. Birds in Kansas. Am. Nat. G. W. Crane and Co., Kans. State Bd. Agric., 1973. Adaptive aspects of the mountain plover social Living Bird 12:69-94. Breeding biology of the mountain plover. ----- , and L. Webster. 1976. Wilson Bull. Breeding status of the mountain plover. Condor 78:265-267. Long, W. S. 1940. Check-list of the Kansas birds. Sci. 43:433-441. Menke, H. W. 1894. Q. 3:129-135. Trans. Kansas Acad. List of birds of Finney County, Kansas. Kans. Univ. �176 Miller, G. C. 1983. Mountain plover transplant - experimental. Job Prog. Rep'9 Colo. Div. Wildl., Wildl. Res. Rep. Jan. Pp. 63-86. Rising, J. D., and D. L. Kilgore, Jr. 1964. Notes on birds from southwestern Kansas. Bull. Kansas Ornith. Soc. 15:23-25. Prepared by Gax~ c_~m~ Gary C. :hiller Wildlife Researcher C � https://cpw.cvlcollections.org/files/original/b4fb851097e90d3328f69c282822545f.pdf 05f948ab48f526a4176a42ed17122f77 PDF Text Text Colorado Division Wildlife Research January 1984 177 of "lHdlife Report JOB PROGRESS State of REPORT Colorado Project Fishes of Colorado: An analysis Distributional Change N-2-R-2 Work Plan of 1 --------------------- Job 1 Period Covered: Author: 1 January - 31 December 1983 C. M. Haynes and Sara E. Whetstone Personnel: J. Bennett, C. Haynes, Division of Wildlife. S. Whetstone, J. Woodling, Colorado ABSTRACT An analysis of eastern slope fish distributional information in 1983. An annotated bibliography was prepared. was conducted This Job Progress Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the author. ��179 FISHES OF COLORADO: Charles M. AN ANALYSIS OF DISTRIBUTIONAL CHANGE Haynes and Sara E. Whetstone P. N. OBJECTIVES The long-range objective of this project is a comprehensive publication detailing the historical distribution and current status of the fishes of Colorado. Intermediate goals are 2-fold: (1) the preparation of a bibliography of information sources concerned with fishes in each of the major river drainages of Colorado, and (2) a data bank containing a summary of the information in the bibliographic sources. To achieve these goals a data collection that would answer 2 objective questions: and storage system was designed 1. Where in Colorado has this fish species been recorded? 2. What fish species have been found in this water (may be 1 of 7 designated drainages, a subdrainage, or a specific location in Colorado). In the 1981-83 seasons these goals were applied to the San Juan, Rio Grande, Colorado, Platte, and Arkansas river drainages in Colorado. Approaching the collection of data on a drainage-by-drainage basis permitted a more systematic search and a finished product, i.e., a bibliography of these regions. A method of data collection was required that would effectively summarize a source of information in terms of the objectives. Relevant data were specifically defined so that a document was efficiently and consistently evaluated. Two methods of information storage were desired. A manual filing system at the Fort Collins Nongame Research facility would permit quick access to the data and would help answer relatively simple questions. Storage of the data in a computer system would permit more sophisticated queries and data evaluation. We tried to keep both storage and retrieval systems simple so that researchers interested in the information would not need an elaborate set of instructions to retrieve the information. SEGMENT OBJECTIVES 1. Document the historical and present distribution of both native and non-native fishes in the Colorado River drainage, with particular emphasis upon nonharvested and threatened and/or endangered species. 2. Provide a data-base which may serve as a predictive tool for recognizing future potential negative impacts upon segments of the �180 fish fauna as a consequence of energy development, stream and river modification (e.g., dams, withdrawals), agricultural development (e.g., irrigation returns, livestock damage, etc), acid-precipitation, etc. METHODS A thorough description of information retrieval, compilation, and different filing systems was given by Haynes and Galat (1983). For 1 January-31 December 1983, emphasis was placed on locating information relative to eastern slope drainages (i.e., Platte and Arkansas basins) and ponds and reservoirs of eastern Colorado. Small private ponds and reservoirs which could be identified on U.S. Geological Survey topographic maps but which have historically received little or no agency attention presented problems as well as vague historical references to small ponds which have undergone one to several common name changes over time. Reconciliation of some of these situations were possi11e following discussions with R. Nittmann (Northeast Region) and W. Wiltzius (Fish Research) and inspections of Northeast Region stream and lake survey files. A number of obscure private ponds and reservoirs, as well as general descriptions of their fisheries, were identified using general references such as Kelley (1983). However, an unknown number of private standing bodies of water will clearly remain unidentified as will their fish faunas. RESULTS Fifty-seven information sources relating to eastern slope fishes were identified and compiled during 1983. This included a number of previously accessed sources for western slope drainages. An annotated bibliography of east slope citations, as well as distribution maps, was completed and will be presented in the final report. LITERATURE CITED 1983. Fishes of Colorado: an analysis Haynes, C. M., and K. H. Galat. of distributional change. Job Prog. Rep., Colo. Div. Wildl. Nongame Investigations, pp. 87-106. Kelley, T. Wyoming Prepared by fishing guide. Official Colorado 1., Inc. Denver, Colo. 386pp. and �18 \ Colorado Division of Wildlife Wildlife Research Report January 1984 'Pc.<, 2JOB PROGRESS REPORT .:·~tate of Colorado ~ - 1roject Work Plan Job N-2-R-2 (SE-3) 2 ~~~~~~~- Identification of Habitat Requirements and Limiting Factors for Colorado Squawfish and Humpback Chubs .. 1 Period Covered: Authors: Personnel: 1 July 1983-30 June 1984 C. M. Haynes and R. T. Muth T. Beck, C. Haynes, S. Platania, T. Pojar, G. Skiba, and S. Steiner t, Colorado Division of Wildlife; D. Bowden, C. Carlson, G. Dean, R. Muth, and J. Simpson, Colorado State University. ABSTRACT A total of 703 seine and ichthyoplankton samples was collected in the Colorado and Yampa River study areas combined i n 1983. Three YOY and 2 juvenile Colorado squawfish (P t ychocheilus lucius) were collected in the Colorado River study area while 331 YOY were collected in the Yampa River (228 by seine and 103 by drif t-net). Estimated spawning dates for 1983 were 27 July-10 Aug and / / July-11 Aug for the Colorado and Yampa Rive r study areas, respectively. Based upon comparable captureper-unit-effort (C/E) values, 1983 yielded the highest number of YOY squawfish in the Yampa River study area (6.02/100 m2) whil~ capture in the Colorado River (0.15/100 m2) was below that of 1982 but greater than 1981. This Job Progress Report represents a prel iminar y analysis and is subject to change. For this reason, information pr esented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the Researcher. 1~ii~~i~Uil~11~1l~i1ii11 BOOW027831 ��] 33 IDENTIFICATION OF HABITAT REQUIREMENTS AND LIHITING FOR COLORADO SQUAWFISH AND HUMPBACK CHUBS FACTORS Charles H. Haynes and Robert T. Muth As a consequence of documented declines in numbers and ranges, the Colorado squawfish {Ptychocheilus lucius} and humpback chub (Gila cypha) have been listed as endangered by both the federal government and the State of Colorado. Suspected causes of decline have been discussed by Haynes and Muth (1982), Holden and Wick (1982), Valdez and Clemmer (1982), Haynes et ale (1984), and others. Since 1977, the Colorado Division of Wildlife has been investigating the distribution, ecology, and status of these rare fishes via systematic sampling in selected reaches of the Colorado, Gunnison, White, and Yampa rivers. Aspects of study have been conducted cooperatively with the U.S. Fish and Wildlife Service (Colorado River Fisheries Project). Nongame Research (Colorado Division of Wildlife, Fort Collins) investigations have been directed toward evaluating reproductive success for both spec-ies via collection of young-of-the-year (YOY) and juven~les, evaluating distribution and timing of spawning in selected reaches of the Colorado and Yampa rivers, and identifying habitat electivity for early life-history stages. P. N. OBJECTIVES The overall objective of this investigation is to identify the physical and biotic factors which limit the distribution and reproduction of Colorado squawfish and humpback chubs in Colorado. Similarly, this project is designed to develop field and laboratory methods for evaluating squawfish and humpback chub habitat, and reproductive success which may be used by management personnel for future habitat enhancement and/or reintroduction programs in accordance with overall recovery efforts. Additionally, information derived from this and other state and federal studies may be used to evaluate the impacts of several upper Colorado River water development projects and enhance the probability of meaningful mitigation where such development projects are believed to have deleterious effects upon rare fish populations. Aspects of this investigation relating to the humpback chub will be incorporated into a doctoral dissertation in 1984-85 (R. T. Muth, Dep. Fish. and Wildl. BioI., Colo. State Univ.). SEGHENT OBJECTIVES 1. Identify, measure, and analyze habitat distribution and abundance of Colorado chubs in the upper Colorado River. parameters which limit the squawfish and humpback 2. Identify macro- and microhabitat features (e.g., flow, temperature, substrate, depth) which are associated with presence - absence of �18 ; (YOY) squawfish, humpback chubs, and other associated species. 3. Determine probable time of squawfish and humpback chub spawning and correlate with radiotelemetry data developed by the Northwest Region, Colorado Division of Wildlife and the U.S. Fish and Wildlife Service. 4. Develop methodes) for quantitatively evaluating squawfish reproductive success in terms of space and time within and between study areas. 5. Investigate 6. Devise methods for the identification and juvenile specimens of Gila spp. drift as a dispersal mechanism for YOY squawfish. and differentiation of YOY STUDY AREA AND METHODS The Colorado and Yampa River study areas have been previously described (Wick et aI , 1981; Haynes and Huth, 1982; Wick et al., 1983; Haynes et al., 1984). Seine sampling was conducted in both areas between June and September 1983. Habitat descriptors and sampling methods were defined by Haynes and Muth (1982). In the Yampa River, drift-net sampling for YOY fishes was conducted between 11 June and 21 August 1983. Net design, deployment methods, and rational were described by Haynes et ale (1983). Three nets each were deployed at two sites in Dinosaur National Monument. Station 1 was located at the lower end of Stratum 2 (km 32.5) a short distance above the Harding Hole area. Station 2 was located at the lower end of Stratum 1 (km 3.1) near the Yampa-Green River confluence. Since 1980, squawfish spawning in Dinosaur National Monument has been presumed to be limited to a reach between km 28.8 and 0.2 (Stratum 1). This was based upon the collection of YOY within this reach exclusively (Haynes and Muth, 1982; Haynes et al., 1983, 1984) and observations made using radiotelemetry (Tyus et al., 1982; Wick et al., 1983). The Harding Hole (Station 1) site was selected to evaluate drift input from upper Yampa Canyon (Stratum 2) while the lower site was selected to evaluate both similar trends in Stratum 1 and possible drift into the Green River. Nets were deployed along the shoreline just beneath the surface. The quantity of water filtered by each net was measured at each sampling time with either a Marsh-McBirney (Model 201) or Pygmy Gurley (Model 625 F) current meter. Water temperature (C) was measured during each collection period. In order to evaluate potential diel drift patterns, samples were made at sunrise, noon, one-half hour after sunset, and at midnight. Sampling duration at each period ranged from 1 to 2 hours depending upon the suspended solids load. Sampling design permitted an evaluation of fish drift periodicity both within a particular growth phase (e.g. protolarvae) and between growth phases (e.g. protolarvae vs mesolarvae). Two-way analysis of variance (2-AOV) and Hotelling's �185 t2 tests were used to compare drift densities (response variable), collectively and for predominant taxa, among the four sampling times and between day and night samples (i.e., combined sunrise and noon = day, combined sunset and midnight = night) on individual sampling dates over the sampling period. To estimate probable predictive equations (Fig. 1). spawning dates of collected were derived using Hamman's For TL < 22.0 mm: Age (days) = -76.7105 + 17.4949 L - 1.0555 L2 (r2 = 0.99) YOY squawfish, two (1981) hatchery data + 0.0221 L3 For TL = 22.0 - 47.0 mm: Age (days) -26.6421 + 2.7798 L (r2 = 0.99) Where L = TL (nearest O. 1 mm) These predictive equations differ from previous equations used by Haynes et al. (1983, 1984) due to reinterpretation of Hamman's (1981) data and, in general, result in a narrowing of previous ranges of estimated spawning dates. The usefulness of these equations is perhaps best evaluated when their resulting data estimates are compared with actual on-site spawning observations. For example, Wick et al. (1983) concluded, based upon observations of ripe radio tagged adults over a previously documented Yampa River spawning site, that spawning in 1982 appeared to peak in intensity during 23 July-8 August. Our estimate for the same year was 24 July-8 August. Practical methods for direct YOY abundance estimates in the Upper Colorado River ecosystem do not presently exist; however, it is critical that methods be developed which permit the evaluation of abundance trends within both space and time. An index of relative YOY abundance (i.e., capture-per-unit-of-effort, C/E) was developed using basic tenets of Ricker (1975), Caughley (1978), Lackey and Hubert (1978), Southwood (1978), and Tanner (1978), i.e., that C/E is related in a constant or predictable way to population size and that changes in absolute abundance over space and time will be reflected by changes in c/E. A disadvantage of C/E methods in the difficulty of rigorous statistical analysis due to the lack of sample replication. Further, conditions that are not consistently possible to achieve are: (1) conditions of capture are consistent (e.g. time, weather), (2) capture efficiency is consistent (e.g. personnel, gear, coverage of habitat features), (3) the capture of one animal is unrelated to the capture of another, and (4) the animals do not learn either avoidance or attraction. Such conditions may be met in closed systems; however, it is rarely possible to meet these assumptions in an open system with the complexity of the Upper Colorado River. The above authors suggest, however. that �136 DEIiRU:.3 IS Z Z.:sa "" !( :z: ·Z•• : ~ ~ ~ (·Z.O ~YS) 1.:58 1•• ~. So -76.7105 of" 17.49<t'L -1.0555 L"+ O.oW I}' (ftW' TL6 22",,...> 1JJ.O IS.O 10.0 TL(MM) 1.88 8.:58 £+2 1.:58 2.88 E+l D£GREE .1 1.18 1.88 8.98 " : Z 8.88 ~ -e ::z: 8.78 (••70 DAYS) ~ 8.68 ~ 8.:58 AGE· 8.<48 -U,.64ZJ + Z.77"13l. (for TL > llrnm) 8.38 3.8' 4.8' £+1 TL(MM) Fig. 1. Relationship of TL (nearest 0.1 mrn) to age for YOY Colorado squawfish. For TL 2 22.0 mm, age is best predicted by a 3rd degree polynomial (upper). For TL ~ 22.0-47.0 mm, age is best predicted by a linear regression (lower). �187 meaningful year-to-year comparisons, as part of a long-term monitoring program, are possible with such indices when time, human, and economic resources are limiting. The following methods are developmental and subject to further evaluation and testing. C/E values (number of YOY squawfish/l00 m2) were derived from seine samples of known area from known specific habitat features using standard 1.6-mm square-mesh nets. Principal personnel have remained consistent since 1981 when these methods were intially employed. Because of a variety of sampling methods in 1979-80, C/E was calculated for 1981-83 exclusively. For these years, YOY squawfish were captured exclusively in recognizable habitat types characterized by Haynes and Muth (1982) as backwaters, embayments, shoreline, concavities, pools and isolated pools. C/E values were thus determined using seine samples from these features only. Further, for any given year, only those samples collected on or after the earliest estimated spawning dates were considered. Flow and temperature data wpre derived both from U.S. Geological Survey Water-Data Reports (1982, 198.1) and unpublished USGS data (R. Ugland, pers. comm.) , in addition to temperature measurements made at time of sample collection. The thermal regime of each study area is characterized in terms of 22 C degree-days (22 C-days) following concepts described by Hayes (1949), Andrewartha and Birch (1954), Arnold (1960), and Baskerville and Emin (1969). 22 C-days, for the purposes of this report, are the summation of maximum water temperatures beginning with the date of the first recorded 22 C maximum and continuing until the date of the final 22 C maximum. A water temperature maximum of 22 C was selected for degree-day summation for two reasons. First, several investigators have provided strong evidence which suggests that spawning is stimulated by water temperatures of 20-22 C (Vanicek and Kramer, 1969; Toney, 1974; Holden, 1980; Haynes et aI, 1984). Second, incomplete USGS records for Deerlodge Park - the closest reference station to documented squawfish spawning areas - precluded the selection of temperatures <22 C. The reference site for the Colorado River study area was USGS station 09163500 near the Colorado-Utah state line, while the refence site for the Yampa study area-was USGS station 09260050 at Deerlodge Park, Colorado. Data for the Colorado River study area in 1981 were incomplete for early-September, leading to a probable underestimate in that instance. Similarly, data were incomplete for Deerlodge Park in 1983; however, modal values for downstream seine temperature measurements were inserted when USGS data were incomplete after it was found that these field values consistently matched USGS data. On 13 July, two ripe female and four ripe male roundtail chubs (~. robust a) were collected by gill net at km 174.0 near the Roundbottom area (Moffat Co.). The ojective of this effort was to provide a known-age larval series of this species for morphomeristic comparison with other Gila spp. and cultured hybrids. Since G. robusta was apparently allopatric in this reach of the Yampa River, we assumed that �188 roundtails from this area would have a high degree of "genetic purity". Eggs were fertilized and water-hardened on-site afterwhich they were transported to a flow-through incubator (Heath Tecna Corp., Kent, Wash.) at Colorado State University. Culture conditions, sampling procedures, and developmental events will be presented in Muth, et al. (in prep.). RESULTS AND DISCUSSION A total of 206 individual seine collections was made in the Colorado River study area in 1983 between 6 June and 10 September. In the Yampa River, 497 samples were collected between 11 June and 23 August, of which 245 were seine samples and 252 were by drift net. Preliminary laboratory processing has been completed with the exception of final measurements of species other than squawfish and Gila. Data will be stored on the MANAGE computer database at Colorado State University, Fort Collins as analyses are completed. Morphomeristic analysis of field-collected Gila spp. is underway as well as statistical comparisons of field and hatchery specimens. The results of this aspect of the overall study will be presented in a doctoral dissertation by R. T. Muth (Dept. Fishery and Wildlife Bio., Colo. State Univ.) in 1984-85. Colorado Squawfish 1983 Seine Collections Three YOY squawfish were collected on two dates in the Colorado River study area (Table 1). All specimens were collected in low-velocity mainchannel backwaters between km 236.4 and 237.3. Age at time of capture is estimated to be 11-32 days post-spawning with a resultant 1983 spawning period of 27 July-l0 August. Two Age I squawfish were collected in the Colorado River study area in 1983. One juvenile (TL = 65.0 mm) was captured at km 218.1 on 2 August in a tributary stream embayment. A second (TL = 76.2 mm) was captured at km 244.6 on 10 September in a mainchannel backwater. Both were released. In the Yampa River study area, 228 YOY squawfish were collected during 7-21 August at 17 sites between km 1.1 and 19.8. All specimens were captured in low-velocity habitats i.e., backwaters (198), embayments (23), concavities (4), isolated pools (2), and a single pool (1). Age is estimated to range from 10 to 24 days post-spawning with a resultant spawning period of 24 July-II August. No yearling squawfish were collected in the Yampa study area in 1983. 1983 Drift Net Collections A total of 4495 individuals representing 13 species was collected at both sites combined in the Yampa River study area in 1983 (Table 2). The total catch at Station 1 was nearly twice that at Station 2. Twelve species were collected at Station 1 of which sand shiners (Notropis stramineus), white suckers (Catostomus commersoni), and mottled sculpins �Table 1. Young-of-year Colorado squawfish seine collections, Colorado and Yampa River study areas, 1983 Study area Date Colorado River 11 Aug 9 Sep Total and range Yampa River TL (rnrn) River Km N 236.4 2 9.0,10.3 24 MC BA 11-16 1 3 21.5 9.0-21.5 22 MC BA 32 11-32 237.3 - Temp (C) Habitat 20 10 Estimated age (days) 7 Aug 15.5 4 9.0-9.5 28 MC BA 11-13 8 Aug 12.9 11.6 10.8 5.5 2.6 2.1 1 3 4 2 2 1 9.8 9.0-9.9 9.4-10.5 9.5, 9.6 9.5, 10.3 10.0 25 26 25 27 30 33 sr MC MC MC MC MC PO EM EM EM CO EM 14 11-14 13-16 13-14 13-16 15 20 Aug 19.8 15.9 5.0 5.0 4.2 1 2 1 2 2 12.6 12.8, 14.4 11.6 12.0, 12.4 8.6, 12.5 23 25 25 25 25 MC SC MC MC HC BA IP BA CO BA 20 21-22 19 19,20 10,20 21 Aug 3.5 3.2 2.6 1.6 1.1 1 l3 141 19 29 228 12.2 11.0-13.2 9.3-16.6 10.8-13.7 9.9-15.0 8.6-16.6 25 24 25 25 26 MC MC MC MC MC BA EM BA BA BA 20 17-21 13-24 17-22 15-23 10-24 Total and range v Estimated spawning period (dates) 27 Jul-10 Aug 24 Jul-ll Aug I-' CJ) ~ �Table 2. Number per species and % of total, 1983 drift-net collections, Yampa River Study Area. Station 1 Species Cyprinidae Speckled dace (Rhinichthys osculus)a Unidentified chub (Gila spp.)a Colorado squawfish (Ptychochielus lucius)a Common carp (Cyprinus carpio) Redside shiner (Richard~onius balteatus) Fathead minnow (Pimephales promelas) Sand shiner (Notropis stramineus) Red shiner (~. lutrensis) Catostomidae Bluehead sucker (Cat~stomus discobolus)a Flannelmouth sucker (C. latipennis)a White sucker (f. commersoni) Ictaluridae Channel catfish (Ictalurus Eunctatus) Cottidae Mottled sculpin (Cottus bairdi)a Total Number 829 578 1 40 1 1 1 Station 2 Number % 28 19 *1 * * * 1269 52 1 42 2 211 7 2 * aNative species. *Less than 0.5% of total number. 2731 255 1 661 18 Number % 7 6 7 * * * * 43 1 * 2986 Natives Exotics 108 94 102 7 2 2 Total 514 34 1509 91 9 983 526 % 937 672 103 47 3 3 1 1 21 15 2 1 1930 70 1 43 1 725 16 2 * * * * * * 4495 65 35 3714 781 83 17 f-' \.0 0 �191 (Cottus bairdi) were exclusive to this site. Ten species were identified from samples collected at Station 2 where a single red shiner (~. luterensis) represented the only species exclusive to this locality. Overall, larval forms predominated in most samples, while juveniles and adults constituted <5% of the total combined catch. Native species were far more abundant at both sites and comprised 83% of the total combined catch. Bluehead suckers (~. discobo1us), speckled dace (Rhinichthys osculus), and unidentified chubs (Gila spp.) were the most common natives and collectively comprised nearly 80% of the total catch at both sites combined. Channel catfish (Ictalurus punctatus) were the predominant exotic species at both sites but were collected in substantial numbers only at the lower site (i.e., 514 individuals 34% relative abundance). No other exotic species was collected in numbers exceeding 1% of the total catch at either site. A total of 103 YOY Colorado squawfish was collected by drift net between 20 July and 21 August (Tables 2, 3). Water temperature at the sampling sites ranged from 21.5 to 28.8 C. Mesolarvae (range 8.0 - 9.3 mm TL) comprised approximately 56% of the total catch. All remaining specimens were protolarvae (range 7.1 - 9.2 mm TL). A single YOY (TL =7.3 mm) collected at Harding Hole indicates at least limited reproduction some unknown distance above this site in 1983 and is the first documentation of reproduction above km 28.8 in Yampa Canyon. Drift densities at Station 2 ranged from 0.4 individuals per 1000 m3 on several occasions to 26.0/1000 m3 in a midnight sample on 9 August (Table 4). Squawfish drift densities were clearly higher at Station 2 during 8-10 August when mean water temperatures ranged from 24.5 to 25.3 C. At this time, both protolarvae and metalarvae were present in the water column. Table 3. YOY Colorado squawfish drift net collections, Yampa River, 1983. Date 20-21 Jula 22-23 Jul 23-25 Jul 3-4 Aug 8-9 Aug 9-10 Aug 15-16 Aug 16-17 Aug 20-21 Aug Totals aStation 1. N TL (mm) Estimated age (days) Estimated spawning period (dates) 1 5 3 3 63 11 2 11 4 7.3 8.2 - 9.2 8.8 - 9.0 7.6 - 8.0 7.4 - 9.3 8.1 - 9.2 8.5, 9.2 8.0 - 9.0 7.3 - 8.4 3 8-12 11 5-7 4-13 6-12 9,12 7-11 3-9 19 Jul 11-15 Jul 13 Jul 29-31 Jul 27 Jul-5 Aug 29 Jul-4 Aug 5-8 Aug 7-11 Aug 13-19 Aug 103 7.3 - 9.3 3-13 11 Jul-19 Aug (All remaining captures were made at Station 2.) �~ 1.0 r<) Table 4. Densities of drifting Colorado squawfish YOY (no/1000 m3) at two sites in the Yampa River study area, 1983. Values equal densities determined from 1-2 hr samples over a 24-hr period (dawn/noon/dusk/midnight). Date Mean Temperature (C) Station 1 Protolarvae (7.3)a - / Protolarvae Station 2 (7.4-9.2) Mesolarvae Total (8.0-9.3) - 20-21 Jul 21.5 22-23 Jul 22 23-25 Jul 22 3-4 Aug 23.5 8-9 Aug 25.3 9-10 Aug 24.5 - / - / 1.1/ 1.2 15-16 Aug 24.8 - / - 16-17 Aug 25.3 - / - / 1.5/ 1.5 20-21 Aug 28.8 - / 0.9/ - aTotal length range (mm) of the particular - 0.4/ - / - / - 0.8/ 1.6/ / - / 0.5/ - 0.7/ - / - / - 0.7/ / - / 0.8/ 0.4 - 1.2/13.2/ 2.9/10.2 / - - / - / - - / 0.5/ - / 0.8/ 0.4 14.6/ 1.7/ 5.7/15.8 15.8/14.9/ 8.6/26.0 2.8/ 3.3/ 1.1/ 3.5 2.8/ 3.3/ 2.2/ 4.7 - / 1.3 0.4/ 1.0/ 0.5/ larval phase. - / 0.9/ - / 0.4/ 1.6/ - / - / - / 0.7/ 0.7 - / 1.0/ - / - - / - / - / - / 2.2/ 2.2 - / 1.3 0.4/ 2.0/ 0.5/ - �193 Overall, no significant diel periodicity in drift densities was observed for either protolarval or mesolarval squawfish; however, die 1 periodicity tendencies in drift abundance were noted. Density-drift rates of protolarvae tended to be higher in noon and midnight samples with a sunrise:noon:sunset:midnight abundance ratio of approximately 1:8:4:9 (Hotelling's t2, p = 0.28; 2-AOV, P = 0.33). Mesolarval squawfish exhibited somewhat higher drift densities in sunrise-midnight samples, wi th an abundance ratio of 3: 1: 1:3 (Hotelling' s t2, p == 0.64; 2-AOV, p = 0.60). Analysis of YOY Squawfish, 1979-83 Since 1979, 105 YOY Colorado squawfish have been collected in the Colorado River study area between km 212.2 and 245.7 (Table 5). Using revised predictive equations, spawning in this river reach apparently occurred as early as 3 July in 1981 and extending as late as 6 September in 1980. For the Yampa River study area, 317 specimens have been collected by seine/dip net sampling since 1980 (Table 6), while an additional 103 were collected by drift net in 1983. Spawning in this area appears to have occurred as early as 2 July in 1980 and as late as 11 August in 1983. Duration of spawning during these years would appear to have been as long as 2 months (Colorado River, 1980) and as short as 1 day (Colorado River, 1981); however, in general, spawning in both rivers apparently lasts about 2-2~ weeks. Prior to 1983, the relatively low numbers of individuals collected prohibited realistic evaluations of spawning "intensity" between the earliest and latest dates. When the combined 1983 Yampa River seine/drift net collections are considered, however, there is strong evidence that limited spawning activity occurred as early as 11 July when mean water temperature at the drift net sites ranged from 18-20 C, but that a peak of activity occurred between 25 July and 10 August when mean water temperatures ranged from 22-25 C (Fig. 2). Spawning after 10 August appears to be sparse. In general, spawning dates for both study areas for 1979-83 coincide with decreasing flows and rising water temperatures (Figs. 3, 4). These observations support similar observations by Vanicek and Kramer (1969), Seethaler (1978), and Holden (1979, 1980). It is not possible, at this time however, to state with any degree of certainty a quantitative relationship between reproductive success and flow characteristics e;ccept in general terms. In the Yampa study area, the number of discreet samples collected in confirmed YOY habitat between the earliest and latest estimated spawning dates was greater in both 1981 and 1982 (i.e., 166 and 168, respectively) than in 1983 when 144 samples were collected. Yet, the relative abundance of YOY in 1983 (C/E = 6.02/100 m2~ was 7-30 times greater than 1981 or 1982 (C/E = 0.86 and 0.19/100 m , respectively). Clearly, the production of YOY squawfish was greater in 1983 than the two previous years and, similarly, peak flow in 1983 (i.e., ~ 650 m3/sec in late-May and early-June) was substantially greater than during the same period in 1981 and 1982 (i.e., ~ 300 and - 450 m3/sec, respectively). If water temperatures are examined as 22 C-days, it appears that 1983 was roughly comparable with 1982 (1,358.0 and 1398.5, respectively) and that both were exceeded by 1981 (2,147.5). These �Table 5, yay Colorado squawfish collections, Colorado River, 1979-83. 1979-80 values include collectLons made with dip nets 1981-1983 collections were made exclusively with seines. Year No. of samplesa Datesb Localitiesc (km) N C/Ed 1979 155(2) 24, 26 Aug 217.7, 223.6 8 1980 114(11) 13 Aug - 2 Nov 212.2-245.7 1981 319(1) 27 Jul 1982 384 (10) 1983 206(2) TL range (nun) Est. spawning period (dates) - 15.0 - 24.5 16 Jul-4 Aug 77 - 13.0 - 38.0 16 Jul - 6 Sep 243.6 1 0.05 18.0 3 Jul 27 Jul, 23 Aug 216.1-244.4 16 0.31 7.8 - 20.1 15 Jul - 11 Aug 11 Aug, 9 Sep 236.4, 237.3 3 0.15 9.0 - 21.5 27 Jul - 10 Aug aNumbers in parentheses represent number of collections which .contained bDates of earliest and latest YOY collection. cDistance from confluence with Green River in Utah (km 0) dYOY/I00 m2 specific habitat. > 1 YaY. I-' \0 +:-- �Table 6. YOY Colorado squawfish collections, Yampa River, 1980-83. 1980 values include collections made with dip nets, 1981-83 collections were made exclusively with seines. Year No. of aampLes f Datesb Localitiesc (km) 1980 215(9) 23-25 Aug 0.2 - 13.9 46 1981 519 (11) 24 Jul-15 Aug 0.2 - 28.8 1982 310 (10) 7-25 Aug 1983 245 (17) 22 Jul-21 Aug TL range (mm) Est. spawning period (dates) - 14.0 - 29.0 2 Jul-8 Aug 23 0.86 9.0 - 22.0 0.5 - 19.6 20 0.19 9.9 - 21.0 24 Jul-8 Aug 1.1 - 19.8 228 6.02 8.6 - 16.9 24 Jul-ll Aug C/Ed N - aNumbers in parentheses represent numbers of collections which contained> 6-30 Jul 1 YOY. bDates of earliest and latest YOY collection. cDistance from mouth of Yampa River (km 0). dNa. YOY/100 m2 specific habitat. f-' 1.0 l.I1 �r-' \.0 0"\ 40· 1 5, t to 30· to o z 1983 N=331 5 t982 N-20 , ,no"lQOn )L" , n " N -48 5 ,O,r] pD , r111',11 on )L-!ln to 20 30 , , to AUG JUly 10 30 10 20 JULY 30 i t98t N-23 "l"~ 20 fl, , 10 20 AUG PREDICTED SPAWNING DATES Fig. 2. Number of YOY Colorado squawfish and predicted spawning dates for 1980-1983, Yampa River study area, Colorado. For 1983, black bars c drift-net captures. Open bars represent seine captures. 20 �197 0 ~ 30 26 22 18 14 10 6 2 1\ ~ ~¥> ~ AMJJAS AMJJAS AMJJAS -jA ~ AMJ J AS 30 26 22 18 1. 10 9 2 AMJJAS 1500 1500 1000 1000 500 500 0 (]) <Il 1 W o ex: « J: o en 0 100 100 ~ en z o i= o AMJ J AS 1979 W N 8 -l -l C/E; - o AMJJAS 1980 77 AMJJAS 1981 AMJJ AS 1982 1 16· 0.05 0.31 AMJJAS 1983 3 0.15 o Fig. 3. Relationship of flows and temperatures to predicted spawnin~ dates of Colorado squawfish, 1979-83, Colorado River study area, Colorado. N = total capture; C/E ; number/100 m2. �198 0 2- I- 30 28 30 26 22 J/ i 18 14 10 / /~~/ / / e 2 / / 22 /1 18 14 ./.1/ / . _.i / .I 10 8 2 600 600 500 500 400 400 300 300 200 200 100 100 50 50 T o Q) .,til §. W c:l a: -c ::I: o en C en z o AMJJAS i= o w _J _J o o N C/E AMJJAS AMJJAS AMJJAS 1980 1981 1982 1983 48 23 20 331 0.88 0.19 6.02 Fig. 4. Relationship of flows and temperatures to predicted spawning dates of Colorado squawfish, 1980-1983, Yampa River study area, Colorado. N = total capture; C/E = number/100 m2. �199 observed trends strongly suggest that, for the Yampa River spawning areas, a relatively high 1983 spring flow in combination with an accumulation of 22 C - days roughly comparable with that for a lower flow year (1982) served to enhance overall spawning success. Further, given that 1983 was a poor reproductive year for most exotic species in the Yampa study area (based upon drift net captures). the high flows of that year served additionally to reduce potential interspecific interactions in the low-velocity shoreli.ne habitats used by YOY squawf Lsh , In the Colorado River study area, fewer samples were collected between the earliest and latest spawning dates in 1983 than in the two previous years (i.e. 97, in 1983 vs. 139 and 166 in 1981 and 1982, respectively). YOY squawfish relative abundance in 1983 was less (C/E = 0.15/100 m2) than 1982 (C/E = 0.33/100 m2) but greater than 1981 (C/E = 0.05/100 m2)o As in the case of the Yampa. 1983 Colorado River peak flows were relatively high (~1700 m3/sec); however, 22 C-day accumulation in 1983 (879) was substantially less than the two preceeding years (i.e., 1981 = 1,759 + and 1982 = 1,147), and only approximately 65% that of the Yampa in the same year, Similarly, the duration of water temperatures > 22 C on the Colorado River in 1983 was only 38 days (6 Aug-12 Sept) in comparison with 57 days on the Yampa River (19 July-13 Sept). Based upon observations for the years 1981-83, it is possible to suggest that, in general, squawfish reproductive success is enhanced by "high" late-spring flows which provide high quality/quantity spawning substrate while, at the same time, serve to limit non-native reproductive success. At the same time, "optimal" flows must be followed by a sufficient accumulation of degree-days. At this time. data are too sparse to meaningfully define "high" and/or "optimal" in a quantitative s~nse; however, it is clear that flows of the magnitude of 1981 Colorado River flows are inadequate, as were 1983 degree-days on the same river. Conversely, it would appear that the 1983 flow/temperature regime on the Yampa provides insight into "optimal" reproductive conditions for this species. Razorback No YOY or juvenile razorback suckers in either study area in 1983. Fourth-Year Sucker (Xyrauchen texanus) were observed (1984) Investigations Seine/drift-net studies will be limited to the Yampa River study area in 1984. A presently undetermined number of areas which have yielded YOY squawfish since 1980 will be seined periodically following the first recorded water temperature of 18 C at Deerlodge Park. Seine samples w i.Ll.be collected every 7-10 days through early-September. DrHtnet sampling will be conducted on approximately the same time scale at Station 2 near the Green River confluence. To investigate YOY movement in the Green River, a drift-net station will be established in the vicinity of the Colorado-Utah state line in Whirlpool Canyon. �200 LITERATURE CITED Andrewartha, H. G., and L. C. Birch. 1954. The distribution and abundance of animals. Univ. Chicago Press, Chicago, Ill. 782pp. Arnold, C. Y. computing 1960. Maximum-minimum temperatures as a basis for heat units. Proc. Amer. Soc. Hort. Sci. 76: 682-692. B~skerville, G. L. and P. Emin. 1969. Rapid estimation of heat accumulation from maximum and minimum temperatures. Ecology. 50(3) :514-517. Caughley, G. 1978. Analysis of vertebrate & Sons, Ltd., New York, N.Y. 234pp. populations. Hamman, R. L. 1981. Spawning and culture of Colorado in raceways. Prog. Fish-Cult. 43:173-177. John Wiley squawfish Hayes, F. R. 1949. The growth, general chemistry and temperature relations of salmonid eggs. Quart. Rev. of BioI. 24:281-308. Haynes, C. M., and R. T. Muth. 1982. Identification of habitat requirements and limiting factors for Colorado squawfish and humpback chubs. Prog. Rep., Colo. Div. Wildl., Endangered Wildl. Invest. SE-3-4. 43pp. Haynes, C. M., R. T. Muth, and G. T. Skiba. 1983. Identification of habitat requirements and limiting factors for Colorado squawfish and humpback chubs. Prog. Rep., Colo. Div. Wildl., Endangered Wildl. Invest. SE-3-5. 18pp. _____ , T. A. Lytle, E. J. Wick, and R. T. Muth. 1984. Larval Colorado squawfish Ptychocheilus lucius Girard in the Upper Colorado River basin, Colorado, 1979-1981. Southwest Nat. 29 (1) :21-33. Holden, P. B. 1979. Ecology of riverine fishes in regulated stream systems with emphasis on the Colorado River. Pages 57-74 in J. V. Ward and J. A. Standford, eds. The Ecology of Regulated Streams. Plenum Publ. Corp., New York, N.Y. _____ 1980. The relationship between flows in the Yampa River and success of rare fish populations in the Green River system PR--31-1, BIO/WEST, Inc., Logan, Utah. 32pp. ____ , and E. J. Wick. 1982. Life history and prospects of Colorado squawfish. Pages 98--108 in W. H. Miller, and C. A. Carlson, eds. Fishes of the upper Colorado present and future. Proc. Symp. Annu. Mtg. Am. Fish. Albuquerque, N. M. for recovery H. M. Tyus, River system: Soc. 1981, �201 Lackey, R. T., and W. A. Hubert. 1978. Analysis of exploited fish populations. Rep. VPl-SG-76-04, Sea Grant, Va. Poly. lnst. and State Univ., Blacksburg 97pp. Muth, R. T., C. M. Haynes, and C. A. Carlson. chub Gila robusta (in preparation). 1984. Culture of roundtail Ricker, W. E. 1975. Computation and interpretation of biological statistics of fish populations. Bull. Fish. Res. Bd. Can. 191:382pp. Seethaler, K. 1978. Life history and ecology of the Colorado squawfish (Ptychocheilus lucius) in the upper Colorado River basin. M. S. Thesis, Utah State Univ., Logan. 156pp. Southwood, T. R. E. 1978. Ecological methods with particular reference to the study of insect populations. Chapman and Hall Pub., New York, N.Y. Tanner, J. T. 1978. Guide to the study of animal populations. Tenn. Press, Knoxville, 186pp. Univ. Toney, D. P. 1974. Observations on the propagation and rearing of two endangered fish species in a hatchery environment. Proc. West. Assoc. State Game and Fish Comm. 54:252-259. Tyus, H. M., E. J. Wick, and D. L. Skates. 1982. A spawning migration of Colorado squawfish Ptychocheilus lucius in the Yampa and Green rivers, Colorado and Utah, 1981. Annu. Symp., Desert Fishes Counc., Death Valley Natl. Monument., Calif. 13:3. United States Geological Survey. 1982. Water resources data. Colorado. Water year 1981. Vol. 3. Dolores River Basin, Green River Basin. and San Juan River Basin. USGS Water-Data Rep. CO-81-3. 436pp. 1983. Water resources data. Colorado. Water year 1982. Vol. 3. Dolores River Basin, Green River Basin, and San Juan River Basin. USGS Water-Data Rep. CO-82-3. 366pp. Valdez, R. A., and G. H. Clemmer. 1982. Life history and prospects for recovery of the humpback and bony tail chub. Pages 109-119 in W. H. Miller, H. M. Tyus, and C. A. Carlson, eds. Fishes of the upper Colorado River system: present and future. Proc. Symp. Annu. Mtg. Fish. Soc., 1981, Albuquerque, N. M. Vanicek, C. D., and R. H. Kramer. 1969. Life history of the Colorado squawfish, Ptychocheilus lucius and the Colorado chub, Gila robusta, in the Green River in Dinosaur National Monument, 1964-1966. Trans. Am. Fish. Soc. 98:193-208. �202 Wick, E. J., T. A. Lytle, and C. M. Haynes. 1981. Colorado squawfish and humpback chub population and habitat monitoring, 1979-1980. Prog. Rep., Colo. Div. Wildl. Endangered Wildl. Invest., SE-3-3. 156pp. ----- , D. L. Stoneburner, and J. A. Hawkins. 1983. Observations on the ecology of Colorado squawfish Ptychocheilus lucius in the Yampa River, Colorado, 1982. Nat. Park Ser., Water Res. Field Support Lab Rep. 87-7, Colo. Div. Wildl. Endangered Wildl. Invest. SE 3-5, 55pp. �Colorado Divisior Wildlife Research January 1984 203 of Wildlife Report JOB FINAL REPORT State of Colorado Project N-3-R-2 Work Plan, 1 --~-------------1 Job Period Covered: Author: Natural History of the Brazilian Free-tailed Bat in the San Luis Valley, Colorado 1 July 1983-30 June 1984 Peggy Svoboda Personnel: C. Braun, W. Graul, and C. Haynes, Colorado Division of Wildlife; J. Choate, Fort Hays State University, Hays, Kansas. ABSTRACT The natural history of a colony of Brazilian free-tailed bats, Tadarida brasiliensis mexicana (Saussure), in an atypical, predominantly male roost in south-central Colorado was investigated in 1982 and 1983. The colony inhabits the Orient Mine which is located at the base of the Sangre de Cristo Mountains in the northern end of the San Luis Valley. The valley is in the Upper Sonoran Life Zone. Iron ore was removed from the mine (elevation, 2800 - 3000 m) between 1881 and 1932. Colonization by freetailed bats almost certainly occurred no earlier than 1900 and most likely occurred after the mine closed in 1932. The colony roosts in a cavernous underground stope in which ambient temperature varies from 60 to 1Zoc when bats are present from mid-June through October. At peak numbers, the colony consists of about 100,000 bats. Outflights in both years lasted an average of nearly one hour. Bats emerged earlier relative to sunset in late summer and autumn than in early summer. Timing of outflights was not affected by percent cloud cover. Outflights were dispersed in June and became more concentrated, often with serpentine components, later in the year. Capture samples taken with mist nets indicated that composition of the colony was 98% adult males until mid-August, when adult females and young-of-the-year became more common. ��205 THESIS NATURAL HISTORY AND GEOGRAPHIC RELATIONSHIPS OF THE BRAZILIAN FREE-TAILED BAT IN THE SAN LUIS VALLEY OF COLORADO Submitted by Peggy L. Svoboda Department In partial fulfillment of Biology of the requirements for the Degree of Master of Science Fort Hays State University Hays, Kansas Spring 1984 �206 ACKNOWLEDGMENTS I thank Colorado work on its property in company Dr. W. D. supervisor 1983-84, field and for access Within the Colorado Graul helped initiate in 1982, C. M. Haynes and S. J. Bissell who helped valuable mining. observations in Wildlife A. Bogan, Fort Wildlife park for frozen Monument, assisted made Munger, technical came E. J. Field D. Carnahan, assisted life in the field S. Smith, The at records of bats U.S. Fish techniques. and Dr. M. of the U. S. Fish provided and space K. Fors, and B. Riddle field more work. pleasant: R. Upham, M. Ploch, manuscript and superintendent Station, Encouragement from F. and G. Svoboda, much prel iminary A number of L. Hirsch, M. B. Upham, was from home J. Upham, much and abroad and especially reviewed of Museum N. Seitz and T. Seitz, who also provided assistance. Paradice. with and geology and C. Chase of the Denver History and especially about provided Station in to K. Balleweg, made with marking Fort Collins K. Sewell, discussion J. Cummings, Field Fort Hays State University people Wunder R. Reynolds, museum. specimens. of Natural L. was the project helpful and information and Collins Service and Fig. 2, and provided at the mine. Service, supervised provided assistance the project mines, prepared Sand Dunes National housed the of Wildlife, grateful Freeman J. Division to information I am especially explore field for permission to historical files. assistance. Great Fuel and Iron Company Dr. W. by Dr. J. R. �207 Choate Ely, and other members E. D. Fleharty, Zakrzewski the Colorado by the Division of my graduate M. E. Nelson, of Fort Hays State Division Nongame of wildlife. F. W. University of Wildlife). Research committee Program, (Drs. C. A. Potter, R. J. and W. D. Graul This project Project N-3R, of was funded Colorado �208 INTRODUCTION The distribution of brasiliensis) extends (Tadarida the West the and north coast to southern Oregon (Hall, 1981; north localities as Nebraska and Iowa (Glass, (Farney £. States--!. b. mexicana cynocephala is the smaller important distinction be non-migratory movements. caves summer wintering areas mexicana populations, colony of !. £. (Cockrum, in 1955). T. b. mexicana differ (Krutzsch, local, T. seasonal roosts mainly long 1969); Texas. distances an in to exception males of T. b. Southwestern from the Pacific coastal to Mexico 1955). but a more and is thought (1974) involved eastern and T. and migration. only migrates and LaVal in one of the migratory b , mexicana. South in the United (Schwartz, T. b. mexicana which do not migrate in caves exist 1970), and in the Southeast structures make usually overwintering populations not live to in Mexico by Spenrath (Walley, to roosting in man-made or and 1975), of the two subspecies, In the Southwest, in reported (Le Conte) pertains roosts Jones, are recognized in the West me~icana Schwartz, 1960). presently (Saussure) cynocephala and to along for the species 1982), Illinois (Smith and Goodpaster, Two subspecies to of record bat and Chile States as £. from Argentina United Isolated Ohio free-tailed and southern 1955). Dakota Brazilian Indies west far the and typically This study pertains southwestern populations do to a of T. �209 The Brazilian researchers free-tailed et al., 1968; Davis R. and Cockrum, in maternity Mexico Creek Cave, cited by numbers et economically quantities two of Carlsbad the 1970; of these bats areas New and Eagle Reidinger, declines mortality, by large Caverns, et alG' 1979), These Villa- was stimulated in 1972, focused especially regularly feed and where pesticides of guano that nearly be triple fertilizer. that free-tailed One presence environmental Altenbach,1973). are insects 1926). are used year. 1 g of insects nightly Guano bats been has guano of mercury contaminant they deposit to assess and consume large and produce thick Davis et ala tons of insects are This estimate per night; consumption amount. stratified ecologically In Texas, 6,000 metric bats every bat eating bat might of flying (Nelson, by free-tailed bats in part because of nocturnal (1969) thought this causes farmed and Davis for the bats 1981). because important on each colonies interest 1969, free-tailed (1962) estimated based al., link 1970; Gel uso et al., 1981). Brazilian consumed these possible in intensively deposits of (Cockrum, poisoning (Cockrum, of its possible 1962, 1967; Constantine in the Southwest--in Arizona on Recently, 1937; Altenbach Geluso attention drink 1962). (Allison, (Constantine, interested et al., 1962; Eads et al., 1955; colonies pesticide initially in the United States because to the spread of rabies declines bat of mined Barbour insects produced by and has been per large used as analyzed the persistence in a food chain was (Petit of and �210 Armstrong brasiliensis regular (1972) reported in Colorado resident reported that, brasiliensis abandoned edge the in August of the San state. a Luis Meacham of mine records 1968, 9,000 Valley in the a however, or that is located of T. the species (1971), in Colorado This mine since 1932. occasional and did not consider of occupied only more had T. been at the northeast Sangre de Cristo Mountains. Records of T. brasiliensis from this mine have Garfield 1973, one, male sex unknown; 1977. Durango, (T. Lytle, Verde one Mesa male National Co., Monte comma Park, Vista, litt.); Saguache Springs, 20 Reynolds, pers. comm.). The colony However, biology 1981 suggested few and in Colorado casual breeding 1965, View one ecology of was not studied observation females and unknown, one male Rio Grande (Cockrum, near a Hot R. isolated before of males in 1972; between maximum 1969. Mineral apparently intensively mostly reaches pers. comma to J. (Armstrong, the Co., two males; of the colony that it consists Plata Co., Mesa Resort male La 9 Montezuma 1964, one male Co., Valley July 1936, 1973. one Co., Trinidad, (P. Somers, date 20 October 16 July 1978); to J. Freeman); 20 July (Armstrong, Co., Gunnison, Junction, 23 August (Fig. 1): 1907, five males (Ellinwood, one male than those individuals west of Gunnison, Co., Grand pers. other pers. comm.); Las Animas date unknown, Freeman); 16 July 1908, 1910); Gunnison (J. Freeman, August been of scattered Co., Newcastle, 1972: Warren, in Colorado 1982. 1978 and and only size a of about �211 _1~O~7 ~1;O~5~ ~1,O~3 o o ••• o 150 km 108 101 Fig. 1. _ Distribution of Tadarida brasiliensis 103 in Colorado. �212 100,000 individuals colony would be regarded were a male northern as unusually summer roost. to compare in press). large The location features investigate the the histo~y by free-tailed bats; to document describe objectives of the colony 2) to locate activities; to document its composition; reproduction in the colony: 5) 8) 1) the roost; of the colony; to assess to locate to to colonization to estimate 7) marginal were: and describe chronology on the farther south. study of the mine relative seasonal its daily this it a unique of a geographically of This if, indeed, with those of the better known colonies Accordingly, 3) and Wunder, margin of the range of the species provided opportunity colony (Freeman 4) to its size; 6) the amount and describe of areas where the bats feed in the San Luis Valley. STUDY AREA Location.--The about colony 32 km southeast is in Saguache of Poncha (380 13' N, 1050 48' W). County, Colorado. Pass in the San Luis Valley This valley is in the south-central part of the state and is a major basin in the Rio Grande system. The valley of 2285 from to 2440 maximum width Mexico (Ramaley, northern covers m, of 80 km. 1942). about 10,360 km2 at an elevation is about 240 It extends Poncha end of the valley, Pass is at the Juan and Sangre de Cristo mountains Range from the Sangre de Cristos. rift km about long, and has 24 km into (2,746 m), at a New the junction of the San and separates the Sawatch The Sangre de Cristos form �213 the eastern Mountains edge roosts lens-shaped limestone the valley, form the western The colony where of whereas stope of limonitic (Litsey, 1960; Stone, iron ore the hillside and to a large pit by openings 3000 2 to 10 m. de Cristo Main ffi. tailing piles 1934). The stope The hillside Mountains openings Juan in Paleozoic is connected ranging the mine on a southwest-facing of the of about are marked slope to in size is at the base at an elevation to Mine, 1934) was mined at least from 1931 Sangre (Stone, of the Orient 1881 through from about San edge. in a large bodies the 2800 to by large and overlook the flat expanse of the San Luis. Valley. Many Luis mines Valley. are located However, iron-producing in the hills the Orient Mine mine in the valley. in this formation for use by bats before mining began (J. B. Stone, Company). (six main large in Using stope mining litt. inaccessible sub-levels) and natural The full extent the this technique, and seven stopes to caves long 1934:321). were not open to the surface and, therefore, Open underhand solution of feet" (Stone, the San was the only Natural "60 or 70 feet wide~ and "hundreds present encircling major cavities also These were cavities were unavailable in 1881. was used to extract Colorado miners developed of tunnels (K. Balleweg, of the mine is difficult Fuel the ore and Iron 13 levels that connected pers. comm.). to assess because of tunnels. Geo!~gy.--Hiddle basin-and-range to late style block Cenozoic faulting, volcanism, and erosion uplift, formed the �214 q eoriorph i c features major al.,1976). from 'l'hevalley the Cristos San years Cretaceous Mountains ago The Laramide and were disrupted period Poncha Pliocene, valley. accompanied Pass a fault and present-day scarp San Luis Valley the of In the Oligocene. by Miocene. flowed southward Luis Valley. In the blocked the San Valley withdrawal River drainage River about 70 in the early south of Poncha the Arkansas Arkansas by de This was followed and faulting into occurred Colorado. the Arkansas (Epis et of the Sangre uplift by volcanism. of uplift in the Miocene, through was Valley that dips e a s t we rd graben to the base seas in south-central drainages Late is a half (Tweto, 1978). million another Juan in the San Luis Springs was diverted into (Epis et al., 1976). now is part of the drainage the The basin of the Rio Grande. Climate.--The short summers climate and cold, 320C in summer. exceed is sunny dry winters. From drainage basin Monthly average January to l6.60C in July 1978). At Saguache 1931 to 1960, temperature temperatures ranged from (31 km than the colony), about SoC in 1982-83. -300C on 7 February -28°C on 29 December States southwest the average Extreme temperatures to 290C on 23 July to 29°C on several Dept. Commerce, 1983, rarely of 4.80C. about -8.20C in Dept. Commerce, of and annual cool, the Rio Grande annual (United with Temperatures had an average lower (United States generally about 1750 temperature m was in 1982 were from and in 1983 were from days in June and July 1984). �215 In Sa0uache, above the average UOC from April last spring the mean (United spring States temperature October. Dept. autumn Commerce, usually 1960) was 4 June freeze 1978). was Mine, the last In 1982, was on 15 June of autumn Dept. Commerce, frost I experienced in 1982 was not recorded before the last freeze was 1983). in spring and in 1983 was on 13 June. and 18 September freeze was on 18 June and the first autumn (United States is The mean date of the (from 1931 through of the first on 29 September Orient through freeze date monthly At the of 1982 The first mid-August frost and in 1983 was on 15 September. Annual precipitation 1931 through with 1960. monthly (United lowest season Dept. humidity is in (from June through 35.4 cc average standard white cylindrical valley in evaporation late June daily Hydrographic streams the early slightly is on 1942). valley. Average 40% and rarely during is high. evaporation from an open surface respectively Precipitation is about atmometer and and August. Evaporation August) 126.7 cm for 1927 through valley, low 1942). are July and increases for June and July 70% (Ramaley, 26.8 cm from 1978). (Ramaley, humidity averaged 4.2 cm, slopes recorded was 3.6 cm and Commerce, and eastern Relative exceeds of months in the center of the valley the western relative The wettest averages States in the valley the growing Ramaley (l942) from a Livingston at the south end of the July. Average in the middle annual of the valley 1931. fea tures .--At the northern end that flow from the Sangre de Cristos of the quickly �216 disappear into the porous, In the center shallow of the valley, is the driest for agriculture irrigation systems on the hillside Hot surfaces reappears flooro as springs because or about mostly is confined wastes produced Hot 2630 Springs on 10 km to the southwest and canal Hot springs surface m). Mine at Valley Hot the water west side of the mine. also of the This water ponds holding at a nearby hog farm. in some of the year. The hydrology wells to a series of open settling flows Much tunnels of this of the water Canyon level at the point was flooded mine at freezes of the mine in 1983 differed the fifth from Orient been built. the San it has adequate 6f numerous (elevation, at Mineral in 1982; area of Colorado, that have Springs Water Although about 2 km south of the Orient valley times water of the valley of the north end of the valley. Luis Valley View soil lakes. 'l'heSan Luis Creek flows from Poncha Pass down the middle water sandy certain in winter. from that observed of the main entrance in the summer of 1983, whereas it was dry in 1982. Vegetation.--The desert scrub Li fe Zone is the association (Cary I dominant (Chrysothamnus (Atriplex corresponding Greasewood 1911). shrub. sppo) are Common smithii). Several is canescens) r a b b i t.br u sh , (Agropyron floor of the San Luis Valley blue common. present grasses grama to the Upper (Sarcobatus species and but are is an open vermiculatus) of rabbitbrush fourwing 1 ess sal tbush abundant western (Bouteloua Sonoran wheatgra.ss gracilis). inland than alkali saltgrass �217 (Q!~!!£~!~~ ~!~i£!~), and a i ro i de s Lr occur sedge (Care~ in patches. alkaline floor association. (Anderson The (~E~~~e~!~~ sacaton spp.) associations Scattered ponds and and sloughs grassland and areas of present. Riparian vegetation is a cottonwood (Popu!us) and willow soils valley alkali are The tallest cottonwoods and Owensby, 1969: Nelson, zone the around floor of the is sarothrae) are conspicuous. between the grasses include (Aristida sagebrush association and juniper coniferous forests lodgepole pine menziesii), predominant this zone (~bies steeper and Colorado species include blue at lower Engelmann concolor), the northern, (Anderson pine (Pinus the open woodland pine spruce pine Big undershrub- of pinyon elevations. limber threeawn 1942). are (pinus Eonderosa), Douglas-fir spruce Other spp.), dropseed in the above Above contorta), grows (Stipa comata). is open woodland in which ponderosa (Pinus stands. 1964; Ramaley, (Juniperus). grama of the valley slopes soapweed spp.), fendler occurs sections 1969: Harrington, on Blue (Muhlenbergia tridentata) and broom Small in pure (Agropyron muhly and southeastern Vegetation fir sometimes xeric (Opuntia polyacantha) of this zone. wheatgrass (Artemesia grassland pear spp.), and needleandthread and Owensby, edulis) and fendleriana), (Sporobolus eastern, components shrubs are common. prickly 1942). valley snakeweed and plains (Sa!l~) 1969: Ramaley, Species of short rabbitbrush (Yucca glauca) the are 12 to 15 m tall undershrub-grassland. (Gutierrezia on (Picea Other (Pseudotsuga pungens) conifers (~. en~lmannii), (Pinus are in white !.!.exi_!is), and �218 foxtail (f. pine elevations aristata). on more mesic base of the mountains. occur in mesic areas Ramaley, this in the coniferous grows together Oak chaparral at the tremuloides) (Nelson, 1969; , (chiefly Sagebrush with Quercus also mountain just grows mahogany snowberry (Harrington, 1964; Ramaley, occurs arou~d the main entrances to the Mine. San Luis (Armstrong, fauna.--Although Valley, Plains elevation districts level. seven Species access to the valley Cristo Range in New Mexico, and Jemez Grande. access around around mountains. Mexico is not volant species adapted effective barrier the southern corridor dispersal are with at the subspecific from the south Great Plains have rim of the Sangre de rim of the San Juan Valley in northern for movement to open country. to endemic species from the Colorado The Rio Grande an effective the the southern whereas in and with the lower- Slope of be resemblances species reach the valley the have species occur might level of the Western along Plateau faunal at the specific Xeric-adapted Hio no endemic subspecies The closest 1972). the Great an Oak from 2 to 3.5 m tall. and rabbitbrush Mammalian the forest woodland. association, vaccinioides). orient of hills occurs at the north end of the valley montanus) -------- 1942) . exposures Stands of aspen (Popul~ the pinyon-juniper gambellii) in northern forests grow at lower 1942). Oak chaparral above Coniferous of New of non- Also, Poncha Pass is most arid-adapted �219 species from the Upper Arkansas River Valley into the San Luis Valley. Land use.--Land influenced ulfalfa, by irrigation. sweet clover, such as cabbage, center-pivot northern and potatoes, crops cauliflower, are used organophosphates systox-R, and chlorinated the (malathion, Orthene), organic the pyrethroid, Pydrin and 1942). Both used. The are incl ude Cygon, Meta- (Lanna te (Thiodan several These parathion, carbama tes (T. Harvey, beans, development, valley. Monitor, compounds wheat, are used for grazing. agricultural in been vegetables spinach, systems end and edges of the valley pesticides are barley, use (Ramaley, irrigation Because of intensive have and cool-climate and domestic canal in the valley Chief lettuce, t o r commercial peas use patterns and Sev in ), and toxaphene), and pers. comm.; L. Stevens, in litt.). Most of the land on the floor of the valley Public owned. Monte Vista National lands national Monument, in the area wildlife Bureau the edges of the valley, Forest around Mine is owned surrounded the valley refuges, of Land around the Alamosa the Great Management Fuel elevations. and Sand Dunes land and the Rio Grande at higher by the Colorado include is privately mostly National The Orient and Iron Company and is by land managed by the Bureau of Land Management. ~ffiTERIALS.ANDMETHODS Arrival observing and departure outside of the colony the roost for evidence were documented of an outflight by and �220 catching the bats opening in 5.5 and 9.1 m mist to discovered the mine. on 12 June roost was checked nets set at the edge of A. passage 1982, after roost was the bats had arrived. The for the presence into of bats the in the spring of 1983. In 1982, the mine was visited (range, In from 2 to 7) per week 1983, (range, the mine was 16 May an observing sunrise. ambient wind temperature, percent meter, cloud recorded. After first bat began and ended, when the last recorded. formation, direction and the bats appeared each of size photoestimation with was humidity in described was measured moon, were times when the outflight to the mine, and the morning in terms estimated by were of density, described by Humphrey was placed As the emerging rock wall, a modification bats photographs 2 min using 400 ASA Tri-X black-and-white bats the from the mine. flash attachment the edge of the mine. of the main returning pit with precipitation when and with a Dwyer hand- in spring, the measured phase of evening, technique camera and a vertical (MDST) of sunset wind, arrived were and direction 35-mm camera relative amount entered Outflights Colony times when bats began bat 3.0 nights 22 Hay and 5 November; wind speed measured cover, of in the first week in August of 1983. at the mine~ psychrometer, of 3.4 nights and 21 August. average from 0 to 6) per week between When held between visited the mine was not visited a sling an average for every 2-min (1971). a A on a tripod at flew between the were taken every film. interval of Speed of the to the nearest �221 0.01 s wi th a digi tal stop watch. were measured negatives to the nearest were processed, inspecting with a magnifying was multiplied length of any gaps across the frame. number of bats The numbers the number The number by the quantity was divided of bats in front by table when 2 min minus the speed gave of the photographic of the every were summed the the bats an estimate of the camera in 2-min intervals of bats passing on a light of bats per photograph obtained This calculation passing photographic in each frame were counted by and negatives glass. in the outf 1 ight After second. bats the photographs Any gaps 2 min. to obtain site during the main outfl ight. The outflight from the mine was sampled nets at the edge of the openings were set at ten sites between 17 May concentrated three, and and 4 November (Fig. 2) for a total 13 November. for a total (Table 1). twice in 1983 by standing flew and catching Ba ts caught and age. pesola to the mine. In of 41 netnights efforts were sites (numbers one, of 56 netnights In addition, between samples on a rock arch over 7 May were obtained which the bats bats in a handnet. in nets were Bats were weighed scale. In 1982, nets 1983, at the three most productive and five) by setting mist iden tified as to species, to the nearest sex, 0.5 g with a 50 g The right forearm of each bat was measured the nearest 1 mm with a metric ruler. identified by metacarpal and the proximal transillumination the right wing and inspection Young-of-the-year of the phalanx joint for presence were between of the fourth to digit the of of a cartilaginous �222 t! ORIENT \ \ GUANO MINE CHUTE LOWER MAIN \ PIT / / / / / / / / m I ~- o o 15 30 50 100 ....• f1 Fig. 2. Locations of net sites 1 through 8 at the Orient Mine. Sites 9 and 10 are at the edge of an open stope that is upslope from site 5. Contour lines are in feet above sea level. Dotted lines denote underground features. �223 Table 1.--Netnights per net site at the Orient part of one night are indicated A. Net Sites (1982 ) as 0.5. May Jun Jul Aug Sep 1 2.0 5.0 LO 2.5 1.0 2 0.5 1.0 Oct Nov Total 11.5 1.5 3 3.0 4 2.0 2.0 4.0 5 0.5 1.0 1.5 1.0 0.5 1.5 3.0 ' 1.0 5.0 6 7 B. Mine between 1.0 3.0 3.5 2.0 1.5 1.0 14.0 8 1.0 1.0 9 0.5 0.5 10 0.5 0.5 Total 2.5 12.5 10.0 10.5 Net Sites (1983) May Jun Jul Aug 1 1.0 2.5 2.5 1.0 3 2.0 7.0 4.0 4.0 5 1.0 4.0 4.0 3.0 Sep 1.5 Oct 1.0 Nov 4.0 13.5 11. 5 a Bats caught with handnet 41.0 Total 2.0 2.0 11.0 .3.0 4.0 2.0 26.0 3.0 4.0 19.0 LOa arch Total 3.0 1.0 8.0 6.0 10.0 4.0 for parts of two outflights. 57.0 �224 zo ne r elongate joints not totally ossified indicated a bat .,.,as your.q, Bats caught bands applied marked with in 1982 were banded to was applied light and In pigment 1983, obtained parts bird bats were u.s. from the in Denver, Colorado. Each sample This material of the forearm and at the base was coded by color and by a unique on the body. The roost was entered site conditions, take inaccessible. observe of of the roost pictures, describe humidity safe vantage direct population However, of the bats on the roost, relative therefore, the to take activity The nearest the roost; estimation was periodically readings temperature. below forearm. paint Service to various of the ears. area right fluorescent Fish and Wildlife location the with white plastic and point was about observation size ambient on the 30 m of young roost was or not possible. Mist nets free-tailed were bats set periodically the mine 20 May set periodically to capture In 1982, nets were from 0.3 to 85.7 km from and 12 August. of guano were obtained cups covered Samples In 1983, from nets were 2.1 to 85.7 km from by confining with aluminum were placed Bats from which free-tailed foil until they in 10% formal in or alcohol. were taken from 18 free-tailed 10% formalin. Valley 22 May and 31 October. bats in styrofoam Stomachs fed or drank. at 29 localities the mine between defecated. as they in the San Luis at 22 localities between Samples set samples bats and preserved were obtained in were �225 caught late at night at the mine. presumably to the roost drank. after Samples Colorado, were Boulder, Much the upper Freeman, produced amount This pile of guano 1982. 16 by 6 In. was measured to the nearest of November guano The of was was a surface inside in the bottom cdvered dimensions with of black of the pile were of guano accumulated on in November of 1982 and December of on the plastic 2.54 cm at 1.5 m intervals, estimated to the nearest was cleared estimate were and the litero In in 19 1 increments on which guano could and to serve as another bats The amount of 1982, the plastic to prepare fed or University by free-tailed 1983: width and depth of guano deposited measured as they for analysis. on 28 June the plastic to J. down a chute and accumulates pit. approximately or in the valley given of the guano the roost falls plastic feeding, as they returned accumulate in 1983 of volume. RESULTS History discovered Railroad of the mine.--Ore bodies at the Orient in 1880. In 1881, built a narrow-gauge km to the northwest, the Orient Mine was an important Fuel known the Colorado C.F.&I. stopped abandoned was leased and developing operations to a private Mine. source Iron Company Coal Denver spur from villa to the Orient Colorado as the Iron the Orient From contractor. Rio Grove, From Gran.de about 13 1881 to 1903, of iron ore for the (C.F.&I.); this and in 19050 and Mine were Company Mine company before was 1892. and eventually 1906 to 1921. the mine In 1922, C.F.&I. resumed �226 development of the Orient Mine and new ore bodies were found. This ore was mined from 1926 through bodies were discovered almost to this ore in 1931; ore was shipped least through the orient 1931. Mine and a sixth Further was closed In 1930. new ore 1930. level tunnel was driven from the mine at mining was anticipated; in 1932 B. Stone. (J. however, in litt. to C.F.&I.) • In the supported late in Buildings the Orient in the store. the canyon that were built Most town Little today mine still during are visible the surface emerge hole indicate area that, where All about midway and exists a in the at Many free-tailed above Mine through bats is located; of to also the large pit. into the stope mining leave up a 60 m wall that has broken and opens little the bat roost mine. of development of these openings in 1890, the was of buildings bats at the Orient of the hillside. roosts. town the mine to the west. from a hole on the hillside the colony Mine library foundations periods an opening 10 m across a of of these buildings this pit is a stope is about The south included Some later the Orient people. just evidence below 19005, hundred of Orient of the free-tailed pit: early Canyon (Fig. 3A). their roost through a large and a town of several located general 1800s This in which are man-made. Maps had been done in the in 1892, tunnels had been dug into the ore body, which was found near the surtace p but the large to C.F.&I.}o excavated pit had not been formed Extensive in the area underground of the present {R. C. Hills, openings roost in litt. had by 1893. been Maps �227 A 8 Fig. 3. Orient Canyon, looking east from the mine (A); main pit of the Orient Mine (B). Both photographs taken in 1983. �228 drawn in 1902 indicate (Fig. 3B) had broken through in two places. One stope in the area of the bat roost sti 11 was being worked 1902 (T. McNamara, open stope that what now is the largeg manager of orient Mine, as of 24 October in 1itt. to J. D. Gi 1chr ist, supervi sor). Description roosted was 1983. of entered Readings (Table stope taken and therefore -20C times Ambient large underground in the third stope. All surfaces rust-colored stope through numerous openings the hillside above stope 1983 and illuminated recessed of ranged bats appear to avoid flying of the mine Light inside enters of light entered the to the the open stope The bats roost inside a 1982 at 1420 hand on the hi 11 side above in a dark, f ree- the stope and did not the light. The pit from which free-tailed of a knife-edged 1983. inside this stope are On 5 August around by from to the pit and one opening most of the area. were the from 20% on 4 June area on the cei 1 ing; howe ver, on 4 September tailed before humidity 1982 and July dust. at 1415 h a beam from the opening in experienced temperature level a fine, the pit. times ceiling conditions bats 1982. with on 4 September the inside the roost ranged The bats roost the and relative to 120C in June 1982 which and seven 30 m below 1983 to about 95% on 14 November covered in in 1982 temperature about roost. humidity stope do not reflect on their in November Relative four of ambient 2) were the bats roost.--The ridge of Harding the sun sets behind bats fly is in the shadow quartzite about the San Juan Mountains 25 min on the west �229 'l'able 2.--Ambient below Date the free-tailed Year temperature and relati~~ humidity bat roost in the Orient Mine. Time (MDST) Ambient temperature Relative humidity (h) (oC) (% ) Bat Activity 13 Jun 1982 1300 12 41 Active 29 Jun 1030 11 35 Active 28 Jul 1715 11 74 Active 14 Nov 1155 -2 95 Quiet 8 May 1983 1000 2 53 Quiet 4 Jun 1807 9 20 Quiet 17 Jun 1000 6 55 Quiet 18 Jul 1500 10 56 Active 28 Jul 1515 12 64 Active 4 Sep 1415 11 63 Active 22 Sep 1500 10 23 Few active �230 side of the valley. the ridge direct as seen sunlight for 3784 Higher depth 1982 and mine in a room roosted in bats, of the 1982 and tunnels found roosting caves the mine of the mine December between of 1983; only!. the genus Myotis at intervals--6436 1 of 1982, the another level stope Although of the where the the pile was presumably here on 5 August is just below were above by!. brasiliensis. level of the mine for evidence Townsend's in two P. townsendii The lower tunnels of Dead were found active two that for presence two and three. workings; townsendii the hole the pit. searched townsendii, were present. to by in early May on another sub-level searched 1 dead bats were "found hanging levels the mine. 19 obtained found on the second the main within were than 28 July were was observed which in from on the third 1983. Several bats, Plecotus outside roost in this room was blackened, of the room, bats were measured On 5 August but none was used regularly big-eared of receive 15 min longer vol ume side no activity and on a sub-level humid the of the pile opposite areas free-tailed south the entrances (as the stope to the hillside Other bats, below was discovered 1982 or on 24 May 1983. on the wall guano 1 for 1983. large and the ceiling by roosting and for about estimates 7527 of bat guano connects of and width pile colony 1 accumulated November. measuring the pit, the sun sets in the year. 1982, increments) from in the evening they do earlier In By 3 September, were and in cool, and caves of of T. brasiliensis and unid~ntified in bats of �231 Seasonal at this chronology.--The roost through varies 1981 colony (Freeman arrives observations When tailed and 3). Records Wunder, in mid-summer, the mine bats kept in press) somewhat were was visited observed arri ve from 1978 indicate later periodically between bats were observed before in March flying No T. brasiliensis evening. free-tai led the than shown by made in 1982 and 1983. bats 12 June (Table time were of first were the caught 17 May and 15 June to be present they out of 1982, mine in mist captured in the set however, on the roost on 16 June Long, dispersed began on 16 June, and the. first serpentine free- nets of 1982: and active nets at the side of the main pit. no on in mist outflights flight occurred on 2 Ju1 y. In 1983, no bats were active on 8 May and 4 June. in mist nets beginning June, roost outflights around on 8 June on 16 June. the had inside Occasional and numbers occurred the stope October. and none density September of bats caught peak numbers and bats were long, active evening and flying in mid-afternoon. of the colony also had departed continued in the net. of the outflight after larger By 18 July, No bats were observed was bats were caught in increasingly quiet. the main part of the colony and free-tailed When the stope was entered again on 17 was Time of departure when the roost was visited to by early decrease In 1982. September. through mid- at the mine on 13 November, In 1983, was reduced outflights varied. the length by the third in the first week week of and of the �232 Table Tadarida Year 3.--Approximate brasiliensis Arrival 1978 Before 17 Jul 1979 After 1980 Between 30 Jun and 3 Jul 1981 Between 26 Jul 1982 Before 1983 Between 8 and 20 Jun 27 Ju1 2 and 12 Jun arrival at the orient and Mine, Departure departure times of 1978-83. Reference Mid- to late Sep Freeman and Wunder, in press Mid- to late Sep Freeman and Wunder, in press Mid- to late Sep Freeman and Wunder, in press Mid- to late Sep Freeman and Wunder, in press Reduced numbers by early Sepi gone by mid-Nov Svoboda, this study Reduced numbers by late Sepi gone by Nov Svoboda, this study �233 month. Free-tailed decreasing nights bats numbers. were caught Mist in November nets were of 1983, leibii) and Plecotus through 27 October set at the mine and small-footed townsendii for two myotis but no free-tailed in (Myotis bats were caught. Dail~ a tunnel over activity.--The connecting a rock arch west pits) early into t~en over a flat, grassy remained bats later scattered from the mine, and some dropped as opposed to flying straight over the valley. 20.0 - 78.4 outflight 1983. bats kmph) flew an average 861 timings for on 25 nights later in the season in the pit outside min before average began. (Fig. 4). the entrance the main of the outflight emerged flight earlier Often during westerly (range, the main 14 October estimation. in relation to sunset a few bats would circle for from 1 to 9 In 36 of 79 observations in 1982 and 1983, one bat of 5 min (range, over the rock arch across to the roost began. in 1982 through the field of view used for the photographic bats of the down the hillside taken from 19 June Free-tailed of 41.9 kmph Bats were timed as they flew over Free-tailed area on the direction directions Free-tailed sections the elevation in a southwesterly emerging two the valley. at about from Most bats flew the pit pit (2877 m) as they flew valley; in the evening divided side of the pit and out over bats emerging the emerged the stope to a large pit. which (upper and lower colony 1 - 20 min) before left the pit an the main flight �234 A ., ; ,. ,. 22 ., ., " ': ". ". -•... - 21 .: : C/) , I 'I :' ., ., ,, ,, . ''.,, ' I, ' I, ' .. .... .~.. ' c :?i 20 a: ::> 0 J: 19 18 JUN JUL SEP AUG OCT NOV 1982 B 22 i I " ,1 -::E , I !i I; 0, :!:" • I::, 1. ... • 21 t- en C , I I I "' , I' " I I I' , •' , " " "II " ., 20 a: :::> 0 :x: 19 l'J 18 JUN JUL AUG SEP OCT NOV 1983 Fig. 4. Times of emergence of free-tailed bats from the Orient Mine relative to sunset in 1982 (A) and 1983 (B). Solid line, serpentine- flight; dashed line, concentrated flight; dotted line, dispersed flight. �235 In 36 observations 1982, free-tailed from 25 min after emerged cloud after from sunset 5 cover 15 June and 15 October an average to 46 min as observed before September: bats emerged before sunset made between of 18 min (range, sunset; the mine. evenings on those SD, 15.5 min) Free-tailed between nights 6 July ranged bats and from 4 25 to 100%. In 43 observations. 1983, bats before emerged 28 August sunset: cloud cover Percent difference cloud and cover, the next night, sunset began began cover, (it was the earliest observed cloud when flights began 14 min after 1982, sunset: the flight began 1 min flight cover before before On 5 July as of that date). of 1982 began was 0% on 5 August, 100% with rain on 6 August, and 25% on 7 August. on nights flights to make a consistant on 5, 6, and 7 August 3 min of each other: Ten 54 min from 0 to 75%. of the outflight. the flight from 5 to 54 min ranged did not appear with 30% cloud Three outflights (range, 24.7 mi n). SD, on those nights cover 20 June and 27 October sunset 6 October in the timing with 80% cloud within 9 min after to 44 m i n af ter sunset: between before made between sunset Cloud in·1983 cover ranged from 0 to 75%. Average length June and 15 October 30.0 min). 20 June SDo Average for 30 outflights 1982 was 59 min (range, length and 27 October 2303 min). observed for 38 outflights 1983 was 50 min b.tween 16 6 - 107 min: SD. observed (range, between 2 - 114 min: �236 Three types serpentine; left of outflights concentrated. and exhibited A less common for over a distance coherence the the valley of existed least during described floor these flights A flights, type of column component lower and Davis of about level and few bats (1966). 1. exiting of the pit. serpentine gaps became In 1982, more began group strayed Free-tailed Gaps began was a concentrated they dropped before edg~ Outflights became 600 m of to appear intermediate more outflights concentrated component to appear in between flight in which flights but a Many bats did not exit from the upper from Most nets were caught during concentrated autumn. bats flew about 5 m above mist nets set also was present. rather, pit southwestern of is the same as the serpentine outflight, and serpentine, in the column: the bats the rock archo most bats left the pit like those in serpentine dispersed the about 7 km from the mine. third dispersed The A high at the edge of the mine at location in the column over at a height 7 km. by Herreid bats in serpentine bats was one in which mine. from the col umn; this pattern flights flights, group coherence. out of the roost, from dispersed; from 1 to 10 m above type of outflight southwestward undulated observed: dispersed directions little flew in a dense column and During the pit in westerly ground were 1 to bats 10 m over captured and dispersed complex in pit into late the in mist flights. summer and were dispersed in mid-June, late June, often after early July. in the outflight: and Beginning four had a on 30 July, of 14 flights �237 between 30 July and 16 October had two or more periods of outflight separated by gaps of from 2 to 25 min. This pattern was more pronounced ·in 1983; flights were dispersed between 20 June and 6 July and more concentrated, often with a serpentine component, between 13 July and 10 October. Between 28 August and 10 October, 13 of 20 outflights had two or more periods of serpentine or concentrated separated by gaps lasting up to 37 min. outflight occurred on 3 October; One such complex it began 38 min before sunset and consisted of the following segments: (3 min); dispersed dispersed serpentine (2 min); gap (23 min); serpentine (8 min); (2 min); gap (1 min); dispersed min): dispersed flights (10 min); serpentine (10 min); and dispersed (4 min): gap (2 (4 min); concentrated (13 min); total, 56 min of flight time in a period of 82 min. Serpentine flights averaged SD, 5.7 min) per night 13 min (range, 6 - 23 min: for 11 nights in 1982 and 18 min (range, 4 - 48 min: SD, 11.8 min) for· 21 nights in 1983. Concentrated flights averaged 41 min (range, 12 - 85 min; SD, 25.4 min) for 14 nights in 1982 and 44 min (range, 10 - 74 min; SD, 17.6 min) for 17 nights in 1983. lasted an average Dispersed flights of 39 min (range, 4 - 107 min; SD, 2604 min) for 27 nights in 1982 and 26 min (range, 2 - 47 min; SD. 14.9 min) for 30 nights in 1983. The return flight began beginning of the outflight. dawn. about 2 hours after the Bats returned sporadically until Most returning bats dove into the upper pit from an altitude higher than the mine, and few were caught in mist �238 nets. At location within 1 hour of 3, most the free-tailed beginning bats were caught of the outflight; occasionally, capture rate would increase between 0330 hand 0600 h. In early summer at location 5, free-tailed bats were caught at a constant rate throughout the night: however Q in July bats were more active at this location between midnight and 0500 h. Just before dawn, many free-tailed bats were mi 11 ing around the opening to the stope; most of these bats were spiralling hillside out of the opening into the upper pit. then diving Activity down the invariably ceased abruptly by 0630 h. Size and Composition--Photoestimations of the colony in the Orient Mine indicated that fewer bats were present in 1983 than in 1982 (Table 4). The peak estimate for 1982 was 154,242, whereas that for 1983 was 107,240. Composition of the colony in 1982 and 1983, based on total catch per year at the mine, were: adult males; 11.1% and 14.4% adult male young-of-the-year; year (Table 5). caught females; 2.1% and 9.3% 0.5% and 3.6% female young-of-the- Adult males comprised 88.6% of all adults captured in 1982 and 83.5% in 1983. were 86.3% and 72.6% in August, Young free-tailed September, and October when bats the proportion of adult females caught in the initial outflight increased (Figs. 5 and 6). Increased proportions of adult females and young in the outflight (Table 6) occurred in 1982 at about the same time as an overall reduction in colony size and a more dispersed flight; however, in 1983, females and young were present at the time of peak numbers in late August. �239 Table 4.--Photographic tailed bats in evening estimation flights of numbers from the Orient Mine, 1982 1982-83. 1983 Date Number of bats 19 Jun 11,396 24 Jun 17,542 30 Jun 26,442 9 Jul 88,771 21 Jul 154,242 5 Sep of free- 14,280 Date Number of bats 2 Jul 5,525 23 Jul 37,934 28 Ju1 24,130 12 Aug 46,541 21 Aug 75,742 28 Aug 34,114 4 Sep 107,240 �240 Tab~e brasiliensis 5.--£~~£~~!!!~~ ~!~a~£!!~ ~!Tadarida from the orient month. Mine, 1982-83, ~ Adult Month Year Jun Jul Aug Sep oct Total Male Young ·Female Male Female Total 1982 63 1 64 1983 118 6 124 1982 151 1 152 1983 601 5 606 1982 348 9 1 1 359 1983 549 127 175 74 925 1982 126 57 15 2 200 1983 401 171 50 .13 635 1982 19 23 ·1 1 44 1983 98 41 2 1 142 1982 707 91 17 4 819 1983 1767 350 227 88 2432 �241 150 · 100 · 50 · en le( III u, 0 II W III ~ :> z - .I. JUN • I I JUL AUG • • SEP • Ff . OCT 1982 Fig. 5. Composition of capture samples of Tadarida brasiliensis at the Orient Mine, 1982. Solid bar, adult males; open bar, adult females; dotted bar, young-of-the-year. Sampling dates are given in Table 6. �242 200 · 150 · 100 · 50 · en ~ 4( m u, 0 a:: w m ~ :::> z II JUN ~ I I JUL I I. I ~~ AUG I SEP I • OCT 1983 Fig. 6. Composition of capture samples of Tadarida brasiliensis at the Orient Mine, 1983. Solid bar, adult males; open bar, adult females; dotted bar, young-of-the-year. Sampling dates are given in Table 7. ~ �243 Table 6.--Summary at the orient Mine, of captures of Tadarida 1982 • .Adult Date 16 Jun 17 21 22 25 29 1 Jul 5 10 11 18 23 28 5 Aug 6 7 10 16 18 20 4 Sep 25 15 Oct 16 Total Location Male 3 2 1 3 10 4 43 20 27 14 1 69 19 1 33 1 46 102 5 109 52 36 90 1 18 707 1 1 5 4 1 s 3 3 1 &. 8 3 7 10 1 &. 1 3 3 &. 3 5 3 &. 3 3 3 3 3 7 7 4 4 brasiliensis Young Female Male Female Total 2 1 3 10 4 1 44 20 27 14 1 70 19 1 33 1 47 105 5 115 53 60 140 3 41 819 1 1 2 5 1 14 43 2 21 91 .1 1 9 6 1 1 1 17 4 1 �244 and early period September. In 1983, of peak numbers September this influx of young and occurred between (Table 7): in 1982, no observations the mine during that period. September 1983, adult young were detected. difficult By 4 September (N 164) were made at 1982 and 14 In autumn, young are fully grown and to id~ntify = and 3 females were present but only a few based on degree of ossification finger joints: 73% of young females (N males 20 August = of 63) and 78% of young that were caught between 22 August and 3 October 1983 had cartilaginous zones on only one side of the finger joints. Bleaching of fur on free-tailed bats at the Orient Mine was not common. Twelve bats caught between 27 June and 3 October 1983 had bleached spots: five bats had spots on the head and neck: four bats had sp6ts on the shoulders: two bats had scattered spots on the back: one bat had a spot on its rump. Mean weights for free-tailed bats captured at the Orient Mine are as follows: adult males weighed 11.6 9 (range, 9.0 - 16.0 g: SD, 1.17 g) in 1982 (N 8.5 - 15.•0 g: SD, 1.09 g) in 1983 (N weighed 11.4 g (range, 8.5 - 90) and 11. 1 g (range, 8 •5 350): young males weighed 0.95 g) in 1982 (N 0.75 = g) in 1983 = (N 637) = and 11.4 9 (range 1765): Q adult females 15•5 g: SD , 1.17 g) in 198 3 (N = 9.3 g (range, 8.0 - 11.5 g: SD, 16) and 9.6 9 (range, 8.0 - = 227): = 15.5 g: SD, 1.40 g) in 1982 (N young females (range, 805 - 9.5 g: SD, 0.41 g) in 1982 (N 11.5 g; SD Q weighed :::I 9.0 9 4) and 9.7 9 (range, 8.0 - 12.0 g: SD, 0.82 g) in 1983 (N = 88). �245 Table at the --- 7.--Summary Orient Mine, of - captures 1983. Young Adult Date 8 Jun 10 16 20 23 25 27 30 3 Jul 6 9 13 15 16 17 18 22 23 25 29 8 Aug 12 13 15 20 22 27 29 3 Sep 14 16 19 21 25 1 Oct 3 6 10 15 17 26 27 Total Location Male 3 5 5 1 1 29 19 15 3 5 1 3· 5 3 1 5 3 5 1 arch arch s 3 5 1 3 5 3 1 5 1 & 3 5 3 5 3 5 3 5 3 5 3 5 3 5 3 5 1 & 3 5 1 & 3 brasiliensis of Tadarida Female Male Female 1 1 29 20 17 1 46 9 44 10 140 89 24 3 3 76 33 1 2 1 46 6 44 10 136 89 24 3 4 3 3 76 33 12 59 112 17 2 174 82 92 49 95 38 122 45 106 45 46 37 4 14 18 7 no catch Total l2 59 1 1 6 11 32 20 43 14 43 29 59 15 15 10 3 2 9 1 48 19 1 6 1767 7 350 4 59 57 24 31 45 2 2 23 28 9 12 13 1 2 ·2 227 1 88 113 17 3 180 99 206 154 171 95 223 76 165 61 63 47 7 19 27 8 67 1 13 2432 �246 Eight other (Table 8). species of bats were caught at the mine ~!~cot~ townsendii, Myotis leibii, and 10ng- legged myotis (Myotis ~~!ans) were the most common species other than free-tailed years, P. townsendii November ~~!~ bats. Combining was caught records from both between June and early and M. leibii between July and early November. and biy brown bats between June and August. (Eptesicus M. fuscus) were caught Male hoary bats (Lasiurus cinereua) appeared between late June and mid-July. A little brown bat (Myotis lucifugus) was caught in June of 1982, and a longeared myot is (Myot is evo t rs) ~as caught in August of 1982. Silver-haired bats (Lasionycteris the sample noctivagans) appeared in late August and October of 1983. in When the roost was occupied by the colony, species of bats other than T. brasiliensis were caught after the main flight of free- tailed bats had ended. Most P. townsendii (21 of 29) were caught after midnight at the mine: of the remainder, two were caught before 1900 h in late October and early November 1983, and six were caught between 2030 hand 2400 h at other About hal·f of the M. leibii midnight, were Six E. fus~ captured (11 of 20) were caught after ~. volans was captured after 2245 were captu~ed before midnight, after midnight. after midnight. and four Three of five L. cinereus appeared between 2245 h and about 2315 h. was caught in the summer. seven were caught from 2200 h to 2400 h, and two were caught before 2000 h. h. times The one M. evotis ~. noctivagans about 2130 h and just before dawn. was caught at �247 bats Table brasiliensis caught at the Orient other Mine, than Tadarida 1982-83. Adult Species Year lucifugus 1982 2 2 M. evotis 1982 1 1 M. volans 1982 4 1 5 1983 9 4 13 1982 3 1 4 1983 14 2 16 1983 2 1 3 1982 1 1983 8 1982 3 3 1983 2 2 1982 12 12 1983 14 3 17 75 13 88 Myotis M. 1eibii Lasionycteris Eptesicus Lasiurus P1ecotus Total noctiva~ans fuscus cinereus townsendii 1982-83 Male Female Total 1 1 9 �248 Violet-green whi te-throated swallows swi fts (Tachycineta (Aeronautes the mine and nested in crevices Birds usually less their outflight. were and free-tailed swallow wer'e great would predators horned observed owls regurgi tated pellets Mine, the mine. Only contained remains July: bat was Sometimes an owl 11 beneath a great 12 pellets from other horned from the pit sections on the forearm the only that of a captured freeon 28 of a banded bat. bones contained bird was collected re~overy contained of all taken from the pit, One of the white-plastic in a pellet found on 4 September at the mine 18 May and 4 September, collected of bats. pel 1 et found on 14 August of one bat I A and a the remains of at least bats. Reproduction.--A the Orient Mine. few females superficially netted might be bear ing young Of 216 free-tailed 1982, two were female: appeared bats swallows of three of the pellets, this constitutes pellet began the outflight and four pellets found bats before of the mine, that had been placed tailed July catching Between were nest 8 km southwest bands at 1983, when a violet-green enter the roost in the evening at the Orient the bats between (Bubo virginianus). bats began. eight contact common with the first bat out of the roost. free-tailed owl direct and of the main pit. by the time bats was on 18 July collided The only saxatal is) were in the wall active The only thalassina) in June bats caught one female to be pregnant. and July 1983, at in June and caught on 29 June 1982 of 730 free-tailed 11 were female. On 23 �249 June 1983 two females with a crown-rump were caught, locations growing caught in the San Luis Valley near water. Adult at 12 locations females four young males young bats were caught females male T. brasiliensis (N = 75) were from 2 to 40 km from the mine were caught (Table 3 and 45 km from the mine: were caught 2 km from the mine. Mist nets were from 2 to 40 km from the mine where other. than T. brasiliensis caught were among trees were caught 2 and 33 km from the mine; three set at 11 locations bats in mist nets set (Fig. 7); 11 of these were over water, and two locations 9); two adult an embryo length of 26 mm. Foraging.--Free-tailed at 13 locations one containing were caught. in mist nets set at 16 locations No bats from less only were than 1 to 86 km from the mine. DISCUSSION Colonization.--Free-tailed they are fast, long-distance al., 1973): variety bats fliers they are flexible of man-made al., 1962), cracks structures summer present Slope, 1910). (Glass, in Colorado, in addition along a tendency 1982). the Colorado fuscus behind If free-tailed the San Luis Valley the shutters bats also before to caves (Davis et 1955), and hollow to scatter as they River bats found roosting used buildings were on the West of a building development et and use a Free~tailed at least by 1907; these bats were Eptesicus selection and crev ices (Krutzsch, roosts colonizers: (Glass, 1982: Williams in roost trees (Lowery, 1974); they have leave are good with (Warren. as roosts of the Orient in Mine, �250 ,",onch. P ••• SAN LUIS BASIN 20 K", LIlV.tll. COLORADO . NEW ~ -----~ MEXICO < Pails Cb --- ~- ~ ~~ ~ 6(~ ~~ Fig. 7. Localities at which mist nets were set in the San Luis Valley, 1982-83. Black stars indicate localities at which Tadarida brasiliensis was netted; circles indicate localities at which T. brasiliensis was not netted; white star indicates location of the Orient Mine. �'Table 9.--Tadarida brasiliensis caught away from the Orient Mine, 1982-83. Location Type hot spring Distance from mine (km) Direction from mine 2 179 (0 ) pond 14 282 trees pond trees 10 26 33 244 163 207 Date 15 21 20 23 1 18 22 27 31 Ju1 Ju1 Ju1 Ju1 Aug Aug Ju1 Ju1 Aug 1982 1982 1983 1983 1982 1983 1982 1982 1983 Age Sex No. A A A A M 14 10 1 y y M F M A A A A A y pond pond pond creek pond lake tank lake Total 6 8 3 168 255 200 24 Ju1 1983 15 Ju1 1982 16 Aug 1983 8 4 13 40 45 280 167 230 225 182 31 2 17 24 1 Ju1 Aug Aug Aug Sep 1982 1982 1983 1983 1983 A A A A A A A A A M M M M M M M M M M M F M M M M F 3 3 3 10 2 10 . 7 6 1 4 3 1 1 1 1 1 1 1 Mean weight (g) 13.1 13.7 11. 0 10.8 10.3 10.3 13.7 10.5 11. 4 13.3 10.8 10.0 12.8 1203 11.0 12.5 14.0 11. 0 11. 5 11. 0 12.5 Range of weight (g) (12.0-15.5) (12.5-15.0) (10.5-11.5) (l0. 0-10.5) (10.0-10.5) (12.5-16.0) (11.0-12.0) .(12.5-15.0) (9.5-11.5) (12.0-13.5) (11.5-13.0) Time of capture (h) 2200-0430 0150-0405 0400 2400-0545 2400-0230 0130-0230 2325-2425 2300-0330 2045-2245 2150-0155 2400-0600 2400-0600 0300-0500 0300-0500 2130 2400-0230 2415 0200-0400 2200 2125 2300-0200 83 N \..r1 f-' �252 the bats would have been present to exploit the man-made roost in the mine. Discovery of the mine by free-tailed bats might have occurred during normal foraging flights or during dispersal in spring or autumn. After it was found, the Orient Mine probably became a permanent roost. Odor has been suggested as a .eans by which migrating free-tailed bats locate summer roosts (Glass, 1982). However, the odor of free~tailed bats at the Orient Mine did not seem strong until the colony arrived in June. The large size of the main pit on the open hillside potentially serves as a visual cue for bats trying to locate the mine. how colonization Exactly was initiated, what bonds are present to maintain the integrity of the colony, and what cues are used by migrating bats to locate the roost site each summer are unknown. Roost selection.--The Orient Mine is unique among mines in the San Luis Valley and perhaps in Colorado. No other mines that were explored had such large entrances, cavernous rooms, ~nd proximity to a large area of open country; other mines usually valleys were located in coniferous forests in narrow at elevations higher than the Orient Mine, and had small entrances and narrow tunnels. Structural features of the orient Mine are similar to those described for caves used as maternity roosts in central Texas (Davis et al., 1962: Eads et al., 1957). The main difference between caves used as maternity roosts and the Orient Mine is that the mine is cooler during summer. Relatively low temperatures in the Orient Mine �253 possibly are responsible for the paucity of ectoparasites on free-tailed bats at the mine and lack of dermestid beetles in the guano below the roost. Ambient temperatures of maternity roosts in Arizona (Constantine, (Cagle, (Constantine, 1958L Oklahoma 1950; Constantine, between June and September. 1958), New Mexico (Twente, 1956), and Texas 1958) range from 180 to 340C Ambient temperatures in summer in the Orient Min. are similar to those at Eagle Creek Cave, Arizona, Ok L ahoma in April g (Constantine, in spring (Twente g 1958) and Merrihew Cave. 1956). The cooler temperatures in the Orient Mine might retard development of young (Tuttle~ 19750 1976) and thereby minimize.suitability of the mine for use as a maternity roost. Seasonal chronology.--Free-tailed bats arrive at the Orient Mine about four months later than they arrive in Texas (Davis et al., 1962; Eads et al., 1955), three months later than they arrive Cockrum, Oklahoma in Arizona (O'Shea, 1976; 1962), and two months (Glass, (Constantineo 1958; 1967). Villa-R. and later than they arrive Twente, 1956) and New in Mexico An increase in number of adult males occurs in June at Carlsbad Caverns, New Mexico (Constantineo 1967); at the same time, groups of adult males disappear from roosts under bridges near Carlsbad Caverns, and predominantly male groups appear at "higher elevations" (elevations not reported) (Constantine, 1967), possibly corresponding to the elevation of the Orient Mine. At the Orient Mine in 1982 and 1983, free-tailed had dispersed outflights early in the summer. bats Serpentine �254 flights began in early July and might have corresponded to either a sudden influx of new bats or a change in flight behavior of bats already present in response to some factor such as weather. A substantial increase in population at the Orient Mine was noted (Freeman and Wundero between 30 June in press). and 3 July Sudden arrivals 1980 of large groups have been reported for a cave near Carbo. Sonora. in late March (Cockrum, 1969) and for Eagle Creek Cave in early June (Cockrum, 1969). pattern was observed uncertain whether in early July of 1982 and 1983. it is this was accompanied increase in numbers. was noted dispersed large in late A gradual June, and difficult colonies of (Constantine, 1967), 1956L Although a definite change of flight and central when by a substantial increase in ,number of bats outf1ights were, long to estimate by photography. free-tailed .no rt hern Texas bats in New and Many Mexico Oklahoma (Glass, 1958: Twente, (Davis et al., 1962) increase gradually in size rather than suddenly. The colony at the Orient Mine reaches a maximum size of about 100,000 bats and is 98% male until mid-August. formation of such a large male summer The roost is unusual. Cockrum (1969) reported that male summer roosts in Arizona usually consist of 10 to 300 bats: the largest male roost described in his study consisted of 1300 bats. It also is unusual that a large colony of males would congregate far to the north of wintering grounds in Mexico and adjacent areas of the United States. Davis et ale (1962) thought that most males remain in Mexico after breeding because few are present �255 at colonies males in Texas are known and Cockrum, from to the transients, 1967: fronts roosts 15 September. from through observed southern 1962; Eads residents south of et a L, , 1957: departure 1967: Davis and of bats et al •• 1962). at the Orient and 16 September. per samples from 1 to 67 bats per night. Mine, therefore, of their summer is later, whereas roost have than departure a do is to the south. the Orient Similar roosts activity capture night; Mine, free-tailed in late morning and afternoon et al •• 1957)0 California in in caves and buildings 76 to 223 bats Arrival September. rhythm of diurnal of Mine in the in size of the colony of occupancy in maternity 1962: Eads arrival bats at the Orient and vocal departure to trigger 13 August Dai!.l acti~itl.--At June (Villa-R. The onset of freezing weather to that at colonies were active summer of at the same time as the first frost on ranged to the south. similar with (Constantine, had ranged period with et al., 1974). Between Free-tailed bats Davis decrease 16 September shorter colonies from the Orient from the north, in 1983 occurred samples and has been suggested The first marked after south and LaVal, from summer Mine small during bats coincides presumably (Constantine, cold in Mexico of free-tailed and October caves Spenrath to remain indeed, 1962). Departure September in summer; daytime in central Krutzsch Texas from late has been (Davis et al., (1955) thought of free-tailed was influenced activity bats that the bats in an attic in by temperature; on warm �256 days, bats afternoon were active and aroused Events observed, pit in to full the roost but outside before the in the morning main or merely pre-flight to the pit, prior flight light sampling (1966). activity to emergence began. This as described restlessness light sampling bats to know what conditions Free-tailed foggy evenings Mine might bats have been observed 1956). in the San Luis during Valley the study period; rarely clouds on the west side of the valley across the valley began near modified which to emerge passed. Allison Carlsbad Caverns to emerge later is not known whether Duration over storms that traveled generally consistantly circled when The the mountains q evening from overcast was thunderstorms during back into the the that thermal triggered the evening storm had currents in flight: it this is a factor at the Orient Mine. of the outflight 2 to 114 min in 1982 and 1983. the Orient cover. Outflights factor (1937) suggested at sunset emergence was completely but quickly in the on cloudy or (Herreid and Davis" or localized only for the roosto of cloud developed the time of outflights bats began roost The by Herreid and is near openings However, and then dissipated. sunset. (1955, 1956) earlier the orient Mine seemed to be independent sky could not be necessary exist outside 1955; Twente, not the upper behavior as described than when the sky is clear 1966: Krutzsch, were by Twente The roost in the Orient and in late just before sunset. the roost a few bats entered represent Davis and quiescent Mine was similar at the Orient Mine Total ranged from time of outflights to that (17 - 99 min) observed at at �257 Vickery I Cave, Oklahoma photoestimations outflights, I Cave flight Texas speed 42 kmph. present in the from Vickery from the Orient speed of bats leaving at 40 kmph (range. et al., 1973), Bracken 7 - 105 kmph: that of bats departing '. I Cave kmph) (Humphrey, maternity was estimated colonies for the beginning in central later, Mine, 600 m over the valley later in flew the immediately altitude bats floor straight evening after from Outflight similar drop the southwest. restricted ground that the 1962)0 011 •• Many lower The of about bats flying elevations average maximum Bracken Cave was 2300 level, observed reported or about 800 - does colonies in the over the valley and are to th~ east of the mine, of Mine are that bats at the Orient not seem number at the Orient for other fly in one direction cover to 56 kmph (Wi11 iams et al., 1973). by steep mountains cloud outflight, pit. The major differences generally to leaving characteristics to those to range at an altitude the mine. would leaving sea level) outflights (Davis et flew from large (or about 2880 m above sea level) 600 - 3100 m above 3JOO m above (range, 24 - 46 was estimated dispersed reached by bat flights m (range, percent Texas of the outflight At the Orient they at 38 kmph and that of bats departing 1971), from 32 kmph during Mine Average (Williams from Vickery were bats of bats departing was estimated 25 kmph) when of based on than from the orient Mine when at peak numberso Mine averaged SD, - numbers However. about six times as many bats emerged Average Cave, of (Humphrey, 1971). to make bats a difference exiting during that in the �258 outflight is less outflight requires bats than at other a similar do not spiral upward colonies period even of time, to gain altitude though and the that the for the foraging flight. Size of co!ony.--Outflights northern Oklahoma (Glass, 1958). reach maximum Although nursery colony, young in samples early July with the Orient Mine the appearance in is not a typical six weeks 1961). then young would in late August of of adult females and If young are born in taken with mist nets. and it takes colonies size when young become. volant peak nu~bers there occurred 1983 and coincided (Short, at maternity for them to become be in the outflight volant by about mid-August. Using photography had several which sources they could across the became darkness: dispersed and did access the roost to possible population constant method bats to observe size over was the magnitude outflight outflight in timing of bats as they flew with that later more emerged fly limited in front and of were the therefore estimated; the stream photoestimation camera: it was not the probably roost probably numbers are of bats period and gives indicated only of bats caught better was not (2 min). underestimates of the number of bats at the mine. and from if all bats left the roost each time the of estimation present two openings difficult always was had increasingly bats not numbers at the Orient Mine bats of error: exit the roost: frame increasing to estimate This the number of an of order Periods of peak by in mist length nets. of Peak �25' numbers in capture September early of 1983: September samples occurred periodic in mid-August observations in late of 1982 were not conducive ant eaLly August and to determinations of the period of peak population. The 1968 colony and actually conservative 1968 was estimated (Meacham in press); to contain These Composition.--On nets, the predominantly of the colony 100.000 that, if young are born More difficult outflight 1982 (N early = both years, females nets summer use the Orient and autumn observation of taken with feature in mid-August 8 June was adult Mine early until in July, mid-August. in the 16 June and 20 August and 13 August males; 1983 after mid-August in samples. southward. = (N in One explanation during as a transient as they migrate af It is logical is the lack of adult females Between was in striking Mine is that females bear their young elsewhere merely (Freeman is an atypical. The most in mist were common the and Wunde:co samples Mine in the Orient between 930), 98% of the sample 1978 among observers. appearance in the summer. 575) and (Freeman in the outflight. to explain of bats on casual roost. not be captured in August of 50,000 at the Orient and young A of the basis of capture females would bats sudden s Lz e , in in August which can vary greatly the since of 1980, the outf1ight are based adult they not declined increasing in August summer was has to consist colony male be 1971); g estimates the main outflight. Mine of the size of the colony was estimated and Wunder, mist might estimate 9,000 outflight press). at the Orient summer, and roost in late Substantiation �260 of this explanation will require colony near the Orient Mine. during location of a maternity No other colonies were found this study, but many mines and buildings were not searched. Another explanation for the increase of females in the samples is that, before mid-August, the sampling method was biased against females~ Pregnant or lactating females might employ an exit or flight pattern that was not sampled with mist nets early in the summer. The mine was searched for other roost locations, but only one abandoned roost which was in the same Serpentine stope flights as the active consistantly roost avoided was the located. mist nets; therefore, two serpentine flights were sampled with a handnet in mid-July of 1983 to ascertain if females were emerging in these flights. Humphrey 50). = All bats in these samples were males (N (1971) reported that composition of samples changed at Vickery I Cave as the outflight progressed; adult females emerged earlier than adult males, and lactating females emerged earlier than pregnant females. Another feature of the composition of the colony was the high proportion of young males females: the ratio was 4.2:1 315) in Other 1983. proportionately (N caught relative = more young in 1982 and 2.6:1 21) workers males also (1.23:1 (1974) found a significant reported a 1:1 sex ratio (N = reported females in but only Pagels and (p < young males to young females). however. have than young samples (Cagle, 1950; Constantine, 1967), Jones to young 0.05) difference Davis et ale (1962), for volant young. �261 Explanations the Orient female for the skewed Mine young: females: southward roost. than young is nearer young females were is limited. were captured, crown-rump 23 June length of females fly lower to the ~round caught in mist nets. by these data. evidence On 3 July of reproduction 1980 four pregnant of which two contained females as a transient rates were caught during it was near of fetuses at or just before 28 mm in southern California this term. caught Crown-rump term in other studies (Krutzsch, thus free-tailed bats Mine or in a different or early does (1957) The on lengths are 29 and 1955), from 23 to 27 F 1 0 rid a 1937). Female Orient Only study. mm in T e xa s (Da vis eta 1 ., 1962 ), and abo u t 25m min (Sherman, females in press). (26 mm) of the fetus in a female 1983 indicated in the fetuses with crown-rump of 16 and 19 mm (Freeman and Wunder, two pregnant males sex ratio of young at birth probably Reprodu6tion.--Direct lengths that growth to 1:1 than is indicated Orient Mine mortality than young Mine than in mist nets a higher females and are more easily the actual male in females to catch the Orient found quickly have more young using bear more more or that young males females Nevertheless. ossify are easier without differ females males males: No reports and males joints young rate than young fly include: finger than in males: than young might sex ratio found in young bats at July. not Approximate differ for Texas, from Glass in Colorado, roost, either bear young time of parturition dates (1958) reported by for Oklahoma, at the in late June in Colorado Eads or et ala Krutzsch �262 (1955) for southern California. a period of peak parturition to 19 June) reported in Texas than that parturition about one month earlier Foraging.--No of free-tailed disperse over brasiliensis km from the Orient bats were Orient feed from Mine chaparral meters California of in Florida where groups large of vicinity flying as far as 40 Free-tailed flew quickly of the mine: that these bats over the valley within feeding at the and did not several times 10 m of the ground over oak or drinking. bats fed over T. individuals. it is uncertain Mine. numbers described 10-13 in Nevada to 25 m above the ground. from is bats seemed to (1967) Ross chaparral Simmons et ale insects Free-tailed bats in and streams 1955). most foraging of less than 200 m above floor capturing hillsides 5 to 10 m (Krutzsch, et al. (1973) observed at heights In Texas, flight behavior the ground where insects common. Free-tailed Vaughan, implies al., bats free-tailed Mine; however, free-tailed usually at heights were in not observed (1978) observed Williams (1937) and water sources on the margin of the valley but were several and Sherman groups of bats were caught in the immediate bats were observed located Rather, the Orient usually was valley. or small (8 June than in Colorado. feeding Individuals in Colorado, for free-tailed fed. the earlier that is slightly one area bats et ale (1962) reported Davis 1970; bats are fast fliers Williams more direct 1972). Ross et al., (Findley 1973), and less maneuverable (1967) suggested that and et al., 1972: greater flight speed (Findley et free-tailed bats �263 forage on swarms of insects as a "filter feeder." Foraging in this manner might be enhan6ed by the wrinkled free-tailed (1966) thought expanded bats, and used which as Vaughan funnels somewhat like lips of could the bristles around the mouths of caprimulgiform birds. Simmons T. et ale (1978) observed __ __brasiliensis . flying slowly and with great control. more flexible than other species be rictal Howevero in Nevada Free-tailed bats are in modification of echolocation pulses to accommodate cluttered and uncluttered situations exploit (Simmons et al., 1978) and probably a variety are able to of 'feeding areas. One locality valley at which a free-tailed alkali flat bordering bat was captured a large. shallow around the lake was less than 1 m tall. lake: in the was on an vegetation The locality at which most bats were captured away from the mine was under a canopy of trees at a hot springs. The free-tailed predators bats at the Orient Mine are important on night-flying insects in the San Luis Valley. Guano and stomach samples collected in this study presently are being Colorado, analyzed Boulder. by J. Freeman Free-tailed at the University bats reportedly of take prey from 2 to 25 mm in length: in 88 bats, Ross (1967) found 34% Lepidoptera (moths), 26.2% Hymenoptera Coleoptera (scarabs (leafhoppers), Neuroptera brasiliensis and chrysomelids), 6.4% Hemiptera (ant1 ions). (flying ants), 16.8% Bailey (true (1931) 15% Homoptera bugs), reported and 1.6% that To eats 95% moths and 5% mixed insects; Storer (1926) reported more than 90% moth material: Sherman (1939) �264 noted that six of eight free-tailed eaten Lepidoptera, HYmenoptera, six contained four cbntained Diptera, Coleoptera, Odona ta, and had six contained two contained Homoptera, one Neuroptera. Freeman (1981) found Lepidoptera in five fecal samples. con ta ined bats he examined one con ta ined When bats numbered more than 25,000,000 at Eagle Creek Cave, they consumed more than 36 metric tons of insects nightly (Cockrum, 1970). By comparison, a colony of 100,000 bats, such as that irtthe Orient Mine, would eat over 0.14 metric tons of insects per night. At least 3800 1 (approximately 13 kg) of guano were deposited beneath the roost in the Orient Mine in each of the two years of the study. This amount does not approach the size of deposits of guano in the large maternity colonieso In the 1950s at Bracken Cave, more than 54 metric tons of guano were mined each year in December and January (Eads et al., 1957). Guano deposits in Carlsbad Caverns between 1901 and 1921 were enormous; from September through March more than 36 metric tons were mined each day (Bailey, 1928). Cockrum (1970) was causing a decline Creek. Samples roosts) collected J. Freeman ppm. concerned that pest icides in numbers of free-tailed of guano were bats at Eagle (one fresh and two from below from the Orient Mine on 19 August 1980 by contained DDE residues of 0.48, 0.33, and 0.31 These levels are similar to those in guano from Eagle Creek Cave and lower than those in guano from Bracken Cave (Clark et al., 1975). guano samples Clark et ale (1975) reported that 15 taken at core depths ranging from 5 to 76 cm �265 from Eagle Creek Cave and analyzed by Reidinger contained from 0.22 to 0.42 ppm DDE, with amounts inversely with depth. Two surface samples (1972) varying of guano from Bracken Cave contained more DDE (0.61 and 0.58 ppm) and also DDT (0.14 and 0.12 ppm). The colony at the Orient Mine does not seem to be declining, and it is difficult with so few data to understand what trends are occurring in reproductive success. The level of pesticide residue in bats at the Orient Mine likely does not pose a threat to the colony at this time. �266 LITERATURE v. Allison, C. Caverns. Altenbach, Evening 1937. J. S., K. N. Geluso, size Caverns in Trans. Ser., Nat 1. and Park Mountains D. M. 1972.. Williams and Wilkins 1931. eds.). Proc. Common 1969. names of a Kansas State Univ. Agric. Exp. Distribution of mammals in Colorado. Mus. Nat. Hist., 3:1-415. Animal 1928. Park, 117:1-62. Univ. Kansas V. National Ser v., 4: 1-442. list of plants. Sta. Tech. Bull., 1979. Biological R. J. Baker, K. L., and C. E. Owensby. Monogr., Carlsbad at Carlsbad in 341-348, in the Guadalupe (H. H. Genoways Armstrong, from brasiliensis Pp. Texas selected flight and D. E. Wilson. of Tadarida 1973. investigations Anderson, bat J. Marnm., 18:80-82. Population Bailey, CITED life of the Co., Baltimore, Mammal s of New Mexico. Carlsbad Cavern. 195 pp. N. Amer. Fauna, 53: 1- 412. Barbour, R. W., and W. H. Da~is. Univ. Press Kentucky, Cagle, F. R. mexicana. Car y, M. 191 10 Fauna, Clark, 1950. J. Lexington, A Texas Mamm., 1969. Bats of America. 286 pp. colony of bats, Tadarida 31:400-402. A b i0 log ical sur v ey 0f Color ado. N. Am e r 33:1-256. D. R., Jr., C. O. Martin, Organochlorine insecticide and D. M. Swineford. residues in the 1975. free-tailed 0 �267 bat (Tadarida brasiliensis) at Bracken Cave, Texas. J. Mamm., 56:429-443. Cockrum, E. L. 1969. brasiliensis. Migration in the guano bat, Tadarida Misc. Publ. Mus. Nat. Hist., Univ. Kansas, 51:303-336. ------ Insecticides 1970. and guano bats. Ecology, 5lg76l-762. Constantine, D. G. 1958. by the atmosphere ------ 1962. Rabies Heal th Rept., 1967. bat. in caves. Public D. G., E. patterns 39:513-520. by nonbite route. Health s, of the Mexican Publ. BioI., Tierkel, Rabies 1968. Public free-tailed 7:1-79. M. D. Kleckner, in. New Mexico and D. M. cavern bats. Rept., 83:303-316. R. B., C. F. Herreid Mexican Mamm., J. in bats 77:287-289. Activity Hawkins. of hair pigment transmission Univ. New Mexico Constantine, Davis, Bleaching free-tailed II, bats and H. L. Short. in Texas. Ecol. 1962. Monogr., 32:311-346. Eads, R. B., Banding Mexican 1957. tailed S. Wiseman, J. and free-tailed Observations bat, Tadarida G. bats. Menzies. 1955. J. Mamm •• 36gl20-12l. concerning mexicana, C. the Mexican in Texas~ free- Texas J. Sci., 9:227-242. Ellinwood, s, Colorado. R. 1978. Unpub1. Gree 1ey, 77 pp. A survey of bats in M.S. thesis, Univ. Northern southeast Colorado. �268 Epis, R. C., G. R. Scott, R. B. Taylor, 1976. Cenozoic volcanic, tectonic, features of central Colorado. in Colorado eds.). field geology Prof. Co n t r i b ,, and C. E. Chapin. (R. and geomorphic Pp , 323-338, in Studies C. Epis and colorado a. J. Weimer, School of Mines, 8:1- 552. Farney, J. P., and J. K. Jones, Jr. records of bats from Nebraska. Findley, J. S., E. H. 1975. Noteworthy Mammalia, 39:327-330. s t ud Ier , and D. E. Wilson. Morphologic properties of bat wings. 1972. J. Mamm., 53:429- 444. Freeman, J., and L. Wunder; colony of the in press. Brazilian Observations free-tailed brasiliensis in bat southern at a (Tadarida Colorado. Southwestern Nat. Freeman, P. W. 1981. A multivariate study of the family Molossidae (Mammalia, Chiroptera): morphology, ecology, evolution. Fieldiana-Zool., Chicago Mus. Nat. Hist .• New Ser., 7~1-173 Geluso, K. N., J. S. Altenbach, Organochlorine and D. E. Wilson. residues in young Mexican bats from several roosts. 1981. free-tailed Amer. Midland Nat., 105:249- 257. Glass, B. P. 1958. in Oklahoma. ------ ·0 1982. Returns of Mexican freetail bats banded J. Mamm., 39:435-437. Seasonal movements of Mexican freetail bats Tadarida brasiliensis mexicana banded in the Great Plains. Southwestern Nat., 27:127-133. �269 Hall, E. R. 19810 John Wiley Harr ington, and Sons, H, Do Second ed. Herreid, The mammals of North 19640 of bats. J. the Cristo the R. 1960. geology E. W. Bull.. I R. A. Stuart T. Arizona Mine. Denver, state to Haun, Do 310 pp. Louisiana Univ. de in Guide Weimer and J. Geo L, , free- Sangre 129-131. The mammals of J. Entry. and Press. its Baton the Bats Bat Res. News, 12:37. in relation destruction of to the insects production .• u.S. of Dept. 1395: 1-12. 1969. King, 1976. Brazilian Handbook Publ., Fat of Tucson, content free-tailed bra s i 1 i ens i s (Mol 0 s s i da e ) • 326. Mexican Mamm., 36:236-242. Orient Pp , Louisiana 1926. Agr ic. o Shea, J. (R. J. Tadarida 86:220-2230 on the near Assoc. of population bat, Nat., Colorado. 1974. 1971. and Dale patterns 565 pp. guano Nelson, Geo16gy waters. Meacham, J. W. Nelson, Flight free-tailed of Colorado G. H., Jr. Rouge, of Colorado. estimation mexicana. Rocky Mountain adjacent plants 1966. Observations Tadarida Mountains; eds.), Lowery. 1955. bat, L. + 90. 666 pp. Photographic Amer. Midland Po H. Li tsey, Denver, Mexican brasiliensis. tailed of the and R. B. Da v i s , 1971.· of 1:1-600 Second ed , Mamm., 47g78-86. Humphrey, S. R. Krutzsch, Manual Sage Books, Co Fog II, size New York, America. Rocky Mountain Arizona, in 331 pp. migratory bats, plants. central Tadarida Sou t h we s t ern Nat.. 21: 321- �27e Pagels, J. F., and C. Jones. of the free-tailed cynocephala Petit, 1974. bat, (Le Conte). Growth Tadarida Southwestern M. G., and J. S. Altenbach. record of environmental stratified vertebrate envi r onmen t , Ramaley, F. 1942. southern 1973. 19:267-276. A chronological from deposited analysis of in a sheltered Res., 6:339-343. Vegetation Colorado. brasiliensis Nat., chemicals excretion Environ. and development of the San Luis Univ. Colorado Valley Studies, in Ser. 0, 1:231-277. R. F., Jr. Reidinger, populatio~ Arizona, Ross, A. 1972. 1967. influencing Unpubl. levels. Tucson, Factors Ph.D. disserto, bat Univo 186 pp. Ecological aspects bats. Proc. Western insectivorous Arizona of the food habits Found. Vert. of Zool .• 1:205-263. Schwartz, A. 1955. brasiliensis The group status of of the genus the species Tadarida. of J. the Mamm., 36:106-109. Sherman, H. B. bat. 1937. J. Mamm., 1939. J. Mamm., Short, H. L. tailed Simmons, bats Breeding habits of the free-tailed 18:176-187. Notes on the food of some Florida bats. 20:103-104. 1961. bats. Growth and development Southwestern J. A., et ale (Tadarida). 1978. J. Compo of Mexican free- Nat., 6:156-163. Echolocation Physiol., by free-tailed 125:291-299. �271 Smith, E., and W. Goodpaster. in Ohio. Spenrath, J. Mamm., 1960. eastern population Texas. bat found 41:117. C. A., and R. K. LaVal. of a resident A free-tailed 1974. An ecological of Tadarida Occas. Papers Mus., study brasiliensis Texas in Tech Univ., at the Orient Mine, 21:1-14. Stone, J. B. 1934. Colorado. Storer, Limonite Econ. Geol., T. 1. 1926. Bats, deposits 29:317-329. bat towers and mosquitoes, J. Hamm., 7:85-90. Tuttle, M. D. 1975. Population (Ml'otis grisescens): and development. Kansas, ecology factors Occas. influencing Papers Population ecology factors influencing newly volant young. J. W., JL 1955. cavern-dwelling Mus. Nat. Hist., Univ. Tadarida 57:587-595. Aspects of a population bats. mexicana. 1978. J. Northern observations Mamm., rift guide New Mexico and Colorado Bur. Mines and Mineral Gale on a colony of 1, Denver-Alamosa. to Rio Grande (J. W. Hawley, Resources of Commerce. Research of 37:42-47. Pp. 13-32, in Guidebook States. study of J. Mamm., 36:379-390. Colorado. United States Department growth and survival Ecology, Ecological 1956. o. growth of the gray b~t (Myotis grisescens): Tweto, early bat 36:1-24. 1976. Twente, of the gray ed.). rift in New Mexico Circ., 163:1-241. 1978. Co., Detroit, Climates 1:1-606 pp. of the �272 1983. 1982. ------ 1983. Climatological '1'.A. 1970. 216, Villa-R., Colorado summary: Colorado Flight patterns of bats New York, B., and Mamm., and flight characteristics Mamm., 47: 249-260. J. bats. and aerodynamics. (W. A. Wimsatt, ed.). Pp. 195Academic 1:1-406. Eo L. Cockrum. guano bat Tadarida Migration 1962. brasiliensis mexicana in the (Saussure). J. 43:43-64. H. D. Illinois taken Acad. E. R. Colorado. A Brazilian 1970. brasiliensis) Warren, data monthly Morphology 1966. in Biology Press, Walley, summary: 88(1-12). of molossid ------ data annual 87(13):1-28. 1984. Vaughan, Climatological in north-central ScL, bat (Tadarida Illinois. Trans. 63:113. Further 1908. free-tailed Colorado College notes on Publ., the mammals of Gen. Ser. 33, Eng. Ser., 1:61-89. 1910. The Sons, New York, Williams, High mammals of Colorado, 300 pp. T. Co, L. C. Ireland, altitude brasiliensis, G. P. Putnam's flights observed and J. M. Williams, of the free-tailed with radar. bat, 1973. Tadarida J. Mamm., 54:807-821. �273 GEOGRAPHIC RELATIONSHIPS AMONG SOUTHWESTERN OF THE BRAZILIAN FREE-TAILED POPULATIONS BAT INTRODUCTION The Brazilian ~x ica~~ States free~tailed (Saussure), to wintering R. and Cockrum, New Mexico, Mexico grounds Kansas, in Mexico Populations Oklahoma, to (Cockrum. 1969; Constantine, 1958, 1959, 1982), whereas New Mexico across Sonora Sinaloa Sierra Madre migrate but apparently (Cockrum, California throughout (Benson, The Colorado and colony is in the San Luis of males few adult females fall only (Svoboda Q from June and young 1984). west of the Migratory These bats southward into in other probably are local, flyway habits southern probably western areas of resident seasonal movements p e r s , comm.). documented primarily 1969). go Oregon the year and have only of Mexico Populations southern winter and eastern in western Arizona, not of et al., 1962: Glassa California. 1969). for in central to areas do 1947; P. Leitner, Mexico parts have a well-documented known for populations and southeastern Mexico eastern (Cockrum, Nevada, and eastern and adjacent populations and western Occidental in central 1967: Davis Arizona are poorly brasiliensis (Cockrum, 1969~ Villa- Texas. migrate and Tadarida mi qra tes from the southwes tern Uni ted 1962). apparently bato of T. brasiliensis Valley. through The colony mid-August are present Free-tailed in consists although in late summer bats are absent a and from the �274 roost from November northern edge of situated between through the May. range the region of The colony is at the ~£~!i!i~~!i!D a~d is T. from which free-tailed bats fly west of the Sierra Madre Occidental for winter and the region from into which winter free-tailed (Cockrum, When was discovered to be about 9,000 bats It now consists of about reaches size maximum colony until roosts 1932. excavation, surface of mine eastern in the Orient Most of Mexico for or stope, has summer bats that occurred (Meacham, bats (Svoboda, congregate was 1902. when it The from 1881 in a cavernous not Therefore, relatively was 1971). 1984). which was worked probably before its size free-tailed Mine, the in 1968, in August 100,000 in late of the hillside this fly 1969). the colony estimated bats open to the colonization recently (Svoboda, 1984) . The free-tailed bats corridor for dispersal (Freeman and Wunder, reported specimens Grande in northern state line. banded Also, on 5 June to in Cockrum basin at Monte in western Vista, San Grande Luis et from a Valley a L, along as (1975) the Rio 95 km of the Colorado Creek Cave in Greenlee 1964 within Co., the Rio Grande Rio Grande Co., Colorado, just Mine. suggested Mexico the Rio (1969, in litt.) noted that a male on 20 June 85 km south of the Orient the Findley within 1963 at Eagle drainage winter from press). New Mexico was recovered (1969) and use of T. brasiliensis Arizona, Cockrum might have that, if free-tailed breeding areas bats separate that from �275 those of free-tailed bats that winter western populations The and eastern objectives of this genic differentiation populations the colony study those populations were saved Springs males, (50 males); 3. 1/2 OKLAHOfvlAg 5 females); Kansas Woods Greer prepared Bats and as Pinal N, N, 6 mi. 2 1/2 2 mi. W Reed Cave, Bats were prepared of the High study from skins; Cimarron both collected Coo, Picacho COLORADO: E Mineral mi. Hot (1 E Hooper (5 males, (2 25 7 mi. S, 2 mi. W Aetna, from Colorado. Arizona, as study skins and are Plains, Fort from Greer Co., Oklahoma, those bats Co., 3 mi. S, 2 mi. E Kenton Co., 20 females). in the Museum skeletons mi. Cimarron and Woods Co., Oklahoma, University. to one of 2 females). 3 mi. Co., Merrihew (29 males, housed ARIZONA: (25 males, Mine, females); genetically whether AND METHODS localities. Co., orient female). of distinct to determine from 164 fully-grown 30 mi. NW Tucson Saguache the extent than the other. at the following Peak, existed, then genetically. to assess is more similar MATERIALS Tissues differ the two migrationally and, if differences in Colorado might were between in eastern Mexico, Co., samples Hays State were prepared Oklahoma, are housed as were in The Museum, Texas Tech University. The locality free-tailed for winter in Arizona bats migrate (Fig. 1). is in the region west of the Sierra Madre The localities in Oklahoma from which Occidental are in the �276 40 30 120 1 10 100 Fig. 1. Approximate limits of migrationally distinct groups of Tadarida brasiliensis in the southwestern United States, after Cockrum (1969). Circles indicate localities from which samples of T. brasiliensis were obtained for this study. Solid line denotes northern limit of species' range, after Hall (1981). �277 region from which The localities by Cockrum Colorado free-tailed in Colorado (1969). nitrogen. together and liver of saline grinding g Trizma base, adjusted 20 Heart the region two were analyzed solution g 0.37 tissues Tissue and and stored to 608 (1.21 water, were centrifuged -80oC at ground in about 1 ml 1 1 deionized homogenates in and stored were with the pH adjusted EDTA, as mapped separately. tissues were removed and kidney Mexico. localities tissue was ground separately with HCl). minutes from other samples kidney, and liver in liquid to eastern are from neither Specimens were pooled: Heart, bats migrate until pH for used for electrophoresis. Horizontal allozymic starch-gel variation. electrophoresis Gels were made starch gel and 11.5 g electrostarch 8.0 buffer and milliamperes electrode were buffer. Stains of presumed polymorphic (a locus the 1,2,3 allele samples): isomerase about loci were were mannose as of which six if the phosphate et al., 1973): phosphoglucomutase- superoxide dimutase (SOD-I) 1977). These loci were scored for all and in statistical the pH 8.0 than 0.99 in any Hopkinson, used 65 prepared polymorphic of less (EST-2): at noted otherwise. examined, was considered esterase hours with Tris-citrate buffers had a frequency (MPI-l) (Nichols (PGM-l,2,3); and 38.9 g Connaught seven et al. (1971) unless Thirty-eight most common for with in 420 ml Tris-ci trate pH and 100 to 130 volts de'scrLbed by Selander were run was used to assess analyses. (Harris and individuals EST- 2 had four �278 alleles: MPI-I, PGM-I,2 and SOD-I each had three alleles; PGM-3 had two alleles. The following loci adenylate kinase-I,2: creatine kinase: fumarase-I,2: transaminase-2 phosphorylase: peptidase dehydrogenase: dehydrogenase. One sufi iciently most common slow allele. "C." All allele, grea test from the were Statistical and Selander, individual genotypes Genotypic and mean frequencies but this xanthine survey locus numbers of was not anodally. was end were of than The locus with "1" and to loci moving gel. Side-by-side scoring of alleles. performed 1981) software. each slower des ignated the "Au as the and "C" a rarer that was were assigned to check for with fast allele anodal I y analyses (Swofford frequencies dehydrogenase: alphabetically loci moved anodal used nucleoside 6-phosphogluconate had a "D" allele larger malate to score for all indi v i dua Ls , mobil ity consecutively isocitrate in the preliminary "B" a rarer EST-2 glutamate diaphorase: B: for GOT-I, were designated polymorphic comparisons A, individual polymorphic Alleles phosphate hexokinase: NADH sorbi tol was heterozygous glucose dehydrogenase-I,2; enzyme; SOD-2; dehydrogenase; dehydrogenase-I,2: lactate. malic farther glucose (GOT-2); dehydrogenase-l,2; aconitase; dehydrogenase-l,2: dehydrogenase: dehydrogenase-l,2; the alcohol glycerol-3-phosphate oxaloacetate enzymes monomorphic: albumin: glucose-6-phosphate isomerase; were using Data were entered as variable heterozygosities for variable loci the BIOSYS-l locus. were Allelic calculated. in each sample were �279 tested for Chi-square goodness-of-fit to expected genotypic frequencies under Hardy-Weinberg equilibrium; Levene's (1949) correction for small sample sizes was used to compute chi- square values. Chi-square values also were figured with data pooled into three genotype classes: most common allele; 2) 1) heterozygotes homozygotes for the for the most common allele and one of the other alleles: 3) all other genotypeso Exact probabilities were calculated for pooled data. An index (FISi) was calculated for all loci in each sample to indicate in which direction the observed deviated from the expected. A negative !ISi indicates that there are more observed heterozygQtes heterozygosities than expected and a positive !ISi fewer observed heterozygotes than expected at Hardy-Weinberg equilibrium (Wright, 1965). Nei's Rogers' (1978) unbiased (1972) similarity distance (QT) (Wright, quantify the relationship identity (I) and distance (S), and the modified 1978) indices between were pairs (DL Rogers' calculated of samples. to F- statistics were calculated to analyze genetic differentiation of individuals relative to the sample they comprise (!IS)o of individuals relative to all samples pooled (!IT), and among all samples amount (FST). !ST can be considered of differentiation among maximum amount of differentiation samples a measure of the relative possible--that ·samples are fixed for different alleles. to the is, when This index measures the extent to which allelic differentiation has become fixed rather than the absolute amount of differentiation among �280 samples; E:.ST= 0 represents no differentiation among samples and EST = samples. 1 represents fixation for different alleles among ~IS is a coefficient that gives the probability of two identical alleles from a common coefficient alleles at a locus being derived by descent ancestor that gives within the sample: the probability ~IT is a of two identical at a locus being derived by descent from a common ancestor within all samples pooled. are positive when conditions population; a negative, heterozygotes. inflated recognized increase value However, positive not necessarily These two coefficients indicates subpopulations when the samples an in a excess of values of !IS and !IT do indicate inbreeding. if genetic homozygosity These values exist but can be are not are taken: this situation is known as the Wahlund effect (Wahlund, 1928). RESULTS Allelic frequencies for each of the variable loci in the samples are given in Table 1. in any sample. Variability NO unique alleles were found for PGM-3 was not observed bats from Picacho Peak, Arizona, or Cimarron Co., ok Lahoma Chi-square values used to test deviations in , of allelic frequ~ncies from the Hardy-Weinberg model are given in Table 2 together with corresponding (!ISi) vaiues that indicat~ in which direction the observed heterozygosities deviated from those predicted by the modelo (p < 0.05) from the model heterozygotes were observed PGM-2 deviated significantly in all of the samples: than were predicted. fewer The only �281 Table 1. Brazilian for Allelic free-tailed localities identified of frequencies bats for six variable loci in from the Southwest. samples. Abbreviations for loci are in text. Samples Locus EST-2 Allele (N) A C D PGM-l (N) A B C PGM-2 (N) A B C PGM-3 (N) A C MPI-l (N) A B C SOD-l (N) A B C Orient Picacho Merrihew Reed Cimarron 51 0.725 00176 0.098 27 0.833 0.130 0.037 . 49 0.816 0.153 00031 30 0.700 0.200 00100 7 0.786 0.214 00000 47 00915 0.085 0.000 27 0.963 0.037 0.000 49 0.918 0.071 0.010 30 00950 0.033 0.017 7 00786 00143 0.071 49 0.867 0.041 0.092 27 0.815 0.056 0.130 46 0.859 0.033 0.109 30 00917 0.083 0.000 7 0.429 00429 00143 51 0.980 0.020 27 1.000 0.000 49 0.969 0.031 30 0.983 0.017 7 1.000 0.000 ,49 0.939 0.000 0.061 23 0.913 0.022 0.065 44 0.955 0.011 0.034 28 0.929 0.000 0.071 6 0.833 0.000 0.167 51 00775 0.059 0.167 27 0.741 0.056 00204 49 0.776 0.031 0.194 30 0.800 0.050 0.150 7 0.857 0.000 0.143 �282 Table from 2.--Chi-square Hardy-Weinberg !~~~~ (FISi)' correction otherwise. for (X2) values equi 1 ibrium ~£le Exact probability for deviation wi th corresponding ~!!E~~~~~= d~~E~~~ ~ll to test size level 1. used ~ fixation ~~~~~~'~ ~l~ (l9!2) indicated used for significance tests. Samples Locus Orient Picacho EST-2 X2 -~ISi . PGM-1 X2 ~ISi 11.221*a 0.453 0.020b -0.038 PGM-2 X2 28.554*** 0.571 17.033*** 0.649 ~ISi PGM-3 X2 ~ISi MPI-1 X2 ~ISi SOD-1 X2 ~ISi 0.790 0.185 0.010b -0.020 0.172b -0.065 4.073 0.150 0.945 -0.160 Merrihew Reed Cimarron 0.311 -0.056 4.439 0.420 0.327b -0.273 2.038 0.191 0.055 -0.040 00327 -00200 41.013*** 0.826 22.778** 0.782 0.032b -0.032 O.OOOb -0.017 6.477 0.462. 13.488 0.481 8.148 0.462 0.111b -0.200 1.694 0.180 0.262 -0.020 4.799 0.303 0.091b -0.167 ___ c 8.229* 0.533 c * P < 0.05, ** P < 0.01,. *** P < 0.001 a Chi-square value using Levene's (1949) correction for small sample size with no pooling of data. b No homozygotes for rare allele in sample (one cell 0). c No variation. = �283 other locus to deviate significantly (P < 0005) from the model was PGM-l for the Colorado sample: this locus also had fewer heterozygotes than predicted. ~-statistics are presented in Table 3. The high mean value for E:IS indicates an excess of homozygous bats within each sample and the high mean FIT indicates a greater number of homozygous individuals relative to the number expected under Hardy-Weinberg equilibrium for all samples pooled. mean E:ST of 0.052 indicates that the populations The have diverged 5% of the way toward total genetic differentiation (all populations being fixed for different alleles). This (f < 0.005) level of divergence from zero. (5%) differs significantly The high KIS' E:IT'and KST values there might be inbreeding there might be genetic occurring indicate that within populations subdivisions within samples, or thus causing a Wahlund effect. Genetic similarity is high and genetic distance is low between pairs of samples Nei's (1978) ! ranges (Table 4). from 0.970 (Reed to Cimarron) to 1.000 (Picacho to Merrihew: Orient to Picacho, ranges from pairings Merrihew, and Reed). 0.943 to 0.972 for all with Cimarron, which Rogers' (1972) ~ combinations range except from 0.860 to 00866. Nei's (1978) Q ranges from 0.0 to 0.002 for all pairs except those from Cimarron, which range from Modified Rogers' D (Wright, pattern: all pairs except those with Cimarron 0.034 to 0.072, and Cimarron 1978) values 0.022 pairings show to 0.0310 the same range from range from 0.179 to �284 Table 3.--F-statistics as calculated ~!! ~ polymorphic degrees of individuals, localities BIOSYS-! loci. freedom r = = (Wright, (Swofford Chi-square of and Selandero (X2) = 1981) for _!NFST(r - 1:) with where N = number of ~.!.!~.!.~, and s = number of (r - !)(s number 1965, 1978, Neio 1977) - !), sampled. X2 do f. 0.014 13.776 12 0.061 0.036 23.040* 8 0.633 0.689 0.153 97.308* 8 PGM-3 -0.024 -0.014 0.011 7.216 8 MPI-l 0.148 0.169 0.025 15.000 8 SOD-l 0.106 0.113 0.007 4.592 8 Overall 0.208 0.249 0.052 Locus ~IS ~IT ~ST EST-2 0.065 0.078 PGM-l 0.026 PGM-2 * P < 0.005 299.832* 72 �285 Table 4.--Matrices coefficients for samples diagonal, Nei's diagonal, Nei's Below diagonal, 1978), above of genetic of Tadarida (1978) unbiased (1978) unbiased modified diagonal: similarity Roger'~ brasiliensis.~. genetic distance genetic Rogers' ~nd distance (1972) (D)i identity distance (DT) genetic Below above B. (1). (Wright, similarity (S) • A. Population 1 2 3 4 5 1 Orient ***** 1.000 1.000 1.000 0.972 2 Picacho 0.000 ***** 1.000 0.998 0.979 3 Merrihew 0.000 0.000 ***** 0.999 0.974 4 Reed 0.000 0.002 0.001 ***** 0.970 5 Cimarron 0.028 0.022 0.027 0.031 ***** 1 2 3 4 5 B. Population 1 Orient ***** 0.955 0.972 0.969 0.865 2 Picacho 0.051 ***** 0.967 0.943 0.866 3 Merrihew 0.038 0.034 ***** 0.947 4 Reed 0.040 0.072 0.062 ***** 0.860 5 Cimarron 0.187 0.179 0.187 0.196 ***** �286 0.196. The difference other (N samples = between the sample might be an artifact from Cimarron of its small sample and size 7). DISCUSSION Migrationally tailed not bats conservative distinct; genotypic allelic samples This mixing colonies among indicate and of = Armitage, separated fST of the has reported where dispersed 1980). bats samples for differentiation to that is with no (0.052) young are there 0.07) in Colorado because free-tailed that among populations slight (fST was occurring samples indicate mean free- in this study, is similar of marmots (Schwartz geographically The differentiation for nine colonies genetic results variability study of Brazilian as sampled differences. in this occurred. populations in the Southwest, genetically fixed distinct among Differentiation was less fuoose from than for Scandinavia that had an fST of 0.096 (Ryman et al., 1980). Brazilian roosts free-tailed throughout In general, New Mexico Mexico the year, show little by Cockrum (1969) evidence of North Great Plains Great America, free-tailed 1952 bats banded and autumn. and western to eastern to September recovered was found regions. and change New Of 162,892 bats banded 539 were but none or in adjacent Arizona of movement Plains. regions, mobile in spring in eastern from September and adjacent areas 220,000 especially free-tai led bats or the southern Arizona bats are highly in in other on the southern Likewise, in northern 1967 of more than Oklahoma (Glass, �287 1958, 1959, 1982}, southern Texas central Texas (Eads et al., 1957; eastern Texas (Spenrath was recovered (Short Short and Laval, in western et et al., 1974), none Mexico, ale, Arizona, 1960)' 1960), and subsequently or western New Mexico. These data indicate within a particular banded Constantine distinct Caverns foreign recoveries Arizona, western in years recovered Creek Cave, same subsequent eastern .of genetic to maintain the 27,000 and in areas Springs~ Caverns record mixing suggested high Southwest. 1957~ females of eastern Three bats Nevada, three Largo females in Eagle recovered New Mexico. later 1967; and recovered in One bat that Glass, for a free-tailed Intuitively, in to the west, was recovered exists bats of 41 and in Monte (Constantine, pers. comm.). to adjacent females 'recovered western 1967) the same year include: two Mexico bats either Mexico. recovered five stay free-tailed in 1956 to banding, near Ely, such within New Mexico, Arizona; in Jalisco, (P. Leitner, over and western were Sonora; and Radium cross 13 were bats (Constantine, Other foreign recoveries 1959). An unpublished Kansas than vicinity Caverns Mexico. that was banded bats more New Mexico, in 1952 at Carlsbad year Caverns regions and in Carbo, Lordsburg banded free-tailed and near Socorro, Cave, Sinaloa, but However, reported, at Carlsbad Las Cruces free-tailed free-tailed (1967) banded at Carlsbad most at Carlsbad that some migrationally banded flyway. or recovered document that bat in western the low level by these records seems inadequate genetic identities and low genetic �288 distances among flyways; might be indicative of a much greater at the time and place of breeding. and east from maternity reported by Glass populations however,. the few records (1982), does in the Southwest and KIT values populations primarily of homozygosity probably (Chesser, male when is a maternity September: southeastern transients Peak 1959). Oklahoma might is a small was mostly autumn as bats comm.) and possibly that consist Cave present feeding does primarily are composed of bats units presumably move was in early (Glass, taken or drinking grounds in becau~e roost spring 1959). it when in late August was obtained (R. Chesser, a colony. breeding in Picacho in pers. Colonies in spring and autumn several was sampled of one sex or that are situated are formed in February from wintering within Merrihew is a maternity not represent from KIS of this bat on the Great from Cimarron they might be used by transients of in Colorado and and the sample The sample were inbreeding in mid-August. that was sampled colony, among by positive colony 1958) colonies been as the vagility should occur at Merrihew Reed have male. The (Glass, few transients (Glass, indicate it was sampled roost movement as indicated 1983). Oklahoma, such movement. does not is one of the northernmost Plains not indicate over to the north in northern but reflects these bats that might facilitate The excess amount of crossing Scattering colonies avai Lab Le units. where likely These and March as the bats to summer roosts. �289 The high fIS in free-tailed time and location aggregation different sampled of sampling, of individuals allelic bats, therefore, likely from different frequencies. at the locations in this known where constitutes tailed are winter and ~any et in Mexico al., roosts the however, at did through almost and as a maternity no males found together Laval, and female congregating for several subsequently summer. reported in that roost. No free-tailed in night weeks were such P. Leitner bats used as likely for free-tailed mating as a are when were for mating having (pers. comm.) observed roosts California winter These nursery serves (Spenrath and in northern February. de roost and females roosts away from diurnal in any when males in eastern Texas was presented the locate through August, Some males spring 1974), but no evidence taken place male in early March, roost April are present. to as a multi-purpose transient present, maternity that Cueva This cave probably from November free- thereafter not with bats present year-round. roost what means that mating he observed la Tigre, near Carbo, Sonora, serves and migrate segregate (1969) it is Female arrive do not sexes Cockrum in Mexico: to mate This presumably and 1962). they the had been Unfortunately, gather males to units with units for this species. when with the females. place (Davis unit the F1S theoretically study, bats pregnant in Texas, colonies takes a breeding bats colonies free-tailed due breeding If breeding would be lower and fST would be higher. not is given night in congregations bats in the Southwest. roosts roosts spring have and been �290 The highly United States not free-tailed and northern of a panmictic suggests mobile reproductively exist, however, sedentary between southeastern differentiation characteristics of genic differentiation populations of !. brasiliensis Reproductive isolation southwestern popUlations in the Pacific coastal United States. would be expected southwestern thus have Absence isolated. populations '.:.han among Mexico population. that southwestern bats of the southwestern If so, between are might and the more region and the greater those genic populations populations. ACKNOWLEDGMENTS I thank use of his from Dr. R. K. Chesser, laboratory Oklahoma. provided Yar the bats the manuscript, R. J. Zakrzewski) was funded facilities, Petryszyn from Arizona. as did other (Drs. C. A. Ely, E. D. Fleharty, Tech University, and specimens (University of Arizona) Dr. J. R. Choate of my thesis M. E. Nelson, at Fort Hays State University. Division for guidance, members in part by the Nongame N-3R, Colorado Texas Research of Wildlife. reviewed committee F. W. Potter, This Program, study Project �291 LITEHATURE Benson, S. B. 1947. in Tadarida Chesser, R. K. Comments mexicana. 1983. populations CITED on migration J. Mamm., Genetic and hibernation 28:407-408. variability of the black-tailed within prairie dog. and among Evolution, 37~320-331. Cockrum. Eo L. 1969. brasiliensis. Migration in the guano bat, Tadarida Misc. PUbl. Mus. Nat. Hist., Univ. Kansas, 51:303-336. Constantine, D. G. free-tailed Davis, 1967. bat. Activity Univ. New Mexico R. B., C. F. Herreid Mexican patterns free-tailed II, bats and in of the Mexican Publ. Biol., 7:1-79. H. L. Short. Texas. Ecol. 19620 Monogr o. 32:311-346. Eads, R. B., J. S. Wiseman, Observations Tadarida Findley, concerning mexicana, Mammals Albuquerque, Freeman, of Glass, B. P. L. Wunder. the Brazilian mexicana) -------- 1957. free-tailed bat, D. E. Wilson, and C. Jones. Univ. New Mexico Press, in press. Observations free-tailed in bat southern at a (Tadarida Colorado. Nat. 1958. in Oklahoma. Menzies. Texas J. Sci., 9:~27-242o of New Mexico. brasiliensis -----------Southwestern C. 360 pp. Jo, and colony G. the Mexican in Texas. J. S., A. H. Harris, 1975. and Returns of Mexican J. Mamm .• 39:435-437. freetail bats banded �292 Additional 1959. banded in Oklahoma. 1982. Tadarida Great Plains. E. R. movements of brasiliensis Southwestern 1981. The mammals bats in human Mexican freetail banded in the Nat., 27:127-133. of North America. H., and D. A. Hopkinson. electrophoresis free-tailed mexicana Second ed. 1:1-606 + 90. John Wiley and Sons, New York, Harris, from J. Mamm., 40:542-545. Seasonal bats Hall, returns 1977. Handbook genetics. of enzyme North-Holland Pub!. Co., New York. Levene, H. 1949. On a matching Ann. Math. Stat., arising in genetics. 20:91-94. M~acham, J. W. 1971. Nei. M. 1977. F-statistics in subdivided problem Entry. Bat Res. News, 12:37. and analysis populations. of gene diversity Ann. Human Genet., London, 41:225-233. 1978. genetic distance Genetics, Nichols, Estimation from a small heterozygosity number and of individuals. 89:583-590. E. A., Polymorphism isomerase V. M. Chapman, and linkage in Mus musculus. 1972 . .~:"":~-::t)C of. average Measures distance. and for F. H. Ruddle. 1973. mannosephosphate Biochem. Genet., 8:47-53. of genetic Stud. Genet. similarity and VII, Univ. Texas Publ., and T. Nygren. 19800 7213:145-153. ~~~?n. N., C. Reuterwall, Genetic variation K. Nygren, and differentiation in Scandinavian �293 moo se (A1 c e sal c es ): ---- ---- Evolution, Schwartz, O. Selander, J. A., in the K. B. Armitage. social Gentry. the in the genus mouse VI, Univ. Movements S. Y. Yang, Biochemical Genetic marmot model. of Southwestern and polymorphism and I. Variation polionotus). in Stud. Texas Publ., 7103:49-90. and t.he Mexican C. F. Herreid free-tailed 1960. II. bat in Texas. Nat., 5:208-216. C. A., and R. K. LaVal. of a resident eastern W. E. Johnson, Peromyscus. (Peromyscus H. L., R. B. Davis. Spenrath, monomorphic? 1980. mammals: 1971. old-field Genet. mammals 207:665-667. systematics Short. and R. K .• M. H. Smith, B. large 34:1037-1049. variation Science, are population Texas.' Occas. 1974. An ecological of Tadarida Papers Mus., study brasiliensis Texas Tech in Univ., 21:1-14. Svoboda, P. L. relationships San Luis Natural 1984. of the Brazilian Valley of Colorado. Hays State University, Swofford, D. L.g and computer program in genetics. Villa-R., guano , B., and geographic bat in the M.S. thesis, Fort 69 pp. 1981. for the analysis E. L. Cockrum. 1962. brasiliensis BIOSYS-l: of allelic Urbana, Mamm •• 43:43-64. I ,I Unpubl. Univ. Illinois, / and free-tailed R. B. Selander. bat Tadarida \ history a variation 65 pp. Migration mexicana in the (Saussure). J. �294 Wahl und, S. Zusammensetzung 1928. korrelationsercheinungen vererbungslehre Wrighto s. by 1965. mating. Vol. 4. populations. Approved The interpretation with Evolution, 1978. vom aus betrachtet. F-statistics special populationen standpunkt Hereditas, der 11:65-106. of population regard und to structure systems of 19:395-420. Evolution and the genetics Variability Univ. von within Chicago by Charles M. Haynes Wildlife Researcher C Press. and of populations. among 580 pp. natural �295 JOB PROGRESS State of Project Work Plan Job REPORT Colorado N-4-R-2 Lowland Riparian Studies Cottonwood Community 1 __::-----1 Period Covered: Author: Personnel: 1 July i983 - 31 June 1984 Warren D. Snyder Gary C. Miller Wildlife. and Warren D. Snyder, Colorado Division of ABSTRACT A preliminary compilation and analysis of inventories of the status and trends of cottonwoods (Populus spp.), their regeneration, and of land use changes (under contract with the Colorado State Forest Service) along the South Platte, Rio Grande, and Colorado rivers was completed and disseminated in an interim report. A contract was initiated to complete inventory of the Arkansas River and a segment of the South Fork of the Republican River. Sampling of cottonwood seedlings, resulting from natural regeneration along the South Platte River in 1983, was initiated to identify factors affecting survival. Literature and other information were compiled and analyzed as a basis for preparing a program narrative study plan which was submitted for review and approval. Trial test plots using stem cuttings of 10 woody species (trees, shrubs, and vines) were initiated in late winter to evaluate stem cutting propagation in riparian sites. This Job Progress Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the aur hor . ��297 LOWLAND RIPARIAN COTTONWOOD COMMUNITY STUDY Warren D. Snyder P. N. OBJECTIVE To prepare a study plan evaluating techniques for regeneration of cottonwood stands in riparian streamside ecosystems in lowland sites of Colorado. SEGMENT OBJECTIVES 1. Review literature and consult with personnel the cottonwood-willow riparian ecosystem. 2. Select regeneration 3. Survey cottonwood-willow ecosystems and select study sites for implementing and evaluating regeneration techniques. 4. Prepare a detailed techniques knowledgeable about to be field tested. study plan to evaluate regeneration techniques. RESULTS AND DISCUSSION The Colorado State Forest Service was previously contracted under a nongame study plan to inventory and quantify condition and trend of cottonwood stands, other vegetation, and land use over an approximate 30-year span. They used photo interpretation methods on stratified random samples of linear miles along the lower portions of the South Platte, Rio Grande, and Colorado rivers. Data provided in this initial inventory were compiled and summarized in an interim report distributed to interested personnel within the Colorado Division of Wildlife within this work segment. The Colorado State Forest Service, under a new contract, began inventory of the lower Arkansas River and a short segment of the South Fork of the Republican River below Bonny Reservoir. Work is to be completed during spring 1984. Since the initial analysis did not use statistical methods and data are not yet available from the 2nd contract, inventory information is not included within this segment report. It will be included as soon as complete analysis and comparision of all drainages are available. Surveys of natural regeneration of plains cottonwoods (Populus sargentii) were begun in summer and fall 1983. Several intensive test transects were established to monitor seedling survival on grazed and ungrazed sites in relation to water table fluctuations and other variables. These will be supplemented with additional transects in late spring 1984. Additional extensive transects will be established to monitor the �298 Table 10 Stem cutting propagation trial plantings on the check station meadow, South Platte Wildlife Area, February-March 1984. N Species Plains cottonwood (Populus sargentii) planted Site Date 25 1 2 21 Feb 24 Feb Sandbar willow (Salix interior) 12 1 05 Mar Coyote willow (~.exigua) 25 10 1 2 14 Feb 02 Mar 6 1 21 Feb Golden willow (Salix sp.) 12 12 15 1 2 3 23 Feb 01 Mar 02 Mar Indigobush amorpha (Amorpha fruiticosa) 15 1 21 Feb Elderberry (Sambucus spp.) 12 12 1 2 01 Mar 02 }far Cotoneaster (Cotoneaster spp.) 15 12 1 2 01 Mar 05 Mar Frost grape (Vitis vulpina) 12 1 02 Mar 4 1 05 Mar Peachleaf willow (~.amygdaloides) Virginia creeper (Parthenocissus quinquefolia) Totals 6 205 �299 frequency of occurrence of cottonwood seedlings within the South Platte River floodplain. Cottonwood-willow ecosystems were surveyed in eastern Colorado to examine potential study sites. Surveys of West Slope riparian areas will be conducted in 1984. Literature was compiled and reviewed covering riparian ecosystems, natural and artificial regeneration of cottonwoods and willows, and vegetation sampling methods. Contacts were made with personnel of the U.S. Department of Agriculture, Soil Conservation Service, in New Mexico concerning stem cutting propagation, and with other Soil Conservation Service personnel concerning sources for certain species. Based on available data, 2 techniques, stem cutting propagation and soil scarification were selected for testing in riparian sites. A study site was selected on the South Platte State Wildlife Area near Crook and preliminary stem cutting propagation trials were initiated. A tractormounted earth auger was used to bore holes into the ground water table which was approximately 0.5 m below the soil surface. A summary of species and numbers of stem cuttings planted is provided in Table 1. Total stem cutting length, length extended above the water table, and length extending above ground were measured on each planting. This permitted determination of the distance the stems were submerged below water level for subsequent comparisons with survival. Capped plastic tubes were inserted into the water table at 4 sites within the meadow monitor changes in ground water level. Additional effort during the segment centered on preparation of a detailed program narrative (study plan). This plan is under review and will be approved for implementation beginning 1 July 1984. Prepared by Warren D. Snyder Wildlife Researcher ��301 APPENDIX A FINAL REPORT Colorado State Forest Contract, and Interim Analysis 1983 ��303 AERIAL PHOTO INTERPRETATION PLANIMETERING AND OF RIPARIAN VEGETATION or COLORADO By Thomas Owens Dan Teska Colorado State Forest Service Fort Collins, Colorado For Colorado Division of Wildlife June 30, 1983 �304 INTRODUCTION During the past few years Division of Wildlife (DOW) researchers have noticed, in the course of other work, an apparent lack of cottonwood reproduction in riparian resources in Colorado. (Riparian: "lands bordering fresh water bodies and the resources they support", Schmidt in Management of Cottonwood-Willow Riparian Associations in Colorado.) Since riparian areas .have more wildlife than any other in Colorado trend. species in them (Graul, Ibid.), this could be a serious The DOW requested the Colorado State Forest Service (CSFS) Map Shop do a vegetative survey using aerial photos to document the actual trend of riparian vegetation in Colorado. CSFS personnel working on this project were Dan Teska and Carolyn Krupp, photo and map work, with Thomas Owens as Supervisor. AREA DESCRIPTION Three rivers were chosen to be mapped: Rio Grande, Colorado, South Platte. and These are the largest rivers in Colorado and contain the most riparian habitat (the Arkansas River may be mapped at a future date). The South Platte is located in central and northeastern Colorado. The study area starts east of Greeley and runs 156 miles east to where the river flows into Nebraska. The western end has an elevation of 4,640 feet above sea level and drops to an elevation of 3,435 feet at its easternmost point. The South Platte is ;'n a broad gentle valley, which creates a broad band of riparian vegetation along the river. The major cottonwood species is plains cottonwood (Po~--sargentii). �305 .The Rio Grande is located in southern Colorado. The study area starts at South Fork and runs 60 miles south and east to the confluence of the Conejos River and the Rio Grande. The elevation at South Fork is 8,160 feet above sea level, and drops to 7,480 feet at the southern end. In the northern and western end of the study area there are cliffs on the northern bank of the river, which constricts riparian vegetation development on this bank, but otherwise the river flows in the very broad and flat San Luis Valley which creates ~ broad band of riparian vegetation. is narrowleaf cottonwood The dominant cottonwood species (Populus latifolia). The Colorado River is located in western Colorado. The study area starts at the western edg~ of Glenwood Springs and runs 104 miles west to the entrance of Horsethief Canyon, two mil~s west of Gilsonite. Easternmost elevation at Glenwood Springs is 5,720 feet above sea level and drops to 4,450 feet above sea level at the western end. The Colorado River has a fairly narrow, deep valley which restricts riparian vegetation narrowleaf development. The major cottonwood species is cottonwood. ~'ETHODS 1. Detennined the availability aerial photography and type of 9"x9" vertical stereo of the rivers from the ASCS Aerial Photo Field Office in Salt Lake City, Utah (the holder of Agriculture Department photos) and the EROS Data Center in Sioux Falls, South Dakota (the holder of Interior Department 2. o photography). Prepared maps (see Figures 1-3) which show the river miles (with river mile being the upstream end) and the earliest and latest photography available (a 25-year separation was the minimum difference used). between earliest and latest �(!) s, ~ en l.J... ..J ~ i '" ~ ....• '-c" In C) C) C) a lu V) ~ -J \u c::. \:: >- l.l ~ -.II I.r, ~ ~ t- :t: ~ 0 Vi ( c~ .,.o ~ ..•e ~ >...; W ::::L Y'" .• .,_ -e .,._ .! '"."0 s % ,.. ~ "'- .;, .. I'" .... CI ~ ~- 0 0 0 ,..... '"t «C r.:- ::i .., ...••...- ..,..,~ l- '" 0 f- i 0.. 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Z UI 3 ••• IW ••• Z <E '" ~ I><' >- 0 ~ -c ':I: '" )< ~ )- x 0 -s ";:} I0 e: �309 3. The DOW randomly selected 29 rlver miles on the South Platte, 21 river miles on the Colorado, and 20 on the Rio Grande. 4. Aerial photos were ordered for the sample river miles. 5. A vegetation classification was developed in discussions between CSFS and DOW (see Figure 4 which follows). Figure 4. DOW -- Cottonwood Classification Hay Meadow/Emergents Grassland Up 1 and Shrub Comnun i ty (Xeric Shrub-Grassland & Mesic Shrub Complex) Wetland Shrub Complex Conifers Agricul tural Developed H GR S J AG D C Co t tcnwocds Size Classes: <6" dbh 6"-16" dbh 1 16"-30" dbh >30" dbh 3 4 2 Crown Density Classes: 10°-35;: 35~>55:~. over 55'! A B River Unvegetated/Sandbar R (Bare soil, sandbars, C NV gravel pits) Lakes, Ponds, Reservoirs Tamarisk Minimum polygon size for all types is 1 hectare. ---Classes to be expressed in Ha. E.g., C2B is cottonwood, 35-55~ crown density class. L T 6"-16" dbh, �310 6. Vegetation types were delineated using Bausch and Lomb SIS 95 Stereoscopes and Old Delft Scanning Stereoscopes using drafting pins. 7. Field trips were taken to verify delineation on acetate overlays accuracy on all rivers. Core samples were taken from a broad range of sizes of cottonwoods to try to determine ages of the size classes. Delineated vegetation types from the old photos were transferred onto 8!;i"xll"paper using a Bausch and Lomb Zoom Transfer Scope. The recent photography was transferred onto mylar overlays on top of the old vegetation types. Details which had remained such as road 8. intersections were used as controls onto each other. 9. Areas were planimetered using a Numonics Electronic Areas were measured in hectares. for each type on each map. 10. to properly register the maps Planimeter. A form was devised to record hectares Totals for hectares for each type and date on each river were calculated. 11. Severa 1 di fferent methods were used ina ttempts to count the ages of cottonwoods. RESULTS Overall areas for each river and each date are presented in Figures 5, 6 and 7. .Specific maps and area figures are presented Attempts possible to age cottonwoods to accurately cottonwoods in Appendix was not successful. It was not count rings in sample cores taken from in winter and spring 1983. 2. �311 DISCUSSION The aerial photo, mapping, and planimetering phases were routine and presented no special difficulties. The specific dates and types of photos used can be found with each map in Appendix B. One unexpected situation was encountered on the Rio Grande. South of Alamosa large trees were identified in initial photo interpretation as cottonwood, but were identified on the ground as willows. This situation was confined to the Alamosa National Wildlife Refuge, which runs for 10 miles along the river. The aginq phase presented difficulties. Cores were taken in January 1983 and in May 1983 on the South Platte. These were glued into molds and sanded. Some cores were stained with phloro-glycinol in alcohol and 25% HCL to try to bring out the age rings. Others were sanded with differently grained sandpaper and varnished. These cores were examined under fluorescent lights with a binocular microscope. It was not possible to accurately count rings with these techniques. �312 APPENDIXES Appendix A Photointerpretation Appendix B Maps and Area Statistics Key �313 Appendix A Photointerpretation Key for DOW Cottonwood Haymeadow/Emergents - H Study - Draft This type covers the largest area. Emergent wetlands have a bright red/bright pink appearance or a dark one; haymeaJows and seasonal wetlands sometimes have a blue-green color (color IR imagery). On true-color photography, the color is green to dark green. On black and white photography, haymeadow/emergent wetlillldsappear a dark shade of gray (indicating a lot of moisture). On all types of photography, there is a smooth texture corresponding to this class. Confidence level high. Grassland - GR Grassland has a variable blue-green color on CIR photogr~phy, green to brown color (and all variations in between) on true-color, and a very light tone of gray to white on black and white imagery. The texture is smooth, but slightly rougher than haymeadows, and not as wet. Confidence level high. Shrub Community - S Xeric shrubs (sages) are found mixed with grasses from crown cover percentages of near. a to over 50. It is darker than the surrounding grasses and has a mottled "salt and pepper" texture. Hesic shrubs are found in narrow draws, along the streambeds, on islands, on floodplains of rivers, and at the foot of slopes. On CIR, they have a bright pink color in wet areas (e.g., Willows) to a nearly black color on slopes (e.g., gambel oak). Height can be detected under stereo viewing. Confidence level very high, although in some cases tree-like willows may be mistaken for cottonwoods. On true-color photography, mesic shrubs appear green in color, and are much shorter than surrounding trees. On B&W photography, they have a light gray signature, and a more even texture than cottonwoods; this type produces a billowing effect on the photo (wetland or mesic). Conifers - J Conifers are found in the higher elevation areas of the Colorado River. This type has a black signature (CIR), dark green (true-color), or a dark gray to black (B&W). Height obvious under stereo viewing. Confidence level high. �314 Agricultural - AG Agricultural lands are found on level to gently sloping areas. They are easily recognized by cultivation marks. Center pivot agriculture is circular (or nearly so) on 160 acres. Flood irrigated agriculture is found along streams, with an irrigation ditch sometimes discernable. For these two types of agriculture, the signature is bright red (CIR), green to dark green (true-color), or a light to dark gray (B&W). Dryland agriculture has a blue-green to white color (mature winter wheat or fallow field) on CIR; green or brown on true-color; and light gray on B&W. Confidence level very high. Developed - D Roads have a dark appearance. Buildings, trees and disturbed soil are visible on ranch yards. Excavated areas have a signature similar to bare soil (bright), but are clearly dug out. Maps provide collateral data. Confidence level high. River - R Rivers are obvious, along major and minor channels; same applies to canals. Has a dark signature on eIR and B&W, and a blue to dark blue on true-color. ~~ite areas also apparent, depending on reflectance to film. Confidence level high. Non-vegetated/Sandbar--- NV Usually found along river beds and islands. On all three types of photography the signature is white, and highly visible. Confidence level high. Lakes, etc. - L This type also obvious. level high. Cottonwoods Very dark signature. ,- Confidence - C On CIR signature is dark red; on true-color, green to dark green; on B&\~, light gray. On late fall B&W imagery cottonwoods have a "ghost-like"appearance (leaves gone). On fall true-color, cottonwoods may be yellow. For size classes, cro~~ size is directly proportional to trunk size -- the larger the crown, the larger the DBH �315 of the tree. Density classes are distinguished by spacing of trees and crown cover. Over 55% density class, for example, appears to have over 75% crown cover. �316 Appendix B River Miles Mapped River miles were measured with an electronic planimeter in 1)1> mid-channel. Th~, a sinuous river, such as the Rio Grande, many more river miles in it than might be expected. South Platte River: S tra tum River Miles 1,2,17,30,31,33,37,38,41 54,58,67,68,74,81,83 II 91, 95, 104, 105, 106, 113, 116, 124, 137 III 141, 142,149, IV Rio Grande 163 River: Stratum River Miles 7, 11, 13, 14, 17 21 , 23, 30, 32, 33, 35, 43, 44 II III Colorado Stratum 47, 55, 58, 60, 65, 70 River: River Miles 3, 5, 8, 11 II III IV 20, 22, 32, 36, 37 54, 56, 59, 67 72, 75, 83, 92, 93, 94, 99, 100 has �317 Figure RIVER: M r L E: 5A South P1actc S t rat ura I qUAD: MAP SCALE: TOTAL AREA: 1627.65 ha - Date:01J Photos Scale: Oate: Re c e n t Photos Scale: Chang. + or CIA ClS -- I I 15.52 I 11Ll 1Q_:Q_~_Jl.a ______________ -_------------ ---- 10.5] -- 9.82 _:::.5_,,__il___ 71 - Cle 13. 19 4.!d C2ft. 103.83 80.84 2L_9_:: C2B (:4.87 h9.26 1 C2C jl).O:', 55.83 16 :3A to 1.41 94.0!. C3B 9b. 16 ._"_ r?r Y4.77 \... ..."-- C4A I --_- --- hR __ U·22 -- 76 ?1 r zq _--=..!±._J_2._ -R 1 _____ no R. 73 C4B 3.62 11. 7 8.08 C4C 6.63 l. 09 -5.54 H 594.94 334. 13 -260.8 GR 72.67 171.49 98.82 S [25.34 129.71 -_- 4.37 J ------ 143.Y7 257.7 D 12.7J 44.15 31.42 R G_).86 136.59 70.73 67.81 45.81 31.9 29.69 AG I lV 22.5 L 2.21 T --- I -- -- _:::_]_,_J]__ __ 9_1~4__________________ 1'\ 4.4Y 0 I) e- - - 113.73 �318 Figure RIVER: MILE: South PIJtte St r a t um II QUAD: MAP SCALE: Da te: OlJ 5]) TOTAL AREA: ['110 t os Scale: Date: Recent 1165.07 Photos Scale: Change - + or CIA 8.07 27.86 19.79 ClS 5.78 3.86 -1. 92 CIC .n! 11. C2A 59.21 52.04 .:': .17 f:?R 33.01 64.44 '11.43 C2C 25.85 43.63 17.71:5 ,C3A 143.(J 82.94 34.5 71. 64 27. 1LI llL-_ .- 13.65 28.36 J 4.7 C4A .63 10.45 9.82 C4B 4.0:'1 C4C 1. OY 3 1f 482. 73 61 10.94 -60.66 -----~-- J - -4.04 1.44 .35 --- 210.77 -271.96 GR 7.8 75.4 67.6 5 107.27 58.68 -48.59 131.48' 239.37 107.89 5. 12 39.63 34.51 R 69.9Y 87.4 17.41 'IV 32.07 16.83 -15.24 1.9 38.72 36.82 J '\G 0 --- . ... T - - �319 F i gu RIVER: South FLltt(' MILE: Stratum III r e 5c QUAD: TOTAL AREA: MAP SCALE: Date: Old PhL) t os Sea 1e : Date: Recent: 1704.24 Photos Scale: Cbc c + CIA /, ' ~.~:' CI8 77 •r (, Cle ')(, :.(, C2lL 1{_6 q') 12.75 I ', -25 . :, -2(). \ 7", Iif) 1'1 -86. r: !,(, (:l -4.2 lb. hI -]0. :' ~() . ') 15.5 36.C - {I ,C; C2C ',!', 8') C3A ")[)I! C38 Oil 'IS n.17 C3C '\ nn S', 'J 2().1 1 -7.0 (),29 n. 1 " C48 C4C q '\ " -30. 74 C2B _CAA ( i 7 -.19 '(I .35 -2.8· 369.31 -289 20.01 16. 7: 1 106,82 -23. AG 101-+. 8~ 520.31 0 'i R Btl NV 7. 16 1 I 'J s==- .• ~ H 6')8 41 GR .l" S 130 'J lJ 'J J L T Jq \ (,9 , s: ., 341. 5.99 .6 157.72 68.8 14.86 7. 7 12. II 11.4 �320 Figure SO RIVER: MILE: South Pl atte St r a tum IV QUAD: TOTAL AREA: 947.75 MAP SCALE: Date: 01 d Photos Scale: Date: Recen t Pho tos Scale: Change + or ._ :lA 9.21 11.56 2.35 CIS 6.66 20.25 13.59 ClC 14.45 5.6 ·-8.85 C2A 90.05 101.82 11.77 C2B 76 73 58.92 -17.8: C2C -- 22.37 15.09 _·7.28 C3A 51. 27 39.02 -·12.2: C3B 6.56 28.08 21. 52 -- - 5.38 -5.63 ' 03 .25 -.78 C4C .03 4.02 3.99 H 387.26 262.91 -124 .. GR 44 7? 126.05 81.33 S 127.98 93.66 . -34.2: ft.G 32.35 45.08 0 8.34 14.26 5.92 R 32.47 100.28 67.81 NV 24.77 13.51 -11. 1 I .56 2.0 1.44 '3_C_ C4A 1 1 01 C48 - -J - T ., 12.73 �321 Fi gu re 5E RIVER: MILE: South Platte QUAD: Total MAP SCALE: Date: Old TOTAL AREA: Photos Scale: Date: Recent Photos Scale: Chang + or C· 1n 75.75 62.26 C13 50.43 35.67 CIC 54.77 22.08 - 32. E C2A 399.84 295.05 --104. ill __ 245.06 239.25 -5.8] C2C 114.08 131. 16 17.0t C3A 501.21 436.5 -64. ! (38 174.57 264.53 89.96 _';3C_ 152.52 100.63 -51.2 C4A 19.48 30.21 10.73 C48 7.85 11.7 3.85 C4C 10.96 6.9 -4.06 H 2123.36 1177.12 -946. GR 128.48 392.95 264.4 S 490.8 388.87 -10l. .AG k92.64 1068.46 D 31.58 104.03 R 257.19 481.99 224.8 NV ~6.43 113.01 26.58 84.73 79.37 J -13 .. -14. ; '-- -- -- .. 575.8: __ o_ 72.45 I _~.36 -T �322 RIVER: MILE: Fi gUl'e 6A Ri 0 Grande Stratum QUAD: MAP SCALE: TOTAL AREA: 916.42 ha .-Date: Old Photos Scale: Date: Recent Photos Scale: Change + or - _. CIA .. CIS 6.02 8.36 2.34 1.78 5.92 4.14 --- . elC 0.95 _Q_A 2.95 14.07 . 13.26 C2B - 13.12 9.83 6.88 10.94 -2.32 C2C 9.56 11.49 C3A 22.27 6.31' C38 12.98 12.06 .. l.93 .- ,-._ ......_.- ..15.96 ---~~-,-0.92 · ..lli 35.2 56.15 C4A 0.56 0.37 C4B 1.54 C4C 1.57 7.21 5.64 679.15 457.34 -221.8 GR 23,11 85.91 62.8 S 48.66 34.45 -14.21 H 20.95 - -0.19 -1.54 · - . J --.- •. AG -_.- 2.28 155.41 D 7.92 6.54 R 133.75 29.79 -3.96 NV \q.21 2.75 -6.46 l :3.77 5.96 2.26 ~1____ 1 153.13 ,- -- __ ._-_. -1.38 · -- �323 Fi gu re 68 RIVER: MILE: Ri 0 Grande Stratum II QUAD: HAP SCALE: Date: Old TOTAL AREA: 1577.79ha Photos Scale: Date: Recent Photos Scale: Change + or - 14.61 25.89 11.28 OS 11.26 11.24 -0.02 eIC 16.15 5.51 -le.64 C2A ~ 53.94 52.17 - L ZL_ CZEl 47.37 38.21 -·9.16 C2C 23.44 21.02 ~.~~. ~ '? C3A 50.75 34.15 --1h.f. ~....,_------ C3B 77.46 126.12 L'.8 Jf --~.•. "3C 77.97 91. 27 13 ...1._. CdA 2.8 0.5 -2.3 C4B 1.48 4.19 2.71 C4C 8.44 4.29 -4.15 H 1002.26 759.41 -242.8. 9.22 9.2? 42.61 -32.54 ~ GR 75. 15 S J 259.79 AG .- ..•... 259.79 -_. 10.77 23.67 R 1::1.66 43.08 -36.58 ~IV II). 11 3.39 -5.72 ..S.17 22.26 7.09 0 12.9 i , T �Fi gure 324 sc RIVER: Rio Grande MfLE: Stratum III QUAD: MAP SCALE: Date: Old Photos TOTAL AREA: Date: Recent Scale: 749.63 ha Photos Scale: Change + or ClA 2.53 1.49 elS 1.1 ele 3 n 0.46 -3.27 C2A 7.13 8.67 1.54 C23 12.8 1.53 -11.27 C2C 16.97 3.97 -13.0 C3A 5.48 2.66 -2.82 C3B 2.88 19.56 16.68 ilL 3.06 7.78 4.72 1.26 1.26 -1.04 -1.1 ., .csa. C48 C4C H 527.14 536.8 9.66 GR 4.76 4.74 -0.02 S 86.91 81. 53 -5.38 J . 18.64 AG \1. 20 19.84 D 13.59 2.35 R •S2 .48 45.41 -7.07 NV 14.23 1.08 -3.15 L ;13.64 9.12 -4.52 T - .. __ . _. -1. 24 �325 Figure 60 RIVER: Rio Grande MILE: Stratum Total qUAD: MAP SCALE: Date: Old Photos TOTAL AREA: 3243.84 ha Scale: Date: Recent Photos Scale: Change + or C1A 23.16 ha 18.65 ha -4.51 ClB 14.14 17.16 3.02 CIC 20.83 20.04 -0.79 C2A 64.02 70.67 6.65 _G2_B 73.43 50.68 -22.75 C2C 49.97 36.48 -13.49 C3A 78.50 43.12 -35.38 3B 93.32 157.74 64.42 C3C 116.23 115.20 38.97 C4A 3.36 2.13 -1. 23 C4B 3.02 4.19 1.17 C4C 10.01 11.50 1.49 H 2208.55 1753.55 -455.0 GR 27.87 99.87 72.0 S 2l0.72 158.59 -52.13 .. J 3.48 AG 435.04 431. 56 - ._ .. - 0 22.28 32.56 10.28 R 165.89 118.28 -47.61 NV 22.55 7.22 32.51 37.34 --T I .- -15.33 4.83 -. �326 Figure RIVER: MILE: Colorado Stratum I QUAD: MAP SCALE: Date: lA Old Photos TOTAL AREA: Scale: 114.4 ha Date: Recent Photos Sca1e: Chang + or ClA 0.34 ha -0.32 0.02 11a CIS ..- - -._--_ .. CIC C2A 0.6h _ill_ __ _...:L..li 0.04 -0.62 -._-,-- O. 12 -4.22 C2C .._.- -5.92 -~-~, ..-- C3A 5.g4 0.02 C3B 0.51 0.02 -0.49 0.80 -L 35 ____Cl_k_ ') 1'i __C!L C4B - o -0.54 'i4 C4C I I H 33. 1 1 14.20 -23.91 GR 1 1 68 26.81 15.13 S i5.29 11. 63 -3.66 J 0.5!+ 1. 17 0.63 t-LI. !r I " AG 0.51 19.95 19.44 R 33.08 37.77 4.69 i'IV 1.00 0.42 -0.58 L 0.25 T -0.25 �327 Figure RIVER: MILE: 7B Colorado Stratum II QUAD: 20 - 37 TOTAL AREA: MAP SCALE: Date: Old Photos Miles Scale: 676.63 ha Date: Recent Photos Scale: Ch anj + or CIA IB.68 ha 9.83 ha -8.85 CIB IB.18 8.51 -9.67 eIC 11.54 6.52 -5.02- _Q.A 19.64 49.30 C2B 15.55 C2C 5.94 9.24 C3..I\ 41. 28 29.35 C3B 19.72 B.39 --1l.]] C3C 7.93 9.53 1.60 - -- 29.66 23.26 ----7.71 -.._.~.,._-.•.-- --- ..•. - 3 ~~~n .---..~- -1 J .93 --- .~-~--------.•.. ---- C4A C4B 9.22 -9.22 C4C 9.15 -9.15 H 217.19 232.49 15.30 38.62 11.06 -27.56 S 107.40 122.63 15.23 J 2.34 GR - AG - ' -2.34 ~ I -1.34 '" 1 •. j'-t 24.B4 0 2. 70 27.54 R 55.65 115.34 59.69 11.05 3.28 -7.77 6. 19 6. 19 NV I l .•.I \, ] - ..- _- I • �328 Figure RIVER: MILE: 7C Colorado Stratum III QUAD: MAP SCALE: Date: Old Photos TOTAL AREA: 252.11 Scale: Date: Recent Photos Scale: Char. + or CIA 4.45 ha 3.01 ha -1. 41 CIS 2.77 0.45 -2.32 2.49 2.49 CIC - C2A 6.47 9.94 3. L;7 C2B 9.34 1. 15 -8.19 2.21 2.21 C2C - .•....~ - C3A 5.22 13.83 8.61 C3B A.59 4.73 -1.86 C3C_ 11. 46 -11.4( C4B 2.20 -2.20 C4C 1. 73 H 43.42 GR 8.45 20.97 12.52 S 80.24 81. 03 0.79 C4A - 52.96 J 6.13 AG 9.54 - .. 6.13 -2.15 0 8.26 6.11 R 44.98 39.83 5. 10 NV 12.53 3.27 9.26 L - -0.80 0.93 T --- - �329 Figure 70 RIVER: Colorado MILE: Stratum 4 QUAD: Miles 72-100 MAP SCALE: Date: TOTAL AREA: 10/22/54 Scale: 782.77 Date: Scale: Chang + or - -6.18 CIA 16.22 ha 10.04 ha ---_._- ---_._- CIS 7.21 ha 2.43 ha - ClC 19.81 4.20 C2A 17.43 10.57 - -- ~-----. -6.:36 . ._---- C2B 11. 17 10.04 --I-13 C2C 16.58 17.18 C3ft. 7.86 18.78' 10.92 '313 7.34 22.49 15.15 13.44 2.55 -10.89 1.19 1.38 0.19 0.89 0.89 -4.78_.-- --15.61 __ ---- 0.60 -i------- r - r;3C C4A C48 C4C 1.29 0.84 -0.45 H 196.46 78.34 -118.12 GR 37.56 78.37 40.81 S 118.06 126.66 8.60 -- - . J 170.43 -8.43 10.88 62.59 51. 71 R 91.24 89.81 ~llf 36.01 7.18 2.46 71.42 AG 178.86 0 I' ~ T --- -r--' -- -1. 43 -- -28.83 -- 68.96 _._-- ..•... 1--- �330 Figure RIVER: Co lorado MILE: Stratum Total QUAD: MAP SCALE: TOTAL AREA:1825.91 Date: Scale: Date: 7E ha Scale: Chan; + or CIA 39.69 22.90 CIB 28.16 11.39 -·16.: --- CIC 31. 35 13.21 -18. : ---_. C2A 44.20 r?B 40.40 34.57 C2C 22.52 69.62 47. ..lC~.. .---; C3A 60.30 61.9B 1.68 C38 34.16 35.63 1.47 .•... _._ C3C 34.98 12.88 -22. ._: C4A 1.19 1.38 0.19 C4B 11.96 .89 -11. ( C4C 12.17 1.77 -10. 495.18 377.99 GR . go ') 1 137 21 40.9C S 320.99 341.95 20.9E J 2.88 1.17 AG 180.20 176.56 0 22.35 116.19 69.85 . 1??4 ss 25.6 f -5. 8~ _--- ... ---~ ' ~ . L -117. -1. 71 -3.6~ -., 282.75 -16. ; 1---.-_._-- 1--.-""--" "." H R ---- 93.8[; ._--....- ...__ 57.8e NV 60.59 14.15 -46.£ -- '_'_'--' L 2.71 77.61 -- ._--- T 74.9C ~'------ �Divis ione l Correspondence . fr:h~~f~},~L .i.V~:J~~~\;~, . ... ,. Only ..~':'~::;.2~~f;~;[tl~;( "" ','. "'::,1:').;/,',. ~~~.~ :\:~~i~-:;:,~)~;· :',..'~~~" ?f;Jt ~~m;~;(>,:: "STATE 'OF COLORADO .,.~ c'~~ :~l~;J:~ J,<r::~;;~'~lr,l)j~~;l-i,it)t: .•'_'.4$~'i-.'~~i$~~; '~~:; c ; .' ".,;i,'."";,',~,,:,,l.', P)'v t?,I:QN. /!' I L D L I F.,.~.~:..~~:i~:~ii~'i~r;F~~:;<;)L~::, "·~':·}·;iii: 0.,0 F " .-')~~ OEPARTMENTOF NATURAL '/"~~\",J1,;~/'- :L~<;.;.,··.;(".;~' '. -, ";~" ';;'. RESOURCES .;' -, TO: "',' ,.. ... , FROM: '~". ,I. ;./~ ••.• "\,- .. ' .. :,,' I·"~·' ,,~. ,~~'•.:;.,."-t'fl~,r7..::;.r . ..,..c ,~~ ''''..:,''<'~'..~ "; '. ..;..... ":.,".\.::,:'l~ SUBJECT: Analysis . .',>: ~,' Warren Snyder" and -eTa,11 tt., ',E:"iBraun, 1'(.!"i:;11"~'-:,A't~~(;i':'.M~,-1.I(I~:~t, ~"••. ''', •...:~!. "*~~~~r~.~I:,t~'1·~\rl~:. .•. it:.~~11r1 .••1~ .~ . ~~ ..'(\3~~· , ,," ,: _ f/.. r7 ,:';";,: !~;,,-:;:, ':"," ~ :':~\' ':,;'~ :i~'d;:{ i~" t~'1f."·I: &0'''-;. l"'i. \,.,rl\¥'&:;tl-,~}'''~iI·~.¥ ••I,,' ~ t· " ~ ; ~,1.. . .''''.' Iotl'~'l{);~,- ..•. ~. (~~ '~Ir~ ·:i','l. Forest Service ;;, ,:,.. '1~' I,\,C'~ ~b! \~'fi·: .'$ .;, '.".;~'~"':,~,"'\~"'H'i,,;.:!\"':'·"h)f.,l?tlb.~,~:.:, .. :,\r.(i\~,_,~.~ ••.••• ,.~~. of. Colorado State .",-~. -','.','.i,:",.'<· .. ·i··:.",:·:~- ",~ •. :,~:;>"y':j\~!\"\N.~' f .. )~, ' ':~:. "~~~:v }\' ." f;~~i~~Ji~t~~~~ ~~~?, ~/ ~~) ... : '.: Regional Hanaze r s " iiii"!-!; ':-it .. "":',:1, ~,,!i"', "i .:.: :~K·~:,ct,>1tl,,1" '1't'-,C--~~'p "f;~ /.:' ,.>~t~::J~j~~~~f~)~:···,i~~t~J;,',~\i:"~:~' ':~i ~;:~:;;, ~,~;: ~~'b'~;~;~:~ ~ - .~_,~,\:~ : ~~~~,.'.::'i,l,~,'~:::,:;.~J,.,i~,,~.~,~:::,:},:,~,l:,;,~,::',~,:.',),.iij,~.~~~).!;,},·~,',:",.",'.:.i,:,:~,,~.:,".~~,;~j,;,l.·~.:,~,';\._:'~.'-.;,:~<,:,·.i~;!r;;·:},::, ~:.~'.~',-.;~: .'.' t.,~,",.t~lt~'~·I"':'t .•• t",t:tt ,v~ l~,f;""l:.~=-t:~.r.. "'r.~!':'_ I ., kiY1 !..'f.'i1.•J· •..•d~ . \. , .. ..j .,s..:c;... _'.j :':"~""~.l ,I' ..." .~I -. .,•. ~.~ .•• ' y. .)~.!"\~ J" J '. , "":....6', ti(~;j:~~ Report o{i Cottonwoods' .,:'\' ,,~.,.;~~' ..} {·i!'~t,.•:~ , .. !~~ 'f;/,.d$~jlf~;;iji'1~~11;t~~ .~ :by'·'th~"t~i~~·~cid't,s~ift~;,: f}~:,~;\r! ~ ,n; k'bd.:~'f "~~~ly'~is':of the tabular data p~ese~ted Forest Service on the Cottonwood photo interpretation contract" has beer;:;,~~~~~1:~<',tJ,{ ~::~ preparE;d. , The data presented in· this analysis indicate that there are .::f,-1!,')(, "", " .( some changes occurring in the cottonwood type (lack of reproduction;~::'\j2,'~~~~);' ..fr' opening of stands, increa~es in shrubs, etc) .. ;Weobvious~y ShO'ul~".9~ -;.:i~:~:~·;'~;:_:.\,/;_,~h~,, .( concerned with the data from the South Platte. We further anticipate ,ii,{!/?ti·bi.)t:iV' , 'J that the data from the Arkansas and Republican drainages will Jl).imic.'.- ;:>:\. ''i,,:::,;.,> •..... those from the South Platte. Once these data are available from this yearrs···t~,!" contract with the Colorado State Forest Service, we will- make them :;,:r..#2.~~~/.j!i:"I~~'i~:~\\i~> i'\ rl", " a~~i~,~:~b,le. Please make this report .;',<;}::;S~'~'~~N;:~:;:~f~~~~~;:;~ _.~~;;)" :',:;. '~~'>D;:: \. ',;;' to the nongame.and 'habitat' available :::;!~ your region, •. R. Hopper , .- -, ...: , .Y;.'~ ·."" .. ,-: "",,3:~~,~!r~f~~~ \,'~;~~t~t~ • biologi~ts'\ .;;_.,\:;. . !.' tIl' II" ',t.-",'''. ,~I.ltif.. ,:,~~/" -i~<'¥" 'i,r"".t,rr,(,..;:, l~_' if,,,, ,",.'. : ~. ~:~~'" ~h~ .~:~~ ;;\)!~/~;'~·t~ ~~{~'\~~~~:::~'.;);' {'t'· ~..~: G ~ Miller J Torres . R: Towry " ~', .r .!,l,'., } '.1"' f· • '';~~' "1, {,' ;~;r;~;~; :~.~.~~:, ... ,~''!: .0;". '~~·lr,;,.q~~V'·J ~i~"~ ,.r....;. ...1I'f,... ~"'>;.I~ •• , " . _"~\:V"', I·~;.( I ,"?':":':;/( ';:,?:;/t:~;!~,.'~:;;;\!J~t;,:~~:·;~;;~4;~~;~ (i::'i.;:t~~,~: :::'!:'~~.( L\?~:_.:'~)~'1': ',.t, ," " ~,),;~}·.;i'>\:~'v:;'~';-il~"i.Vi.:it1!~~:-;~.f:~yr; . ·t~,,:.~~$. ;:~l.l\·;,'<;l<'4(:;;';' ~,r;~:<;;~, '., .'.'; ,;,.1 ;. " :,"', .•.' ~'. :..,.... ' ... '.~ ...':./,'«:( !:~~:~.;..;' ...: ,..; " "', ,- ':.: J: i i):·.·::·.:t:. '., ;.>. ', . ~ . ,-. .. . '.';"": . ~ ~:.' " .~::;:t~i (_ OOW-A-F·8 '. ...•..•... ~, . "'Zlr.,~~~r:fi!l~i,;,'_".1.'..,•,.~;'..',tf,',:,:.~,:,:,1,:,[,f.;.1~i '~';;"~J,:;::_;.~~~,)!i~:' .;,-.: )~~J§f'::>~~>.:'" or "t ,".._ �332 1:~TERn1 COTTONhlOOD IWENTORY Warren REPORT D. Snyder The Colorado State Forest Service was contracted to use aerial photo interpretation to quantify the condition and changes of cottonwood stands, other vegetation, and land use along major river drainages in Colorado over an approximate 30-year span. Inventory has been completed along the South Platte River from Greeley to the Nebraska line (104 mi.), along the Colorado River from Glenwood Springs to the Utah line (104 mi.), and along the Rio Gr ande River from South Fork downstream through the San Luis Valley (73 mi.). Inventory is continuing along the lower Arkansas River and a small segment of the South Republican below Bonny Reservoir. This report surmnarizes preliminary findings obtained on the 3 drainages completed to date. Statistical analysis has not been conducted. South Platte River An inventory of 5,4!!5 h3 of riverbottom within 29 linear miles (17.7%) was completed using stratified random sampling. Original year aerial photos were mostly Crom 1941 (a few from 1948 and 1949) and recent photos were from 1979 (2 from 1978) yielding an average time span of 36.4 years. The change in area and proportion of sampled cover types is surnmarLz ed in Table 1 arid shows a modest decline in total area occupied by cottonwoods. Shrubs, which occupied less than 10% of the inventoried area, showed a more dramatic decline as did hay meadow. Agricultural farmland, grassland, and river channel and sandbars showed major increases. Cottonwoods.:. 6 in. dbh showed the most marked decline in area (33.6%) while those up to 30 in. dbh decreased less (Table 2). Sites dominated by trees> 30 in. dbh, which comprised only 2-3% of the tree-occupied area, increased 28.2%. Thus, the shift from young to progressively older deteriorating stands of cottonwoods was pronounced. Stands with < 35~~ and > 55~s canopy cover decreased whereas those in the Lnr.ermed i a t « stand densities increased. Colorado River An inve!1tory of 1,826 ha of r i ve rbo t t cm within 21 (20.2%) linear randomly selected miles was completed. The original aerial photos dated from 1951-57 and recent photos were from 1978-80 yielding an average time spand of 25.1 years. Cottonwoods occupied less than 20% of the inventoried habitat and showed a modest decline in occupied area (17.4%) during the time span (Table 3). Shrubs occupied a near equal proportion of the land and increased slightly during the interval. Since tamarisk is a major component of the shrub community along the lower Colorado, the increase ~r:sh rub s should be viewed ,.. dth interest and concern. Prop'o-rtions and ~hanges of other occupied cover types are summarized in Table 3. Major declines in quantities of cottonwood < 6 in. dhh and> 30 in. dbh ~ere noted during the time span (Table 4). Little change in the 2 middle ;tge-cl3sses was noted. Cottomvood stands tended to become more open as indicated by an increase in area of < 35% canopy cover and a decrease �333 of higher density stands. Although young age-classes of cottonwoods declined, comparison of stands with those in the other 2 drainages show a much higher proportion of young trees along the Colorado. In recent photos, 16.4% were < 6 in. dbh along the Colorado, in contrast to 10.4% along the Rio Grande, and 7.3% along the South Platte. Rio Grande River Twenty miles (27 .4~O of the Rio Grande River we re inventoried representing 3,263 ha. Initial photos were nearly all from 1941 (1 from 1944) and the interval of recent Jates ranged from 1973 to 1983 yielding an average time span of 36.3 years. As along the Colorado River, cottonwoods occupied < 20% of the sampled riverbottom, but increased by about 9% in total occupied area during the interval. Shrubs occupied a much smaller proportion of the land (Table 5) and decreased in quantity during the interval. Hay meadmv, the major component of inventoried land, was partially converted to farmland. Quantities and changes of other inventoried lands are presented in Table 5. Co t t onwo o d tree age composition appeared to be relatively stable over the interval (Table 6). Area of ttees < 6 in. dbh and > 30 in. dbh were stable and losses of stands in the 6-15 in. dbh range were compensated by those in the 16-30 in. dbh range. Low density stands decreased in quantity whereas those in the 35-55% canopy cover and > 55% canopy cover ranges increased (Table 6). Lack of cottonwood regeneration does not appear to be an important problem along the Rio Grande River at present. To summarize, t he primary concern lies ,.. lith deterioration of both cottonwood and shrub plant con~unities along the lower South Platte River. High water during 1983 caused extensive regeneration of trees, shrubs, and vines potentially at least temporarily reversing this trend. However, continued efforts to construct the Narrows Reservoir and other dewatering efforts along the South Platte in the future must be viewed with concern. Preliminary findings of this inventory raise many questions that presently cannot be answered. Future research and monitoring will address the important questions and will permit better management of these vital riparian habitats in Colorado. �334 Table 1. Proportions and changes (in hectares) of vegetation types over a 36-year interval along the South Platte River, northeastern Colorado, 1941-79. Vegetation Early interval ha % type Cottonwood Shrub Hay meadow Grassland Agriculture Developed River Unvegetated Standing r..:ater Totals 1,806.5 490.8 2,141.0 128.5 '~92.6 31.6 257.2 86.4 5.4 5,440.0 33.2 9.0 39.4 2.4 9.0 0.6 4.7 1.6 0.1 Recent ha Change (ha) interval % 1,638.8 388.9 1,177.1 392.9 1,068.5 104.0 482.0 113.0 80.8 5,446.0a 30.1 7.1 21.6 7.2 19.6 1.9 8.9 2.1 1.5 -167.7 -101. 9 -963.9 +264.4 +575.9 + 72 .4 +224.8 + 26.6 + 75.4 8Differences between early and recent totals apparently occurred during planimetering of vegetation types and are not believed important enough to warrant correction. Table 2. Proportioll and changes by age-class and canopy cover (in hectares) of cottonwood stands over a 36-year interval along the South Platte River, northeastern Colorado, 1941-79. Age-class (in. dbh) Canopy cover <6 <35 35-55 >55 Subtotals 6-15 <35 35-55' >55 Subtotals 16-30 <35 35-55 >55 Subtotals >30 <35 35-55 >55 Subtotals All ages Totals <35 35-55 >55 en Early interval ha % Recent ha 75.5 50.4 54.8 180.9 10.0 62.3 35.7 22.1 120.1 42.0 295.0 241.8 131. 2 668.0 45.9 436.5 264.5 100.6 801.6 2.1 30.2 12.0 6.9 49.1 399.8 245,1 114. 1 759.0 501. 2 174.6 152.5 828.3 19.5 7.8 11.0 38.3 996.2 477.9 332.fl 1,806.5 824.0 554.0 260.8 1,638.8 interval al Change ha /0 al i, - 13.4 - 14.7 - 32.7 7.3 - 60.8 -33.6 40.8 -104.8 3.3 + 17. 1 - 91.0 -12.0 48.9 - 64.7 + 89.9 - 51.9 - 26.7 - 3.2 3.0 + 10.7 + 4.2 4.1 + 10.8 +28.2 .----:.. 172 .2 + 76.1 - 71.6 -167.7 - 9.3 �335 Tab.:e 3. a 25-year Proportions and changes (in hectares) of vegetation types over interval along the Colorado River, western Colorado, 1955-80. Vegetation Early ha Cottonwood Shrub Hay meadow Grassland Juniper Agriculture Developed River Unvegetated Standing water Totals interval % Recent ha 20.7 17.6 27.1 5.7 0.2 10.1 1.2 12.9 4.3 0.2 377.5 320.9 495.2 104.3 2.9 184.3 22.3 235.6 78.0 4.5 1,826.0 interval 311.6 341. 9 378.0 137.2 1.9 170.8 108.8 282.7 14.5 77.6 1,825.0a % Change (ha) 17.1 18.7 20.7 7.5 0.1 9.4 6.0 15.5 0.8 4.2 :_ 65.9 + 21. 0 -117.2 + 32.4 l.0 - 13.5 + 86.5 + 47.1 - 63.5 + 73.1 aDifferences between early and recent totals apparently occurred during planimetering and are not believed important enough to warrant correction. Table 4. Proportion and changes by age-class and canopy cover (in hectares) of cottonwood stands over a 25-year interval along the Colorado River, western Colorado, 1955-80. Age class (in. dbh) <6 6-15 16-30 >30 :'.11 Ages C2nopy cover (%) Early ha <35 35-55 >55 Subtotals 39.7 28.2 3l.3 99.2 dS J5-5S >55 Subtotals 44.2 40.4 36.0 120.6 <35 35-55 >55 Subtotals 58.3 39.0 35.0 132.3 <35 35-55 >55 Subtotals 1.2 11.9 12.2 25.3 <35 35-55 >55 Totals lld.4 119.5 114.5 377 .4 interval % Recent ha 26.3 26.4 11.4 13.2 51.0 32.0 69.8 34.6 28.6 133.0 35.0 62.0 21.0 40.5 123.5 6.7 1.4 1.8 0.8 4.0 159.6 68.8 83.1 311. 5 interval Change % ha % 16.4 -13.3 -16.8 -18.1 -48.2 -48.6 42.7 +25.6 - 5.8 - 7.4 +12.4 +10.3 + 3.7 -18.0 + 5.5 39.6 - 8.S .' + 0.2 1.3 - 6.6 -10.1 -11.4 ::.-2"1-;-3-84.2 +16.2 -50.7 -31.4 -65.9 -17.5 �336 Table 5. Proportions and changes (in hectares) of vegetation types over a 36-year interval along the Rio Grande River, San Luis Valley, Colorado, 1941-83. Vegetation type Cottonwood Shrub Hay meadow Gra.ssland Agriculture Developed River Unvegetated Standing water Totals Early interval ha % Recent ha 17. 1 6.5 67.7 0.8 0.1 0.7 5.4 0.7 1.0 559.8 210.7 2,208.5 27.9 3.5 22.3 175.7 22.5 32.5 3.263.4 611.2 158.6 1,753.5 99.9 435.0 32.6 118.3 7.0 37.3 3.253.8a interval % Change (ha) 18.8 4.9 53.9 3. 1 13.4 1.0 3.6 0.2 1.1 + 51. 4 - 52.1 -455.0 + 72 .0 +431.5 + 10.3 - 57.4 15.5 + 4.8 aDifferences between early and recent totals apparently occurred during planimetering and are not believed important enough to \.arrant correction. Table 6. Proportion and changes by age-ciass and canopy cover (in hectares) of cottonwood stands, over a 36.S-year interval along the Rio Grande River, San Luis Valley, 1941-83. :\ge clClss (in. dbh) ,6 6-15 16-30 >30 All ages Canopy cover (%) Early interval ha % <35 35-55 =--55 Subtotals 23.2 14.1 20.8 58.1 <35 35-55 >55 Subtotals 70.5 73 .4 50.0 193.9 <35 35-55 >55 Subtotals 78.5 96.6 116.2 291.3 <35 35-55 >55 Subtotals 3.3 3.0 10.0 16.3 <35 35-55 >55 Totals 175.5 187. 1 197.0 559.6 Recent ha 10.4 35.7 17.2 10.6 63.5 34.7 70.7 50.7 36.6 158.0 52.0 172.4 155.2 370.7 2.9 2.1 4.2 12.7 19.0 Interval % Change ha % 10.4 +12.5 + 3.1 -10.2 + 5.4 + 9.3 25.8 + 0.2 -22.7 -13 .4 -35.9 -lS.5 60.7 -35.4 +75.8 +19.0 +79.4 +27.2 3.1 - 1.2 + 1.2 + 2.7 + 2.7 +16.6 -23.9 +57.4 +18.1 +51.6 + 9.2 Id.l 151.6 244.5 215. 1 611 .2 �337 APPENDIX B FINAL REPORT Colorado State Forest Service Contract, 1984 ��339 AERIAL PHOTO INTERPRETATION PLANIMETERING AND OF RIPARIAN VEGETATION OF COLORADO (ARKANSAS AND REPUBLICAN RIVERS) By Daniel Teska Thomas Owens Colorado State Forest Service Fort Collins, Colorado For Colorado Division of Wildlife June 29, 1984 �340 L:;TRODUCTIO:l' During the past few years Division have noticed, cottonwood of Wildlife (DOW) researchers in the course of other work, an apparent reproduction "lands bordering in riparian resources in Colorado. fresh water bodies and the resources .-1chmi dt in ~Ianagement of Cottonwood-Willow (Riparian: they support", Riparian Associations i:,llurado.)Since riparian areas have more wildlife l)ther in Colorado lack of in species than any (Graul, Ibid), this could be a serious trend. U1e DOW requested Shop do a vegetative the Colorado State Forest Service (CSFS) Map survey using aerial photos to document the actual trend of riparian vegetation CSFS personnel working in Colorado. on this project were Dan Teska, Jerry Bresnan, and Diedre Thiebaud. with Thomas Owens as supervisor. AREA DESCRIPTION The Arkansas and South Fork of the Republican five rivers chosen to be mapped (the Colorado, Grande Rivers were mapped earlier). rivers in Colorado The Arkansas Rivers were two of South Platte. and Rio The Arkansas is one of the largest and contains vast riparian habitat. River is located in central to southeastern The study area starts on the west edge of the Nepesta Colorado. 7~' quadrangle. 22 miles east of Pueblo and runs 162 miles east to the Kansas border. The western end has an elevation to an elevation of 4440 feet above sea level and drops of 3350 feet at the Kansas border, Colorado and its easternmost point. The Arkansas, the lowest point in like the South Platte River, is in a broad ge~tle valley which contains a broad band of riparian vegetation cottonwood along the river. The major cottonwood (Populus sargentii); tamarisk (Tamarix chinensis) major shrub species along the river, but willow also very prevalent. species is plains (Salix~) is the is �341 The South Fork of the Republican Colorado, near the Kansas border. Reservoir in Yuma County, proceeding River is located in northeastern The study area is below Bonny three miles west from the Kansas state line (located on the Hale Ponds 7~' quadrangle). averages approximately th Republican cottonwood The elevation 3550 feet above sea level. The South Fork of is in a gentle valley. Plains cottonwood is the major species. METHODS 1. Determined the availability aerial photography and type of 9"x9" vertical stereo of the rivers from the ASCS Photo Field Office in Salt Lake City, Utah ( the holder of USDA photos). 2. A prepared river miles map of the Arkansas (with 0 river mile being the upstream and latest photography earliest the Republican greatest available and latest photography separation River (see Figure 1) shows the end) and the earliest (a 25~year separation between was the minimum difference used). On a 7~' quad was used to show river miles; a l~~ye~r between the earliest that could be obtained 3. The DOW randomly on the Arkansas, selected and selected and latest photography was the on this river (see Figure 2). 22 river miles and one non-randomly 3 river miles on the South Fork of the Republican. 4. Aerial photos were ordered 5. A vegetation classification for the sample river miles. was developed in discussions between CSFS and DOW (see Figure 3). types were delineated using Bausch & Lomb SIS 95 and Old Delft Scanning Stereoscopes on acetate overlays 6. Vegetation Stereoscopes using drafting pens. �342 ~ ~ ~ ::> \A.l H V1 ~ Vl Z. "«. :ltc:: § 0- - ~ \u ...J ~ ••... ••• i I �~ -.;_ .-_ "" I I ~, ::. -:::---- . ~/..' '.... q,.r(' .--._ ''r-.. .-' ~ ~-e-~~ ':/ , ~ _'--. '~:".. /j. -:-- ~ ~ "'\... -, i- -._) "•. ,...__ �344 Figure 3 DOW -- Cottonwood Classification Hay Meadow/Emergents H Grassland GR Upland Shrub Community (Xeric Shrub-Grassland Wetland Shrub Complex & Mesic Shrub Complex) ) ) ) S Agricultural AG Developed D Cottonwoods C Size Classes: 1 6" dbh 6"-16" dbh 16"-30" dbh 30" dbh Crown Density 2 3 4 Classes 10%-35% 35%-55% over 55% A B C River R Unvegetated/Sandbar (Bare soil, sandbars, NV gravel pits) Lakes. Ponds, Reservoirs Classes to be expressed L in hectares, E.g., C2B is cottonwood, 6"-16" dbh, 35%-55% crown density class. �345 7. Field trips were taken to verify delineation accuracy on both rivers. 8. Delineated vegetation types from the old photos were transferred & Lomb Zoom Transfer Scope. The onto 8!:l"xll"paper using a Bausch recent photography was transferred the old vegetation types. Details which had remained intersections onto mylar overlays were used as controls to properly on top of such as road register the maps onto each other. 9. Areas were plaimetered Areas were measured hectares using a Numonics in hectares. Electronic A form was devised Planimeter. to record for each type on the map. 10. Totals for hectares for each type and date on each river were calculated. RESULTS Overall in Figures Arkansas areas for each river and each date are presented 4 and 5. An inventory and results of shrubs was taken along the are shown in Figures 6, 7, and 8. Specific maps and area figures are presented in Appendix B. DISCUSSION The aerial photointerpretation, phases were routine and presented mapping and planimetering no special difficulties. The specific dates and types of photos used can be found with each map in Appendix B. All photography ~hotography used was black and white. was 1:20,000; ~;40,000. On the Arkansas, scale for the recent photography photography was dates for the old photos ranged between 1940 to 1950; recent photography Fork of the Republican, Scale for the old was taken in 1980. On the South old photography was taken in 1980. was taken in 1961; recent �346 After extensive determined species field checking from the photography approximately the same height interspersed. With black and white difference between An inventory with zone. Sample available. photos the two vegetation of shrub species types. Therefore, tamarisk for interpretation was purposes, was taken along the Arkansas dominance 1/100 of an acre and results were taken as follows. shrub areas on the maps interpreted I, 2 •• 0., no Areas in in the riparian can be found table. Points within also randomly determined was noted. the number I and II sampling was denied. (3) A sampling of points errors area were the desired I and II was not achieved. error of 20% was taken. However, due to the is privately owned, of 25% and 23% were reached, since it was not possible Therefore, selected the radius of that area was fact that most of the area along the Arkansas respectively, were were then chosen with a each randomly at each point was taken. Each shrub inside used to determine photography the (2) A fixed plot sample of 1/100 of an acre 0 and the species (1) In each stratum, with recent to be sampled random number Strata are there was no visible the extent of tamarisk plots were Sample points on Strata Both are 6, 7. and 8. in Figures counted, shrub the DOW. 1984 to determine numbered it was and size, and in some instances in the shrub classification after consultation June, truthing, that it was impos~~?_1~_!:9._distinguish between__willow and tamarisk included and ground to survey points where access number of sample points on �347 APPENDICES Appendix A Photointerpretation Key Appendix B Maps and Area Statistics �Appendix 34 ~ A Pho t o Ln t erp re t a t Lon Key for OOW Cottonwood Hay Meadow/Emergents Study - H inis type covers one of the largest areas. Hay meadow/emergent wetlands appear a dark shade of gray on black & white photography, indicating a lot of moisture. this class on the photos. Grassland There is a smooth textue associated Confidence level high. - GR Grassland has a very light tone of gray to white on black imagery. The texture is smooth, but slightly and not as wet. Confidence Sh rub Community than hay meadows, level high. (sages) are found mixed with grasses cover percentages surrounding rougher from crown of near 0 to over 50%. It is darker than the grasses and has a mottled "salt and pepper" texture. Mesic shrubs are found in narrow draws, along streambeds, on floodplains light gray signature, Tamarisk and willow produces a billowing On B & W photography, shrubs have a and a more even texture than cottonwoods. are the main species in this class. This type effect on the photo. - AG Agricultural They are easily ture is circular lands are found on level to gently sloping areas. recignized by cultivation marks. Center pivot agricul- (or nearly so) on 160 acres. Flood irrrgated ture is found along streams, with an irrigation discernable. on islands, of rivers, and at the foot of slopes. Height can be under stereo viewing. Agricultural & white - S Xeric shrubs detected with Signature agricul- ditch sometimes is light to dark gray. Confidence level high. �349 Developed - D Paved roads have a dark appearance. Buildings, soil are visible on ranch yards. Excavated trees and disturbed areas have a signature similar to bare soil (bright), but are clearly dug out. Maps provide collateral data. Confidence level high • -------.-~,,_ .. . River - R Rivers are obvious. along major and minor channels; to canals. They have a dark signature also apparent, depending on B on reflectance same applies & W photos. White areas of film. Confidence level high. Non-vegetated/Sandbar - NV Usually found along river beds and islands. and highly visible. Confidence Signature is white level high. Lakes, etc. - L This type also obvious. Very dark signature. Confidence level high. Cottonwoods - C On B & W photography, cottonwoods signature have a "ghost-like" For size classes, is light gray. On late fall imagery appearance (leaves gone). crown size is direclt proportional to trunk size -- the larger the crown, the larger the dbh of the ~ree. Density classes are distinguished by spacing of trees and crown cover. Over 55% density class, for example, 75% crown cover on the photo. appears to have over �3.50 Appendix B River Miles Mapped River miles were measured mid-channel. Thus a sinuous more river miles Arkansas with an electronic planimeter river such as the Arkansas, in has many in it than might be expected. River Stratum I II III Non-random River Miles 004. 008, 012, 015, 030 045, 047, 053, 055. 058, 059 081, 082, 089, 095, 106, 129, 136. 143. 150, 089A South Fork of the Republican 1R, 2R, 3R Shrub Inventory ISS, 162 �Figure 4A 351 RIVER: Arkansas MILE: Stratum I QUAD: }1AP SCALE: Date: Old Photography TOTAL AREA: 1036.04 ha Scale: Date:Recent Scale: Photography Change + or - CIA (7) 6.03 (12) 31.76 25.73 CIB (7) 6.26 0 -6.26 CIC (6) 12.83 (2) 1.86 -10.97 C2A (26) 27.42 (24) 76.66 49.24 C2B (16) 39.28 (11) 0.46 - 39.74 C2C (8) 16.06 (5) 12.43 -3.63 C]A (30) 88.75 (22) 61.72 -27.03 C38 (25) 69.45 (l0) 24.41 -45.04 C3C (15) 26.26 (1) -22.19 C4 (8) 7.42 (2) 2.31 -5.11 -< 4.07 - C4B (8) 12..8 (2) 3.0 -9.8 C4C (1) 1.27 (1) 0.62 H (15) 236.08 (14) 223.06 -13.02 Gft. (17) 93.38 (23) 122.46 29.08 S (77) 225.52 (31) 251.85 26.33 AG (2) 13.06 (9) 57.26 44.2 D (3) 2.37 (5) 7.95 5.58 (7) 72.57 (8) 90.4 (35) 79.03 (4) 2.56 (1) 0.20 (4) 20.65 R NV L 1.89 . 17.83 --_._--- .. -76.47 20.45 �Figure 4B 352 RIVER: Arkansas MILE: Stratum II QUAD: MAP SCALE: TOTAL AREA: 1011.6 ha Date: Old Photography CIA Cia CIC Cll -. C2B C2C C3A C3B C.3C C4A C4B C4C H GIl S AG D R NV L Scale: 1:20,000 B&W Date: Recent Scale: 1:40,000 Photography B&W Change + 011:' - (3) 3.6 (4) 2.3 -1.3 (3) 2.8 0 -2.8 (1) 0.5 0 -0.5 (22) 17.8 (12) 18.6 0.8 (9) 8.8 (4) 8.5 -0.3 0 (2) 3.9 3.9 (43) 52.0 (29) 64.0 12.0 (22) 20.8 (9) 14.6 -6.2 (7) 13.1 6.6 0 -4.0 (9) 6.5 (5) 4.0 (6) 8.6 (1) 0.9 -7.7 (5) 1.9 1.5, (23) 453.5 (17) 337.2 -116.3 (23) 83.1 (23) 91.2 8.1 (70) 195.0 (46) 227.5 32.5 (1) 3.0 (1) 95.4 92.4 (31) 31. 7 (21) 55.4 (7) 79.6 (6) 71.8 (19) 39.0 (2) 1.9 -37.1 (2) 0.9 (1) 2.5 (l) 0.4 " 3.4 . 23.7 -7.8 - ~ �353 Figure 4e RIVER: Arkansas MILE: Stratum III MAP SCALE: Date: Old Photography CIA QUAD: TOTAL AREA: 1529.2 ha Various Scale:l:20,000 B&W Change + or - (3) 3.9 -5.0 0 -2.1 0 0 0 (29) 65.5 (14) 24.8 -40.7 C2B (12) 62.9 (3) 6.5 -56.4 C2C (9) 10.0 0 -10.0 (16) 48.5 (11) (8) 13.4 (4) 12.6 -0.8 ClC (2) 6.0 0 -6.0 cu. (3) 3.9 (2) 2.8 -1.1 0 -3.8 (l) 0.2 0 -0.2 (13) 81.0 (10) 110.8 29.8 (56) 378.8 (36) 281.8 -97.0 (80) 464.7 (46) 658.1 193.4 (6) 17.9 (13) 273.6 255.7 (5) 2.5 (3) 9.3 6.8 (13) 69.8 (14) 72.3 (71) 288.2 (23) 48.8 (6) 1.1 (2) 0.3 CIB CIC ell C3A C3K ClJD C4C II GI. S Ai; D a. NV L (9) 8.9 Scale: 1:40,000 Date: Recent Photography B&W (3) (3) 2.1 3.8 <, -24.9 23.6 . 2.5 __ . .-. -239.4 -0.8 - �354 Figure 4D Arkansas RIVER: MILE: Strata Total QUAD: MAP SCALE: Various Date: Old Photography CIA CIS CIC Cll C2B j C2C ClA C3B C3C C4A C4B C4C II GR S laG D R. NV L TOTAL AREA: Scale:1:20,000 3598.62 ha Date: Recent Scale:1:40,OOO Photography Change + or - (19) 18.53 (19) 37.96 19.43 (13) 11.16 0 -11.16 (7) 13.33 (2) 1.86 -11.47 (77) 110.72 (50) 120.06 9.34 (37) 110.98 (18) 54.74 -56.24 (17) 26.06 (7) 16.33 -9.73 (89) 189.25 (68) 149.32 -39.93 (55) 103.65 (23) 51.61 -52.04 (26) 38.76 (8) 17.17 -21059 (16) 15.32 (4) 5.11 -10.21 (3) 3.9 -21.3 (3) 1.87 (6) 3.79 1.92 (51) 770.58 (41) 671.06 -99.52 (96) 555.28 (82) 495.46 -59.82 (227) 885.22 (123) 1137.45 252.23 (9) 33.96 (23) 426.26 392.3 (39) 36.57 (29) 72.65 (27) 221.97 (28) 234.5 (125) 406.23 (29) 53.26 -352.97 (9) 2.2 (13) 24.35 22.15 (17) 25.2 "- . Numbers in parentheses indicate number of areas for each class. 36.08 12.53 - �I:..LOULt:..:. 355 J RIVER: S. Fork Republican QUAD: MILE: Total MAP SCALE; Date: Old Photography CIA C1B CIC CZA C2B - C2C C]A C3B C3C C4A C4B C4C H GR. S AI; D R NV L TOTAL AREA: 526.51 Various Scale: 1:20,000 Scale: I:40,000 Date: Recent Photography Change + or - (5) 5.63 0 -5.63 (3) 1.85 0 -1.85 0 0 (18) 22.22 (B) 42.21 19.99 (9) 17.41 (4) 21.48 4.07 (2) 3.6 (1) 2.0B -1.52 (18) 25.2 (13) 24.56 -0.64 (15) 26.04 (14) 41.02 14.98 (7) 23.95 (9) (1) 1.27 0 -1.27 . (1) 8.09 6.82 0 (1) 2.63 2.63 (1) 0,87 (1) 6.B4 5.97 (16) 288.73 (17) 168.02 -120.71 (17) 21.75 (6) 37.17 15.42 (5) 60.42 (5) 113.21 52.79 0 0 0 (5) 12.3 (6) 7.07 (10) 14.0 (4) 5.69 0 0 0 (l) - 1.27 .•... 22.49 46.44 . -5.23 . --- -8.31 0 �356 Figure 6 Stratum I tl Shrubs SEecies 1 3 Tamarisk 2 9 Tamarisk 3 1 Tamarisk 4 15 Willow 5 20 Willow 6 1 7 3 8 1 Willow 1 Tamarisk 9 8 Tamarisk 10 23 Tamarisk 11 22 Tamarisk 12 24 Tamarisk Pt It 16.7% - Willow 75.0% - Tamarisk 8.3% - Mixed Mean - 10.92 shrubs/pt Standard deviation - 9.31 Standard error - 2.69 Limit of error - 25% Coefficient of variation - 85% - Tamarisk Tamarisk �Figure Stratum 357 7 II Pt 1/ II Shrubs Species 1 19 Willow 1 Tamarisk 2 24 Tamarisk 3 1 Russian 18 Tamarisk 4 11 Tamarisk 5 17 , Olive Tamarisk 8 Hawthorne 6 62 Willow 7 3 Tamarisk 8 17 Tamarisk 9 12 Tamarisk 10 5 Tamarisk 11 17 Tamarisk 12 11 Tamarisk 13 2 Tamarisk 14 2 Tamarisk 15 4 Tamarisk 16 10 Tamarisk 17 4 Tamarisk 18 9 Tamarisk 19 6 Tamarisk 15.7% - Mixed Mean - 13.84 shrubs/pt 78.9% - Tamarisk Standard deviation 5.4% - Willow Standard error - 3.16 - 13.79 Limit of error - 23% Coefficient of variation_-=-_29.6% �358 Figure 8 Stratum III Pt II II Shrubs SEecies 1 21 Tamarisk Russian Olive 3 2 16 Tamarisk 3 20 Tamarisk 4 16 Tamarisk 5 17 Tamarisk 6 24 7 35 Tamarisk .8 33 Tamarisk 9 18 Tamarisk 10 14 Tamarisk 11 18 Willow 12 17 Willow ., Tamarisk 8.3% - Mixed 16.7% - Willow 75.0% - Tamarisk Mean - 21.0 shrubs/pt Standard deviation - 6.8 Standard error - 1.96 Limit of error - 9.3% Coefficient of Variation - 32.3% - �359 RIVER: Arkansas MILE: 004 QUAD: Nepesta MAP SCALE: 1:7,500 TOTAL AREA: 148.46 ha Date: 9-3-55 CtA Scale: 1:20,00( Date: 10-24-80 Scale: 1:40,000 Change + or - 0 (3) 16.07 16.07 (1) 1.02 0 -1.02 (1) 0.57 -1.04 (3) 2.67 (8) 24.2 21. 53 C2B (3) 8.07 (2) 5.08 -2.99 C2C (1) 1.15 (1) 2.78 1.63 C3A (8) 28.54 (2) 16.21 -12.33 (6) 18.45 0 -18.45 (1) 3.12 0 CIB elC e2A C3B C3C C4A C4B C4C H Ga S AI; D I. NV L - (1) 1.61 .. -3.12 '" 0 0 0 0 0 0 0 0 0 0 0 0 (1) 2.26 (4) 4.8 2.54 (7) 67.31 (6) 64.75 -2.56 0 (1) 0.83 0.83 0 0 (1) 8.7 (1) 12.87 (3) 5.56 (1) 0.3 0 0 0 . 4.17 .. --- -5.26 0 �w (J\ o t\ NV md~ ~ .... 001/ N"re.stif 01" phlt+o,r. ptly I:JlJJ()~ 8+w .5c.,rt. 3..1 195I m., Sc.a/e.: Arkansas I: 7S()o River �..... . '. ) . s (!.~ s C.~A ~ QA . ii:":"'~: mile 00". N.,..t. ~w m., ~ ,hot,,~r.,"Y I : '111,00' 0,,+. C.JA 0' f-' a.,.W ~"J./9'1J SUM: I: 1S0P A,. It. ."'is R iYer ::0. �362 RIVER: Arkansas QUAD: MILE: 045 MAP SCALE: 7-14-51 Date: 1:9,700 Rocky Ford TOTAL AREA: Scale: 1:20,000 Date: 7-18-80 43.9 ha Scale: 1:40,000 Change + or - - CIA 0 (1) 0.7 0.7 CIB (1) 2.0 0 -2.0 CIC 0 0 0 0 -0.9 (2) 1.7 0.8 (1) 2.1 2.1 (2) 1.6 -1.0 CZA (2) 0.9 C2B (1) 0.9 , -_ C2C 0 C3A (2) 2.6 C3B 0 0 0 ClC 0 _0 ._ 0 C4A 0 0 0 C4B 0 0 0 C4C 0 0 0 (2) 14.8 0 -14.8 GI. 0 0 0 s (4) 7.1 (4) 27.4 20.3 0 0 0 0 0 0 I. (1) 15.3 (1) 10.4 NV (1) 0.3 0 -0.3 0 0 0 B AI; D L • '. . -4.9 �< s z 363 �s z . ~. �365 RIVER: Arkansas Date~ MILE: 081 QUAD: Cornelia MAP SCALE: 1:9,600 TOTAL AREA: 124.5 ha 11-2-50 Scale:1:20.000 Date: 7-18-80 Scale: 1·40.000 Change + Oli." -- ClA (4) 2.7 0 -2.7 CIS (2) 2.0 0 -2.0 C1C 0 0 0 ell (4) 4.7 (2) 2.9 -1.8 C2B (1) 1.1 0 -1.1 0 -2.5 -. C2C (2) C3A (3) 4.2 - . -_ 2.5 0 Q (2) 0.8 -~3.4 C3B (2) 4.8 (1) 0.5 -4.3 C.3C (1) 5.2 0 -5.2 C4A (2) 1.8 0 -l.8 C4 (2) 0.7 0 -0.7 C4C 0 0 0 H (4) 13.6 (2) 47.6 34.0 ca (7) 21.4 (6) 8.3 -13.1 s (7) sa !ill NV L I I I 44.4 <, "- - (9) 53.3 8.9 0 0 0 (4) 1.9 (2) 4.5 2.6 (1) 4.3 (1) 4.1 (5) 8.8 (2) 2.5 (1) 0.4 0 . -0.2 " _-"._ " _"_- -6.3 -0.4 ~ =-- �W 0' 0' 't 6ft H s \J ·N MILE. O~ I (_oftNf;...lIA o.» p~oroG~A'H,( j: 2"1000 5 Oc.... 70(3 G.I( M 1\ , ,8."w "). I I q >0 5c AI.-t: : /:"G,oo c.,,4 A ~k: A.fV SAJ R.. Iv {;./J.... �- 367 �368 RIVER: S. Fork Republican MILE: QUAD: Hale Ponds lR MAP SCALE: TOTAL AREA: 147.47 ha 1:18,500 Date: 6-18-61 Scale: 1:20,000 Date: 10-7-75 Scale: 1:40,000 Change .{- orc CIA ClS CIC ell C2B C2C ClA C31& CX 0 (1) 1.15 0 ". (3) 4.04 - (3) 9.77 0 0 0 -1.15 0 0 (3) 16.32 12.28 (2) 10.49 0.72 (1) 2.08 -1.52 (3) 7.62 -0.07 (4) 3.46 (2) 6.11 2.65 (3) 7.06 0) 16.11 9.05 (2) ~ -_ 306 (6) 7.55 IJ" = - = C4A C4 C4C B GI. S AI; (1) 1.27 -, 0 0 NV L -1027 0 0 0 0 0 0 0 (4) 90.24 (6) 77.73 -12.51 (8) 9.35 0 -9.35 (1) 2 15 (1) 2.79 0.64 0 0 (2) 5.21 (3) 6.64 1.43 (3) 2.62 (2) 1.58 -1.04 0 0 0 0 D 0 . 0 d �.» .-- ". " _ . ,':' .~ ~.- ,". _ .. _ .. - .•. _- .... . ~ .~. .-~ ��371 JOB FINAL REPORT State of Project Colorado ----------------------------N-5-R-2 ------------------------------ Work Plan -------'---1 Nonconsumptive Use of Wildlife in Colorado 1 Job Period Covered: Authors: 1 July 1983 - 31 June 1984 JoAnn Profera, Personnel: Janet L. Schreur Clait E. Braun and Bert Widhalm, Colorado Division of Wildlife; Eugene Decker, JoAnn Profera, Ron Ryder, and Janet L. Schreur, Colorado State University. ABSTRACT The objectives of this preliminary were to (1) examine public demand for wildlife viewing, using a test species, (2) examine and evaluate existing data on nonconsumptive values of wildlife in Colorado, and (3) prepare a detailed study plan on an appropriate and approved research topic. All objectives were met and the reports on objectives #1 and #2 are attached. Objective #3 was met by the development of an approved study plan for wildlife viewing at Russell Lakes State Wildlife Area by Janet L. Schreur. This study plan has been incorporated into the project documents for N-5-R-3, Work Plan I, Job 2 for fiscal year 1984-85. Approved by: Clait E. Braun ��j/j DEVELOPMENT OF SAGE GROUSE PUBLIC VIEWING TOURS NORTH PARK, COLORADO Spring 1984 JoAnn Prof era Colorado Division of Wildlife Fort Collins June 1984 �374 Wildlife management has typically been directed toward consumptive users of the wildlife resource as most state wildlife agencies derive the bulk of their income from the sale of user (hunting, fishing, trapping) licenses. However, most state wildlife agencies are legally mandated to manage all species of wildlife for the benefit of all citizens who reside within the state. All wildlife includes both migratory and resident species, but because migratory species are jointly managed with the Federal Government, most state efforts have been directed toward resident species. If legal mandates are to be met, wildlife management must also provide for nonconsumptive users and the wildlife species that provide nonconsumptive use opportunities. Population trends are shifting away from more traditional consumptive uses of wildlife (hunting and fishing) toward nonconsumptive uses such as wildlife viewing and photography (Hendee 1969). A significant number (54%) of the U.S. population participated in nonconsumptive wildlife activities in 1980 (U.S. Dep. Inter. 1982~). In Colorado, over 2 million residents participated in nonconsumptive recreation in 1980, outnumbering consumptive users 2 to 1 (U.S. Dep. Inter. 1982Q). In the past, the Colorado Division of Hildlife's management plan has included strategies to provide the public with information regarding wildlife viewing and nonconsumptive use areas, but few organized viewing opportunities were available to these nonconsumptive users (Colo. Div. Wildl. 1983). Thus, a pilot program was initiated by the Colorado Division of Wildlife in coordination with Colorado State University to identify the demand for and feasibility of wildlife viewing tours. Guided tours to sage grouse (Centrocercus urophasianus) leks were conducted to investigate public demand for organized tours to see this species. Sage grouse were selected for program development because their displays are unique, and lek locations and attendance are predictable, increasing the probability of successful viewing opportunities. Sage grouse reaction to human approach was studied to determine viewing distances minimizing disturbance to the birds. The objectives of the study were to identify public demand for organized wildlife viewing, specifically sage grouse viewing, and to establish guidelines for viewing tours minimizing disturbance to sage grouse. STUDY AREA The study was conducted in North Park, Jackson County, Colorado. North Park is a large intermontane park at an elevation of about 2,500 m surrounded by mountain ranges rising sharply to 3,800 m. Several small streams in the park flow in a northerly direction and are the headwaters of the North Platte River. The climate is characterized by cool temperatures, low precipitation, and a short grow i.ng season. Sagebrush (Artemisia spp.) is the dominant sb rub w i t h herbaceous vegetation consisting primarily of low growing perennial forbs and bunchgrasses (Beck 1977). The study was conducted at Coalmont Lek, 2 km southwest of Hebron. The lek is approximately 200 m wide and 300 m long and is on the northwest side �375 of the bench rising to the east of Pole Mountain Reservoir. Har baceous vegetation dominated the relatively flat display area with dense sagebrush stands on the north, east, and south sides. The western edge was bounded by a steep hill sloping to an abandoned coalmine. A north-south jeep trail bisected the lek about 10 m from the observed center. A 2nd north-south jeep trail occurred about 90 m west of the center. Sage grouse arrived at the lek from the east-northeast. Displays c.entered around a few sagebrush bushes with males distributed on territori.es in a general north-south direction. As the season progressed, males were frequently observed displaying in the dense sagebrush surrounding the lek. Birds departed the lek generally to the west, southwest. METHODS Lek Selection.--Several leks were considered for developing opportunities for sage grouse viewing. Those considered were Coalmont, Delaney Butte, and Perdiz. Criteria for lek selection included vehicle access in early April, expected number of males, and viewing distance to the birds. Due to unusually late snowmelt and poor road conditions, Coalmont was the only lek meeting the criteria, therefore it was used for sage grouse viewing tours. Tour~.--Public viewing tours were scheduled for Saturdays and Sundays beginning 5 April and ending 27 May for a total of 16 days. Press releases were distributed 3 weeks prior to the first scheduled tour (Appendix A). An information handout containing general information on sage grouse mating behavior and a questionnaire measuring user satisfaction were developed by 1 April to be distributed during tours (Appendices B and C). Reservations for tours along with the participant's name, phone number, ~nd group number were taken in advance. Pre-tour contacts confirming reservations were made by phone the week of the tour detailing meeting time, place, and anticipated grouse activity. A count of all birds present was made 1 day prior to tours to assess grouse activity and road conditions. I arrived at 0400 hours at the assigned meeting place to greet participants. Tours began promptly at 0430 with a brief introduction (15 min.) that included the itinerary and proposed tour, a brief physical and behavioral description of sage grouse, and distribution of information handouts. On the weekends of 17 and 26 May, tours left at 0435 due to limited activity of grouse. Participants drove their own vehicles to the lek 27 km from the meeting point. Vehicles were led single file to the lek along the jeep trail 90 !1 west of the lek center. I moved between the vehicles answering questions and relaying information (Appendix D). Participants observed grouse displays from vehicles with binoculars or spotting scopes. Windows could be down to facilitate viewing but participants were asked to remain in their vehicles or, if necessary, leave vehicles only from the west side, a, ~y from the grouse. Vehicles were parked at the same point on each tour. �376 Tour length was dependent on display intensity and length. As grouse activity diminished, questionnaires were distributed and completed at the lek. Tours were concluded with the collection of the questionnaires. Vehicles were backed around leaving the lek by the same trail used for entry. Experiment.--An experiment was conducted the day preceding and following the guided tours beginning 27 April and ending 28 May. The lek was flagged at 10 m intervals in a straight line from the viewing point to the center of activity. Two trials were conducted as birds were approached along this line at marked intervals in a vehicle and then on foot. Birds were observed for 20 seconds and the distance to the nearest male and female along with their activity was recorded. Bird activities were recorded as either strutting for males or as walking away (males and females). Distances that males and females first flushed were also recorded. Birds were not forced to flush if still displaying when approached to 10 m. The study was conducted approximately !z hour before sunrise to obtain maximum counts (Jenni and Hartzler 1978). RESULTS Tours.--Ten tours were led beginning 28 April and ending 27 May. Tours scheduled for 5 through 21 April were cancelled due to excessive snow on the lek and poor road conditions. Reservations were made by 95 people, 66 of which actually attended tours. Thirty percent of the people making reservations and contacted by phone failed to attend. Tour length ranged from 80 minutes on 20 May to 155 minutes on 14 May. Birds were present on the lek at time of departure on all tours dates except 20 May when birds flushed at 0526 due to movement of a participant from the vehicle. On this tour date, participants were led to the center of the lek for a 30-minute discussion that included description of lek characteristics. Distance from vehicles to the center of sage grouse activity ranged from 90 to 120 m due to slight movement of the activity center. Questionnaire.--Results of questions 1-9 were combined into responses expressing satisfaction or dissatisfaction (Table 1). Participants were given a range of responses on questions 1 through 5. "For questions 1 through 4 satisfaction was indicated by marking response 3. Responses 2 and 3 indicated that the participants were not satisfied. Satisfaction on questions 5 and 6 was indicated by a response of 1. Questions 1 through 6 were answered by 57 participants. The remaining 9 participants were part of a private group that accompanied the tour on 28 April. These people only completed questions 7 through 16. Eighty-nine percent or more of the participants were satisfied with the length of the tour and the way the information was presented. A high percentage (82%) was satisfied with the amount of information presented, while 16% of the participants would have liked more information. Thirtyseven percent of the participants would have liked to have been closer to the birds and 67% felt the information handout was useful. When viewing sage grouse, 78% of the participants preferred to remain at a single viewing area. Participants indicated they would take a selfg' ded tour (87%) but 88% preferred a guided tour when directly asked. Thc_:ystated self-guided tours would lack control and would disturb the �377 sage grouse. Seventy-eight percent of participants would pay for the opportunity to view sage grouse, 83% of these people would pay $5.00 or less. Of the 36 participants who indicated purchasing a hunting or fishing license, 27% purchased both with 24% purchasing only a fishing license and 3% purchasing a hunting license only (Table 2). Forty-four percent of those participating did not buy either a hunting or fishing license. Seventy percent of the participants had contributed to the Nongame income tax check-off program. Demographic questions 14 through 16 are summarized in Table 3. Participants traveled an average of 307 km with 50% traveling from the DenverBoulder area. Grouse Disturbance.--Distances at which grouse reacted to approach varied (Table 4). Approach distance to males seem to be related to the number of hens present. A maximum of 59 hens was counted on 7 May and males continued to display when approached on foot to 20 m. On 25 ~~y no hens were observed and males flushed at 90 m when approached on foot. Females began walking and flushed sooner than males on all dates. Both males and females began walking and flushed sooner when approached on foot than when approached in a vehicle. No males or females flushed upon arrival or during the initial lek count. On 18 May, all birds flushed from the lek at 80 m when approached in the truck; consequently a 2nd trial was not conducted. Not all birds began walking when approached. Late in the season (14 May), all females immediately flushed from the lek when approached in the truck. Early in the season, if individual birds were approached either on foot or in the truck, the individual bird would remain in the lek area. Cn 21 and 28 May when the individual bird that was approached flushed, the rest of the birds on the lek flushed and left the lek area. On 28 April a northern harrier (Circus cyaneus) was observed over the lek while males were present. Males stopped dis~laying and crouched but did not flush. When the northern harrier landed on the lek, the grouse remained on the lek and the males farthest from the raptor began displaying within 30 seconds. DISCUSSION Most participants were satisfied with the sage grouse tours. The area where participants expressed a concern was the viewing distance to the birds with 37% wanting to be closer. The major displaying activity occurred at least 80 m from the viewing point with a few birds displaying as close as 35 m. Most participants wishing to get closer were photographers. Although the majority of the photographers had some type of magnification, they were still interested in being closer. Early in the season from early to mid-April, it may be possible to drive onto the lek to facilitiate photographing the birds. However, with a large group, disturbance to t' , birds may occur. As the season progresses substantial distance to t.:.2 birds must be maintained or birds will flu~h as exemplified on 25 May when all birds present flushed during the tour. Another problem �378 expressed by photographers was the position of the vehicles when viewing the lek. Because vehicles were parked west of the lek, the morning sun backlit the grouse making it difficult to obtain detailed pictures. Due to the position of the lek and the heavy sage surrounding the displaying area to the east, it would be difficult to drive east of the lek. Another lek might be used or participants could be advised in advance of photographic opportunities and lek positioning. Thirty-five percent of the participants felt that the information handout was not necessary. The information presented in the handout was general and was reviewed by myself with the participants during the tour. If the handout provided specific information on the area and was highlighted by the guide, it might be more useful. As 86% of all participants had completed college or graduate school, the information handout might be directed to a higher level of audience training. The number of participants was greatly affected by the poor weather conditions in early April. Eleven of the no-shows were directly attributed to the weather either through tour cancellations or poor road conditions. The observed reaction of the sage grouse to the northern harrier is similar to the grouse reaction towards a great horned owl (Bubo virginianus) documented by Lumsdem (1968). Grouse will flush from the Lek if approached by a golden eagle (Aquila chrysaetos), but will remain and resume displaying when other raptors approach (Lumsdem 1968). If signs are used in conjunction with viewing tours, rap tors including golden eagles, may perch on these signs increasing sage grouse disturbance. RECO}l}!ENDATIONS Based on information study, the following presented. obtained from the questionnaire and disturbance recommendations for sage grouse viewing tours are 1. Guides should activity. be used for tours 2. Tour groups activity. 3. A reservations system to 8/lek tour. 4. Participants 5. Alternatives to guided tours should be investigated including a study to determine the effectiveness of viewing from blinds and self-guided tours. 6. Alternative 7. Placement and use of signs on the lek should be carefully considered to prevent their use as hunting perches for avian predators. should maintain should to control a 80 m distance should be used remain leks should in vehicles be studied participant approach and from the center of lek to limit the number when viewing as possible of vehicles grouse. tour sites. �379 LITERATURE CITED Beck, T. D. I. 1977. Sage grouse flock characteristics selection in winter. J. Wildl. Manage. 41:18-26. and habitat Colorado Division of Wildlife. 1983. Today's strategy ...tomorrow's wildlife. Colo. Div. Wildl. Denver, 96pp. Hendee, J. C. 1969. Appreciative versus consumptive uses of wildlife refuges: studies of who gets what and trends in use. Trans. North Am. Wildl. and Nat. Resour. Conf. 34:252-263. Jenni, D. A., and J. E. Hartzler. 1978. Attendance at a sage grouse lek: implications for spring censuses. J. Wildl. Manage. 42:46-52. Lumsden, H. G. 1968. The display of the sage grouse. Onto Dep. Lands and For., Res. Rep. (Wildl.) 83. 94pp. u.s. Department Department survey of U.S. Govt. of the Interior, Fish and Wildlife Service and U.S. of Commerce, Bureau of the Census. 1982a. 1980 national fishing, hunting and wildlife-associated recreation. Print Off., Washington, D.C. U.s. Department of the Interior, Fish and Wildlife Service and U.S. Department of Commerce, Bureau of the Census. 1982b. 1980 national survey of fishing, hunting and wildlife-associated recreation. Colorado supplement. U.S. Govt. Print. Off., Washington, D.C. �380 Table 1. Response to questions 1-9, for sage grouse tour questionnaire determining participant satisfaction, North Park, Spring 1984. Question 1. 2. 3. 4. 5. 6. 7. 8. 9. N responses % Tour length Satisfied Too long Too short 57 51 4 89 7 2 4 Information presented Satisfied Too technical Too simple 56 51 91 Amount of information Sa t Ls'fi.ed Too much Not enough o 5 9 57 9 82 2 16 Viewing distance Satisfied Too far Too close 57 36 21 63 37 Car caravans Effective Not effective 57 Information handout Useful Not necessary 55 37 18 Sage grouse viewing Remain at 1 lek Visit 2 or more leks 64 50 78 14 22 Take self-guided Yes No 63 55 8 87 13 Pay to view Yes No 47 1 o 46 11 tour sage grouse 64 50 14 81 19 67 33 78 22 �381 Table 2. Age, sex, and education North Park, Spring 1984. of sage grouse tour participants, Question % N responses 14. Sex Hale Female 66 34 32 51: 49 <16 17-19 20-29 30-39 40-49 50-59 >60 66 0 0 9 22 10 11 14 14 33 15 17 21 66 1 8 24 33 12 36 50 15. Age 16. Education Elementary High School College Graduate School 2 ------- Table 3. licenses, Sage grouse tour participants North Park, Spring 1984. Activity Hunted or fished Hunted and fished Fished did not hunt Hunted did not fish Did not hunt or fish who purchased N hunting or fishing % 36 18 16 2 29 55 27 25 3 45 �w ex N Table foot, 4. Distance (m) at which North Park, Spring 1984. male and female sage grouse Approached Date 27 30 04 07 11 14 18 21 Max. males /1ay 32 25 31 35 37 32 34 40 25 May 28 /1ay 29 30 Apr Apr May May May May May Max. females 52 42 47 59 22 4 2 1 0 2 Con t inued to display Males Began to wa Ik with Flushed -- -- 30 30 20 20 25 --- -- -- 50 40 -- 75 70 --- -- 40 80 50 20 --- 70 60 90 55 40 showed signs of disturbance when approached Approached truck Females Began Flushed to ",aIk Cont inued to display Males Began to lialk in a vehicle and on on foot Flushed Females Began to via Ik Flushed 20 -- -- 30 -- 70 -- -- 60 60 20 40 -- 20 -- -- 30 50 70 40 30 50 35 -- 30 50 80 70 75 -- -- 100 -- -- -- -- -- 40 -- 90 90 80 3D 40 20 20 40 -- -- 65 20 30 �AFPENDIX A. 383 Press release for sage grouse tours 1984. 6060 Broadway, Denver, Colorado 80216 CONTACT: IMMEDIATE RELEASE Ft. Collins -- If you are interested behaviors GEOFF TISCHBEIN in observing one of the most unusual in the animal kingdom, now is your chance. of Wildlife, in cooperation The Colorado Division with Colorado State University, ing trips to sage grouse "strutting" grounds beginning will start at 4:30 a.m. at the intersection where a guide will lead the participants 484-2836 of highways will be conduct- 7 April. The trips 14 and 125 in Walden to the viewing areas. Every spring, for thousands of years, male sage grouse have congregated on historic "strutting" attempt grounds where they stake out territories to attract female sage grouse. the most curious behaviors and The strutting of the males is one of found among North American wildl ife and continues every spring for as long as two months. Interested persons should contact Clait Braun at the Colorado Division of Wildlife Research Center in Ft. Collins, 484-2836. required and will be accepted on a first-come, suggests taking binoculars first-serve basis. and/or spotting scopes and recommends be prepared for cold, rainy-snowy able in Walden. Reservations weather. Motel accommodations are Braun participants are avail- �384 APPENDIX B. Information handout for sage grouse tours. WHAT'S IT ALL ABOUT? The display of male sage grouse people have witnessed. grouse, the Colorado University is a spectacular To improve the a~preciation Division has developed of Wildlife this tour. grouse react to your presence activity without disturbing with Colorado State and how you can best enjoy their courtship them. By participating in this tour, you will and enjoyable to the people in future years. Sage grouse return annually display grounds A STORY OF DOMINANCE to open areas called leks. These communal provide a focal point for spring activity. showy white breasts that contain Hales, with large two greenish-yel low air sacs, return from areas in late March to establish territories. A hierarchy achieved with the dominant males near the center of the lek. activity of sage This study will investigate how sage SAGE GROUSE COURTING: wintering of the displaying in coordination help make sage grouse viewing more accessible of Colorado spring event that few as males are busy defending attract females. acquired territories is soon There is much and displaying to Can you identify the mating display and the territory defense display? Smaller and more mottled are attracted in early April and to the center of the lek and the dominant males. males wil I mate with> lek and the dominant successful than males, females arrive 90% of the females. males? in attracting Subdominant These older Can you spot the center of the males also display but are seldom females. IS IT ONLY A NAME? The yellow-bellied not suck sap! sapsucker does not have a yel low belly and it does Does the sage grouse too have a misfit name? life cycle provides us with a clue. begin nesting. Camouflaged overlook Sagebrush within and free of snow. and snow. provide a protective the sagebrush, the grouse. After mating, Wintering predators vival of these birds. named. the sagebrush from wind provides a vital food 95% of its winter Thus, this shrub is essential to the sur- Sage grouse, unlike the yel low-bellied sapsucker, The existence of this unique grouse of sagebrush-dominated its life requirements. is left unexposed protection Over half of its summer diet and> diet are the leaves of sagebrush. the maintenance like the golden eagle can easily This shrub is excel lent in providing source for the grouse. leave the lek and site for both hen and eggs. areas are where sagebrush More than just protection, appropriately females A look at its plant communities is dependent are upon that provide al I of �385 APPENDIX C. SAGE GROUSE QUESTIONNAIRE Thank you for joining the sage grouse tour. more about your preferences questionnaire, for observing you will provide valuable sage grouse viewing This questionnaire sage grouse. information is to learn By completing 1-6 by circling the appropriate 1. The length of the tour was number.) too short too long 2 2. The informat ion presented was too technical 5. The amount of information presented was The viewing distance 5 not enough 2 3 4 2 3 4 2 3 5 too close 5 not effective effec t ive 6. The informat ion handou t presented was valuable 4 5 not necessary 2 7. 4 3 too far Viewing grouse from car car avans was 5 too simple too much to the grouse was Please check the appropriate 4 3 2 4. ve in the future. (Please answer questions 3. t-;s that will help imp" 3 4 5 blank: When viewing sage grouse, would you prefer: to remain at a single viewing site for an entire morning? to vis it two or more areas in a morning? 8. Would you take a self-guided Yes tour to a sage grouse strutting area? No 9. Would you pay for the opportunity Yes No to enter a wildl ife area to view sage grouse? If yes, how much? --_$ 10. Have you purchased a fishing license in Colorado within the past 5 years? Yes No 11. Have you purchased a hunting Yes I icense in Colorado w i t h I'nthe past 5 years? . No 12. Have you contributed Yes to the Nongame income tax check-off program? No 13. What is the town of your residence? 14. Your sex? 1 ;;. 1 .~ • Your age? Male Female 16 or below 40-40 What is the highest 17-19 50-59 20-29 over 60 level of formal schooling you have completed? Elementary High school or vocational College or technical school Do you have any suggestions? school Graduate school Please explain, use the back if necessary. 30-39 �386 APPENDIX tours. D. Outline of information presented by guide during sage grouse Sage grouse description Males/females -- physical characteristics Family characteristics Displays Courtship display/territorial defense display Sounds/movements/length of displays Lek System Dominance hierarchy Territories -- size - arrival of males, females Adults vs. juveniles After Breeding Season Males - life expectancy Females - nest - young (life histories) Wintering Lek Characteristics Sagebrush use Size, history Predators Golden eagle Band ing Hunting information/population Trapping Scientific study size �387 LITERATURE REVIEW THE DEMAND FOR NONCONSUMPTIVE WILDLIFE USES Janet L. Schreur Colorado Division of Wildlife Fort Collins, Colorado June 1984 �388 There is evidence recreation predicted that participation in nonconsumptive is greater than has been previously to continue participation to wildlife on an upward trend. in nonconsumptive managers. of their efforts realized wildlife and it is The amount of interest and use of wildlife Most state wildlife is of particular agencies interest have dedicated most to hunting and fishing interests. This has been justifiably so because is becoming clear that there is another use group, wildlife managers should consider sportsmen have "paid the bills". using the resources Now that it of this group for the benefit of all wildlife. Participation Arthur Surveys (1979), using data from the 1975 survey conducted U.S. Fish and Wildlife Service estimated that 49 million 9 years of age took special trips to view wildlife billion days in wildlife survey, he estimated 75 million that 15 million hunted or fished. tivew wildlife recreation observation. Americans over and spent 1.6 Using data from the same people photographed Arthur's estimates wildlife and indicate the apprecia- user group makes up a large proportion group. by the of the wildlife �389 The 1980 National Recreation (U.S. Dcc. population participated group, Survey of Hunting, Inter. 1982~) estimated in nonconsumptive 36Z observed wildlife inciiciJte,contrary t he hun.ters and fishermen lower estimate wildlife, Fish ilnd Wildlife proportion wildlife (1979) estimates, population is La rger activities. vs. 49 million Service's These data than the proportion of of This survey had a much that took at least 1 trip to observe by Arthur and Arthur's of the U.S. popul~tion Of this that the proportion and 29%, respectively). (15 of the number of people 29 million that 54~{ of the U.S. and 10% llhotugraphed wildlife. to Arthur's w i.LdLi f e v i ewe rs in Fishing and Wildlife-Associated (1979). estimates Both the U.S. indicate a significant take trips away from home to observe w i l d l:i f e . Surveys of nonconsumptive done at the s t a t e level for wildlife several recreation states, demands have also becn Lnc lud ing Colorado. The U.S. De par t.me n t of Ln t er lor (1. 982.!?)e s t i ma t ed that non cons urnotive w i Ld l i f e users in Co Lo r.rdo Coloradoans million ou t nurnbered participated there of over 1 million 2 to 1. by consumptive high participation wa s not higher Active w.iLdLi f e p eopLe wildlife and v i.ewi ng '.Jildlife. In v i cw i ng (bird 331:: of all Tn an Oregon r ecrea ri.ou rate feeding and birding Oregon of tile rcs i dcn t s we re consumptive for a total recreationists. done in o t l.er states. consumptive hunters 373,000 rate was found for nonconsumptive than the or been fishermen Over 2 million recr~;ltion in 1980 and ~ in nonconsumptive we re 925,000 Su rvey s have also Il7;' users of them tuok trips for the main pur posc of comparison, 505,000 consumptive residents. use rs . In recreation, (Aney survey, a but it and Cowan 1975). trips) was enjoyed by c omp.arLson , 7J9,000 or �390 A survey indicated of of :!assachuset ts demand wildlife a high Demand Intensity (19,6) found ~linnesota indicated wild!if~ tlia 1978). that surveyed it a select wildlife t i.on Colorado the (U.S. Dep. in "i1e! by traveling demand for (U.S. in nature 5.6%, but and llowev e r , tilt: authors already walks less 6% of fishing 79Z wildlife chan the 10% UI w ild Li.f e the people recreationists on 3 large was estimated that wate1ling ill nonconsumptive survey than bird 1982.9) or wildlife da v demonstrated Lda h o , less Li.k ewis e , only In a different their recreation Dep . Inter. nonconsumptLve of indicated nonconsumrtiv~ In in surveyed respondents to a major participated were the l.ime Area point were calculation 1975). Canoe high outdoor 1982~) surveys. 1973). VS. enjoyed Nati o na I Waters the by the (Gray wa t c h i ng areas SOl of l>e111 1977). participation Boundry w il.d Li.Le was that a lower t hc Saskatchewan hun ti ng , 7.3 the r cup 3 choices. actively and Dem~nd Indi~es ind iv i dua Ls that. of Inter. residents jpg most interest e i t hc r than (Sc hwe i t z e r e t a1. than t hoi indicate r e c r c-at i.o n (Fazio Forests, in viewing state's surveyed tllcir )',roup Som~ surveys r e cr ca to found was \,'25 v i.ewi.n g when analyzed t.o m.i j o r bird They co n s e r va t t on c ornrm s s i ouc r s Rt'lHtive ob s e r. t visitors observation indicated and 34% of visitors In Arizona, th0ir Scores that (Shaw e t a1. for muni c i paI to N;.ticmal be more (47.1/;) (Tyre w i LdLt f e professionals and i:Jportant a ud James 1971). \.Jitter 69::~ were recreatioll most and bird Shaw (1979) wa t c he r s . by 67% and important. surveyed Hunting 39% indicated was indicated that bird as the most and wildlife found that enjoyed outdoor watching ~ere . �391 on several Based est i rnated observation, as 0 f 5% were concluded life 27% t hat surveys that were participants and U. S. serious these there the including pop activities appealed sex ratio of and exclusive and Recreation In s po r tsme n also the Colorado w i Ld I i.fc named of This l'n~;aged Der. in (U.S. indicates wer e in that: this Wildlife Prefcrenccs t nc lud ed found thac mammals were birds were the fished. In game species witii and eagles observing Fazio than this and that !,.J1ldlife65% of rare, reported the also game species that largest are, reporting participation preferences. Gray grouping endangered high most on the (Bolli only group. people; too. [or or 54/~ user Sp o r tsruon non co ns ump t i ve users son.gbircls BcLl.L (l977) the taxonolnic general, nonc.o n s urnpt i ve not just observation Idaho, big support and and us e r s are r e c r c a t.i o n . preferred 111 preferred. preferred cd or farther 2nd choice. an i maLs were users t wildlife the mutually 1982_1.~), 41% of pur c Ly nonconsurnptive l.av c gone as them as 19H2~) estimated Inter. between users Hun t i.ng , Fishing c a t ago r y mak Lng it nonco ns umpt i ve w i LdLiI.e r a t c s and have do not "c omb i na t i ou users" enjoying Some surveys of of 50:50. of wildlife surveys of walks differences think He noric o ns umpr Lvc wi l d li f e activities. hun ove r La pp i ng group residents to Dep . Inter. r e c r e a t Lon i.s t s also this Idaho Survey (U.S. survey tend all was about think Recent 1980 National Associated all they non-overlapping. The participants wildlife. from demographic managers noncollsumptive, separation. or ,"i 1d 1 i f e 2% photographed to people (1979) e a r Li e r , :'lore t i c i pat e din 11 pal and few socioeconomic the til) birders, IJhen many na t ur a L resource consumptive 1.1 1a t ho s e cited (1975) viewing and unusual p r e f c rr e d big list. Combination 1977, Lyons 1982). �392 Ec on.nni c Significance It is a difficult and controversial va lue for nonconsumptivc recreatioll and other and Wildlife wildlife this Service activities is almost n was noncol1sumptive tive wildlife basis by households (Payne and DeGraaf Hountain Rocky Houn t a i,n $70.00 ships to organizations. worth of equipment, that $89 birds binoculars, that $500 million in 1974. The authors field guides, Audubon their enjoyment Society that appr2ciative and a portion owned concerned members of nonconsumptive Audubon wildlife obtained of the and Rocky they had invested wildlife in uses. $90. 00 wo r t h of equipment Society subscription members spcn t $~:80.00 per year and donated in issues was and cameras. National National impact of nonconsumr- on an industry-wide per year for such things as magazine time to involvement evidence They estimated Na t .i o naI Park visitors spent and more U.S. on the economic Park vis:itors to learn how much for enhancin~ For comparison, ($17.3 billion) total expenditures of birdseed, (1976) surveyed equipment consumptive The U.S. Fish (1974) estimated to estimate of nongame scopes, National 1982b). on fishing Horvath 1975). this f i gu re f rom the sales Proctor Inter. ill the southeastern use by estimating SPL~;itfor the enjoyment demands. with activities. study attempted sale of spotting resource in 1980 (U.S. Dep. wildlife Nonc+t he+le ss , it is necessary that $14.8 billiO!l was spent on nonconsumptivc ($8.5 billion). spent Another estimated a monetary is to be commensurable natural as much as was spent than on hunting mi.LlLo recreation. w iLd Li f e recreation if nonconsumptive wildlife wildlife task to calculate with wildlife. recreation and member- owned $20.00 and $700.00 of their Thes~ is big business. figures arc �393 As another cxamp le of the economic wildlife recreation, importance a Tillage in India reaps large benefits as a result of a nearby developed nonconsumptive wildlife TherL is a nesting colony of waterbirds State, India. of non cons urnp t Lv e The villagers area (Spillett near Vedanthangal of Vendantlwngal pests and by furnishing the India government miles each in Vedanthanga1 (0 t owe r , predicts \.;i11continue (1969) educated obs~rved (1) t he spent money loss of accessable Our society has been shifting faster than llunting land for where hunting is (4) more organizations. users are typically more urban backgrounds. Conversely, and have less education towards more urbanization levels which indicates Lime (1976) in the quality of hunting, that nonconsumptive hun t ers come f rom rural backgrounds wildlife (Shafer and future. and (5) more people in conservation and come ~rorn predominantly educational a day but it does appear that nonconsumI,tive to increase in the primary w i Ld Li Ie use, (3) decline Hendee recreation , (2) decline in the rural population pressure, a and other items. use will grow proportionally or fishing for several reasons: anti-hunting These visitors form of wildlife That mil} be optimistic f i sh i ru; visitors 5,000 that by the year 2000, nonconsumptive that nonconsumptivc hun t. i ng and By 1966, see the colony. recreation will be the primary w i Ld Ld f c recreation In 1962, Use It has been predicted 1974). to their agriculture a source of fertilizer. for food, transportation, Trends of Nonconsumptive ~oeller the paved the road to the colony, put in restrooms, parking Lo t and an observation traveled >100 in Madras have been protecting colony for decades becallse the birds were beneficial by controlling 1970). rhan average. and higher there may be a corresponding decrease �394 in hunting and an increase Another indication in appreciative of the direction wildlife wildlife use. recreation is headed can be found in the participation levels of young people in the different Results of one study indicate types of wildlife that childhood outdoor of adult recreation (1977) results year-old recreation. recreation activities activities are important (Yeosting and Burkhead show a high level of nonconsumptive nonconsumptive interest Audubon Combining the of a growth trend for Audubon (1975) suggest that the trend in membership Society can be used to indicate in bird watching, reasonable, Belli's use. Payne and DeGraaf the National 1973). use in the 15-19 group and the reverse was true of hunting. results of these studies support the prediction indicators a nonconsumptive the trend is definitely Society had 41,000 members. the trend in activity. upward. If this is In 1963, the National By 1970, membership 142,000 and by 1975 to 321,000 members. of had risen to This is an 8-fold increase in 12 years. Conclusions There is substantial wildlife recreation population. evidence in the literature is a popular past-time These recreationists of this activity wildlife is upward. enthusiast's for a large segment of the u.S. contribute money to the U.S. economy in their pursuits Wildlife that nonconsumptive a significant amount of and the trends for the growth managers should plan to use these support for the benefit of all wildlife species. �395 LITERATURE CITED W. \.J., Aney, and C. D. recreational L. H. Arthur, resource. 1979. the American values. L. A. public G. C. Univ. value of wildlife: Trans. T. C. Assessing For. Servo Gen. Tech. 1977. preciptions i.:Q_ 32-3LI Pages Coordinators. Idaho, Moscow. important 30(2) :8-9. of nonconsumptive in Ldaho, 1975. Nonconsumptive commissioners Massachussetts, He nd e e , J. of Daniel, amenity E. H. resource Rep. RH-68. wildlife users in Idaho. 1IOpp. Characteristics of nonconsumptive North Am. l.Jildl.and Nat. Resour. J. oriented versus of who gets what and Nat. 1974. for wildlife in Massachusetts. Appreciative studies Am. \Hld1. demand by municipal M.S. Thesis, Univ. Amherst. 1969. C. refuges: Horvath, Wildl. shows wildlife 42:117-l28. conservation j Tech. J. R., and L. A. Belli. Conf. Survey and sportsmen. 1977. Survey w i Ld Li f e users Gray, The esthetic U.S. Dep. Agric., M.S. Thesis, Fazio, 1975. Oregon and B. L. Driver, Zube, Belli, Cowan. Resour. Economic recreation. Conf. Survey Trans. consumptive and trends uses of wildlife in use. Trans. North 34:252-263. of southeastern North Am. Wildl. wildlife and wildlife and Nat. Resour. Conf. 41:533-546. Lime, D. I·i. Lyons, J. R. 1976. Wildlife 1982. Nonconsumptive U.S.: identifying and Nat. Resour. More, T. A. 1979. the other Conf. too. J. For. wildlife-associated constituency. Trans. 74:600-604. recreation in the North Am. Wildl. 47:677-685. The demands of the literature. 16] p . is for nonhunters, for nonconsumptive U.S. Dcp. Agric., w i.Ldl.Lfe uses: For. Servo Gen. Tech. a review Rep. NE-S2. �396 Payn~, B. R., and R. M. DeGraaf. trends associa[~d Proc. Symposium 1975. Economic values and recreational with human ~njoyment on management of nongame birds. Pages 6-10 in of forest and range habitats for nongame birds. U.S. Der. Agric., For. Servo Gen. Tech. Rep. WO-l. Proctor, B. R. 1976. in Colorado. Recreational surnp rLve wildlife activities Unpubl. Rep., Colo. State Univ., Fort Collins. preferences for birds in Saskatchewan. C. Hendee and C. Schoenfeld, programs. Shafer, on noncon D. H., D. A. Scott, A. W. Blue, and J. P. Secter. Sc~~eitzer, in J. Expenditures Wildl. Manage. E. L., and G. H. Moeller. 1974. 1973. Pages 42-49 eds. Human dimensions lnst., Washington, 24pp. in wildlife D. C. Wildlife priorities and benefits: now , 2000, and beyond. Trans. North Am. \Hldl. and Nat. Resour. Conf. 39:208-215. Shaw, W. W., D. J. Witter, D. A. King, and M. T. Richards. Nonhunting wildlife enthusiasts and wildlife \\cst. Assoc. Fish and IHldl. Agencies. Spillett, J. J. 1970. uses of wildlife 1971. on recreation of Interior and Commerce. Lnt er ., FLSll U.S. Departments of fishing, species. Int. sites and areas. For. Servo Res. Note. SE-161. 4pp. 1982a. of fishing, hunting and wildlif~-associated DL'p. values Length and rate of individual in various activites U.S. Der. Agric., U.S. Departments conservation: Nat. Publ., New Ser. 17:121-129. Tyre, G. L., and G. A. James. participation Proc. 58: 255-263. Economic aspects of wildlife of consumlltive and non-consumptive Union Conserv. management. 1978. and ihldl. of Interior and 1980 National recreation. survey U.S. Serv., \.Jashington,D. C. 156pp. Cornrne rce . 1982b. 1980 National hun t i ng, and w i LdLi f e+a ssocLa ted recreation: survey Colorado. �397 De p . Ln t c r ., Fish U.S. D. hTjtt<:r, J., w i Ld Li.f Yeosting, and \':. \,J. and Sll:!I,), l-lildl. Serv., 1.97<). Bel ic f s ,)C e pro f essi.ona Ls about wildlife D. R., and D. L. Burkhead. recreation analysis. exp~rience J. l.ci su rc on adult ]{e:s. L973. leisure 5:25-36. h'ashin)',ton,D.C. b i r d c r s , hun man agcmc n t 'lr . Significance behaviour: te rs, :.:.nd ans , North of childhood an exploratory � https://cpw.cvlcollections.org/files/original/4254994e75588567a4e0d72ac1269c4c.pdf 2249013bfef52359dda54db3a1de4258 PDF Text Text Colorado Division of Wildl ife Wildlife Research Report Apri 1 1984 JOB PROGRESS Colorado State of Project REPORT W-37~R-37 Work Plan Job: Job Title: Evaluation Period Covered: Author: Personne I: (45-01-504-15050): Game Bird Survey 23 of No-Till Wheat Farming 1 July 1983 through 30 June 1984 Warren D. Snyder Warren D. Snyder ABSTRACT Background information concerning alternate wheat farming systems was collected and evaluated. Efforts to implement and evaluate no-till biennially-cropped winter wheat farming on the South Republican State Wildlife Area were coordinated with management personnel of the Divisionis Southeast Region. A detailed program narrative (study plan) was prepared and incorporated into the project documents. ��3 EVALUATION OF NO-TILL WHEAT FARMING Warren D. Snyder P. N. OBJECTIVE To prepare a detailed study plan as a basis for evaluating the effects of changing cropping systems on r lnq-riecked pheasant (Phasianus colchjc~) nesting success and production. SEGMENT OBJECTIVES 1. Review literature and contact agriculture research personnel knowledgeable about reduced tillage and no-tillage summer fallow systems and their appl ication to the central High Plains of eastern Colorado. Consult with wildlife agency personnel in neighboring High Plains states to learn if similar research efforts are planned or in progress. 2. Survey eastern Colorado using personnel of the CSU Extension Service and the U.S. Department of Agriculture, Agricultural Research Service and Soil Conservation Service to identify the type, extent, and location of new cropping systems currently in use. 3. Confer with CDOW management personnel to learn if a cooperative effort can be made to obtain a suitable study area either on CDOW land or on private land in eastern Colorado. 4. Prepare a detailed study plan to evaluate the impact of a reduced or no-tillage system on pheasant nesting success and production assuming a suitable study area can be obtained. RESULTS AND DISCUSSION Literature concerning reduced tillage and no-tillage small grain farming methods was acquired and reviewed. Contacts were made with George Nason of the Nebraska Game and Parks Commission concerning development and implementation of a study on the impacts on ecofallow farming on pheasants. Recent information concerning toxicities of certain herbicides was provided by Randy Rodgers of the Kansas Fish and Game Commission. A summary of conservation tillage practices by county in Colorado prepared by the Conservation Tillage Information Center showed 612,273 acres (247,970 ha) of reduced tillage, 318,642 acres (129,050 ha) of mulch tillage, and 16,647 acres (6,742 ha) of no-tillage small grain applied during 1983. Nearly one-half (3.240 ha) of the no-till treatment was in Kit Carson County in east-central Colorado. Mulch-tillage uses a combination of herbicides and cultivation and reduced tillage uses any system, primarily sweep tillage, so that at. least 30% residue cover is retained on the soil surface after planting. These data show that no-tillage has yet to gain major acceptance by wheat farmers in Colorado. �4 Dr. Darryl Smika, Agronomist, U.S. Department of Agriculture stationed at the Great Plains Experiment Station at Akron, Colorado, was contacted concerning selection and use of herbicides, equipment, and methods for notill summer fallowing in a biennial winter wheat-fallow crop rotation. Local CSU Extension Service personnel were contacted concerning recent developments in wheat farming methods. Contacts menting Division Wildlife concerning equipment acquisition and potential sites for impleno-till farming and an evaluation study were made with Colorado of Wildlife management biologists. The South Republ ican State Area evolved as the only site available for initial evaluation. Subsequent meetings were held wl th Tom Lytle, Habitat Management Biologist, concerning study design, field selection, herbicide selection and acquisition, and other factors in treatment initiation. A detailed program narrative concerning evaluation of the impact of no-till winter wheat farming on ring-necked pheasants was prepared and currently is under review. Prepared by ~lt~~I)J~~d_V Warren D. Snyder Wi ldl ife Researcher ~'~~- C �5 Colorado Division of Wildlife Wildlife Research Report April 1984 JOB PROGRESS REPORT State of Colorado Project No. W-37-R-37 Work Plan No. Job Title: Job No. 3 Responses of Sage Grouse to Vegetation Period Covered: Personnel: Game Bird Survey (15053) 13 Fertilization 1 July 1983 - 31 December 1984 Blaze Czerniakowski, JerrYHupp, JoAnn Profera, and Tom Remington, Colorado State University; CIa it Braun, Jack Corey, Ed Frankonis, Keith Kahler, Steve Porter, Steve Steinert, Joe Timchak, John Wagner, and J. Allen White, Colorado Division of Wi ldl ife. ABSTRACT Long-term sage grouse (Centrocercus urophasianus) baseline data collection continued in North Park, Colorado in 1983-84. The 1983-84"wiAter was exceptionally harsh with heavy precipitation and late snow cover. Timing of breeding events was delayed and counts of males on leks decreased by almost 50%. This decrease was attrib~ted to poor body condition and not poor survival as recoveries of birds banded prior to 1984 indicated normal survival. Sage grouse hunter success was normal (53%) as was birds per hunter (1.3). Percent chicks in the fall harvest (57) was among the highest on record. Nestinq success (63% of al 1 females were successful) was the highest recorded in the 1974-84 interval. Overwinter survival of yearlings (young produced in 1983) was good (23% of the total harvest). Hatching in 1984 was delayed by 2-3 weeks from the "ave raqe!". Annual turnover est imates were 52% for adult males and 42% for adult females. The direct (harvest rate) recovery rate· for 234 hens banded in 1984 was 5% wh i 1e the direct recovery rate for 194 males banded in 1984 was 10%. About 1,000-1,200 sage grouse were harvested in 1984. . ��7 RESPONSES OF SAGE GROUSE TO VEGETATION FERTILIZATION Clait E. Braun This long-term study to examine the responses of sage grouse to vegetation fertilization in North Park, Colorado was initiated in 1980-81 when it appeared that surface mining of coal would be expanding. The study was scheduled in 2 phases. The objectives of Phase I prior to fertilization/ mitigation by coal companies related to collection of baseline data against which the effects of treatment (fertilization) would be evaluated. Thus, the feeding preferences of sage grouse in January-April including the dietary composition of sagebrush (Artemisia spp.) by species and/or subspecies at feeding and random sites was examined. This portion of the study was completed in 1983 (Remington 1983). With the virtual shutdown of the Kerr and Wyoming Fuels mines in 1983, only the Walden mine has continued to operate. Also, the State Mined Land Reclamation Board has failed to recognize vegetation fertilization as a possible mitigation tool. Because of these 2 events, Phase II (fertilization) has not been undertaken and the study has been placed in a baseline monitoring basis. This will continue until funds are available for fertilization of selected study sites. P. N. OBJECTIVE The objective of this study between Phase I and Phase II is to monitor the sage grouse population throughout North Park through banding, lek counts, and collection of harvest data. Segment Objectives: 1. Use night-lighting techniques to capture adult and yearling male sage grouse roosting on leks during March-May. Trap and band 300 males and at least 100 females within North Park from March through May. Obtain body weight and other measures of body size from sage grouse at time of capture. 2. Make counts (4/lek) of males and females on all known leks in North Park. 3. Survey areas in North Park to locate new or relocated leks. 4. Collect 10 adult male sage grouse from 4 to 5 leks during the early (20 Mar - 10 Apr) period of the display season. Collect an additional 10 adult males from leks during the latter period of display (15-30 May). Estimate body lipid reserves of collected individuals. Evaluate depletion of lipid reserves during the display season by comparing carcass composition of early- and late-collected males (reported in Work Plan 3, Job 15). 5. Clear established and June. pellet group transects (30) during September-October 6. Collect population data through use of wing barrels and check stations during September-October. �8 7. Prepare progress reports. Present pertinent findings at appropriate scientific meetings. METHODS Sage grouse were located where they roosted along roads, trails, and on leks and captured in late winter and spring using night-lighting techniques as described by Giesen et al. (1982). Considerable success in capturing females was achieved by locating flocks immediately prior to darkness and then returning for trapping attempts after several hours had elapsed. All birds captured were classified to age and sex (Beck et al. 1975), weighed to the nearest 10 grams, and selected birds were measured. A sample of males was collected by asphixiation during 2 time intervals for development of physiological indices (Work Plan 3, Job 15). Counts of birds present on leks in April and May were made following procedures outlined by Braun and Beck (1976). Pellet group transects were cleared and all pellets were classified following Schoenberg (1980). Harvest data were collected through use of hunter-check stations and volunteer wing collection stations (Hoffman and Braun 1975). RESULTS AND DISCUSSION Lek Counts Number of active leks and males counted per lek decreased (p < 0.05) in 1984 from levels documented in 1983 (Tables 1, 2).· The winter of 1983-84 was extraordinarily harsh and extensive snow cover remained into early May. There was little if any bare ground on lek sites in April and sage grouse remained in flocks where exposed sagebrush was available. Attendance of leks by grouse was delayed and leks did not become active until late April to mid-May. Nine leks that were active in 1983 were not active in 1984. While vehicle access to leks was exceedingly difficult in 1984, all leks were eventually reached. While it is possible that some leks classified as inactive may have had some activity, this was highly unlikely because most were completely snow covered into early to mid-May and none had fresh sign when checked in late May-early June. Mean number of males counted on leks in 1984 was only 54% of the mean in 1983 (Table 2) and was the lowest on record in the 1973-84 interval. If number of males counted on leks in spring reflects size of the spring population, then population size decreased significantly from 1983 to 1984. The decrease in males counted per lek may have reflected an actual ch~nge in population size or it may have reflected reduced attendance of males at leks in 1984. In either case the record snow depths in winter 1983-84 were the ultimate cause. Evidence to support a conclusion that sage grouse died over winter could not be found either through observation or searches for carcasses in spring (i.e., on pellet transects). There was conclusive evidence that body weights and fat levels were significantly lower in spring 1984 than in spring 1983 (J. Hupp, unpubl. data). These data suggested that adult males were more affected than yearling males and that yearling hens were more affected than adult hens. In general, all males were affected more than all females. Thus, a hypothesis could be formulated that counts of �9 Table 1. Counts of sage grouse on leks, North Park, 1984. Lek Aspen Al ka 1 i Lake Arapahoe Bighorn Boettcher Jet. Canuck Cherokee Cheyenne Coalmont Deer Creek Delaney Butte Denmark Fish Hatchery Hound Lost Creek #1 Migan Ortega Perdiz Peregri ne Prague Pronghorn Ptar Ra i 1road Ram Raven Ridge Road Ri ley Spring Creek #1 Spring Creek #2 Spring Creek #4 Turkey Ute Walden Wattenburg #2 '.. Counts 1 4 1 1 4 1 1 3 4 2 5 3 5 2 3 1 1 3 2 1 1 1 3 1 2 3 2 3 3 1 3 1 3 3 Number of Males· Females 11 36 18 0 22 0 0 38 43 14 54 8 10 19 2 0 0 8 11 0 0 0 11 0 5 31 0 50 8 0 47 2 0 18 0 24 1 0 8 Males 01 Jun 17 May 29 May 5 0 34 8 0 6 0 0 2 Females 08 May 29 May 16 May 17 May 18 May 22 May 19 May 0 0 1 49 0 10 5 1 7 0 0 0 7 0 0 0 0 0 0 0 Hi~h count No birds 23 13 17 24 02 10 09 16 May May May May May May May May 19&23 May 28 Apr 08 02 17 09 May ~1ay May May 18 May 29 May 02 May 11 May 30 May 27 May 02 May < 10 May 16 May 04 May 04 May 23 May 23 May 18&22 May 08 May 08 May 23 May 08 May 15 May 02 May 11 May 29 May 1,10, 18 May 23 May 23 May �10 Table 2. Trends in peak counts of male sage grouse. North Park. 1973-84. Years Lek 1973 1974 Alkal i Lake 81 72 1,,76 1977 1')78 1981 1 q!32 1083 36 39 62 29 68 32 60 116 56 61 1979 72 43 1980 39 21 2~ !S 21 24 12 10 )8 27 22 J J 15 100 22 (1977) Arapahoe Aspen 1975 68 (1977) 6 ighorn 50 €-2 52 76 47 36 31 107 14 113 22 30 27 21 21 4 46 (1976) Boettcher Lake Jet. 59 49 26 27 106 22 92 13 16 33 31 8 a o 127 137 94 80 101• ° 101 25 32 36 52 62 a a a I) 28 52 23 109 66 47 60 72 (1981) Buteo Canuck (1974) Case Flats (1978) (1978) Cheyenne Coalmont #5 (1973) Cowdrey Deer Creek De laney Bu tte 47 30 37 27 19. 29 28 21 21 9 5 1 II 27 5 36 31 41 4 13 o o (1977) Oenmark Fish Hatchery 67 (1974) Hound Lost Creek 111 13 Lost Creek 112 10 Monahan Draw Owl Creek II o 4 12 28 80 136 144 11 2 a o o o I) 97 82 67 23 50 63 10 71 81 64 69 27 21 28 2,5 78 27 20 22 o 18 33 47 113 30 20 29 36 26 o o o o o a o o o 70 53 27 I' o o 53 o a a o o 8 23 27 13 21 3 6 (1976) o o (1979) Higan Q 19 69 33 62 )0 58 (1978) Eagle ° 1'3 ~- 1 1 o 14 16 15 9 16 Perd iz (1979) o o o o Peregr ine (1978) 17 20 (1978) Pronghorn (1977) 35 34 26 7 (\ 10 43 10 a o a G 50 35 55 20 19 24 62 41, 33 64 11 63 40 43 67 f,o 41 31 54 48 30 58 53 41 40 o o a o 59 Prague Ptar 15 (1978) Rai I road (1975) Raven 86 87 32 (1977) 41 73 32 9 Ridge Road 36 33 27 Ri ley (1973) 12 15 12 18 10 14 9 49 45 11 23 8 3 Roth (1974) Spring Creek #1 46 33 Spring Creek #2 39 15 49 14 Spring Creek #4 9 10 3 Thrasher 65 8 (1978) 2 76 63 12 11 o 49 o 59 73 f.3 47 53 54 56 37 a a o o o o 65 84 51 50 55 59 NC 7 a o 17 13 26 II 50 3 Tu'rkey (1982) Ute-North (1979) Ute-South (1978) 26 18 30 17 42 42 40 18 12 21 21 o 16 27 27 30.9 31.9 31.1 39.5 43.5 38 37 34 Wattenburg 24 22 33.1 27.7 Average/Lek 5 37 16 14aIden Reservo i r (1973) #2 o 40. I I: 1.2 18 21.2 �11 males on leks in spring 1984 were low because some males had inadequate body reserves to support breeding activities. Therefore, an unknown proportion of the male population, while present, did not attend leks or attended leks only sporadically in spring 1984. This hypothesis is supported by observations of flocks of males foraging in early morning in May when they should have been on leks, sporadic and weak display by those males on leks. few males roosting on leks at night, excellent production of young in 1984, and no change in the proportion of 2nd and 3rd year recoveries of males banded in 1982 and 1983 (harvested in 1984). If this hypothesis is correct and the 1984-85 winter is average, the mean number of males counted per lek in spring 1985 should be similar to the average in 1983. Pellet Transects All 30 pellet transects were counted and cleared as desired except that transects were not counted nor cleared in fall 1984 due to time constraints. To date, no apparent patterns are available except that use, as measured by pellet counts, is higher in winter than in summer and that use in northwest North Park (north and west of Independence Mountains) is significantly less than in the mining area (Tables 3. 4). It is also apparent that transects 174 (mining area), 19, and 40 (northwest area) do not intercept areas used by sage grouse. At present, the large variability in pellet counts by season, year, transect, and area suggests that randomly located O.6-km transects have little merit as an index to sage grouse use or population size. Harvest Data Collection The sage grouse hunting season in North Park opened one-half hour before sunrise on the 2nd Saturday in September (8th) in 1984 and closed at sunset on 7 October, the same length (30 days) as in 1982-83. The daily bag limit was 3 with a possession limit of 6 (TabJe 5). Table 5. 1973-84. Years 1973-74 1975 1976 1977-80 1981 1982-8'" Sage grouse season length and bag limits, North Park, Colorado, Season length (days) 3 9 9 16 23 30 Bag/possession 2/4 2/4 3/6 3/6 3/6 3/6 Iimit �Sage grouse droppings/day, Table 3. Transect 3 4 48 57 60 74 78 81 83 88 99 115 129 136 166 174 179 194 196 197 Average aND = Summer 1979 "Ii nter 1979-80 Summer 1980 Winter 1980-81 0 0 0.01 0.02 " .27 0.36 0.50 0.79 0.84 0.09 0.56 0.05 1.61 0.71 0.40 0 4.62 0.03 0.39 0.03 2.06 0.76 O. 11 0.07 0.03 2.20 0.51 0.59 0.06 3.24 0.48 0.02 0.93 0.29 0 0.41 0.21 1.02 0.08 0.03 0.24 1.43 0.06 1.02 0.08 0.60 0.69 0.02 0.06 0.84 0.25 o. 11 0.52 1 .22 0 0.05 0.05 0.21 0.12 0.06 0.04 0.07 0.46 0.39 0.03 0.32 1.47 0.13 0 0 0.27 0.12 0.69 0.03 0 0.55 0.12 0.07 0.59 0.61 0.65 0.38 0.27 no data. O. 11 ...• mining area, North Park, Colorado, 1979~84~ N \.Jinter Summer Summer 1981 .. 1981-82 1982 \.J inter 1982-83 Summer 1983 Winter 1983-84 0.09 0.01 0.01 0.28 0.01 0.05 0.27 0.36 0 0 0 0.18 0 0.53 0.04 0 0.44 0.01 0.01 0.05 0.21 0.04 0.01 0.81 0.54 0.10 0.71 0.25 0.68 0.01 0.03 0.24 1.68 0.45 0.01 0 0.48 0.02 0.09 0.17 0 0 0 0.13 0 0.09 0.14 0 0.10 0 0 0.87 0.05 0.09 0.01 0 0.13 0 0 0.01 0 0.04 0 0 o. 11 1. 13 1.22 0.05 0.07 0 0 o. 11 0.89 3.44 0.07 0 0.10 0.44 0.20 0 0 0.02 0 0.87 0 0 0 0.18 0 0 0.74 0 0.05 0.02 0 0.01 0 0 0.02 0 0.03 1.31 0.06 0.35 0.58 0.18 0.02 0.14 0.22 0.01 0.21 0.17 0.55 0 0 0.01 0.84 0.19 0.39 0.12 0.33 0.08 0.39 0.10 0.26 0 Summer 1984 NOa �4. Sage grouse droppings/day~ Table Transect 12 17 19 40 49 50 63 76 80 95 Average a NO = Northwest North Park. Colorado~ 1979-84. ~Iinter 1983-84 Winter ~980-81 Summer 1981 Winter 1981-82 Summer 1982 ~J inter 1982-83 Summer 1983 0.01 0.27 0 0 0 0.01 0 0 0.06 0 0 1.64 0 0 0 0.02 0 0 0.63 0 0 0.15 0 0 0.05 0 0 0 0.38 0 0.52 1.54 0 0 0 0.18 0 0.71 0.06 0 0.30 0 0 0.21 0.04 0 0 0·39 0 0 0.13 0 0 0 0 0 0 0.01 0.01 0 0.04 0.23 0.06 0.25 0.07 0.01 0.15 Summer 1984 a NO 0.23 0 0 0 0.08 0.01 0.03 0 0.38 no data. - w �14 Two check stations were operated, one at Willow Creek Pass on Colorado 125 and the other east of Gould on Colorado 14, during both days of the opening weekend versus both days of the opening weekend and the 2nd Sunday in ear-lier years (1975-83 except neither the Gould nor Stateline check stations were operated on the 2nd Sunday in 1976; the Stateline check station was not operated on the 2nd Sunday in 1979. and the Muddy P~ss check station was operated only on the first Sunday in 1980). Neither the Muddy Pass (operated only in 1974 and 1980) nor the Stateline (operated from 1974 through 1979) check stations were operated. As In previous years of operation (1974-83 for Willow Creek, 1976 and 1979-83 for Gould), each station was open from about 1000 to 1800 hours MDT depending upon traffic volume. These stations were staffed with 2 research and at least 3 regional personnel. In addition, either 2 or 3 students from the Wildlife Management Techniques class at Colorado State University assisted at each check station. Data obtained per party were: county of origin, number of hunters, hours hunted (total of all hunters in each party), birds observed, birds bagged, birds lost, number of banded b j rds and 1ocat ion where each was harvested, and area hunted within North Park. In addition, hunters were questioned about their previous sage grouse hunting experience in North Park. One wing was obtained from all birds possible. Data collected from grouse hunters who hunted in areas other than North Park were tabulated and analyzed separately. Volunteer wing collection barrels and signs were placed along Colorado 14 northeast of Muddy Pass and near the top of Cameron Pass. north of Threeway on Colorado 127, and at Willow Creek Pass on Colorado 125. Volunteer wing collection barrels were available to hunters during the entire season near Three-way and Muddy Pass and for al J days that check stations IrJerenot operated east of Gould and at Willow Creek Pass. Barrels were not placed at Arapahoe Creek (only in 1982-83) nor Seymour Reservoir_(1982). Some wings were also obtained through field checks by Arapahoe National Wildlife Refuge and CDOW personnel. Hunter and Harvest Data. Check Stations.--Numbers of hunters checked continued the decl ine that started in-1980 and were significantly less than the high of 730-738 hunters checked in 1974 (3-day season, 2-day check) and 1975 (9-day season, 3-day check) (Table 6). Numbers of birds harvested and birds per hunter also decreased in 1984. These data suggest that the sage grouse population in fall 1984 was lower than in 1983 but not as low as in 1974-77 or 1980. Reported crippl ing J05S was low (2.6%) in 1984 while hunter efficiency increased. Thus, it appears that when number of birds observed decreases. crippl ing Joss also decreases while hunter efficiency increases (Table 6). While the estimate of crippling loss may be low because of reporting bias, some birds reported as crippled and lost are retrieved by other hunters and are included in their bag. It would thus be inappropriate to assume that crippl ing loss is substantially higher than that reported. A high estimate would be 10%. The hunter efficiency calculation (bird .retrieved divided by birds observed) indicates that sage grouse are not as easily harvested as commonly believed. �15 Table 6. Sage grouse harvest statistics, N Year 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 hunters checked N North Park, Colorado, 1974-84. birds ·observed N birds harvested Hunter efficiency (%) Cr ipp 1 i ng loss (%) 6,062 5,735 3.393 3,303 4,922 6,910 5.000 6,189 3,886 4,961 2.814 785 551 459 385 480 982 794 695 570 792 517 12.9 9.6 13.5 11.7 9.8 14.2 15.9 11.2 14.7 16.0 18.4 5.1 7.1 5.7 10.6 5.7 4.7 4.3 5.4 4.0 4.8 2.6 730 738 .595 353 350 521 567 523 515 474 396 Birds per hunter 1. OJ 0.7 0.8 1.1 1.4 1.9 1.4 1.3 1.1 1.7 1.3 Sage Grouse Hunter Experience.--The percentage of sage grouse hunters who normally hunt sage grouse in North Park has not markedly changed from 1974 to 198L} (Table 7). Despite more conservative (16 days, 2 and 4 or 1 and 2 bag and possession Iimits) season length and bag and possession limits outside of North Park, the percentage of hunters who normally hunt sage grouse elsewhere did not markedly increase in 1984 (14% vs. 13% in 1983). However, from 1974 to 1984 the percentage of hunters who normally hunted sage grouse elsewhere has increased from 8 to 14% of the total while firsttime sage grouse hunters have decreased by almost one-half(2~28% in 1974-79 to 13% in 1984). However, there has been substantial turnover in hunters each year as measured by the percent of first-time hunters provided that overall hunter numbers have remained stable. This suggested hunter turnover could be caused by (1) a lack of success in bagging a sage grouse or (2) a disl ike of sage grouse once they are harvested and eaten. Table 7. Previous sage grouse hunting experience of sage grouse hunters, North Park, Colorado. 1974-84. Year 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 Norma 11y hunt Norma Ily hunt First time sage grouse hunters in No rt h Par k elsewhere N N % % % .lL 68 65 65 65 20 4"1 40 32 251 )07 360 340 346 351 73 66 26 8 297 73 73 100 48 56 62 56 16 18 9 11 13 14 183 432 366 211 63 68 69 71 8 7 7 10 65 187 159 82 66 87 107 115 103 79 52 24 28 28 25 19 18 19 23 20 16 13 Regulations Bag/ Season possession length 2/4 2/4 3/6 3/6 3/6 3/6 3/6 3/6 3/6 3/6 3/6 3 9 9 16 16 16 16 23 30 30 30 �16 Hunter Success.--Hunter success decreased in 1984 but was well within the range measured in the 1974-83 interval (Table 8). Birds per hunter and hunter success have correlated well in all years as has the percent of hunters who have achieved at least a 1-day bag limit~ Of interest are the data that indicate that no more than 30% of all hunters achieve the bag limit in 1-day and that only 1-5% of all hunters achieve a 2-day limit on the opening weekend. Table 8. Year i974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 a Sage grouse hunter success • North Park, Colorado. Successful (%) 59 46 40 54 56 68 60 62 52 65 53 b Achieved bag limit (%)c 1 day 2 days 27 20 9 15 19 30 24 20 16 27 20 limi t ----- 1 1.1 0.7 0.8 1 1.1 2/4 3/6 3/6 1.4 3/6 1.9 1.4 1.3 1.1 1.7 1.3 3/6 3/6 3/6 3/6 3/6 3/6 5 1 < Birdsper hunter 1974-84. 5 5 2 2 2 3 3 2/4 aDerived from check station data. bHarvested at least 1 sage grouse. cPercent of total hunters checked. Hunter Origin.--Vehicle 1 icense prefixes were recorded at check stations;to ascertain origin of hunting parties. About one-third to one-half of all hunters originated in the Denver metropolitan area each year with Larimer County being the singJe most important point of origin (20 to 33% of all hunting parties). Boulder and Weld counties were also important contributors to sage grouse hunters in North Park (Table 9). It should be noted that few hunters were checked from Jackson County even though they compr ised about 10% of all persons obtaining hunting permits during the 1974-77 special seasons in North Park. Local hunters were not checked as the check stations were placed at exits to North Park. �17 Table 9. Origin of sage grouse hunting parties, North Park, Colorado, 1974-84.a County (%) Year Denver Metrob Lar imer 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 50 52 53 47 52 33 42 39 43 50 50 24 20 20 23 21 31 26 33 27 21 22 aAscerta ined from vehicle b Adams, Arapahoe, Boulder 10 11 13 16 12 17 13 11 12 8 6 Weld 5 6 6 5 4 9 7 7 6 6 8 All other 11 11 8 9 11 10 12 10 12 15 14 license prefixes. Denver, Douglas, and Jefferson counties. Time Distribution of Harvest.-~Sage grouse wings were obtained at check stations, wing barrels, and field checks. In 1984, 70% were obtained during the first weekend and 3% from the last weekend (Table 10). With liberalization of the season starting in 1977 (9 to 16 days), the proportion of the harvest that occurred in the opening weekend decreased from 92-99% to 61-73% (Table 10). This decrease reflects reduced hunter pressure during the opening weekend as there is no evidence that total harvest or hunter numbers have markedly decreased. There is some evidence that total harvest and hunter numbers increased slightly with more liberal seasons. It is important to note that some sage grouse hunters are active throughout the entire season even though hunter pressure is quite light after the 3rd weekend. Harvest and Hunting Pressure.--Leading hunting and harvest areas in North Park in 1984 were near Lake John. Petersen Ridge-MacFarlane Reservoir, Ridge Road, and Spring Creek-Owl Ridge. These 4 areas had 77% (77.4) of the hunters and 81% (81.2) of the total harvest on opening weekend (Table 11). While these regions have been the major hunting areas each year since 1974, their contribution has fluctuated over time (Table 11). However, hunter pressure and harvest has not paralleled changes in sage grouse populations (as measured by lek counts) in North Park during 1974-84. �18 Table 10. Time distribution Colorado, 1974-84. Year 1 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 99 94 92 67 62 73 69 72 61 72 70 of sage grouse wings received, North Park, Weekend 2 (% received)a q 3 5 .Season 1eng th (days) 3 4 9 7 13 6 11 12 8 13 6 7 9 16 8 10 16 6 16 9 6 7 3 6 16 2 4 2 2 2 4 3 23 30 30 30 a Totals do not equal 100% as wings collected during the week are not included. Age and Sex Structure.--Immatures comprised 57% of the fall harvest in 1984, similar to the excellent production in 1979 (57.7%) and 1983 (57.4%) and above the long-term average of 51% (Table 12). The percentage of yearlings in the harvest was also high (23.4%) while the percentage of adults in the harvest was the 2nd lowest documented (19.6%). These data indicate that nesting success and survival of young in 1984 was good to excellent, and that survival of young produced in 1983 was good. However, survival of adults may have been low (if the population was stable) or appeared low because of the excellent production of young and good overwinter survival of young from 1983. The latter explanation is most plausible as there was no indication of disproportionate mortality of either adult males or females (22:78 not significantly less than the 22.5:77.5 ratio in 1983) (Table 12). The long-term data from North Park indicate that in the average year about 51% of the fall population of sage grouse is comprised of young of the year, 22% are young of the previous year, and 27% of the birds are at least 2 years of age. With this age structure, it is obvious that poor production in 1 or more consecutive years would result in a markedly reduced population of sage grouse. The data (Table 12) also document that sex ratios of yearl ings and adults strongly favor females (yearl ings = 63:37, adults = 72:28). Thus, poor production would most noticeably affect males as yearlings comprise about 52% of the spring population of this sex each year. Poor production should be immediately noticed in counts of males on leks the following spring. Without recruitment, numbers of males on leks could decrease by 50% in 1 year. �.•. Table Sage grouse harvest and hunting 11. lndependence Mountain Lake John pressure Wa 1den Reservoir (% of total) within North Park, Colorado, Rid~e Road 1974-84.a,b Peterson Ridge-HacFarlane Res. Pole Mountain Spring Creek-Owl Ridge Year Hunt. Harv. Hunt. Harv. Hunt. Harv. Hunt. Harv. Hunt. Han! . Hunt. Harv. Hunt. Clarv. 1974 4.7 6.0 18.1 16.4 12.4 15.4 9.6 11.3 4.7 3.8 20.3 19.6 17·9 1975 3.1 3.8 24.3 24.7 14.2 16.7 11.1 15.6 1.8 2.0 17.2 14.3 1976 2.3 3.0 28.2 21.3 13.0 12.0 13.0 21.1 2.9 4.5 18.5 21.3 Michigan River SE Ea~le Hill Hunt. Harv. Hunt. Harv. 13.4 2.2 1.3 10.0 12.8 13.6 6.2 2.6 3.8 10.4 9.8 10.9 6.3 3.1 5.9 8.0 5.0 2.0 2.1 7.4 13.2 9.6 1977 1.4 0.5 28.9 23.9 15.8 12.2 16.9 20.8 3.2 6.5 16.3 12.7 8.0 8.1 1978 0.9 0.0 29.4 18.1 18.3 10.6 14.6 13.7 2.0 2.5 21.1 36.7 6.0 3.1 1.7 5.6 6.3 1979 3.5 2.7 23.0 19.7 8.3 7.5 10.2 13.8 4.0 4.2 30.0 33.3 12.5 9.11 2.5 3.7 6.0 5.7 13.0 2.8 1.5 1980 2.0 2.3 16.6 15.5 10.2 14.9 14.0 8.1 1.4 2.6 29·0 32.7 14.1 9.9 9.4 1981 3.3 1.7 . 15.9 14.4 4.2 4.0 14.8 27.6 1.7 0.7 27.7 26.3 14.9 12.9 1.1 1.7 11.5 10.5 1982 2.3 3.2 19.7 20.6 7.8 8.2 18.4 22.1 1.6 0.1 21.9 21.2 18.6 12.1 1.4 2.7 8.4 9.3 26.9 9.6 12.0 11.9 8.8 0.2 0.8 18.9 21.5 15.7 13.9 3.6 3.4 8.7 7.2 22.5 6.9 5.0 15.3 20.6 1.0 0.0 20.6 24.7 15.5 13.4 1.8 1.7 9.4 6.7 1983 4.7 5.5 26.6 1984 ·1.0 5.2 26.0 aData from check stations only. bTotals may not approximate 100% as in most years some hunters and birds harvested could not be allocated to a particular zone. I..D �N o Table 12. Age and sex composition of the sage grouse harvest, North Park, Colorado, 1974-84. Males Year N 197~ 1975 1976 171 101 10~ 1977 1978 136 184 1979 1980 1981 1982 1983 1984 306 207 23~ 197 295 236 Avg. 48.9 47.6 119.5 ~6.9 47.8 ~9.0 4~.9 47.9 50.1 45.6 48.5 47.7 Immatures Females N % 179 111 106 154 201 318 25~ 255 196 352 251 51.1 52.11 50.5 53.1 52.2 51.0 55.1 52.1 ~9.9 54.~ 51.5 52.3 Total N :(; N 350 212 210 50.1 42.0 42.3 49 52 46 290 385 624 461 ~5.9 53.2 ~7 62 102 80 ~89 393 647 487 Yearlings Females Males 57.7 49.1 ~7.~ 50.6 57.~ 57.0 51.3 :(; 35.5 46.8 78 39.3 38.2 ~3.~ 39.8 32.0 34.1 53 90 68 39.8 36.7 34.0 37.4 -N------z 89 611. 5 59 71 53.2 60.7 61.8 56.6 60.2 68.0 76 81 154 170 151 80 155 132 65.9 60.2 63.3 66.0 62.6 Males Total N % 138 111 19.8 22.0 117 123 1113 23.5 19.5 19.8 256 250 229 133 245 200 23.7 26.6 22.3 17.1 21.7 23.4 21.9 N % Adu Its Females N % ~5 21.~ 165 78.6 55 49 30.2 28.8 127 121 1,8 21.9 3~.2 171 129 129 158 227 170 183 131 69.8 71.2 78.1 65.8 6~.2 67 72 70 87 81 53 37 35.8 30.7 27.7 32.2 22.5 22.0 28.0 69.3 72.3 67.7 77 .5 78.0 72.0 Total N % 210 182 30.1 36.0 3~.2 3~.6 27.1 18.6 170 219 196 201 228 31~ 251 236 168 24.3 30.4 32.3 20·9 19.6 26.8 Sample size 698 505 ~97 632 724 1,081 939 1,032 777 1,128 855 �21 Nesting Success and Production.--Primary feather molts of hens harvested were classified (number of primaries from the previous year still retained) and compared to the primary feather molt patterns of males of the same age-class harvested in the same time interval. Estimated nesting success of all hens increased in both 1983 and 1984. Of importance is the good success of yearling hens in both 1983 and 1984 (Table 13). The data available indicate that at least one-half of the adult hens are successful in most years (except 1981) while success of yearling hens is highly variable (> 50% in only 3 of 11 years). In years when at least 50% of the yearlings are successful (1979, 1983, 1984), chicks per hen increase as does hunter success as measured by birds per hunter data from check stations (except in 1984). It is probable that the percentage of successfully nesting females was overestimated in some years (1974, 1975, 1977, and possibly in 1984) and underestimated in others (1980, 1981). Because this estimate is based on the molt patterns of primary feathers which are affected by timing of breeding (males) and nesting (females), it is logical to assume that timing of breeding ~vents (weather controlled) is the most important factor involved. Thus, underestimates are derived following "early" springs whi Ie late springs (for example, 1984) may provide the most reliable estimate. The evidence indicates that, at least in North Park, the percentage of juveniles in the fall harvest closely approximates overall nesting success. Also, hunter success (birds/hunter) closely follows the young per hen ratio in the harvest. Table 13. Sage grouse nesting success and production Colorado, 1974- 84. Year 1974 1975 1976 1977 1978 1979 1980 '1981 1982 1983 1984 Estimated nesting success Yea r+ All Adults lings hens 65 53 53 59 60 65 56 37 59 68 71 46 39 27 33 38 56 31 22 38 51 55 59 49 43 50 51 60 43 31 52 58 63 Young in harvest(%) 50 42 42 46 53 58 49 47 51 57 57 rates, North Park, Young per hen Young per successful hen 1.4 1.1 1.1 1.2 1.8 2.2 1.4 1.3 1.5 1.9 1.9 2.3 2.3 2.5 2.0 3.6 3.7 3.3 4. 1 3.0 3.2 2.9 Birds per hunter 1.1 0.7 0.8 1.1 1.4 1.9 1.4 1.3 1.1 1.7 1.3 Timing of Hat ch lnqv-r-He t ch lnq was delayed in 1984 and over 80% of all chTdks hatched after 28 June. This was about 2 weeks later than in 1983 and 3 weeks later than in 1982 (Table 14). Timing of nesting events was later each year from 1982 through 1984 because of heavy and late lying snow cover, In contrast, 1981 was an "early" year and over 80% of the hatch �22 occurred prior to 21 June. Data in Tables 13 and 14 suggest that nesting success and production are better in "Ja te" years (1979. 1982, 1983, 1984) than in "early" years (1976, 1977, 1981) although 1975 appears to be an exception. This relationship appears to be related to moisture conditions which could affect (improve) vegetative cover during the nesting period and/or provide adequate forage (succulent forage, insects) near nesting sites for young chicks. Theoretically, early chick survival should be enhanced if successful hens did not immediately travel to meadows along streams to forage following hatching of their clutches. Table 14. Estimated Colorado, 1974-84. timing of hatching of sage grouse eggs, North Park, Year (Percent of all chicks) 74 interval 75 i6 77 78 79 80 11-17 May 81 82 83 <1 18-24 May" 2 <1 (. <1 25-31 May { <1 2 14 6 12 21 25 22 4 2 4 31 30 20 9 3 35 28 9 21 24 27 12 27 19 19 26 22 20 14 10 8 34 43 6 19 27 16 21 11 8 8 17 21 8 8 19 51 15 6 2 7 9 11 2 8 15 26 13-'19 Jul 10 <1 4 2 3 <1 3 5 3 20-26 Jul 7 <1 <1 <1 <1 2 2 2 <1 <1 <1 <1 <1 <1 <1 <1 3 1- 7 Jun > 16 8-14 Jun > 18 15-21 Jun 22-28 <1 16 6 6-12 Jul 6 27 Jul-2 Aug Annual Turnover.--Estimates of annual turnover rates were calculated for adult males (Table 15) and adult females (Table 16). The estimated annual mortal ity rate was 52% (1974-84 average) for adult males and 42% for adult females. These estimates are based on the premise that the percentage of yearlings in the population should equal the annual mortality of adults if the population is stable. However, if the population was increasing then the percentage of yearlings in the population would overestimate adult mortal ity rates. Because the sage grouse population in North Park obviously increased in the 1974-84 interval (except 1984), the calculated mortal ity rates for both adult males and females may be biased upward by several percentage points. �23 Table 15. Estimated annual turnover of adult male sage grouse, North Park, Colorado, 1974-84.a In harvest Year lings Adults Year N % N % 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 Avg. 45 55 49 48 67 72 70 87 81 53 37 48 51 52 50 52 41 47 53 60 37 35 48 49 52 46 47 62 102 80 78 53 90 68 52 49 48 50 48 59 53 47 40 63 65 52 Sample size 94 107 95 95 129 174 150 165 134 143 105 Estimated annual mortal ity of adult males in a stable population 52% aData from wing collections only. Table 16. Estimated annual turnover of adult female sage grouse, North Park, Colorado, 1974-84.a In harvest - Year N 165 127 121 171 129 129 158 227 170 183 131 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 Avg. Adults Yearl ings N % 89 59 71 76 81 154 170 151 80 155 132 35 32 37 31 39 54 52 40 32 46 50 42 % 65 68 63 69 61 46 48 60 68 54 50 58 Sample size 254 186 192 247 210 283 328 378 250 338 263 Est imated annual morta Iity of adult females in a stable population = 42% a Data from wing collections only. �24 Banding and Band Recoverl es -r-Dur lnq 1984, 428 sage grouse were banded in North Park of which 23lf were hens (Table 17). The high number of hens banded (the same as in 1977) was the result of the delayed spring and deep snow which concentrated hens and kept them in flocks later than in most years. Additional effort was also placed on locating and banding hens in 1984 prior to flock breakup. The reduced number of males banded in 1984 was the result of poor access, low lek attendance by males, and the delay in breeding activities which resulted in males roosting on leks at night for only a short period. i Table 17. Sage grouse band ing data, North Park, 1973-84. Number banded Females Year 1- 2+ AI I 1- Males 2+ 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 41 22 62 71 101 68 27 68 74 133 22 52 94 38 69 32 111 109 49 130 145 234 54 100 166 69 102 65 234 80 54 138 120 183 106 111 127 110 110 152 102 99 88 153 114 123 98 146 173 190 190 157 92 32 48 72 31 42 33 123 All 179 142 291 234 306 204 257 300 300 300 309 194 Recoveries were received from 59 banded sage grouse in 1984 (20 females, 39 males) of which all but 6 were shot by hunters. Recoveries represented bandings rn 1977-84 for females and 1982-84 for males (Tables 18 and 19). Recoveries of 2nd year and older birds (banded prior to 1984) occurred in the same proportion as in earlier years indicating that the severe winter of 1983-84 did not noticeably affect survival of older grouse. This strongly suggests that the low counts of males on leks in 1984 was not the result of 10\1\1 survival of adults. Older males were in the population at the rate experienced in earlier years of the study. Thus, it can be reasonably concluded that the low counts of males on leks in spring 1984 was because males failed to attend leks. This failure to attend leks was probably the result of poor body condition caused by the harsh winter of 1983-84 (J. Hupp, unpubl. data). Direct recovery rates (= harvest rates) of sage grouse banded in 1984 varied from 5 (f emel es ) to 9-12% (adult-yearl ing males). These rates are within the range documented in the 1973-83 interval (Table 20). These data indicate that liberalization of season length and bag limits have not markedly affected the overall harvest rate of sage grouse in North Park. �25 Table 18. Male sage grouse banding and recovery data, North Park, Colorado. 1973-84. N recovered {year} 2 3 ~ 5 6 3 7 6 7 4 0 3 4 3 1 1 2 1 2 1 1 1 0 0 0 0 1 1 0 0 0 1 1 2 2 3 3 3 2 2 2 0 0 0 0 3 0 3 1 0 N Age Year banded 6 Yearl ings 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 80 6 4 54 6 18 16 19 13 13 13 14 7 16 12 5 6 138 120 183 106 111 127 110 110 152 102 5 10 4 4 5 5 1 10 1 0 1 1 1 0 0 Adults 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 99 88 153 114 123 98 146 173 190 190 157 92 7 8 10 16 12 9 13 9 16 19 15 8 4 5 4 3 2 9 10 5 5 6 4 1 0 1 0 0 0 0 0 0 0 1 0 0 0 0 �26 Table 1.9. Female sage grouse banding and recovery data, North Park. Colorado, 1973-84. N recovered N Age Year banded {yead 'g-- q 5 6 7 4 1 1 1 0 2 0 0 0 0 0 0 2 0 1 1 1 1 0 0 0 0 2 0 0 0 0 1 0 2 3 Year 1ings 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1.• 1 2 22 62 2 71 2 6 2 1 2 4 5 101 32 48 72 31 42 33 123 6 6 5 0 1 0 7 8 4 2 0 6 4 4 3 0 2 0 1 0 2 0 3 0 5 2 1 0 1 1 1 1 0 2 0 0 4 1 2 4 3 0 3 6 1 0 0 0 1 0 0 1 1 0 0 0 0 0 Adults 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 68 27 68 74 133 22 52 94 38 60 32 111 6 2 13 1 3 8 5 5 0 6 2 3 1 2 0 0 2 0 0 1 0 1 0 0 0 1 0 0 1 0 0 0 0 1 0 0 0 0 0 1 �27 TabJe 20. Direct recovery rates of banded sage grouse, North Park, Colorado, 1974-84. ----Year 1- 1973 1974 5 9 1975 10 1976 1977 1978 1979 1980 1981 t 982 1983 1984 3 6 16 15 Females (%) 2+ All 7 7 9 3 10 4 6 6 8 9 11 13 12 13 7 0 5 5 7 10 10 14 10 11 9 5 7 9 8 13 8 7 11 7 14 13 0 All 3 'i1 5 2+ 13 13 10 10 0 Males (%) 1- 6 9 10 9 6 8 10 11 10 12 9 10 13 12 10 8 10 8 10 10 All birds (%) Season Length Limits 2/4 7 9 10 9 9 11 10 8 16 16 11 23 3/6 3/6 8 30 30 30 3/6 3/6 8 7 3 3 2.1 If 9 9 16 2/4 3/6 3/6 3/6 3/6 16 3/6 __ .·__....__ ._._~._u Estimate of Total Harvest.--Total harvest in 1974-77 was estimated from a 100%-survey of all sage grouse hunters in North Park. This was possib'le because all sage grouse hunters were required to obtain a free permit prior to hunting. Because the ltJillowCreek Pass check station was the only one that was operated each year from 1974 through 1984, the number of hunters checked at it on opening weekend (2 days) was used to estimate total hunters from 1978 through 1984. From 1974 through 1977, an average of 30.7% of all North Park sage grouse hunters were checked at Willow Creek Pass. Thus, 0.307 was divided into the number of hunters checked at Willow Creek Pass each year from 1978 through 1984 to derive an estimate of total sage grouse hunters in North Park (Table 21). This number was then multiplied by the birds per hunter value calculated from check station data each year to derive an estimate of total harvest (Table 21). These data indicate that tot~l number of hunters declined from about 1,000 in 1974-76 to about 600-70. This is in agreement with the total number of hunters contacted at all check stations (Table 6) as total number of hunters contacted at check stations has decl ined from about 700 to 400-500. The data also suggest .rha t total harvest has decl ined (Table 21). However , s1 ightly more wings were received from hunters in 1979, 1981, 1982, and 1984 than the estimated harvest. Thus, the harvest was underestimated by the method used. This underestimate could be in the magnitude of 200-300 birds each year. It is reasonable to assume that the total harvest of sage grouse in North Park approximates 1,000-1,500 birds per year. �28 Table 21. Estimated total number of sage grouse hunters and total harvest, North Park, Colorado, 1974-84. Na hunters contacted Year 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 378 374 .317 149 226 173 234 208 203 216 200 Total hunters Birds per hunterC 966b b 1,187b 1.1 0.7 0.8 845d 736d 56J~d 762d 678d 661d 70~d 651 1.1 979b 1.4 1.9 1.4 1.3 1.1 1.7 1.3 Total wings received Total harvest b 1,193b 1~075b 847 1,070b 1,030e e 1.072 e 1,067 e 881 e 727 e 1 ,197 e 846 698 505 497 632 72.4 1" O[H 939 1.032 777 1,128 855 •• ____ a >r ••••••.. ______ • . Wi 110w Creek Pass Check Station only during opening weekend. bDerived from questionnaire North Park. cOerived checked) • survey of all sage grouse hunters in from check station data (total birds checked ~ total hunters dOerived by dividing percent of all hunters checked at Willow Creek Pass in 1974-77 (0.307) into number of hunters checked at Willow Creek Pass in each year. eOerived by multiplying total number of hunters by birds per hunter. Estimated Population Size.--Population size in spring was estimated assuming that peak lek counts represented 60% of all males associated with each lek and that 90% of all leks were located and counted. The estimated total number of males was multiplied by 2 to derive the total number of hens as harvest data (Table 12) indicate that females comprise 68% of the total adults and yearl ings. Fall population size was estimated by dividing the percent of adults and yearlings in the fall harvest into the spring population estimate. ~ The data available (Table 22) indicate that spring sage grouse populations in North Park increased from about 3,000 birds in 1974-75 to 7,000-8,000 in 1978-79 and were relatively stable at 6,000-7,000 birds from 1980 through 1983. While it would appear that the population significantly declined in 1984, harvest~ production, and recovery data strongly indicate that this did not happen. It: is expected that counts of males on leks in spring 1985 will return to levels similar to that in 1983. Size of the fall sage grouse population in North Park has fluctuated from about 6,000 to 20.800 birds. This reflects the relatively short 1ife span and high annual turnover (52% for males, 42% for females) of sage grouse and the importance of nesting success and production. �29 Table 22. Estimates of spring and fall sage grouse population Park, Colorado, 1974-84. N Year leks 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 19 19 21 26 34 35 30 31 31 31 22 Males per lek 28 31 32 31 40 44 40 41 41 39 21 Total a males 987 1,092 1,219 1.508 2,479 2.774 2,211 2.312 2.312 2.210 840 Estimated Spring Total b c population females 2,961 3,276 3,657 4,524 7.437 8.322 6,633 6,936 6,936 6,630 2,520 1.974 2,184 2.438 3,016 4.958 5.548 4.422 I~, 624 4.624 4.420 1,680 size. North Young in harvest (%) Est imated fall d popu 1at i0"__ 50 42 42 46 53 58 49 47 51 57 57 5,922 5.648 6,305 8.378 15.823 19,814 13.006 13.087 14, 155 15,419 5,860 aOerived by dividing males/lek by 0.6 (percent of males present at high count) and multiplying by number of leks (number counted plus 10%). bDerived by multiplying number of males by 2 as females comprise 68% of the adult and yearling population. c Total males plus total females. dDerived by dividing percent adults and yearlings the spring population estimate. LITERATURE in fall harvest into CITED Beck, T. D. I., R. B. Gill, and C. E. Braun. 1975. Sex and age determination of sage grouse from wing characteristics. Colorado Div. Wildl., Game Inf. Leafl. 49 (Rev.). 4pp. Braun. C. E., and T. D. I. Beck. 1976. Effects of sagebrush control on distribution and abundance of sage grouse. Final Rep., Colorado Div. Wildl. Fed. Aid Wildl. Rest. Proj. W-37-R-29. Pp.21-84. Giesen, K. M., T. J. Schoenberg, and C. E. Braun. 1982. Methods for trapping sage grouse in Colorado. Wildl. Soc. Bull. 10:224-231. Hoffman, R. W., and C. E. Braun. 1975. A volunteer wing collection Colorado Div. Wild1., Game Inf. Leafl. 101. 3pp. station. Remington. T. E. 1983. Food selection, nutrition, and energy reserves of sage grouse during w(nter, North Park. Colorado. M.S. Thesis, Colorado State Univ., Fort Collins. 89pp. Schoenberg. T. J. 1980. Potential impacts of strip mining on sage grouse movements and habitat LIse. Job Prog. Rep., Colorado Div. Wild!. Fed. Aid Proj. W-37~R-33. Pp. 245-264. �30 Prepared by ---::.::;~....::!~~~:?=:..::-~..:..:~::....::.:::.:....:......:-=-__ Clait E. Braun Wi ldl ife Research leader �Colorado Division of Wildlife Wildlife Research Report April 1984 31 JOB FINAL REPORT State of Project Colorado W-37-R-37 Work Plan 3 Job Title: Period Covered: Author: (45-01-504-15050): Job: Game Bird Survey 14 Dispersal and Recruitment of Juvenile Sage Grouse 1 July 1983 through 30 June 1984 Peter O. Dunn Personne I: ABSTRACT Natal dispersal, led fidelity (attendance within and between years), and summer habitat of sage grouse (Centrocercus urophasianus) were studied on Cold Spring Mountain, northwestern Moffat County, Colorado, from July 1981 through June 1983. There was no difference between male and female natal dispersal distances, although the sample size was small (N = 25 individuallymarked year1 ings). Sixty percent of marked yearling grouse identified in spring attended the lek closest to where they were marked as juveniles. Sixteen percent of all individually-marked juveniles (25/157 birds) were known to have attended leks as yearlings. There was no difference between yearling and adult female lek attendance rates; however, yearling males attended leks less often than adult males. These differences may be related to yearlingsl inexperience with breeding. Fall sage grouse dispersal was analyzed from 118 relocations of radio-marked sage grouse (N = 8) during July-November 1981 and 213 relocations (N = 10) 'dur i nq August-=-November 1982. Grouse steadily moved away from capture sites until November each year when they moved to wintering areas. Movements to winter range in late November were related to snowfall and subsequent availabil ity of sagebrush (Artemisia spp.) cover. Maximum one-way distance to wintering areas was 30.3 km (N = 4 radio-marked grouse). There were no differences between males and-females. Summer habitat use by 14 radio-marked grouse (5 juveniles, 7 unsuccessfullynesting hens, 2 hens with broods) was analyzed from 192 transects at sage grouse locations and 40 randomly located transects in 1982. Univariate and stepwise discriminant function analyses (DFA) indicated that sage grouse used sites closer to the edges of cover types with greater horizontal and canopy covers, and more homogeneous shrub densities and intercept distances than average. Within these sites, grouse roosted beside shrubs which were larger than average. There were no differences in habitat use among juvenile and adult female categories (unsuccessful and brood hens) when tested with DFA. In areas sprayed with herbicide, grouse did not use sites with greater shrub canopy cover than average probably because of a Iimited amount �32 of available shrub cover. Although shrub cover and structure may be important correlates of sage grouse habitat use during fall to spring, in summer, the importance of forbs in grouse diets alters the basis of habitat selection. Sage grouse summer habitat should be managed for more homogeneous than average sagebrush structure and density characteristics within sagebrush stands. Among stands, these shrub characteristics, and also cover types, should differ, and they should exist in close proximity. Prepared by Q~k.-tJ. /!U-"P--r</ Peter O. Dunn ~~ Graduate Student App roved by __,(t=C2=d,,--=l;_. -"-~_-,-,-.,J.~, __ Clait E. Braun rfl3 Wildlife Research Leader �33 COLORADO STATE UNIVERSITY Spring. 1984 WE HEREBY RECOMMEND THAT THE THESIS PREPARED UNDER OUR SUPERVISION BY PETER OWEN DUNN ENTITLED AND RECRUITMENT OF JUVENILE SAGE GROUSE FULFILLING DISPERSAL BE ACCEPTED AS IN PART REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE. Committee on Graduate Work cRt~/S:- a Department Head ~ ii �34 ACKNOWLEDGMENTS Financial support for this project Division of Wildlife. Federal and the Rob and Bessie The support ledged. of these Foundation for serving James Teer, grateful to Drs. appreciate Dr. about Clait Braun field supervision sincere interest Several J. the assistance support, this study. and editorial people assisted thank Jerry assistance, enthusiasm, acknow- Welder Wildlife and Fred Samson acquired review of this thesis, funding. and professional of literature. and provided appreciate his growth. C. Braun. S. Steinert, this Ronald Ryder I especially Hupp , Tom Olsen, reviewing and donations me in the field: R. Ryder, I especially of Dr. critical assistance. in my personal Hupp , T. Olsen, Director, W-- 37- R. Texas. is gratefully Philip Lehner the local avifauna, initiated Sinton. committee and for critically for academic and administrative many discussions Project and hospitality. on my graduate I sincerely and organizations Dr. for his interest I am particularly thesis. Welder Wildlife Foundation. thank by the Colorado Aid to Wildlife Restoration agencies I especially was provided K. Giesen. S. Vana , and D. Ward. and Don Ward for their and good humor during many long days in the field. I thank their help: the following Colorado Division of Wildlife personnel J. Black, C. Brown, W. Rusaell , and C. Woodward. even after endless typing M. Bauman. Jann corrections. v J. Haskins. Black was especially The interest for B. Mangus, courteous and assistance of �35 John Parks and Herb Conley of the Bureau acknowledged. my thesis. Dr. David Bowden reviewed His attention Wright Dickinson, area. provided frank discussions access I also thank the statistical aspects of to detail is appreciated. the principal to his land. about ranching Lisa Bennett, They all encouraged private landowner I thank on the study him for his help and for and sage grouse; Brian Wakelyn for somewhat less tangible, ance. of Land Management are Cade vi Curt Mack. and Leslie but nevertheless me with their discussions. D friendship, important assist- interest. and �36 TABLE INTRODUCTION. 1 Study 3 Area LITERATURE II. CITED 7 NATAL DISPERSAL AND LEK FIDELITY SAGE GROUSE •.•••••• Study OF 8 10 Area and Methods ....•. 12 Natal Dispersal 12 Recruitment 13 Results Discussion. . . LITERA TURE III. OF CONTENTS 19 CITED 23 LATE SUMMER-SPRING MOVEMENTS JUVENILE SAGE GROUSE. • Study Area and Methods Results OF 26 . 27 .. 29 Discussion LITERA TURE IV. 40 • • • • CITED SUMMER HABITAT USE BY ADULT FEMALE JUVENILE SAGE GROUSE.. ••••• Study Results Area and Methods Univariate Multivariate Analysis AND . • • . . . . • . • • • • . Analysis 47 49 . ••. . . . . • • . . . • . . 44 53 53 60 vii �37 TABLE OF CONTENTS Discussion • • . . . . . . Management LITERATURE Implications CITED • 0 • (Cont'd) 63 70 72 • r r viii �38 LIST OF TABLES Table 2.1 Page Sage grouse lek observations, Moffat County, Colorado. 2.2 Sage grouse Mountain. Cold Spring March-May Moffat County, 1982-83 (%, ~ days), lek attendance Colorado, Mountain. 14 •.•.• Cold Sprin g • March-May 1982-83 . 2.3 Inter-lek movements Mountain, 3.1 16 of sage Moffat County, R 2 and probability (~)a sion models of sage sites date, Variables November parametric test grouse distances Variables random June-September 1981 and use sites locations Moun- 1 August32 differing among ran- for parametric Cold Spring (!: < 0.05) 18 regres- on Cold Spring June-September significantly and grouse 1983. from capture from previous (!: < O. 05) statistics, Coun ty , Colorado. March-May . . . • . . significantly dom and sage 4.2 grouse 1982. Cold Spring of simple and multiple and age of bird 15 July-14 14 November 4.1 Colorado, (km) with movement rate (m/day), tain. grouse, or non- Mountain, Moffat 1982 • differing use sites. a Cold Spring 54 between Mountain. 57 1982 ix �39 LIST OF TABLES (Cont'd) Table 4.3 Page separating use sites following stepwise analysis. Cold Spring June-September 4.4 Canopy cover forb ground juvenile r r (!: < Variables brood discriminant Mountain. function Moffat County. Colorado. 1982 . (%) cover 62 and height (%) sage grouse, September 0.05) random and sage grouse (ern) of sagebrush. at relocations Cold Spring 1982 and results of radio-marked Mountain. of previous habitat and June- studies of 67 x �40 LIST OF FIGURES Figure 1.1 Cold Spring Mountain study Moffat County. 3.1 Distance marked (N = Summer to spring sage grouse (3 males; function Colorado, 31 sage grouse Colorado, July- 1982 .• 1 female; Moffat County, Discriminant use sites. Colorado 36 movements of 4 radio-marked May 1982-83 County. juvenile Mountain, 9 August-28 grouse • in 1982 moved (krn) from 1981 and August-November Cold Sprin g Mountain, 4.1 Moffat County, and mean distances 18) on Cold Spring juvenile sites by 8 radio- in 1981 and 10 grouse Mountain, 5 . sites of radio-marked November 3.3 Colorado sage grouse Mean angles capture northwestern (krn) moved from capture on Cold Spring 3.2 area, #452) on Colorado, .....••.... scores Cold Spring of random and sage Mountain, June-September xi 39 Moffat 1982 •••• 65 �41 INTRODUCTION In the western United States, land by agricultural ~ r r and energy development upon sagebrush (Braun species dependent grouse are dependent populations upon sagebrush are indicators development techniques will become increasingly Research is being on mitigating adverse conducted Although generally adquate , there habitat in summer local populations sage grouse maintenance seasonal To better ducted 9 of habitat understand to sage grouse Colorado; this thesis habitat habitat states on sage requirements as to whether sage concerning or if they contribute Without a better is grouse how to the knowledge it will be difficult on populations the importance Mountain. research northwestern of that of to pre- of sage grouse. of summer habitat biology and management, the results western It is not known if juvenile patterns. disturbance reports on sage grouse developments is no information natal area. in 1981-83 on Cold Spring local are accelerating. in Colorado and other and there use and dispersal Since sage useful. of sage grouse from available affect Because alterations is little information of leks in their diet the impacts rangelands are formed or maintained. select habitat dispersal knowledge 1976). quality. impacts of energy-related grouse. select et al. of sagebrush range- may adversely range of western to mitigate impacts of sagebrush for food and cover. of sagebrush and disturbance populations disturbance research use and was con- Moffat County. in 3 sections. �42 The overall juvenile objectives sage grouse of leks in their dispersal of this research produced natal area. were: characteristics identify summer habitat (1) determine in one area contribute (2) determine of juvenile sage sage grouse if to the maintenance if differences movements of male and female juvenile identify Study project occur between grouse, (3) fall movements. and (4) use by sage grouse. Area The sage grouse studied population from 1978 through been banded Counts Cold Spring grouse range prior of sage grouse were not established Cold Spring Mountains. away from the surrounding attending to this study. before The topography therefore, hills and mesas (Fig. 1.1). yearly relief. west to east. precise study of the eastern range Spring Three area boundaries extension from mountainous mountains. include: Mountain. of the Uinta to rolling Cold Spring Talamantes (2,622 m) , major topo- Creek which flows (Fig. 1.1). north which flows into the Green River; River which flows through Mountain Mountain was of Cold Sprin g Mountain : Vermillion Creek. and east of Cold Spring and the Green 1978. in the Western Hemisphere. area varies Major drainages north since on Cold Spring Middle (2,904 m) , and Diamond Peak (2,909 m) provide graphic 160 krn ", field work began. east-west of the study have known leks on and surrounding breeding Mountain is part the largest Mountain has been From 120 to >300 chicks/year Mountain have also been conducted The winter unknown 1983. with no recoveries of male sage on Cold Spring Browns Park south of Cold �43 • r t - - 11-1 o COLORADO ,KILOMETERS Fig. 1. 1. Court ty. Cold Spring Colorado. Mountain Con tour study intervals area. northwestern are 305 m, Moffat �44 Northwestern cipitation annually Cold Spring which Colorado's climate is semiarid with 20-51 cm of pre- depending on elevation Dep . Inter. Mountain receives 11% falls during the driest months occurs from November often remain snow-covered January (U.S. being Cold Spring occupies Snowfall at lower eleva- from late October (Artemisia juniper Quaking aspen tridentata) (Juniperus (Populus (Picea spp.) - Douglas tremuloides) least of sagebrush cultivation 74% of the former 1979), rangeland The Bureau fir grouse Mountain, 1968. but of about owned by W. Dickinson. on. Sagebrush ranges aspen cover or types with herbicide, has occurred in Moffat County to on at (Hoffman 20% (9,583 ha) of the big (Hoffman 1979). 1,166 ha on Cold Spring discontinued all spraying 2.6 km2 now occurs the principal most of the (Hoffman 1979). by spraying sprayed shrub. men ziesii) grazing at least pine woodland occurs primarily range altered cover mountain or to improve of Land Management spraying respectively while non-sagebrush (Pseudotsuga rangelands, has been Mountain in 1966 and Herbicide sage On Cold Spring sagebrush with July an d and mountainsides. 40% (4,926 km+) of the total land area land under area The mean and pinyon osteosperma) Mountain and in most canyons Alteration bring to mid-May. rangeland 60% (7,358 km ") of Moffat County, comprise in the study months of the year. consistin g of pinyon -juniper , rock outcrop, spruce and November are while mountains and coldest of 1978). edulis)-Utah area. 1978). to April. the warmest Big sagebrush study October, 1978). annually, is 4. 4 C on Cold Sprin g Mountain, Dep , Inter. (Pinus March, Dep . Inter. tions ann ual temperature 46-51 ern of precipitation August; (U .S. (U .S. rancher yearly in 1971. on land on the study area. ,, • �45 LITERA TURE CITED Braun, C. E., M. F. Baker, Schroeder. alteration 1976. r t Hoffman, Conservation of sagebrush Wilson Bull. D. M. of sagebrush Pp. U . S. Department . Colorado Bur. J. S. Gashwiler , and M. H. committee report communities on effects on the associated of avifauna. 88: 165-171. 1979. Investigations and sage grouse Rep •• Colo. Div. Wildl., Job 10. R. L. Eng, of the distribution in the Moffat County Fed. Aid Proj. \\1-37-R, and status area. Final Work Plan 3, 95-162. of Interior. coal regional Land Manage. 1978. A supplement environmental DES 76-21. statement. 555pp. to the northwest U.S. Dep , In ter . , �46 NATAL DISPERSAL AND LEK FIDELITY OF SAGE GROUSE Dispersal Krebs has a major role in population et aL 1976) and distribution also have a role in the evolution 1978), mating systems (Taylor regulation and Taylor of song dialects (Greenwood (Lidicker 1977). 1962. It may (Baker and Mewaldt 1982). and the stability of local 1 populations (Reddingius importance, studies investigated fall movements. ing areas. and den Boer 1970). of avian dispersal Dispersal greater species of species with lek mating systems and to a limited extent. might show exceptions female dispersal (Centrocercus in birds urophasianus developed (attendance movements between leks) Colorado. the hypothesis typical dieted I tested avian pattern that yearling philopatry Iek fidelity. and that. because dominance hierarchies and Iek fidelity that of Sage grouse to study the effect among lekking describes of sage grouse species natal dis- than males. and in northwestern female sage grouse farther (birds follow the I also pre- 8-10 months old) would show natal among males. yearlings because pattern within and between years. of dispersing presumably even though they have one of the This paper of a population grouse 1980). ) are an ideal species (Wiley 1978. Stiles and Wolf 1979). persal has has been to the general (Greenwood of a Iek mating system on dispersal most highly are few; most research not movement from natal to initial breed- examined in only a few birds polygamous Despite its potential would have a low rate of they are first-time breeders. 1 �47 Study Area and Methods The study was conducted Moffat County. July Colorado and adjacent 1981 through (Artemisia stands. and Douglas-fir t t area. pine area is semi-arid quaking {Pinus edulis)-Utah and meadows. Lodgepole menziesii) 1978. July Cold Spring occur Mountain of which Sage grouse juniper pine sagebrush aspen above (Po ulus (Juniperus (Pinus contorta) 2.620 m on Middle and August studies with numbered colored plastic grouse Drive traps. with long-handled (Giesen et al . 1982). Captured wing molt and primary length through 1983, searches May 1982 and of leks with spotting and previous were recorded nile banding ing (attended) scopes and unique Dep , on the area have been banded nets birds were used 1975). used as an approximation cannon individually Observations dispersal were calculated. of a juvenile's natal grouse to sex and age by During late March 5 days /week banded during of marked birds distances of net. to capture were made at least and the lek on which the bird as a yearling combinations were classified for birds summers. and individually- a bumper-mounted (Beck et al. and straight-line location grouse were captured aluminum bands bands. and spot-lighting sage 46-51 cm (U .S. have been conducted edge each year. 1982. juvenile marked (2,622 m) receives 11%falls in August From 120 to >300 juvenile In 1981 and spring of Wyoming and Utah from with interspersed annually, 1978). during The study (Pseudotsuga of precipitation since parts Mountain in northwestern (2.904 in) and Diamond Peak (2.909 m) on the northern of the study Inter. 1983. and pinyon osteosperma) Mountain June sppv ) rangeland tremuloides) on Cold Spring between on leks the juve-: was observed Banding that locations displaywere area since most were �48 marked before long-distance 1971, unpubl. data this 1 lek, the Iek attended calculating dispersal Lek attendance was observed depending 100. was calculated breeding adult the number was estimated by dividing of birds rates per hour per male" refers entering capture from the grouse and banded to banding = banded males (6) Lek. attendance 58%. Total recruit-· the breeding of yearlings from the number population. seen on leks each the previous by subtracting of banded summer. hunter juveniles. to each Iek were expressed for differing by the maximum number recruitment females. the total days that for known mortality of yearling per hour to correct divided express rates 5 adult 6 days of lek ob servations Therefore, of juveniles and known predation Recruitment then was adjusted on the lek captured male #9947 was captured the number by the total number Recruitment grouse to the bird's #9947 was 18 days seen on the lek .;-31 as the number harvest males. (31) in the denominator. men t , defined spring for birds (37) minus the days prior of days males or females. were observed (26 adult 1983 after which males were seen. were seen on Gee Flats of days that a bird the number of days of the respective prior For example. bird. was corrected on Gee Flats Lek on 11 April for bird as the number season days of lek attendance equaled more than was used in by the total days that males) by subtracting denominator. which attended distances. Attendance each spring during For birds most or where mating occurred on a lek divided by 5 yearling sex's study). on the sex of the particular multtp'Iied during (>2 km) movements took place (Wallestad observation times at leks and of males observed on a per male basis. to this relative on leks to References recruitment as rate. to "grouse It was �49 not possible to standardize data reflect differences interpreted cautiously. the 0.1 probability natal or non -natal in observation Statistical lek attendance so those times among leks and should be tests were considered significant at level. Results Natal Dispersal. --There from natal females years to breeding (Kruskal areas were no differences between were combined. The rate was similar since the sex ratio was nearly equal each spring consequently of resighting of males and females (males Ifemales) on leks of marked males or data from both of juveniles (0.94 in 1981 and 0.87 in 1982; X2 and almost equal numbers distances 1982 and 1983 for either ~ > 0.1); - Wallis test, in dispersal individuals = at banding 2.76, !: = 0.09) of each sex were seen (3 females and 2 males in 1982, 10 females and 10 males in 1983). Dispersal distances (Mann-Whitney (N = U test, 25; 12 males. also did not differ the closest lek). of yearling P = 0.26), 13 females). in tendency males and females did not differ although Male and female yearling (Fisher's lek to the location of their Sixty ment attended percent (15/25) to their times on leks were not equal because of new leks late in the breeding Determination tions. First. were assumed other leks. nests exact banding of all yearling the lek closest the sample size was small P >0.1) as juveniles grouse natal area. to attend (natal area with known recruithowever. of a lack of access observation and discovery season. of natal area lek attendance where individually-marked to be closer test, sage grouse depended juveniles on 2 assumpwere produced to the lek where the female mated than to This assumption was deemed valid because only 1 of 8 �50 radio-marked violated it. female grouse 5.4 ± 1. 2 krn (~± Nests averaged the female was observed. 12.4 ± 1.4 km (~± were captured SE) apart. sites to be correct juvenile 12 September 1982. remained prior grouse natal area. 18 radio-marked to 5 September sites). each and because <2.1 km from its nest Only 5 of 157 juveniles sites minimum of where juvenile because <2.2 km from capture radio-marked a to be in that bird's small home ranges averaged and nest SE) from the Iek where leks averaged Second, was thought had relatively fall (grouse while active and marked were assumed This assumption juveniles with known lek attendance 1 site until were banded after 11 September. Recruitment. --Eight known leks were on the study 'on 2 previously-used were not observed area. Grouse leks in 1982 or 1983. Observa- tions were made on 2 active leks in 1982 and 6 active leks in 1983 (2 leks were found late in May 1983 and only observed (Table 2.1). Direct 17% (20/116) for 1982 and 1983, respectively, individually-marked recruitment juveniles) and 1983 did not differ years. ~ age-class than (!_ rates test without (yearling or adult), was no difference ance rates ficantly Lek attendance data for 1982 data from birds seen both for females. less attendance Recruitment Within each females had a lower attendance between difference yearling test [LSD], (8%) and adult but among males, yearlings than adults and and 16% (25/157 > 0.1) so the data were combined (Table 2.2). males (ANOVA. least significant There were 12 (5/41) of yearlings pooled. on 2 days) rate ~ < O. 1). (11%) at ten d-: (33%) showed signi- (43%) (ANOVA, LSD, ~<0.1). among leks was examined with data from 1983 because in 1982 all 5 yearling grouse with known recruitment attended Gee Flats �Table Sage 2.1 grouse lek observations. Gee Flats Cold Spring Mountain. Moffat County, Beaver Sugar- Basin a loaf Colorado, Whiskey a Draw March-May 1982-83. Swede Cold Flatsa Spring a 1982 Number of males 13(7) ~ (SD) 30(13 Apr) Max !'::!.(date) Total Number days b 26(9) 39(26 Apr) 27 23 23(23) 20(22) of females ~ (SD) Max!! (date) Total days Observation time (hrs) 66(08 Apr) 82(21 Apr) 21 20 27.2 21.1 0.5 1983 Number of males ~ (SD) Max!'::!. (date) Total Number days 22(15) 42(23 Apr) 37 30(8) 39(07 May) 9 15(8) 26(10 Apr) 31 12(7) 20(20 May) 16(7) 21(25 4(1) May) 5(25 May) 13 2 2 1(2) 3(1) 0 of females ~ (SD) Max ~ (date) 9( 11) 42(23 Apr) 7( 5) 14(07 May) 10{lS} 88{10 Apr) 8(28 Apr) 3(25 May) 0 V1 �\J1 N Table 2.1 Cont'd. Total days Observation time (hrs) aSugarloaf, Two other Gee Beaver Sugar- Whiskey Swede Cold Flats Basin loat' Draw a Flatsa Spring 35 8 26 4 2 0 29.4 5.7 18.9 3.7 1.3 0.75 Whiskey Draw. leks had no birds bOnly includes days and Swede Flats leks were found in. 1983. in 1982 or 1983. when observers were present at leks. Cold Spring Lek had no birds in 1982. a �53 Table 2.2 tain, Sage grouse Moffat County, lek attendance Colorado. (%, ~ days), March-May Cold Spring Moun- 1982-83. Age (yrs)a <1 2 3 x - 28 46 51 SD 28 31 2-50 3-74 19b 6 10 6 6 3 Sex and parameter All adults 4 Male attendance Range N 63 40 42 26 16-96 1 3-96 3 52 6 7 Female attendance x SD Range N aUnknown included under bSeven observed 12 13 11 age birds were excluded 3-25 1 30 from the age columns, but all adults. yearlings 1983 are included later 3-25 5 which were initially with the as yearlings. 12 males initially banded banded in spring 1982 or as juveniles and �54 Lek , In 1983. most yearling grouse Lek (0.14 grouse/hour/male) approximately leks trend Basin to Sugarloaf 2.1). between These yearling recruitment the related in the next case) mum numbers Jackson were analyzed hypothesis Colorado during to determine the peaks I examined that of males. spring in both (6 males, 1982 and ments were known to have been individually-identified Interlek between ent pairs grouse movement rates (Table unpubl , data) to leks methods in North intensive counts periods Using these 2.3). ( ~8 including data. maxi- with maxi- ~ > 0.5) • were seen on leks Of these, 1 male (17%) and between years. No interlek move- made in 1982. while 14 of 91 (15%) were seen at more than generally leks when movement rates of leks Maxi··' 4 females) 1983. leks attended (females. Park. females per male was not correlated grouse I female (25%) changed tance leks in North these grouse of males. were made at 21 leks during of attendin g males (!:.::: -0.16. 1983. numbers used in this study: mum number during sage more birds Research of attending males Because 1974-79 (C. E. Braun. mum number sage other relatively of male and female lek attendance. Ten adult of attending if more females per male recruit maximum numbers in any year) did not follow any size of a Iek . leks with larger Park were similar to those countsllek and relative of is no relationship of males and females attending County. with lar-ger there statistically. attended This sequence or maximum number that Basin and Gee Flats leks had and Gee Flats) data suggest data were too few to test to test at Beaver (0.04 grouse/hour/male). among leks in average (Table data while Sugarloaf equal rates (Beaver were recruited Yearling decreased 1 lek in spring with increasing were compared grouse between dis- differ- made more interlek �Table 2.3 Interlek movements of sage grouse, Cold Spring Mountain, Interlek Gee Flats Sugarloaf Interlek ments to Sugarloaf to Gee Flats Whiskey Draw to Beaver Basin Moffat County, Colorado, March-May 1983. movement Gee Flats Beaver to Basin Sugarloaf Whiskey to Draw Beaver Basin to Gee Flats move(~) 11 7 1 1 35.1 35.1 1 1 Sum of obser vation time on both leks Interlek (hrs) 48.3 48.3 22.6 9.4 move- ments/hour of observation time (in decreasing order) Distance 0.23 0.15 7.5 7.5 0.11 0.09 0.04 0.03 be- tween leks (km) 10.9 13.1 12.2 13.1 V1 V1 �56 (20%, 5/20 movements than adults Among sex and age-classes average number 13%, 9/71 for yearlings; of grouse. there of in ter lek movements for adults). were no differences (ANOVA. P> 0.1), in however the sample size was small (N ::; 25). Discussion In birds, females generally between natal areas 1980). Explanations and initial breeding leads to greater amous species 1980). generally (resource leads to greater ary maintenance of sex (Shields (Dobson I could not refute grouse do not disperse farther tendency led me to refrain from accepting biased of all yearling sage grouse adequate by differences grouse attended will require unfortunately, larger dispersal sample sizes by Greenwood have been based and mate defense male dispersal) males. however, the evolutionfor mates that female sage the small sample farther than males or rejecting the null hypothesis. in observation time among leks. natal area lek suggesting Both the dispersal require long-term 60% that and philopatry sample sizes for adequate studies in poly g'- (Greenwood and subsequently the null hypothesis their on: in monogamous species for females to disperse may be philopatric. hypotheses (reviewed 1982), and (3) competition than size and a slight Although than males defense female dispersal (2) the promotion of inbreeding 1982). sites distances for the sex bias in dispersal (1) the type of mating system generally move greater testing; research to acquire (see Kepple 1979. Jamieson and Zwickel 1983. Zwickel 1983). After breeding natal dispersal. area in successive have discussed birds years the establishment are generally (Greenwood of and fidelity faithful to the initial 1980). Few investigators to lek territories by �57 sage grouse. Studies by Jenni and Hartzler examined the effect of variable lek attendance censusing 1952). technique most females breed even though performed their first year than yearling by adult than yearling than adult at or near males. success the mating center and Braun of these 1978). them toward the mating center reproductive fitness as opportunities then retain peripheral success territories expensive territories arise territories for territory on that lek until reaching territories 1978). although may be greatest leks to find one suitable of yearling this hypothesis. (Beck and move may have greater than males which are non-territorial male reproductive visit several defend Iek lek attendance which occupy (Davies that in their percentage may be energetically leks and only establish Average greater of male grouse Males which regularly suggest investment movements support may be high mate. to the same Iek and In this study. territories males rarely Wiley's findings reproductive males making irrter-lek maintenance o. that Almost all matings are yearly males and the greater The reproductive several mature. within the lek (Wiley 1978) territori.es Wiley (1978) suggested males which return males may have a greater on a commonly used lek while yearling they are physiologically by adult territory adult (Patterson (1978) and Emmons (1980) or visit (Davies 1978). when yearlings establishment and the mating center as an adult. The only other reported rates radio-marked ence in results Iek atteridarice study of male lek attendance of 85%for 13 radio-marked adults; these between rates studies by triangulation (Emmons 1980) yearlings did not differ and 92%for 17 statistically. The differ- may be due to Emmons' determination of radio signals, which allowed him to of �58 identify ever, grouse grouse have been ences in sagebrush at the periphery recorded between near hidden as attending. st udies After within Yearling other of Gee Flats to investigate several they that yearling 14 of 27 (52%) yearling in North male yearlings made more interlek yearlings Park. females and the greater they movements need females several than and may need Other move-- 1976) males made in ter-: adult spend behavioral Because between of yearling 2 leks suggests leks prior (Wiley 1978). ait tiu g displaying. W-37--R-29, than in Iek attendance may be greater to receive ling and adult seen and T. D. I Beck Emmons (1980) movements percentage (7%. 2/27) attending to examine on a reported that males (all 13 2 or more leks). time on 2 leks as adults interlek presence males make most interlek Aid. Proj. Colorado. visited The lack of difference total differ- were not a territory. and 7 of 48 (15%) adult lek movements females they at breeding C. E. Braun r ep , , Colo. Div .. Wildl. Fed. radio-marked physical were not seen establishing (Dalke et al , 1960, 1963). found the large were regularly are inexperienced leks before have also Indicated (unpubl. may not held a territory Lek but they how- males may be moving among leks more frequently because ments a bird 1983. grouse the Iek because grouse studies cover However, mean that mid-May in 1982 and as attending In my study. This may explain in Iek attendance. 200 m of the periphery counted leks. of a lek and in heavy a lek does not necessarily Iek , near (20%. 2110) that on 1 lek , than that yearling yearlings Yearling of adults to mating because and adult than adult spend as much attendance and if yearlings need of inexperience or if cues for mating by wa.tching older there were no differences females in lek at ten darice in this study. between or a study yearwith �59 radio-marked appear females (Petersen to lead to a lengthening percentage of yearling ing in interlek be examining observed (20%, 1980), of lek attendance 2/10) vs . adult movements suggests prior cues does not by yearlings, females (7%. 2/27) that yearling more leks than adults 2 of 14 (14%) yearling need for behavioral The engag- females may act ual ly to mating. Petersen and 1 of 11 (9%) adult (1980) females at ten d-: ing 2 leks. One hypothesis for the initiation that clumping is a means of increasing be heard or the amount of time during duced (Model 2, Bradbury study refute from Cold Spring on a relative basis. of males. because there over which males may which display it predicts on a per male basis observations numbers the range and Gibson 1983: HI). that hypothesis females must increase of male clumping on leks suggests signals The results that numbers with increasing are proof this of lek size. Mountain and North Park indicated. were not more females on leks with larger Lek that �60 LITERATURE Baker, M. C •• and ~uttaPi). Beck, T. L. Evolution D. 1., grouse. and Game Inf . Leafl, Pages 109--138 ~_ Press. New York, P. D •• D. B. Nat. of sage Resour. 2111: 4' 1978. C. ~'. :F ·:·aun. (r"v···".:i). Weighb, :? 19,(S. Pages B a teSO:1, ~Y(; -:.',L~r.rrJi.ni;.- 1983 .. T_,ek ..~ and matt' c noroc . (:.d. Pv rab., D. 1960. :" .::tcmtcn .T. y Seasonal r;. movements g rout.e in ...Iabo , Trans. Crawford, arid and breeding North Am. Wilen. s.nd ·?'5::)91~·'~:J7. 19f 3. Ecology, oi sage grou.se in Idaho. and management J. Wildl. 27: 81.1~84L 317-350 in J. R. Kr ts F. S. F N.Y. ecology. Dobson, Color-ado Se: " an d age , and Manage. 0.: 4pp. H. M. Ci'c;-;on. ConL productivity, .. in mamma! and 0 N. B. 0 .. Of f'J 1982. male dispersal to 32:712·722. E. F. Sc hla'tter-er . behavior -r,:' 80: 241··243. J. W., and Bradbury, r·. , C. E .. r:r.;.:\,',. Condor , R. B. Gill, Dalke. rl. CITED An im. . ,1 ,..•. , Da Ties, ed.:>. _ 0 Behavioral K. rr·+'1: :.chav. 30; 118.3- .11.;2. �61 Emmons, S. R. Park. 1980. L '·k ;:".rtl:,t , :,r :'e 0:: r{F,.1e ,:;;:).g'~ grouse Colorado. M. S. Thesis, Colo. North in [)1:<~teUniv , , Fort Collin s . 69pp. Giesen, K. M., T . .1. Schoen her'g for tr-apping sag,:: grouse and .; '01. 1 C" C ·~d,). Br".un. Wildl. ]982. S···. Methods i~t.'H. 10:224" 231. Greenwood, P. J. 1980., birds Jamieson, and Jenni, mammals. 1. G., in blue and Krebs, Ariim . BeLe< v , Can . J. grouse C. J., . R. Hilbor-n , J, 1976. the r egula'tion Nat. 1°18. .• Ma"lO.f,C. Lefl1'(, W;leI. Mana~,,~. 2':; ':'1. fP',)US6 '1~:4b·'Si~. -:13:n.';-"i(:7. J. A. Rodficld , M. Tc.i !..<?~,vn.s:~:r:..:}L. of popula tion an d sitE fidelity At t endartc e at c. s::.ge C'.cl·'·.l~.:'~'>, Fm:i.r),:~'·'";:i.O/l Dispersal biolog:'; mec and disp':!r;;al in fluct u- Cm . .J'. Zool , porsaib Ie t, 54:79-95. hanxsm pe'rrnittin g :'(:n;',ity' b elow c-ar-rvin g capaclty. Am. 96:29-~? Denver, grousf' .' ort Wi1dl. and disper-sal in 28: Jl4!_)' 1.l62. M;.~:r:oh.•.. populatkm atin g popu le.tions of M. 1962. '.r. prL, ... 1. Winga,;.::;, J, Lidicke r , W. Z. phtlopa.try 61:570"57:). ZfX.l .• J. F. Hart" imp lica tion s for' of sp ruce sv-vtemo , and F .. C. Zv.:,~kf:,1,. 19"j. grouse. D. A., lek: Ma':irlf_ Colo. 'HIpp. in North Collir r:. • Pa.rk , Col:··;"do. ;l( . ~ . M"S. 'I'besrs , Colo. State Urriv •• �62 Reddingius. J .• and P. J. illustrating stabilization Oecologia 5: 240-_~8.1. Shields. Stiles, Stat ~ Urriv, F. G., mating i 27: 0 L. R., Bur. Wallestad. Am. animal numbers 0(- New Y k -j by sfn:eading - R. O. . i, Y r , of r-isk. r,:-:-mit 2·1 p. ':.' 1. .-.·on __ ~(:: h l'-)mi-,l::"b:irci., Oi-nithol, 97 t . A~' gr :,a m~ g_:';C\hon jo::._, mct:inn:c;" reg. Man se- ] )11. 'J. 1 .l_p 9. of Int. r or coal Land C in the Lmg-·t.:<.lE;d dud L. A. Department Color-ado Sivnulation cxper iment 78pp. and population U S. 19'10. n d L. 1.. W behavior Monogr. Taylor. (!~. 1982. W. G. sex. b dt;.'D n r ,"_nen+ , .tat mei me t n .hwe ..': :p. In··er .• 1 .• DES 76 .:.J. Surmncr rno verr.er 1::5 a:.. 1 bi-,-,t •..• : '__ L _, ~')y ~;;J ge 238: 114-125. Zwick el , F. C. to breeding 1983. range. Factors affec tin g thE: r etur'n Car .. J. 2':.001. of young 6J: 1:'28--1132. blue r;:;rouse �63 LATE Sl!MMER--SPRING JUVENILE Spacin g b eh.avior-s movement rates which greater A.lthough rai;:,l ranges sage areas dynamics habitats. This paper ments of juvenile of f;\.clI,mer t he hn;ed'ir' ~~poptrla.tion movemen t s 0 (_ in ._1<1....-0. and fan (Watson an c b::o:.n cxamin ed ic'r and t::~)!:i~.?._~?J!_l~_~~.3!:~~:j~~~::;:_:.:;) (Kep pie 1979). :\l~i.VC: not may be impor-tant population ~3:i~'-? {reviewed Zwick.ei 1983), movements (q"~!2:~!f!._C(C~E::::::::!!_:,_.t:~?:()l?.l~l;:':~~~!3~~:-!g) from E'lunmer grouse to breeding arid P::d grou s« {!,::,:.ggp'~l.~.~~:.g~?P~':~ (I~.• o~~c~!..·~~.~~) C;a.mieson and ments of juveniles r.H;,::U [,tudi.j~d" Knowled ge of for understanding sage and for rnitiga.r~~.1ga,"J·'.rer3e hurns n impacts deac.r-ib es the sage late summer to early rnove-: grouse on spring mov e-: grou se in nor-thwes terrr Color'ado , Area and Methods The study Moffat County, 1981 through land n. death, (XL~::'P<lr!~lS:~:..~~C:; cl_):_~:d? E~;~)!:l:~~.t:=t:~J (Bowman sp:cuce grolls grouse of juvenile Study into p ra'ir-ie+c hi ck ens and blue ll' ex :..nple of the impo'-tance to r-ecr-uitment Robel 1977). SAG.!:.. GROUSE :sub:>,-:ql1'~nt1y lin it population scoti.£~~_) are a classic Moss 1980). OF may ~)1'oduc.:! cha ..n;f~F_::s in birth, Watson and Moss J 970, i-. rebs movements MOVEMENTS with wa.s con due. ec on Cold S;n-ing Colorado and in adjac ent Wyoming .June 1983. hrter-sper-sec' Mountain L ...1'11' .. dudy :c,ea :>c' and Ut ..·h from July A',~ .:ri···:'.:4!.ct ·,1d.kin,_ . P =n '~:P!~~!~l~' rn northwestern sc:.g,"bl twh r'an ge-: i:_i_er::~~~()_l<i~::J and pinyon �64 menziesii) occur above Diamond Peak (2.909 Spring Mountain of which m) on the northern (2,622 m) r eceives 11% falls in August >300 juvenile July 2,620 m on Middl e Mountain sage grow:;e were banded each year In 1981 and 1982, juvenile marked with numbered colored plastic spotlights et al. mitters Drive on juvenile October tl'2,pS, length (heck 1982. 'I'hrrteen to cap·.u:ce grouce ond August Forry-xwo 19f ed radio were used in 1982. b ir ds were ber 1982. Relocations to 15 November marked juveniles relocations mid-day ing each year. During were relocated (~4 hours after before cide with most feeding during mid-day, marked grouse during January time periods. morning ground. were us eci in 1 August 3 times to 11 Sept em- 11 :.1eptember to mid Apr-il 1983, 4 radio- (~4 hour" before WeekJy radio after sunset), sur rise) , ana e veri-: T'heae time periods and evenings. A 3-·elem(;nt yagi ant en ria was used on the ·~io jr,'d1S·- r-adio tr-arrsrrdtter-s 1 tirne Zmont.h, to >4 hours and 1980) were placed monrh l y after into morning sunvIse sunset) least at least were evenly divided (>4 hours 3.1: net, (Gi.~!3erl r eloc at.ed a.t least 1981 and from wert'; made 1'0. transmitters o eolar+power-ed from 15 July to 31 August of an d JUly through 23 ba tter-y+power ed and Radio- marked during 1:")~'ey. and ;:~!e by wing 1981. and weekly comb ine.tions m;:d:·ke:rt.: (Arns t r up solar+power Mountain b urrmer+rnour, t ed cannon et al . J9'i'~». duz-in g July annually, weee captur ed ~.nd indivldu3.11y-· wer-e uaed to poncho -type grouse 3. Cold ~1_98:~, b ir ds wer e CL3.ss!f:ieci Captur-ed attached grouse nets area. From 120 to on Cold Spring train J 978 through and lon g+hanclled molt and primary 1978). alumin um b an ds and unique bands. 1982)., edge of tbe study Dep , Inter. and n) 46··51 em of precipitation (U .S. and August (2.904 coirr- and roosting to relocate An airr::l··,,:"t with 2 strut-r;,ou.lted radioyagi �65 antennas 1983. was used to locate birds on 3 November Radiolocat.ion s made on. the ground lation at close range locations were plotted maps using distance on other from their were analyzed Test results of variance capture date, with non+paramctr-ic were considered topographic sites If the interaction was signifi-- was individually or movement rate). statistical significant fi:n:;t wer-e (ANOVA) for interactions site for each bird (age of bird, Bird coordinates. bit-cis and time periods. variables birds, 0 moved by grouse from capture by txian gu-: U S. Geological Survey Mercator among individual cant. were determined Transverse with 2-way analysis 15 January 200 m) or by flushing on 7, 5-minute Universal Distances analyzed (approximately 1981 and tests at the 0.05 regressed Directions (Bat.schelet 1978)" probability level. Results Fifteen and 43 juvenile 1982. respectively. Locations 1982 were used for analysis mittel's. predation. were analyzed sites hunter both years during August effects and time periods, increases in distances increases from capture sites birds 3.1). were r-adiorna rk ed in 1981 and in 1981 and 10 birds of loss of r-adio signals or incomplete in 1981 and steadily (Fig. data" Movements 213 locations in 1982. moved away from their 3.1). Because it was not possible were gradual sites capture of unequal during with date in either was most highly 'I'hz-ee birds to determine occurred, correJated sample individ- when However , the data both years. year in or trans-- (2--way ANOVA, P ::: 0,01) between from capture did not vary (Table harvest. grouse ual birds (km/day) b ecause to November sizes and interaction that grouse of only 8 birds from 118 locations During indicate sage (Table Movement rates 3.1). with date for showed no relationship Distance 12 of 18 between distance �o - I ~ ~ ...",... U) W 8 iii n 5 1981 0' (1'\ ~982 ~ ~ 5 [I- m w iLl: :::> 4 I 3 r t t:-c 0 ~ 5 15 I 0 a: H w o Z 27 2 ~ ~55 (5 18 -a 1~ 9 14 '-8 11-24 25 JUL7 AUG JUL 3. L Distance 10 grouse is the mean. number L II 43 I{/) and 6 ~23 ~ Fig. n (km) moved in 1982 on Cold The bar of radiolocations. from capture Spring is 1 SE above Data are 5·1 a SEP 22 AUG, 4 SEP 821 AUG sites Mountain, and from 1 G SCP2 OCT Number in 1981 Hori scr.t al line above (Tab~e 17 OCT· ~4 NOV grouse Mof'fa t Coun t y , Colorado. 12 males an d 6 females ..•.. --- OCT by 8 r-ad io-rnar k,..d sage below the mean. ---_ 3·16 the Lar 1S the 3.1). �Table 3.1 R2 and probability (km) with movement 14 November rate 1981 and (~)a of simple and from previous 1 August locations - 14 November multiple (m/day), regression date. and age of bird grouse distances on Cold Spring from capture Mountain. from capture Movement 15 July sites, Best multiple modelb Variables in Bird Sex Iocations 1981 438 M 14 0.01 (>0.5) n.22 (0.25) 0.22 (0.25) None None 465 F 10 0,01 (>0.5) 0.77 ([L 00) 0,77 (O.OO) 0.77 (0.00) Date or age 486 M 16 0.01 (>0.5) 0.76 (0.00) 0.64 (O.OO) (),76 (0.00) Date 501 M 18 0.11 ( 0.30) 0.38 (0.01) 0,,38 ({LOl) 0.38 (0. Gl) Date 585 F 13 0.00 (>O. 5) 0.66 (O.OO) 0.66 (0.00) 0,66 (0.00) Date 687 M 11 0.00 (>O.5) 0.56 (0.01) 0.56 (0.01) 0.56 (0.01) Date 712 F 18 (),ll ( 0.75 (0,00) 0.75 (D.OO) 0.75 (0.00) Date 785 M 18 0.00 0,57 (0.00) 0.57 (a.OO) 0.57 (0,00) Date 0.44 (0.00) Date 0.60 (0.00) Age All birds 1982 rate (>0.5) - ~2 (~) Year 0.4) sites 1982. Distance N models of sage Date Age 118 multiple model 520 M 28 0.00 (>0,5) 0.59 (0.00) 0.60 562 F 18 0.48 ( 0.00) 0.10 (0.20) 0.08 [D.25} 0.69 (0.00) Movement 576 F 27 0.08 ( 0.15) 0.34 (0.00) 0.34 «(LOO) 0.34 (O.OO) Date or age 669 M 15 0.27 ( 0.04) 0.37 (0.02) 0.45 (G. 00) 0.78 (0.00) Age, 694 M 21 0.06 ( 0.27) 0.00 (>O.5) 0.00 (>O. 5) None 702 M 22 0.14 ( 0.09) 0.74 (0.00) 0.73 (0.00) 0.74 (D.OO) rate ~ age movement rate None (0. 00) Date 0" -.....J �0" co Table 3.1. Cont'd. Distance N Year Movement R2 (P) sites. Best multiple modelb Variables in Bird Sex 712 F 17 0.02 (>0.5) 0.67 (0.00) 0.67 (0.00) 0.67 (0.00) Date or age '734 M 15 0.04 ( 1),47) 0.56 (0.00) 0.56 (0.00) 0,56 (0.00) Date or age 757 M 28 0.02 (>().5) 0.56 (0.00) 0.56 0.56 Date or age 797 M 22 0,02 (>O.S) 0.10 0.09 (0.16) All birds "Probability bStepwise locations from capture rate Date (0.16) Age (0.00) 213 of no relationship mult.iple regression. between distance from grouse capture sites and (0.00) multiple None None 0.59 (G.OO} Date and rate the independent variable. model movement �69 from capture sites and date, age of bird, or movement rate. 15 July to 4 September 1981 and 1 August and 69% of all locations (N capture sites. while during 77 and 78%. respectively. from capture move farther females. 3 October than moved while testing from their the assumption year (Partial capture 1 August. ~ test, sites Because of parallel sex each year. August test, ~ <0.00) when individual coefficient. restriction that variation 1978: 6) N should Also. there lines could not In 1981, females were sites than without moved farther paral- than the other action was to accept moved between 10 August. females on any ANCOVA is inappropriate in distances 3 (~== (Fig. 18) birds the nun males and in 1981 and 1982 had a non- mean direction. 3,2). There means of all birds P > 0.05). of N has little e males and regression from capture or significant correlation between of date to November. Movements of all but random orientation, if females than males on any date after the most parsimonious of no difference females during direction B. 36) were >2 km of the effect ~<0.05). lel lines and males and females alternately (Rayleigh's 1981 and 1982, to determine it allows control for a difference while in 1982 males were farther hypothesis = (N (ANeOV A) was used males because be made for either date after were ~ 2 km from to 14 November. of all radiolocations of covariance However, farther 135, 106), respectively, 1982. 82 sites. Analysis on distance ::= to 4 September During was no significant were combined I recognize that be equal for all birds, eff ect on the statistical VIas no ob vious from capture difference mean (Mardia's I violated however. sites the test a slight analysis (Batschelet between directions of �70 -, 4 KM s Fig. 3.2 ture Mean angles sites Cold Spring November females and mean distances of radio-marked Mountain. 1982. (N = 6). juvenile Colorado. Solid lines are (km) moved from cap- aag e grouse July-November (!:,:!, =: 18) on 1981 and August- males (~ ::: 12); dashed lines are �71 movements differ of males and females in mean direction test, 3.2). moved in either Males and females did not year (Mardia-Watson-Wheeler P >0.05). Except for on 21 January 1 male grouse (#501) found 1982, the last located in exposed seen after sagebrush. northeast following a snowstorm adult hen was located 1982 and January which covered until with known locations 1982. The number in 1982 decreased Mountain increased 14 December, snow depth summer range was covered. lost prior to 3 October. do not know if they whether there were other 4 of the 9 birds November birds earlier to mid-January. grouse 1 female) snow levels By grouse other grouse were relocated. grouse signals. so I or However, were lost. radiomarked 1983 after Maximum distance to 7 on on sage was later than area to leave summer for loss of radio and February to November period 9 of 58 radio-marked None of these (3 males. in January this appeared had moved ~3 km when they Four juveniles were located During Some birds reasons A juvenile of radio-marked from 0-5 to 20-50 cm. were dispersing 5 km wintering was ~1 m and all sagebrush in late October-November; 14 November. from 17 on 30 October and to 0 on 25 November. on Cold Spring from August were all exposed area 1982. to May 1983 moved to its subsequent 23 to 30 October 14 November no grouse on its wintering 3.3) which was radiotracked marked g rouse were 1982, but were radiotracked Mountain grouse 35 other of Cold Sprin g Mountain on 15 December male (#734, Fig. during on 21 January In fall 1982, 7 grouse One radio-marked of juvenile Male #501 and sagebrush 2 February on Cold Spring 2 radiolocations in 1981 were on 14 November. range (Fig. in summer being 1982 lost during from capture sites late- to their �WYOMING WHISKEY COLORADO -....J •• DRAW N Lb. MIDDLE W MTN ~ :r: OlAMON!) W I?EAK <t I- ::> GEE )45~~ 'r...'1L--~ N o CAPTURE '" LEI< o ~ ~ Fig, 111694 2 3 SiTE 4 5 KM 3.3 1 female. 1982-83. Summer to spring #452) on Cold movements Spring of 4 radio-marked Mountain, Moffat County. juvenile Colorado, sage grouse 9 Augus (3 males; t - 28 May �73 locations in January-April The female grouse while the (Fig. 1983 averaged 18.2 km for these moved a maximum of 30.3 km from her capture 3 males moved from 11.4 to 17.3 krn from their 3.3). Color-banded and radio-marked grouse northeast of Cold Spring Mountain; southwest of Cold Spring Mountain along the Utah-Colorado 3.3). 1 area was a ZOO-km:'.sagebrush A radio-marked between 2 February in the same area, flat 5--30 km (25 km+) was 'I km area male (#694) in the latter moved 23 km to the wintering area northeast boundary wintering of Cold Spring and 26 March 1983 and attended from 4 April to 19 May (Fig. sites were found in 2 areas: the other site, capture wintering (Fig. 4 birds. area Mountain Gee Flats Lek , 3.3). Discussion Late summer movements of juvenile sage grouse Mountain were not characterized by rapid, most members of the population, as Godfrey for ruffed grouse (Bonasa umbellus). and female sage grouse September and October, on Cold Spring rates on Cold Spring November because in Idaho km/day. The lack of difference grouse suggests ments of the sexes spring (Keppie that affect similar to the greater 1979). prairie-chickens during (about Sage grouse are differences recruitment, emigration throughout (Bowman and (Dalke et a1. 1963). Movement September1 day) and did not move> O.3 in movements between if there found movements of male of up to 20 days when birds that (1969) Mountain were sporadic movements> 2 km were quick by periods movements by and Marshall Mountain did not change separated sage synchronized Instead, similar to greater Robel 1977) and sage grouse on Cold Spring male and female between the move- then they probably occur in of female than male spruce seem to follow topographic features grouse and �74 avoid areas sage without grouse movements east orientation (80% slope) forest. sagebrush However, during radio-marked indicated mid-November and lower valleys (1952), leks) grouse movements between areas were related of sagebrush. distance during wintering reported moved north to breeding marked grouse color-banded winter Another adults (Beck study and juveniles generally distances may have of most Field observations (1982) Patterson flats Patterson reported (nesting that sage areas and on the availability varied depending (1952) respectively. averaged 28-30 3 males) (Schoenberg (68 males. on the and Dalke 10 females) In Colorado, km for 7 radio- 1982) and 8--12 km for tra ve1ing from breedin g to 1977). of fall movements of radio-marked were tracked left summer range Markham 1983). range in November. probably above snow. (4 females, ranges Mountain when one -way movements from summer to win te r ran ges range grouse cover as to sagebrush and breeding of up to 160 km in Wyoming and Idaho, winter time. to snow level and its effect cover without since locations that and Schoenberg Movement distances to suitable et ale (1960) to mid-January generally (1960). areas Cold Spring as snow levels increased Dalke et al. grouse large movements to winter were unknown sage with pinyon-juniper was snow covered. long-distance birds that may cross male #694 which crossed Synchronized, to south- Mountain and away from its steep grouse on the mountain of summer along the northwest face which was covered sage did radio-marked occurred since mean angles were generally of Cold Spring southwest all sagebrush cover until in October 30 November) and November From 10 July to 7 September, sage grouse found that (Connelly (both grouse and 95% of 0.11 r-adiolocatiorrs �75 (N = 131) were ~2 km from the general locations = (N findings 22) were >2 km during are consistent of 14 radio-marked summer range until after with results grouse prior lower probability This early to November. study while most birds dispersal These Mountain. and Markham's chances of establishing while 82% of all from Cold Spring in Connelly a yearling's area, October to mid-September, 1 October. tion may increase capture moved from did not leave or post-hatching of recruitment a territory emigra- if there in natal Three is a than in other areas (Watson and Moss 1980). During summer have exhibited Moss (1980). 1982, 1 yearling post-hatching female and at least emigration From hatching during as described 25-27 June until 7 August, were ~2. 5 km from the nest and chicks (45-47 days old) were 5.2 km from the nest were 9.6 km from the nest. chick on 17 August 12 September. 3 October. remained mitter and started site on 1 other 2 km of her nest a large radio-marked female with a brood site until loss of the radio trans- and 1 of her chicks, averaged which was radio-marked. site until sage 3.8 km prior grouse into an area covering adjacent the next The hen was last loce.ted 2.2 km from the nest 12 September. females which were unsuccessful Juvenile site; The female was last seen with a moving back towards not move >2.1 km from the nest from nests the female site on on 8 August radio-marked On 7 August the hen the nest In contrast, within may by Watson and and chicks day they site. 2 chicks Utah and Wyoming. area is important to 20 August produced at least nesters, (range on Cold Spring for proper Among 7 maximum distance 1.3-6.5 km) . Mountain dispersed 874 km2 in northwestern Understanding did Colorado and movements of grouse management of populations in such and �76 seasonal areas habitats. In this case, protection of summer b rood-a-ear-in g would not directly benefit wintering found on summer ranges during February-late important to know if grouse as one population populations. birds, or whether From the results and hunter unpubl. Spring Mountain return data), to breed the study effort throughout grouse in the study within interval we.r c It is Mountain can be rnanaged are recruited that as no grouse April each year. of radiotracking, it appears and trapping Mountain during birds band recoveries Braun, ing pressure on Cold Spring grouse into other ne;c;::cby recaptures of' l~G·~)cJ:,,:. Moffat County (C. E. produced area. on Cold However, h r n':·· 30-50 km of Cold Spring were light or lacking. �77 LITERATURE Amstrup c. S. p 1980. A radio CITED collar J. WildI. Manage. for game birds. 44:214-217. Batachelet , E. Pages 1978. 3-24 in K migration. Second order statistical Schmidt-Koenig 0 navigation analysis and W. T. and homing. of directions. Keeton. Springer-Verlag, eds . Animal Berlin, West Germany. T. D. 1. Beck. selection ---- o 1977. in win tel". of sage T J. e d). and mortality 1975. Sex and age deter- of juvenile Colo. Div , WildI. 4pp. and R. J. Robel. p 4 b 18- 26. from wing characteristics. Leafl , 49 (revis Game InI. 0 J. Wildt. Manage. grouse and habitat flock characteristics R. B. Gill. and C. E. Braun. mination Bowman Sage grouse Brood break-up. prairie chickens. dispersal. mobility. J. Wildl. Manage. 41: 27-34. Connelly, J. W•• and nuclide o. concentrations D. Markham. of sage 1983. grouse Movements in Idaho. and radio- J. Wildl. Manage. 47:169-177. Dalke. P. D., D. B. Schlatter'er sage Conf. grouse , Py rah , D. C. Stanton. 1960, in Idaho. 25:396-407. Seasonal Trans. movements North J. E. Crawford, and breeding and E. F. behavior Am. Wild1- and Nat. of Resour . �78 ----- , productivity, Manage. K Giesen. and management 1963. of sage grouse Ecology, in Idaho. J. Wildl. 27:811-841. M.. T. J. 0 and for trapping Schoenberg. sage grouse and C. E. Braun. in Colorado. 1982. Methods Wildl. Soc. Bull.. 10: 224--· 231. G. A •• and W. H. Marshall. Godfrey, persal Jamieson. of ruffed in blue grouse. Keppie , D. M. 1979. of spruce Krebs. 1978. regulation. Denver. U.S. T. J. in North Park, Collins. 86pp. Department Colorado Bur. Watson. and site fidelity 61:570--573. mortality. and recruitment 43g 717-727. hypothesis of population 56: 2463-2480, The sage grouse in Wyoming. Sage Books, 341pp. 1982. Sage grouse Colorado. of Interior. M.S, 1978. movements and habitat Thesis. DES 76-21. in relation 1970. food resources. ed , selection Univ .• Fort to the northwest statement. U.S. Dep. Irrter . , 555pp. Dominance. to population 167-220 in A. Watson, Colo. State A supplement environmental A •• and R. Moss. to their and dis- 33: 609-620. Dispersal overwinter J. Zoo1. Land Manage. Pages Zoo1. Wildl. Manage, coal regional aggression J. 1983. A review of the Chitty 1952. Colo. Schoenberg, J. Can. R. L. Zwicke1. Dispersal. grouse. C. J. Patterson. Can. Brood break-up J. Wildl. Manage, grouse. 1. G .• and F. C. 1969. spacing limitation behavior. in vertebrates. Animal populations Blackwell Sci. and in relation Publ .• Oxford, U.K. �79 ---- • and 1980. Advances in our understanding population dynamics of red grouse from a recent numbers. Ardea 68: 103-111. fluctuation of the in �80 SUMMER HABITAT USE BY ADULT FEMALE AND JUVENILE SAGE GROUSE Sage grouse throughout North sagebrush America. nesting. (~_~!}_!E~<::_e:z:~u~ ~:t.:'5'":e!:~sianus)are widely (~.~!_~m!~i.:'?:. spp ,) -dominated Their and cover y ear+z-currd has been 1969. Eng and. Schladweiler dependence (Patterson reduction and distribution has been reduced or eliminated by agricultural (Aldrich 1963. Martin sagebrush rangeland based numbers 1970, Wallestad will require on an understanding Except for summer habitat, grouse habitat grouse has been examined 1974). (Klebenow which nested differential factor survival 1978) or brood small (~<30) in sage occurs mortality characteristics 1982). (habitat population b :.:tween sexes has an effect of sage use by sage and Schladweile r 1982), and hens However, these use in propoxtion measurement sample sizes. grouse populations habitats. 19'71, Schoenberg (Schoenberg selection amounts of of sage grouse males (Wallestad are limited by inadequate relatively an important habitat Decreasing development Summer habitat 1969. Oakleaf unsuccessfully have not investigated or have seasonal have been well described. broods availability). 1975). the structural for adult 1974, where sagebrush or energy management of their 1952. Klebenow This dep,~ndence has led to a 1982). grouse for food, and Schladweiler Walles tad et al , 1975, Schoenberg in sage r-an gelan de in wes tarri on sagebrush well documented 1972. Wallestad distributed of habitat Summer habitat dynamics in summer on population studies to structure. may be if wei ght+r-elated (Beck and Braun regulation, as in �81 several avian species . of this paper (e g , Krebs and Perrins is to describe ing female sage grouse Study in northwestern was conducted Moffat County. September use by juvenile and nest- Colorado. stands. 1981-82. The study is area meadows and quaking Lodgepole menziesii) on Cold Spring Colorado and in adjacent with interspersed , summer habitat The objective Area and Methods The study ,i 1978). s occur pine (Pinus above 2,620 juvenile sage and August grouse (U.S. with numbered bands. Drive traps, long-handled Captured length birds markers Cold annually of From 120 to >300 Mountain during July and unique and individually-marked combinations cannon net, grouse of colored plastic and. spotlights (Giesen et al. and 1982). to sex and age by wing molt and primary For ty-two radio transmitters 1981 and July through radio transmitters birds area. attached (Amstr-up 1980) were placed on juvenile and 6 solar-powered Radio-marked (Pseudotsug~ 1982. were captured to capture were classified July and August solar-powered powered were used 1978). on Cold Spring a bumper-mounted (Beck et al , 1975). poncho-type during nets ~remuloid~s) 46-51 em of precipitation Dep , Inter. grouse aluminum bands (Populus rangeland edge of the study from 1978 through In 1981-82. juvenile sagebrush June- on Middle Mountain (2.904 m) and 111 were banded each year semi-arid ~o~_!orta) and Douglas-fir Mountain (2,622 m) receives which 11%falls in August Wyoming and Utah during aspen Diamond Peak (2,909 m) on the northern Spring Mountain in northwestern October 1982. to grouse Thirteen were used in 1981, and 23 battery-- radio transmitters were relocated at least were used in 1982. 3 times weekly from 15 July �82 to 31 August 1981 and from 1 August radiolocations were evenly divided mid-day (>4 hours in g (~4 hours after before sunset) cided with activities antenna sunrise analysis Vegetation board. radio-marked (1968) along north-south tion (hen location for broods) tion meastrremen is included: un foliated sagebrush and other cover from the line transect; bare ground nearest as the center 5%) evenly spaced shrub (weighted the transect [McDonald 1980]); shrub of each side of each transect; measured board; density horizontal Names of plants (1968) cover 0 Transects axes with the bird loca- Vegeta-- and total shrub height canopy and to the of each shrub by its 'width perpendicular and the most common forb species plots. using Daub enmir-e (cover was estimated at 5 and 10 m from the center Daubenmir-e method, of the t'ran sect , species, on both transects; the transect was measured cover of forb s , grasses. plots intercepting used canopy cover of foliated and percent from 5 Daubenmire coin- birds. at each site and east-west percent and ev err-: 3--element yagi and Jones' were measured sunrise). Radiolocatioris grouse (1941) line intercept Two Ifl-rn tr-ansects were oriented by flushing 2 x 5 drn plots, after sunset). .A grouse. of r-adto-rnar'ked a modification of Canfield's Weekly These time periods and roosting. were determined 1982. (~4 hours to >4 hours before such as feeding at locations and Daubenmir'e'a into morning time periods. was used to relocate for habitat to 11 September to (plan ts Zm") within 0.5 m cover at the transect and at 45° using center the cover on a tz-an sec t es timat ed from follow Beetle (1970) and Scott and Wasser (1980), Forty numbers random sites corresponding were located by randomly selectin g 2 5-digit to the last 5 numbers of north and east �83 Universal tain Transverse (250 km2). the resulting Mercator coordinates The distance covering (in paces) point from the nearest Cold Spring and direction Mounof (azimuth) known location on a map was then calculated, Variables chosen possibly based accounting on previous studies for observed of sage grouse 1982) and avian communities in sagebrush berry 1981). Habitat from 192 transects (2 10-m transects use sites were excluded were measured Forty in 1982 were not measured on these were measured at relocations 7 unsucc hens. grouse ssful were excluded Percentages and foliated ground were eliminated because grouse Shrub difference. Statistics which appeared canopy of individual therefore, cover by visual plants estimate) bare by grouse = use sites by differences by this than at in plant 4-8 week for transects with herbicide (BARE), GRASS + FORB) 4-8 weeks earlier separately in areas (~25% un foliated and unsprayed was analyzed juvenile ground random and grouse and to examine habitat data 1 female), of other (FORB), were not biased were calculated Vegetation sample sizes. were biased characteristics grouse measured (4 males. relocations at random sites to have been sprayed tion sample variance in 1981. grouse forbs between locations variables (FOLIATGC; FOLlATGe they were measured and, transects of inadequate from analyses use sites grouse no random transects most habitat (GRASS), covers phenology. shrub because of grass because hens; was analyzed in 1981 at juvenile of 5 juvenile and 2 brood (Wiens and Roten- at sage transects in 1981. and because (Schoenberg sage grouse each) from analysis use were habitat rangeland use by radio-marked and 40 random transects. habitat areas use in these by measuring to parti- areas. height Use �84 (CNTRHT), plants width beside 1982) . (CNTRWD), and intercept which grouse These variables were feeding caused stands birds in aspen regularly August computer (SPSS) ance stands. meadows during analyses programs mornings to Because radiotracking gen- in meadows and 'I tran- were conducted 1975). Habitat (ANOVA) and Duncan's distributions of forb species Discriminant function hen sites) and evenings in July and with univariate Package variables and multivariate for the Social Sciences were individually use sites with analysis multiple range test were compared by chi -s quare analyses use among groups Sage grouse and insects. among random and sage grouse fully nesting were thought 21 transects in the Statiatical (Nie et al. habitat Schoenberg if birds were eliminated from analysis. for feedin g on forbs Statistical pared stands used disturbance. (after to move more while they were in meadows and aspen than in sagebrush sects (CNTRINDT) of or roosting were not measured have moved due to observer erally distance were used (random, because (DMRT). comof vari-- Frequency analysis. to examine differences juvenile, brood many variables hen, in and unsuccess- could be considered simultaneously. Forty habitat variables but 2 considerations First. were available for use in discriminan t analyses. led to the use of a subset some of the variables were highly correlated that they measure similar features variables Second, is necessary of the habitat. for unbiased the goal of the discriminant adequately separated groups of the original interpretations analysis indicating while independence of discriminant of analysis. was to find a model which along discriminant the least complex and most understandable (~> 0.7), 40 variables. subset function (DF) axes with of original variables. �85 To avoid these all variables problems. among random and grouse which did not differ variables then a one-way (~> (~ <0.05) highly correlated variables (after Noon 1981). to eliminate variables resulted among-groups was retained. from the ANOVA, variance Therefore. or ease of only 1 of a pair of was included in the discriminant All tests on If a group of correlated F values with the greatest biological interpretation use sites O. 05) among groups. with significant the variable ANOVA was performed were considered significant analyses at the P <0.05 level. Results Univariate Analysis. --Twenty-one and sage grouse (Table 4.1). use sites for parametric This set was further which significantly differed when ANOVA assumptions than Kruskal-Wallis are not violated. Seventeen geneity or patchiness grouse sites plants beside zontal cover sities variables (CV variables) combined the bird (either aspen. These variable results between structure, cover, differed between the tests. hetero- random and all CNTRWD. CNTRINDT). which had larger HORCOV10)" more homogeneous proximity and homo- and habitat (CNTRHT, and closer test eliminated by this criteria Grouse used sites 9 (DENSITYH). it is a more powerful (Table 4.2). (HORCOV5 variables but not ANOVA; of ANOVA (e. g .• normality describing tests by eliminating Variables and led to different among random or non parametric reduced (Zar 1974: 140). of variances) differed in a Kruskal- Wallis test. may have violated assumptions geneity variables greater shrub to edges of habitat hori- den- types meadow. or pinyon-juniper; TYPEDT) than random sites.· differences among sp rayed , unsprayed. were consistent �00 0" Table 4.1. Variables parametric test (~< O. OS) differing significantly statistics. Cold Spring Mountain. among random Moffat County, and sage Colorado. grouse June-September One-way Mnemonic Variable SAGE Foliated DSAGE Unfoliated TOTALCC Total FOLlATCC Foliated shrub HORCOV5 Visible squares on cover board HORCOV10 Visible squares on cover board MEANWD Mean crown MEANINDT Mean intercept Height width distance of the shrub or shrub Width of the shrub Intercept distance was roosting random or shrub sites. em p p Correlationsa 0.05 0.11 A 0.02 0.00 a at 5 m , % 0.00 0.00 at 10 m , % 0.00 0.00 0.07 0.03 b 0.00 0.01 B 0.00 0,00 0.00 0.00 0.01 0.01 under the line transect. under em the line tran- which the grouse of random which the grouse at center or feeding, Wallis a % % of the shrub ANOVA 0.00 at' center beside Kruskal- 0.00 of shrubs beside 1982. a cm or feeding. CNTRINDT of shrubs or non- 0.04 cover, cover. for parametric 0.06 % cover, canopy % cover, canopy canopy or feeding. CNTRWD canopy sagebrush shrub sect, CNTRHT sagebrush use sites of random beside or shrub was roosting sites. em was roosting sites, em the grouse which at center of �Table 4.1. Cont'd. Variable DENSITYH Coefficient of variation (CV) of shrub 2 10-m transects random SAGEH (pairs) density at each grouse site CV of foliated sagebrush canopy cover within CV of sagebrush crown width within pairs CV of sagebrush intercept distance within CV of forb within BAREH HORCOV5H ground pairs cover board squares among 5 Daubenmire among 5 Daubenmire at 5 m among 3 sides on a transect 0.00 0.00 0.03 0.05 0.17 0.05 O. ot 0.03 0.00 0.09 0.61 0.00 0.01. 0.03 of plots plots a transect CV of visible cover cover Correlationsa of a transect CV of bare within ground p pairs transects FORBH Wallis or transects SAGEIDH ANOVA between of transects SAGEWDH Kruskal- p Mnemonic the One-way of the 00 '-J �00 00 Table 4.1. Cont+d , Mnemonic Variable HRCVIOH CV of visible HRCV45H CV of visible TYPEDT Estimated to interpret; analyses variables were not correlated ANOVA Wallis p P 0.02 0.00 0.19 0.01 0.00 0.00 Correlationsa of the on a transect to edge, with the same letter in the discriminant at 45° among 3 sides minimum distance or distance aVariables on a transect squares cover board Kruskal- at 10 m among 3 sides of the squares cover board One-way to a different with lower-case type, m were highly . because cover they correlated (r- > 0.7). had the largest!:. letters value. were eliminated and were used in the discriminant analyses. Var-iables with an upper-case were more important from discriminant analyses. biologically letter were used or were easiest All remaining variables �89 Table 4.2 sites. Variables a Cold (!: < O. 05) significantly Spring Mountain, June-September Sprayed Random Variable random Random pb x - and grouse use 1982. Unsprayed Grouse ~ between differing All sites ---Random Grouse ~ ~ pb ._--- Grouse ~ pb ~ 4 9 0.1;; 15 26 0.00 13 16 0.25 DSAGE 10 12 0.60 2 3 0.10 3 8 0.00 TOTALCC 15 23 0.57 23 32 0.00 22 26 0.00 FOLIATCC 5 11 0.10 22 29 0.06 20 18 O./H HORCOV5 72 22 0.00 41 19 0.00 46 21 0.00 HORCOVIO 47 7 0.00 21 8 0.01 26 8 0.00 HORCOV45 92 88 0.55 92 82 0.03 92 86 0.07 CNTRHT 26 43 0.02 32 49 0.00 31 45 0.00 CNTRWD 29 54 0.02 44 65 0.00 41 59 0.00 CNTRINDT 16 35 0.05 24 41 0.00 23 38 0.00 SAGE DENSITYH 1.6 0.2 0,05 0.8 0.2 0.00 0.9 0.2 0.00 SAGEH 1.4 1.3 0.74 1.3 1.0 0.03 1.3 1.2 0.53 SAGEIDH 0.9 0.8 0.30 0.9 0.7 0.03 0.9 0.8 0.03 HORCOV5H 0.6 1.1 0.10 0.7 0.8 0.30 0.7 1.0 0.01 HRCVIOH 0.7 0.5 0.61 0.6 0.3 0.03 0.6 0.4 0.23 HRCV45H 0.1 0.3 0.27 0.2 0.5 0.00 0.2 0.3 0.00 TYPEDT 303 158 7 115 N 0.02 495 160 33 77 0.01 461 159 40 192 0.00 --_. --_--------_. aGrouse use bprobability cular variable; cSmal1er sites include juvenile. of no differ-ence 2-tailed numbers between unsuccessful random hen, and and grouse brood use hen sites for use the __ sites. partf- t test. indicate more vegetation obscuring squares on the cover board . �90 and all (combined) geneity variables sites: however, showed differing In sprayed areas. grouse (SAGE. DSAGE, TOTALCC. dom sites (Table 4.2). contrast. had greater (TOT ALCC) canopy sites. foliated covers, were compared near HRCVI0H sagebrush (Table cover between were further canopy cover and more variation in hori- HRCV45H) than at random 9 between in horizontal cover at 10 m random and all grouse was probably of sage grouse due to a threshold became completely to sage grouse; selection analyzed. habitat use effect obscured in of cover in sprayed In unsprayed (SAGE) was primarily canopy cover (TOTALCC) at grouse greater to greater random sites canopy covers). unfoliated shrub Crable 4.1; have cited the impor-tance of consequently, cover buted in sagebrush while variation canopy all sites, m habitats. Most studies ancies areas. use than random site", when all sites 10 m, in which the cover board sagebrush in unsprayed COver at 5 m and 45° (HORCOV5H. This lack of difference most sagebrush heterogeneities HRCV45H) similar to ran-- (SAGEIDH). at grouse 4.2), sites. (SAGE) and total shrub less variation in horizontal 9 and hetero- with canopy covers use locations (HRCVIOH) showed no difference sites. 9 distance HRCV45H) was greater among these sites at 10 m and 45° (HRCVI0H Variation cover FOLIATCC) and habitat Grouse (SAGEH) "and intercept results occupied (SAGEH. SAGEIDH. HORCOV5H zontal cover the remaining responsible greater cover foliated for greater use than random canopy discrep- and un sp ray ed areas areas, sagebrush canopy the apparent cover shrub sites; 9 differences while at (DSAGE) contri- (TOTALCC) at gr-ouse use than TOTALCC :;::SAGE + DSAGE + other Generally sagebrush shrub in SAGE or DSAGE led to �91 differences between random and grouse shrub canopy cover shrub canopy covers) These results select sites Because cover available with areas suggested ttonate results use of sites Therefore. with greater 4.1) Sage grouse composition (DSAGE). shrub density areas female juvenile (Multivariate average r'an-: srtes :;:: areas. in unsprayed showed dh:propor-· rather canopy than covel" alone. analyses rather which were highly correlated use was also examined for differences use sites shrub however. grouse random in had total canopy cover. sagebrush 0 with height sites there and for differences (MEANHT). and variation at female than was no difference Habitat use differed cover at 5 m (HORCOV5) and variation in forb sagein male locations between male and when examined with discriminant ~ te t , ~ >0005). between Canopy cover of un foliated (DENSITYH) were greater P < 0.05); t ha.n as unsprayed sage grouse among daily time periods. among grouse brush (DSAGE). 0 habitat males and females. horizontal in sprayed (DSAGE) grouse may not have been as much total canopy or FOLIATee, DSAGE. TOTALCC (Table (!_ test. cover TOTALCC was used in the discriminant than SAGE. sage 58~~of the random transects (SAGE) or unfoliated foliated overall. at unaprayed in spr-ayed that foliated in all comparisons. canopy there for use by sage grouse cover >21%. compared These SAGE + other foliated of TOTALCC at sprayed range of this difference. in TOTALeC; (TOTALCC) in sprayed of TOTALCC Only 28% of the random transects 0 that. sagebrush (DMRT, P < 0.05; :;;:6-22% and range 0 indicate had less total shr-i b cover areas = was similar- to random sites with more un foliated unsprayed 0-44%) FOLlATCC do not necessarily Random sites dom sites (FOLlATCC. use sites during analysis the day for in sagebrush canopy �92 cover (SAGEH); these were lower in the morning, and for HORCOV5. and lower. again. The 7 most common forb species grouse hen; use sites test in sagebrush common forbs (Eriogonum pussy toes on grouse umbellaturn) in the evening on transects (juvenile. X2 ":.: 9.2. of independence, unsuccessful of juvenile (Lupinus transects dandelion into stepwise Missing data sects. at: and Subsequent variables discriminant of the juvenile of juveniles locations in meadows (14% on 11 sites, qualitatively similar ful hen sites, analysis) 0 covariance TYPEDT); analysis hen sites assumes did not meet this assumption. or not homogeneity exists. sites. (SAGEH. r-emained 59 of 73 unsuccess-- were then that most of the transects the results however, Green without missing however. are homogeneous, in the analysis 24 of 73 unsuccessful were conducted (up to 68 of 84 juvenile Discriminant used of 88 t rarr- (N ::::144 of 232 transects). elimin.ation of these 9 group 40 variables Table 4.1). the elimination per hen sites and 30 of 35 brood matrices (Methods. 65 of 84 juven ile sites. analyses which forced HORCOV5H, HRCVIOH forced of transects 17 of 35 brood discriminant of 18 of the original analysis variables the number 38 of 40 random whether (!...~.£~~£~~.~ ~_:f.f~~i~ was the most common forb - ··A subset for several leaving hen sites, here The rnoe.t (79%). Multi:y'ariat~~nalys~~. entered and brood ammophi!__us). and rose Common dandelion 14 radiolocations locations). hen. amon g were aulpb ur+er-iogonurn , sand lupine At the (DMRT, ~. <0.05). did not differ nale ) was the most common forb at 3% (2/77) in sagebrush. at mid-day df ::.:12. P ::: 0.7). use transects (An_~~~nari~_!os.~~). higher group used in the varfance-: habitat data presented (1974) has proposed discriminant ft ric tions that may be used �93 and interpreted sistently separation than other interpretation. Therefore. unsuccessful analysis examined differences procedure is a function nearest cover type the absolute (MEANINDT. size Variables e ability and were groups and be are presented variable (HORCOV5) and variation SAGEIDH), 9 for (Table 4.3). An F statistic was used to test test the bird is to group The DF for all distance of shrub variation and width of the shr-ub beside this is a significance 0.2) (~> along the DF. (DENSITYH). means (centroids), DF's did value of each coefficient cover ran- Seven varf-: 2 additional of its associated of horizontal (TYPEDT), between 94% of the total explained DF coefficients and ecological interpretation groups was deemed valid. and brood hen sites. Standardized led in each group led to identical both power to separate to the contribution data presented equal to each other a significa.nt discriminatory in Table 4.3; distance groups with univariate probabilities DF (Table 4.3); retaining easily interpreted. separation in were interpretable, amount of sample variance on the first DF contained proportional hen, on the first a significant each variable analysis by the stepwise the smallest subset prior the discriminant dom. juvenile. by habitat results of transects discriminant sample variance DF's separate and were consistent setting The first ab les selected function of groups. or equal to the proportion not explain were satisfied the discriminant In addition" results. and (2) these which result DF's which do not have an ecologically meaningful to wide separation analyses. of groups. These criteria here because intercept (1) DF's con- have ecologically meaningful interpretations significant better biologically if they meet 2 criteria: to the crown in shrub density (CNTRWD) the equality of group for the Ma.halanobis �94 Table 4.3. Variables sites following stepwise Moffat County, (!'_ < 0.05) separating discriminant Colorado, random and sage grouse analysis, June-September Cold Spring Mountain. 1982. Group comparison -----..----_---------------------------Random juvenile V8. unsuccessful hen vs. Standardized grouse VB. brood hen correlation coefficient use R.andom vs. _------ .. all grouae si.tes .-----.~--.--- Cor-relation coefficient Standardized Unstandardizeda Variable HORCOV5 0.791 0.792 0.037 TYPEDT O.6()4. 0.()O2 0.003 MEANINDT 0.541 0.537 0.063 SAGEIDH 0.464 0.466 1. 778 DENSITYH 0.320 0,,330 0,633 CNTRWD --0.255 --0.249 --0.009 -4.334 Constant. 1.5 Eigenvalue Percentage of eigen-- value associated the first Canonical with DF 94. (3 DFa calculated) correlation Significance square 1.5 O. rt 0.77 of chi.- (!C) 0.00 _0. 00 ------_-----------------a 100 (1 DF calculated) A discriminant score for the function ing each unstandardized of each associated -------------------may be obtained corz-elation coefficient variable by multiply - by field-measured and summing the product.s values plus the cons tant , The discriminant score may then be compared along the DF (Fig. 4.1) the mean (group centroid) for habitat fo:," habitat at random Iocations . used by grouse and the mean to �95 distance between unsuccessful hen, but differences dom sites groups. There and brood hen groups did occur between (~< 0.00). use group. Variables cients use categories and associated in that grouse and proximity sagebrush (Table locations to edges intercept 9 centroids of grouse !:'. < 0.00) ~ test. power of discriminating number of correct and grouse distance the bird was determined. for classification), indicating that accurately summer habitat to those from random (HORCOV5) density 4,1). in (DENSITYH). 9 hut greater As in the first differed (Multf- To examine the group separation. transects = claasf fica-: 147 transects this set of variables the into random Overall correct was 90% (N coeffi- less variation (CNTRWD). of individual u.se sites function (MEANINDT) vaziab les in achieving tion of random and grouse use into 1 cover (TYPEDT). along the DF (Fig. use categories identify horizontal use and random groups classifications habitat differed (SAGEIDH) and shrub wicitb of the plant beside analysis variate types intercept and ran- were almost identical Grouse use sites of habitat shrub of grouse discriminant had greater drstance and smaller average average use category ~ > 0.28). was performed. analysis 4.3). test, were reclassified standa.rdized from the 2nd discriminant in the 1st analysis sites each grouse and a 2nd analysis among juvenile. (Multivariate!::. To improve interpretation along the DF, the 3 grouse grouse were no differences used (Table 4.3) of sage grouse. Discueaion Pr-evious studies described broad succulent forbs and height shifts of sage grouse in habitat and subsequent at grouse loc Hens. summer habitat use depending changes have generally on availability in sagebrush of canopy cover' Only 2 st udie s , both of which used can �96 cc cc a.: w w I-! l- C/) CI) I- W w cc cc w 0 cc ::r: 0 ::r: o Fig. CIJ _. 4.1. Discriminant Moffat bar and above JJ ..J ~3 RANDOM (!') Z function N == 106 scores Coun tv , Colorado. below mean i:J GROUSE USE .~ ~ =38 9 C/) Mountain, a: en w C/) 1 SE; of random and June-September ver-tical sage 1982. line is range. grouse use sites. Horizontal line Cold Spring is mean; �97 multivariate statistics grouse in relation dom locations: 9 have examined summer habitat to what may have been available at average Klebenow 's (1969) analysis random and brood sites, primarily sects) he analyzed This is the first report which found selective grouse. Additionally. few (N = (N >30 t ran- study habitat these are the first Summer habitats by measuring height. shrub densities plants) dense areas sagebrush density, did not show disproportionate or studies density been described canopy cover, (1982; only density Although Martin (1970) rep or ed that broods than adults, he did not quantify canopy cover was greater at juvenile (Table 4.4). grouse locations in Klebenow's Although Klebenow reported {Artemisia trid~ntata} than that at random sites. =: 15%). than and lower than random sites that big canopy cover at brood sites was less he did not report if total sagebrush between random and brood hen locations 12%; brood hen x Total but it was similar to (1982) study (1969) study, used less available habitat. random sites in Schoenberg's cover differed did not at brood locations was lower than at at random locations in this study sagebrush and use of available by Schoenberg or Klebenow (1969; total shrub but sagebrush random sites). borders. sagebrush in this study edges in of sage grouse of sage grouse have previously forb cover, Sage grouse sagebrush differ. data on heterogeneity and proximity to habitat by sage (Klebenow 1969. Wallestad 1971) have implied that sa.ge grouse used habitat summer, was 23) brood hen of an intensive researchers between (1982) study use of available summer habitat while other or ran- did not differentiate and since Schoenberg's of winter habitat. transects. use by sage Other researchers (random have reported canopy x ::: a va.riety �\J) 00 Table 4.4. Canopy juvenile sage cover grouse, (%) and height Cold Spring (em) of sagebrush. Mountain, and forb June-September Sagebrush canopy Grouse (1970) Peterson (1970) Klebenow (l96!f) aJuveniles bJune 24 15 84a 16 12 40 28 27 16 23 26 12 80 26 c Montana 12-21b Montana 1ge 47 ~40ntana 6-12b 91 Idaho 12f SD of brood habitat. Forb ground cover N x SD N 17 84 5 5 84 12 23 7 12 23 69 27,17d 18-2Sb 69 studies of radio-marked 33 41-51b 47 22 91 4 only. to September. cBlanks indicate dValues are means for 1968 and eEightv-eight fAll except sagebrush height x Colorado Martin Random N Colorado (1971) Sagebrush SD This study Wallestad cover x ~ (%) at relocations of previous 1'1 Location (1982) 1982 and results cover SD Source Schoenberg ground (~. no value percent 1969, respectively. of radio-located 3 of 98 broods tripa:rtita) reported. found were added broods <6 weeks old were in areas were in areas to obtain 12%. with <31%shrub cover. with average canopy cover Values for big sagebrush of 14%. and threetip �99 of sagebrush ing study reported canopy covers areas height study, did not differ birds and methods sagebrush as in this average Although sagebrush than height of values variables consistently of differ- grouse of shrubs shrub (l982) to random sites; at juvenile but height locations beside on random tr'an-: cover was not analyzed with respect reported values than random locations stru.cture to (Table 4.4) were within who ha.ve found greater forb (Klebenow 1969. Schoenberg and cover variables Mountain were similar to those described other because in relation height given by others These vegetation grouse Only Schoenberg of the center in this study, cover at juvenile 1982). at brood sites forb ground random locations the range (Table 4.4). from random transects. was greater sects. used by juvenile in other on Cold Spring studies; however. such as HORCOV5, TYPEDT, and DENSITYH more segregated sage grouse summe'r habitat from availab le rangeland. Results of uni.variate regard to variables grouse use sites. sites closer shrubs Both types which were larger unsprayed ar eaa , than In sprayed with great ~r shrub amount of available un foliated of analyses shrub sites, that grouse grouse may not be as important grouse used horizontal densities and and intercept roosted. beside between in canopy sprayed and may not be able to use canopy cover than average cover. random and Inconsistencies by differences areas. between with greater shrub average. were similar with indicated types Within these wer'e explained analyses differed and more homogeneous than average. cover results sites which consistently to the edges of habitat canopy covers. distances and multivariate because of a limited Whether cover. is mostly foliated or a habitat correlate to sage grouse �100 as total canopy cover. vide some cover Schoenberg sagebrush. f01" because several even fhe branches years after (1982) also noted that but that treatment sage grouse within sprayed of dead shrubs areas. with herbicide. used. areas grouse total sagebrush habitat may have been more homogeneous than average, were actually available variable types. canopy cover than average. selecting then one would expect than what was generally Klebenow (1969) reported brush stands Sage grouse because the optimal habitat available, 1f from a wide range of to be Iess Wallestad (l9'71) and that sage grouse used a variety but did not comment on the within+stand v of sagebrush of sage-- homogeneity structure. Daily variation patches an optimal habitat with dead use Gites s till had greater birds pro- were used, in sage grouse habitat but unlike other use indicated studies. there that habitat was little within season varia tion on Cold Sprin g Mountain. Durin g morrrin gs. birds fed in relatively which may have had greater forb cover. in areas homogeneous areas, while during the rest with more horizontal canopy cover. cover related reported open. There to brood movements on Cold Spring grouse summer because land if succulent variation the study forbs in sagebrush canopy Mountain as has been remained in sagebrush area was relatively Grouse broods are available, or free water and loafed areas in Montana (Wallestad 19'71). probably summers of the study. precipitation cover and greater roosted was no monthly change in sagebrush for lower elevation radio-marked of the day grouse throughout mesic during Most the both may remain in sageb rus h range-as in areas with >30 (Wallestad 19n. Autenreith ern annual 1981); or broods �101 may move to upland stad 1981) or bottomlands meadows (Autenreith 1971) which support succulent vegetation after forbs (Walle- have desiccated in more xeric areas. Multivariate selected trated analysis was useful by sage grouse.. on shrub zontal cover and habitat for identifying Unlike previous cover and structure, (not correlated interspersion studies with total shrub in summer. may be more important habitat correlates but in su.mmer the importance the basis showed t.hat hor-i-: !:. :.:::'-0.45) canopy cover. (TYPEDT) were 2 of the most important habitat alter correlates which have coricen-: this analysis minants of sage grouse seasons, habitat (habitat correlates) Shrub co tel' and structure for sage grouse of forbs deter- during in grouse diets other may of selection. Management Implications Wildlife biologists have urged lands consider the needs the sagebrush type support 1977, Autenreith geneity et al . 1982). summer sage grouse Differences habitat areas However, habitat in addition in seasonal (Autenreith diversity 1976). these habitat has not be n previously and density diversity of these guidelines of stands, et al , emphasize while this study on habitat has hetero- cover. investigated. have been well Summer use than average char'ac ter-i stic s within sagebrush types (Braun 1982). but within+summer should be managed for more homogeneous structure habitat use by sage grouse 1981, Schoenberg of adds further use is based to shrub r-an ge - obli gate species This study cover and structure, and interspersion. documented et al. of sagebrush and other for managing sage grouse management of sagebrush shown that of sage grouse (Braun to guidelines that managers and cover types sagebrush s tan ds , but a. (meadow, sagebrush, �102 and aspen). averaged to enlarge sagebrush of stand greater This and cover study than average areas, alteration previous on sage grouse components it may be 1981:65); should be managed according reports The effects therefore, sage to gtrid elirrea important et al . for the summer range. that sage grouse cover in both sprayed on sage grouse r'eith et al , 1982) which preserve of live of summer sage have indicated and of the detrimental (Martin 1970, Braun of sagebrush are closely r-elat ed to the availability (Autenr-eith as meadows, types sagebrush 1976. Autenr-eith et 0.1, 1982)" range 150-200 m as a guideline and others and support of sagebrush sage grouse (mostly meadows), the edges of meadows (Braun and for'b s are essential habitat, unsprayed effects around Managers might also uae Sagebrush select type edges Since sage grouse from 100 to 200 m the recommended strip to be retained interspersion grouse exist in close proximity, 159 m from habitat prudent 1977) , should on. of forb s on. aummez grouse (Braun alteration et 0.1. s ummar habitat et al , 19'1'1. Auten- forb producing areas" such �103 LITERA TURE Aldrich, J. W. on idae , 1963. Geographic orientation J. WildL Manage. Amst rup , S. C. Manage. 1980. CITED A radio-collar T. 1981. Sage grouse practices. Idaho. D. I., grouse. ---- Condor, management 9. Bull. 1. E. Braun. A. Agric. C. Exp. E.. grouse 1970. T. maintenance Recommended Britt, of sage Schroeder. alteration from wing Stn , Res. • M. F'. Baker. 1976. Editors. States 1982. Sage Grouse Sage grouse Comm .., Twin 42pp. 1978. Weights J. 31. 1975. of Colorado sage grouse Conservation 88:165-171. Colo. Div , Wi.ldl. 4pp. plant names. Urriv , Wyoming 124pp. habitats. R. L. Eng, Sex and age determina- characteristics. and R. O. Wallestad. of sagebrush Wilson Bull. Idaho 80:241-243 . Leafl , 49 (revised). A. in Idaho. 238pp. • R. B. Gill. and C. E. Braun. Game Inf. Braun. West. Tech. and C. tion of sage Beetle. J. Wildl. for game birds. , W. Molirii , and C. E. Braun. management Beck, 'I'et ra- 27:529-545. Dep , Fish and Game. Wildl. Bull. Falls, American 44:214-217. Auten rerth , R. E. ---- of North T. Wildl. Soc. Guidelines Bull. for 5: 99--106 . S. Gashwil r , and M. H. committee communities 1977 . report on effects on the associated of avifauna. �104 Canfield. R. H. 1941. in sampling Application range of the line interception vegetation. J. For. 39: 388-,394. Dauberrmir-e , R. F •• and J. B. Daubenrni re , of eastern Stn. Eng. Washington Tech. B'ull , 60. R. L .• and P. rnen ts and and northern 1968. Idaho. Forest vegetation Wash. Agric .. Exp . l04pp .. Sc hladwetler , habitat method 1972. use in central Sage grouse Montana winter move- .1. Wild}. Manage. e 36: 141--146. Giesen. K.. M., T for trapping J. Schoenberg, e sage grouse a.nd C. E. Br aun , in Colorado. 198;";. Methods WildL Soc. Bull. 10:224-- 231. Green. R. H. 1974. Multivariate in g environmental Jones. R. E. 1968. factors. Idaho. Krebs. J. R .• and C. Pr aeg er Publ . N. S. grouse eds . occurrence" coverage 5:):73---83. cover 1980. Sagebrush nesting 1978. Pages Population New York, 9 va.r-y- used by pr'an-ie grouse. an d brood Behavior 23-,47 control in ~_I2:. and population F. ,J" Ebling and by social b chavior . N"Y. control related J. Wildl. Manage. to habitat an d sage 34: 313--320. Line-dn tez-cep t sampling and density" habitat 33:649-661. J. Perrins. 1970. McDorra'[d , L. L. than Sage grouse (Pa~~_~ ~~~£). tit with temporally 32:28-31. 1969, D. M. Stoddart. Martin. Ecology J. Wildl. Manage. the great analysis A board. to measure J. Wildl. Manage. Kleb enow , D. A. niche of attributes J. Wild.L Manage" other 4A:530--S3.-·L in �105 a.. Nie, N. Co H. Hull, Bent. 1975. Statistical Hill Book Co. Noon. B R. 0 9 Oakleaf. Ecol. R. J gradient: Monogr. R. L. Inc 0, Denver, Peterson" J juvenile sage of an avian guild along a temper- and expression of sage Thesis, The sage grouse Un iv , Nevada, grouse of competi-- to uplan d Reno, in Wyoming. 73pp. Sage Books , 341pp. The food habits grouse McGr'aw- 675pp. the importance M,S. Colo, 1970, 0 N. Y. The relationship 1952, and D. H. 51:105--124. meadows in Ne vada . Patterson, K. Steinbrenner, for the social sciences. The distribution 1971. 0 package New York, 1981, ate elevational tion. J. Go Jenkins, and summer in cen tr al Montana, J 0 distribution WildL of Marrag e , 34: 147-155. Schoenberg. T. J e tion in North Fort Scott, Park, Collins. T, G., 1982. Sa.ge grouse Color-ado. movements M.S. Thesis, and habitat Colo. seIec'- State Un.iv. , 86pp. and C. H 0 Wasser. plan ts for wildlife biologists. 1980. Checklist of North American The Wildl. Soc , , Washin g ton , D" C . 58pp. U.S. of Interior, Department Colorado Bur. coal regional Land Manage. Walles tad , R, O. grouse broods 1975, in central Di v . 1971, ..t" supplement environmental DES 76-21. Life history U.S, Dep . In te r , , 555pp, Montana. and habitat use by sage J. Wild!. Manage. and habitat Mont. Dep. to the northwest statement. Summer movements in central Montana. 65pp. 1978, requirements 35: 129-136. of sage grouse Fish and Game, Game Manage. �106 ____, • and P. Schladweiler. habitat selection 1974. Breeding of male sage g rouse , season movements J. Wildl. Manage. and 38: 634-' 637 . ---- • J. G. Peterson, grouse in central Wiens, J. A., and R. Montana. J. WildJ. Manage. and J. T. Rotenber.ry. community structure of birds 19'15. L. Eng. 1981. Foods of adult 39:628-630~ Habrta t as socia ti('y"" in shrubsteppa sage environments. an d. Ecol , Monogr.51:21--41. Zar , J. H. Cliffs. 19"/4. N.J. Biostatistical 620pp. analysis. Prentice-Hall, Inc . , Englewood �107 Colorado Division of Wildlife Wildl ife Research Report April 1984 JOB PROGRESS Colorado State of Project REPORT W-37-R-37 Work Plan 3 Job Title: Game Bird Survey (45-01-504-15050): Job: 15 Sage Grouse Distribution and Habitat Use in the Gunnison Basin Period Covered: 01 January Author: Jerry HURP Personne. J : through 30 June 1984. C. Braun, C. Coghill, J. Houston, T. Henry, P. Mason, J. Olterman, D. Reynolds, T. Sherill. Colorado Division of Wildlife; J. Hupp, R. Ryder, Colorado State University; M. 81ymeyer, J. Capodice, B. Kaufman, C. Scheck, U.S. Bureau of Land Management; C. Molitoris, B. Wall 1s, E. Zleroth, U.S. Forest Service; K. Lair, Soil Conservation Service; D. Bruno, J. Hollingshed, Western State Co l l eqe . ABSTRACT Sage grouse (Centrocercus uropnasianus) distribution and habitat use were studied near Gunnison, Colorado during January-June 1984. Sage grouse were disproportionately distributed in the Gunnison Basin during January-March. Areas near South Willow, South Beaver, and Gold Basin creeks may be important wintering regions. Areas northwest and northeast of Gunnison and north of Parlin may also be favored \~intering areas. Sixty male and 13 female sage grouse were captured and marked between 21 March and 28 May. Radio transmitters were attached to 22 individuals (10 males, i2 females). Topography and shrub structure were evaluated at sites used by winter flocks and radio-marked birds. Sage grouse primarily used drainages in early winter but increasingly used sites with south or west aspects as snow cover diminished in late winter. Sagebrush stands at early winter foraging areas and at nest sites were more thickly canopied and consisted of larger plants than sagebrush stands at late winter and spring use sites. Sage grouse strutting acti0ity was surveyed on 16 leks. Mean, peak male attendance (23.3 males) was slightly lower than during 1983 (34.0 males). Reduced .attendance probably was due to poor, early spring body condition among males following severe winter wes the.r. Helicopter surveys for previously unlocated leks were conducted be tween 8 end 10 Flay. Hale flocks were observed in 4 areas where leks were not known to occur. Ten sage grouse nests were located during the nesting season. Mean clutch size of 7 completed nests was 6.9 eggs. Nesting success was 56% with the peak period of hatching occurring during the 2nd and 3rd weeks of June. Distance between leks where females bred and nest sites ranged from 0.7 to 8.2 km = 4.2 km). ex ��109 SAGE GROUSE DISTRIBUTION IN THE GUNNISON AND HABiTAT BASIN USE Jer rv Hupp P. N. OBJECTIVES The primary objectives of this study are to: (1) evaluate winter and spring distribution of sage grouse in the Gunnison Basin, Colorado. (2) descr ibe vegetat ion and s t ruc tura 1 component"; of seasona 1 sage grouse use sites~ and (3) develop a sage grouse habitat management plan for the Gunnison Bas in. - ... - SEGMENT 1. Describe \'J!nter availability spp in the Gunn j son Bas in. i 2. OBJECTIVES and d Is t r l bu t l on of sagebrush (~!_t.E\~~L?!..~_ ) Evaluate sage grouse Basin. 3. Capture sage grouse winter distribution ~.'~. in the Gunnison on wintering of body 5 ize and we iqht. radio transmitters. areas ard on leks. Obtain measures Mark captured 1nd IV lduaI 5 wi th leg bands and 4. Monitor sage grouse movements beb~een wintering areas and leks. 5. Measure and compare shrub structure at winter and spring sage grouse use sites. 6. Locate previously undiscovered sage grouse leks in the Gunnison Basin. Survey sage grouse lek attendance. 7. Obtain lnformat lon on clutch s ize and nesting the Gunnison Basin. 8. Collect male sage grouse for SUCCE;5S of sage grouse in analysis of spring lipid reserves. 9. Analyze data and prepare annual Job Progress DESCRIPTION report. OF SlUDY AREA The study area is an intermontane basin that includes parts of Gunnison and Saguache counties" Co Iorado , ['!evatlon ranges from appr-oxlmate ly 2,300 to 2,850 m. The basin Is semi-arid with a mean annual precipitation of 28 em at Gunnison. ~lost: pr-eclp i t a t lon occurs from .Januarv t hrouqh March. Upland area,-i are moderately ro I '! ing and are dissected by tributaries of the Gunnison River" Broad. fl a t flood plains occur at lower elevations along major streams. Soils are primal'lly gravelly or sandy loams. Big �1'10 sagebrush (A. tridentata) is the dominant shrub vegetation in upland areas. A. nova grovis '(;"il-dry-rrdges. Pur s h Ia tr Idenre te and Amelanchi~r a l n l fo l l a are interspersed \'IIi th sagebrush s ftes. -Rabb i spp.) is a common understory shrub that can become dominant fo l l ow i nq removal of the sagebrush canopy. on-some Understory forbs include species of Phlox, As trag a Ius. ~~ t ~.~~"and 0 t he rs . DomTnaii-t tbr·ushl}~liEYI0t.6~1~!'ii~{ Lupinus, Taraxacum, Polygonum. '9 r as's'e-s' iIl"'upT;.j'na· '<3 r'~;:_;s-i;:Ic"Tud e AgroPXE£.12. and £,oa. Quak inq aspen (p(P~Y..:::, !E~~ml:!..1~i_~,5:':;) usual l v grows above 2.900 m and on moist sites at lower elevations. Sagebrush uplands ar e used as spring and summer pas tur e for domestic 1 ives tock , Large concentrations of wild ungulates also occur in some areas of the Gunnison Basin during winter months. Lower el evat i on flood pl a l ns dre used as irrigated hay meadows. _Fest~. METHODS Distribution Sagebrush Availability 9 regions for' s tudv of ':;'~9C ~J,rOIE;e and Saq eb rus h distribution was eva luated from Land Management (BLt,,1), U.S. Forest Service (USFS), and Soil ConService (SCS) so l 1 survey and vegetation dat a . These d a t a we r e to a map (scale ~ 1:126,700) of the Gunnison region. The Gunnison sagebrush Bureau of servation transferred and Winter Basin was divided distribution (Fig- inc) 1) .. winter' sagebrush ava Llsb ll i ty in the Gunnison Basin was evaluated from transects on 2.L}-25 January. Approx ime tel y 390 km of transects wer e via f ixed-wl nq a i rc r aft at 150 m e l eva t Ion . Air speed was 130 km/hr • .Transects were genera 1 Jy or tented north-south and spaced at 3. 2",krn i nt erv, 1s . An observer on the pas senuer side of the cabin looked through a 25-..c.m2 viewing square marked at shoulder l evel on the window. At '15~5,,;,cond intervals, presence or absence of exposed sagebrush in the vi e\'II i n9 square was recorded. This provided a percentage estimate of exposed sagebrush in each region of the Basin. A 2nd observer marked areas of exposed sagebrush on 7.5' topographic maps and guided the pilot along the correct course. Early aerial flown Ground transects to evaluate saqebru sh ava l l ab i 1lty wer e also conduc t ed , t rans ec ts were established near Chance and Grafl in qu l ches be tween B and 5 February , Tran scct s were stratified between 7,438 and 3,834 rn , Sagebrush exposure above the snow was recorded at 25-m i rrterva 1salon] transects. Snow depth was measured at 100-m intervals along transects. Terrain featu res crossed by transects were a 150 recorded. Twerrty-f'our , 0.5-km Sage Grouse Ground Winter surveys via Distribution snowshoes , skis. and snowmobiles were systematically con- 7 .Ianuarv and 3'j M~H'Ch to evaluate sage grouse d i s tribu ti on in the Gunnison Basin. Bour.darles of survey aree s we.re mar ked on tr-poqraph rc maps so the amount of area 5urv~ycd In each of the 9 regions could be determined. Areas used by sage grouse were markE~ on topogrdphic maps. Due to extensive 5nOIf../ cover, it W(l', P()~.Lb l e to identify use s i tcs f rorn sage qrou se ducted between tracks as of observed we l I as flocks flock were s i qbt i nq s . recot d ed . Whel~c f-'O~:;sibIe, s I ze and sex compos l t l on �,. PARLIN i 1 L- - Fig. 1. Boundarles ~Inter distribution of 9 regions in the Gunnison were conducted. Basin in ~hich systematic ground sur~ey5 of sage grouse �112 Sage Grouse Capture and Marking Sage grouse were captured on w l n t eri nq ar eas and 'leks us l nq n l qh t l i qh t l nq techniques (Giesen e t a l . 1982). In addition, an incubating adult female was nest-trapped and radiomarked following Smith e t a l • (1980) without causing abandonment. Body weight, carpa l , t arsus , and culmen l enq ths were measured. Approx irnate ly 1-2 cc of blood was obta ined f rom each i nd i,. vidual for electrophoresis ana l vs l s and examination for blood paras i te s . Age was based on ou t.er wing primary characteristics (Beck e t 31. 197~,). All birds were marked with aluminum and colored plastic leg bands. Colored band combinations were specific to l nd Iv i dua l s or different regions of the Gunnison Basin (Fig. 1). Radio transmitters were attached to some ind lv i-: dua 1s. Poncho-mounted ~ 50 l e r+powe red tr ansm i t t ers (25-28 g) wer e attached to females while tall-clip transmitters (25 g) were attached tc' central rectrices of males. All radio t ransm itte r s were within 150.850-151.1+00 MHz. Habitat Use and Move:Tlents Winter (7 ,Jan-31 Mar) habitat use was based on sage grouse observations recorded during svs temat i c qround surveys. Use areas included f Iv sh sites and areas of recent foraging activity bJsed on tracks and droppings. Tooographic features were recorded at all use sites. In addition, some winter use sites were marked for veqet at l on analysis. Two, 30-m transects were es t ab 1 i shed a 1onq nor t h+sou th and ees t+wes t axes. Transects I n tersec t ed over the appr ox irnate center of the act IV i ty area. SnON depth was measured at the center point and ends of transects. Exposed sagebrush within 1.0 m of transects ""as examined for evidence of sage grouse feeding. Fed-upon plants were marked with colored vinyl ribbon. Spring (1 Apr=31 May) habitat use was birds" Radio-marked sage grouse were receiving antenna. Radio signals were were flushed, Flush sites were plotted phic features of spring use sites were Vegetation at sage grouse nest monitoring to other rad lo-marked based on flush sites of radio-marked relocated with a hend+he ld , 3-e-lement Fo l l owed until marked individuals on 7,5' topographic maps. Topograalso recorded. sites was described f emales . Some nests were at nests located by also located incidental field activities. Shrub structure at seasonal use sites was measured following snow melt. Modified Canfield (1941) line transects were used to describe shrub vegetation. Two , 15-m transects were oriented along north··south and east-west axes above the use site. Sagebrush canopy intercepted by the transects was measured to provide a percentage estimate of canopy cover at each use site. Height of individual sagebrush plants beneath the transects was measured at the tallest 1 iving stem. Crown length of intercepted plants was measured along the longest crown axis. Crown width was measur-ed at an axis perpendicular to length. TI)'~ combined measures of he iqh t , width, and length were used as es t lma tes of plant size. Schoenberg (1982) found shr ub plant size was a useful measur1 of sage grouse habitat. Sagebrush density was measured within D.S-m of t ransect s , Crown height, width, and l enq r.h of fed-upon plants was also measur-ed c't w l n t e r us~,~si t es . �113 Mean sagebrush height and plant size were calculated for each use site. Analysis of variance was used for a preliminary comparison of percent sagebrush canopy, density. mean height. and plant size among seasonal use sites. During the analysis a distinction was made between early (7 Jan9 Mar) and late (10-31 Mar) winter periods due to reduced snow cover in mid- and tate March. Lek Surveys Locations of known sage grouse leks were plotted on 7.5' topographic maps. Leks were visited during early morning hours (0430-0700 ~1ST) between 1 April and 25 May. Numbers of attending birds were recorded. Al thouqh most leks were visited 2-4 times during the display season. poor access prevented multiple counts on some strutting grounds. Searches for previolJsly unlocated leks were conducted from a he I l copt er during early morning periods (0430-0700 MST) between 8 and 10 May. Radio-marked sage grouse were also monitored during early morning periods to locate previously unknown leks. Nest ing B l o 'logy Radio-marked females were monitored during the display season to evaluate dis t ance between breed ing and nes t sites, and to obta in informat ion on nes ting success and clutch size. Clutch size and fate of incidentally discovered nests WEce also recorded. Li p i d Ana I ys i 5 Ten adult and 5 yearling male sage grouse were trapped and collected on or near leks during 11 April-May, early in the display season. An additional 10 adult males were collected between 14 and 18 ~;ay towards the end of the display season. All birds were frozen within 4 hours for preservation. Lipid reserves of collected individuals will be measured. RESULTS AND DISCUSSION Sag~brush Distribution and Availability Approximately 1,474 km2 of sagebrush-dominated lands are potentially available to sage grouse in the Gunnison Basin (Table 1). This figure is preliminary and subject to change pending verification of map data. The estimate may also be modified foliowing compilation of sagebrush treatment records. Sagebrush availability varied among regions In late January (Table 1). Sagebrush was least available in the Steuben, Cochetopa, and McIntosh regions (0-3.5%). Sagebrush availability was greatest in the Beaver and Parlin regions (11.7-'11.8%). Over a ll, exposed sagebrush was available in only 6.7% of the Gunnison Basin in late January. �114 Table 10 Sagebrush distribution and winter availability in 9 regions of the Gunnison Basin. Distribution data were compiled from f eder-aI agency records. Winter availability was estimated during aerial transects conducted on 24-25 January 1984. . a R eglon '---'---' Sagebrush .__ .__,_~!1:~L_..__ ,.,_ ..__ ~_._. Exposed , 96 Mcintosh Cochetopa Steuben Doy 1ev i 11 e Tomichi Beaver Chance Parlin Sapinero '"". __ ._--_._-- in winter <% L,__ . ..__ .3 .' ".) '108 2.1 119 0 145 7.6 1lf8 6 ..7 172 11..7 4, ,,l-f 11 ,,8 '199 225 263 7.9 1 ,474 Total Average 6, '/ dSoundaries as delineated in Fig. 1. Sagebrush availability also varied among topographic features during early winter (Table 2). Sagebrush was disproportionately available in drainages and draw bottoms (X2. :.= 11.8. p < 0.01). Although only 23% of t he points sampled along ground tr ansec ts were in drainages, 41% of the exposed sage'brush was in drainage sites. Sagebrush was least available on 6·'20% slopes with north or east aspects. Exposed sagebrush in drainages vias likely Ao tridentata vaseyana as growth of this subspecies is favored on moist sites Cfisdale---and-Hironaka 1981). Because of its vigorous growth form, A. l. ~as:~yan~ was available in draw bottoms in spite of deep snow (Table 3). Table 2. Distribution of exposed sagebrush among 4 terrain categories in the Gunnison Bas in. Sagebrush avai lab l1 ity was measured along 2[1. O.S-km ground transects on 3-5 February 1984. Proportions are in parentheses. N ___ . Terrain . . .samp_!!~ p'?.! n t E ___ ~_ exposed s_agebru_~,~.lan t:~ Dr a i nage 108 (0.27.9) 18 (0,,409) 0-5% slope 176 (0.372) 17 (0.386) 56 (0. 119) 5 (0.113) 6-20?6 slope, south or west aspect 6·· 20% slope" north or east aspect Total X2 = 11.8. ~ < 0001 '132 (0.280) II (0.091) , �115 Table 3. Mean snow depth among It terrain categories in the Gunnison Basin. Snow depth was measured along ground transects near Chance and Grafl in gulches on 3-5 February 1984. ,----_._--x Terrain Ora i nage 0-5% slope 6-20% 510 e, south or west aspect 65.0 53.5 SE 9.9 19 ..7 39.1 6-20% slope, north or east aspect 57.3 2,5.1 Sage Grouse Distribution Ninety-five areas of sage grouse activity were observed between 7 January and 31 March. This included 46 s Iqh t inqs of sage gr-ouse flocks. Flock size ranged from 'j to -'SO individuals (x.::: lL~1). Sex of flock members was identified at 34 observation sites.- Sixteen flocks consisted of maJeonly (x == 5.0 individuals) whi l e 5 flocks were comprised sol e l y of females (x = 5:-4 individuals). The remaining 13 sage grouse f Iocks contained both sp..xes(x = 11.7 individuals). The proportion of flocks (47.1%) containing both sexes of sage grouse is greater than observed by Beck (1975) in North Park, Colorado (19%). Sage grouse were observed throughout the Gunnison Basin (Fig. 2). However flocks were not evenly distributed (Table 4, X2 = 35.29 P < 0.001), A disproportionately large number of flocks were o"b-served in the Beaver, Chance , Parlin, and Mcintosh regions. Few sage grouse were observed in the Steuben, Sapinero, Tomichi, Doylevil1e, and Cochetopa regions. Areas near South Willow, South Beaver, and Gold Basin Creek drainages may therefore be impor-tant sage grouse wintering regions. Areas northwest and northeast of Gunnis6n, and north of Parlin may also be favored winter sites. Additional winter studies are needed to confirm sage grouse winter distribution. Sage Grouse Capture. Marking, and Body Size Sixty male sage grouse (42 adul ts , 18 yearlings) were trapped and marked between 21 March and 21 May. Thirteen female sage grouse (9 adults, 4 yearlings) were marked between 29 March and 28 May. One banded yearling and 1 banded adult male, and 2 marked adult females 'V~ererecovered fo l l ow i nq mortality. Radio tr-ansml t ter s were attached to 22 sage grouse (Table 5). Three radio t r ansm i t ters were recovered following predation of marked birds. Six transmitters were recovered from males following breakage and premature molt of central rectrices. The high recovery rate of tail-clip transmitters may indicate that the radio package is too heavy, and causes premature loss of central r ect r lces , �,. 0' -II! N S"''Ie U~(l Gpo ••se S;r~ \ /-f:..j ..• \ o -\ . ~ ) J ~ ( ~ ~. 0, '1\ • o. l.~\-:-.. 0 ~~I.r A\ • ~ \. .~-- , ~~ \~ ?: "\ \1 ~( .,. I \ t r .0. ') ~~ _f< e e. ~! / =) \ i I- Fig.? Distribution of observed sage grouse use sites In the Gunn!son Basin, 7 Janu~ry-31 Locat1ons may represent> 1 observation. March 1984. �117 Table 4. Chi-squaie goodness of fit for distribution of 95 sage grouse observations among 9 regions of the Gunnison Basin during winter 1984 (7 Jan-31 Mar). Proportions are in parentheses. Area • Reglon Use surveyed a Steuben Mcintosh Parlin Tomichi DoylevilJe Chance Beaver Sapinero Cochetopa (ha) Expected 1,800 3 (0.029) 5 (0.048 20 (0.213) 5 (0.056) 2.956 13,110 3.423 Total 10 ~852 17 (0. 177) 13,668 9,085 5.034 1.500 21 (0.223) 14 (0. 148) 61.428 95 Table 5. 2.5 (O ..263) 2 (0,021) 2 (0.021) 27 (0.284) 25 (O.263) 4 (0.042) o (0.0) 2 (0.024) as delineated Radio-marked 1 (0.011) 9 (0.09S) 8 (0.082) P °Soundaries Observed <: 95 0.001 in Fig. 1. sage grouse in the Gunnison Basin, March-June 1984. Radio rrequ Band no. ncy (MH!L_ Males 01 02 03 04 05 06 07 15 23 57 A~e 51.052 151.285 151. 000 151.910 150.939 150.968 151.013 151.358 150.980 151.018 Date OJ captured 21 Mar 10 Apr 11 Apr 11 Apr 13 Apr 17 Apr 21+ Apr 30 Apr '{rIg 'j Yrlg Yrlg Ad Yrlg Yrlg Yrlg Yrlg Yrlg Yrlg 11 May 18 May Date recovered Cause of reco'l:ry 01 Apr Predationb Tai I molt Taj I molt 17 Jun 19 Apr 24 May Tai 1 molt 26 May Tal J melt 29 May 10 Ju I Tai I Molt Tar i molt 11 May Predation 23 May Preda t l on Females 150.876 151.131 150.908 150.863 151.041 150.955 151.343 151.254 151.238 151.159 151.283 151. 311 5701 5702 5703 5704 5705 5706 5708 5709 5710 5711 5712 5704 Yrlg Ad Yrlg 29 Mal' Ad Ad Ad Ad Ad 22 Apr 22 Apr 22 Apr co Apr 211 Apr 01 May 06 May Yrlg Ad 06 May 07 May 28 May Yrlg Ad --.-------.--.--.-----.aAd 10 17 Apr •.-"'-- .... ~-.-~-~-.-~'------"-. edu lt , Yrlg ~.year l ing. bTransmitter recovered following breakage of premature molt of central rectrices. �118 Gunnison Basin sage grouse are physically smaller than other studied sage grouse populations in Colorado. Across all sex and age-classes, sage grouse captured in the Gunnison Basin in 198/ .• weighed 26-29% less than qrouse cap-tured in North Park, Colorado during a similar period (p < 0.01, Table 6). Body structure differences also exist. Carpal length of Gunnison Basin sage grouse is approximately 10% shorter than in North Park sage grouse (p < 0.01, Table 7). Body size variation could indicate there is little gene flow among Gunnison Basin sage grouse and other- Colorado popul a t lons , Isolation is possible due to surrounding alpine habitats that could re"trict dispersal. Table 6. Comparison of body weights between sage grouse captured in the Gunnison Basin and North Park, Colorado, March-May 1984. Significant differences are indicated by f < 0.05. Gun!'i son Ba-STn"---.---.-----NOrti;-p a rT·~·-.-·---<-·-"------·---. ..SE N Males Adult Yearling 42 2,038 1,786 16.6 2].6 97 18 9 4 1 ,210 1,084 23.4 38.4 19.7 2,890 2,502 21.1 0.0001 0.0001 109 1,645 10.7 0.0001 119 1,485 14.6 0.002 95 Females Aduit Yearling -------_._--'--_._-_.-._-------------_._-_------ .. _ •.._-----.Table 7. Comparison of carpal lengths between sage grouse captured in the Gunnison Basin (Mar-May 1984) and birds captured in North Park, Colorado (Mar-May 1983). Significant differences are indicated by f < 0.05. Gunnison N ~(nim) North Park ----------- Basin SE N p .?5..(mm} SE 0.46 0.52 0.0001 0.0001 0.0001 0.004 Males Adult Yearling 42 302.9 18 294.5 0.91 2.31 170 152 337.2 329.4 9 261.9 251.0 3.30 34 28 2.83.7 1.02 3.0 276.7 0.87 Females Adult Year u ns 4 -------- .._---------_._----------_._----------- �119 Habitat Use Topographic features were recorded at 91 winter use sites (Table 8). Sage grouse primarily used drainages and draws during January. Use of drainages declined in February and March as birds increasingly moved onto 6-20% slopes with south or west aspects. Use of these terrain types was consistent with observations of sagebrush availability. Ground transects indicated that sagebrush was primari ly avai lab Ie in drainages in early February (Table 2). As snow settled on south and west slopes. sagebrush became more available in late winter. Grouse responded to improved availability by increasingly using these sites. Table 8. Distribution (%) of sage grouse use sites among 5 terrain categories in the Gunnison Basin between January and March 1984. Jan Feb Terrain (~ '" 29) (!! = 44) Drainage 45 34 22 0-5% slope 27 5 0 14 43 6'1 0 9 a 14 9 11 Mar ----18) (!J.'" .----.- 6~20% slope~ south or west aspect 6-20% slope~ north or east aspect Ridgetop -------Shrub structure varied among seasonal use sites (Table 9). Mean sagebrush canopy, height, and plant size differed among winter. spring, and nest sites (p < 0.01). There was no difference in sagebrush density among seasons-(f> 0.05). Table 9. Comp~rison of 4 measures of sagebrush shrub structure at seasonal use sites of sage grouse in the Gunni~on Basin. Colorado, 1984. Significant difference among seasons indicated by ! < 0.05. Sagebrush density (pJants/m2) Sagebrush canopy (%) Season ~ x SE x SE Early wlntera l.1lte_wlnterb Springe Nests 10 1.0 1.3 0.1 0.3 0.2 32.9 2.8 16.3 1.3 23.9 31.3 1.8 2.7 7 29 9 1.5 1.4 o.s 1 .1 II 0.05 F f_ " a07 January - 09 M reh. b10-31 March. COl April - 30 May. 6.86 P < 0.01 Sagebrush height (em) x Sagebrush plant size (dm3) SE X SE 62.9 26.5 15.0 499 93 34.5 45.7 1.6 156 70.2 23.1 16.7 2.8 251 32.7 -----------29.16 P " o.ot 9.4 22.35 P " 0.01 �120 Early winter- deep snows great'Jy reduced sagebrush avai labi 1 ity in the Gunnison Basin. Only the tall vigorous sagebrush plants growing in moist drainages were available to sage grouse. Low-growing saqebrush on drier' slopes and ridges was covered by snow. As a result, mean sagebrush canopy, height, and plant size were greater at early winter foraging areas than at other seasonal use sites (Table 9). Sage grouse primarily fed on the tallest sagebrush plants within winter use sites. Mean height of fed-upon sagebrush plants was 8g.7 em (SE ~ 9.4) while plants not used by foraging sage grouse averaged 57.7 cm (SE : 3.8). The difference reflects the extent that smaller plants at early winter use sites we re covered with snow, Because sagebrush availability is greatly reduced in severe winters, maintenance of tall sagebrush stands in drainages is important to provide exposed forage and cover for sage grouse. Sage grouse winter preference for vigorous. thickly-canopied (> 20%) sagebrush stands has been previously observed by Eng and Schladweiler (1972), and Wal1estad (1975) in Montana, and Schoenberg (1982) in North Park. Colorado. Sagebrush on late winter use sites was much shorter than on early winter foraging areas (Table 9). Fed-upon plan ts were only 51 ightly taller 29.1 cm, SE = 4.2) than non fed-upon sagebrush (i= 24.9, SE = 3.3) at-late winter use sites. Smaller plant size reflects the more xer ic nature of soils on south and west facing slopes where grouse primarily foraged in late winter (Table 8). South and west facing slopes may have been preferred late winter foraging areas because reduced snow cover allowed easier movement. Also? growth of A. t . wyomingensis is favored on upper slopes and ridges (Ti sda I e and Hi rona"ka T9g-Tr~"-sage"grouse have been observed to pref erent la lly forage on /2. t, ~on:..!_n.gensis due to its higher nitrogen and lower monoterpene content TRemington 1983). ex "" Sagebrush at spring use sites was also shorter than at early winter use sites (Table 9). In spring, shorter sagebrush plants were available due to reduced snow cover. Also sprjng habitat use was modified by proximity of sites to leks. Mean distance of radio-marked males from leks was 1.4 km (SE = 0.3, N = 17) once display activities started. In spring, sage grouse may not use-vigorous sagebrush stands? unless such stands are readily available near leks. Mean canopy cover at spring use sites (2309%) was less than that reported in previous studies by Wallestad and Schl adwe ller (1974), Emmons (1980)? and Schoenberg (1982). Sagebrush plants at nest sites were taller than at spring and later winter sites, but shorter than at early winter foraging areas (Table 9). Nesting females probably used more vigorous sagebrush stands because they provided better nest concealment. Maintenance of vigorous sagebrush stands is important to providing nesting habitat. Kl ebenow ('i969) , Wallestad and Pyrah (1974), Petersen (1980),and Schoenberg (1982) also report sage grouse use of vigorous (> 20%) sagebrush stands for nesting. Wi nte r to Spr i ng Movemen t Four male and 3 female sage grouse were captured and radiomarked between whe re wl nter i nq South Beaver Creek). 29 March and 17 Apr l 1. All birds were captured in areas flocks had been observed (Antelope Gulch, Gold Basin,and �121 Each of these radio-marked individuals moved long distances (12.6-33.0 km) from capture sites following radio attachment ('ri~&-. 3-4).· Birds radiomarked after 17 April generally moved only shor'Zdistances « 10 km) from capture sites. Long distance movements occurred prior to peak lek attendance in late April and represented migration from wintering to breeding areas. Spring movement from Antelope Gulch, Gold Basin, and South Beaver Creek support observations on the importance of these areas to wintering sage grouse. Lek Surveys Sage grouse strutting activity was surveyed on 16 leks (Table 10. Fig. 5). Fourteen leks were active in previous years. Two leks (Mashburn 1 and 2) were discovered during the display season by monitoring radio-marked birds. Persistent snow cover delayed display activities on most leks. Little display was observed until the 3rd week of April. Some leks in areas of deep snow (Needle Creek, Houston, Sapinero 1,2, and 3) did not become active until the 1st week of May. Due to poor access, some leks (Chance, Wood Gulch. North Parlin) were not visited until May. Table 10. Number of individuals and date of peak sage grouse attendance at Gunnison Basin leks. 1984. Females Males lek Gold Basin Chance Gulch Woods Gu l ch Needle Creek South ParI in North Parli n Allen Ohio Creek Mashburn 1 Mashburn 2 Sapinero 1 Sapinero 2 Sapinero 3 Razor Creek Antelope Signal Peak Kezara a Sugar Creek Beavera A 1ka lla N 7 4 5 9 36 30 37 18 22 43 13 4 0 19 53 3 40 13 10 10 Date 19 Apr 11 May 12 May 12 May 28 Apr 21 May 12 May 07 May 07 May 11 May 12 May 12 May 01 May 28 Apr 16 May 09 May 08 May 10 May 10 May N Date 5 19 Apr 0 0 0 49 1 23 5 10 5 1 0 0 0 27 1 28 Apr 21 May 26 Apr 26 Apr 30 Apr 11 May 12 May 28 Apr 16 May aObserved during helicopter survey. Count is approximate and may include some females. Lek not verified through ground survey. �N N Capture Site N 6 Lek Site r .Q~. '0' I) , .:/ ~ ( ~~~ o Fig. 3. spr i n9 Movement of 3 radio-marked female sage grnuse breed ing sites, Gunn i son Bas in> Apr iii 984. between capture 5 sites on wintering 10km areas and fp*'. �~6 Captu re Site 6 Lek Site N .. r ~ £.1', , '" / p~, o I Fig. 4. Movement of 4 radio-marked male sage grouse spring breeding sites, Gunnison Basin, Apr! 1 1984. between capture 5, sites on wintering ", ..,. ~ 10km n areas and N W �N Conf irmed Lek """ Unconfirmed Lek Mashburn 1 . Alkali ~\ '\·Ohio Cr. Il!>\\, ••• .: N \JAllen \ / ~ •.. f '1," I ~\,. <j:e ntelooe (. \/ .oSignat.Peak \J Jld~ ( ON. h ~ . i -~ Sugar Cr. \ / .. \~ \:j,,\"\ \ ~ rig. 5. Sage grouse leks In the Gunnison /\rrii-hay i931f. Unconfirmed rot verified through ground leks were surveillance. during oRazor Cr. . . '" ~\ J 0Needle I~ j" (1It~ 9 Sasin where mnle and female attendance observed Parli~.'\ Oo.,ylQllille .~. \0., he l l c op t er • 1 ..rC" OWOOds Gul. ~ ( 0 So Parlin . Gold Basin~P °Chanc~ (\ Bekver Gut ) 1. ~ surveys on 8-10 ? 1pkm was evaluated, ~~a·.' but itiere Cr. �125 Mean male attendance in 198J-} was slightly lower, than in 1983 (Table 11). However, the difference was not significant (p > 0.05). Peak male attendance at 8 leks was much lower (> 25%) than during recent years. These primarily were leks where deep snow persisted through April. Peak female attendance data are not available for recent years; thus comparisons of female attendance are not possible. The peak of female attendance occurred during the last week of Apri 1 (Table 10). Table 11. Peak male sage grouse attendance at Gunnison 1980 lek Gold Basin Chance Gulch Woods Gulch Needle Creek Parl inb A I 1 en Ohio Creek Sapineroc Razor Creek Antelope Blue Mesa Upper S ix,-t>ll 1e McCabe Signal Peak Lost Canyon Mashburn 1 Mashburn 2 Ned 28 7 10 36 27 if3 72 16 46 n2 43 15 18 itS 6'2 0 0 2 _'-----------Cl NC b :::; 1980-84. J83_,_,~=- 11~81i: NC 38 8 1l.f 88 27 51 38 82. NC 39 NC 26.4 37.9 "j 47 52 NC 7 ~) 33.6 x 0 21 53 60 68 - Peak ma le attendance 19S2 __ -__ -.~- f§!I=-_~ 15 0 Basin leks. 7 4 5 9 66 37 NC 18 17 19 53 NC 10 5 NC NC 11 5 3 NC 22 43 34.0 23.3 NC 62 -,.~-------- No count. Includes Nor-th and South Parlin clncludes Sapinero leks. 1, 2, and 3 leks. Reduced male attendance at some leks probabJy does not indicate a population decline due to severe winter mortality. There was no evidence of excessive winter mortality. Carcasses of winter-killed sage grouse were not apparent in wintering areas. Also, survival of radio-marked individuals from late March through May was good. Only 3 of 22 radio'-marked individuals were killed during the period (Table 5). Attendance declines probably reflected abnormal display behavior, Ma'le displays were of low intensity and males often sat pas s lvel v on leks unless femal es were pres en t , By early May males typically abandoned leks prior to 0500 MST. Display activities had almost totally ceased by 23 May. �126 The erratic display observed in Spring 1984 was probably the result of poor, early spring body condition among males. Energy demands of d i s> playing sage grouse are potentially great. Males lose body weight during the display season due to metabolism of lipid reserves (Beck and Braun 1978; J. W. Hupp, unpubl. data). Because of severe winter conditions, sage grouse in the Gunnison Basin may not have been able to accumulate lipid reserves. Males proba~ly entered the display period with lighter than normal body weight and we r e unable to invest large amounts of enerqy in display. Normal spring body weights of Gunnison sage grouse are not known. Thus, comparison of 1981.f we iqht data with previous years was not possible. Weight data for North Park sage grouse are available (C. E. Braun, unpub l • data). Early spring (19 Mar-09 Apr) 1984 weights of adult male, North Park sage grouse 2.7611, N '"' 20) were lighter (p < 0.001) than during a similar period Tn 1983 (x',:;;- 3.2869, No.:: 18). No'rth Park also experienced severe winter weather--in 1984. Display of North Park sage grouse vias erratic and of low intensity. Peak attendance at many North Park leks was much l owe r than during previous years (C. E. Braun, per s . coomun.); a pattern similar to the Gunnison Basin. Male sage grouse in both the Gunnison Basin and North Park apparently entered the display season Ifli th Jowl ip id reserves. Th is probab 1y affected d isp 1d.' behav ior and could account for low lek counts. ex:= Lek Searches He1 icopter surveys for previously unlocated leks were c.onducted during 8-10 May between 0440 and 0700 MST. Male sage grouse flocks IfJere cb se rv ed in 4 areas If/here 'leks were not known to occur (Fig. 5). Males wer e observed displaying at 3 of these sites (Kezar , Willow, Beaver) prior to being flushed by the helicopter. Birds flushed at each site were counted (Table 11). Poor access and early termination of display prevented ground verification of these leks. Nesting Biology Ten sage grouse nests were located during the 1984 nesting season. Six were of radio-marked birds while 4 were discovered incidentally to other field research. Mean clutch size of 7 completed nests was 6.9 eggs and is comparable with previous sage grouse nesting studies (Patterson 1952, WaJlestad and Pyrah 1974, Petersen 1980). Clutch size of 3 yearling females ranged from 5 to 7 eggs while clutch size of adults varied from 6 to 9 eggs. Nine eggs were collected f r om 2 abandoned nests. Mean egg dimensions (54.0 x 36.2 mm) and weight (39.6 g) are slightly 'less than reported by Patterson (1952) and Petersen (1980). This may reflect the smaller body size of Gunnison Basin sage grouse. Eggs in 5 of 10 nests successfully hatched. Two yearling and 2 adult radio-marked females and 1 unmarked female of unknown age were successful. Three nests were destroyed by mammal ian predators. Pr'edat l on f o l l ow inq abandonment is possible. A r ed lo-rnarked yearling female abandoned her nest for unknown reasons. An unma rked f ema 1e abandoned her nest fo 11owl nq r escar> cher disturbance. Nesting success, not counting the research-induced abandonment, was S of 9 nests (567b). This is comparable to nesting success estimates obtained from wing molt progression of harvested females in the Gunnison Region in previous years (C. E. Braun, and J. W. Hupp. unpubl. rep , , Colo. Div. WildL, Fort Collins, 1983)" �127 Eggs in 4 active nests hatched between 15 and 18 June. In addition, 3 newly hatched broods were observed between 14 and 19 June, suggesting that these dates include the peak hatching period. This is similar to peak hatching periods in recent years (C. E. Braun, and J. W. Hupp, unpubl. rep., Colo. Div. Wildl., Fort Collins, 1983). Distance between breeding and nest sites for 5 radio-marked females ranged from 0.7 to 8.2 km (x = 4.2 km). Mean lek to nest distance was slightly greater than that observed by Wal1estad and Pyrah (1974) in Montana. and Petersen (1980) and Schoenberg (1982) in North Park, Colorado. LITERATURE CITED Beck, T. D. I. 1975. Attributes of a wintering population of sage grouse, North Park, Colorado. M.S. Thesis, Colo. State Univ., fort Collins. 49pp. , and C. E. Braun. ----.8<'"'='"0: 241-243. ___ 1978. Weights of Colorado sage grouse. Condor , R. B. Gill, and C. E. Braun. 1975. Sex and age determination of Colo. Div. Wildl., Garne lnf , sage grouse from wing characteristics. Leafl. 49 (revised). 4pp. Canfield, R. H. 1941. Application of the line interception method sampling range vegetation. J. For. 39:388-394. Emmons, S. R. Colorado. in 1980. Lek attendance of male sage grouse in North Park, M.S. Thesis, Colo. State Univ., Fort Collins. 69pp. Eng, R. L., and P. Schladweiler. 1972. Sage grouse winter movements and habitat use in central Montana. J. WildIe Manage. 36:141-146. Giesen, K. M., T. J. Schoenberg, and C. E. Braun. 1982. Methods for trapping sage grouse in Colorado. Wildl. Soc. Bull. 10:224-231. Klebenow, D. A. 1969. Sage grouse nesting and brood habitat J. Wildl. Manage. 33:649-662. Patterson, R. L. 1952. The sage grouse in Wyoming. Denver, Colo. 341pp. in Idaho. Sage Books, Inc., Petersen, B. E. 1980, Breeding and nesting ecology of female sage grouse in North Park, Colorado. M.S. Thesis, Colo. State Univ .• Fort Co lIins. 86pp. �128 Remington, T. E. 1983. Food selection, nutrition, and energy reserves .of sage grouse during winter, North Park, Colorado. M.S. Thesis, Colo. State Univ .• Fort Co llins. 89pp. Schoenberg, T. J. 1982. Sage grouse movements and habitat selection in North Park, Colorado. M.S. Thesis, Colo. State Univ .• Fort Collins. 86pp. Smith, L. M., J. W. Hupp, and J. T. Ratti. 1980. Reducing abandonment of nest-trapped gray partridge with methoxyflurane. J. Wild1. Manage. 44: 690,-691 • Tisdale, E. Wo, and M. Hironaka. 1981. The sagebrush-grass region: review of the ecological l Lte ra ture , Univ. Idaho, For ,, \AlildL. Range Exp. Stn. Bull. 33. 31pp. a and Wallestad, R. o. 1975. Life history and habitat requirements of sage grouse in central Montana. Montana Fish and Game Dep. Tech. Bull. 66ppo , and D. ----:-in centr~l Pyrah. 1974" Movement and nesting of sage q rou se hens Montana. J. Wildl. Manage. 38:630-633 . • and P. Sch ladwe i ler , '1974. Breeding ---selection of male sage grouse. J. Wildl. Prepared by Approved by season movements and habitat Manage. 38:634-637. �129 Colorado Division of Wildlife Wildl ife Research Report April 1984 JOB PROGRESS State of Colorado --------------------------- Project W-37-R-37 9 --~- Work Plan Popul~tions Personnel: Job: Game Bird Survey 7 and Habitat Preferences of Wintering .-------- of Slue Grouse Period Covered: Authors: (45-01-504-15050): Characteristics Job Title: REPORT 1 July 1983 through 30 June 1984 Brian S. Cade and Richard W. Hoffman C. E. Braun, D. J. Freddy, R. W. Hoffman, T. E. Remington, S. F. Steinert, Colorado Division of Wildlife; B. S. Cade, K. A. Medve, R. A. Ryder, Colorado State University_ ABSTRACT All data have been collected and analyzed in accordance with the objectives of this study. A draft thesis was prepared and reviewed. Revisions are currently being made. The revised thesis will be submitted to committee members for final approval. The approved thesis will be submitted as the final report next segment. A manuscript summarizing blue grouse migration patterns and habitat use during winter will be prepared for publication in a technical journal. ��i 31 CHARACTERISTICS OF WINTERING AND HABITAT PREFERENCES POPULATIONS OF BLUE GROUSE Brian S. Cade and Richard W. Hoffman P. N. OBJECTIVES The primary objectives of this study are to: (1) identify the vegetational and structural components of blue grouse winter use sites and (2) determine the spatial relationship between blue grouse wintering and breeding areas. SEGMENT OBJfCTIVES 1. Compile data, analyze results, prepare final report, 1 popular article, and 1 technical manuscript. METHODS /\ND MP.TER I ALS Methods have been descr i bed by Cade (1982). A de t a i 1ed descr i pt ion of methods will be included in the final report. RESULTS AND DISCUSSION The copious amounts of physiographic and vegetation data obtained in this study along with data related to movements between breeding and wintering areas have been campi 'led, summarized, and analyzed. Results were prepared in thesis format. Several drafts have been reviewed and the final draft is being revised for committee review. Once the thesis receives committee approval, technical manuscripts will be prepared. LITERATURE CITED Cade, B. S. 1982. Characteristics and habitat preferences of wintering populations of blue grouse. Job Prog. Rep., Colo. Div. WildJ. Fed. Aid Proj. W-37-R-36. Pp. 37-66. Prepared by ��133 Colorado Divi~ion of Wild1 ife Wildlife Research Report Apri 1 1984 JOB PROGRESS REPORT State of Colorado --------------------------- Project W-37-R-37 Work Plan _..;.._9 Job Title: Game Bird Survey (45-01-504-15050): Job: 8 Food Selection and Nutritional Ecology of Blue Grouse During Winter Period Covered: 1 July 1983 through 30 June 1984 Author: T. E. Remington Personnel: C. E. Braun, R. W. Hoffman, T. E. Remington, Colorado Division of Wildlife. T. J. Schoenberg, ABSTRACT Winter food habits and preferences of blue grouse (Dendragapus obscurus) l.-jere investigated in Middle Park, Colorado. R.adio-marked birds fed predom i nan t lv on needles of Douglas-fir (Pseudctsuga menziesii) or lodgepole pine (Pinus contorta). Limber pine (Pinus flexilis)was fed upon occasionally while feeding was not observed in Engelmann spruce (Picea Engelmannii) or subaipine fir (Abies lasiocarpa). Most feeding \~rthin Douglas-fir and lodgepole pine (90 and 98% of all ob~ervations, respectively) was in the upper or middle canopy and on 1-3 year-old needles (90 and 63%, respectively). Consumption of needles of both species decl ined progressively with needle age (p < 0.05). Preferences of capti~e birds for 5 conifer species \.-.Jere ranked as Douglas fir> lodgepole pine> limber pine> Engelmann spruce = subalpine fir. Captive birds ate more (p < 0.05) needles from branches from "o ld!' Douglasfir trees than from lIyoung" trees and more (P < 0.05) 1 and 2 year-old needles. than 3 and 4 or 5 and 6 year-old needles (Douglas-fir). ��135 FOOD SELECTION AND NUTRITIONAL ECOLOGY OF BLUE GROUSE DURING WINTER Thomas E, Remington The quality and quantity of winter food resources is generally recognized as a critical factor in the distribution and/or productivity of grouse populations (Gullion 1966; Miller et al. 1966, 1971; Moss 1969; Moss et aI. 1975; Watson et al. 1977; Watson and O'Hare 1979; Beckerton and Middleton 1982). Few reports exist of food habits of blue grouse during the critical Hinter" period (Beer 1943, Stewart 1944, Marshall 1946, Hoffmann 1961, Boag and Kianiuk 1968), Food preferences of blue grouse and the nutritional quality of their diets are unknown during winter. Data on winter food hab lts , food selection, and food quality of blue grouse are lacking from Colorado. Two current theories of herbivore food selection may pertain to blue grouse. Blue grouse may selectively feed to maximize nitrogen intake (Mattson 1980), or to minimize their intake of plant defense compounds which may inhibit their ability to digest foods or otherwise reduce fitness (Freeland and Janzen 1974. Bryant and Kuropat 1980). Blue grouse must contend with a winter diet that is both low in protein and high in fiber, and which also contains d iqest Iblllty+r educ inq compounds that may make significant smount s of nutrients unavailable. The overall objective of this study is to test the hypothesis that during winter, blue qr ou se selectively Feer' f rom arnonq ava Il ab le food i t erns to maximize digestible nitrogen and/or ener"gy. P. N. OBJECTIVES The objectives of this study are to investigate ("I)winter food habits, and (2) winter food preferences of blue grouse, and (3) measure the nutritional quality (protein, fat,minerals) and anti-quality components (tannins. terpenoids, phenolic resins, fiber) of blue grouse winter foods and their relationship to diet preferences. Specific objectives are to: 1. Identify winter foods of b lue grouse. 2. Investigate blue grouse winter use of conifers for food by species, tree ages, and growth forms in relation to their availability. 3, Quantify 4. Measure protein and fiber content of conifer needles from trees fed upon by blue qrou se and f rcrn r-andom lv-Toca t ed trees. tree species composition and physical blue grouse winter feeding sites. char ac ter i s t ics of �136 5. Measure tannin, phenolic resin, and mono-, di-. and sesquiterpene levels in conifer needles from trees fed upon by blue grouse and from randomlylocated trees. 6. Rank blue grouse preference for Douglas-fir, pine, limber pinL, and Engelmann subalpine fir. lodgepole spruce as winter foods. 7. Measure protein, fiber. dnd total digestibility r'andom ly+sel ec t ed needles suba 1pine fir, 8. of Douglas-fir. and Enqe Imann spruce. by b1ue grouse of le pine, limber pine, I odqepo Investigate the possible deterrent effects of specific tannins, phenolic resins, and mono-: , di"", and sesqu lterpenes to b rows Inq by blue qrou se and to blue grouse digestibility of conifer needles. SEGMENT OBJECTIVES 1. Capture and instrument up to 12 blue grouse. 2. Relocate instrumented birds for observation of feeding location of feeding sitej. Record feeding times, tree tree positions wlth In which feeding occurr ed . beh~vior and species USEd, and 3. Measure height and .ibh , snd co llec ': co+es det erm j nat Ion of each f ed+upon t ree .. 4. Collect need le samples from fed-upon trees for analysis of nutrient and monoterpene 1eve 1s . Count browsed and unbr owsed need 1as on browsed branches as an index to needle aSie preferences of blue grouse .. 5. Measure height and dbh , determine age, and collect needle 15 randomly-located tr ees of each of the 5 conifer species by increment borer for age s amp les from on Whiteley Peak. 6. Collect 2 grouse/week for chemica] analysis of crop contents and measurements of carcass fat levels. 7. Conduct trials to rank blue grouse preference for Doug1as-fir. lodgepole pine. limber pine, Engelmann spruce, and subalpine fir and for tree ages (old vs. young) and needle aqes (1-6) within Douglas-fir. DESCRIPTION OF STUDY AREAS ,.,.~-.•..... -.--.-. ..--.---.,~.--~-..- ..- .....• ~.-~-,;~.~ Whiteley Peak and adjacent Burn t Mountain approximately 30 kry.L-'d~.st of Kremml ing in Grand County, Co Ior ado , w re the pr lmarv study areas. In addition, 2 blue grouse originally r.-1diamarkerl on Green ~1ountain were ra Iocated periodically on Blue R~dge a~d above Guthrie Gulch for observations of feeding activity. �13/ Vegetative and physical characteristics of Whiteley Peak have been described (Cade 1983). Vegetative cover on Burnt Mountain consists primarily of homogenous associations of lodgepole pine/subalpine fir or quaking aspen (Populus tremuloides)/sagebrush (Artemisia spp.). Blue Ridge and Gut hri e Gulch are both dominated by 10dgepoTe--p(ne/subalpine fir associations. 150'lated pockets of mature Douglas-fir and limber pine are present along the top of Blue Ridge. .METHODS- Most observations of winter feeding activity were made by locating 8 r ad iomarked blue grouse ,nd observing them or other birds within that flock. Solar (Wi l d l l f e Materials, inc. ~ Carbondale, ! 11.) or 1it.hium battery-powered (Telonies. Inc.~ Mesa. Ariz.) transmitters were attached to vinyl ponchos which 51 ipped over the head of the birds (Amstrup 1980). \'.JE,dghts of the transmitters and harness were about 20 and 30 gms for solar and batterypowered units, respectively. Grouse were captured with a telescoping noose pole (Zwickel and Bendell 1967). Data collected at feeding sites included tree species fed upon, po_ition within t ree where feeding occu rred , and flock composition. Fed-upon trees were marked with oranqe flagging and numbered aluminum t.aq s , Tn':c dbh and heights were measured with a dbh tape and clinometer, respectively. Tree ages were es t lmated by counting annual growth rings from cores obtained by increment borer. Needle samples were collected from browsed areas within fed-upon trees. The age of needles eaten by blue grouse was determined by counting bud scales back from the terminal bud on each branch or branchlet that was fed upon. Preference trials were conducted with 3 wild-captured adult males and 1 juvenile female. The 1st preference trial (tree species) was preceded by an acclimation period of 8,6,7, and 5 days for birds 1-4, respectively. Birds were offered branches of each of the 5 conifer species tested during this acclimation per l od , Branches were changed daily. During the trial, pair wise combinations of Douq las+f i r , lodgepole pine, 1 Imber: pine, Engelmann ,. spruce, and subalpine fir were presented to the birds .. The order of treat·· ments (pairs of conifer species) was randomized; birds were then randomly assigned to treatment schedules. The randomization was r es t r i ct ed so that each treatment was replicated once in the morning (0700 to 1200 MST) and once in the afternoon (1315 to 18'15 MST). Each treatment was thus rep l jcated 8 times, twice with each bird. Branches of the 2 species tested were alternated and wired to 2 pegboards which were hung on the sides of the cage, Treatment response was the amount of each species consumed (gms) determined by weighing each branch before and after a replicate and subtracting sp lllaqe from the difference. Spillage was collected on aluminum trays that slid under the cages and separa:ed to species. Initially, 1,,000 ± 50 gr.c, of each species were cff'er ed to insure ad libitum quant lt les • Sy the 3rd day of the trial it was determined that 600 ± 509015 were adequate r or the morn iog rep 1 kate and 750 ± 50 qms v.... as adequate for the afternoon rep l i cat e . Con t ro ls were weighed during the first 2 days, but there wer e no detec.tab le we iqht losses due to de~;siCdtion. Branches used in the r.r ia ls vJf~n:' qa thered from the middle canopy of 5 r andom ly+l oc at.ed "matur e" t r ee-s of each species. �138 Tree age preference trials were conducted In a similar fashion to species trials. Branches col l ec ted from 3 "o Id" and 3 "younq" Douq l as fir trees were paired. Pairs Here collected from the same area to eliminate site variation. Young and old tree pairs were 27 and 137, '12 and 593. and 27 and 75 years old. Timing of the 2 daily replicates was altered to 0700 to 1445 and 1545 to 18.30 t1ST to reduce the d i spari ty in intakes between morning and afternoon replicates. preference Need 1e age preference t ri a l s were conduc ted by present i ng 3 con te i ner s containing ad 1 ibitum {lOO gms =: male, 75 gms :': female} quantities of '1 and 2, 3 and 4~ and 5 and 6 vear+ol d Douglas-fir needles to the birds twi ce daily .. Containers were attached to 1 side of the cage in a random array. Needles were removed from branches collected from m l d+canopy of randomly-located mature trees. RESULTS AND DISCUSS!ON One hundred and two f'eed l no observations of 8 rad l c-marked blue qrouse were recorded. Most feeding wa; in Douglas-fir or lodgepole pine (Ta~Ie 1). Limber pine was fed upon occasionally by 1 r-ad lo-mar ked bird, but several instances of unbanded birds feed I nq in limber pine wer e recorded. Significant feeding (> 30 sec) within Engelmann spr uce or subalpine fir was not observed. Blue grouse use of DougJas-fir has been previously documented (Beer 1943, St ewar t 19l!4, Marshall 19lf6). Feeding on lodgepole pine has been documented by Boag and Kicenluk (1968). Table 1. Tree species f~d upon (number of observatlon~ blue grouse, Middle Park, Colorado, November 1983-April by radio-marked 1984. -----------Psme 9 6 18 0 0 8 'j 0 10 58 Pico 0 0 0 0 16 4 5 13 0 0 0 0 0 0 3 0 0 0 0 0 0 0 1 38 2 4 9 6 0 2 0 20 16 4 11 6 ·13 11 102 Pifl Psme/Pico Totals b o ---------- •.----~.-------------.----,-.-----..--.,---- ....._----_ .._--aPsme = eouglas-fir, bFed upon both species Pico = during lodgepole 1 feeding pine, Pifl = limber pine. period. Ninety and 98% of the birds observed feeding wi th in DougJas~i_i_~ __and l odqepole p l ne , respectively, were in the upper or middle canopy {Table 2). Boag and Kiceniuk (1968) noted that spruce grouse (Dendraqapus canadensis) and/or blue qr'ousc concentrate br'ows l nq within mid···canopy'of-TodgepoTe---pine. Hoffmann (1961) found t ha t blue grouse fed ex clu s l vel v within the upper port ion of wh i te fir (!~~i.~:_~. ~,~9_~~!:) .. �13~ Table 2. Tree positions Middle Park. Colorado, Lo,wer._·j[L_._., _S..•.. p_e_c_i_e_s_a Psme Pica a Psme = (number of observations) November 1983-April 1984. Douglas-fir, of feeding Mid~...!.e 1/3 _ blue grouse 1/3 Upper _ 11 47 51 1 22 23 Pico = lodgepole In pine. No differences in feeding behavior were apparent among sex and age-classes of blue grouse. Birds of mixed sex and age classes were frequently observed within the same tree. Structural characteristics of fed-upon trees varied within and among stands, but birds usually fed in the largest trees available (TabJe 3). Ages of fed-upon Doug las-f i r trees ranged from 55 to 597 years. OnIy 6 of 33 (18%) fed-upon Doug I as+f irs for wh i ch ages were ob ta ined were l es s than 100 years old. Most of these wer e in a young, mixed conifer stand on the north side of Whiteley Peak where older trees were absent. Ages of fed-upon lodgepole pine ranged from 62 to 'I'll years. Ellison (1976) noted that spruce grouse avoided feeding in saplings of white spruce (Pj cea q l auca}, Boag and IUcenluk (1968) observed that spruce and/or' b lue grou's'efel-alrnost exclusively within "older" (> 15 years) trees, Table Middle 3. Height (m). dbh (cm), and age of trees fed Park, Col orado , November '1983-Apri 1 ·j981f. Desc r ip t ive statistic x SD Range N --=-~: Douglas-fir ··1ft-----Obh Ag~ __ upon by blue grouse, lodgepole pine Ht . Dbh__ ~_~ge-=- 15.9 50.9 219 15.2 28.0 78 5.2 19.5 129 3·2 8.2. 15 9-31 21-90 5-20 9··/.f'9 28 28 36 36 55-597 33 27-111 26 -----.-- ..---.--.-------~---.-----. ----'-'--- ... Blue grouse use and selection of needles by age-class were investigated by counting browsed and unbrows.d Douq l as-f Lr and lodgepole pine needles from 1 to 6 years old. Needle age-classes were defined as to 1 to 6 years for simplicity. Current-year qr owth needles described as "1" year+o Id need l es were from 5 to 10 months old over the period of study; "2" year+o l d needles were 17 to 22 months, etc. Six years was chosen as the cutoff point because needles of both these species are generally per s is tent to 6 ve ars (Harl ow and Harrar 1969), and no browsing was detected on older needles .. One thousand ne edl e s were counted from each fed-upon tree. �140 The percentage of needles of each age-class browsed by blue grouse was used to test for selection. Blue grouse selectively fed among age-classes of both Douq I as-f i rand 1odgepo 1e pine (Hote 11 ings r2. P < 0.001)" Consumption of needles of both species declined progressiveTy with needle age. This was most apparent within Douql as+f lr (Table 4). Each age-class "lithin Douglas-fir was preferred (higher percentage browsed) to those o"lder except needle ages 5 and 6 were equally non-preferred (pairwise t tests, P < O.Os). Within lodgepole pine, 1 and 2 vear+o ld needles were pn:::fe'rred(p <' 0.05) to ages 4=6; 3 year-old needles were preferred to ages 5-6; and 5 year-old needles were preferred to 6 year-old needles (Table 4). a Percentage of to 6 year-old needles of Douglas-fir (N = 18) and pine (~,= 22) browsed by blue grouse, November 1983-Ap"ri I 1984. .._--_. ._. Lodqepole pine Doug] as -f i r .--.----Needi,~age·---------Needle age -1 --2--'3··-'~---5----T 2 3 <t 5 0 Table 4. lodgepole __ -------------_ -'--.--'..---.----r- x 61 25 12 5 2 66 60 56 SD 10 17 l2 8 5 19 13 13 1,000 aN = number of trees for which needles were counted. needles were. counted/tree. 50 15 33 19 Approximately Ell ison (1976) found that during winter spruce grouse browsed aqe+c las ses of white spruce needJes in proportion to their availability; no selection was evident. One vear+o ld needles of lodgepole pine we re the aqe-c lass "mos t heavily used!' by spruce grouse and/or blue grouse in Alberta (Boag and Kiceniuk 1968:28). The net effect of this selection is that 1-3 year-old needles comprise the bulk of needles consumed from both Douglas-fir (90%) and lodgepole pine (63%) (Table 5). These estimates are minimal since usually only branches containing needles of all (1-6) age-classes were selected for counting. Thus, use of 1-3 year-old needles on branches less than 6 years old wouldnlt have been counted. Table 5. Percentage of 1 to 6 vear+o ld needles browsed by blue grouse from Douglas-fir (Na = 19) and lodgepole pine (Na = 22) in relation to total needles brows'ed, November 1983-April 1984.- x 44 32 SD 17 10 1,000 7 6 3 6 It 2 20 24 19 18 12 8 9 6 7 6 aN ~ number of trees for which n0edles were counted. needles were counted/tree. Approximately 7 �141 Douglas-fir was preferred (p < 0.1) over both subalpine fir and Engelmann spruce and lodgepole pine was preferred over Engelmann spruce. A defendable ranking of species preferences is Douglas-fir> lodgepole pine> limber pine> Engelmann spruce = subalpine fir. There seemed to be no reaJ difference in preference between the 2 least preferred species, sUbalpine fir and Engelmann spruce. Except for this pair, all species in the above ranking were apparently preferred to those ranked below it. Since all 5 species were offered to all 4 birds, once in a morning trial and once in an afternoon trial, the total amount of each species consumed can be used as a check of this ranking. The amount consumed was 2,492,2,350, 1,493,927. and 809 grams of Douglas-fir, lodgepole pine, limber pine, Engelmann spruce, and subalpine fir, respectively. It is interesting to note that this ranking compares well to what radiomarked birds fed upon in the wild. All radio-marked birds subsisted pr irnarlly on the 2 most preferred species, occasionally feeding on the 3rd ranked species and avoiding the 2 least preferred species entirely. Total intakes of each treatment pair were similar for all pairs except for subalpine fir/ Enge1mann spruce, which was markedly lower than a·11 others. If blue grouse are selectively feeding among species to maximize nitrogen intake, the least preferred species should be lower in nitrogen. It is reasonab"le to expect intakes to rise to maintain nitrogen intake if the cholce is 2 non-preferred species. If,on the other hand. blue grouse are selectively avoiding plant defense compounds. then non-preferred species should be relatively high in defense compounds and intakes should decline when food choices consist of 2 non-preferred species. The data from this trial support the latter theory. Selection for older trees (or avoidance of younger trees) was tested by offering birds equal, ad libitum amounts of needles from "old" or "young" trees by a 1 ternat ing branches from each w i th j n cages. Preference for older trees was confirmed (p < 0.05) (Table 7). Needles from "o ld!' trees were consumed in greater quantities in 21 of 24 replicates. Table 7. Mean intake (gm) and difference in intake by blue grouse of needles from young and old Douglas-fir trees (~= 24). Old Young 61.0 22.8 17.5 18.8 ------------------------------------------~----------------x - so SOb as' Ignlrlcant ·co at b SD = SO / /"N. p < -- 0 •• 05 Difference 38.2a 33.6 6.8 �142 Apparent selection for certain species and age-classes of conifers and for needles of certain ages (Tables 1, 3.. 4) was tested in trials using captive birds. By presenting ad libitum quantities of each treatment to captive birds, problems of measuring and interpreting use and availability in the field were eliminated. Selection for certain species, ages or growth forms of trees on the basis of perching ability, thermal cover or other factors related to feeding efficiency rather than chemical properties of needles was eliminated in captivity. Blue grouse responded well to captrvity and the experimental designs used. Differences of biological significance were detected in 9 of 10 comparisons of conifer species (Table 6). Because of the small number of birds tested (with correspondingly low degrees of freedom) and multiplicity of t'tests (requiring a probability of error of 0.01 to test significance at '0.1), statistical significance (p < 0.1) was achieved in only 3 of 10 comparisons. Species preference trials will be repeated next year with 4 more birds. The additional degrees of freedom and reduced variability due to "larger sample sizes should (if preferences remain similar) make 9 of 10 comparisons statistically significant. Table 6. Hean total consumption and mean difference in consumption grouse of needles from conifer species offered in pairs. blue by -"-Pai r Total consumed (gm) - b Difference SD-C x x Psme/Alba Psme/Pien Psme/Pifl Psme/Pico Pico/Abla Pico/Pien Pico/P ifl Pien/Abla Pien/Pif] Abla/Pifl between pair members (grn) -il 100 106 102 105 102 d 118 10 lOOd 15 "'4 18 17 15 67 41 104 90 93d 104 88 74 3 23 104 -50 .•46 23 101 aCorrected for morning, afternoon differential ing intake/afternoon intake]). Pifl bPsme ~ Douglas-fir, Abla = subalpine fir, Pien limber pine, Pico = lodgepole pine. = cSD "" "SIT" if ~ s ~_ :::: 10 11 (morning intake -;;[morn- = Engelmann spruce, 8. dDifference between pairs significant to adjust for mUltiple [10J t tests). at P < 0.1 (tested at f < 0.01 �143 Selection may have been even greater than indicated. In all 3 replicates where consumption of needles from young tre s exceeded consumption of needles from older trees, browsing was concentrated on terminal or lateral leaders of branches (from young trees) exhibiting extremely rapid growth. It may be that grouse are selecting needles at a finer level of resolution than just tree age. It is unclear how the captive blue grouse identified branches to feed upon. There were subtle differences in needle color and d nsity between branches from young and old trees that may have served as cues. At this time it is also unclear why blue grouse prefer feeding on needles from older trees. They may be selectively feeding to avoid monoterpenes. Needles from young and old scotch pine (Pinus sylvestris) differ markedly in relative concentrations of monoterpenes (Htivnak et al. 1973). While it is apparent that blue grouse selectively feed on younger needles, especially within Douglas-fir (Tables 4 and 5), it is not clear whether this reflects preference. For instance. it's possible that blue grouse move to the tips of branches to eat buds, and then feed on needles in the vicinity of the twig tip. Buds of Douglas-fir contain substantially lower levels of monoterpenes than needles (Von Rudloff 1971). It is also unclear hOYI blue grouse identify palatable needles to feed upon. Preference for needle ages was tested by randomly ordering 3 containers containing ad libitum quan t lt Ies of 1 and 2, 3 and 4, and 5 and 6 year-old needles within each cage. Blue grouse strongly preferred (F = 450, 2 and 23 df, P < 0.001) 1 and 2 year-old needles (Table 8). Table 8. Mean intake (gm) of 1 and 2, 3 and 4, and Sand Gyear-o'ld Douq las+f i r needles during 6 trials with 4 captive blue grouse (!= 24). March 1984. Statist ic -'j ~~~~--------------. x so Needle aoe "and Ii and2----··----:3 72.5 6.9 3.1 a 3.0 aMeans sharing the same letter do not differ (p > 0.05, Duncan Multiple Range Test), There was no difference (p > 0.05) in intake between 3 and 4 and 5 and 6 year-old needles. It is iignificant not only that blue grouse preferred 1 and 2 year-old needles but also that they were able to identify them ;-Jhen they were removed from the branches. There we re no discerni_!:?li:_._differences in needle size, color, or texture. This is a strong indication that blue grouse use chemical cues to identify palatable needles. �144 The reason for selection of younger needles is unknown at this time. Lowry (1970) and Ellison (1976) reported that nitrogen content of black (Picea mariana) and white spruce dec] ined with needle age. Thus, selectionf'O-r:younger needles may be selection for nitrogen. Sixteen grouse were collected. Organ lengths and weights. and carcass compos i t ion (% wat er , fat) wi 11 be measured. A 11 (75) random trees have been measured and sampled. Needle sempl es from randomly-located and fed-upon trees have been prepared for chemical analysis. Techniques for extracting, quantifying, and identifying monoterpenes have been developed. Most samples have been run but results have not been analyzed. Nitrogen content of needle samples is currently being measured. LI TERATURE Amstrup, S. C. 1980. 44: 21 4- 2.17. A radio-collar C I TED for game birds. Beckerton, P. R., and A. L. A. Midd1eton. levels on ruffed grouse reproduction. Beer, J. R. 1943. 1982. J. Wildt. Manage. Effects of dietary protein Manage. 46:569-579. J. Wildl. Food habits of the blue grouse. J. Wild1. Manage. 7:32-1.,4. Boag, D. A., and J. W. Kiceniuk. 1968. Protein and caloric content of lodgepole pine needles. For. ehron. 44:28-31. Bryant, J. P .• and P. J. Kuropat. 1980. Selection of winter forage by subarctic brows inq vertebrates: the role of plant chemistry. Annu. Rev. Ecol. and Syst. 11:261-286. Cade, B. S. 1983. Characteristics and habitat preferences of wintering populations of blue grouse. Job Prog. Rep~, Colo. Div. Wildl. Fed. Aid Proj. W-37-R-35. April. Pp.37-66. Ellison, L. 1976. Winter food selection by Alaskan spruce grouse. Manage. 40:205-213. J. Wi·ldl. Freeland. W. J., and D. H. Janzen. 1974. Strategies In herbivory by mammals: the role of plant secondary compounds. Am. Nat. 108:269-289. Gullion, G. W. 1966. A viewpoint concerning the significance of studies of game bird food habits. Condor 68:372-376. Harlow, W. M •• and E. S. Har rar , 1969. Textbook of dendrology. McGraw-Hlll Book Co., New York, N.Y. 512pp. 5th ed . Hoffmann, f{. S. 1961. The qual itv of the winter food of blue grouse. Manage. 25:209-210. .J. Wi ld l, �145 1973. Analysis of and Pite~ ~~ls~. und Ho l zverwer tunq Hrivnak, J.~ M. Mahdalik, E. Varadiova9 and L. Sojak. monoterpenes from the needles of _Pinus ~~tris using capillary gas chromatography. Holzforschung 25:24-26. Lowry, G. L. 1970. Variations in nutrients of black spruce Pages 235-259 in Co T. Younqber q and C. B. Davey, eds . and forest soils', Oregon St.ate Univ. Press, Corvalis. Marshall. W. H. 1946. of Richardsonls seasonal Cover" preferences, grouse and ruffed grouse needles. Tree growth movements in southern and food Idaho. habits Wilson Bull. 58:42-52. Mattson, W. J •• Jr. 1980. Herbivory in relation Annu. Rev. Eco!. and Syst. 11 :119-161. Miller, nitrogen content. G. R .• D. Jenkins, and A. Watson. 1966. Heather performance and red grouse populations. I. Vi~ual estimates of heather performance. J. Appl. Eco!. 3:313'-326. , A. Watson, and D. Jenkins. tions to experImen 10:323-335. Moss, to plant ta l improvement 1971. Responses of red grouse populaof their food. Symp . Sr. Eco l, Soc. R. 1969. A comparison of red grouse (Lagopus laqopus sco t l cus ) stocks with the production and nutritive value of'··lleathe-r'-rC"aTfu·;;a-·vi:iTgi:H-is). J. Anim. Ecol , 38:103-112. ------.----.- _____ • A. Watson, and R. Parr. in red grouse cess Stewart. R. E. 1975. nutrition and breeding suc- Maternal (_~..::!.gopu.?_ lagopus.._scot!~~). J. Anim. Food habits of the blue grouse. 19L14. Ecol" 44:233-'244. Condor 46:112-120. Von Rudloff, E. 1971. Chemosystematic studies in the genus Ps eudo t suqa , Leaf oil analysis of the coastal and Rocky Mountain vari"eties----of th-e Douglas-fir, Can. J. Bot. 50:1025-1040. Watson, A., and P. J. O'Hare. treated and untreated Red grouse 1979. bog. Irish J. Appl. I. populations on experimentally Eco!. 16:433-452. , R. ---on red Moss, J. Phillips, and R. Parr. 1977. The effect of fertilizers grouse stocks on Scottish moors grazed by sheep, cattle and deer. Pates 193-212 in P. Pes son , compiler. Ecologie du petit gibier et ameneqement des chas ses . Gauth j er-N i 11 ar s , Par" is France. 9 Zwicke l , F. C., and J. F,. Bende l l • J. Wildl. Manage. Prepared Approved by ~ <:. y,::_~ . Thfj'S). R1i ng by tOI ~_/Mt{_I1{4/ R I chard Wildlife '1967. 31:202-204. lfV/.:!:~~_::::':"__ W. Hoffmy;; Researcher , C A snare for captu.rlilll._blue grouse. ��Colorado Division of Wild1 ife Wildlife Research Report April 1984 147 JOB PROGRESS REPORT State of Colorado --------------------------- Project W-37-R-37 Work Plan 12 Job Title: (45-01-504-15050): Job: Game Bird Survey 15a History and Status of Rio Grande Wild Turkey Transplants Along the South Platte River, Colorado Period Covered: ! January - 30 April 1984 Author: Steven Rosenstock Personnel: Larry Budde, Mike Creamer, Larry Crooks, Ken Dillinger, Marv Gardner, Rick Hoffman, and Tom Kroening, Colorado Division of Wild] ife; Steve Rosenstock, Colorado State University. ABSTRACT The history and status of wild turkey transplants along the South Platte River in northeastern Colorado were investigated from January through April 1984. Sixty wild-trapped Rio Grande turkeys (Meleagris gal Jopavo intermedia) from Kansas and Texas were released at 4 locations along the South Platte River between 1981 and 1983. Turkeys have become established at or near all 4 release sites and between 120 and 150 birds were counted during spring 1984. The 1980 Hil lrose transplant was most successful. Birds from this release have shown significant population growth and have dispersed to occupy approximately 60 km of riverbottom habitat. The 1980 Tamarack release resulted in a small population that has shown negligible growth and dispersal. Turkeys from the 1983 Masters release appear to be doing well. Turkeys along the South Platte are primarily confined to the 0.25-1.5 km-wide riparian zone adjacent to the river. Average home range of 9 flocks located during winter 1983-84 approximated 1.6 km2. Foraging activities were most commonly observed in harvested cornfields, livestock feeding areaSj and hayfields. Supplemental feeding by landowners was a common practice. Turkeys used dense woody and herbaceous vegetation in the riverbottom as loafing and escape cover. No roosts were located. Nests (1983) were reported in an alfalfa field, a fencerow, and an upland pasture. All 1983 brood sightings were in or immediately adjacent to the riverbottom. Potential limiting factors on the South Platte turkey population include: riparian flooding, nest destruction by mowing operations and predators, inbreeding, climatic conditions, and poaching. ��149 HISTORY AND STATUS OF RIO GRANDE WILD TURKEY TRANSPLANTS ALONG THE SOUTH PLATTE RIVER, COLORADO Steven S. Rosenstock Wild turkeys are becoming an increasingly popular game bird in Colorado. To increase future hunting opportunities for this species, the Colorado Division of Wildlife has made several releases of Rio Grande turkeys along the South Platte River in northeastern Colorado. Rio Grandes have b~en successfully introduced into riparian habitats in other states including Idaho, Kansas, and Nebraska. This repor t contains information obtained from January through April 1984 evaluating the results of turkey releases along the South Platte River. P. N. OBJECTIVES 1. Document the history of Rio Grande wild turkey releases in the South Platte River drainage. Obtain the fol Iowl nq information: a. b. c. d. e. 2. dates of releases. locations of releases, number of birds released at each site, sex and age composition of individual releases, and origin of transplant stock for each release. Examine the current distribution South Platte as of January-April of Rio Grande turkeys along the 1984. 3. Obtain estimates of current population size(s). 4. Identify habitat types/components !oafing. 5. Identify problems for future study, and if possible. hypotheses. used for foraging, roosting, and propose testable SEGMENT OBJECTIVES 1. Review Colorado Division of Wildlife records of Rio Grande wild turkey transplants along the South Platte River. 2. Contact Division employees South Platte releases. and other personnel who participated In the �150 3. Contact District Wildlife Managers and landowners along the South Platte River to help obtain current population, distribution, and habitat use information. 4. Conduct aerial surveys along the South Platte River to locate turkey flocks and roost sites. 5. Conduct ground surveys of riparian habitats along the South Platte River known or suspected to be occupied by Rio Grande tur keys . Identify habitat components used for foraging. loafing, and roosting. 6. Contact researchers Grande and managers in other states working turkeys and summarize ava llab le data. with Rio 7. Conduct ongoing review of literature pertaining to the RIO Grande wild turkey, including: habitat use, movements, dispersal, food habits, reproduction, limiting and mortality factors. effects of harvest, effects of Iand-manaqemerrt pract ices. trapp ing and trans" planting. radio-telemetric obcervation, other research/study techniques. 8. Compi l e data, analyze r esult s and prepare a final report to be submitted to the CDOW and the Department of Fishery and Wildlife Biology, Colorado State Un iver s i tv in fu lf lllrnent of special study (FW 1+95) requ lrement s . DESCRIPTION OF STUDY AREA The study was conducted along a 440-km section of the South Platte River in northeastern Colorado (Fig. 1). The dominant plant community along the river is plains cottonwood (£~,'<.?.f>u'lus._~_argentii), which cover's 56% of the naturally vegetated f Ioodp la i n [Lindauer 'T§8:3). Dense stands of narrow-leaf willow (Salix ln t er lor ) occur along watercourse banks and cattail (Typha spp.) is common in marshes, oxbow ponds, and sloughs. Drier sites support mixtures of shrubs and herbaceous vegetation. The riparian zone varies in width from 0.25 to > 1 km. Most land adjacent to the riverbottom is in production of corn, wheat, sorghums, alfalfa, and grass hay. Uncultivated lands are used as pasture for livestock. The riverbottom is also heavily grazed on private lands. METHODS Meetings were held with CDOW research and management personnel the status of Rio Grande turkey introductions along the South and to identify information needs. Management personnel were and CDOW records were reviewed to obtain specific information turkey releases conducted to date" The following information release was collected: date, location, numbers of birds, sex composition, and orIq In of transplant stock. to discuss Platte River contacted on Rio Grande on each and age �151 From January to April 1984 District Wildlife Managers and landowner s were contacted in person and by telephone to help locate turkeys. Reported sightings were confirmed by ground surveys to collect additional distribution, population, and habitat use information. An aerial survey by fixed-wing aircraft was conducted on 6 April 1984 in an effort to locate additional turkey flocks and obtain an overview of riparian habitat along the river. Hunter surveys from the 1983 limited deer season in Big Game Unit 89. 92, and 881 were reviewed. These surveys requested deer hunters to report any observations of wild turkeys. Research and management personnel in Idaho, Kansas, and Nebraska were contacted to obtain information on Rio Grande turkey populations introduced into similar habitats. A review of literature on the Rio Grande turkey was conducted. Emphasis was placed on: habita~ use, movements, dispersal, reproduction, limiting and mortality factors, effects of land-management practices, and techniques of capturing, marking, and instrumenting turkeys. A landowner report card was prepared and will be distributed by CDOW personnel in an effort to collect ongoing information on Rio Grande turkeys along the South Platte. RESULTS AND DISCUSSION Since 1980, the CDOW has released 60 Rio Grande turkeys (Table 1) at In January 1980, 21 turkeys from Kansas \'Jerereleased northeast of Hillrose. On 19 February 1982 20 birds from Texas were released on the east end of the CDO\-l'sTamarack Ranch property. Also on 19 February, 3 birds from the same group \'Jere released at Balzac Bridge east of Hillrose to supplement the 1980 release. On 12 January 1983, 16 birds fran Kansas were released on the Yocam Ranch near Masters. 4 sites along the South Platte River (Fig. 2). Table 1. Age and sex composition of Rio Grande turkey transplants on the South Platte River, Colorado. Toms Date Locat ion Source 1/80 2/82 2/82 1/83 Hi 1 j rose Tamarack Ranch Balzac Bridge Masters Kansas Texas Texas Kansas Totals -"--------,----_. ad. unk. ad. juv. unk. Total 6 9 5 21 20 3 3 6 7 3 16 15 12 10 0 --_. juv. Hens --------- 12 10 11 11 60 �152 The site of these releases is similar to those made in riparian habitats in Kansas and Nebraska in the 1960ls (Suetsugo and Menzel 1963; Capel 1967, 1968a). There is considerable debate over the optimum transplant size. Some-biologists and managers recommend releases as small as 4 birds, in a ratio of 1 tom:4 hens. Others favor releases of up to 50 birds (R. Little, Area Supervisor, Kans. Fish and Game Comm., pers. commun.). Distribution and Numbers Nine winter flocks totaling 110 birds were located (Fig. 3). Flocks ranged from 3 to 27 individuals and included both sexes (Table 2). Composition and location of the 9 flocks remained relatively constant during the observation period. In their native r-ange, Rio Grande turkeys usually form sex-segregated wl nter groups (Thomas et a1. 1966, Bailey and Rinell 1967, Logan i970). However introduced populations in Kansas and Nebraska do form mixed-sex winter flocks (Capel 1967, 1968a; K. Menzel, Biologist, Nebr. Game, Fish and Parks Comm., pers. commun.T. Average winter flock size in Kansas ranges from 10 to 30 birds although aggregations of up to 100 birds have been observed (R..Little, pers. commun.) .. Table 2. Composition of Rio Grande turkey flocks along the South Platte River, Colo. January-April 1984. Flock A B C D E F G H I Location --.------Yocam Ranch, NE of Masters 5 km W of Yocam Ranch Goodr ich ! km W of Snyder Peterson Ranch, N of Hi 1 J rose Balzac Bridge, E of Hill rose Messex Merino Tamarack Ranch Toms 2 Hens -------~.-~.-...-.Total ---~--.~ 1 4 5 2. 2 20 27 3 3 3 7 18 20·-25 15 Total 'V 110 The stretch of river between Snyder and Merino supports the largest number of turkeys. Five flocks, totalling approximately 76 birds were located in this area. These birds originated from the 1980 Hillrose release. To the we s t , between Masters and Weldona, 3 sma lI flocks totalling 14 birds were found. These are apparent survivors and/or offspring from the 1983 release at Masters. A population of approximately 20-25 birds is present on the Tamarack Ranch Property, site of the 1982 release. Numerous other sightings (Fig. 4) were unconfirmed due to the observer's inability to locate birds to verify the sightings or because time constraints prohibited follow-up surveys. It is possible some flocks within the study area went undetected. Ground searches along the riverbottom we re time-consuming and general iy unsuccessful. All flocks located �153 in this study resulted from reports by landowners and area CDOW personnel. Wild turkeys are of prime interest in this area, and landowners are well informed of their whereabouts. The aerial survey did not result in the location of any new flocks. To ascertain if turkeys could be spotted from a fixed-wing aircraft, considerable time was spent attempting to find known flocks. Even with moderate snow cover, this proved difficult. Only 1 known flock was located. Capel (1967) noted that turkeys will freeze when approached by a fixed-wing aircraft. Future attempts at aerial surveys should be from a helicopter when 100% snow cover is present. Turkey sightings were reported by respondents to a survey of participants in the 1983 limited deer season in Big Game Management Units 89, 92. and 881 as shown in Figure 5. These sightings (Table 3) correlate wen with the observatIons of this study. The greatest number of turkeys was seen by hunters in Unit 92, which extends along the South Platte from Fort Morgan to Atwood . Within Unit 92, most turkeys were observed between Snyder and Merino. Table 3. Turkey sightings reported by hunters participating in 1983 Deer Season, Big Game Units 89,92, and 881 (T. Davis, CDOW unpub l, data). ._-_._-_._--------_. Hunters Unit ---Turkey sightings N -----_._------------_. N -------- .. •...-----~-- Turkeys observed N __ .------ ---------- 32 89 92 106 34 881 68 6 aThis figure includes multiple sightings of the same birds. Separation of probable duplications yields an estimated total of 75-175 birds. Dispersal from Release Sites Landowner observations suggest that turkeys transplanted in January or February tend to stay together near the release area through the remainder of their first winter. Dispersal usually occurs the following spring. Following the 1983 ~1asters release, 7 birds moved from 3 to 16 km downriver within 8 months of their release. No specific dispersal information was obtained for the other releases. �154 Rio Grande turkeys released in Kansas have initially stayed close to the release site. Capel (1967) found that most birds stayed within 0.8 km, movements of other birds ranged from 1.6 to 24 km. He sugges ted that post-release movements are effected by environmental factor's and human activities in the area. Rio Grandes released in Nebraska showed similar behavior. Some remained in the vicinity of the release site, others moved up to 40 km (Robertson 1963, Suetsuga and Menzel 1963). £\anq~ Expansion Rio Grande turkey populations have been established from the 1980 Hi llrose release and 1982 Tamarack release. Birds from the 1983 Masters release appear' to be doing we ll, however it is too early to evaluate the success of this transplant. The Hillrose release has shown the greatest success. Birds from this transplant and their offspring have dispersed appr'oxl> mately 20 km upr lver and 40 km downriver from the release site. The minimum population in this area is at least 76 birds. The Tamarack population appears static. The total population is estimated at only 20-25 birds, which have not dispersed from the release site. Varying success has been achieved by other states that have introduced Rio Grande turkeys into riparian habitats. Kansas had excellent initial success wl th transp Iants of Ok Iahoma birds in the Iate 1960 I 5" Reproduction and population expansion occurred at nearly all release sites. Population increases of up to 250% were documented after the first breeding season (Capel 1967, 1968a). These populations did well for' 4-5 years, then began to decline. R.-'-Uttle (pers. commun.) speculated that the decline was precipitated by inbreeding resulting from the small size of initial transplants. Lange (1983) emphasized the importance of genetic diversity in turkey transplants. Larger, more recent transplants, using birds from Oklahoma have continued to show consistent increases (up to 100% annually) and range expansion 4-5 years post release (R. Little, pers. commun.). Nebraska experienced similar success. Transplants during the early 1960ls showed healthy initial reproduction and growth (Robertson 1963). However, after 4-5 years, the populations declined (K. Menzel, pers. commun.). Only 2 of 25 introductions persisted. As a result, the Rio Grande turkey transplant program was terminated. It should be noted that Nebraska made mostly small transplants of 10-15 birds using Texas stock. K. Menzel (pers. commun.) attributed the decline to habitat limitations and/or climatic factors. Recent introductions in Idaho have met with mixed success (F. Bizeau and R. Norrell, pers. cornmun.). Initial results of Rio Grande turkey introductions in Colorado appear similar to those reported from other states, although not enough time has elapsed to observe a post-establishment population decline. Growth �155 and expansion of the Hillrose release suggests that habitat conditiLns along the South Platte are suitable for Rio Grande turkeys. However, the rate of population increase has not been as high as expected when compared to introductions in Kansas. There are a number of potential limiting factors; the most likely being a shortage of undisturbed nesting habitat. For the past 3 years, much of the riverbottom and adjacent riparian zone has been inundated by high water during the nesting season. Minimal cover exists outside of the riparian zone due to intensive farming. Five instances were documented in 1983 where turkeys nested outside the riverbottom. All 5 nests f a iled . Three nests in an alfalfa field were destroyed by mowing and 2 nests in an adjacent fencerow were destroyed by predators. Flooding has been linked to turkey population declines in Kansas (Peabody 1963). Nesting Rio Grande turkeys in Kansas show a strong affinity for alfalfa fieids, even in the absence of riparian flooding (R. Little, pers. commun.). Mowing has caused significant nest destruction. Nest mortality from alfalfa mowing has also been observed in Nebraska (Suetsuga and Menzel 1963; Robertson 1963. 1964; K. Menzel, pers. commun.). A lack of secure nesting cover may also be increasing the incidence of nest predation. Striped skunks (Mephitis mephitis) and raccoons (Procyon lo tor ) are abundant in the riverb-ottom and adjacent areas. Both are--cZ'm':'s'i"cfer-ed major predators on wild turkey nests (Narkley '1967, Baker 19l9). High rates of nest predation have been documented for Rio Grande turkeys introduced into riparian habitats in Idaho (Ogden 1983). One biologist contacted felt that the characteristic narrow width of riparian habitats plus high predator densities will prevent long-term establ ishment of wild turkeys (F. Samson, Colo. Coop. Wild1. Res. Unit, pers. commun.). Preda·· tion on adult turkeys is not considered a major limiting factor (Markley 1967). However, it may be important if heavy predation losses are incurred by a recent small transplant. Many landowners and CDOW management personnel have expressed concern that poachers are taking large numbers of turkeys along the South Platte. To date, only 2 incidents have been documented (L. Budde, pers. commun.). Given the extent of poaching problems involving other species, it is not unreasonable to suspect that more illegal harvest has gone unnoticed or unreported. Turkeys are particularly vulnerable during the winter months, especially to "roas t+shoot lnq" (Markley 1967). In a relatively sparse population, such as occurs along the South Platte, small illegal take could have significant consequences. For the Tamarack population, some genetic characteristics of the birds may be contributing to the lack of growth and expansion. It is possible that the Texas strain does not adapt well to northern climatic/ecological conditions. Kansas has experienced better success with transplant stock from Oklahoma than w i t h Texas birds (R. Little, pers. commun.). Nebraska's transplant program used Texas turkeys exclusively (K. ~1enzel, pers. commun.). Loss of vigor from inbreeding is another factor possibly limiting populat i on qr owth , �156 HABITAT USE General Observations by this investigator, other CDOW personnel, and landowners along the South Platte indicate that turkeys are largely confined to the cottonwood-riparian habitat immediately adjacent to the river. Several sightings up to 2 km from the river-bottom have been reported. Riparian and adjacent habitats along the South Platte are similar in size, structure, and composition to areas of Kansas and Nebraska where Rio Grande turkeys were successfully introduced (Peabody 1963, R. Little and K. Menzel, pe,s. commun.). In these states, turkeys are also confined to the riparian zone and adjacent areas. However, in Kansas, several flocks have become established in large, abandoned farmstead shelterbelts (R. Little, pers. commun.). The most heavi ly used forag lng areas from late fall to early spring are harvested cornfields adjacent to the riverbottom. Heavily used fields had large amounts of waste grain present, and were within 100-200 m of dense cover. On several occasions. turkeys were observed foraging over 300 m from cover. Most turkeys located were also being fed by landowner s , Whole corn and cracked corn were the 2 feeds commonly used. Some land" owners fed only when deep snow was present, others provided food on a regular basis. Turkeys become quite accustomed to these handouts and often appear daily at the same time to be fed. They become tolerant and almost tame to their human providers, particularly during periods of extreme cold and deep snow. However, the turkeys remained quite shy and wary of "s t ranqer s". With onset of spring and flock breakup (Mar), this tameness disappeared and the birds reverted to their natural wildness. Artificial feeding of Rio Grande turkeys is common practice throughout their native range (Glazener 196]) and with introduced populations in Kansas and Nebraska (R. Little and K. Menzel, pers. commun.). This practice is an open invitation to disease and domestication problems (Kennamer 1981). None of the landowners feeding turkeys along the South Platte raised domestic turkeys, but some do raise chickens. Surprisingly, several were awar e of the potential for disease and domestication and distributed food near the riverbottom away from their farmyards. In this situation, artificial feeding may not be entirely detrimental. During winter 1983-84, deep, crusted snow persisted for weeks. "Handouts" may have been the sale food source available and could have prevented exceptionally high overwinter mortality. Feeding may also have enabled turkeys to come through the winter in better condition (Markley 1967). Porter et al. (1983) suggested that a combination of harsh winter conditions and food shortages can increase winter mortality and decrease reproductive performance of turkey hens. �157 This protective attitude towards turkeys expressed by many landowners may have potential benefits. It fosters interest and concern for the wellbeing of turkey populations. Many landowners keep a watchful eye on "their turkeys" to prevent poaching or other harrassment. This interest has also made some landowners more receptive to land management practices to benefit turkeys and other wildlife. Turkeys regularly visited cattle feeding areas to forage for grain in cow manure. When spring greenup occurred, turkeys were observed foraging in hayfields. Feeding mostly occurred in the early morning and early evening hours. Foraging habits of wild turkeys observed along the South Platte were similar to those shown by introduced Rio Grande turkeys in Idaho, Kansas, and Nebraska (E. Bizeau, R. Little, K. Menzel, pers. commun.). Loafing, Roosting and Escape Cover Loafing and escape cover was provided by clumps of dense woody and herbaceous vegetation in the riverbottom. When surprised by a human, turkeys would flee (usually on foot) into the thickest avai lable cover. If none was nearby, they would often fly across the river to seek cover. One flock regularly used a large (300 x 10 m) evergreen shelterbelt as a travel lane and as loafing/escape cover. No regularly used roosts were located duroing this study. None of the landowners contacted had observed turkeys roosting in close proximity to their homes or other centers of human activities. Several felt the birds were using a number of roosts, well removed from human activities. Peabody (1963) reported that in Kansas, Rio Grande turkeys roost in large mature cottonwoods. Such trees are abundant in the riparian zone adjacent to the South Platte. In their native Texas range, Rio Grandes often use several winter roosts (Smith 1975). Da i1y Movements Most winter flocks observed remained within a relatively small range that seldom exceeded 1.6 km2. Flocks were often found in the same general area on repeated visits. Winter home ranges of Rio Grande turkeys in Kansas were ~ 1.6 km2 in size (Lange 1983; R. Little, pe rs . commun.). Nesting and Reproduction Breeding displays and gobbling were first observed during the 3rd week of March. Observations made in this study and by landowners during pl-evious years suggests that the peak of mating occurs in early April which compares favorably with the chronology of breeding activities of wild turkeys in Kansas and Nebraska (Cape" 1968~, 1971; K. Menzel, pers. commun.). �158 Nests and/or broods have been seen near all 3 release sites (Table 4). Nests have been reported in alfalfa fields up to 0.4 km from the river, in an adjacent fencerow, and in an upland pasture 1.6 km from the river. These nest sites are similar to those used by Rio Grandes in Kansas and Nebraska (K. Menzel and R. Little, pers. commun.). All brood sightings have been in or adjacent to the riverbottom. The largest brood reported consisted of 17 poults accompanied by 2 hens. Average brood size is 6-8 in Kansas (Capel 1968a), and 5 in Nebraska (Seutsuga and Menzel 1963). ._ Table 4. Rio Grande turkey nests and broods reported along the South Platte, Colorado, Summer 1983. ~~~~ Sighting __~~~~ Type A Nest B Brood ~~~ Size -=~~~~ Location s. 17 of Riverside H~ab}tat.ty~p_e _ Reservoir Upland pasture 9 km E. of Ft. Norgan Riverbottom 4 km W. of Snyder Riverbottorn (w/2 hens) C Brood 7 D Nest & brood 6 E Nests(5) F Brood G Brood H Brood km W. of Snyder 0.4 km SE of Merino Fencerow(2) Alfalfa field(3) 8 4 km NE of Atwood Riverbottom 5 S. of 11iff Riverbottom Tamarack Ranch --_._----EVALUATION AND RECOMMENDATIONS Population growth and range expansion achieved by the 1980 Hillrose release suggests th3t viable populations of Rio Grande turkeys can be established along the South Platte River. If this popUlation continues to grow, a limited harvest may be possible in 1-2 years. Kansas usually begins with a controlled toms only spring harvest about 3 years after successful establishment and when populations reach 100-150 birds (R. Little. pers. commun.). However, the "recessive es t ab llshment " (Leopold 1933) shown by the Nebraska and first Kansas releases suggests that it is too early to fully evaluate the releases in Colorado. If expansion of turkey populations along the South Platte continues to be a priority in the Northeast Region, further research and management activities will be necessary. The following recommendations are based on information obtained frorn this study, from contacts with other researchers and managers working with Rio Grande turkeys in similar habitats, and from the literature. �159 I. Research A. Continue collecting information on Rio Grande turkey populations along the South Platte River. 1. 2. 3. I I. Distribute landowner report cards. Conduct brood surveys to obtain production information. Continue to request information on turkey sightings on hunter surveys for Big Game Management Units 89, 92, and 881. B. Design and conduct a research project to examine nesting parameters, nest site selection, and reproductive performance of Rio Grande turkeys along the South Platte River. C. Maintain contact with researchers and managers working with Rio Grande turkeys in other states to avoid duplication of effort. Management A. Contingent on the availabil ity of suitable transplant stock, continue turkey releases on the South Platte to increase genetic diversity and fill in sections of unoccupied habitat. 1. 2. 3. B. Avoid any future use of Texas birds. Priority release sites should include: a. the west end of the Tamarack Ranch Property, b. the stretch of river between Goodrich and Fort Horgan, c. the stretch of river between Sterling and Proctor. On all future releases, mark all birds with wing tags to help obtain information on post-release dispersal and movements. Provide information to landowners on enhancing and turkey populations on their lands. turkey habitat LITERATURE CITED Bailey, R. W., and K. T. Rinell. 1967. Events in the turkey year. 73-91 in O. H. Hewitt, ed. The wild turkey and its management. Wi Idl.Soc., Wash., D.C. Pages The Baker, B. W. 1979. Habitat use, productivity and nest predation of Rio Grande turkeys. Ph.D. Diss., Texas A&M Univ., College Station. 46pp. Capel, S. W. 1967. Wild turkey population trends and introductions. Compl. Rep., Kans. For. Fish and Game Comm. P-R Proj. W-23-R-5. Job 41pp. 1968a. Wild turkey population trends and introductions. job Compl. Rep., Kans. For. Fish and Game Comm. P-R Proj. W-23-R-6. 29pp. �160 Capel, S. W. 1968b. Management and hunting of wild turkeys. Job Compl. Rep., Kans. For. Fish and Game Comm. P-R Proi. W-23-R-7. 60p. 1971. Management and hunting of wild turkeys. Kans. For. Fish and Game Comm. P-R Proj. W-23-R-7. Fina 1 Rep., l Spp . Glazener, W. C. 1967. Management of the Rio Grande Turkey. Pages 453-492 in O. H. Hewitt, ed. The wild turkey and its management. The Wildl. Soc., Wash., D.C. Kennamer, J. E. 18-19. 1981. Winter feeding -- good or bad? Turkey Call 8(6): Lange, C. A. 1983. Fundamentals of successful wild turkey trap and transplant operations. Kans. For. Fish and Game Comm. Unpubl. manus c . Lindauer, I. E. 1983. A comparison of the plant communities of the South Platte and Arkansas River drainages in eastern Colorado. Southwest. Nat. 28: 249-259. Leopold, A. s. 1933. N.Y. 481pp. Game management. Charles Scribners Sons, New York, Logan, T. H. 1970. A study of the Rio Grande turkey by radio telemetry. Final Rep., Okla. Dep. Wildl. Conserve P-R Proj. w-86-R. 41pp. Markley. M. H. 1967. Limiting factors. Pages 199-243 in O. H. Hewitt, ed , The wild turkey and its management. The Wildl-.-Soc., Wash., D.C. Ogden, C. 1983. Habitat use, reproduction and mortality factors of an introduced population of Rio Grande turkeys in Idaho. Segment Rep., Idaho Dep. Fish and Game. P-R Proj. WU-129. 2pp. Peabody, W. 1963. Distribution, habitat and population trends of turkeys in Kansas. Job Compl. Rep., Kans. For. Fish and Game Camm. P-R Proj. W-23-R-1. 22pp. Porter, W. F., G. C. Nelson, and K. Mattson. 1983. Effects of winter conditions on reproduction in a northern wild turkey population. J. Wildl. Manage. 47:281-290. Robertson, K. 1963. Evaluation of experimental releases of wild turkey in Nebraska. Quart. Rep., Neb. For. and Parks Comm. P-R Proj. W-15-R-39, 12pp. 1964. Evaluation of experimental releases of wild turkey in Quart. Rep., Neb. For. and Parks Comm. P-R Proj. W-15-R-40. Nebraska. 11pp. �161 Smith, D. M. 1975. Behavioral factors influencing variability of roost counts for Rio Grande turkeys. Pages 170-175 in L. K. Halls, ed. Proc. 3rd Nat. Wild Turkey Symp. Tex. Chapter1Jildl. Soc., San Antonio. Suetsugu, H. Y., and K. E. Menzel. 1963. Wild turkey introductions in Nebraska. Trans. North Am. Wildl. and Nat. Resour. Conf. 28:1-20. Thomas, J. W., C. Van Hoozer, and R. G. Marburger. 1966. Wintering concentrations and seasonal shifts in the range of the wild turkey. J. Wildl. Manage. 30:34-49. �<1' N JULESBURG N •••.E .... W~ S Jackson Reservoir SCALE (kilometers) Riverside Reservoir r-: ---.--..--,--,.--1 10 20 30 40 50 60 70 ao '. FORT MORGAN Figure 1. Location of study area along the South Platte River in northeastern Colorado. �JULESBURG N w. Sedgwick -dt ~E -G S Proclo; iii" Messe. Jackson Reservoir SCALE (kilometers) Riverside Reservo:r I I 10 I 20 t 30 I 40 I 50 I 60 I 70 I 60 -0 FORT MORGAN 1/83 Figure 2. Locations of Rio Grande turkey transplants made by the CDOW along the South Platte River. 0' W �C1' J:- JULESBURG N Sedgwick w s Proctor llill Jackson Reservoir Riverside Reservoir Prewitt SCALE (Idlomelers) Reservoir 10 20 30 40 50 60 70 60 A Figure 3. Winter to data presented flocks located along the South Platte River, January-April in Table 2. 1984. Letters correspond �JULESBURG N Sedgwick E s Iliff Jackson Reservoir Riverside Reservoir Prewill Reservoir SCALE (Idlometers) 10 20 30 40 50 SO 70 80 FORT MORGAN Figure 4. Unconfirmed sightings of Rio Grande turkeys, January-April 1984. 0' Vl �0" 0" JULESBURG N --} Sedgwlctl w s 89 Proctor Iliff 88 92 Messe. Jackson Reservoir SCALE Prewitl Reservoir Riverside Reservoir I ~ I 10 I 20 (kilomelerS) t 30 I 40 I 50 I 60 I 70 I SO fORT MORGAN Figure 5. Boundaries of Big Game Management Units 89. 92 and 881 along the South Platte River. �Colorado Division Wildlife Research Apr iI 1984 167 of Wildlife Report JOB PROGRESS State of Colorado ----~~~~~---------- Project W-37-R-37 Work Plan Turkeys Job: Chronology in Relation Period Covered: Game Bird Survey (45-01-504-15050): 12 Job Title: REPORT 15b of Breeding and Nesting Activities to Timing of Spring Hunting of Wild Seasons 1 July 1983 through 30 June 1984 Author: Richard W. Hoffman Personnel: C. E. Braun, R. W. Hoffman, R. L. Holder, T. J. Spezze, and R. D. Velarde, Colorado Division of Wildlife; Ken Medve, Colorado State University. ABSTRACT Objectives of this study were to (1) examine and evaluate existing data on the Merriam's wild turkey throughout its range to identify management problems and potential research needs, and (2) prepare a detailed study plan on an appropriate research topic based on the problems and the needs identified in objective 1. A detailed study plan was prepared. ��169 CHRONOLOGY OF BREEDING AND NESTING ACTIVITIES OF WILD TURKEYS IN RELATION TO TIMING OF SPRING HUNTING SEASONS Richard W. Hoffman Past research efforts on Merriam's Wild Turkey (Meleagris gallopavo merriami) in Colorado and elsewhere throughout its range have been inadequate and poorly designed. These earlier studies were mostly descriptive in nature and none was designed to test specific hypotheses. The traditional approach to managing turkeys in Colorado has been through trapping and transplanting when, ironically, minimal knowledge exists concerning distribution, habitat preferences, population levels, production rates, survival rates, and harvest levels. Baseline data about these and other population attributes are essential for identifying problems, formulating testable hypotheses, and developing management strategies. P. N. OBJECTIVES 1. Examine and evaluate existing data on Merriam's Wild Turkey throughout its range to identify management problems and potential research topics. 2. Prepare a detailed study plan on an appropriate determined from Objective 1. research topic as SEGMENT OBJECTIVES 1. Rev iew I iterature on wi ld turkey management 2. Review literature and test methods for age and sex classification, locating, trapping, and banding wild turkeys. 3. Interview selected personnel of the Colorado Division of Wildlife concerning wild turkey research needs. 4. Interview selected personnel involved in research or management of wild turkeys in other states to identify potential problems and assist in the selection of an appropriate research topic. and ecology. 5. Visit occupied ranges of wild turkeys in Colorado that are potentially suitable for conducting research. 6. Prepare a detailed study plan on an approved research topic and select study areas to meet research requirements. �170 RESULTS AND DISCUSSION The following study plan has been prepared in compliance with the 1983-84 objectives of Work Plan 12, Job 15. Full implementation of this study plan is dependent upon receiving outside funding or additional CDOW funding. Otherwise, only approaches 1, 2, 10, and 11 can be accomplished in 1984-85 with the present funding level of $5,000.00. A. NEED Merriam's wild turkey (Meleagris gal~o~avo merriami) have been legally hunted in Colorado since 1949(Burget 1957). Only fall hunting was pe r+ mitted until 1964 when the first spring season was held; subsequent spr'lng seasons were held in 1965, 1967, 1968 and 1970 (Colo. Div. I,Jildl.1983). A continuous 23-day spring season, opening in mid-April (range, 14-21 Apr) and extending through early May (range, 6-13 May), has been held every year since 1973. In 1973, an estimated 496 hunters harvested 64 turkeys during the spring season. By 1982 (1983 figures are not available), 2,978 spring hunters were harvesting 632 turkeys. This increase in hunter par-: ticipation and harvest has generated a growing concern among managers about the impacts of the additional pressure upon the resource. Their specific concerns include: 1. What portion of the spring gobbler population without disrupting breeding success? can be safely harvested 2. When is the best time to have a spring season to maXImize harvest of males and minimize disturbance of breeding females? 3. What effects do spring hunting have upon reproductive success of females? 4. How variable is the timing of breeding and nesting activities on an annual and regional basis? Previous studies of the wild turkey in Colorado (Burget 1957; Hoffman 1962,1966, 1968, 1973; Myers 1973) failed to address these concerns. To date, no quantitative data exist upon which to justify the timing and length of spring seasons or to biologically justify these seasons. CDOW personnel in the SE Region feel that acquisition of such information is critical to convincing landowners to allow spring hunting and to meeting strategic plan objectives of increasing hunter opportunity and harvest through improved access. �171 B. OBJECTIVES Major objectives of this study are to: Document the timing of winter flock dispersal, onset of gobbling, -----peaks of qobb llnqi-ne st initiation, onset of incu-bation; and peak of hatch in relation to geographic area, year, and timing of the spring season.- 1. 2. Describe the gonadal -cvc le-of fema-les and compare the reproductive condition of females in relation to timing of the spring season. 3. Measure the abandonment of human disturbance 4. C. females to varying levels hunter activity -end-harvest -of wi ld turkeys on a statewide Monitor basis. EXPECTED rate of incubating around the nest. RESULTS AND BENEFITS Es t ab l ls hmerrt of b lo loq i ce lly sound and soc lal lv acceptable harvest negulations is a recurring pr obl ern facing an game management agencies. It is anticipated that data derived from this study will be instrumental in formulating season recommendations for wild turkeys that max im i ze hunter opportunity without impacting reproductive success. Providing factual information regarding the mechanisms controlling spring gobbling and nesting activities, in terms that hunters can use to improve their hunting techniques, should help increase hunter success and thus better use an available 'resource. If it can be demonstrated that spring hunting has a negligible impact upon reproductive success, then managers can approach landowners with convincing data to allow spring hunt, ing. Increased access is the key to providing more recreational opportunity and attaining desired harvest levels. Successful completion of this study and accompl ishment of all objectives are dependent upon receiving outside funding support. The lack of such funding will necessitate prioritizing the stated objectives into those which are economically feasible to accomplish. D. APPROACH Hypotheses to be tested include: a. Gobbllng activity of wild turkeys in Colorado extends the early May closing date of the spring season. b. Most hens have not initiated spring season opens. c. incubation Gobbling activity, timing of nesting regional and annual basis. by mid-April beyond when the events, and hatching vary on a d. Nesting subjected females exhibit a high to human disturbances. rate of nest abandonment when �172 ¢. Abandonment rates are' greater during early stages of incubation. f. Subadult hens seldom nest and have lower nesting' success than adult hens. g. Spring and fall harvest samples are comprised of mostly adult males (> 60%) and poults (> 50%), respectively. 1. Review literature pertinent to the objectives of this study. 2. Conduct a 100% survey of 1984 spring hunters to determine their in an experimental permit system and mal I wing survey •. Data on hunter activity and harvest will be collected in conjunction with this survey and provided to the \-lildlife Services Section for inclusion in the Small Game Harves t Repor t , It is anticipated that most (> 80%) hunters will be willing to cooperate in the experimental survey. This information will be useful in obtaining approval to implement the permit system wi n9 'co II ee rion pf'ogram. \:lUI ingness to cooperate 3. Select 3 study areas, 1 each in the NE, SE, and 51,.1 r.eqi ons , where adequate numbers (100+) of turkeys exist for trapping, marking, and monitoring purposes. Ccxnparisons of the chronology of breeding and nesting activities among areas within the same year and within areas between years will be used to evaluate whether spring seasons can be set on a statewide basis and whether it is necessary to reevaluate the timing of the season on an annual basis. 4. Using drop nets (Glazener et a l. 1964, Lange 1984) and cannon nets (Dill and Thornsberry 1950) attempts will be made to trap and individually mark 100+ birds on each study area. Captured birds wi] 1 be classified to sex and age (Lewis 1967), and banded with serially numbered, locking, aluminum leg bands, and patagial wing tags (Knowlton et al. 1964). Four adult males on each area will be equipped with tail-mounted radio transmitters (Bray and Corner1972). In addition, 28 adult females will be instrumented with the same package. Instrumentation and monitoring of radio-equipped females will be restricted to the study area in the SE Region. 5. ·Document timing of winter flock breakup. At least 2 winter flocks on each study area will be located and monitored every 3rd day beginning on 7 March and continuing until flock dispersal. Assuming sex segregation (Bailey and Rinell 1967),floeks to be monitored will be selected on the basis of their composition; i.e., at least 1 male flock and 1 female flock will be selected for observation. Additional flocks wi 11 be monitored in the SE Region depending upon the distribution of Instrumented birds. �173 6. Document gobbling activity. a. Starting on] March and continuing through 25 May. gobbling activity will be monitored every other day beginning 30 minutes before sunrise and continuing until 30 minutes after sunrise. This 1-hour period will be divided into 6-10 minute listening periods. Only instrumented gobblers will be monitored to minimize variability associated with using different birds and time necessary to locate birds for observation. Gobbling activity of instrumented birds will be compared with activities of uninstrumented birds to assess tha influence of the radio package on gobbling activity. Gobbling activity will be recorded as the number of gobb Ies/b i rd/l ().-m inu te per iod. Time of f i rs t qobb le and t ime of "fly down" wi l l also be recorded. Two observers each monitoring a different bird wi lI participate in each l-hour observation period. This will require the assistance of area management personnel. Gobb ler s selected for monitoring on a particular day will be located on the roost either the evening before or 1 hour before sunrise on the monitoring morning. Gobbling data will not be recorded unless positive identification is established. individual gobblers wi ll be monitored on a rotating basis. Thus, with 4 birds and 2 observers monitoring every other day, each bird will be observed every 4th day. b. Every 2nd observation day, gobbling activity will be monitored for 3 l-hour periods (30 minutes before to 30 minutes after sunrise, .1130-1230 MST, and 1600-1700 MST). Two lO-minute listening intervals within each of these 1-hour periods will be randomly selected to test responses to tape-recorded calls of a female and another gobbling male. The type of call to be played in the 1st listening interval will be randomly assigned thus fixing the call to be played in the 2nd interval. Every 2 minutes within the selected lO-minute interval, the call will be played and the response recorded. A standardized data form will be prepared that can be used for recording gobbling activity during all types of trials. c. Additional information to be collected during the observation periods inc lud e ; weather conditions (temperature, wind speed and direction, cloud cover, precipitation type and amount if any, precipitation in previous 24 hours, frost conditions, fog conditions)~ behavior at time of trial; distance to subject; presence or absence of other gobblers and hens; harem size; and agressiveness. �174 d. I, Two observers, each monitoring a different bird, will participate in each lvhour observation period. Sample sizes will therefore include 50 l-hour morning observation periods, 24 l-hour noon observation periods, and 24 l-hour evening observation p~riods. Within these observation periods there will be 144 lO-minute listening intervals (48 intervals/morning, noon, and evening observation period) randomly selected to test gobbler responses to tape recorded calls; 72 will involve female calls and 72 gobbling of another ma le , 7. Radio-equipped females will be located biweekly beginning on 7 March and continuing through the breeding, nest Initiation, lncubat ion , and hatching periods. Attempts will be made to approximate the dates of these activities as closely as possible without disrupting the females. Documentation of hatching dates will be given top priority. Timing of the other activities can be approximated by knowing the date of hatch. 8. Six nesting hens will be selected to measure the effects of human disturbance on the rate of nest abandonment. Disturbances will be designed to simulate hunting conditions; i.e., wa lk inq a t d lf f er en t distances from the nest. discharge of a shotgun atdifferent d i st arrce s from the nest, and flushing the female at different stages during the incubation period. Sample sizes (total number of known nesting females) will be a problem. Assuming an 80% nesting rate, and 50% nesting success, potentially, only 11 of the 28 instrumented hens wi 11 provide data through the incubation period. Some of these hens wi] 1 undoubtedly be included in the disturbance experiments and may possibly abandon their nests. Therefore it will be necessary to either instrument more (re, 40) hens per year or conduct th i s portion of the study for 5 years. 9. Collect 2 hens/week (preferably 1 adult and 1 subadu lt ) starting the 1st week of April and continuing through the 4th week of May. Weights will be taken immediately upon collection. Presence or absence of a brood patch will be noted. Ovaries and oviducts will be excised and placed in 10% formalin solution for later examination and measurement. Carcasses will be frozen and stored for use in other studies. After fixation, the ovaries and oviducts will be removed, blotted dry? weighed, and measured. '[\1Jovaries will be inspected under a dissect in9 microscope for ev idence of postovu1atory f o ll i c les , The number of postovu la torv foll ides and d Iameter of the largest nonovulated follicle will be recorded for each ovary. �175 10. 11. Implement a hunter qu~stionnaire-wing permit system. collection program using a a. There will be no ~imit placed upon the number of licenses that can be sold. Anyone wanting to hunt turkeys can purchase a license at any lfcense agent, but before they can hunt, they must obtain a permit. IINot val id without a pe rrn lt " wi 11 be printed on all turkey licenses. b. Permits will be unlimited and free of charge. They will be available on a walk-in basIs at all CDOW regional ~ffices, the D~nver office. and area offices in Pueblo, Monte Vista, Durango, and Gunnison. In addition, permits can be obtained ~hrough mall application to the Denver office. The C1Ppl ication wi 11 be printed on the l+paqe inf orma t lon sheet and distributed to all license agents who sell turkey licenses. Co Separate spring and fall permits will be required. Spring perm its Itli 11 be available from 1 March through the end of the Fa lI permits wi l I be available from 1 Auqus t spr ing season. through the end of the fall season. do Every individual obtaining a permit will receive a wing envelope. Wing envelopes will be mailed along with the permit to those individuals who apply by mail. Complete instructions will be printed on each envelope. Wings and breast feathers ~'Ji 11 be examined to ascertain age (spring and fall) and sex (spring only) composition of the harvest and to estimate hatching dates, productivity rates, and nesting success. Infor-mation provided by hunters on the wing envelopes wi 11 be useful in assessing the geographic and time distribution of the harvest. e. A 100% survey of permit holders will be conducted immediately following the spring and fall seasons to obtain information on hunter activity, hunter success, and total harvest. Non-respondents will be mailed a follow-up survey. A 90% response rate is desired. f. Information resulting from the hunter questionnaire-wing co llec.tion program will be incorporated into the annual small game harvest survey. Compile data, analyze results. and prepare progress and/or final reports,and publish findings in appropriate technical journals. Data will be analyzed using standard statistical tests. Management recommendations will be included in the final report. �176 Time Schedu Ie Approaches Approaches Approaches 1984-85 1985-86 1986-87 Approach Approach Approach Approach Approach Approach Approach Approach Approach Approach Approach Personnel 1-11 1, 4-11 1, 4-11 10 July-June 1984-85, 1985-86, 1986-87 July-September 1984 September-January 1984-85 January-March 1985, 1986, 1987 March-April 1985, 1986, 1987 March-May 1985. 1986, 1987 March-June 1985, 1986, 1987 April-June 1985. 1986, 1987 April-May 1985, 1986, 1987 February-June and August-October 11 January-February 1 2 3 4 5 6 7 8 9 1985. 1986, 198] 1985, 1986, June 1987 Assignments Person-Days 1984-85 Richard W. Hoffman Util ity Worker I Wildlife Technician 198 44 III 88 330 1985-86 Richard W. Hoffman Utility Worker I Wildlife Technician III 198 44 88 330 1986-87 Richard W. Hoffman Utility Worker I Wildlife Technician II I 198 44 88 330 Yearly Costs: 1984-85 $ 67.790 1985-86 1986-87 69,000 70,000 �177 -Supervision and Cooperation Section Chief: Clait E. Braun Principal Investigator: Richard W. Hoffman Statistical Services: David Bowden Other Supervision: Richard Hopper, Chief, Wildlife Research Cooperation: CDOW Regional, Area, and District Wildlife Managers, CDOW Regional Biologist, NWTF, Colorado Chapter NWTF, Private Landowners E. LOCATION OF WORK Study Headquarters: Colorado Division of Wildlife, Wildlife Research Center, Fort Collins, CO. Field Area: SE Region, Lasi\nimas, andHtJerfano counties and statewide. F. RELATED FEDERAL AID STUDIES Colorado Federal Aid to Wildlife 15053), Wo~k Plan 12, Job 16. LITERATURE Restoration Project W-37-R (5505X - CITED: Bailey, R. W., and K. T. Rinell. 1967. Events in the turkey year. Pages 93-111 in O. H. Hewitt, ed. The wild turkey and its management. The Wi ldl . S;c., Washington, D.C. Bray, O. E., and G. W. Corner. 1972. A tail cl ip for attaching to birds. J. Wild]. t·1anage.36:640-642. Burget, M. L. 1957. The wild turkey in Colorado. Dep. Fed. Aid Proj. W-39-R. 68pp. transmitters Colo. Game and Fish Colorado Division of Wildlife. 1983. Colorado small game, furbearer varmint harvest. Dep. Nat. Resour.,Div. Wildl. 210pp. Dill, H. H., and W. H. Thornsberry. 1950. A cannon-projected capturing waterfowl. J. WildJ. tlanage. 14:132-137. Glazener, W. C., A. L. Jackson, and M. L. Cox. 1964. turkey trap. J. Wildl. Manage. 28:280-287. net trap for The Texas drop-net Hoffman, D. M. 1962. The wild turkey in eastern Colorado. Game and Fish. Tech. Publ. 2. 49pp. 1966. Merriam's turkey roost preferences Colo. Div. Wi ldl., Game Info. Leafl. 45. 6pp. and Colo. Dep. on mountain ranges. (reprinted Apr. 1979). �178 1968. .Roost i nq sites and habits of Merriam's Colorado. J. Wildl. Manage. 32:859-866. turkeys in 1973. Some effects of weather and timber management on Merriam's turkey in Colorado. Pages 263-271 in G. C. Sanderson and H. C. Schultz, eds. Wild turkey management: current problems and programs. Univ. Missouri Press, Columbia. Knowlton, F. F., E. D. Michael, and W. C. Glazener. 1964. for field recognition of individual turkeys and deer. 28:167-170. Lange, C. A. 1984. The Kansas drop-net Comm. lOpp , (unpubl. mimeo). turkey trap. A marking technique J. Wildl. Manage. Kansas Game and Fish Lewis, J. C. 1967. Physical characteristics and physiology. Pages LI5-72 O.--H. Hewitt, ed. The wild turkey and its management. The l,.Iildl. Soc., Washington, D.C. Myers, G. T. 1973. The w lld turkey 0:1 the Uncompahgre Plateau. Colo. Div. Wild]. Fed. Aid, Proj. W-37-R. 153pp. --- Prepared by ~~~~~~~~~+.r~(+------ In Final Rep ,, �Colorado Division Wildlife Research Apri I 1984 of Wildlife Report 179 JOB PROGRESS State of Colorado --------------------------- Project W-37-R-37 Work PI an 13 Job Title: (45-01-504-15050): Game Bird Survey 8 Job Population Columbian Period Covered: REPORT Characteristics Sharp-tailed 1 January and Habitat Grouse through 31 December Requirements in Northwestern of Colorado 1983 Author: Kenneth M. Giesen Personnel: Mike Bauman, Clait Braun, Ken Giesen, jim Haskins, Jim Hicks, Rick Hoffman, Mike Middleton, Dan Schaad, Steve Steinert, Colorado Division of Wildlife. ABSTRACT Counts on sharp-tailed grouse (Tympanuchus phasianellus columbianus) on 24 active leks in Routt and Moffat counties in 1983 resulted in 144 males, 28 females, and 311 total birds being counted. There was no apparent peak of male attendance while female attendance at leks peaked during the last week of April and the 1st week of May. A total of 52 sharp-tailed grouse was trapped and banded (41 males, 11 females) and 16 (7 males, 9 females) were fitted with poncho-mounted radio transmitters. Dispersal of radiomarked males from lek of capture was < 1.0 km with 82.5% of the locations occurring within 500 m. Females dispersed farther with 74.4% of the radio locations occurring between 1.0 and 2.0 km from the lek of banding. Habitats used by male and female sharptails differed with alfalfa-hay pastures used proportionately more often by males and mountain shrub communities used more often by females. A sample of 224 wings was received from the hunter harvest of which 150 (67.0%) were from juveniles. Most wings (N = 103, 46%) we re from the initial weekend of the hunting season w i th California Park being the major area of harvest. ��181 POPULATION CHARACTERISTICS AND HABITAT REQUIREMENTS OF COLUMBIAN SHARP-TAILED GROUSE IN NORTHWESTERN COLORADO Kenneth M. Giesen The Columbian or mountain sharp-tailed grouse has declined in distribution and abundance throughout its historical range, including Colorado (Aldrich 1963, Miller and Graul 1980). Reasons for this decline are not well documented although changes in land u e resulting from agriculture, energy development, and human population growth have coincided with population declines (Hart et al. 1950, Kessler and Bosch 1981). Although the Columbian sharp-tailed grouse is a game species in Colorado, little information is available upon which to base management decisions. Baseline data on Columbian sharptail populations, habitats, and harvest are lacking not only in Colorado but throughout its range. Previous studies on distribution of Columbian sharptails in Colorado indicated that the largest populations and apparent best habitats occur in Routt and eastern Moffat counties (Rogers 1969, Giesen and Hoffman 1981) with most of the harvest occurring near California Park and Twentymile Park in central Routt County. Research on the sharptail resource is needed to obtain basic information on breeding densities, nesting success, production and survival of young, fall population numbers, harvest, population turnover, and recruitment to the breeding population. Information on seasonal movements and habitat use is needed to quantify food and cover requirements. Land use changes resulting from human population growth, agriculture, and energy development are expected to further reduce sharptail habitat in the near future while hunting and other recreational uses of sharptails and their habitat are expected to increase. This report covers the 3rd year of a 5-year study. P. N. OBJECTIVES The major objectives are to measure sharptai! breeding density, production, harvest, survival and turnover rates, and to obtain qualitative and quantitative measures of Columbian sharptail habitat in western Colorado. Segment Objectives: 1. Review pertinent literature applicable to the objectives of this study. 2a. locate dancing grounds in March, April, and May by systematic search of suitable habitats using binoculars, spotting scopes, and a parabolic microphone listening device during morning and evening display periods. �182 2b. Count male and female sharptails on known sharptail leks in the 2 study areas twice weekly from mid-March through May during morning display periods. Count male and female sharptails on leks adjacent to study areas at bi-weekly periods during April and May. 2c. Identify individual male and female sharptails on leks using binoculars and spotting scopes to observe color band combinations. 3a. Locate sharptail broods by systematic search of suitable habitats and by using a tape-recorded chick distress call. 3b. Ascertain brood size from counts of broods located in July and August. 4a. Trap adult sharptails using walk-in traps on leks and brood areas and baited walk-in traps in fall and winter. and individually mark with aluminum bands and colored plastic bands. Place solar or batterypowered radios on 2 males and all females captured on the study leks. 4b. Capture sharptail chicks using walk-in and/or drive traps in areas where sharptails are known to occur. 5. Locate all radio-marked sharptails weekly for documentation of home range size and seasonal habitat use. 6. Habitat at sharptail use sites and random sites will be quantified using cover boards, vertical measurements. intercept, and point-centered quarter 7. Estimate nesting success from wing molt of hens captured in summer and from wings obtained from hunter-harvested birds. 8. Ascertain distribution of hunters and hunter harvest from field hunter checks. check stations, and wing barrels. 9. Obtain number and location of marked birds shot through use of field hunter checks, check stations, and voluntary mail reporting. 10. Food habits will be ascertained from crops of sharptails obtained from hunters and systematic collections. 11. Compile data, analyze results, and prepare progress report. METHODS Field surveys were conducted on both study areas (Cedar Hill Gulch, Hayden-North) from mid-March through May using binoculars and a parabolic microphone listening device to locate leks within the study areas. Binoculars and spotting scopes were used to obtain counts of male and female sharptails on leks and flush counts were used to obtain complete counts. Walk-in drive traps were used to capture sharptails on leks. Intensive searches on foot using a tape-recorded chick distress call or a trained pointing dog were used to locate sharptail �183 broods in July and August. Sixteen solar- or battery-powered radios were attached to ponchos (Amstrup 1980) and placed on sharptails to facilitate periodic relocation. Vegetative structure and species composition was measured on leks, grouse use sites, and random sites using a cover board (Jones 196B), vegetative intercepts (Wiens and Rotenberry 1981), and the point-centered quarter methods ~Cottam and Curtis 1956). Distribution of hunters and hunter harvest was measured using 2 check stations, field checks, and 10 wing barrels. DESCRIPTION OF STUDY AREA Two areas (Cedar Hill Gulch, Hayden-North) were selected for intensive study. Cedar Hill Gulch is in Moffat County (T9N R89W, parts of Sections 31,32, and 33; TBN R89W, parts of Sections 4-10 and 15-17) and is primarily dominated by native shrub communities and irrigated hay meadows. Hayden-North is in Routt County (T8N R88w, parts of Sections 3-5, and T9N R88W, parts of Sections 19-21 and 28-33). Hayden-North is do~!nated by native shrub communities, pastures, and wheat fields. A complete vegetative description will be included in the final report. RESULTS AND DISCUSSION Lek Counts From 22 March through 15 June, 196 counts of sharp-tailed grouse on 24 active leks were obtained in Routt and Moffat counties. Twenty counts were made on an additional 10 historic leks (Bear Creek, Cal ifornia Park Road #1, California Park Road #2, Fly Gulch, Green Acres, Pinnacle, Schneiders, Scott Place, Taylors, Wingate) on which no sharptails were seen. Only 3 of these leks were active in 1982 (Bear Creek, California Park Road #1, Scott Place). A minimum of 311 sharptails (13.0/active lek) was counted. The number of active leks counted in 1983 decreased slightly from 1982, primarily due to poor road conditions and inclement weather. Overall, the number of sharptails counted and the average number per active lek has remained relatively stable since research efforts began in 1981 (Table 1). Two leks were relocated and counted for the first time since Glenn Rogers' work in the 1960's (Morapos Gas Field, Noland Ranch). Because of other research activities and poor access, little effort was directed to locating additional leks in 1983. �184 Table 1. counties, Year 1964 1965 1977 1978 1979 1980 1981 1982 1983 Sharp-tailed grouse dancing ground counts 1964-65 and 1977-83. N active grounds counted 7 15 2 6 15 5 24 26 24 N sharpta i is 38 91 16 54 123 36 335 317 311 counted in Routt and Moffat Average/ lek 5.4 6.7 8.0 9.0 8.2 7.2 14.0 '12.2 13.0 ---There was an uneven counting effort among leks. Five leks accounted for most of the counting effort (124 counts, 63.3%) because counts coincided with trapping activities. The optimal number of counts is unknown a l-: though the 5 leks counted 10 or more times averaged 15.0 birds while the remaining 19 leks averaged 9.8 birds. Leks counted at weekly intervals had the most birds prior to or during female attendance. After female attendance, the number of males observed attending leks decl ined and their display activity became sporatic or ceased. The best time to locate leks and obtain maximum counts is in late March or April. Preliminary analysis of count data and evidence fom the Jiterature suggests that not all male sharptails attend leks in spring. This may be especially true for yearl ing males. Although there was high production in 1982 (66% chicks in the harvest), yearlings apparently did not recruit in high numbers in i983. If yearlings survived, they may recruit in 1984 as 2-yearolds. Banding data support this hypothesis as only 6 of 36 males (16.7%) trapped on leks in 1983 were yearlings, yet survival of banded males was less than 50%. Most males recruiting to leks are apparently 2 years of age or older. This suggests that the number of leks in a given area may be a better indication of population trend than the number of males (or total birds) per lek. Trapping and Banding A total of 52 sharp-tailed grouse (41 males, 11 females) was captured and individually marked in 1983. All but 2 were captured using walk-in drive traps on leks; 2 hens were netted on their nests. Over 60% of the maJes attending leks on the study areas were captured and marked (Table 2). Only 4 of 41 males trapped on leks in 1983 were yearl ings (9.8%) suggesting that adult males were easier to trap because of their dominant behavior or that maJes do not recruit into the breeding population until they are 2 years of age or older (Rippin and Boag 1974). �185 Table 2. Trapping success of male sharp-tailed Hill Gu Ich , Pe II y IS, and Sm ith I s leks, 1983. High count of males Lek Cedar Hi 11 Gulch Pelly I s Smith's Telemetry Percent males banded N males banded 19 A II leks grouse on leks at Cedar 9 it?4 9 t 77 .8 18 12 66.7 46 28 60.9 Investigations Sixteen radio transmitters were placed on sharp-tailed grouse (9 females, 7 males) from April through June to obtain data on nesting, movements, and habitat use. Mortal ity and transmitter malfunction resulted in los5 of information beyond 5 weeks for 5 birds although transmitters on 7 birds functioned a minimum of 18 weeks. The largest number of radio-marked grouse were associated with the Smith's Lek study area and resulted in 3 males being located a total of 63 times and 4 females a total of 42 times. Average clutch size of 7 nests located by telemetry in 1983 was 10.4 (range 9-14). All locations of radio-marked males were within 1.0 km of the lek they were trapped on with 82.5% of the observations being within 500 m. Females dispersed farther from the banding location with 74.4% of the locations being between 1.0 and 2.0 km from the lek of banding. Habitats used by male and female sharp-tailed grouse also differed with males using alfalfa-hay pastures most often while females used serviceberrysagebrush (Amelanchier-Artemis~) communities most (Table 3). Table 3. Habitat availability and use by radio-marked male and female sharp-tailed grouse trapped on Smith's Lek, Routt County, Colorado, 1983. Habitat Percent a occurrence type 59.4 32.5 6.3 Mountain shrub Wheat Alfalfa-hay Roads, farms 1.9 Totals a b < 2.0 100. 1 km of lek of banding. Apr-Dec 1983. b ill Ma I es 25.4 1.6 73.0 Percent use = 3) F ema I es C& = 83.7 2.3 14.0 0 0 100.0 100.0 'of) �186 Harvest The hunting season for Columbian (mountain) sharp-tailed grouse in 1983 started one-half hour before sunrise on 10 September and closed on 25 September (Units 14, 16, 18, 20, 22, 24, 26, 28, 54, 58, 60 9 October (Units 10 and 12). Since most huntable populations of Columbian sharptails in Colorado occur in Units 14, 16, and 26 they were exposed to a 16-day season. This was the same season length as in 1981 and 1982 but shorter than the 1980 season in Units 14 and 16 (25 days). Bag and possession 1imits were 3 and 6 and separate from the bag Iimits of sage grouse (Centrocercus urophasianus). This is the 2nd season in modern times having separate bag and possession limits for these 2 species. The sample of wings received in 1983 (224) was greater than that received in any year since 1976 when we began collecting grouse wings in Routt and Moffat counties. The increased sample of wings received in the last 3 years is primarily due to the establishment of 2 sharptail research check stations and 10 wing barrels. Wings were received from 4 check stations (California Park"" 45; Cedar Mountain = 4; Twentymile Park = 3; Gould = 2), 14 wing barrels (149 wings total), and field checks of hunters (21 wings). Of this sample, 121 were from Unit 14, 9 from Unit 16, and 94 from Unit 26. Comparative data from 1981 and 1982 are presented in Table 4. Nearly half (46.0%) of the wings were collected during the initial wee kend of the season. Operation of grouse check stations on opening weekend accounted for 54 (52.4%) of these wings. Harvest on weekdays was less than on weekends. Temporal distribution of the 1983 harvest is given in Table 5 along with comparative data for 1981 and 1982. Table 5. Time distribution western Colorado, 1981-83. of sharp-tailed grouse wings received, north- 1981 Period First weekend First week Second weekend Second week Third weekend Totals 1983 1982 N % 104 26 7 3 2 73 .2 18.3 142 % N 50.6 12.4 10.7 10. 1 16.3 103 29 37 1.4 90 22 19 18 29 50 16.5 2.2 22.3 99.9 178 100. 1 224 99.9 4.9 2. 1 N % 5 46.0 12.9 �18] Table 4. Origin of sharp-tailed grouse wings, northwestern 1981 Location N 1981-83. Colorado, 1982 % N 198::...:3=--_ % N Of /0 Unit 14 Calif. Park Check Station Gould Check Station Black Mountain Wing Barrel Ralph White Res. Wing Barrel Moffat Co. Rd. 29 Wing Barrel Hayden Cog Rd. Wing Barrel Calif. Park Wing Barrel Elk Mtn. Wing Barrel Hahn's Peak Rd. Wing Barrel Field Checks Subtotal Unit 16 Cedar Mtn. Check Station Cedar Mtn. Wing Barrel Moffat Co. Rd. 3 Wing Barrel Subtotal 35 19.1 45 0.0 0.0 0.0 2 0.9 1 o 0.4· 0.0 N/A 11 4.9 5.1 3 16 16 17 10 1.3 "7.1 24.6 0.0 0.0 34 0 0 N/A N/A 3.5 N/A N/A 0 N/A 9 2 1.4 3 N/A NiA N/A 1.7 N/A o 0.0 1 0.6 24 16.9 17 9.6 66 46.5 64 36.0 121 54.0 1 0 0 0 0.0 0.0 0.0 4 5 1.8 2.2 2 0.7 0.0 1.4 o 0.0 3 2.1 0 0.0 9 4.0 38 26.8 4.2 2LI 6 13.5 3.9 3 15 5 3.5 36 23 o 0.0 9 6.3 2 28 20.0 1.1 15.7 6.7 10.3 3 24 1.3 10.7 12 8.5 2.1 13 7.3 9 0.0 0.0 3 3 o o 5 o 20.1 7.1 7.6 4.5 Unit 26 Twentymile Park Check Station Twentymile Park Wing Barrel Hayden Gulch Wing Barrel Milner Wing Barrel Steamboat Springs Wing Barrel Oak Creek Wing Barrel Wolcott Wing Barrel Rock Creek Wing Barrel Field Checks 7 1.3 o o 0.0 o o 0.0 3 1.7 11 4.0 1.3 1.3 4.9 73 51.4 113 63.5 94 42.0 0.0 1 0.6 o Subtotal o o 0.0 1 0.6 o TOTAL 142 100.0 Subtotal 3 Unit 54 Muddy Creek Wing Barrel 178 100.1 224 100.0 �188 Age and Sex Composition Age was ascertained from 224 wings examined and gonadal inspection of 53 sharptails provided sex ratio in the harvest (Table 6). The percentage of birds in the yearl ing age category is a minimum estimate as most adults and yearlings (64.9%) had molted primary 10. Sex was ascertained from 5 birds retaining Pl0; all but 1 were females. Thus, it is likely that most sharptails retaining Pl0 during the hunting season are late or renesting hens. Table 6. Age composition Colorado, 1983. of harvested sharp-tailed Males Age class N % a a Females ----_._-_. % N grouse, northwestern 'N % --.Adults Yearl ings Chi cks 64 10 150 28.6 4.5 67.0 5 0 18 21.7 0.0 78.3 Ii 3 23 13 .3 10.0 76.7 __ ._----- . aOnly those for which gonads were examined. While we cannot assume a random sample was obtained from the sharptail population we can draw some conclusions from our data. 1. Production of young was excellent in 1983 (67.0% chicks) and was the highest measured in 8 years of wing collections. 2. Because the percentage of juveniles in a stable population is a measure of total year-to-year turnover, we can be confident that even in years of poor production (i.e., 1981) more chicks are produced than can be recruited into the following year's breeding population. Our long-term harvest data suggests that annual population turnover exceeds 50%. 3. The sex ratio in the adult and young-of-the-year age-classes is sl ightly skewed toward females although the 1976-83 average indicates an even sex ratio. Other studies of prairie grouse have also indicated an even sex ratio in all age-classes. Data on age and sex composition of the hunter harvest from all years for which data are available are presented for comparison in Table 7. �189 Table 7. Age and sex composition western Colorado, 1976-83. Adults Year N 1976 1977 1978 1979 1980 1981 1982 1983 10 47 13 32 25 83 60 74 a % 71.4 65.3 46.4 40.5 39.7 58.4 33.9 33.0 Totals 345 8-year average Young % N 4 25 15 47 38 59 117 150 28.6 34.7 53.6 59.5 60.3 41.6 66.1 67.0 453 of harvested sharp-tailed Total sample Adults Males grouse, north- b Young Females 14 72 28 79 63 142 177 224 5 3 14 9 5 798 37 4 Males Females ~--•..-- 6 4 18 7 3 13 7 24 7 18 10 3 15 19 23 41 71 7Lf 5 1 56.8 43.3 alncludes yearlings. b Known sex only. LITERATURE CITED Aldrich, J. W. 1963. Geographic orientation J. Wildl. Manage. 24:529-545. Amstrup, S. C. 1980. 44:214-217. A radio-collar of American Tetraonidae. for game birds. J. Wildl. Manage. Cottam, G., and J. T. Curtis. 1956. The use of distance measures phytosociological sampl ing. Ecology 37:451-460. in Giesen, K. M., and D. M. Hoffman. 1981. Distribution and status of mountain sharp-tailed grouse. Final Rep., Colo. Div. Wildl., Fed. Aid Proj. W-37-R-34. Apr. 1981. Pp. 183-189. Hart, C. M., O. S. Lee, and J. B. Low. 1950. The sharp-tailed grouse in Utah. Its life history, status, and management. Utah Dep. Fish and Game, Fed. Aid Div. Pub]. 3. 79pp. Jones, R. E. 1968. A board to measure cover used by prairie grouse. J. Wildl. Manage. 32:28-31. �190 Kessler, W. B., and R. P. Bosch. 1981. Sharp-tailed grouse and range management practices. Pages 133-146 ~ Proc. Wildl./Livestock Symp., Univ. Idaho, Moscow. Miller, G. C., and W. D. Graul. 1980. Status of sharp-tailed grouse in North America. Pages 18-28 in P. A. Vohs, Jr., and F. L. Knopf, eds. Proc. Prairie Grouse Symp., Okla. State Univ., Stillwater. Rippin, A. B., and D. A. Boag. 1974. Recruitment to populations of male sharp-tailed grouse. J. Wildl. Manage. 38:616-621. Rogers, G. E. 1969. The sharp-tailed grouse in Colorado. Game, Fish, and Parks, Tech. Publ. 23. 94pp. Colo. Div. Wiens, J. A., and J. T. Rotenberry. 1981. Habitat associations and community structure of birds in shrubsteppe environments. Ecol. Monogr. 51:21-41. Prepared by ~jjl.~~ K~G i'esen--- Wildlife Researcher C �Colorado Division ife Research Apri I 1984 Wildl of Wildlife Report 191 JOB PROGRESS REPORT Colorado State of --------------------------- Project W-37-R-37 Work P Ian 17 Job Title: Period Covered: Game Bird Survey (45-01-504-15050): Job: Population 7 Dynamics 1 January of White-tailed through 31 December Author: Clait Braun and Ken Giesen Personnel: Clait Braun and Ken Giesen, Ptarmlqan 1983 Colorado Division of Wildl ife. ABSTRACT Long-term studies of populations of white-tailed ptarmigan (LagoDus leucurus) were continued at hunted (Mt. Evans) and unhunted (Rocky Mountain National Park) areas in Colorado in 1983. Densities of breeding ptarmigan continued to decrease from the high in 1976 (Rocky Mountain National Park) but stabilized at Mt. Evans. Nesting success was average (Rocky Mountain National Park = 54.5%) to better than average (Mt. Evans = 60%). Average brood size to 1 September was 2.8 (Mt. Evans) and 3.5 (Rocky Mountain National Park) chicks/successful female. Hunters at Mt. Evans in fall 1983 removed at least 13% of the estimated fall population of ptarmigan. ��193 POPULATION DYNAMICS OF WHITE-TAILED PTAR~IIGAN Clait E. Braun and Kenneth M. Giesen Long-term studies of trends in population size and investigation of reasons for fluctuations in size of tetraonid populations are lacking. Studies on the population dynamics of unhunted and hunted populations of white-tailed ptarmigan were initiated in Colorado in 1966 and have continued essentially uninterrupted at 2 sites. Studies of the unhunted population (Rocky Mountain National Park) have identified possible short-term cycles of 7-8 years with an amplitude of 25-30% between high and low breeding densities. Conversely, studies of the manipulated population (hunted) at Mt. Evans through 1980 have not indicated any cyclic pattern and it would appear that controlled hunting may mask any long-term trend that may occur. This report covers the 3rd year of a 5-year study designed to examine the question whether white-tailed ptarmigan are truly cyclic and whether hunting affects the apparent oscillations. P. N. OBJECTIVES The goals of this investigation are to be able to predict the length and amplitude of cycles in white-tailed ptarmigan in Colorado, to examine the impact of hunting on cycles, and to clarify underlying causes of the apparent cycles. SEGMENT OBJECTIVES 1. Conduct breeding (May-Jun) and brood (Aug-Sep) censuses of whitetailed ptarmigan using tape-recorded calls of males (breeding) and chicks (broods). 2. Censuses will be conducted on prev lous lv established, defined study areas at Mt. Evans (hunted) and at Rocky Mountain National Park (unhunted) . 3. Capture (noose poles) and band (aluminum and plastic color-coded bands) all unmarked white-tailed ptarmigan encountered on study areas at Mt. Evans and at Rocky Mountain National Park. 4. Individually identify all ptarmigan observed on study areas at Mt. Evans and Rocky Mountain National Park through use of binoculars. 5. Make hunting season and bag limit recommendations collect hunting data through use of volunteer hunter field checks. for Mt. Evans and wing barrels and 6. Compile data, analyze results and prepare progress reports. �194 STUDY AREA AND METHODS Areas investigated were Mt. Goliath-Mt. Evans in Clear Creek County and at Tombstone Ridge-Sundance Mountain to Fall River Pass in Rocky I~ountain National Park in Larimer County. The physiography, geology, location, and vegetation of these study areas has been previously described (Braun 1969. 1971; Braun and Rogers 1971; Giesen 1977). Ptarmigan were located through use of tape-recorded calls (Braun et al. 1973), captured through use of telescoping noose poles (Zwickel and Bendell 1967) as described by Braun and Rogers (1971). and classified to age and sex and banded following Braun and Rogers (1971). Age of chicks was estimated following Giesen and Braun (1979). Numbered plastic bandettes were not used as in earlier years (Braun and Rogers 1971) as a color-code system using up to 4 different colored plastic bandettes was instituted in 1977-78. A check station was operated on the Mt. Evans highw2Y during the opening weekend (24-25 Sep) of the ptarmigan season in that area. A volunteer wing collection station was available to hunters in the area from 25 September through 9 October when the season closed. RESULTS AND DISCUSSION Mt. Evans Surveys of breeding white-tailed Spring 1983 (May-Jun) revealed mated males. This is a density (Table 1). Pairing and breeding early June, similar to 1982 but snow cover in 1982 and 1983. Rocky Mountain ptarmigan on the Mt. Evans study area in the presence of at least 11 pairs and 4 unof 6.5 birds/km2, the same as in 1982 activities were initiated in late May and later than in 1981 because of the extensive National Park Surveys of ptarmigan present on the Rocky Mountain National Park study units during May and June 1983 indicated that 23 pairs and 5 single males were present. This is a density of 6.7 birds/km2, lower than the 7.8 birds/km2 recorded in 1982 (Table 1). Nesting Success and Brood Size Mt. Evans During the July-early September interval, 6 different hens were identified with broods (average size to 1 Sep = 2.8 chicks/successful hen) while 4 hens were observed without broods. Thus, 6 of 10 hens (60.0%) were known to be successful in nesting. Estimated hatching dates for 10 chicks for which data are available ranged from 11 to 27 July, Six chicks hatched from 6 eggs between 16 and 23 July in 1 known nest. \-Jhi le nesting was later in 1983 than In 1982 and 1981, nesting success was higher. �195 Table 1. 1966-83. White-tailed ptarmigan breeding densities (birds/km2), Colorado Study area Year Rocky Mountain National Park (5.5 km2) 1966 1967 1968 1969 1970 1971 1972 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 11.3 9.8 11.5 12.0 9.6 9. 1 8.7 7.8 8.0 1] .1 13·5 12.9 10.7 8.7 8.4 8.2 7.8 6.7 Mt. Evans (4.0 km2) 3.0 2.7 2.7 2.2 2.0 4.2 7.5 6.2 6.2 6.2 6.7 >6.0 7.5 10.3 9.5 9.0 6.5 6.5 Rocky Mountain National Park Six successful hens (6/11 = 54.5%) were observed between 1 September with an average brood size to 1 September of 3.5 hens were observed without broods in 1983 (5111 = 45.5%). occurred during the last week of July, about 10 days later term average. August and early chicks. Only 5 The peak of hatch than the 10ng- Harvest Ht. Evans In 1983, the ptarmigan season in the Mt. Evans area (Unit 52 south of Interstate 70 and east of the Guane11a Pass Road between Georgetown and Grant) was delayed 2 weeks after opening of the statewide season on 10 September. This delay was similar to that in the 1977-80 period following the experimentation with season length and limited closure from 1972 through 1976. This experimentation followed the documentation of overharvest in this area in the late 1960's and the 2-year closure in 1970 and 1971. In 1983, the season opened one-half hour before sunrise on 24 September and closed at sunset on 9 October. �196 A wing collection of the season was the road remained highway department was operated both barrel placed along the Mt. Evans highway prior to opening available until after 9 October. Because of the mild fall, open until 1 October when it was closed by the state because of an agreement with the CDOW. A check station days of the opening weekend. During the 2 days of check station operation, 20 hunters with 6 ptarmigan were checked. This compares to 16 hunters with 5 ptarmigan in 1982, 20 hunters with 18 ptarmigan in 1981, and 12 hunters with 9 ptarmigan checked in the same period in 1980. No additional wings were deposited in the wing barrel. Three banded birds were reported by mai 1. In all, at least 9 ptarmigan were harvested in 1983 of which 5 were banded. The fall population resident in the area was estimated to be about 40 birds. The total fall population (residents and nonresidents) in the area hunted was estimated at 50-60 birds. Thus the harvest of 9 birds represented about 13% of the fall population. LITERATURE CITED Braun, C. E. 1969. Population dynamics, habitat, and movements of whitetailed ptarmigan in Colorado. Ph.D. Thesis, Colo. State Univ., Fort Co IIins. 189pp. 1971. Habitat requirements of Colorado white-tailed Proc. West. Assoc. State Game and Fish Comm. 51:284-292. and G. E. Rogers. ----=-Colo. Div. Game, Fish ___ 1971. The white-tailed ptarmigan and Parks Tech. Publ. 27. 80pp. ~' R. K. Schmidt, Jr., and G. E. Rogers. 1973. white-tailed ptarmigan with tape recorded calls. 37:90-93. ptarmigan. in Colorado. Census of Colorado J. Wildl. Manage. Giesen, K. M. 1977. Mortality and dispersal of juvenile white-tailed ptarmigan. M.S. Thesis, Colo. State Univ., Fort Collins. 55pp. ----;-- , and C. E. Braun. juvenile white-tailed 1979. A technique for age determination of ptarmigan. J. Wildl. Manage. 43:508-511. Zwickel. F. C., and J. F. Bendel1. 1967. A snare for capturing blue grouse. J. Wildl. Manage. 31:202-204. Prepared by: Clait E. Braun Wildlife Research Leader Kenneth M. Giesen Wildlife Researcher B �197 Colorado Division of Wildl ife Wildl ife Research Report Apri I 1984 JOB PROGRESS REPORT State of Colorado --------~~~------------- Project W-37-R-37 Work Plan 21 Job Title: Period Covered: Game Bird Survey (45-01-504-15050): Job: 2 Dynamics of Cottonwood Regeneration 1 July 1983 through 30 June 1984 Author: Warren D. Snyder Personnel: Gary C. Miller and Warren D. Snyder ABSTRACT A preliminary compilation and analysis of inventories of the status and trends of cottonwoods (Populus spp.), their regeneration, and of land use changes (under contract with the Colorado State Forest Service) along the South Platte, Rio Grande, and Colorado rivers was completed and disseminated in an interim report. A contract was initiated to complete inventory of the Arkansas River and a segment of the South Fork of the Republican River. Sampling of cottonwood seedlings, resulting from natural regeneration along the South Platte River in 1983, was initiated to identify factors affecting survival. Literature and other information were compiled and analyzed as a basis for preparing a program narrative study plan which was submitted for review and approval. Trial test plots using stem cutting of 10 woody species (trees, shrubs, and vines) were initiated in late winter to evaluate stem cutting propagation in riparian sites. ��199 DYNAMICS OF COTTONWOOD REGENERATION Warren D. Snyder P. N. OBJECTIVE To prepare a study plan evaluating techniques for regeneration of cottonwood stands in riparian streamside ecosystems in lowland sites of Colorado. SEGMENT OBJECTIVES 1. Review literature and consult with personnel cottonwood-willow riparian ecosystem. 2. Select regeneration 3. Survey cottonwood-willow ecosystems menting and evaluating regeneration 4. Prepare a detailed techniques knowledgeable about the to be field tested. and select study sites for impletechniques. study plan to evaluate regeneration techniques. RESULTS AND DISCUSSION The Colorado State Forest Service was previously contracted under a nongame study plan to inventory and quantify condition and trend of cottonwood stands, other vegetation, and land use over an approximate 30-year span. They used photo interpretation methods on stratified random samples of linear miles along the lower portions of the South Platte, Rio Grande, and Colorado rivers. Data provided in this initial inventory were compiled and summarized in an interim report distributed to interested personnel within the Colorado Division of Wildl ife within this work segment. The Colorado State Forest Service, under a new contract, began inventory of the lower Arkansas River and a short segment of the South Fork of the Republican River below Bonny Reservoir. Work is to be completed during spring 1984. Since the initial analysis did not use statistical methods and data are not yet available from the 2nd contract, inventory information is not included within this segment report. It will be included as soon as complete analysis and comparison of all drainages are available. Surveys of natural regeneration of plains cottonwoods (Populus sargentii) were begun in summer and fall 1983. Several intensive test transects were established to monitor seedling survival on grazed and ungrazed sites in relation to water table fluctuations and other variables. These will be supplemented with additional transects in late spring 1984. Additional extensive transects will be establ ished to monitor the frequency of occurrence of cottonwood seedlings within the South Platte River floodplain. Cottonwood-willow ecosystems were surveyed in eastern Colorado to examine potential study sites. Surveys of West Slope riparian areas will be conducted in 1984. �200 Literature was compiled and reviewed covering riparian ecosystems, natural and artificial regeneration of cottonwoods and willows, and vegetation sampling methods. Contacts were made with personnel of the u.s. Department of Agriculture, Soil Conservation Service in New Mexico concerning stem cutting propagation, and with other Soil Conservation Service personnel concerning sources for certain species. Based on available data, 2 techniques, stem cutting propagation and soil scarification were selected for testing in riparian sites. A study site was selected on the South Platte State Wildlife Area near Crook and preliminary stem cutting propagation trials were initiateJ A tractor-mounted earth auger was used to bore holes into the ground water ta~)le which was approximately 0.5 m below the soil surface. A summary of species and numbers of stem cuttings planted is provided in Table 1. Total stem cutting length, length extending above the water table, and length extending above ground were measured on each planting. This permitted determination of the distance the stems were submerged below water level for subsequent comparisons with survival. Capped plastic tubes were inserted into the water table at 4 sites within the meadow to monitor changes in ground water level. Additional effort during the segment centered on preparation of a detailed program narrative (study plan). This plan is under review and will be approved for implementation beginning 1 July 1984. Prepared by Warren D. Snyder Wildlife Researcher C �201 Table 1. Stem cutting propagation trial plantings meadow, South Platte Wildlife Area, February-March on the check station 1984. N p Ianted Species Plains cottonwood (Populus sargentii) 6 25 Sandbar wi 1low (Sa Iix interior) 12 Coyote wi llow (~..•ex igua) 25 10 Peachleaf wil low (~. amygdaloides) Site 1 2 Date 21 Feb 24 Feb 05 Mar 1 2 6 14 Feb 02 Mar 21 Feb 1 2 3 23 Feb 01 Mar 02 Mar Golden wi llow (Sa I ix sp.) 12 12 15 Indigobush amorpha (Amorpha fruiticosa) 15 Elderberry (Sambucus spp.) 12 12 1 2 01 Mar 02 Mar Cotoneaster (Cotoneaster 15 12 2 01 Mar 05 Mar spp. ) Frost grape (vine) (Vitis vulpina) Virginia creeper (Parthenocissus guinquefolia) Totals 21 Feb 12 02 Mar 4 05 Mar 205 ��203 Colorado Division of Wildlife Wildlife Research Report April 1984 JOB PROGRESS State of Colorado -------------------------- Project W-37-R-37 Work Plan REPORT Game Bird Survey (45-01-504-15050): 22 Job: Job Title: Upland Game Publ ications Period Covered: 1 July 1983 through 30 June 1984 Author: C. Braun Personne I: C. E. Braun, K. M. Giesen, R. W. Hoffman, J. W. Hupp, and W. D. Snyder, C910rado Division of Wildlife. ABSTRACT Publications accomplished under this job in Segment 37 are: Braun, C. E. 1984. Attributes of a hunted sage grouse population Colorado, U.S.A. Int. Grouse Symp. 3: In press. in 1984. Biological investigations of white-tailed ptarmigan Colorado, U.S.A.: a review. Int. Grouse Symp. 3: In press. in ------.,.. , and T. D. I. Beck. 1983. Effects of changes in hunting regulations on sage grouse harvest and populations. Game Harvest Manage. Symp., Texas A&I Univ., Kingsville. In Press. Giesen",,_K._M. 1983. Know your grouse. Colorado Wildl. 10(8):7, 10. 1984. Identification of grou~e species by hunters in northwestern Colorado: implications for management. Colo. Div •.Wildl. Game Inf. Leaf I. 111. 2pp. , and C. ------recruitment E. Braun. 1983. Changes in survival, productivity, and of white-tailed ptarmigan during population fluctuations. Central Mountains and Plains Sect., The Wildl. Soc. 28:17. Abstract. , and ------tailed ptarmigan Abstract. 1983. Reproductive performance of female whitein Colorado. Colorado Field Ornith. J. 17(3) :7. -----. � https://cpw.cvlcollections.org/files/original/bfab47857dff2333b4ee1172f2b17057.pdf 2f92d6a256bc6b371c1707ba44987210 PDF Text Text 1 Colorado Division of Wildlife Wildlife Research Report July 1984 JOB PROGRESS REPORT State of Colorado Project No. 45-01-502-15050 Big Game Investigations - Cervids Work Plan No. Multispecies Investigations Job. No. Period covered: Author: Publishing Big Game Research Results 7 7/1/83-6/30/84 L. H. Carpenter Personnel: R. B. Gill, L. Lovett, N. McEwen, S. Torbit, and all Big Game Researchers ABSTRACT ~, ! During the 1983-84 Segment, the Big Game Research Section had 12 manuscripts published, 10 others accepted for publication, and 5 manuscripts in the review process. ! ��3 BIG GAME RESEARCH PUBLICATIONS Len H. Carpenter P. N. OBJECTIVE To publish the results of research- conducted under the auspices of Federal Aid Projects 45-01-502-15050 and 45-01-503-15050 in a variety of professional journals and other indexed publishing media to insure widespre~d dissemination and availability of this information to natural resource managers and ecological scientists. SEGMENT OBJECTIVES 1. Job Progress Reports 2. Baker, Dan L. 1984. Digestion of grass hay by mule deer and elk. J. Wildl. Manage. 3. Bartmann, R. M. 1983. Composition and quality of mule deer diets on pinyon-juniper winter range, Colorado. J. Range Manage. 4. Bartmann, R. M. 1984-. Estimating mule deer winter mortality in Colorado. J. Wildl. Manage. 5. Bartmann, R. M. 1984. Estimating breeding dates of mule deer from fetal age-length relationship. J. Wildl. Manage. 6. Bartmann, R. M. Colorado. 7. 1984. Winter severity and mule deer mortality in Wildl. Soc. Bull. Bear, G. D. 1984. An expanding telemetry collar for elk calves. Outdoor Fact Sheet, Colo. Div. of Wildl. 8. Bear, G. D. 1984. Seasonal distribution and population dynamics of the Estes Valley, Colorado, elk herd. Colo. Div. of Wildl. Tech. Rep. 9. Bear, G. D., and G. Wagner. 1984. Mark-recapture method applied to elk population estimates. J; Wildl. Manage. 10. Beck, T. D. I. 1984. Black bear harvest analyses for Colorado, 1979-82. Colo. Div. of Wi1d1. Spec. Rep. 11.- Beck, T. D. 1., and M. A. Haroldson. 1984. Estimation of weight in Colorado black bears from chest and neck girth measurements. Southwest Nat. 12. Carpenter, L. H., R. B. Gill, and D. L. Baker. 1984. Tests of nutritionally based habitat evaluation system. Colo. Div. of Wil dl. Rep. �4 13. Dailey, T. V., and N. T. Hobbs. 1984. Niche separation and feeding ecology of mountain goats and mountain sheep in Colorado. J. WildL Manage. 14. Freddy, D. J. 1983. A "Butte" of an investment. Colo. Outdoors. 15. Freddy, D. J .• R. M. Bartmann, L. H. Carpenter, and R. B. Gill. 1984. Measuring mule deer sex-and-age ratios in juniper-pinyon woodland. Wildl. Soc. Bull. 16. Freddy, D. J .• W. M. Bronaugh, and M. C. Fowler-. 1984. Reactions of mule deer to human harassment during winter. Wildl. Soc. Bull • 17. Freddy, D. J., and K. K. Karrow. 1984. western Colorado. J. Mammal. Breeding white-tailed deer in 18, Freddy, D. J .• and K. K. Karrow. 1984. Fidelity of mule deer to seasonal ranges. Colo. Div. of Wildl. Rep. 19. Green, R. L., and G. D. Bear. 1984. Daily activity patterns of elk in north central Colorado as related to vegetation types. J. Wildl. Manage. 20. Haroldson, M., and T. D. 1. Beck. 1984. of black bears. Southwest Nat. Extensive fall movements 21. Haroldson, M., and T. D. I. Beck. 1984. Habitat use by female black bears in Colorado. J. Wildl. Manage. 22. Hobbs, N. T., T. E. Remington, and R. Sayre. 1984. Effects of sagebrush on in vitro digestion of grass cell wall. J. Range Manage. 23. Hobbs, N. T., and D. S. Schimel. 1984. Fire effects on nitrogen mineralization and fi~ation in mountain shrub and grassland corrmunHies. J. Range Manage. 24. Hobbs, N. T., and R. A. Spowart. 1984. Effects of prescribed fire on nutrition of mountain sheep and mule deer during winter and spring. J. Wildl. Manage. 25. Hobbs, N. T., and R. A. Spowart. 1984. Fire effects on the distribution of nitrogen and digestible organic matter in herbage. J. Range Manage. 26. Kufeld, R. C. 1984. Effects of burning, spraying and anchor chaining of elk, mule deer and cattle use and vegetative yields of Gambel oak range land. Colo. Div. of Wildl. Spec. Rep. 27. Lance, W., and T. M. Pojar. 1984. Diseases of pronghorn antelope. A literature review. Colo. Div. of Wildl. Rep. 28. Miller. M. W., and N. T. Hobbs. 1984. Efficacy of invermectin in treatment of lungworm in bighorn sheep.' J. Wi-ldl. Diseases. �5 29. Nelson, R., and T. D. I. Beck. 1984. Natural history of season changes in behavior and biochemistry in wild bears. Science. 30. Pojar, T. M. 1984. Hematological indicators of stress in pronghorn. J. Wildl. Manage. 31. Pojar, T. M., and D. Nash. J. Wildl. Manage. 1984. Genetic variation in pronghorn. 32. Reed, D. F. 1984. Highways, deer fencing and associate structures specifications, maintenance, and effectiveness. Colo. Dep. of Highways or Colo. Div. of Wi1d1. Spec. Rep. 33. Reed, D. F. 1984. Introduced mountain goats: interactions with indigenous alpine species. Proc. North Wi1d1. Sheep and Mountain Goat Council. 34. Spowart, R. A., and N. T. HobbS. 1984. Fire effects on feeding ecology of mule deer and mountain sheep. J. Wildl. Manage. 35. Spowart, R. A., N. T. Hobbs, and D. M. Swift. 1984. Modeling fire effects on energy and nitrogen balance of ungulates during winter. Ecological Modeling. 36. Tiedeman, J. A., R. E. Francis, C. Terwilliger, Jr., and L. H. Carpenter. 1984. Steppe habitat types of Middle Park, Colorado. Colo. Div. of Wild1. Rep. 37. Torbit, 5., L. H. Carpenter, A. W. Alldredge, and D. M. Swift. 1984. Mule deer body composition - A comparison of methods. J. Wi1dl. Manage. 38. Torbit, S., L. H. Carpenter, D. M. Swift, and A. W. Alldredge. 1984. Winter dynamics of mule deer body composition. J. Wildl. Manage. PUBLICATION PROGRESS 1. Job Progress Reports - all studies these preports have been printed in: Colorado Div. of Wildl. 1984. Wildl. Res. Rep. 2. Baker, Dan L. 1984. Digestion of grass hay by mule deer and elk. J. Wildl. Manage. This manuscript will be published by J. Wildl. Manage as: Baker, D. L., and D. R. Hansen. 1984. Digestion of grass hay by mule deer and elk. J. Wild1. Manage. 48(4). 3. Bartmann, R. M. 1984. Estimating mule deer winter mortality in Colorado. J. Wild1. Manage. This manuscript was published in J. Wi1dl. Manage. 48(1):262-267. 4. Bartmann, R. M. 1983. Composition and quality of mule deer diets on pinyon-juniper winter range, Colorado. ~. Range Manage. �6 This manuscript was published in J. Range Manage. 36(4) :534-541. 5. Bartmann~ R. M. 1984. Estimating breeding dates of mule deer from fetal age-length relationship. J. Wildl. Manage. This manuscript was not accepted for publication. 6. Bartmann, R. M. 1984. Winter severity and mule deer mortality in Colorado. Wildl. Soc. Bull. This manuscript will be published in the Wildl. Soc. Bull. 12(3). 7. Bear, G. D. 1984. An expanding telemetry collar for elk calves. Outdoor Fact Sheet, Colo. Div. of Wildl. This manuscript is still in the review process. 8. Bear, G. D. 1984. Seasonal distribution and population dynamics of the Estes Valley. Colorado. elk herd. Colo. Div. of Wildl. Tech. Rep. This manuscript is still in the review process. 9. Bear, G. D .• and G. Wagner. 1984.· Mark-recapture method appl ied to elk population estimates. J. Wildl. Manage. This manuscript is still in the review process. 10. Beck, T. D. I. 1984. Black bear harvest analyses for Colorado, 1979-82. Colo. Div; of Wildl. Spec. Rep. No progress was made on this manuscript. 11. Beck, T. D. 1., and M. A. Haroldson. 1984. Estimation of weight in Colorado black bears from chest and neck girth measurements. Southwest Nat. No progress was made on this manuscript. 12. Carpenter, L. H., R. B. Gill, and D. L. Baker. 1984. Tests of nutritionally based habitat evaluation system. Colo. Div. of Wildl. Rep. No progress was made on this manuscript. 13. Daney, T. V., and N. T. Hobbs. 1984. Niche separation and feeding ecology of mountain goats and mountain sheep in Colorado. J. Wild1. Manage. This manuscript will be published in J. Wildl. Manage. 48(3). 14. Freddy, D. J. 1983. A "Butte" of an investment. Colo. Outdoors. This manuscript was published in Colo. Outdoors 32(5):26-28. �7 15. Freddy, o. J., R. M. Bartmann, L. H. Carpenter, and R. B. Gill. 1984. Measuring mule deer sex-and-age ratios in juniper-pinyon woodland. Wi1dl. Soc. Bull. No progress was made on this manuscript. 16. Freddy, D. J., W. M. Bronaugh, and M. C. Fowler. 1984. Reactions of mule deer to human harassment during winter. Wildl. Soc. Bull. This manuscript will be submitted in September, 1984. 17. Freddy, D. J., and K. Karrow. 1984. western Colorado. J. Mammal. Breeding white-tailed deer in This manuscript was published as: Karrow, K. K., and D. J. Freddy. 1984. Mountain whitetails. Colo. Outdoors 33(1):10-11. 18. Freddy, D. J., and K. K. Karrow. 1984. Fidelity of mule deer to seasonal ranges. Colo. ·Div. of Wildl. Rep. No progress was made on this manuscript. 19. Green, R. L., and G. D. Bear. '1984. Daily activity patterns of elk in north central Colorado as related to vegetation types. J. Wildl. Manage. No progress was made on this manuscript. 20. Haroldson, M., and T. D. 1. Beck. black bears. Southwest Nat. 1984. Extensive fall movements of No progress was made on this manuscript. 21. Haroldson, M., and T. D. 1. Beck. 1984. Habitat use by female black bears in Colorado. J. Wi1dl. Manage. No progress was made on this manuscript. 22. Hobbs, N. T., T. E. Remington, and R. Sayre. 1984. Effects of sagebrush on in vitro digestion of grass cell wall. J. Range Manage. This manuscript will be published as: Hobbs, N. T., B. L. Welch, T. E. Remington, and R. Sayre. 1985. Wild1. Shrub Symp: The biology of Artemisia and Chrysothamnus. USFS Intermt. For. Range Exp. Sta., Miscell. Publ. 23. Hobbs, N. T., and D. S. Schimel. 1984. Fire effects on nitrogen mineralization and fixation in mountain shrub and grassland communities. J. Range Manage. This manuscript has been accepted in J. Range Manage. 37(6). 24. Hobbs, N. T., and R. A. Spowart. 1984. Effects of prescribed fire on nutrition of mountain sheep and mule.deer during winter and spring. J. Wildl. Manage. �8 This manuscript has been published in J. Wildl. Manage. 48(2): 551-560. 25. Hobbs , N. T.s and R. A. Spowart. 1984. Fire effects on the distribut-ion of nitrogen and digestible organic matter in herbage. J. Range Manage. This manuscript will be published by the J. Wiidl. Manage as: Hobbs, N. T .• and D. M. Swift. 1985. Incorporating explicit nutritional constraints. in estimates of carrying capacity. 49 26. Kufe1d R. C. 1984. Effects of burning, spraying and anchor chaining of elk, mule deer and cattle use and vegetative yields of Gambe1' oak range land. Colo. Div. of Wild1. Spec. Rep. This manuscript was published as: Kufeld, R. C. 1983. Responses of elk, mule deer, cattle, and vegetation to burning, spraying. and chaining of Gambel oak rangeland. Colo. Div. Wild1. Tech. Pub1. No. 24. 47pp. 27. Lance, W. R., and T. M. Pojar. 1984. Diseases of pronghorn antelope. A literature review. Colo. Div. of Wildl. Rep. This manuscript will be published as: Diseases and parasites of pronghorn; a review. Colo. Div. of Wildl. Spec. Rep. No. 57. 28. Miller, M. W., and N. T. Hobbs. 1984. Efficacy of invermectin in treatment of 1ungworm in bighorn sheep. J. Wi!dl . Diseases. The first draft of this manuscript has been completed. ,29. Nelson, R., and T. D. I. Beck. 1984. Natural history of season changes in behavior and biochemistry in wild bears. Science. This manuscript will be published as: Nelson, R., T. D. I. Beck, and D. L. Steiger. 1984. Serum Urea-creatinine ratios in wild black bears. Science. 30. Pojar, T. M. 1984. Hematological indicators of stress in pronghorn. J. Wildl. Manage. The first draft of this manuscript has been completed. 31. Pojar, T. M., and D. Nash. J. Wildl. Manage. 1984. Genetic variation in pronghorn. No progress was made on this manuscript. 32. Reed, D. F. 1984. Highways, deer fencing and associate structures specifications, maintenance. and effectiveness. Colo. Dep. of Highways or Colo. Div. of Wildl. Spec. Rep. The first draft of this manuscript has been completed and will appear as a highway fi na1 rep. 33. Reed, D. F. 1984. Introduced mountain goats: interactions with indigenous alpine species. Proc. Northern Wildl. Sheep and Mountain Counc. �9 34. Spowart, R. A., and N. T. Hobbs. 1984. Fire effects on feeding ecology of mule deer and mountain sheep. J. Wi1d1. Manage. This manuscript has been submitted to J. Wi1dl. Manage. as: Fire effects on diet overlap of mule deer and mountain sheep. 35. Spowart, R. A., N. T. Hobbs, and D. M. Swift. 1984. Modeling fire effects on energy and nitrogen balance of ungulates during winter. Ecological Modeling. . The first draft of this manuscript has been prepared. 36. Ti~dema~J. A., R. E. Francis, C. Terwilliger, Jr., and L. H. Carpenter. 1984. Steppe habitat types of Middle Park, Colorado. Colo. Div. of Wildl. Rep. This manuscript is still in the review process. 37. Torbit, S., L. H. Carpenter, -A. W. Alldredge, and D. M. Swift. 1984. Mule deer body compos-i t ion - A comparison of methods. J. Wi 1d1. Manage. This manuscript will be ,published by J. Wild1. Manage. 48(4): . 38. Torbit, S.~ L. H. Carpenter, D. M. Swift, and A. W. Alldredge. 1984. Winter dynamics of mule deer body composition. J. Wild1. Manage. This manuscript will be publ,ished as: Torbit, S. C., L. H. Carpenter, D. M. Swift, and A. W. Alldredge. 1984. Differential loss of fat and protein by mule deer during winter. J. Wild1. Manage. 48(4): Additional publications which were not scheduled but which were published or accepted for publication during the 1983-84 Segment are listed below: Bartmann, R. M. 1983. Appraisal of a quadrat census for mule deer in pinyon-juniper vegetation. Outdoor Facts, Game Infor. Leafl. No. 109. Colo. Div. of Wi1dl. Gill, R. B., L. H. Carpenter, and D. C. Bowden. 1983. Monitoring large animal populations: the Colorado experience. Trans. N. Amer. Wildl. Conf. 48:330-341. Kufe1d, R. C. 1983. Preferred deer habitat: 32(5):24-25. What is it? Colo. Outdoors. Nelson, R., and T. D. I. Beck. 1984. Hibernation adaptation in the black bear: Implications for management. 7th East. Black Bear Workshop. Homossala, FL. Nelson, R.~ D. L. Steiger, and T. D. I. Beck. 1984. Physiology and biochemistry of hibernation in the bear. Sixth Int'l. Conf. Bear Res. and Manage. Grand Canyon, AZ. �10 White, G. C., and R. A. Garrott. 1984. Portable computer system for field processing biotelemetry triangulation data. Outdoor Facts, Game Infor. Leafl. No. 110. Colo. Div. of Wildl. Prepared by ---;--_~~-=--I/~, ~--:~-{..f-~---'----Len H. Carpenter Wildlife Research Leader �11 Colorado Division of Wildlife Wildlife Research Report July 1984 JOB PROGRESS REPORT State of Colorado Project No. 45-01-502-15050 Big Game Investigations - Cervids Work Plan No. Multispecies Investigations Job. No. 1 ---------------- 8 ---------- Period covered: Author: __ ------ < . Big Game Publication Editing and Library Services 7/1/83-6/30/84 M. Hershcopf and L. Carpenter Personnel: R. B. Gill, N. McEwen, A. Moreki1l ABSTRACT During the 1983-84 Segment. 14 books were purchased for permanent reference by DOW personnel. Thirty-two additional publications were located, ordered, and obtained free of charge for use. Twenty-two theses were purchased, obtained on interlibrary loan or given to the ltbrary. An additional 985 individual references requested by Big Game Researchers were located by library staff and made available for reference. About 40 of these requests were not available locally and were obtained through interlibrary loan. ��---", 13 BIG GAME PUBLICATION EDITING AND LIBRARY SERVICES Marian W. Hershcopf and Len H. Carpenter , P N. OBJECHVE t To provide a centralized support program for big game research technical editing and library services so that Big Game Research Scientists can allocate additional time to the conduct uf actual research. SEGMENT OBJECTIVES To provide coordinated, efficient, and economic editing and library services to all Colorado Big Game Research programs (Federal Aid Projects 45-01-502-15050 and 45-01-503-15050). SUMMARY OF SERVICES Publications purchased with 45-01~502~15050 and 45-01-503-15050 Funds and placed in Research Center Library Chadwick. D. H. 1983. A beast the color of winter: the mountain goat observed. Sierra Club Books, San Francisco. 208 p. Crawley , M. J. 1983. Herbivory; the dynamics of animal-plant interactions. Un iver-s lty of California Press, Los Angeles. 437 p. Goss, Richard J. 1983. Deer antlers: regeneration. function, and evolution. Academic Press, Inc., New York. 316 p. Gray, Peter. 1982. The dictionary of the biological sciences. Krieger Publishing Co., r~alabar,-FL. 602 p , Robert E. Jones, J. K. Jr., et al. 1983. ~lammals of the northern great plains. University of Nebraska Press, Lincoln. 379 p. 9 Kiddy. C. A. and H. D. Hafs, eds. 1971. Sex ratio at birth-prospects for control; a symposium, July 31 and August 1,1970 at Pennsylvania State University, University Park, PA. American Society of Animal Science. 104 p. Kurten, Bjorn.' 1971. The age of mammals. York. 250 p. Columbia University Press, New National Research Council. Committee on Animal Nutrition. Subcommittee on Feed Composition. 1982. United States-Canadian tables of feed composition; nutritional data for United States and Canadian feeds. 3rd review National Academy of Sciences, Washington, D.C. 148 p. �14 Orskov, E. R. '1982. Protein nutrition in ruminants. Academic Press, New York. 160 p. Robbins, C. T. 343 p. 1983. Wildlife feeding and nutrition. Academic Press, New York. Van Soest, P. J. 1982. Nutritional ecology of the ruminant. Ruminant metabolism, nutritional strategies, the cellulolytic fermentation and the chemistry of forages and plant fibers. 0 & B Bookss Inc. Corvallis, OR. 374 p. Walther, F. R. 1984. Communication and expression in hoofed mammals. University Press, Bloomington. 423 p. Indiana Weed Science Society of America. 1983. Herbicide Handbook of the weed science society of America. Weed Science Society of America, Champaign. 515 p. Whitaker, J. O. 1980. Tha Audubon Society field guide to North American mammals. Alfred A. Knopf, New York. 742 p. Publications obtained free or at lo~ cost In addition to books purchased with Federal Aid Funds, about 32 free reports and short publications from state or federal agencies or from private sources, were located, ordered and obtained for use by Big Game Research personnel. Theses purchased, obtained on interlibrary loan or as gifts for use by res_e_a_r_c_h_e_r_s _ Barber, K. R. 1983. Use of clear-cut habitats by black bear in Pacific Northwest. , M.S. Thesis, Utah State Univ., Logan. 169 p. Barlow. J. P. 1982. Methods and applications in estimating mortality and other vital rates. Ph.D. Dissertation, Univ. of California, San Diego. 194 p. Carr, P. 1983. Habitat utilization and seasonal movements of black bears in the Great Smokey Mountains National Park. M.S. Thesis~ Univ. of Tennessee, Knoxville. 95 p. Davis, J. L. 1970. Elk use of spring and calving range during and after controlled logging. M.S. Thesis, Univ. of Idaho, Moscow. 51 p. Hughes. B. A. 1978. Factors affecting wildlife mitigation choices in the oil shale region. M.S. Thesis, Colo. State Univ., Fort Collins. 199 p. Knight, J. E., Jr. 1980. Effect of hydrocarbon development on elk movements ,and distribution in northern Michigan. Ph.D. Dissertation, Univ. of Michigan, Ann Arbor. 79 p. Lentz, W. M. 1980. Aspects of habitat and denning requirements of black bear in northeastern Georgia. M.S. Thesis, Univ. of Georgia, Athens. 82 p. �15 Lloyd, Kevin A. 1979. Aspects of the ecology of black and grizzly bears in coastal British Columbia. M.S. Thesis, Univ. British Columbia, Vancouver. 151 p. Manv i 11e, A. M. 1982. Human impact on the black bear in Michigan's lower peninsula. Ph.D. Dissertation, Michigan State Univ., East Lansing. 188 p. McKell, C. M. 1950. A study of plant succession in the oak brush (Quercus gambelii) zone after fire. ~.S. Thesis, Univ. of Utah, Salt Lake City. 79 p. Mould, E. D. 1980. Aspects of elk (Cervus canadensis nelsoni) nutrition and associated analytical procedures. Ph.D. Dissertation, Washington State Univ., Pullman. 72 p. Mubanga, G. 1983. Use of fecal indices to predict forage quality and intake of mule deer. M.S. Thesis, New Mexico State Univ., Las Cruces. 44 p. Parker, K. L. 1983. Ecological energetics of mule deer and elk; locomotion and thermoregulation. Ph.D. Dissertation, Washington State Univ., Pullman. 128 p. Rominger, E. M. 1983. Bighorn sheep food habits and gambel oak manipulation, Waterton Canyon, Colorado. M.S. Thesis, Colo. State Univ., Fort Collins. 125 p. I ) Schleyer, B. O. 1983. Activity patterns of grizzly bears in the Yellowstone ecosystem and their reproductive behavior, predation and the use of carrion. M.S. Thesis, Montana State Univ .• Bozeman. 130 p. Sejkora, K. J. 1982. Polonium assi~ilation and retention in mule deer and pronghorn antelope. M.S. Thesis, Colo. State Univ., Fort Collins. 36 p. Severson, K. E. 1964. A description and classification, by composition, of aspen stands in the Sierra Madre Mountains, Wyoming. M.S. Thesis, Univ. of Wyoming, Laramie. 94 p. Sizemore, D. L. 1980. Foraging strategies of the grizzly bear as related to its ecological energetics. M.S. Thesis, Univ. of Montana, Missoula. 67 p. Teeter, R. G. 1981. Indigestible markers: methodology and applications in ruminant nutrition. Ph.D. Dissertation, Oklahoma State Univ., Stillwater. 160 p. Williams, K. R. 1972. The relationship of soil temperature and cytokinin production in aspen invasion. M.S. Thesis, Univ. of N.M., Albuquerque. 39 p. Wright, K. L. 19 Elk movements, habitat use, and the effects of hunting activity on elk behavior near Gunnison, Colorado. M.S, Thesis, Colo, State Univ., Fort Collins. 206 p. Yarrow, G. K. 1979. Comparison of tame and wild deer food habits. M.S. Thesis, Mississippi State Univ., State College. 82 p. �16 Reference document location and delivery The Research Center Library staff also located and delivered about 985 individual articles on request for the Big Game Research Section during this segment; about 40 were not available locally and were obtained through interlibrary loan procedures. Prepared by ('i~C-dA tJ. H~-st-..CoFf;_ Marian W. Hershcopf ~ Librarian Len H. Carpenter Wildlife Research Leader �17 Colorado Division of Wildlife Wildlife Research Report July, 1984 JOB PROGRESS REPORT State of Colorado Project No. 45-01-502-15050 . . "'- Big Game Investigations - Cervids Work Plan No. 1 Multispecies Investigations Job No. 9 Cervid Research Administration Period covered: Author: 7/1/83-6/30/84 L. Carpenter Personnel: R. B. Gill, L. Lovett, and N. McEwen ABSTRACT Employee evaluations, completion of Program Plans, and development of the 1984-85 budget highlighted Cervid Research Administration during the 1983-84 Segment. ��.1:.1 CERVID RESEARCH ADMINISTRATION Len H. Carpenter P. N. OBJECTIVE To supervise and administer research on deer and elk in Project 45-01-502-15050. SEGMENT OBJECTIVE Supervise and administer all deer and elk research studies in Project 45-01-502-15050. METHODS AND MATERIALS These have been described previously (Carpenter 1983). RESULTS AND DISCUSSION The Wildlife Research Leader was assigned to the big game feeding evaluation in January and continued until segment end. This assignment seriously impacted the Leader1s time for administering Cervid Research. Consequently, only the routine duties of employee evaluation, planning, supervision, and budgeting were completed. LITERATURE CITED Carpenters L. H. 1983. Cervid Research Administration. ~~ildl. Game Res. Rep .• July, Part 1. 25-26. Prepared by Len H. Carpenter Wildlife Research Leader Colo. Div. ��21 Colorado Division of Wildlife Wildlife Research Report July 1984 JOB PROGRESS REPORT State of Colorado Project No. 45-01-502-15050 Big Game Investigations - Cervids Work Plan No. Deer Investigations 2 Job No. Quantifying Capacity of Winter Ranges to Support Deer--Evaluation of Thermal Cover Used by Deer Period covered: Author: 7/1/83-6/30/84 D. J. Freddy Personnel: E. M. Rominger, D. C. Bowden ABSTRACT Comparative changes in body weight, intake of digestible energy (DE), and behavior were monitored for 6 mule deer having, and 6 mule deer not navinq, access to thermal cover between 9 December 1983 and 17 March 1984. Neither changes in body weight nor intake of DE indicated that thermal cover provided an energetic advantage. Final conclusions await comparisons of behavioral responses of cover and no-cover deer. ��23 NUTRITIONAL BASIS FOR QUANTIFYING CAPACITY OF WINTER RANGES TO SUPPORT DEER David J. Freddy P. N. OBJECTIVE To determine if a system can be developed to estimate number of deer winter ranges are capable of supporting. SEGMENT OBJECTIVES 1. ~Complete data summary and analysis of 1982-83 cover experiment and summarize results in manuscript format. 2. Compare body weight loss, intake, and behavior in 2 groups of mule deer having different access to thermal cover during winter. ACKNOWLEDGMENTS The U.S. Forest Service~ Rocky Mountain Forest and Range Experiment Station, kindly loaned instrumentation for measuring weather conditions. Eric M. Rominger provided diligent assistance in collecting and summarizing data and Lynn L. Stevens provided timely laboratory analyses of rations fed to deer. METHODS AND MATERIALS See Program Narrative (Appendix A). RESULTS AND DISCUSSION 1982-83 Experiment Results to date for the cover experiment in 1982-83 have been summarized in a manuscript. However, several more analyses are planned and, therefore, this manuscript is not complete. 1983-84 Exoeriment Effects of the presence and absence of thermal cover on deer behavior, weight performance, and intake of pelleted ration were monitored for 100 days from 9 December 1983 - 17 March 1984. Behavior of 12 deer, of which 6 had access to thermal cover, was monitored during 6, 6-day, 24-hour trials conducted about every 2 weeks. Intakes of pelleted ration and changes in body weight were measured every 3 and 17 days, respectively. Average intake and percent change in body weight were determined for each deer for e ch of the 6, 17-day time periods. �24 Weather conditions were severe as snow and cold temperatures persisted for most of the experimental period and high winds often occurred. Deer were probab Iy metabo 1ica lly stressed duri ng 4 of 6 tri a 1s assumi ng a lower critical temperature of -23C (Mautz et al. 19849 in press)(Tables 1 ,2) . Deer again adjusted well to isolation pens, especially since the animals were confined for 100 continuous days (Fig. 1). t~inimal nervous behavior was exhibited, even by deer not having access to cover structures. Pe 11 eted Rat ions Deer were maintained on 3 different pelleted rations from October, 1983, through April, 1984 (Table 3). During the 'IOO-day experiment , two rations were fed: low energy-I with 64% in vitro di qes t tble dry matter (IVDDM) and 17% prote in was fed from 9-26 December 1983, and low energy-II with 59% IVOOM and 13% protein was fed from 27 December 1983 - n March 1984 (Tab'le 3). Af ter the exper iment , deer were maintained on the low energyII ration through 29 April. Fawns were supplemented with 50 g of alfalfa leaves mixed into the pelleted ration evet'y 3 days from 23 January - 29 April because both fawns began chewing and clipping hair on their ribs and flanks. Although hair removal subsided after supplementation, it was not totally averted. Whether thi s behavi or by fawns was due to the reduced protet n and energy levels in the Iow energy II ration is not known. Adult deer had no apparent problems with the rations. Snow was available to deer as a source of water from 29 November April. 1984. Water was provided to deer from 20-29 April. 1983 - Ch~!I~s .In.BqQ,LVJeLghts There was no difference in wei ght gain or loss between cover and no-cover deer over the lOO-day period {P > O.5d~ Fig. 2, Tables 4,5}. Percent weight loss was 6.2 ± 0.7 (SE)-for adult deer having cover and 7.1 ± 3.4 (SE) for adults not having cover. Both fawns gained weight (2-10%) with the no-cover fawn gaining the most weight. From 9 December 1983 - 27 January 1984 all deer exhibited some periods of wei ght ga -i n , but there was no difference between cover and no-cover deer (P > O.50~ Table 5). However, in 4 of 5 pairs of adult deer, deer with cover either ga-ined more or lost less weight than their paired counterparts. From 28 January - 17 March all adult deer lost weight but there was no difference between cover and no-cover deer (P > 0,50, Table 5). Inter-est inq ly , from 26 December 1983 .. lI January 1984~ deer with cover lost less weight (P < 0.02, Tables 4.5). This is the only time interval where differences in weight performance occurred and it is possible that this time interval contained the highest average wind speeds of the 6, 17-day periods. The data suggest, then, that cover had minimal effect on changes in body weight over the lOa-day period, but the pos sib il i ty remai ns that cover did provide a measurable energetic advantage during shorter time intervals. I f �25 Body weights of 4 deer common to experiments in 1982-83 and 1983-84 generally declined similarly (Table 6). All 4 deer had access to cover in 1982-83 but only 1 deer had cover in 1983-84. These weight performances also suggest that deer were' minimally effected by the absence of cover. even though ambient conditions were more severe in 1983-84. Of interest also is the effect of different pelleted rations on changes in body weight. In 1982-83p IVDDM of the ration was 74% and protein content was 18%~ whereas the ration in 1983-84 was 59% IVDDM and 13% protein. In spite of the reduced quality of the ration, these 4 deer did as well or better on the poorer quality ration. Intakes of Digestible Energy During the Experiment Intakes of digestible energy (DE, Kcal/kg BWO.75/day) for cover and no-cover deer were not different for the lOa-day period or within several shorter time intervals (P > 0.20, Table 5). Differences in intake within pairs of deer did occur Tp < 0.10, paired t-test, Tab1e 7) but showed no cons istent re1at i on to the presence or absence of cover. In the 4 cases where intakes differed within· pairs of deer for the lOa-day period (p < 0.02), higher intakes were associated twice with both cover and no-cover deer. The inability of the measurements of intake and body weight to reveal advantages of cover suggests that the artificial cover did not provide an energetic advantage or that both parameters are' insensitive to measuring net energy flow in deer. Estimated daily intake of DE voluntarily declined for each deer during the lOa-day period (r > 0.79) P < 0~06). Overall, intake declined from 216 to 135 kca1/kg BWO~75/day with declines averaging 35% for fawns and 38% for adults (Table 7). Percent reductions in intakes for cover deer .averaged 36.0 ± 3.1 (SE) and for no-cover deer 38.5 ± 3.9 (SE). Intake of DE was above maintenance for fawns (180 kcal/kg BWO.75/day Baker et a1. 1979) for most of the experimental period wh'ile adults were below maintenance (160 kcal/kg BWO.75/day, Ul1rey et ale 1969) from late January through the end of the experiment in mid-March (Fig. 3). It is not surprising, therefore, that fawns gained or at least maintained. body weight while adults lost body weight. Several analyses relating intakes to ambient conditions await completion. There is the possibility that when interactions between individual daily intakes and ambient conditions are examined that differences between cover and no-cover deer will emerge. Seasonal_ Intakes of Digestible Energy Intake of DE by fawns and adults voluntarily declined from December into March and April, respectively, and were below maintenance for adults from 28 January - 12 April and for fawns only from 1-18 March (Fig. 3). Intakes of fawns and adults increased during late March and late April, respectively (Fig. 3). The changes in intakes for adults corresponded with declines and increases in resting hear~ rates of a tame adult female mule deer during winter and spring (Freddy 1984) and presumably reflect seasonal changes in metabolic rates, �26 Unfortunately. intakes of fawns increased during the time interval immediately following the relocation of all deer to nutrition isolation pens adjacent to the pasture where the cover experiment was conducted and intakes of adults increased when all deer had access to water instead of snow. Ullrey et al. (1969) indicated there was a direct relationship between amounts of feed and water consumed by white-tailed deer. These 2 factors~ and not intrinsic changes in metabolic rates, could have been responsible for the increase in intakes. Behavioral Patterns and Cover Use Behavioral patterns of cover and no-cover deer and the use of cover by cover deer have not been analyzed as all data have not been placed on computer. CONCLUSIONS Ne-ither changes in body we iqht nor intake of digestible energy indicated that cover provided an energetic advantage for deer during winter. Fur-ther exper-imerrtat lon may involve feeding deer an extremely low quality pel1eted ration and altering cover structures to maximize the effect of cover. Under these extremes of energy input and outfl ow, an energeti c advantage of cover may be measurable. Final conclusions from work in 1983-84 await analyses comparing behavioral responses of cover and no-cover deer. LITERATURE CITED Baker, D. L., D. E. Johnson, L. H. Carpenter, O. C. Wallmo, and R. B. Gi 11 . 19'79. Energy requirements of mule deer fawns duri ng wi nter. J. Wildl. Manage. 43:162-169. Freddy. D. J. 1984. Heart rates for activities of mule deer at pasture , J. Wildl. Manage. 48: in press. Mautz, W. W., P. J. Pekins, and J. A. Warren. 1984. Cold temperature effects on metabolic rate of white-tailed, mule, and black-tailed deer in winter coat. Proc. Int. Conf. on Biology of Deer Production. Dunedin, New Zealand. In press. Ul1rey, D. E., W. G. Youatt, H. E. Johnson, L. D. Fay, B. L. Schoepke. and i~.T. Magee. 1969. Digestible energy requirements for winter maintenance of Michigan white-tailed does. J. Wildl. Manage 33:482-490. Prepared �27 Table 1. Trial 1 Experimental Dates conditions Conditions 13 Dec 18 Dec 1983 100% snow coverage in all pens; high winds with blowing snow 3 Jan - 2 100% snow coverage in all pens; cloudy days, foggy· nights; reduced solar radiation 8 ,Jan 1984 16 Jan - 3 for cover trials in 1983-84. 100% snow coverage in all pens; coldest temperatures; minimal wind 21 Jan 1984 4 1 Feb 6 Feb 1984 100% snow coverage in all pens; strong chinook wind 1 night; intense solar radiation; some fog at night; snow melting 5 17 Feb 22 Feb 1984 100% snow coverage in all pens; periods of high winds; intense solar radiation; snow melting 6 7 Mar 12 Mar 1984 100% snow coverage in all pens; periods of high winds; snow melting; warmest temperatures Table 2. Tria 1 1 2 3 4 5 6 for cover trials in 1983-84. Tempera ture C -------Ambient Black Globe tvli n Max Min Max Ambient conditions v Dates Dec Dec Jan Jan Jan Jan 1 Feb 6 Feb 17 Feb 22 Feb A SE SE X SE X SE X SE -15 1 17 2 184 17 -22 2 14 3 208 18 -16 -30 1 11 1 258 20 2 -5 -24 2 19 2 338 6 -20 2 -3 -23 2 23 2 384 25 -10 1 2 -11 1 28 4 428 36 1983 -13 1 0 1984 ~ 1984 1984 -21 2 -8 -30 1 -22 1984 7 Mar 12 Mar 1984 13 18 3 8 16 21 X Total Solar Radiation cal/cm2/day 2 1 �Table 3. Characteristics April 1984. and ingredients of 3 pelleted rations and alfalfa leaves fed to mule deer October 1983 Ration Special Mix - fed 13 Oct - 21 Nov 1983 Ration Characteristics Ingredient Wilea. t mi dd 1 i ngs Brewers dry gra i n Cottonseed hulls Suncured alfalfa pellets Dehydrated alfalfa pellets Corn starch Molasses Vitamin A,D,E mix Trace rni nera 1 In vitro DOM GE Kcal/g DE Kcal/g Protein Dry matter AOF NOF Lignin Low Energy-I - fed 22 Nov - 26 Dec 1983 % Inqredi ent Cottonseed hulls Suncured alfalfa ~Jhole corn ~!ho1e wheat Sunflower meal 7.0 Brewers dry gi~a in Corn s ta rch 5.0 Molasses 3.0 Biophosphate 2.0 0.002 Calcium phosphorous 0.005 Vitamin A,D.E mix 43.0 25.0 15.0 73.60 4.55 3.37 18.00 94.60 20.60 32.70 5.20 % 35.0 25.0 12. ~l 8.0 7.9 4.1 3.7 2.0 1.3 Low Energy-II - fed 27 Dec 83 - 29 Apr 84 Ingredient Cottonseed hulls Suncured alfalfa ~~ho1e corn ~~ho1e whea t ~101asses Bentonite Biophosphate Caldum phosphate Vitamin AsD,E mix ot' ,0 47.0 23.3 .! 2.4 8.0 5.0 2.5 Alfalfa leavesa - fed 23 Jan - 29 Apr 1984 Ingredient % Alfalfa leaves 100 1.3 0.5 0.002 0.5 0.002 63.80 3.98 2.54 17.10 91.00 23.90 38.00 6.30 a50 g mixed wi th pel leted ration and fed every 3 days to f'awns only 58.90 4.08 2.40 12.60 86.70 25.20 38.90 7.00 69.80 4.47 3. 12 24.10 91.50 19.30 32.70 4.10 N 00 �- -- - ...•. ------- Table 4. Changes (%) in body weight for mule deer for the entire 100-day cover experiment and for 6, 17-day time intervals within the 100-day experiment. 1983-84. Deer are listed in order of experimental ~irs subject to cover (c) and no-cover (nc) treatments. Deer Time Interval 177c 176nc 162c 168nc 150c 191nc 153c 156nc 154c 194"C 144c 203nc Dates 9 Dec 1983 17 Mar 1984 -5.0 4.3 -7.3 -2.9 2.2 -0.2 4.3 2.6 1.0 1.2 1.5 -0.8 -3.1 0.3 -0.6 -0.9 -0.7 -2.2 -5.4 0.6 2.0 -1.7 0.8 3.6 0.4 -1.3 -0.6 -1. 6 -1.9 -6.4 -5.0 -3.9 -0.8 -1.1 -5.1 -1.3 -3.0 -3.6 -4.7 -2.5 -1.1 -2.8 -3.4 -1.0 -0.4 -2.2 -'1.3 -1.8 -2.8 5.5 0.4 -3.7 -1.4 -2.4 -4.2 -1.3 -2.2 0.8 -0.2 -2.6 0.8 9.5 -1.5 7.6 2.3 3.0 0.0 26 Dec 1983 10 Jan 1984 3.8 0.0 -0.4 -4.1 11 Jan 27 Jan 1984 1.4 -1.8 -2.1 28 Jan 12 Feb 1984 -3.9 3.1 13 Feb 29 Feb 1984 1.5 1 Mar 17 Mar 1984 1.1 9 Dec 25 Dec 1983 -4.2 -11.4 -7.9 -11.8 -6.5 -13.7 aAges of deer were 6 mos (176,177), 18 mos (162,168,150,153.154,156), and >42 mas (144,191.194,203). I" \.{ �30 Table 5. Results of l-sample t-tests comparing average differences in percent weight gain or loss and intake of pel1eted ration between cover and no-cover deer during several time intervals, 1983-84. Time Interval Dates Body weight Intake t-value P-1evel t-value P-leve1 9 Dec 1983 17 Mar 1984 0.09 P >0.50 -1. 11 P >0.20 9 Dec 1983 27 Jan 1984 -0.48 P >0.50 -0.71 P >0.50 28 Jan 17 Mar 1984 0.32 P >0.50 -1.34 P >0.20 9 Dec _. -1.63 P >0.10 -0.96 P >0.30 26 Dec 1983 10 Jan 1984 ·-3.43 P <0.02 -0.34 P >0.50 11 Jan 27 Jan 1984 -0.59 P >0.50 -0.52 P >0.50 28 Jan .. 12 Feb 1984 0.67 P·>0.50 -1. 16 P >0.20 13 Feb 29 Feb 1984 -0.72 P >0.50 -1.35 P >0.20 1 Mar 17 Mar 1984 0.34 P >0.50 -1.27 P >0.20 25 Dec 1983 Table 6. Percent weight loss in 4 adult mule deer (2 M, 2 F) common to ~ove~~eriments in 1982-83 and 1983-84. Percent weight loss Time interval Deer 10 Dec 82 - 20 Mar 83 203 M 144 M -13.9C 194 F -lO.Oc 191 F -12.5c c = access to cover nc = no access to cover -7.1 c Ti me i nterva 1 9 Dec 83 - 18 Mar 84 _13.7nc -6.5c _2.9nc _11.8nc �--- - .•...•. -.._, -- Table 7. Ad libitum average daily intakes of in vitro digestible energy (dry matter basis, Kcal/kg BWO.75/day) for mule deer on pelleted ration during 8 time intervals beginning 9 December 1983 and ending 29 April 1984. Intakes determined over 3-day intervals (n) and converted to daily intakes. Beginning dates (M/D/Y) of each time interval are shown and deer are listed in order of ex~erimental eairs. Interval Deerb Average Time a 177c 176nc 203nc (All deer) Interva 1 162c 16Snc 150c 191nc 153c 156nc 154c 194nc 144c 1 12/_9/83 x SE N 229.4 * 260.0 8.2 7.6 5 5 241.6 18.9 5 233.5 9.7 5 259.5 18.3 5 178.1 9.9 5 201.2 20.5 234.6 11.4 6 256.5 17.5 6 189.6 7.6 6 209.5 15.2 6 176.4 29.9 6 143.7 6.3 220.5 205.1 10.3 147.0 * 181.1 8.7 6.4 6 6 163.8 32.0 6 138.3 211.4 * 202.7 2.4 2.5 5 5 165.2 * 177.2 4.8 6.3 5 5 216.1 * 178.4 7.6 9.0 6 6 151.1 7.0 6 207.5 6.0 5 187.5 7.6 5 205.1 * 171.5 16.4 7.3 5 5 216 5 219.2 18.4. 5 162.2 7.2 6 176.6 11.4 6 184.8 8.9 6 168.7 3.9 6 166.3 * 139.9 7.6 7.1 6 6 184 176.4 iO.O 6 193.2 7.0 176.3 * 161.1 6.8 5.9 6 6 150.9 * 124.3 5.9 4.3 6 6 170 175.9 * 106.5 13.1 3.2 5 5 153.7 8.6 5 ·165.8 10.8 5 155.3 10.0 150.6 3.9 5 140 .4 ~(11 8 2.1 6.1 5 5 160 151.7 * 115.1 16.5 2.3 6 6 147.4 * 159.8 4.7 5.8 6 6 129.2 * 140.3 2.5 2.8 141 .0 * 100.3 2A 5.0 6 6 149 2 12/26/83 x SE N 6 3 1/11/84 x SE N 3.0 6 6 6.5 6 6 4 1/28/84 x S N 5 °. 5 2/13/84 x SE N '155.5 3.7 6 6 6 w I-' �Tab 1e 7. 6 3/ _, /84 x SE N (continued 173 . 0 * 140 . 2 11. 5 5 15.2 - - - - - - - - - 7d 3/l8/84, SE 207,0 5. 1 N 8 v 1\ * w - ~age 2) 5 - - - - 183.8 3. 1 8 N 10.1 139.1 8.5 5 5 139.4 - - - - - - - 152.1 8.5 8 151.9 * 100.6 12.3 4.4 - - 5 8.4 7.8 5 120 . 7 8.2 * i 44 . 7 129.2 3.1 5 * 91.6 4.4 135 4.9 5 5 5 5 5 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 1 ~.t. 5 151 .8 G.4 ,.. 8.8 ,~ 150. 1 143.6 * 8 120.0 c ,.., 126. 1 128.9 0.,) 4,6 2.9 8 8 8 110 . 5 "~ 159.0 3 .....? 4.8 8 13tL 3 8 >'< 81.0 4.3 2.8 8 0 liB () 3d 4/]_2/84 243,2 * 2'10.2 8.2 4.7 x SE a v N 6 188.7 5.9 6 192.2 ',67.4 6.1 6 4.1 6 'k 155.8 o, r- 6 FJ.3.7 '1 i * 165,4 3.7 6c4 6 6 141.9 ~, 196.3 6e3 5.8 6 144.8 h u b"oes ," of deer were 6 mcs (176,177); . t erva Hi 18 mas r 1 i 7 , 2ro nays , • ; and • trrterva (162,168~150~153,154.156)~ 6 6 ------------------------ -------l"8 's -ib"" cays J .' *Means different (E <0.10, paired t-test). --- _--- ., tenqth , . H1 and >42 mcs (144,191~194.203). -Deer had access to cover during intervals 1-6. nc"[Jeer m c nor. nave access t 0 cover our ,.. i ng 11:t' r: terva ,lSi , -0. , ". deer P 'I aceo'. HI 150 . 1 at4o'i on pens prOV1Q'lng • •• •• 1 .dA',II s imi tar env t• rcnment.••at' conditions . ...I' 111 ,3 6.3 .. r nterva t s i- 6 were '6 I) or 'I 7' days; Ch. * 305 172 �33 .;--.. / :, "!~,)'.,,, ~ -r ! J A· " . Fig. 1. Deer bedded within a no-cover isolation pen (TOP) and deer lying on dark soil within a cover isolation pen (BOTTOM). �34 flO 80 ~ ~ •......• l- If) :r. - '0 ~ l&.J ~ ..•~ _-- >- so a ","" ..:..~ - - -6-- ~A~"" '0.- -6...,.•.•. 0 Q) I/O .,. S£P ", ' ""l I oc.r NOV DEC. JAN FEB MAY MONTH Fig. 2. Changes in body weight between September, 1983, and May, 1984~ for male ( ) and female (-----) mule deer having cover (solid symbols) and not having cover (open symbols). �35 2W ~ ~ FAW S 2'10 ~, ~ N e ~ ~ 220 200 lro - -- -.,..._ -- -- --- ....,..... ta'OlD ___ ~ _. \Ih :.':c: "-..J q U ~ ~ tu ~ ,"0 12D eso AOULTS -c l- fl'..oo <' H " I~() 160 I<l/O !UJ ~~=r 9 DEC ~~ Ot!C, ---,- i H JAN :ta J ill I.! I 18 FIi. MAlt NV\~ , l' 111. DATi! Fig. 3. Average daily intakes of in vitro digestible energy for 2 fawns (TOP) and 10 adults (BOTTOM) from 9 December 1983 _,29 April 1984. Values shown for adults are means ± SE. Dashed lines represent estimated maintenance intakes of digestible energy. A I( ��37 APPENDIX I PROGRAM NARRATIVE State of Colorado Project No. Work Plan 45-01-502-15050 ---, 2 Deer Investigations Job No. A~ Big Game Investigations - Cervids Nutritional Basis for Quantifying Capacity of Winter Range -Evaluation of Thennal Cover Used by Deer NEED Cover experiments conducted in 1982-83 revealed that tame mule deer adjusted well to experimental isolation pens and used artificial cover sign ificant1y more than non-cover control areas (Freddy 1983). 9 However 9 use of cover types did not appear predictable~ as differen- tial use of cover was primarily a function of individual deer choice. Weather conditions were mild throughout the 1982-83 winter and therefore there was quite possibly no strong driving force causing deer to use cover predictably. Our ability to discern preferential use of cover may be ltmited , but it is a concept that should not be abandoned. AdditionallY, the energetic advantage of thermal cover, if any. needs to be quantified. As stated by Peek et al. (1982) the need for thermal covey' by mule deer may be minimal. This concept must be explored. Hence, in this second year of cover experimentation, I have chosen to examine the effects of the presence and absence of cover on deer energetics throughout the winter (Dec - Mar). ConceptuallY9 deer having cover should have an energetic advantage over deer not having cover. �38 The sirnpl i s t measurement of energy balance in deer is the gain or 'loss of body weight. although body composition would Ii kely provide a more complete understanairrg nf animal condition (Torbit, 1981). The rate of body wei-ght loss determines the time interval-over which-a deer can survive a' negative energy balance. body wet ght, d-eath is- probable. Once a deer- reaches a critical Both food and cover conceptual ly influence the rate of energy outflow and therefore weight loss. Moen (1966) felt that the role of thermal cover was negligible pro- vided deer had access to high qual ity forage. Swift et a1. (1980) hypothesized that cover was also not necessary even when deer were on characteristically 10V'1 qual ity w-inter diets. The underlying basis for these hypotheses is that when deer have access to forage, the combination of insulative qualities of their pelage, behavioral adjustments to reduce activity and thermal stresss and heat of fermentation of low quality forage are sufficient to render cover a negligible role in deer energetics. To state that thermal cover is unimportant is grossly contrary to many habitat improvement and mitigation proqrams , Thus, there is a basic need to examine the role of thermal cover separately from the concepts of escape and security cover. B. OBJECTIVES The objective of this research will be to compare body weight loss or gain, food intake, and activity patterns between deer having access to thermal cover and deer not having thermal cover. This study is sequential to work completed during the 1982-83 winter. �39 C. EXPECTED RESULTS OR BENEFITS Completion of this r-esearchwill aid in developing a habitat evaluation system by establishing the role of cover in maintaining energy balance of deer during winter-.-"The habitat evaluation system will provide a .. quantitative basis for the Division of Wildlife to assess and improve winter ranges-used by deer. D. APPROACH The approach of this study will be to monitor the effects of the absence and presence of thermal cover on deer energetics during winter. There will be 2 groups of 6 tame deer. While being confined to individual isolation pens (Freddy, 1983), deer in one group will have access to thermal cover while deer in the second group will not have access to thermal cover. t I I Twelve isolation pens will be divided into 6 pairs of pens so that pens within pairs are as identical as possible (Fig. 1). Artificial cover structures (Freddy, 1983) will be randomly placed within 1 pen J of each pair. Twelve deer ranging in age from fawn (6 mos.) to adult (4+ yrs.) and in weight from 30 kg to 90 kg, will be grouped into 6 pairs. Pairing will be based on similar body weights, age, and sex (Table 1). One deer from each pair will be randomly assigned to be housed in an isolation pen with cover and the other deer of each pair to a pen without cover. Deer will be housed in pens during a 100-day period from 9 December 1983 through 18 March 1984. Three factors within each pair of deer will be measured: 1) body weight at 17-day intervals, 2) food intake in 3-day intervals for the entire 100 days, and 3) activity patterns, cover use by those �40 deer having cover, and postural positions when lying during 6 6-day 24-hour behavioral trials (Freddy, 1983) conducted at approximately l7~ day intervals. Rationale: Compairing physiological and behavioral responses . of deer havinq access to thermal cover with deer not having cover should provide a strong experimental test of the role of cover in deer energetics during winter. Conceptually. deer without cover may compensate by increasing food intake, reducing activity. and spending more time lying in an energy conserving posture. Further monitoring of cover used by those deer having access to covey' may allow more accurate pred ictions of covel" use when combined with data collected in 1982-83. Hypotheses. tha t wi 11 be tested: 1. Rate of body weight loss will not be different between • ( I IIcover" and "no-cover" deer. 2. Food intake will not be different between "cover" and IIno-coverlldeer. 3. Activity patterns and frequency of postural positions when lying wi 11 not be different between "cover" and "no-cover" deer. Additionally. for those deer having access to cover: 1. Deer will exhibit no preference for cover types. 2. Deer will choose cover types independent of time of day. 3. Deer will choose cover types independent of ambient conditions. 4. Deer will choose cover types independent of time of day (December - March). , �41 Maintenance of Experimental Deer Throughout the 100~day experimental period, deer will be fed ad libitum a deer-elk pelleted ration having 45-50% dry-matter digestibility. Intake of each deer will be determined over 3-day intervals throughout the 100-day period. each 3~day period. Orts will be weighed after Samples of arts will be obtained and frozen to 1ater correct for mo i sture content. Snow will be provided to assure each deer a source of water or water will be provided if snow is not available. Deer will be weighed at l7-day intervals using a platform scale. If deer reach a degree of body weight loss considered potentially lethal, deer may be removed from the experim~nt. For adult deer, body weight loss of 20-25% of the initial December body weight will be considered critical (Carpenter, 1979) and for'fawns, 15-20% of the initial December weight (Baker et al., 1979). Any deer having exces- sive hair loss due to ingestion (Carpenter, 1979) will be removed from the experiment. Deer reaching critical body weights and removed will be refed using a highly digestible pelleted ration mixed with the low quality ration at an incremental rate of 10% per day along with small amounts of alfalfa. Isolation Pens and Artificial Cover Isolation pens and artificial cover within those pens providing cover will be the same as in 1982-83. Pens without artificial cover will have all vegetation removed from within pens and for a 5-m area around pens. A small feeder located at the center of all pens will be the only structure within "na-cover" pens. Management of snow �42 within and around pens will follow procedures used in 1982-83 (see 1982-83 PROGRAM NARRATIVE) . The 6 pairs 0 f deer wi 11 be rotated through all 6 pairs of pens in a Latin Square design so that deer are in a different pen during each of the 6 behavioral trials. Deer will be placed in new pens at l7-day intervals. DATA COLLECTED Animal Behavior: Behavior of deer and their use of cover treatments will be monitored as in 1982-83. An exception will be that for deer without covet', deer will only be assigned to a pen quadrant and not to a treatment. Ambient Conditions: Measurements of ambient conditions will also be the same as in 1982-83. Exceptions will be that ambient temperature, solar radiation, and wind speed and direction will be measured throughout the lOO-day period (see 1982-83 PROGRAM NARRATIVE). DATA ANALYSIS Th is experi men t fo11ows a paired compa risons design. Pr irnary compa risons between pa irs of "cover" and "no-cover" deer will be: 1) percent change in body weight. 2) food intake in kg/kg BWO.75• 3) frequency of activity, primarily time spent lying and active, and 4) frequency of head-up and head-down lying postures. These variables can be com- pared within each l7-day interval and pooled over the entire lOO-day experimental period. For those deer having access to cover, analyses will be similar to 1982-83. This part of the experiment can be considered a randomized complete block design with deer treated as blocks (random effect) and each block subject to 4 cover treatments (fixed effect). Analyses �45 E. LOCATION Field work will be conducted at the Division of Wildlife Junction Butte Research Center located 5 kmsouth of Kremmling, Colorado. F. RELATED FEDERAL AID PROJECTS 45-01-502-15050, Work Plan 2, Jobs 6 and 10 45-01-503-15050 LITERATURE CITED Baker, D. L., D. E. Johnson, L. H. Carpenter, O. C. Wallmo, and R. B. Gill. 1979. Energy requirements of mule deer fawns during winter. J. Wildl. Manage. 43:162-169. Carpenter~ L. H. 1979. Colo. Div. Wild1. Game Res. Rep.~ July Part II. 69-74pp. Freddy, D. J. 1983. Evaluation of thermal cover used by deer. Colo. Div. Wildl. Game Res. Rep., July Part I. 31-68pp. Moen, A. N. 1966. Factors affecting the energy exchange and movements of white-tailed deer, western Minnesota. Minnesota. Ph.D. Thesis, Univ. of 121pp. Peek, J. M., M. D. Scott, L. J. Nelson, and D. John Pierce, and L. L. Irwin. 1982. Role of cover in habitat management for big game in northwestern United States. Trans. North Am. Wildl. Nat. Res. Conf. 47:363-373. Swift, D. M., J. E. Ellis, and N. T. Hobbs. 1980. Nitrogen and energy requirements of North American cervids in winter--a simulation study; Pages 244-251 in E. Reimers, E. Gaare, and S. Skjenneberg, eds. Proc. 2nd Int. Reindeer/Caribou Symp., Roros, Norway. �46 Torbit, S. C. 1981. In vivo estimation of mule deer body composition. Ph.D. Thesis, Colo. State Univ., Fort Collins. 98pp. , 1 �47 Table l. Deer Probable ~airings of deer for 1983-84 cover ex~eriments. Pai r Deer Sex Age Weighta 1 176 M 6 mas 36.0 177 M 6 mas 34.0 162 F 1 yr 54.0 168 F 1 yr 52.0 191 F 3 yrs 64.5 150 F 1 yr 61. 5 153 M 1 yr 65.5 156 M 1 yr 64.0 194 F 3 yrs 70.5 154 M 1 yr 74.5 144 M 6 yrs 92.5 203 ~1 3 yrs 91. 5 2 3 4 5 6 aBody weights (kg) as of 23 November 1983. �48 6 pai rs of isolation of 6 patrs o_f d~_er(1:::~ Table 2. ) :thro_u9_~ - ._-_ --- Rotation .. .. ", .. . ~ens (A-F}a Time ..... ... .. -- ... _ during 6 17-day time ~eriods duri!19 '1983-84. Deer Pai rs (see Table 1) -_ .. _-- Per-iod -" --- - ._- . -_ .--. 4 2 3 6 1 5 Dec A B C 0 E F 26 Dec B C D E F A 11 Jan C D E F A B 28 Jan D E F A B C 13 Feb E F A B C 0 1 Mar F A B C 0 E Beginning 9 ---------- aA ::: pens 2 & 11; B - pens 5 & 12; C E == pens 6 & 10; F .- pens 7 & 8. ::: pens 1 & 4; D .- pens 3 & 9; �49 ,.-..----~.-~=---"'!I----.--~--~~----~-.,~-, ~ ~i " Q; \.'cJ ~.., \1", (", Observation Tower o \ \~eather Instrumentation ....,_. 10 approx. In Fip,.l. 2.2-ha Loc at i on of cover p a s t ur o of s arre bru s (C) and no-cover h , Lett(~rs C,\-F) (:\C) denote i so l at ion pe ns v i t hi n a i t i pairs of sol a on pe ns • ��Colorado Wildlife July Division Research 51 of Wildlife Report 1984 JOB PROGRESS REPORT State of Colorado 45-01-502-15050 Project No. lNork Pl an No. 2 Job. 6 No. BiG Game InvestiGations - Cervids Winter H3bitat Selection and Act iv i ty Patterns of i~u1e Deer in Front Range Shn.lb i and and Forest Habitats Period Covered: Author: Personnel: 7/1/83-6/30!8~ R. C. Kufeld 8. Parmenter. D. Schrupp ,£l.8STR4CT Triangulation points of approximately 1,500 locations and 250 hours of 24-hr activity data involving 20 radio-collared mule deer were collected during the November 14, 1983, through Match, 1384, per-iod. These data are cur rent ly undergoi n9 ana Iys is. Two of 25 ;~cd-j o-co 11a red does mi grated f'rorn Lory State Park (1 in ,June and 1 in Sep tembe r , 1983) to a mountain valley 26 km west of the study area. One returned in November. The other was believed killed during hunting season. Another migrated from Lory State Park to the east side of Hor se too th Re servo i r in June, 1983, and returned in November', 1933. In fYlay, 1984, it ,:igain migrated to the east side of the reservo t r-. /\11 remaining jee\" have remained within 1.6 km of their capture point as of May 7, 1984. ��53 WINTER HABITAT SELECTION AND ACTIVITY PATTERN OF MULE DEER. IN FRONT RANGE SHRUHLfl.ND AND FOREST HABJTATS Ro Jan d - C " Ku f ('1 d 1. To test Te Ion i cs te l emetry equipment to determine its accuracy at various distances in Iocatinq a t ransmi t ter on the Horsetooth Mountain study area and the ability of an observer using that equipment to correctly detect deer acti vi ty patte rns by habi tat type. 2. To determine habHat within habitat: types • • se lect.ion and cct'iv-ty pattern of mule deer in the Horsetooth ~t)lmti.l"in area during winter . SEGMENT OBJECT~VES 1. Monitor activity 2. , radio-collared mule deer to determine habitat selection patterns throughout 24-·hour' pi~y'-iods . and Develop computer methodoloqy tor ana lysi s of te leme try data on location and activities of mule depr. B. Parmenter as si s ted in collection, field data. D. Schrupp coordinated habitat and telemetry data. tabu l att on and computerization of creation of a system for computerizing t t j'lETHODS J-\ND ~1ATFRIALS Nineteen radto-co l l ared adul L periodically monitor_d fro~ 2 before the 1983 separate deer and during the 3r'd day of the responses of deer to hunting. Lory State Park by State Park anyone time. female deer in Lory State Park were ~st2blishe0 triangulation points 1 day season, dur~ng the 2nd day of that season 1983 combi ned deeY'-e 1k season to determi ne Deer hij~~ey~ were checked in and out of Personnel) and 60 hunters were allowed in at Twenty rad-io-collared adul t f'ema l e dee r VJere peri odi ca l ly monitored from the same 2 triangulation points from November 14, 1983. through March, 1984~ to determine h-bitat selection ~0d activi~y. Equipmen~ procedures. and monitoring schedules were described by ~ufeld (1981, 1982). The monitoring schedule included 3 sun~isp, 3 daytimea 3 sunset, and 3 nighttime �54 peri ods duri ng November, period during December. During the rest February, and 11at'ch, - nd 2 of each Each. eriod lasted 6 hours. January, of the year. per-odic checks using hand-he l d telemetry qeneral equipment were made at Intervals of 1 week to <l:J days to determine location of all radio-collared deer. 2 days 0f the 198: separate Jeer :eason, 82 hunters, respect ive ly , hunted at Lory Stilte Park. Duri nq the season, 12. 15, and 14 hunters were checked in on the first. third days. All 19 deer stdyed within their r~spective home despite the hunting pressure. During the first and 59 comb-ined second and ranges Tel eme try data showtnq approximately 1,SOO deer Iocat t ons and 250 hours of recorded deer sctivity involving ('0 r-adi o-co l l.ared deer' were collected during the November -14-" 19CL, -;hrou;h n rch , 1934, period. These data are current ly under~~oit1g erulyst s -1tld wi 1"1 b':, presented in a future repcrt Only 3 of 28 deer- have left thei r home ran~!l';c; since being instrumented. The rest appear to be vear-uround res idents . The I1rigrational h-istory of 4 , • 4 these 3 is as follows: 1. Deer' #"J./.!-9.641, an adul t doe W<; -irt::;trumented in Lory State Park 1·-19--82. It 1eft its home ranqe betl:/{-:f:n 6--9-·82 and 6---18-82. I twas located 7-"13··82 in a valley 0;-' about ::.? ha , 3.2 km north of Pennock Pass~ which lS 29 km west of its heme range. It spent the entire summer there and W3~~ s tii l there lO·-?b-1.J2<. On 11-1 ..82 it was back in its home ranqe in Lory ,::,tat€' Park. It remai ned in its home range until somet-ime between 8-28-·83 and 9-21-83 when it di sappeared , On 9-21-83 it was located 1n the above-described valley. It was still in the valley on 10-26-83. O~ 11-14-83 it returned to its home range at Lory State Park where it ;'f~maine{\ unt i l H died as d. resul t of a fan on 4-·5·-83. Aqinfj by dental cementum t ndi ca ed this deer was "l4 year's old at time of death. Thus, it woul d neve been 12 years old when ins trumented. 2. Deer #·149.671, an adult doe, \'Ja~; instrumented in Lory State Park 1-27·-82. It left its home ranqe be tween 5·..27--8~~and 6-9-82. It was located in the vel Iey 3.2 km nOY'tJ' o-f Pennock Pass , (see deer' frJ49.64·1 above) on 7--B-S;;: '-J;··,tXt~ it spsnt the entire summer. It was s t i l l there on 10--25--82. On Ij-<-(32 it was back in t ts home range in Lory State Park. It remained in .j t,; horne range urrtl l some time between 6-1 and 6-10-83 when it diseppe~red. It was located in the valley 3.2 km north of Pennock Pass i~-?2-b3 where it again spent the summer. It was in the val l ey or 10--;:::6-,1.),:_). Th~s was 3 days before the 1983 separate deer season. It had not recurned to Lory State Park as of 5-7--84. Ail aeriaI search on -i-<!O--84 f the ent ire Poudre and Big Thompson drainages ;m;"Jud'jy,q the Vi"] ;r-w where located 10-26·-83 revealed no tr-ace of i ts Yddiej sicne l . The author suspects "it was killed by a. hunter dur i iq the 1983 f:i\~er seasc nand the co l Iar transpor-ted out of the area. t f f �55 ,,< Deer 149.64·1 and 14:1.671 were almost constant companions when on the study area at Lory State Park. AHhough they left a week apart in the spring of ·1982, they both migrated to the same valley 3.2 km north of Pennock Pass where together' they spent the summer. They returned, together, to Lory State Park between 10'025-82 and 11-1-82. The following year #149.671 migrated to the valley in June but #149.641 di dn It fol l OvJ unti l September, Deer li149. 641 subsequently returned to Lory State Park in November. ·l983, but ifl49.671 may have been killed in the va l l ey or' enroute to Lory State Park. 3. Deer #149.921. an adult coe, was Estimated at about 5 years of age based on tooth wear when instrumented "it] Lory State Park 2-3-83. Between 6-10--83 and 6-?0-83 it swam eas tward across Horsetooth State Park and spent the ent i re summer and part of the fall -j n the vi clnt ty of the "A" wes t of Hughes ~tad·ium, It gdVI2 bir-th to and raised 2 fawns in that area. It swam wesLward across the reservoir on 11-24-84, and spent the rest of the winter in its regular home range in L.ory State Park. It was observed there in the company of 2 fawns 4·-18-84. Sometime between 4- °13··-84 and 5-7-84 it crossed the reservoi rand returned to the vi ci ni ty cf the LITERAT[JFT Kuf'el d, R. C. ·1981. Wintf: Y' hab-itat AU .. CITED se lecti on and activ i ty patterns mule deer in fron t range shrub land and fovest Wildl. Game Res. R2p. July (1):97-110. --Tn !. habitats. of Colc , Div. 1982. ~J"inter hab itr.t se Iec ti on and act ivi ty patterns of mule deer f'ront ranqe shrub l and and forest habi ta ts . Colo . Di v. ltIi"ldl. Game Res. Rep_ July (1):29-34. Prepared ��57 Colorado Division of Wildlife Wildlife Research Report July, 1984 JOB PROGRESS State of Coiorado Project No. 45-01-502-15050 Work Plan No. Job. No. Personnel: 2 ._------10 Period covered: Author: REPORT 3io Game Investiqations - Cervids Deer Investigations Mule Deer Habitat Use in Piceance Basin 7/1/83-6/30/84 R. M. Bartmann A. W. Alldredge, L. H. Carpenter, J. Depperschmidt, L. Dickerson, D. A. Garrott, R. A. Garrott, J. Graham, G. C. \~hite ,I\BSTRA.CT Gl~aduate student ,John Lee completed his IlLS. thesis "f'iuie Deer Habitat Use and Movements on Piceance Basin Winter Range as Estimated by Radiotelemetry." Preparations for a study of summer habitat use by mul e deer were completed and collection of field data was begun. ��59 ~lULE DEER HABITAT USF IN PICEANCE BASIN Richard M. Bartmann P. N. OB,JECTIVE ee Segment Objective. SEGME~r OP,]E~TIVE Administer funding, supervise graduate students, and provide field assistance in conduct of studi~~ to &_sess mule deer habitat use in proximi ty to oil shale development pro.i ac ts throuqh use of redt ote leme try methods. RESULTS ~ND JISCGSSION results Laboratory. includes the prepa red by The of the entire coopeY-:,-.ive deer' s tudy wi th Los Alamos National Colorado St.at.e i.Jr1"iver.;-jty_ and the Irivi s i on of Wild1ife, which summer dee)' hab i tat wor+., ;<, 'i nc lu ed in the 1984 repor-t Los AIamos personne 1 And . nc I u,,>::d as Appendi x IL /--7 Prepared /' (!/]_... by L .~ .'"?-; -:>--- :__ _}L-_~=- , ----, ~:, ~_:.::=::_:.:,-:_-:~-=-::~::::: Richard M. Bartmann Wildlife Researcher " ��61 AFPENL:i X A. 3A.f .~ MU1~EDEER STUDY~, PICEANCE ISCAL YEAR 1984 RF'"> ,)PT Robert A. Garr-itt an.d Gr·n.j C, White Environmental Science Group, MS-K495 p, O. Bo.: l()63 Los Alamos National t08 laboratory Alamos, NM 875-15 AD::IHAC'T An additional 126 deer were radio collared instrumented population total strong between fidelity to individual "i; wintering dug during in the timing October. lated with weather Timing and summering Adult doe winter of spring migration, mortality 19% for the Little Hills population, previous years. Fawns, instrumented animals consecutive year coyotes the C<·b Tract in starvation of fall mivr-tion and may be delayed area, deaths of cold temp ratures however, W3.:3 throughout. and each migration, traveled Little varia- with most movement appears 15% for the C-b Tract which is comparable !31'.l:Terf'd occur- to be cone- the winter with the winter. and rates of 95% of the For the fourth half of the radio collared fawns in high mortality Persistent population to mortality hea ..•:,' mortality rilled approximately on both areas, All deer showed areas however, the up to ::i. month when the winter has been from cad! area dying during The extremely 1983, bringing the onset of winter. these areas along the same routes during bility was observed severe, to 192 m December rates were due to an increase deep snow ana extended undoubtedly periods were rerponsible for �62 Mule Deer Study Progress Report 2 ~!9E:4. these deaths. This report mule deer study. Laboratory, the project summarizes the results of the: fourth year of field work on the Piceance This is the hst J'(~:~.r the project Pending approval of fiscal yt;r -ill be administered L',' .• fundins will be adrninist .:r,!d til rough Cobr~., (I Basin by Los Alamos National or Energy, from the U. S. Department Sta .',' Un iversity . Trapping Mule deer were trapped ill in the C-b '1 r3!:' <.;.rf~from 11 N rvember to 2 December Decemb ~r t·.., the Little Hills area from?' total years. fawns as adeouate animals numbers Only adult does witt the 4 years or this study instrumented IPl33 of r:.•die-collared ."Mlio (OlI~tJ~ that Drop nets were used to capture .). adult Emphasis U able 2) ~,TI::;' I orcy-three survived Basin to 192 as of December Table 1. Mule deer. trapped, during December 198:~. instrumented, .Af~~-s.e?ZcL.~_ c-s instrumented captured To ~'s Littl ': dills .F3.WEI Ie rna .'::: F2';Wll male Adult female Adult. male Totals Tot.als on instru- During with 408 of these adult does from the G·b Tract trorn ]98:·~<~;3bringing a from previous ".~:'. about to expire wert reinstrumented. the instrumented population area ill 198~j. and released on two study areas in the Piceance Basin ,_,IL.!:.r:'U?IJJ.1. N.c:....j~l.gX... sm , i.A Tr::>.ct Fawn female Fawn male ..b_dult. female Adult I Hk was placed does were available a. tot: .•, of ',I!'ll"c.-.;iHlat :,1:)-900 d •.s hap:. been and 21 from the Little Bilk the Piceance December JJB were ivA.rume I~~.d('rank of 283 deer of which menting i:j 1983 and so 30 30 E,l 30 s o llG 5'".:. 2(l'ec<":tDtUl'es) 27 ,,~ ;14 3:5, 4~) 4 J11 ••I 4( recaptures) 0 6;1 126 �63 Mule Deer Study Progress Report 1984 Table 2. Summary 0 of the number .A~a__ 3 _--_._-__ __ mule deer instrumented as of 15 December . -. ...••-.- ..,--.•.. ~-_._."" ......""' ..•.•..•-..,.. ..._... .... ),ota! AQJ!lt d_oes Ys~I fl~'§' ., C-,b Tract 1980 1981 1982 198:~ Little Hills :! 98:2 1983 28 25 53 66 1)1 117 61 51 43 112 60 59 21 60 29 80 89 of each year . 103 }'all Migration The Call migration the previous 2 years. peak of migration of G"b Tract All deer migrated occurring routes All deer returned used during deer requiring ranges Ninemile deer followed similar November migration north of the migration Meeker did not leave her summer range. Another years. and Little Hills study range. in less funneled 1983 through through they had used during 23 November, All instrumented fall deer. the 1983 to be longer with most One adult summered to the wintering their m the G·b Tract 10 November. doe that range unt.il. approximately areas returned 2.'5 deer tended approximately range some time during the first 2 weeks in December. Tract from 20-60 km the same migration Two deer also remained onto the winter until HI 2~'3 observed for individual the migration. of Meeker she was on her winter patterns to the Little Hills area most deer had moved area with the dr, mage to eacn C-h Tract, 2-3 weeks to complete Gap October during the movement Most of these animals Little .Hills deer were followed for the first. time Duration long after completing to the G"b Tract area along approximately All deer used the same routes to return spring migration. range during 1). C-b Tract. deer traveled the 1921 and 1982 fall migrat.ions. Instrumented These (Fi!. t.o winter range with most animals the Twin Gulches area of the Stuart migration. from summer in mid-October (12-35 miles) to reach their winter than 2 weeks. deer was €,c}sentially the same as we had observed on their summer doe stayed However, in the by 23 on LO 7 Hill south of arriving on the winter deer from both the C-b areas they had used in previous �64 Mule Deer Study Progress Report 1984 Fhll Migrat.'. 20 3 L lt1 C -b 'li"act N,,-..::U U, 12 8 "o ...f<._""'''''''~A_-'''~'''' ~-4 ~_-~ ••.•.• -16 0_20 -+----'''_ s... Q) 16 '8 ~ ..,_.,.-------'-------r----'----··I-~~--'-·'--~, ,----,.------ LiUJE;Hills start. -19 12 : o -4 '.-8 =·12 -16 -20 ,.-I-·-··-'~·-"-'·-·""'·-"-'''1'''-'oct.. Oct.. nisb Oct. is-a 8-~1.4 1-7 Figure L Chronology ceance Basin, ... "" '-'1''''-'-'''''- ---'-'-~"~'I--------1 ---. t'- '---- "-r of the fall 1983 :n'gratioD Oct, Nov" Nov. 2'}'31 1-7 6-14 for instrumented deer on 2 study areas in the Pi- Winter 'Movements Instrumented those documented deer remained Movements deer from the C b Tract for the previous dispersed early January. resulting the same resulting However, movements routes s occupying the north ,~';!ler?ll}did several major to areas probably ':~ll],~edthe {'C: to area in October, areas used during previous years. and northwest began in late December not begm during the previous snowstorms In m)..110 em of snow pack snow accumulation Oil-tract area toward similar winter movement, patterns After ret-urning to the C-b Tract the study area These off..tract movements until the end of January. early January ~!winters. throughout off the C-b Tract z.rea displayed on the study occurred areas. 2 winters in late December These storms and and and the rlier olI..tract movements. of the instrumented deer were :11::;0 predictable they had occupied during previous winters.' in that deer followed After this mid ..winter �65 Mule Deer Study Progress Report movement, deer remained relatively area just prior to initiating Deer wintering observed 5 1984 sedentary animals. Instead the trap sites where they were originally porarily moved during Several deer that previously approximately Despite sedentary, movements- occupying areas near were more mobile than A group of deer from the Rattlesnake Ridge onto a south-facing Creek. mid-winter Some deer, however, slope in the Piceance area tem- Creek drainage. high on North Ridge moved to lower elevations and several deer moved from North Dry Fork of Piceance Ridge captured. had remained the extensive most deer remained the winter of 1982-83, across North the White River, to the C-b Tract the spring migration. in the Little Hills area did not display in the C-b Tract we had observed until spring when most returned Ridge onto south-facing slopes along the deep snow on the top and north-facing half of the instrumented deer remained in these along slopes of North areas throughout the winter. Spring Migration Due to time and log-istic constraints, during May. the spring This migration. was one month approximately with the severity later migration of the winter. were very light with generally periods than the onset .reserves of fat and protein observed During during mild temperatures condition in spring :"2 winters additional body reserves before beginning 1981 appears throughout deer undoubtedly more than in a mild winter. due to severe winter migration until early :/ODd 1982, but to be correlated of 1980-81 and 1981-82 snow accumulations persisting During such winters were followed (Fig. 2). The one month delay in the the previous the winters begins in spring with the onset of warm weather obtain of migrations of 1982-83 and 1983 ..84 were more severe with persistent of cold temperatures" physiological deer from C-h Tract C-b Tract. deer did not begin their spring the same time as the 1983 spring migration onset of the spring the winters only instrumented Reversal and production weather In contrast, deep snow and extended deplete their body energy of this negative energy balance of new veg ration. may delay their migration. the winter. spring migration Deer in poor in order to �66 Mule Deer Study Progress Report 1984 Spring MlgI-diion 1981-84 1981\ 1\ I. I I i \ 60- I / I 50 1 30.'\ ! ,'( 20 " I 10 / f I " I ," tion of migration migration \' 1982 \\', ' \ JI I;' I i)' I I \ \ \ '-// \\ Apr. 4.-30 _'f_._-,. - \ \. \ --.., . ,y 1-7 of instrumented migration deer on C-b Tract, were similar the same routes they had used during for individual \ \ " \ . __'~' of the 1984 spring along essentially I \\ /'" Figure 2. Timing of spring migration traveling \ ...j,,\ 1983 \,o. __ ...._. ~--·'"-r·-r··..--··,.........·-· ._ 1/ Jh ~ aspects , •••• L-\\ /,/ \\ oW "1 o· 19B4(,. \ 1 I Other \ \ \ \ deer ranged 1981.-84. to previous years with all deer earlier migrations. The dura- Irom 1 to 4 weeks with most deer completing the in 2 or 3 weeks. Summer Movements Thirty-two instrumented area where periodically bad used during Tract . years. animals located durmg the summer. previous summers. and 19 adult does from the Little Hills Ail does returned This site-specific as 12 does have been tracked Nineteen same fidelity. adult does from C-b Tract fidelity has been well documented for 4 years, 9 does for 3 years, does from the Little Hills area have also been tracked In addition to these animals currently have either died or had their radios expire, provide time. Once had ranges does the summer, arrived on their occupying summer an area no greater for C-b and 11 does tor 2 for 2 years and show the being monitored, which throughout to the same area that they approximately 22 does, fidelity data for various lengths they remained than 1.5 km".'" relatively of' sedentary The only exceptions to this �67 Mule Deer Study Progress Report pattern have been infrequent pancy. These movements 1984 temporary 'were usually 7 use of areas up to 4 km from the "normal" made by yearlings; however, area of occu- adult does were occasionally involved. WAortali.ty Six of .58 instrumented additional 9 transmitters to battery depletion large proportion expected adult call be expected of the transmitters 2 transmitters 2: apparently starved, cause of death or Iell off the animals. to become more Irequent and 1 became (excluding rates documented Five of 28 instrumented additional tangled transmitter for (;'''0 Tract for the fifth doe On both the Cob Tract W3;S areas without contacting As we had observed predominant were attributed dator caused. because a for 2 to 4 years, which is the ll.was hit by :JI These deer represent which is comparable rate. The fate (if during These deaths and Little Hills areas 56 rate observed or 60 an additional throughout the hunting represent car, 15% to the 8- 21 2, predator, season, and the 19% winter mortal- during the 1982-83 winter. instrumented fawns died during the Iawn on each area was undetermined These 2 radios probably were conducted failed transmitting the Piceance prematurely Basin and the surrounding the signals. last year, {awn survival causes of mortality were dissimilar. to coyote predation In contrast" failures due or the 5 does that died, :1.was killed by undetermined. with the signal. aerial searches An animals during the past 3 years. ity rate which is slightly higher than the 10% mortality as repeated failures), 1 was illegally shot and abandoned because we lost contact Transmitter as the study progresses a fence. ill! the winter. adult does from the Little Hills area died during the winter with an falling off animals. winter for a 95% mortality area died during Of the 6 does that died, :2 were killed by coyotes, population 16% winter mortality C-b Tract in the field have been operating died from starvation, of the instrumented d.IY~ either ceased operation life of most units. 2 apparently does hom in the 2 study Fifty percent while only 12% or the areas was similar [Table 3), but of the G-b Tract fawn mortalities Little Hills fawn mortalities 88% of the Little Hills fawn mortalities were pre- were due to starvation while �68 Mule Deer Study Progress Table 3. Summary Basin. Report ).984 8 of survival rate of instrumented ,:}l~!l __ ~_ _.);~fl:.L 1980·81 1981·82 19S2~83 19'83·84 !:2:Y!:!!1. .:1.<:b..1tQ.Q.f.& 0.22 0.38 0 ..05 0.92 0.90 0.84 0.85 Li.We HiHs J98:Z-83 036 0.90 ___ ._. !.~~~:~! .__ QQ~.~ __ ..__._Q.8L_ C-b Tract, only 46% of the C-b Tract fawn mortalities Table 4. Causes of winter fawn mortality -~_,_:':.~";!""=:~ observed dation winter of 1983-84, however, on G··b Tract animals fawns that bad starved Carcass winters increase succumbing remained to death Tract population 3ft 75 ~! 112 56 deer during. this past winter was similar to that fawns suffered little mortality due to starvation during December while off tract. occurred to death throughout coyote predation throughout During the the rest of' the winter with over half the (Fig. 3). Intact carcasses of 55 and January were weighed when they were found (rom 12 to 36 hours after Host (awns had lost. between 20 and 30% from 14.1 to 29.1 kg. in the number including similar: 7 49 0 body weight. area have consistently ing the study, Tot~ls While deer remained on the C-b Tract area, coyote pre- heavy mortality The high overall {awn mortality all JJ)lh k_tle Basin. as soon as deer moved off the area in mid-winter, weights ranged of their December (Table 4). the Piceance ill - 56 (Fig. 3). Litt.le Hills fawns starved instrumented to starvation ,) i:.• the previous 3 winters. In previous due to !]-2 ...L.:" ~t:. 26 sharply. death. ...- 28 however, the winter on 2 study area" Winter kill Undetermined Tot.als was heavy: declined were attributed Predator of coyote predation during 0.:35 ........... "...__.. ..~- --~._...--~.- M.__U_ll._.~~ ..L_ The pattern mule deer from 2 study areas in. the Piceance remained observed Oil of fawns starving both areas during to death. Predator losses in the C-b Tract in the range of <if, to 50% of the instrumented the win ter of 1983--81. Predator 8% in 1982-83 and 12% in 198:)-,84. have normally the winter of 1983-84 was been less than population dur- losses on the Little Hills area have also In contrast, .)~':; during previous starvation losses in the C-b winters while 43% of the �69 Mule Deer Study Progress Report 1984 Mortalities 1983-84 Figure 3. Timing of instrumented instrumented bled ill fawn mortalities fawns died of starvation the Little Hills population during major throughout winter tures storms during the study low mortality late December cold temperatures. red uced deer mobility Most adult mortality does had sufficient rates; however, reserves the snow pack during 01 snow the remainder of the of deep snow and cold tempera- and increased to maintain 1982-83 to 82% during left deep accumulations This combination and food availability or. losses also dou- is due to the severity of the winter. and early January ill Star-vation the winter undoubtedly areas with little decrease due to persistent reserves. the winter of 1983-84. going from 41% during the winter of 1983-84.. This increased Several in 2 study areas, winter 1983-841. the depletion themselves of body energy as evidenced by their fawns did not have the needed reserves and suffered heavy mortal- ity. The experiment was completed died during ote predation to determine this past winter. if radio-collared Fifty-eight the 3 winters the experiment (Table 5). Transmitter fawns are subjected of 91 instrumented war; conducted, to increased male fawns were known to have with most mortalities and collar Iailures resulted predation ill censored attributed to coy- data for 20 fawns, �70 Mule Deer Study Progress Report with these data eliminated Table 5. Summary transmitters. 1984 10 from the analysis. of the fate of male fawns instrumented with ear tag and collar mounted _ ..__ ._.._---_.. Animal's Year 1982 1983 1984 Total Total Radio A t.tl'l'hmf"nt Fate Died Predatio'l Lived ~----- Died Other Device Failure Ear tag Collar 0 3 4 7 2 1 12 6 18 17 Ear tag CoUar 4. 4 5 7 4 2 0 J3 0 13 Ear tag Collar 1 1 11 8 2 5 1 1. 15 15 Ear tag Collar 5 8 20 8 8 13 7 46 45 20 91 22 Total 13 16 42 ----.-.-.------.-~-.-_,. ...--------..---, .•........ We were unable between fawns hypothesized and/or to detect a difference instrumented with tag transmitter, a difference not subject in either predator-induced ear tag that. the collar may predispose providing in mortality rates fawns to increased the last 2 winters, between radio to determine changes for a second field season. Dr. Carpenter (CDOW) both possibilities. We had by a visual cue or the Failure fawns indicates ear- to detect collars do coyote predation .. in the body composition of deer throughout This study is being conducted "with cooperation from Los Alamos National In addition, 8 instrumented Data on a variety were ground Laboratory. (CSU) and During the water, in the vicinity of the Little fawns that had died of starvation of carcass measurements up and total by Dr. Torbit the winter for the study, and during the winter of 1983-84 fawns To date, 24 adult does and 24 fawns have been collected in the samples. transmitters. Size and placement the 2 groups of instrumented winter of 1982-83 adult does were collected Hills Wildlife Area. or overall mortality coyote predation an attack. precluded mortality Study was continued the carcasses collar-mounted &-11 month old mule deer fawns to increased Deer Body Composlulou were used. and a secure grip on the animals during at least during The study Total were included and fat indices were collected fat, protein, and ash content ox before each animal �71 Mule Deer Study Progress determined. cipated 1984 11 analysis of many of the samples is not yet complete; Laboratory that these data will prove invaluable deer undergo data Report during the stressful winter also will allow us to correlate instrumented in understanding period when most mortality nondestructive deer with body composition the dynamic carcass however, physiological OCCUl!S. measurements it is anti- taken changes Hopefully, these currently on all and winter survival. Road Reflector Experiment The experiment along 4 l.6-km covered weeks from October of road-killed of the Swareflex deer found Olll through 19 when the reflectors data are insufficient for at least another of reflectors were An additional 9 d er were killed To date, 31 deer have been killed on the 4 test were covered and 12 when the reflectors were operational. to determine year. Two sections was continued May when deer were on the winter range and each section recorded. on the test sections during the winter of 1983-84. sections, wildlife reflector of the Piceance Creek Road. (1 mile) sections on alternate the number to test the effectiveness the effectiveness of the reflectors These so the study will be continued , I abitat Studie Graduate student S. degree during State University, the summer of 1984. The abstract Locations hour during of adult winter radiotelemetry (SD=25.1, John Lee, Colorado triangulation. N--32). Areas during or use during Does showed fidelity to certain 12-hour graphics and a If>-mm movie were used to enhance of oil shale development were estimated technique Basin, periods to an oil shale development as an investigative for the M. every 1/2 Colorado averaged using 24.7 ha areas, moved readily across roads, and areas in close proximity impacts hemionu8) 1981-82 in Piceance occupied Radiotelemetry requirements from his thesis follows. female mule deer (Odocoileu8 1981 and hourly completed display is discussed on mule deer in northwestern site. Computer-generated and interpretation with reference Colorado. of data. to monitoring �Mule Deer Study Progress Accuracy bearings standard 12 1984 of the telemetry radio-transmitters sidered Report system placed at surveyed signal bounce Bearings by taking errors. bias. Bearing replicate with absolute error bearings on > 10(1 were con- of bias and precision. from calculations for tower orientation of bearing deviation points. and excluded were corrected was quantified System precision Remaining was measured by the in size f om 7.0 - 20.6° at arcs ranged P=O.05. There was no difference in precision (p.,,",O.38) between observers but there difference (P=:O.002) in precision polygon area are displayed. Mule deer habitat accuracy defined habitat Chained Results areas «.) snow accumulation rangeland months Estimated at P 0.05. and use-availability was the most em) while pinyon-juniper ing periods with snow accumulation. months. deer-use The first year of data collection on the Roan Plateau the study are 1) to estimate of error Telemetry parameters. Habitats In 1981 there WM ill types used habitat woodland Sagebrush of habitat Deer tended conducted Isopleths system were used no difference in (P=.::O.15) but there was a difference in 1981-82 (P<O.OOl). months pinyon-juniper signals. from point location estimates. (P<O.OOl) in all months. to availability use between to modulating are discussed with respect to system and study design. use was estimated monitoring disproportionate of bearings the vicinity summer-long was the least used habitat was dependent 011 the summer of the Colony habitat periods selection with little was the most used habitat to spend nights in chainings was initiated during on activity systems were erected tested on 2 study in May of this year; Approximately an estimated marized and movement patterns areas during summer data collection 670 hours of tracking 7000 locations in a separate report involving that of adult began all on time of day during all and days in woodlands. habitat study which is being Shale Oil Project. and avoidance Objectives and activity 18 instrumented will be compiled of patterns dis- female mule deer. Permanent 1983. The accuracy of these systems was in June will be completed dur- type during of adult female mule deer over 24··hour periods and 2) to test for the effects of construction turbance a W:a\S and will continue until telemetry October. by the end of the field season, generating does. Results and circulated of this work will be sum- after the field season is corn- �73 Mule Deer Study Progress Report 198'~ 13 pleted. Aeknowledgmenttl We gratefully acknowledge Shale Office, Bureau Wildlife, Colorado local ranchers. 7405-ENG-36 the encouragement, of Land Management, State University, Financial support to Los Alamos vided by Exxon Company advice, Cathedral National Division of USA, Rio Blanco Oil Shale Co., and many by the U. S. Department Laboratory. USA, Cathedral of the Area Oil Bluffs Shale Oil Co., Colorado Exxon Company was provided and cooperation Supplementary Bluffs Shale Oil Company, of Energy, financial contract support w- was pro- and the Colorado Division of Wildlife. Anderson, D. R., K. P. Burnham, from recovery Bartrnann, 19841. Estimating J. Range Manage.:. age-specific survival rates J. Anim. &01.:. In Press. data of birds ringed as young. R. M. 1983. Mule deer diet composition Colorado. Bartrnann, and G. C. White. and quality on pinyon-juniper winter range, In Pre3S. R. M. 1983. Winter foods of mule deer in Piceance Basin. Colorado Division OM Wildlife Game Info. Leaflet 107:1-4. Bartmann, tion. Bartrnann, R. M. 1983. Appraisal Colorado of a quadrant census of mule deer in pinyon-juniper vegeta- Division of Wildlife Game Info. Leaflet 109:1-4. R. M. 1984. Estimating mule deer winter mortality. J. Wildl. Manage. 48(1):262- 267. Bartmann, R. M., A. W. Alldredge, tame mule deer. J. Wildt and P. H. Neil. 1982. Evaluation Manage. 46(3):807-812. of winter food choices by �74 Mule Dee. Study Progress Report 1934 Bartmann, R. M. and D. C. Bowden, data. 14 1984. Predicting mule deer winter mortality from weather Wildt Soc. Bull. 12:. In Press. Bartmann, R. M. and L. H. Carpenter. 1982. Effects of foraging experience Oil food selectivity of tame mule deer. J. Wildt Manage. 46(3):813-.813, K. P., G. C. White, and D. R. Anderson, Burnham, annual Garrott, survival Fates of adult mallards. R. A. and R. M. Bartmann, wnei. 1984. Estimating the effect of hunting on J. Wildt Manage. 48(2):350-361. 1984. Evaluation of vaginal implants for mule deer. J. Manage. 48(2):646-.648. Garrott, R. A., R. M. Hartmann, and G. C. White. 198-5. Effects of radio-transmitter J. Wildt Manage..'. on fawn deer mortality. Garrott, R. A. and R. W. Hayes. packages Submitted. 1984. A radio-controlled device for triggering traps. Wildl, 1982. Age and sex selectivity J. Wildt Soc. Bull. 12:. In Pr~ss. Garrott, R.. A. and G. C. White. in trapping mule deer. Manage. 46(4):1083-1086. Garrott, R. A. and G. C. White. 1983. A comparison instrumented of predation rates between mule deer fawns with collars and ear tags. in D. G. Pincock eds. Proc, Fourth International Wildlife Biotelemetry Conference, Applied Microelectronics Institute and Technical Universi- ty of Nova Scotia, Halifax. Garrott, R. A. and G. C. White. 1984. Methodology Ior assessing the impacts of oil shale development Ox; the Piceance eds. Issues and Technology logical Institute. Basin mule deer herd. in the Management Boulder, CO. Pages 228-231. of Impacted Western in R. D. Comer et al. Wildlife, Thorne Eco- �75 Mule Deer Study Progress Report Gill, R. 8., L. H. Carpenter, analysis to estimate 15 1984 R. M. Bartmann, to estimate mule deer locations in Piceance 232-23'r. in R. D. Comer et al, eds. Issues and Technology Western Lee, Wildlife, Thorne Ecological Institute, Boulder, J. E., G. C. White, R. A. Garrott, R.. M. Bartmann, the accuracy Manage.:. Torbit, of a radiotelemetry S. C., L. W. Carpenter, Pages of Impacted CO. mule deer locations. 1984. Assessing J. Wildl. A. W. Alldredge, and D. M. Swift. 1984. Mule deer body compo- J. Wildl. Manage. 48:. In Press. of methods. 1984. Winter dynamics of mule deer body com- J. Wildl. Manage, 48:. In Press. position. data. in the Management and A. W. Alldredge. system for estimating S. C., D. M. Swift, and A. W. Alldredge. White, G. C. Basin, Colo. Submitted. sition - a comparison Torbit, 1983. Fecal J. Wildt Manage. 47(4):902-915. mule deer diets. Lee, J. E. 1984. Radio-telemetry D. L. Baker, and G. G. Schoonveld. 1983. Numerical estimation of survival rates from band recovery and biotelemetry J. Wild I. Manage. 47(3):716-728. White, G. C. 1084. Optimal Wildl. Manage. 48(4):. locations ceance Basin mule deer herd. Inventories vallis, OR. J. H183. Estimation of survival rates from band recoveries of J. Wild!. Manage. 47(2):506-511. White, G. C. and R. A. Garrott. Resource studies using biotelemetry. In Press. White, G. C. and R. M. Bartmann. mule deer in Colorado. of towers for triangulation 1983. Assessing impacts Pages 483-486. for Monitoring Changes of oil shale development in J. F. Bell and T. Atterbury and Trends, on the Pieds, Renewable Oregon State University, Cor- �76 Mule Deer Study Progress Report 1984 White, G. C. and R. A. Garrott. 19841. Portable computer system for field processing biotelemetry triangulation data. Colorado Div. OK Wildl. Game Information Leaflet 110:1-4. White, G. C. and L. J. Lane. 1982. Comments on an example of simulation models as decision tools in wildlife management. 16 Wildl. Soc. Bull. 1O{3}:285-286. �77 Colorado Division of Wildlife Wildlife Research Report July, 1984 JOB PROGRESS REPORT State of Colorado Project No. 45-01-502-15050 BiG Game Investi~t-ions - Cervids Work Plan No. 2 -------- Deer Investiaations Job. 11 Testing of Mule Deer Census l1ethodology No. Period covered: Author: 7/1/83-6/30/84 R. M. Bartmann Personnel: A. W. Alldredge, A. Alvord, S. W. Alvord, T. A. Blocker, D. C. Bowden, L. H. Carpenter-, D. J. Freddy, D. A. Garrott, R. A. Garrott, R. Green, A. Markill, D. Weybright, G. C. White, D. Whittaker, and numerous CSU student volunteers. ABSTR,l\.CT Mule deer census evaluations were made in pastures at the Little Hills Wildlife Ar-ea in December, 1983, and January and February, 1984. Forty-eight deer were radio-collared and p[aced in 4 pas tures of 58-70 ha at densities of about 12 and 25 deer/km£, Some unmarked deer were also in the pastures and increased densities by amounts yet to be determined. Quadrat size, 65 and 130 ha, had no effect (P > 0.05) on number of deer counted, but count direction, east-west an~ north-south, did with counts on east-west transects being lower (P < 0.05). There were no measurable short-term or long-term effects of harrassment of deer from repeated counts on proportions of deer counted. However, behavior changes were noted during each month's surveys. The lower proportions (P < 0.05) of deer counted in Pasture 2 compared to the other 3 pastures during December and January were attributed to characteristics of individual deer rather than to counting conditions in the pasture, The range in deer densities had no measurable effect on counting accuracy. Excluding the December and January data from Pasture 2, the mean proportion of marked deer counted was 0.69 ± 0.04 at the 95% confidence level. ��79 TESTING OF MULE DEER CENSUS METHODOLOGY Richard M, Bartmann P. N•.. OBJECTIVES 1. Assess effects of search area size on number of deer counted. 2. Assess practicality of altering the number of deer in the pastures between monthly counts. 3. Measure the effects of repeated aerial surveys on numbers of deer counted. 4. In cooperation with the Los Alamos National Laboratory~ increase the effective transmitting range of the deer defecation monitor implant to about 1.5 km and tnccrporate data storage capability to allow noncontinuous monitoring of animals. SEGMENT OBJECTIVES Same as P. N. Objectives. ~11ETHODSAND MATERIALS Fences of Pastures 29 3, 4, 5, and 6 were repaired and modified as required to hold deer for the census evaluations. Pastures 5 and 6 were combined by removing a division fence· to form a single 62-ha pasture. Sizes of the 4 pastures, 58p70 ha. were then within 11% of the 65-ha size of deer census quadrats in Piceance Basin. Pasture Stocking An unknown number of deer got into the pastures before radio-collared deer were stocked. Records were kept of mortalities and removals of live unmarked deer throughout the winter to help arrive at a minimum number in each pasture. Pastures were stocked with marked deer at 2 densities: about 12 and 25 deer/km2. Densities were randomly assigned with Pastures 2 and 5 selected as low density and Pastures 3 and 4 as high density. Forty-eight deer were caught with drop-nets, radio-collared. and placed in the pastures between 18 November and 8 December 1983. Radio-collars were white and 5.l-cm wide. Each transmitter included a mortality sensor that doubled the pulse-rate of the signal after an animal was still for 4 hours. The numbers and kinds of deer (bucks:does:fawns) put in each pasture were: Pasture 2 (2:3:3), Pasture 3 (2:8:6)~ Pasture 4 (2:6:8), and Pasture 5, (1:4:3). �80 All radio-collared deer were located by triangulation at appropriate times to be sure individuals were in the proper pastures during each census evaluation. In addition. all radio-signals were moni tored 3 times daily during surveys to verify if a.nimals were alive or dead at the time of each count. Between monthly surveys, dead an imal s in all but Pasture 2 were replaced with adult does trapped on winter range. Replacements in Pasture 2 were recruited f om unmarked deer already in the pasture. Except in Pasture 2, fawns were not used as replacements as they were assumed poor survival risks due to the severe winter. Prior to the February flights, 11 unmarked deer already in Pasture 2 were trapped with Clover traps and redi o-co l l ared. Three were replacements for radio-collared deer that died and 8 were added to double the marked deer popul at lon. Al so , prtor to the February flights, an radio-collared deer and an unkown number of unma.rked deer escaped from Pasture 5 into PasturE: 4. This increased the marked deer population there from 16 to 24. Capturing deer for removal from the pastures and retrieval of radiocollars was done wi th Clover traps 'in March and April. However , roads washed out before all deer could be captured and deer are still present in 3 pastures. Snow Measurements Data on snow conditions in the pastures were recorded during each monthly census eva luation. Percent snow cover was estimated from aerial photos taken from a f ixed-winq aircraft. Bare ground tn each pasture was outlined on the photos and measured with a planimeter. A 5-stake transect , with stakes about 10-m apart, was non-randomly es tabl ished on a south slope 9 a nor-th slope, a r idgetop 9 and a bottom site in each pas ture. Information was recorded for snow depth (em), snow consistency, and snow SUPPOy·t capab i1i tv. Snow cons is tency was ra ted as CI) dry, wi 1"1 not form a snowball or (2) wet~ will form a snowball. Snow support capability was rated as (I) none, wi l l not support a person, (2) var-iab l e , will support a person in some places and not in others, or (3) high, will consistently support a person. Census Evaluations Objective 1 - Assess Effects of Search Area Size on Number of Deer Counted --------------------------------------------------------------------------The Ia rqer the quadrat. the more time deer have to redistribute themselves between successive passes of the he li copter , The longer this time-lag, the more diffi ell l t it may be to keep track of deer. Thus, some deer coul d be counted more than once VI/hi'I e others are never counted. Two quadrat sizes were tested in Decemoer , 1983: 65 ha and no ha , Each pasture represented a 65-ha quadrat and also comprised 1/2 of a 130-ha quadrat. The larger quadrat was formed by adding a second 65-ha parcel of unfenced vJinter ranqe adjacent to the south boundary of each pasture (Fig. 1). Corners of these 4 unfenced areas were delineated with 30 x 60-em orange markers. Deer counts on the unfenced half of each 130-ha quadrat were not used in any analyses. Instead, the only purpose for searching these areas was to create the t ime-Laq mentioned above. �81 Six series of counts were made over a 3-day interval from 13-15 December 1983 with an AM and PM count each day. A series included a count of the 4, 65-ha quadrats (pastures) and the 4, 130-ha quadrats. The counting sequence was randomly determined for each series with the constraint that counts on the large and small quadrats within the same set must be separated by counts on at least 2 other quadrats. This allowed deer time to settle down before being counted the second time. The same search pattern was used for all quadrats,"beginning in the northeast corner and proceeding south along the east boundary fence. Transecting then continued in north and south directions until the entire quadrat was searched. Transect spacing was determined by the 2 observers based on terrain and cover density considerations. Objective 2 - Assess Practicality of Altering the Number 2f_Q~~~_!~_!~~_~~~~~r~~_~~!~~~~_~2~~~!~_~2~~E~ _ Exchanging deer densities among pastures between monthly census evaluations would help reduce pasture effects on count results. Thus, it was planned to trap 8 deer in each high density pasture and transfer them to the 2 low density pastures. This was not accomplished due to access problems caused by the deep snow. However. the trapping of deer in Pasture 2 in early February indicated this is a viable option. Object; ve 3 - MeasUl~e the Effects of Repeated Aer"la 1 ?~~~~~~_9~_~~~~~~~_2f_Q~~~_~2~~!~9 _ Results of deer counts for each month were graphically plotted to reveal any trends over time with repeated counts that may indicate short-term effects of harrassment. Proportions of deer counted in each pasture were compared among months with ratio estimate procedures to check for longerterm effects of harrassment. Objective 4 - In Cooperation with the Los Alamos National Laboratory, Increase the Effective Transmitting Range of the Deer Defecation Monitor' Implant to about 1.5 km and Incorporate Data Storage Capability to Allow Non-continuous ~9~!!9r!~g_2f_~~!~~!~ _ Entire responsibility for pursuit of this objective now lies with the Los Alamos National Laboratory. Q!b~~_~9~e~~!~2~~ The original plan was to repeat the comparison of deer counts on 2 sizes of quadrats in January. After examining the December results, I considered this unnecessary. Instead, the effects of search direction on count results were evaluated. Quadrats in December were searched in north-south directions approximately parallel to major drainages. In January, results of counts made in these directions were compared to those made in east-west directions that crossed major drainages. Six series of counts were made with all pastures searched in both directions mornings and afternoons for 4 days. The extra day was needed to complete the AM count missed on Day 2 due to weather. Procedures used in December to select pasture counting sequences were applied here. All counts began in the northeast corner of each pasture. Observers were the same as in December but the pilot was new. �82 The February census evaluations, not originally planned, were made to check for longer-term effects of deer harrassment and the effects of late winter snow conditions on count results. In addition, we wanted to identify if the low proportions of deer counted in Pasture 2 during December and January were due to pasture conditions or to characteristics of individual deer. To facilitate this, the radio-collared population in Pasture 2 was increased from 8 deer to 16. Six counts were made mornings and evenings for 4 days with pasture sequences randomly determined each time. Helicopter problems forced rescheduling the AM count for Day 2 to Day 4. Pastures were flown once each time and only in north-south directions. The pilot and observers were the same as in December. RESULTS Snow Measurements Snow cover was essentially complete in all pastures in December. In January, only Pasture 4 had measurable bare ground (3%) while percentages of bare ground in February ranged from 1-10% in all pastures. These low percentages primarily reflect the small amount of south exposures in the pastures. They do not 'include bare areas under trees which cannot be seen on an aerial photo but are quite obvious to observers searching for deer. Snow depths were greatest on north slopes and bottoms and lowest on south slopes (Table 1). Maximum snow depths on ridgetops were similar to those on north slopes and bottoms, but shallower depths under trees caused a Iower average depth. Snow depths increased about 4-fold from December to January. Snow was even deeper on all but south slopes in February because cold temperatures did not allow melting of the re1atively 1ittle snow that fell after the January surveys. Snow accumulated rapidly in December and was dry and powdery in all areas. There was a slight crust on south slopes in January and by February it would support a person. Partially supportive crusts occurred in other areas exposed to sunlight with snow consistency underneath dry and granular. As the snow became deeper and more crusted, the deer began to restrict their use of the pastures. Census Evaluations Objective 1 - Assess Effects of Search Area Size on Number of Deer Counted A radio-collared buck escaped from Pasture 2 before the December surveys but was recaptured and placed back in the pasture for the last 4 series of counts. The only mortality of a radio-collared deer was a fawn that died in Pasture 3 between the fourth and fifth series of counts. There was no difference (P > 0.05) in the proportions of deer counted in the pastures when they were searched as individual quadrats and when they were searched as part of the larger. 130-ha quadrats (Table 2). If there were any counting errors due to size of search area, they were either too small to detect or were compensating, i.e., duplicate counts of the same �83 animals approximated the number of deer missed. These results suggest counting accuracy need not be a major consideration in selecting 1 of these 2 quadrat sizes in pinyon-juniper habitat. Since no significant differences were found, it was decided not to repeat this test in January. Objective 3 - Measure the Effects of ~~P~~~~9_~~r!~1_~~r~~~~_Q~_~~I!!~~r~~Qf_g~~r_~Q~~!~9 Plots of successive counts each month reveal no consistent trends for either marked deer or total deer (Fig. 2). Considerable variation among counts, particularly for total deer, was evident, however, and could obscure any short-term effects of harrassment that exist. Proportions of deer counted in Pasture 2 in December and January were lower (P < 0.05) than in February. Proportions were similar (P > 0.05) among months in the other 3 pastures. The results for Pasture 2 are not considered due to repeated disturbance, so there were no measurable longer-term effects of harrassment. While no harrassment effects were quantified, there were noticeable changes in deer behavior over time each month. In December deer seemed more reluctant to move during successive approaches of the helicopter. In extreme cases deer under trees refused to flush even when the helicopter hovered overhead and deer in open areas would run under the nearest trees and refuse to leave. This behavior seemed more pronounced in January and was especially obvious in February. Q~b~r_~Ql!!e~r!~Q~~ Proportions of marked deer seen in all pastures when searched in eastwest directions was lower (P < 0.05) than when searched north-south (Table 3). The mean proportions of deer counted on east-west flights were also lower in every pasture, but the difference was significant (f < 0.05) only for Pasture 4. Proportions of marked deer counted in Pasture 2 in February was significantly higher (P < 0.05) than in December and January. Therefore, the reason for the lower counts the first 2 months seems more likely attributable to peculiariti~s of individual deer than to counting conditions in the pasture. Effects of deer density on counts is difficult to assess quantitatively at this time. While numbers of marked animals in each pasture were known, numbers of unmarked deer have yet to be determined. Estimates based on capture-recapture analyses indicate there were 69, 65, and 43 unmarked deer in the pastures in December, January, and February, respectively. (Tables 29 3, and 4). The lower total for February is due, in part, to the trapping and radio-collaring of 11 unmarked deer in Pasture 2. The death of an injured buck in Pasture 2 was the only loss of a marked anima 1 between the December and January surveys. Therefore, I assume the loss of unmarked deer was also negligible and the total number of deer in the pastures was similar both months. Estimates based on capture-recapture techniques support this assumption. �84 Mortality of marked deer did occur between the January and February surveys and was composed entirely of fawns. Therefore, some mortality of unmarked deer was also expected and was reflected in the capturerecapture estimates. Accuracy of these estimates is unknown at this time, but a minimum number for each pasture will be obtained after deer removals are completed. Excluding the December and January-data for Pasture 2, the proportions of deer counted in each pasture over'winter ranged from 0.55 to 0.77 (Table 5). The overall mean was 0.69 ± 0.04 at the 95% confidence level. This small confidence interval indicates deer density, at least within the range experienced, was not an important influence on counting accuracy . Prepared .... -- ~.... .. �85 Table 1. Show depths (cm) in the pastures at the Little Hills Wildlife Area in December, 1983, and January and Februar~, 1984. North South Month slope Ridgetop Statistic slope Bottom December January February x Range x Range x Range - 13 7-19 47 31-57 55 42-71 1 0-5 29 13-37 20 0-32 9 3-15 38 23-48 41 19-53 10 8-14 45 41-52 49 28-60 �Table 2. Number of deer counted in the eastures at the Little Hills Wildlife Area, 13-15 December 1983. co O"l Day 1 Pasture ]. M 2 3 No. of marked deer '2 2 8 3 16 3 Quadrat size (ha) Day 2 AM PM 1 U M 6 0 8 10 19 5 14 4 9 9 12 12 9 14 13 6 14 12 M U M 64.75 129.50 64.75 129.50 4 2 14 1 2 6 19 23 8 3 15 18 23 11 7 4 16 64.75 129.50 13 13 10 12 13 8 9 14 13 . 12 5 8 64.75 129.50 5 8 4 7 8 2 6 5 4 AM M U 3 PM AM M 9 Day 3 = marked deer; U = unmarked deer Only 7 marked deer were present during the AM and PM counts on Day 1. Only 15 marked deer were present during the AM and PM counts on Day 3. U 1 2 PM M U 13 4 20 4 8 12 4 13 14 22 9 9 1 4 14 12 7 13 8 13 11 14 11 11 16 17 5 3 6 4 1 2 6 4 9 U 3 7 5 3 3 �- - - -- - Table 3. Number of deer counted in the ~astures at the Little Hills Wildlife Area, 10-13 Januar~ 1984. Day 1 Day 2 Day 3 Pasture No. of marked deer 2 7 3 1 16 AM PM Search direction M U N-S E-W 3 1 N-S E-W 1 PM M~ AM PM M U M U M U M U M U 15 6 2 0 12 14 4 2 4 14 16 3 3 19 16 8 14 2 0 17 9 11 8 7 6 6 9 6 5 12 8 9 6 12 7* 6 4 16 N-S E-W 11 9 13 12 13 7 17 10 10 5 5 8 N-S E-W 7 6 6 2 5 1 7 5 8 2 3 11 12 13 10 6** 5 4 2 8 6 3 5 3 9* .4* 7* 11 3 12* 7 1 7* 9 11* 13 2** 9 8** 11 5 4 1 0 4 5 4 = marked deer; U = unmarked deer * There were 17 marked deer present for these counts. ** There were 15 marked deer present for these counts. M o .... �~ Table 4. Number of deer counted in the pastures at the Little Hills Wildlife Area, 21-24 February, 1984. Day 1 Day 2 Day 3 Day 4 Pasture 1 M No. of marked deer AM r~?--u t~ AM PM PM AM PM U M U M U M ~ U M U "10 3 2 16 11 16 11 4 15 9 16 9 5 3 16 13 8 11 5 13 7 10 9 15 7 1 4 4 24 10 10 16 20 17 21 18 14 22 22 12 18 = marked deer; U = unmarked deer �89 Table 5. Proportions of marked deer counted in the pastures at the Little Hills Wildlife Area during winter 1983-84. Month Pasture P SE N December 2 0.40 3 5 2 Pooled 0.71 0.71 0.70 0.71 2 0.38 0.048 42 3 0.58 0.68 0.77 0.068 0.049 0.088 98 94 0.67 0.036 0.71 0.100 96 0.66 0.68 0.69 0.127 0.070 0.052 96 144 4 January 3 4 5 0.067 0.054 0.046 0.043 0.032 92 188 192 96 48 2 Pooled 2 February 3 4 Pooled 4 All months 0.69 ± 0~04 (95% conf. limits) - 1 Proportions based on combined counts on 65-ha and 130-ha quadrats. 2 3 Calculated without Pasture 2 data. Does not include counts made in east-west directions. 4 Calculated without Pasture 2 data for December and January. �90 Figo 1. Pastures 20 ), 4p winter range (dashed lines) 12905-ha quadratsa and 5 and the areas of unfenced added to each to appr-oxdmat e �91 N~D _- IS'{) 1¥tJ 111) pJt1 IUJ 0 ,/ I nO I I I 1fJ() .tt.. to ~ o ~ 7f) 0 I ! I \ I V 0 4 &" ". Sf) " 'to 3@ Fe. h. 2,f) If) r- I ----T-----~~----_r~-I 3 t"" S Figo 2. Trends in num.bers of marked deer (solid lines) and total deer (dashed lines) counted on successive flights in the pastures each month. ��93 Colorado Division of Wildlife Wildlife Research Report July 1984 JOB PROGRESS REPORT State of Colorado Project No. 45-01-502-15050 Work Plan No. Job. No. 12 Period covered: Authors: Personnel: 2 ------------------ Big Game Investigations - Cervids Deer Investigations Determination of Mule Deer of Body Composition 7/1/83-6/30/84 S. C. Torbit L. H. Carpenter, R. f~. Bartmann, A. W. Alldredge, R. A. Garrott, and S. Boyle P. H. Neil, ABSTR.L\CT Chemical analysis was completed for 24 adult female mule deer collected from Piceance Creek winter ranges during 1982-83 winter. Fat content ranged from 15.1% in December to 3.8% in April samples. Significant decreases were detected in fat/ash ratios between December and February collections and February and April collections. Wet protein/ash ratios declined significantly between October and December samples and increased significantly between February and Aprl l samples. Left kidney fat index was found to be positively correlated with total fat reserves. Stepwise multiple regression provided equations to predict body composition from external measures of body size, but correlations were weak. Body composition analysis of fetuses removed from pregnant does showed little variation over time, since water comprised the greatest percentage of fetal components (i = 85%). Thirty-two fa.wns were collected during 1983-84 winter from Piceance Creek winter ranges. Thirty-two fawns were collected in October, December, February and April (8 each month). Eight additional fawns that died during late winter were retrieved by R. A. Garrott. All 40 fawns were processed for body composition analysis, however, chemical analysis is not yet complete. Mule deer fawns entering Piceance winter ranges (October) were shown to be composed of little fat (i = 5.32%). Measures of external body size appear to be less variable than those for adults collected the previous winter. Chemical and statistical analysis \'1i11be completed next segment. ��95 DETERMINATION OF MULE DEER BODY COMPOSITION Stephen C. Torbit P. N. OBJECTIVES 1. Detennine body composHion and describe the relationship between body water~ body fat and lean body tissue of mature female and fawn mule deer inhabiting the Piceance Basin winter range of western Colorado. 2. Compare chemica1 estimates of total body fat to kidney fat and bone marrow fat to determine validity of these indices as indicators of animal condition. , ,t 3. Evaluate variances of measured parameters of mule deer and if results warrant, develop a detailed study plan to investigate winter dynamics of body composition in free-ranging mule deer. SEGMENT OBJECTIVES 1. Complete chemical and statistical analysis of 24 adult female collected during 1982-83 winter. 2. Collect 32 mule deer fawns during 1983-84 winter to establish relationships between body water. body fat and lean body tissue. METHODS AND MATERIALS The 4 components of mammalian body composition are water, fat~ protein and ash (Torbit 1981). Since water is considered a separate component. fat, protein and ash components make up total dry matter and when discussed individua lly ~ are expressed on a dry matter basis. However, dry protein can be converted to a wet protein component by utilizing the relationship of Pace et a1. (1947) that considers mean water content of protein to be 72.6%. Total body water for females collected in April ~ 1983, was corrected for amniotic fluid volume. All deer collected in April were pregnant and amniotic fluid was included with the dam when samples were prepared for chemical analysis. During late wi nter , 1984, pregnant females were collected from several winter ranges in western Colorado as part of another study. Reproducti ve tracts from these fema 1es were collected. fetal measurements made and amniotic fluid measured (Table 1). Mean values of amniotic fluid ~<Jereused to reduce total body water values for females collected during April, 1983. Thirty-two male and female mule deer fawns were collected f rom the Piceance Basin during the 1983-84 winter. Eight deer were collected in October, December. February and April. Deer were either trapped in Clover traps (Clover, 1956) and euthanized with a drug overdose or shot. All eight deer collected in October and April were shot with a small caliber rifle. �96 Shooting was necessary since deer were not attracted to Clover traps at those times, Jnmedi ate ly after death deer were weighed to the nearest 0.5 kg and the following measurements taken: total body length, chest girth, and left hind foot length. Animals were wrapped in plastic bags, frozen and transported to Fort Collins, where they remained frozen until processed for chemical analysis. Carcasses were reweighed, thawed and prepared for analysis as described elsewhere (Torbit 1981). Kidney fat index (Anderson and f~edin 1965) was determined for both kidneys on all 32 deer and bone marrow was removed from the left femur and analyzed for fat content from an deer. Samples from the minced carcass were collected to assay for water, fat, protein and ash (To~bit 1981). Eight additional fawns were collected soon after death from starvation by R. A. Gar rot.t from Piceance Basin. All fawns were frozen, processed and submitted for chemical analysis as outlined above. Differences in body components between months were analyzed by 2-sample t-Les ts . External measurements of body size were related to size of protein and fat s tores by s tep-wise mul t t var i ate regression procedures. RESULTS AND DISCUSSION Adult Females Chemical analysis i s now complete for all adult female mule deer collected during 1982-83 winter. Maximum fat content (15.1%) occurred in deer collected in December, 1982, and fat was depleted by April, 1983 (Table 2). Protein concentration appeared to remain constant throughout winter collections (Table 2). To correct for differences in skeletal sizes among deer, dry protein, wet protein and fat components were divided by the ash component (Table 3), that procedure provided a more detail~d view of catabolic processes. Statistical analysis showed that significant differences occurred in Fat:Ash ratios between December and February samples (P = 0.09) and February and April samples (P ::;0.004)(Fig. 1). ~Jet Protein:Ash ratios were found to be signif-jcant'ly different between October and December samples (p == 0.07) and February and April samples (p = 0.004). No differences occurred between December and February sampl es (p = 0.54) or October and April samples (p = 0.79)(Fig. 2). The dynamics of fat and protein stores during winter (Figs. 1 and 2) revealed a pattern of catabolism character i zed by an early protein loss and delayed fat response to starvation. That general pattern supported results of our studies of catabolic process with hand-reared mule deer (Torbit 1981). Additionally, results showed a significant increase in size of protein stores during spring (Fig. 2). This increase may indicate a period of compensatory growth of protein stores during later winter and early spring. �97 Kidney fat indices declined throughout 1982-83 winter reflecting general loss of fat (Table 4). A significant relationship was detected between total body fat and left kidney fat inde . That relationship (Fig. 3) (% Total Body Fat = -6.84 + 4.3 1n LKFI, r2 = 0.85) was found significant at the 5% probability level. The form of the equation relating left kidney fat index to total body fat was incorrectly reported on page 92 of the 1983 progress report. The equation reported in the present progress report is correct. A highly significant relationship was not detected between total body fat and right kidney fat index. Bone marrow fat values (Table 4) were highly variable over time. Procedures used to assay and calculate bone marrow fat are currently being reviewed to ascertain the validity of reported values. t , t I Step-wise multiple regression analysis was performed to develop a non-destructive method of estimating mule deer body composition. Body size measurements (Table 5)(Field Weight, Total Body Length, Chest Girth and Left Hind Foot Length) were used to estimate condition (% Fat and % Protein). Regression analysis revealed that a predictive equation could be developed but the equations listed below may not provide sufficient precision or accuracy for detailed body composition studies. The inability to generate highly correlated regressions may be due to the dynamic nature of both fat and protein during winter. a) ~b Body Fat = 540 + 4.5 Chest Girth - 2.8 Total Body Length - 66.5 Total Body Length Field Weight + 109 Chest Girth Field Weight _ Chest Girth 866 Total Body Length r2 = 0.52 b) % Protein = -55.4 = 0.9 Field Weight + 0.8 Chest Girth + 0.8 Total Body Length + 48 C~est Gi~th _ Fleld Welght 0.6 Total Body Length + Chest Girth Field Weight r2 = 0.42 Fetal body composition (Table 5) provided little definitive information regarding changes in fetal body composition. Little variation occurred in fetal fat and protein components since water comprised the greatest proportion of fetal body components Cx = 85%). fvluleDeer Fawns Thirty-two fawns (16 males, 16 females) were collected during 1983-84 winter. Eight additional fawns that died during later winter and were subjects of a companion study, were also processed for body composition ana 1ysis. ' Chemical analysis has not been completed for all samples, however, fawns collected in October were composed of very little fat Cx.= 5.32%) upon entering Piceance winter ranges (Table 6). �98 Body size measurements and kidney fat indices (Table 7) for fawns collected during 1983-84 winter appear to be less variable than those for adult females (Table 5). Regression analysis will be performed upon completion of chemical assays to establish relationships between total body fat and kidney fat index and total body fat and bone marrow fat. Additionally, attempts will again be made to develop a procedure to predict body composition from external measures of body size. Body size and kidney fat index data for fawns found dead on Piceance Creek winter ranges are presented (Table 8). Again, chemical analysis for these fawns have not been completed. The starved fawns lost an average of 23.58% of their body weight before death and kidney fat indices of these animals showed more variability between left and right kidney than did fat indices of collected fawns. LITERATURE CITED Anderson, A. E., and D. E. Medin. 1965. Two condition indices of the Cache La Poudre mule deer herd and their application to management. Colo. Game, Fish and Parks Game Infor. Leafl. No. 23. 3pp. Clover, M. R. 1956. Single gate deer trap. Calif. Fish and Game 42(3):199-201. Pace, N., L. Kline, H. K. Schachman, and M. Horfenist. 1947. Studies on body composition. IV. Use of radioactive hydrogen for measurement in vivo of total body water. J.B.C. 168:459. Torb i t , S. C. 1981. In vivo estimation of mule deer body composition. Ph.D. Diss., Colorado State Univ., Fort Collins. 98pp. I t I �Table 1. Fetal mule deer weights (g) and amniotic fluid volume (ml) retrieved from adult female collected from western Colorado winter ranges February and March 1984 Fetal wt. (g) left Fetal wt. (g) right Total fetal wt (g) Amniotic fluid vol (ml) 248.4 . 144.8 508.5 315.6 3750.0 1658.0 19 20 260.1 170.8 231.1 225.6 202.9 231.1 428.5 975.0 2520.0 21 25 98.4 157.4 211. 6 322.9 26 27 28 31 494.7 479.6 213.3 168.0 92.7 1072.0 1021.8 213.3 168.0 1780.0 1410.0 1905.0 1850.0 720.0 760.0 95.0 187.7 1255.0 235.6 261.2 16546.0 128.6 31335.0 178.0 379.9 109160.0 330.4 1691.8 690642.0 831.0 Deer ID 17 18 t J t 32 - x s2 s 113.2 165.5 577 .3 542.2 �100 Table 2. Chemical composition (%) of female mule deer collected from Piceance Basin October, 1982 - A~ri), 1983 Deer No. % % % % Water Fat Protein Ash 63,60 65.57 65.34 61.07 64.06 63.70 10.38 10.01 9.61 11.82 11. 74 10.96 19.88 19.88 19.55 20.77 19.74 20.08 6.15 4.58 5.50 4.45 4.90 63.89 2.17 1.47 10.75 0.69 0.83 19.98 0.15 0.39 5.18 0.35 0.59 64.20 60.15 65.39 55.63 63.67 63.48 9.02 14.06 9.80 15.1:0 9.91 10.76 20.59 19.64 19.96 21.90 19.91 19.90 6.15 5.60 4.76 7.24 6.48 5.49 62.10 10.66 3.27 11.44 5.27 2.30 20.32 0.58 0.76 5.95 0.62 0.79 66. 13 65.61 69.39 67.33 63.43 67.10 9. 15 10.34 6.32 7.75 11.28 19.29 18.06 5.40 5.92 5.86 5.50 5.69 5.74 66.50 3.29 1.81 8.87 2.67 1.64 lB.95 0.32 0.57 5.69 0.03 0.18 69.92 5.99 1.68 2.25 5.64 1.63 5.57 lB.72 18.94 20.12 19.66 17.29 19.B7 5.40 5.22 5.13 4.51 3.83 5.29 3.79 3.82 2.14 19.10 0.90 1.04 4.90 0.31 0.61 October 1 2 3 4 5 6 x s2 s December 1 2 3 4 5 6 ---x2 s s 5.51 I I t February_ 1 2 3 4 5 6 x s2 s 8.3B lB.44 19.44 19.67 18.80 A~ril 1 2 3 4 5 6 - x2 s s 72.20 72.49 70.18 77 .25 69.29 71.B8 7.13 2.92 4 �101 Table 3. Chemical composition (kg), protein and fat depot size corrected for body size for adult female mule deer collected from Piceance Basin October, 1982 - A~ril, 1983 Protein Wet prot. Fat Deer Field Empty Total Total Total Total 1. D. ash wt. body wt. H?O fat protein Ash Ash Ash (kg) (kg) (kg) (kg) (kg) (kg) t I t 1 3.30 2.10 3.10 2.16 2.03 2.49 3.23 4.33 3.55 3.77 4.43 4.10 4.06 5.63 4.44 5.03 5.78 5.20 1.69 2.18 1.75 2.14 2.64 2.24 54.32 48.57 18.94 32.72 4.35 5.72 31. 10 5.19 16.82 0.17 4.10 0.42 9.69 1.24 1.11 2.53 0.25 0.50 3.90 0.18 0.43 5.02 0.37 0.61 2.11 0.12 0.35 66.80 64.60 67.20 69.90 57.10 56.70 59.18 57.21 59.71 63.93 49.42 49.39 38.00 34.41 39.04 35.63 31.47 31.35 5.34 8.04 5.85 9.66 4.90 5.31 12.19 11.24 11.92 14.00 9.84 9.83 3.64 3.21 2.84 4.63 3.20 2.71 3.35 3.50 4.19 3.02 3.08 3.63 4.10 4.35 5.16 3.61 3.93 4.63 1.47 2.50 2.06 2.09 1.53 1.96 63.72 56.47 25.61 28.98 5.06 5.38 34.98 8.59 2.93 6.52 3.02 1.74 11.50 2.50 1.58 3.37 0.41 0.64 3.46 0.15 0.39 4.30 0.25 0.50 1.94 O. 15 0.38 Feb. Feb. Feb. Feb. Feb. Feb. x2 s s 1 73.30 61.84 2 52.90 45.64 3 72 .60 68.14 4 69.70 65.82 5 68.80 60.47 6 66.40 62.63 40.89 29.94 47.26 44.32 38.36 42.02 5.66 4.72 4.31 5. 10 6.82 5.25 11.93 8.24 12.57 12.-79 11.90 11.77 3.34 2.70 3.99 3.62 3.44 3.59 3.57 3.05 3.15 3.53 3.46 3.28 4.39 4.06 3.83 4.29 4.25 4.04 1.69 1.75 1.08 1.41 1.98 1.46 67.28 60.76 46.71 52.24 6.83 7.23 40.47 29.81 5.46 5.31 0.63 0.80 11.53 2.77 1.66 3.45 O. 15 0.39 3.34 0.04 0.20 4.14 0.03 0.19 1.56 0.08 0.29 Apr. Apr. Apr. Apr. Apr. Apr. 1 2 3 4 5 6 57.30 62.50 59.20 60.50 54.90 62.00 32.91 35.69 33.49 35.33 34.05 35.47 2.82 0.80 1.04 2.84 0.72 2.85 8.81 9.36 9.29 9.90 7.62 10.17 2.54 2.58 2.37 2.27 1.69 2,71 3.47 3.63 3.92 4.36 4.51 3.75 4.54 4.69 5.07 5.56 6. 12 4.76 1.11 0.31 0.44 1.25 0.43 1.05 34.49 1.85 1.13 0.99 1.17 0.99 9.19 0.32 0.91 2.36 0.11 0.33 3.94 0.14 0.38 5. 12 0.31 0.55 0.77 O. 17 0.42 Dec. Dec. Dec. Dec. Dec. Dec. 1 2 3 4 5 6 x s2 s • t 53.63 45.79 56.36 39.18 45.59 50.86 10.66 9.10 11.02 8.14 9.00 10.21 2 3 4 5 6 - ~ 56.40 54.40 59.50 46.80 50.80 58.00 5.57 4.58 5.41 4.63 5.35 5.57 Oct. Oct. Oct. Oct. Oct. Oct. x s2 s - x2 s s 47.07 49.43 46.17 50.34 44.08 51.19 59.40 48.05 7.05 6.21 2.65 2.73 34.11 30.02 36.83 23.93 29.20 32.40 �102 Table 4. Body size measurements (cm), left and right kidney fat indices and bone marrow fat (%) for adult female mule deer collected from Piceance Basin October; )982 - April, 1983 Left hind Deer Chest Length foot RKFI LKFI Bone marrow 1.D. girth (cm) (cm) length (%) (%) fat (%) Oct. Oct. Oct. Oct. Oct. Oct. 1 2 3 4 5 6 x2 s s Dec. Dec. Dec. Dec. Dec. Dec. 1 2 3 4 5 6 - x2 s s Feb. Feb. Feb. Feb. Feb. Feb. 1 2 3 4 5 6 v "2 s s Apri 1 April April April April April x2 s s 1 2 3 4 5 6 88.00 90.00 91.00 87.00 89.00 95.00 149.00 143.00 142.00 130.00 140.00 144.00 46.00 46.00 46.00 45.00 46.00 45.00 90.00 6.67 2.58 141. 33 33.22 5.76 45.67 0.22 0.47 94.50 91.50 90.50 97.40 90.50 87.00 157.00 154.50 155.50 151.00 147.00 139.00 49.00 48.30 47.00 45.00 48.00 48.00 91.90 10.85 3.29 150.67 37.97 6.16 98.00 89.50 93.00 91.00 90.00 87.50 a 87.65 44.03 34.19 78.16 72 .61 88.24 88.61 83.75 86.46 86.02 89.08 88.24 67.48 439.38 20.96 87.03 3.38 1.84 28.57 65.05 57.73 103.00 63.54 34.03 34.55 71.98 55.45 123.40 74.09 55. 14 9.02 14.06 9.80 9.91 10.76 47.55 1.65 1.28 58.65 590.60 24.30 69. 10 759.43 27.56 10.71 3.11 1.76 155.00 141 .00 153.00 147.50 147.50 149.50 47.00 46.00 47.00 47.00 44.80 46.50 41.38 57.32 16.25 31.47 44.69 61.97 31. 58 42.56 14.63 31. 58 35.52 51.95 b b 6.32 7.75 11.28 8.38 9-1.50 11.17 3.34 148.92 20.12 4.49 46.38 0.63 0.80 42.18 235.81 15.36 34.64 130.38 11.42 8.43 3.26 1.81 87.00 90.50 89.50 88.50 88.00 93.00 148.00 160.00 146.00 153.00 159.00 159.00 45.00 47.00 45.50 48.00 46.00 46.50 24.28 20.21 9.30 19.27 10.98 25.00 22.03 8.98 10.80 23. 16 6.13 26.75 91. 93 4.93 3.07 72.50 0.25 86.91 89.42 3.78 1.95 154.16 31. 14 5.58 46.33 0.97 0.98 18.17 36.62 6.05 16.31 62.73 7.92 43.27 1677 .00 40.96 aRight kidney not collected in October blmproper leg bone collected - data unavailable b ,, f I • • �~ ~- -- - - ._.~ - - Table 5. Total weight (g) and chemical composition (%, kg) of fetal mule females collected from Picenace Basin October, 1982 - A~ril, 1983 H20 wt /0 Total % % % wt (g) Dam 1.D. H 20 Protein Ash (g) Fat Feb. 1 194.0 86.77 9.30 2.50 168.33 1.43 Feb. 2 87.5 1.22 1.70 88.54 8.54 77 .47 Feb. 3 257.5 87.85 8.36 2.49 226.21 1.31 Feb. 4 164.0 89.67 1.85 147.06 1.00 7.48 Feb. 5 261.0 2.20 229.50 87.93 1.20 8.67 Feb. 6 201.0 180.52 89.81 1.18 7.25 1.76 01 x 2 s s April April April April April April -- x2 s s 1 2 3 4 5 6 deer removed from adult Protein wt (g) Ash wt 2.77 1.07 3.37 1.64 3.13 2.37 18.04 7.47 21.53 12.27 22.63 14.57 4.85 1.49 6.41 3.03 5.74 3.54 Fat wt (g) (9) 194.17 3468.70 58.90 88.43 1.13 1.06 1.22 0.02 O. 13 8.27 0.49 0.70 2.08 0.11 0.33 171.52 2648.00 51.46 2.39 0.66 0.81 16.09 27.90 5.28 4.18 2.80 1.67 1676.00 1830.00 1251.00 1930.00 1453.00 954.00 82.68 87.13 85.88 86.37 85.15 83.15 1.67 1.15 1.35 1.51 1.65 1.58 12.26 8.71 10.39 9.84 11.06 12.91 3.39 3.01 2.38 2.28 2.14 2.36 1385.72 1594.48 1074.36 1666.94 1237.23 793.25 27.99 21.05 16.89 29.14 23.97 15.07 205.48 159.39 129.98 189.91 160.70 123.16 56.82 55.08 29.77 44.00 31.09 22.51 1515.67 114.30x103 338.00 85.06 2.67 1.63 1.49 0.03 0.18 10.86 2.01 1.42 2.59 0.20 0.45 1292.0~ 9.00xl0 3.00xl02 22.34 27.53 5.25 161.44 868.00 29.47 39.88 169.40 13.01 I-' a w �Table 6. Field weight (kg) and chemical composition (% and kg) for mule deer fawns collected from Piceance Basin, October, 1983. kg Field Empty body Deer % wei qht (kg) weight (kg) % H2O kg H2O kg Fat Protein I.D. % Fat Protein % Ash Oct. 1 Oct. 2 Oct. Oct. Oct. Oct. Oct. Oct. 3 x2 s s 4· 5 6 7 8 29.60 29.80 26.60 24.70 24.70 25.00 29.80 27.20 25.4·0 26.30 23.70 21.30 21.00 20.70 25.60 24.30 66.4·0 70.80 69.80 73.00 74.00 27.20 4.62 2.15 23.50 4.43 2.11 a Chemical analysis being repeated. kg Ash 0.55 0,47 4.67 4.42 4.34 3.03 3.54 1.07 1.13 1.02 17.80 16.30 1. 17 1. 76 4.58 4.55 1.40 1. 20 16.70 1. 13 1.06 1.30 0.40 0.63 4.16 0.34 0.58 1.17 0.01 0.12 9.58 5.08 5.93 2.60 2.23 18.38 16.79 18.30 14.23 16.85 4.50 4.79 4.50 5.29 4.88 16.90 18.60 16.50 15.50 15.50 69.40 67.00 4.56 7.26 17.90 18.74 5.45 4.95 70.10 6.85 2.62 5.32 5.70 2.39 17.31 2.07 1. 44 4.91 O. 11 2.43 1.34 !-' 0 ..j:.;. 1.41 1.14 1.26 a 0.34 �..•.. ,...- -- - ...._....._.. - -._.. Table 7. Field weight (kg), empty body weight (kg), body size measurements (cm) and left and right kidne~ fat indices for mule deer fawns collected from Piceance Basin October~ 1983 - A~ril, 1984 Right kidney Left kidney Left hind Empty Chest Total body fat index fat index ft length body girth length Field (%) (%) (cm) wt (kg) (cm) (cm) Sex wt (kg) Oct. Oct. Oct. Oct. Oct. Oct. Oct. Oct. 1 2 3 4 5 6 '1 8 F M F F M M M M - x2 s s Dec. Dec. Dec. Dec. Dec. Dec. Dec. Dec. 1 2 3 4 5 6 7 8 M F M M M M F M - x2 s s Feb. Feb. Feb. Feb. Feb. 1 2 3 4 5 F F M F M 29.60 29.80 26.60 24.70 24-.70 25.00 29.80 27.20 25.40 26.30 23.70 21.30 21.00 20.70 25.60 24.30 68.60 71.00 66.50 65.40 65.50 68.80 70.50 69.30 115.50 115.50 112.00 114.50 111.00 111.40 115.70 110.90 38.50 39.50 38.00 38.20 39.30 36.80 40.50 39.30 55.86 29.03 30.34 13.83 10.84 23.15 15.70 35.71 36.04 34.78 32.05 18.18 14.63 26.47 12.07 40.40 27.20 4.62 2.15 23.50 4.43 2.11 68.20 4.11 2.03 113.30 4.15 2.04 38.80 1.12 1.06 26.81 187.61 13.70 26.83 100.13 10.01 33.60 30.40 31.80 35.00 37.00 32.70 32.10 31.80 29.40 26.50 27.40 31.20 32.20 28.80 28.30 28.10 74.50 74.00 78.60 80.00 80.00 76.00 74.50 75.50 129.00 123.00 114.30 125.10 129.50 123.40 123.50 121.00 40.00 39.50 41.80 . 42.40 42.50 40.00 40.00 40.40 11.38 27.87 08.33 20.14 11.21 8.47 19.53 52.10 13.49 30.09 10.08 21.54 6.50 13.16 24.19 37.01 33.10 3.86 1.96 29.00 3.17 1.78 76.60 5.53 2.35 123.60 19.99 4.47 40.80 1.28 1.13 19.88 189.10 13.75 19.51 97.37 9.87 16.60 29.90 32.30 30.60 29.00 13.20 25.20 28.00 24.80 23.10 60.00 75.00 74.00 77.80 70.60 100.00 123.00 126.40 128.00 125.50 35.50 42.00 42.00 40.50 41.60 5.63 12.88 15.04 12.17 4.34 8.00 10.85 16.92 16.16 7.91 •.....• 0 U1 �Table 7. Feb. 6 Feb. 7 Feb. 8 (continued - page 2) F 28.80 22.20 M 24.00 18.80 M 32.90 28.00 x? s~ s April Apri 1 April April Apri1 April April April -- x2 s s 1 2 3 4 5 6 7 8 F F t~ F F F F F I-' 0 71.80 70.20 75.80 124.50 119.30 125.30 41.80 41.50 43.20 4.92 3.42 6.82 7.83 5.26 7.35 28.00 25.10 5.00 23.30 23.90 4.89 71.90 26.30 5.13 121. 50 71.90 8.48 41.00 4.82 2.20 8.15 17.68 4.21 10.04 16. 14 4.02 24.40 25.60 27.50 26.20 22.50 31.50 26.50 28.60 19.30 23.20 24.90 21. 10 20.00 29. "10 22.60 25.00 72.50 74.00 68.50 70.80 65.00 74.00 72.50 70.00 123.00 118.00 127.00 123.50 116.00 127.00 116.50 122.00 40.00 40.00 42.00 41.00 38.00 42.00 40.00 41.50 6.42 5.21 5.17 4.95 4.04 5.60 3.06 6.21 3.00 8.70 8.77 5.36 6.00 6.77 3.77 6.25 26.60 6.46 2.54 23.20 8.89 2.98 70.90 8.22 2.87 121. 60 16.80 4.10 40.60 1.59 1.26 5.08 1.07 1.03 ~ _ ..• -- - .•. . 6.08 3.74 1.94 0"1 �- Table 8. Body weight (kg), empty body weight (kg), and left and right kidney fat indices for dead mule 1984. Empty Change Deer Capture Death body in body wt (kg) wt (kg) wt (kg) wt (%) 1.D. 149.450 193.510 193.610 193.630 193.650 193.690 193.720 193.820 - x s2 s -- -- change in body weight (%), body size measurements (cm) deer fawns collected from Piceance Basin during late winter Chest Girth (cm) Total body length (cm) Left hind . Left kidney Right kidney fat index foot length fat index (cm) (em) (cm) 26.60 33.40 33.70 30.70 31.10 29.50 28.60 33.30 20.00 25.20 23.80 23.20 22.70 25.30 21.20 27.30 16.60 23.50 19.60 21.40 18.90 23.60 17.30 25.50 24.80 24.50 29.40 24.40 27.00 '14.20 26.20 18.00 75.80 75.50 74.50 70.00 70.50 75.00 77.00 73.50 118.00 128.00 127.50 116.00 122.00 117.00 123.50 129.00 40.00 40.50 42.50 40.00 40.00 39.00 40.80 43.00 7.10 4.70 6.80 7.80 11.90 11.30 11.20 8.80 10.20 27.20 8.30 14.60 9.60 14.30 9.80 6.60 30.90 5.70 2.40 23.60 4.50 2.20 20.80 9.10 3.00 23.60 21.70 4.70 74.00 5.50 2.30 122.60 23.90 4.90 40.70 1.60 1.30 8.70 5.70 2.40 12.60 37.10 6.10 ,_. C> --.J �108 2.0 1.5 1.0 I , t 0.5 Oct Dec COLLECTION Feb Apr il DATE Fig. 1. Ratio of mean fat weight (kg) and mean ash weight (kg) plotted against collection date for 24 female mule deer collected from the Piceance Basih during 1982-83 winter. Oct. = Dec. p = 0.54 Dec. f Feb. p = 0.09 Feb. r April p = 0.004 x x x x x x �109 5.5 ...- --- -._ - 0' .::t:. .x c: <1> +- .J:: 5.0 ~ 0 0' Q_ ~ +- .c: ~ « 4.5 L.. <1> CI) I ) t c: 0 Q) ~ c 0 <1> ~ 4.0 Oct Feb Dec Collection Apr Date Fig. 2. Ratio of mean wet protein weight (kg) and mean ash weight (kg) plotted against collection date for 24 female mule deer collected from the Piceance Basin during 1982-83 winter. X Oct. f x Dec. x Dec. = x Feb. p = 0.07 p = 0.54 x Oct. = x April P x Feb. f x Apri 1 p = = 0.004 0.79 �110 • 12 ~ 0 ..•...•• +- if 8 ~ "'0 0 OJ f c 1 -e- 0 ~ 4 o----~----~----~--~----~--~ 40 80 120 Left Kid ne y Fat Index (%) Fig. 3. Relationship between left kidney fat index (%) and total body fat for 24 female mule deer collected from the Piceance B s!n during 1982-83 winter. (% TBF = -6.84 + 4.3 ln LKFI r 2 - 0.85). �111 Colorado Division of Wildlife Wildlife Research Report July 1984 JOB PROGRESS REPORT State of Colorado Project No. 45-01-502-15050 Big Game Investigations - Cervids Hork Plan No. Elk Investigations Job. No. 2 -------------------- Period Covered: Author: 3 Elk Population and Ecology Study 7/1/83-7/30/84 G. D. Bear ABSTRACT A total of 78 elk (36 cows, 33 calves, 6 spikes, 3 bulls) were captured in the North Park ar·eaduring January and February, 1984, with Clover traps. Forty-five yearling and adult elk were fitted with telemetry collars. Telemetered elk will be monitored on a weekly basis to determine seasonal ranges and migration patterns. A mixture of nyanzol dye applied with a portable sprayer produced numbers on the sides of the elk which could be read from an aircraft. These numbers provided a short-term marker for identifying individual elk from the air. Only two calves were captured and marked during the calving season. ��113 ELK POPULATION AND ECOLOGY STUDY George D. Bear P. N. OBJECTIVES 1. Develop techniques to more accurately and precisely estimate elk population levels. 2. Define natality and mortality characteristics of elk in North Park, Colorado. 3. Evaluate the relationship of the hunter harvest to the population structure. SEGMENT OBJECTIVES 1. Capture and place 50 radio collars on 50 adult elk in North Park then monitor movements on a weekly basis to determine seasonal home ranges and migration patterns. 2. Capture, weigh, measure, and mark 20~30 calves with mortality collars to derive an estimate of mortality rates and cause of mortality. METHODS AND MATERIALS For a complete description of methods used. refer to the Program Narrative (Bear 1983; DOW Files). RESULTS AND DISCUSSION A total of 78 elk (36 cows, 33 calves, 6 spikes, 3 bulls) were captured on seven different locations in the North Park study area during January and February. Forty-five of the yearling and adult elk were collared with telemetry units equipped with mortality sensors. Telemetered animals will be located on a weekly basis throughout the year to establish seasonal ranges and determine interchange between the large winter herds. UTMs are determined for all locations plotted for each of the collared elk. These data will be summarized later. Mixtures of nyanzol dyes and leather dye was used to mark the identification number on the elk. A l-quart (.95 1) portable sprayer, a l-pint (.47 1) portable sprayer, and a weed sprayer were used to apply dye solutions. The quart sprayer was found to be most suitable. It was easy to transport and contained sufficient dye and air to mark several elk. and had the proper size nozzles to apply the dye. The pint sprayer was a very handy size and had the same nozzles, but it contained only enough dye and air to mark a couple of elk before being recharged. The weed-sprayer was too large and delivered too much spray so there was a problem with the numerals running together. However, it could be easily recharged in the field. �114 Two colors of nyanzol dye (black and orange) and a black leather dye were tested. The leather dye was not sufficiently durable. It provided a very good number during the first month after application but by the second month, about one-third of the numbers had faded to the extent that they were difficult to read. The orange nyanzol dye did not provide enough contrast with the color of the elk to be legible. The black nyanzol dye produced durable and legible numbers. A solution of 600 cc alcohol, 400 cc of dye solution, 250 cc of gum arabic solution was mixed, allowed to stand 1-2 days~ then 250 cc of 30% hydrogen peroxide added before using this solution produced acceptable results. More gum solution plugged the sprayer nozzle and best results were achieved when the gum solution was added after the dye solution cooled. Successful marking of the elk depended on application procedure as well as composition of the dye. The dye needed to be applied with quick even strokes. If the hair were saturated, the dye ran through the underfur and resulted in a blotted number. Numbers needed to be made larger than 12 in (30.5 cm) so they would not run together. The number should be placed high on the side and back of the elk to make it visible from an aircraft. Due to the size of-the numbers, they extended far down onto the sides of a calf. Preliminary observations indicated this system of marking individual elk has merit. However, it may be difficult to read the numbers when flying directly over an elk and when only quick glimpses are made as elk run through heavily forested areas. Also, it is not always possible to mark elk on the same side when handling them. A double marking system will be attempted when implementing this technique next winter. Possibly, the large numbers can be supplemented by ear-tag-streamers. Plastic and cloth streamers were used this year. Some plastic ribbons were lost during the two-month period, so a more durable material should be used such as vinyl or cloth. The plastic ribbon was tried in an effort to find a marker that would not carry-over into the next year; however, it was not durable enough. Only two calves were captured and collared during June. Several factors may be the cause of such poor success: poor weather conditions, inadequate helicopter time, poor calf crop, and investigator1s lack of knowledge of the calving areas. This phase of the study should be suspended for the present time due to budget constraints and poor success. /-f A Prepa red by --=-<_/_--::.~_---"'--;"::-c-c_._-'r-::---_---_·-'C-_-_' George D. Bear' Wildlife Researcher 0:)-(,,---'-------_ �115 APPENDIX A ELK TRAPPING - NORTH PARK January 5, 1984 - February 16, 1984 Location of Traps Independence Mt. - along the Loop Road (5 traps) Sentinel Mt. - basin on north side above railroad tracks behind Davis Ranch (4 traps) Buffalo Ranch - along side of road in litheGap"; before ranch house (4 traps) Swift Ranch - approximately 1 1/2 mi (2.4 km) north of Lake John Road and 1/2 mi (0.8 km) west of North Platte (3 traps) Baller Ranch - 1/4 mi (0.4 km) south of main ranch house on Michigan River and Hwy 14 (2 traps) Owl Ridge - Speck's Draw approximately 1 mi (1.6 km) SW of Junction of Gould-Rand Road with Owl Creek (5 traps) Owl Ridge - State property on east side of road when Gould-Rand Road crosses Owl Ridge (4 traps) Marking of Elk Elk were marked with metal ye11o~ colored eartags. The Fort Collins address was on the tag with year and animal number (84-XXX). Blue or orange flagging was attached to the tag. The animal indentification number was painted on the side or back of the elk using a dye mixture. Forty-five elk were fitted with white telemetry collars. Trapping Crew DOW George Bear Steve Porter Rick Riser USFWS Gale Brewer Dave Johnson Mortality None John Wagner Steve Steinert Richard Byrd A1 White �116 Elk Trapped Independence Mt. Sentinel Mt. Buffalo Ranch Swift Ranch Ball er Ranch Owl Ridge - Speck Owl Ridge - State Total IS Recaptures: 9 E1 k co 11ared: 45 Cows Calves Spike Bull Total 1.3 16 1 32 5 5 2 2 2 1 10 2 4 8 1 7 3 8 1 1 7 2 1 3 3 36 16 33 6 3 78 �117 Number 1 2 35 Date 1-5 1-5 1-6 1-6 1-6 1-6 1-6 1-9 1-9 1-9 1-9 1-10 1-10 1-10 1-10 1-10 1-10 1-10 1-17 1-17 1-17 1-17 1-18 1-18 1-18 1-18 1-18 1-20 1-20 1-24 1-24 1-24 1-24 1-24 1-24 36 1-25 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 . 23 24 25 26 27 28 29 30 31 32 33 34 Elk Tra~~ed - North Park p 984) Age Sex location Ca M Buffalo F A Buffalo A F Sentinel Yr M Sentinel Yr Sentinel M Ca F Sentinel M Sentinel A Ca M Buffalo F Buffalo A Buffalo A F Sentinel A F A F Buffalo Buffalo A F Buffalo Ca M Sentinel Ca F Sentinel A F Sentinel F A Sentinel F A Independence F A Independence Ca F Independence Ca F Independence A F Independence Ca F Independence Ca M Independence A F Independence Ca F Independence A F Independence F Ca F Independence A M Independence A Ca F Independence Yr M Independence Ca M Independence Yr M Swift Ca F Swift Ca M Swift ._,"_ "'., ..•.. -~. Remarks Collared Coll ared Coll ared Collared Collared Coll ared Co11ared Collared Collared Co11ared one-tag Collared Collared Coll ared Call ared Call ared Call ared Call ared Collared Collared Call ared Call ared .... �118 Elk TraQQed - North Park n_9.~_4)- continued Number 37 38 39 40 41 42 43 44 45' 46 47 48 49 50 51 52 53 54 55 56 51 58 59 60 61 62 63 64 65 66-67 68 69 70 71 72 73 74 Date 1-25 1-25 1-25 1-25 1-26 1-26 1-26 1-27 1-27 1-27 1-27 1-27 1-27 1-27 1-31 1-31 1-31 2-2 2-2 2-2 2-3 2-3 2-3 2-7 2-7 2-7 2-10 2-10 2-10 2-10 2-10 2-10 2-10 2-16 2-16 2-16 2-16 Age Yr Ca Ca Ca Yr Yr Ca A Sex F ? M F F F F F A F Yr Ca Ca A A Ca M A F Ca M A M A F A F Ca Ca Yr F M A F Ca Ca F A F Ca ? A F A F A F Yr F A F Yr Ca M A F Ca M F M F F F F F F .Location Independence Independence Independence Independence Independence Independence Independence Independence Independence Independence Independence Independence Independence Independence Independence Independence Independence Owl Owl Owl Baller Owl Owl Owl Owl Owl Owl Owl Owl Owl Owl Swift Swift Ball er Owl Owl. Owl Remarks Co11ared Call ared Collared Collared Collared Collared Coll ared Coll ared Coll ared Collared Co11ared Collared Collared Collared Collared Coll ared Collared Coll ared Coll ared Collared Coll ared Co 11ared �119 '£1k Tra ~Qed - No rth Park {l984} - continued Number 75 76 77 78 79 'Date 2-16 2-16 2-16 2~16 2-16 Age Ca Sex A F A Ca Ca F F M M .Location Owl Owl Owl Owl Owl 'Remarks one-tag Col1 ared Collared ��121 Colorado Division of Wildlife Wildlife Research Report July 1984 JOB PROGRESS REPORT State of Colorado Project No. 45-01-502-15050 Big Game Investigations - Cervids Work Plan No. 3 Elk Investigations Job. No. 3 Evaluation of Factors Influencing Elk Nutritional Status and Population Performance Period covered: Author: 7/1/83-6/30/84 D. L. Baker ABSTRACT Due to reassignment to the supplement winter feeding evaluation study, no progress was made on 1983-84 Segment Objectives. ��123 EVALUATION OF FACTORS INFLUENCING ELK NUTRITIONAL STATUS AND POPULATION PERFORMANCE Dan L. Baker P. N.-OBJECTIVES 1. To develop and test a system for evaluating the potential of habitats to support elk. 2. To improve the predictive capability of this system by identifying and quantifying variables influencing the range-supply-animal demand model of nutritional carrying capacity. SEGMENT OBJECTIVES 1. Complete all nutritional analyses for deer-elk intake and digestion trials. 2. Analyze comparative voluntary intake, rate of passage and apparent digestibility data for mule deer and elk fed native forages. 3. Summarize and begin manuscript preparation for in vitro extent and rate of digestion studies of native forages. 4. Summarize tannin analysis for deer-elk forages. Prepa red by ~fj~:=--~~£~ . ...!...iXL--=-.::c=...::.~-=-Dan L. Baker _ �� https://cpw.cvlcollections.org/files/original/6c28f1151d0fa09318ed19feadb54ea7.pdf a70734926f37e6ee523f3d20fc93db37 PDF Text Text 125 Colorado Division of Wildlife Wildlife Research Report July 1984 Pt..1.. JOB PROGRESS REPORT State of Colorado /,§ Project No. 45-01-503-15050 ,~it'" Big Game Investigations - Nohcervids Work Plan No. 1 Multispecies trivestigations Job. No. 1 Animal and Pen Support Facilities for Big Game Research Period Covered: Author: 7/1/83-6/30/84 P. H. Neil ABSTRACT A total of fifteen pronghorn fawns and 8 mule deer fawns were and are presently being hand reared and trained during the fiscal year. A different formula was used in an effort to simplify temporary holding and rearing for field personnel. Twelve mule deer were involved in a winter range thermal cover study in Kremmling, Colorado, and returned to Fort Collins in good condition other than minor weight loss. Three mule deer were used in a study to evaluate the supplemental feed used in the State winter feeding program. No adverse effects·from the feed were revealed. Training of bighorn sheep, Rocky Mountain goats, elk, antelope and mule deer for upcoming studies is progressing smoothly and on schedule. .. I~ilj~llij'ijl~'~il~lrlilli~~m~~~ml BDOW025831 ��127 I1.NlMAL AND PEN sur=osr FACILITH:.S FOR BIG GAME RESEARCH Pau H. Neil P. N. OBJECT!VES To provide and maintain populations 0f captive big game animals and pen facilities to support big game research proqraas , SEGMENT OBJECTIVES 1. Continue to develop and maintain facil'ties at CSU Foothills Campus and Wildlife 2. 3. Research Center. Coordinate rearing. training. and research activities with captive wild and tame big qame animal s for' an big game cerv t d and noncervf d research projects. Integrate personnel, bi q game an ima 1 and phvs i cal p 13 lit support f'ac il Hi es, and fiscal resources into a single ~udget. r~ETHODS .AND 1~,Li.TER1!·'.L"S Routine neonate r"'aY'ing procedures were used to compIete the hand rearing and training of 3 orphaned mule deer fawns received during June, 1983. These animals were reared and trained for future nutrition studies and to supplement the big game research herd. Twelve adult mule deer were transpor-ted via horse t ra i le r to the Junction Butte Research Facility near Kremmling. Colorado, in November in support of the winter ranqe thermal cover s tudy , Dave freddy, Principa-I Investigator. A three-YEar-old female elk and a yearling male elk were transported to the Foothills Research Fact li ty f rom Washinqton State. Both are fistulated and have been tnccrpore ted into the research herd. These animals are presently undergoing training on the scale. isolation pens. and digestion cages. A supplemental feedirg trial u51ng three adult ~ule deer was conducted to deter ine the effects of the supplemental feed that was used in the western portions of the state in the Division of WildlHe's supplemental feedi n9 program that ~vas 'j ni t i eted due to s,:,ven::, \_or; nter condi t t ons , Fifteen pronghorn arte10pe fawns and five mule deer fawns are being hand reared and trained at the present time for future game damage and nutritional stud"es. Nine of the pronghorn faw~s were born at the Foothills Facility and 6 fawns were orphans brought in by field personnel. All five mule deer fawns were orphans brougnt in by field personnel from various parts of the state. �128 Routine rearing and training procedures are being used with the exception of the milk formula. All animals are being reared on undiluted canned evaporated milk rather than the diluted f'ormul a previ ous ly used. This method is being used in an effort to '(educe diarrhea prob lems and simplify temporary ho·iding situations for field personnel. Intens ive training in metabolic cages of three bighorn sheep and five Rocky f't1ountain goats and the scale t s present lv being conducted in prepara- tion for upcoming intake and digestion trials. be-ing catne red and prepared Materials and equipment are also RESUL~S AND DJSCUSS!l~ The three mul E deer fawns that welAe ra i sed in the fi rst part of the year us i ng the routi ne procedure with the except i on of the formul a. The deer W,'2Y'e fed undiluted canned evaporated mi l k and no diarrhea problems were encountered tr.r-ouqh the ent i re rear ing process. were reared The twcl ve adu l t mule Q(~er' transpor-ted to Fort Collins with no ill effects to other KreHliil·; i n9 '>'Jete transpor-ted than mlncr weight The two el k brought in from \~aSrd{l~it(jrl ::iT:i'te al'ie, respono inq well t ra irrinq, which wi 11 .ont inue for th(~ remainder of tne SUI1lIneL The three mul e deer used in the supp lemental feeding tt'·ial back loss. were to gradually ct ~::Opercent we iqht 1035 was attained. They were then taken completely off feed for three days. Each deer was then fed 2~OOO gms of the same waffer that was being used in the supp Iementa 1 feed inq program to de termi ne H any t l I effects wcul d be observed. Intakes W2l"e low tne fi rs t few days. aVEraging 592 gms per day for all three deer. After eighteen days, intakes were averaging 1,178 gms per daj for all three deer. Weight increases for the same period averaged 8.6 percent. No adverse e-;f.ects were observed. fed succes s ively lo,.; qual i ty d-iets until To this point in time. minimum dia.rrhea problems have been encountered in the -15 pronghorn fawns and r; mule deer fawns that are bei ng hand reared on the undi luted canned evaporated milk formula. The few prob lems that did appear clea red up with-in 1··2 days. Training is progressing smoothly and on schedule .. If the undi Iuted fornul a works well this year, I plan to writ an amendment to Colorado D'ivi s ion of ~:i-ldnfe Game Fact Sheet #106 for fi e Id pe rsonne l to simplify the temporary hold-ing and rear-inc of orphan~d ungula~es until the animals can be sent to a suitable holding faciiitv. research herd presently consists of 27 mule deer, 7 elk, 10 Rocky Mountain goats, 9 bighorn sheep, 23 pronghorn antelope. and one domestic CON. The total Preparec by �129 Colorado Division of Wildlife Wildlife Research Report July 1984 JOB PROGRESS REPORT State of Colorado Project No. 45-01-503-15050 Work Plan No. 1 ----------------- Job. No. 3 Period covered: Author: Personhel: Big Game Investiaations Multispecies Noncervid - Noncervids Investigations Research Administration 7/1/83-6/30/84 R. B. Gill L. E. Lovett, L. H. Carpenter ABSTRACT Despite a work year fraught with disruptions, dislocations, and distractions, most of the work which was programmed for Noncervid Research was accomplished. Three studies ;... Wo\~k Plan 1, Job 3, Work Plan 2, Jobs 3 and 4 - were not completed because the principal investigators were reassigned to evaluate the 1983-84 winter feeding pr·ogram. Chronic difficulties continue with the administration of the Noncervid Research program. It has been exceedingly difficult to develop and maintain an effective check-book accounting system because many of the fiscal impacts to program budgets occur at the state office level and a re not communi cated di rectly, c1 early and timely to cost-center supervisors. The most nettlesome problems involve within-fi~cal-year changes in personal services rates, accounting for warehouse withdrawals, charges for vehicle costs, contracts, year-end ro ll forward procedures, and dynamic purchasing rules and regulations. ��131 NONCERVID RESEARCH ADMINISTRATION R. Bruce Gill P. N. OBJECTIVE Supervise and administer research activities within the scope of Big Game Noncervids research project. SEGMENT OBJECTIVE Supervise and administer all activities conducted within Project 45~Ol~503~15050 for FY 83-84. RESULTS FY 83-84 was an extremely difficult year to maintain momentum and productivity of a diverse research program w-ithin CDOW. Director Grieb retired in April. but really was on terminal leave from January. Director Ruch did not take the he1m until 1ate Apri 1. The trans iti on period was not smooth. Rumors ran rampant wi tbin CDOW of pending changes in organization structure and job duties. - This, coming on the heels of a disruptive research audit~ made it very difficult to maintain morale and productivity. Fortunately, the Noncervid Research staff is largely self-motivated and all are high quality employees so most of the work was accomplished according to standards established in their respective FAPAS·s. Budget and accounting problems continued to be chronic sources of exasperation. A final allocation of fiscal resources was not accomplished until FY 84~85 and already commenced. This caused considerable confusion concerning the direction and scope of individual research projects. In the future, these problems could be resolved by linking the resource allocation and program planning processes so that activities are identified in the program plans according to priorities. Administrative Directive A.".26set in motion a process of designating certain individuals as cost center supervisors. These cost center supervisors were given responsibility for administering fiscal resources assigned to projects. Cost center supervisors were instructed to establ-ish check-book accounting systems so they could track expenditures charged to their cost centers. The purpose of the check-book accounting system was to prevent overexpenditures of allocated fiscal resources. From the onset, this system has been plagued with difficulties. Foremost is that there are no direct links between cost center check-book accounting systems and the centralized statewi~e accounting system in Denver. In addition, the statewise system does not provide sufficient resolution to be most useful to the cost center supervisor. �132 This year problems were encountered when salaries increased within midfiscal year because of an occupational study which upgraded most research positions and because of delayed implementation of the July salary survey cost-of-living adjustments, There was poor communication between accounting and cost center supervisors concerning the precise amount to which these changes would affect salary increases. Again~ this problem could be resolved if there were better conmun ica t ion links between cost center supervisors and the central accounting staff. Additional check~book accounting problems were attributed to delays in assigning costs to warehouse withdrawals and a mystical system for assigning vehicle costs. A potential solution to all of these problems would be to redesign the accounting system literally from the bottom up. Design an interactive computerized system that provides the degree of resolution required for responsible cost center supe rvis i on , but which could reduce by summation the degree of resolution provi ded to higher order supervisors. In my opinion, a complete restructuring of CDOWls entire accounting system is a high pr i ori ty concern and should be accomplished with input from all users of the system. A completely computerized system seems to be the only vi able option for the divers t ty of needs extant in CDOW. Prepared by --=R-. -::BO"'"t'uce Gi11 Wildlife Research Leader �Colorado Division of Wildlife Wildlife Research Report July 1984 133 JOB PROGRESS REPORT State of Colorado Project No. 45-01-503-15050 Big Game Investigations - Noncervids Work Plan No. Multispecies Investigations Job. No. 4 Period covered: Author: Big Game Forage Selection Dynamics 7/1/83-6/30/84 R. B. Gi11 Personnel: S. C. Torbit, W. A.,Alldredge, P. H. Neil ABSTRACT Very little progress was made toward the objectives of this job because the principal investigator was reassigned to evaluate CDOW's winter feeding program. The emphasis of this job was shifted to address 2 priority strategy objectives in CDOW's Program Plans: 1) 6-1-1: Develop specific quantified habitat management guidelines for pronghorn; 2) 6-1-2: Investigate the impacts of antelope grazing on winter wheat production and bindweed spread. ��135 BIG GAME FORAGE SELECTION DYNAMICS R. B. Gill P. N. OBJECTIVE Prepare a Program Narrative describing experiments to test big game forage selection theory. SEGMENT OBJECTIVE Same as P. N. Objective. RESULTS The objective of this initial job - to prepare a Program Narrative - was not accomplished during the Segment because I was reassigned for 6 months (January-June) to participate in an evaluation of CDOW's big game winter feeding effort. During the Segment this study was redirected towards a more practical end. CDOW entered into a cooperative agreement with Colorado State University's Depar-tment of Fishery and Wildlife Biology and the Agricultural Experiment Station to investigate the impacts of pronghorn grazing on winter wheat yields. A study plan was prepared and approved by CSU, the CDOW's NE and SE Regional staff, the Wildlife Services Section and the Noncervid Research Section. The first winter season of study revealed that pronghorn switch habitat use rather abruptly in the early winter from grassland habitat to winter wheat fields and abruptly switch again in spring from winter wheat back to grass lands. I hypothesize that these switches are occasioned by changes in nutritive quality of grassland forages relative to winter wheat. The cell contents of forages are good indexes to changes in nutrititive quality of forages because cell contents are nearly 100% digestible and contain the majority of that forage's digestible nitrogen. So, as cell contents of a forage increase so does that forage's digestibility and soluble nitrogen (Van Soest 1982). Cell contents of pronghorn diets from native shortgrass prairie increase in spr+no and peak early in sumner , Then they decline as plants mature and enter winter senescence (Schwartz 1977). Cell contents of winter wheat, in contrast, peak in later winter and decline to summer harvest (Fig. 1). There are two threshold periods. The first occurs in December when cell contents of winter wheat plants surpass cell contents of grassland forages. The second threshold occurs in May when cell contents of winter wheat decline below cell contents of grassland forages. These 2 thresholds probably chronicle the timing of changes in pronghorn habitat preferences. �136 Research is now being planned to test thi hypothe i and to evelop methodology to remotely sense changes in cell contents of both winter wheat and grassland forages (Colwell 1974; Ashley and Rea 1975~ Tucker 1979, 1980). LITERATURE CITED Ashley~ M. D., and J. Rea. 1975. Seasonal vegetation differences ERTS imagery. Photogram. Engineer. Remote Sensa 41:713·-719. Colwell, J. E. 1974. Vegetation Environ. 3:175-183. Schwartz, C. C. 1977. prairie, Colorado. CO.113pp. canopy reflectance. from Remote Sense Pronghorn grazing strategies on the shortgrass Ph.D. Thesis. Colorado State Univ. Ft. Collins, Tucker. C. J. 1979. Red and photographic infrared linear combinations for monitoring vegetation. Remote Sense Environ. 8:127-150. 1980. Remote sensing of leaf water content Remote Sense Environ. 10:23-32. Van Soest, P. J. Books, Inc. 1982. Nutr'itional ecology of the ruminant .. Corvallis, OR.' 374pp. ".':'J Prepared by \~ in the near infrared. ."~Q \\ 1-J~), R. Bruce Gi 11 Wildlife Research Leader o 8! B �137 100 90 80 ./ ,,-.- ...., 70 '\ I 60 Cell contents (%) Sl;o .----!:!s;_,.. I \ 50 \~_,. 40 \~ (l) G: 30 I -:s (l) 20 / ~ / '\ 10 / \ J F M A Ma Ju Jl Month I Au S 0 N D Fig. 1. Seasonal fluctuations in cell contents of winter wheat and pronghorn diets from shortgrass prairie forages. ��139 Colorado Division of Wildlife Wildlife Research Report July 1984 JOB FINAL REPORT State of Colorado Project No. 45-01-503-15050 Big Game Investigations - Noncervids Work Plan No. Bighorn Sheep Investigations Job 2 ----------------- No. 2 Period covered: Author: Comparative Bioenergetics of Rocky Mountain Goats and Bighorn Sheep 7/1/83-6/30/84 T. V. Dailey Personnel: N. T. Hobbs ABSTRACT We measured energy expenditure of mountain goats and mountain sheep in 180 thermoregulation trials, 93\now .locomotion trials, 134 hard-surface locomotion trials, and 100 treadmill locomotion trials. Because data analyses are incomplete, results are not presented here. Results will be submitted for publication in scientific journals as follows: Comparative Thermorequl at i on in Mountain Goats and I~ountain Sheep, Energy Expenditures by Mountain Goats and Mountain Sheep for Locomotion, Comparative Oxygen Consumption and Thermoregulation by Exercising Mountain Goats and Mountain Sheep. ��141 COMPARATIVE BIOENERGETICS OF ROCKY MOUNTAIN GOATS AND BIGHORN SHEEP Thomas V. Da t ley P. N. 'OBJECTIVES 1. Formulate and test hypotheses that elucidate intra- and interspecific energetic relationships of mountain goats and mountain sheep and their environments. 2. Describe seasonal cycles in voluntary food intake, body growth and fasting heat production for mountain goats and mountain sheep. 3. Describe the energy costs incurred by mountain goats and mountain sheep for bedding, standing, locomotion on level and steep slopes and in deep snow, and for thermoregulation when they are exposed to wind, low ambient temperatures and natural radiation conditions. SEGMENT OBJECTIVES 1. Rear and train 5 bighorn sheep lambs and 5 mountain goat kids for use in energeti cs experiments. 2. Estimate energy costs for thermoregulation and locomotion of bighorn sheep and mountain goats during winter. ACKNOWLEDGMENTS For their essential assistance I thank M. Miller, M. Bowser, S. Alvord, D. D'Amico, M. Nelson? G. Sullivan, M. Gabardi, P. Creeden, E. Winicov, D. Johnson, B. Wunder, J. Stager, and T. Hobbs. Appreciation is extended for help by L. Adams, D. Baker, R. Gill, D. McGlasson, R. Reiner, A. Orabona, D. Reed, R. Sayre, S. Pagano, J. Gross, J. Ha, L. Lovett, and D. Wynne. For animal and facility support I thank T. Spraker, Cheyenne Mountain Zoo, University of Denver, C. Miller and CSU Physiology and Biophysics Department, Gene and Barb Day, Arapaho Wildlife Refuge. U.S. Fish and Wildlife Service, City of Idaho Springs, Clear Creek Ranger District, U.S. Forest Service, and the U.S. Bureau of Reclamation. METHODS AND MATERIALS Methods and materials are described in the Program Narrative for this study. �142 RESULTS AND DISCUSSION Results and Discussion will Prepared by -.f} t rn be submitted in FY 84-85 for publication. J--'7/: ~ I~~~ -~7 Thomas V. Dailey �143 Colorado Division of Wildlife Wildlife Reseach Report July, 1984 JOB PROGRESS State of Colorado Project No. 45-01-503-15050 2 WOl"k Pl an No. -------_ Job. No. 3 Period covered: Author: Personnel: REPORT Big Game Investigations .. . - Noncervids Bighorn Sheep Investiaations Comparative Digestive Physiology of Bighorn Sheep and Mountain Goats 7/1/83-6/30/84 N. T. Hobbs R. B. Gill. A.BSTRl!.CT A study plan was prepared and reviewed. Further progress by reassignment to studies on supplemental feeding. was precluded ��145 COMPARATIVE DIGESTIVE PHYSIOLOGY OF BIGHORN SHEEP AND MOUNTAIN GOATS N. Thompson Hobbs P. N. OBJECTIVES 1. Exami ne diges tion of grass and browse di ets in b i ghorn sheep and mountain goats. 2. Incorporate data in model of interspecific competition. SEGMENT OBJECTIVES 1. Prepare a detailed study plan for experiments on comparative digestive physiology of bighorn sheep and mountain goats. 2. Conduct digestion trials to compare digestive efficiency and rate of passage of grass and browse diets in bighorn sheep and mountain goats. -METHODS AND MATERIALS , Literature was reviewed and a study plan prepared. RESULTS AND DISCUSSION After completing the study plan for diqes tf on experiments (Appendix 1), personnel were reassigned to studies of supplemental feeding. Prepared by f/J ~ N~ Thompson Hobbs /.JJL_ < ��147 APPENDIX 1 11/83 PROGRAM NARRATIVE State: Colorado Project No. 45-01-503-15050 Big Game Investigations - Noncervids Work Plan No. 2 Bighorn Sheep I_nv~st~tions Job No. 3 Comparative Digestive Physiology of Bighorn Sheep and Mountain Goats A. _ NEED The Strategic Plan for Managing Wildlife in Colorado identifies 2 high priority tasks for management of bighorn sheep (avis canadensis) populations and the habitats they occupy. The undertaking of greatest importance is to increase the carrying capacity of bighorn ranges through various methods of habitat alteration. Another important strategy calls for defining the habitat requirements of bighorn populations. Implementing both of these strategies requires a thorough understanding of the value of various food plantsto bighorn sheep. In addition, the primary strategic task for managing mountain goats is predicting the outcome of competition between mountain goats and bighorn sheep. Many workers (Bell 1970. Janis 1976, Hoffman 1973, Jarman and Sinclair 1979, Kay et al. 1980, Hanley 1982) have demonstrated that a fundamental mechanism of ecological separation among sympatric ungulates is interspecific variation in efficiency of digestion of diets differing in physical and chemical composition. That is, ungulates possessing different digestive capabilities are compelled by those differences to use different food resources, and, as a resul t , those species rarely compete fo;~food. With a single exception (Hebert 1973), no studies have examined the ability of �148 bighorn sheep and mountain goats to digest the foods they consume. We propose to investigate that ability. B. OBJECTIVES 1. To compare digestive efficiency of mountain goats and bighorn sheep consuminq grass, forb, and browse diets. 2. To compare rate of passage of grass, forb, and browse diets consumed by mountain goats and bighorn sheep. C. EXPECTED RESULTS AND BENEFITS Knowledge of digestive physiology of mountain goats and bighorn sheep will assist resource managers in making decisions on animal habitat requirements and efficacy of habitat treatments aimed at meeting those requirements. Such information will illuminate the potential for competition between mountain goats and bighorn sheep when they share limited food resources. D. APPROACH 1. Hypotheses Ha: Bighorn sheep will digest test diets more completely than mountain goats, but digested energy intake per kilogram of body weight will be greater for bighorn sheep. H: o Digestive efficiency and energy intake will be similar for bighorn sheep and mountain goats. Ha: Rate of passage of all diets will be faster in mountain goats than bighorn sheep. H·o· Rate of passage will not differ on any diet for mountain goats and bighorn sheep. �149 .2. Rationale Dailey et ale (1984) compared the botanical composition and nutritional quality of diets of mountain goats and bighorn sheep grazing on alpine ranges during winter and summer. Diets of mountain goats contained more dicots and were more lignified and less digestible than diets eaten by bighorn sheep. Many workers have found parallels between animal diet choices and their digestive capabilities. That is, species choosing dicot domi nated diets appear to be best adapted to rapidly ferment cell solubles and rapidly excrete fiber; species choosing graminoids are better able to more completely digest the unlignified fiber fraction. These observations coupled with the report of Dailey et al. (1984) suggest that bighorn sheep should digest their food more completely than mountain goats, but mountain goats should digest and excrete ingesta more rapidly. 3. Methods a. Test Diets and In Vivo Digestibility Four diets will be fed to experimental bighorn sheep and mountain goats: 1) Diet 1 will consist of chopped, third cutting, alfalfa stems. Chemical composition of this diet will be similar to forbs consumed by bighorn sheep and mountain goats during summer and fall. 2) Diet 2 will consist of native grass harvested mature during early fal·l. At this time. chemical and physical �150 characteristics of Diet 2 will be similar to graminoids eaten by mountain goats and bighorn sheep during winter. 3) Diet 3 will consist of native grass harvested and cured during mid-summer. Composition of Diet 3 win be similar to graminoids eaten by mountain goats and bighorn sheep during summer. 4) Diet 4 will be composed of 75% blueberry (Vaccinium spp.) stems and 25% native grass hay (the same as Diet 3). Composition of this diet will be similar to browse dominated diets consumed during winter. Three adult bighorn sheep and three adult mountain goats will be conditioned to digestion cages and fed experimental diets during January, February, March and April of 1984. A complete balance trial and rate of passage estimate will be made for each animal during each month. Each diet will be fed in four periods (Table 1) as follows: !3ui~: The purpose of the buildup is to adjust animals and their rumen microflora to new test diets and to void the diet previously fed. Buildup periods, in conjunction wi th pretri als , prevent carryover effects from previous diets. Because high quality diets will be alternated with low qual ity ones (Table 1), the buildup will also allow recovery of animal condition followinq feeding of low qual i ty rations. Pretrial: The purpose of the pretrial is to measure �151 intake for use in the balance trial. During the first 5 days of the pretrial period, the test ration will be fed ad libitum until daily intakes can be predicted. Daily rations of the test diet will then be adjusted until no orts remain after each 12 hr. feeding. Table 1. Feeding schedule for bighorn sheep-mountain goat digestion trials. Day Locationa Diet Trial Phase Intake High quality IP Buil dup 1-14 Gradual increaseb grass IP Pretrial Ad libitum 15-21 22-24 DC Pretrial Ad libitum 25-31 DC Trial 32-45 IP Buil dup 90% of Ad libitum · b Gradual lncrease 46-52 IP Pretrial Ad 1ibitum 53-55 DC Pretrial Ad 1 ibitum 56-62 DC Trial 63-76 IP Bui 1dup 90% of Ad libitum · b Gradual 1ncrease 77-83 IP Pretrial Ad libitum 84-86 DC Pretrial Ad 1ibitum 87-93 DC Trial 94-107 IP Buil dup 90% of Ad libitum · b Gradual lncrease 25% Grass 108-114 IP Pretrial Ad libitum 1 115-117 DC Pretrial Ad libitum 118-124 DC Trial 90% of Ad libitum Low quality grass Al fa1fa stems 75% Vaccinium alP = Isolation pen DC = Digestion cage bNew ration will be added to previous diet at rate of approximately 10% per day until new ration is being fed ad libitum. �152 Trial: Buildup and pretrial will last 21 days. Following this period, feces and urine will be collected daily for 7 days. Subsamples will be taken from each day's collection and frozen. Urine will be collected in polyethylene jugs and kept at a pH of approximately 3 by addition of 10% solution of sulphuric acid. Feed, orts and feces will be freeze-dried, then processed through a Wiley mill with a l-mm screen and analyzed for dry matter(DM) , organic matter (OM), cell wall constituents (CWC), acid detergent fiber (ADF), lignin (LIG), nitrogen (N), and gross energy (GE). Retention time and ruminal turnover rates will be estimated for each animal and each test diet using 3 stable rare earth elements. to mark all test diets. Vherbium (Vb) will be used This will allow comparisons among all diets using the same marker. Cerium (Ce) will be used to independently mark the grass portion of the grass-browse mixture. Vaccinium spp. will be marked with Samarium (Sm). These additional elements will be used to assess possible associate effects of animals consuming diets containing different proportions of browse and grass. Methods for marking meals and dosing animals will follow (Allen 1982). This method requires the soaking of a por- tion of a meal in a rare earth marker solution for 24 hrs, rinsing in an acid rinse, and air drying prior to feeding. The amount of marker to be administered is calculated according to the following formula: dose (~g/lOO kg weight) = MAL x F/100 kg body weight x 1/K2-dayx DX (1/.5) �It53 where MAL = minimal analytical level for the marker, (llg/g DM), F = dry matter fi11 for segment of interest (g/lOO kg body weight), k2 = approximate turnover expected for the marker expressed in days, D = days post dose of last collection. This calculation assumes (1) a rumen dry matter fill of 2% of body weight, (2) a daily turnover of 1/K2-day, and (3) a serial diluting effect of 0.5. Fecal collections will be made just prior to dosing and every 2 hrs following dosing for the first 24 hours. Subsequent collections will be made every 6 hours for the following 6 days. The entire fecal sample will be col- lected, weighed and stored for later analysis. Concen- tration of marker in feed and feces will be measured using neutron activation analysis (Young et al. 1975). Calculation and measurements of ingesta turnover rates and total mean retention times in the gastro-intestina1 tract (GIT) will be calculated according to method of Grovum and Williams (1973). Fecal excretion of markers will be used to fit a 2 compartmental model described by the foliowing equation (Brandt and Thacker 1958): y = Ae-Kl(t-TT) _Ae-K2(t-TT) where y and A = constant rations of marker in feces dry matter = rate constant describing marker kinetics in the reticulo-rumen, �154 K2 = rate constant pertaining primarily to digesta kinetics in the caecum and proximal colon, t = ~he period after ingestion of marker, TT = transit time of marker through the omasum, abomasum, and the small and large intestines. Previous studies with sheep suggest that these digestion parameters are biologically significant (Grovum and Williams 1973). The advantage of this model is that ruminal turnover rates of ingesta can be calculated based on fecal excretion of markers. 4. Analysis Effect of diet and animal species on forage digestion and rate of passage will be analyzed with a factorial analysis of variance for a randomized complete block design with 4 repeated measures on 3 replicates (Table 2). diet effects. Effects of month will be confounded with Replicates will be considered to be random effects; diet and species, to be fixed effects �155 Table 2. Analysis of variance table for comparisons of mountain goat and bighorn sheep digestive function. OF Source Total 23 Species 1 Animal/species ] F test 4 Animals/goats 2l Animals/sheep 2- test for equal variances Diets Species/diets F test Animals x diets/species Animals x diets/ goats 6 6~ test for equal variances Animals x diets/sheep 1/ SCHEDULE I October - November 1983/ Prepare study plan, train animals for digestion studies January - March 1984 Conduct digestion trials ~. ; April - June 1984 Analyze digestion data August - October 1984 Prepare manuscript and final report E. LOCATION The study will be conducted at the Colorado Division of Wildlife Foothills Research Facility, Colorado State University Foothills Campus, Fort Collins, Colorado. �156 F. RELATED FEDERAL PROJECTS Non-cervid 45-01-503-15050 WP2, J2 Comparative Energetics of Bighorn Sheep and Mountain Goats During Winter WP4, Jl Seasonal Habitat Selection and Activity of Sympatric Mountain Goat and Bighorn Sheep Populations LITERATURE CITED Allen, M. S. 1982. Investigation into the use of rare earth markers as gastrointestinal markers. M. S. Thesis 9 Cornell University. Ithaca, NY. 107pp. Bell, R. H. U. 1970. The use of herb layer by grazing ungulates in the Pages 111--123 J..!!. A. Watson, ed. Animal populations in Serengeti. relation to their food resources. Symp. Br. Ecol. Soc. 10. Blackwell Sci. Publ., Oxford, UK. Brandt~ C. S., and E. J. Thacker. 1958. A concept of rate of food passage through the gastro-intestinal tract. Dailey, T. V., N. T. Hobbs, and T. N. Woodard. J. Anim. Sci. 17:218. 1984. Experimental comparisons of diet selection by mountain goats and mountain sheep in Colorado. J. Vlildl. Manage. Grovum, W. L., and V. J. Williams. sheep. In press. 197~. Rate of passage of digesta in 4. Passage of marker through the alimentary tract and the biological relevance of rate constants derived from the changes in concentration of marker in feces. Hanley, T. A. Br. J. Nutr. 30:313-329. 1982. The nutritional basis of food selection by ungulates. J. Range Manage. 35:146-151. �157 Hebert$ O. M. 1973. Attitudinal migration as a factor in the nutrition of bighorn sheep. Vancouver. Ph.D. Diss., University of British Columbia. 357pp. Hoffman, R. R. 1973. The ruminant stomach. E. Afr. Monogr. in Biol. 2. E. Afr. Lit. Bur. Nairobi, Kenya. 350pp. 9 Janis, C. 1976. The evolutionary strategy of the Equidae and the origins of rumen and cecal digestion. Evolution 30:757-774. Jarman, P. J., and A. R. E. Sinclair. 1979. Feeding strategy and the pattern of resource-partitioning in ungulates. Pages 130-163 in A.R.E. Sinclair and M. Norton-Griffiths, eds. Serengeti: dynamics of an ecosystem. Kay, R. N. B.~ W. V. Englehardt, and R. G. White. physiology of wi1d ruminants. 1980. The digestive Pages 743-761 in Y. Ruckenbush and P. Thivend, eds. Digestive physiology and metabolism in ruminants. AVI, Westport, Conn. Young, M. C., F. E. Haskin, M. E. Wacks, B. Theurer, and P. R. Ogden. 1975. Neutron activation analysis of dysprosium (a potential inert marker) in hay and feces. J. Anim. Sci. 41 :178-184. ��159 Colorado Division of Wildlife Wildlife Research Report July 1984 JOB PROGRESS REPORT State of Colorado ------------------ Project No. 45-01-503-15050 Work Plan Period covered: Author: - Noncervids 2 Bighorn Sheep Investigations 4 Plan of Nutrition and Bighorn Sheep Population Performance ------------- Job No. Big Game Investigations 7/1/83-6/30/84 N. T. Hobbs ABSTRACT Initial development of study plans and experimental protocols were completed. I hypothesized that disease outbreaks in bighorn populations result from endocrine-mediated effects of stress on immune functions. Two important stres ses , popu lat ion density and plane of nutrition, were identified for experimental t rea trnents , Methods for challenging sheep with lungworm, and for reversing those challenges, were developed. Reassignment to studies on supplemental feeding precluded further progress. ��161 PLANE OF NUTRITION AND BIGHORN SHEEP POPULATION PERFOR~1ANCE N. Thompson Hobbs P. N. OBJECTIVES 1. Investigate relationships between stress and disease resistance in bighorn sheep, 2. Determine causes of tmmune failure in high density populations. 3. Determine the most effective way to quantify immune responses of bighorn sheep. 4. Determine effect of nutr i t ion and stress on immune competence. SEGMENT OBJECTIVES 1.. Prepare a detailed study plan to investigate relationships between nutritional plane and parasite resistance in bighorn sheep. 2. Conduct pilot experiments to determine the most effective method to challenge bighorn sheep with Prot~str9J.')gy"lus sp. larvae. 3. Rear and train bighorn lambs for controlled nutrition-parasite experiments during 1984-1986, METHODS AND MATERIALS _Preparation of Study Plan for Controlled Experiments Literature was collected and reviewed. Infection of Snails with Protostrongylus sp. Larvae Three hundred dry-land snails were collected from the Spring Creek area in Fort Collins, CO. Snails were placed on petri dishes (20 per dish) ~-~containing sand and bits of rotting wood. Water (approx. 3 ml) was added dail~~ 10 gm oatmeal and 1 gm Ca C03 was added biweekly. Dishes were maintained at approx. 20 C. Bighorn sheep feces with known counts of Protostrongylus sp. larvae were Baermanized (Beane and Hobbs 1983) through f ilter paper. Fifty snails were placed on filter paper containing 1500 larvae and were left there for 8 hrs. Another group was given the same treatment for 12 hrs. A third group was exposed to 4000 larvae for 12 hrs. Larve1 infections were detected by viewing snails at 50X magnification. �162 Treatment of Protostrongylus Infection wHh Ivermectin Three adult male and 1 adult female b iqhorn sheep were treated with Ivermectin admin-istered subcutaneously at a dose of 200 mg kg-l. Counts of Protostrongyl us 1arvae in the feces VJeY'econducted (Beane and Hobbs 1983) every 3 days for 16 days post-dose. RESULTSAND DISCUSSION Preparation of St_ll"~l.PlaY!__!or Contro lled Experiments_ Based on revi ew of 1i terature (Append; x "I), two hypotheses to explain catastrophic die-offs of bighorn sheep resulting disease: were developed from epidemic Hypothes is I: Disease outbreaks resul t from effects of chroni c stress due to tnappropr+ate ly high popu lation density. The primary mechanism for this relationship was hypothesized to be compromise of the immune system as a consequence of endocr-i ne responses to Ionq-term stress of crowding in preferred habitats. Hypothes -j s II: Disease outbreak results from ma1nutriti on due to poor range conditions and hi qh popu lati on density. The primary mechanism for this relationship was hypothesized to be compromise of the immune system resulting from inadequate protein and energy nutrition leading to supression of antibody synthesis. Long-term management of bighorn populations requires defining a fundamenta -I objecti ve for that management. C1 early, the object; ve of over r i di nq tmpor tance in managing for thrifty bighorn populations is preventing excessive mortality f rom disease. Hm" that objective can be met, however. is not c lear , If Hypothesis I proves to be true, then bighorn management should emphasize reducing the ecological density bighorns experience by (1) harvest and transplant of excess animals and (2) increasing areas of preferred habitat. If Hypothesis II is correct, then management efforts should be directed at enhancing food supplies within bighorn habitats. An experiment is being designed to address these hypotheses and to allow development of indices of stress and immune f'ai lure in bighorn populations. Treatments include 2 Ieve ls of population density and 2 planes of nutrition in a factorial arrangement. Several responses to these treatments will be observed: 1. Immune competence 2. Blood cortisol 3. Fecal i ndt measured by b 1as togenes is techni ques. levels. ces of blood cor-t i so l levels. Dr. Terry Spraker agreed to cooperate in blood and immune system assays. Dr. Elizabeth Williams provided details on blastogenesis techniques she is currently using to examine effects of chronic stress on bighorn immune competence. �163 Treatment of Protostrongylus Infection with Ivermectin Experiments revealed Ivermectin administered subcutaneously at 200 mg. kg -1 body weight can substantially reduce lungworm (Protostrongylus spp.) burdens in bighorn sheep, even when the initial infection of treated animals is low (Table 1). Table 1. Lungworm 1arvae counts .(no. individuals/gm drymatter in feces of Yeate2__9nd cont ro1 bi.9l!_orn _shee~ __ ,., .. Day Post Dose Control Treatment 1 5 8 10 12 14 16 180 160 180 180 180 170 170 140 260 90 65 50 30 >10 We observed no detrimental side effects of treatment. The ability of IVermectin to reduce lungworm to near 0.0 despite relatively low burdens initially will be extremely useful in developing means to challenge uninfected animals with known lungworm burdens. Infection of Snails with Pl"'otostn:ingylussp. Larvae Fifty to 70% of snails infected with Protostrongylus survived after 5 weeks following treatment. Assuming 3 larvae per snail, \~e anticipate feeding 100 snails to infect bighorns with high levels of Protostrongylus. Further progress was precluded by reassignment to supplemental feeding studies. LITERATURE CITED Beane, R. D., and N. T. Hobbs. 1983. The Baermann technique for estimating Protostrongylus infection in bighorn sheep: effect of laboratory procedures. J. Wildl. Diseases 19:7-9. Prepa red by -o-:--(})-=k,,----=--t~~)~-'-----''-----N. Thompson Hobbs Wildlife Researcher �164 APPENDIX I LITERATURE REVIEWED FOR STUDY PLAN PREPARATION Alban, S. D., B. Mitchell, and B. W. Staines . 1983. Fertility and body weight in female red deer: a density--dependent relationship. J. Anim. Ecol. 52:969-980. A 11 ison> A. C. -1982. Coeval ut ion between has ts and infecti ous di sease agents, and its effects on virulence. Pages 245-267 in population biology of infectious diseases. R. M. Anderson and R-. M. May, eds. Springer-Verlag. 314pp. Anderson, R. M., H. Jackson, R. M. May. and T. Smith. 1981. The population dynamics of fox rabies in Europe. Nature 289:765-771. , and R. M. ~lay. 1978. ---popul ati on interactions. Regulation and stability of host-parasite J. Anim. Ecol. 47: 219-247 and 249-267. _---:~ and 1979. Popu1 a tion bi 01 ogy of i nf'ect i ous di seases. Natur 280:361-367 and 455-461. -- , and 1981. The population dynam-ics of microparasites and their invertebrate hosts. Phil. Trans. Roy. Soc. B 291:451-524 . --8'5: • and 41'--426. 1982. Coevolution of hosts and parasites. Parasitology , and 1982. Directly transmitted infectious diseases: --contro TbYvacci nati on. Sci ence 215: "1053-1060. Arave, C. W. 1977. Effects of dominance rank changes, age and ody weight on plasma corticoids of mature dairy cattle. J. Dairy Sci. 60:244-248. Batzli. G. O. 1983. Responses of artic rodent populations to nutritional factors. Gikos. 40:396-406. Burchfield, S. R., S. C. Wouds, and M. S. Elich. 1980. Pituitary adrenocortical response to chronic intermittent stress. Physiol. Behav. 24:297-302. Caughley, G., and C. J. Krebs. 1983. Are b-ig mammals simply l i tt le mammals writ large? Oecologia (Berl.) 59:7-17. Chandra, R. K. 1975. Antibody formation in first and second generation offspring of nutritionally deprived rats. Science 190:289-290. Christian, J. J. 1980. Endocrine factors in population regulation. Pages 55-117 in M. N. Cohen, R. S. Malpass, and H. G. Klein, eds. Biosocial mechanisms of population requ lat i on. Yale Univ. Press, New Haven, Conn. 406pp. Clutton-Brock, T. H .• F. E. Guinness. and S. D. Albon. 1983. The cost of reproduction to red deer hinds. J. Anim. Ecol. 52:367-383. �165 Dantzer, R., and P. Mormede. 1981. Can physiological criteria be used to assess welfare in pigs? Curro Topics Vet. Med. Anim. Sci. 11:53-73. --J. , and 1983. Stress in farm animals: Anim.-Sci. 57:6-18. DeForge, J. R. 1976. Stress: Bighorn Counc. 20:30-31. a need for reevaluation. is it limiting bighorn? Trans. Des. Ewbank, R. 1973. Use and abuse of the term "stress" in husbandry and welfare. Vet. Rec. 92:709-710. Feuerstein, V .• R. L. Schmidt, C. P. Hibler, and W. H. Rutherford. 1980. Bighorn sheep mortality in the Taylor River-Almont Triangle Area, 1978-1979: a case study. Colo. Div. Wildl. Spec. Rep. No. 48. Forrester, D. J. 1971. Bighorn sheep lungworm-pneumonia complex. Page 158-73 in parasitic diseases of wild mammals. J. W. Davis and R. C. Anderson. Iowa State Univ. Press, Ames. 364pp. Frazer, D., J. S. D. Ri tchi e , and A. F. Frazer. 1975. The term "stress" is a veterinary context. Br. Vet. J. 131:653-662. Friend, T. H.• et a1. 1977. Adrenal glucocorticoid response to exogenous adrenocorticotropin mediated by density and social disruption in lactating cows. J. Dairy Sci. 60:1958-1963. __ , F. C. Gwazdouskas, and C. E. Polan , 1979. Change in adrenal response from free stall competition. J. Dairy Sci. 62:768-771. Garrett, W. N., and D. E. Johnson. 1983. Nutritional energetics of ruminants. J. Anim. Sci. 57:478-497. Halas, E. S., M. J. Hanlon , and H. H. Sandstead. 1975. nutrition and aggression. Nature 257:221-222. Intrauterine Hansen, C. G. 1971. Overpopulation as a factor in reducing desert bighorn populations. Trans. Des. Bighorn Counc. 15:46-52. Hassell, M. P. 1982. Impact of infectious diseases on host populations (group report). Pages 15-35 in population biology of infectious aiseases. R. M. Anderson and---itM. May, eds. Springer-Ver 1ag. 314pp. Hicks, L. L., and J. M. Elder. 1979. Human disturbance of Sierra-Nevada bighorn sheep. J. Wildl. Manage. 43:909-915. Holmes, J. C. 1982. Impact of infectious disease agents on the population growth and geographical distribution of animals. Pages 37-51 In . population biology of infectious diseases. R. M. Anderson and R. M. May, eds. Springer-Verlag. 314pp. Hopkins, P. S., C. J. Nolan, and P. M. Pepper. 1980. The effects of heat stress on the development of the foetal lamb. Aust. J. Agric. Res. 31:763-771. �166 Hudson, R. J. 1973. Stress and in-vitro lymphocyte stimulation by phyto-hemagglutinin in Rocky Mountain bighorn sheep. Can. J. Zool. 51:4·79-482. Jensen, M. M .• and A. F. Rasmussen, Jr. 1970. Audiogenic stress and susceptibility to infection. Pages 7-19 in physiological effects of noise. B. L. Welch and A. S. Welch, eds. Plenum Press. New York. 365pp. Johnson, H. D .• and W. J. Vanjonack. 1976. E Effects of environmental and other stressors on blood hormone patterns in lactating animals. J. Dairy Sci. 59:1603-1617. Jung, H. J. G. and G. C. Fahey~ Jr. 1983. Interactions among phenolic monomers and in vitro fermentati on. J. Dairy Sci. 66: 1255-1263. 9 Kattesh, H. G., et al. 1980. Glucocorticoid concentrations, corticosteroid binding protein characteristics and reproduction performance of sows and gilts subjected to applied stress during mid-gestation. J. Anim. Sci. 50;897-905. Keith, L. B. 1983. 40:385-395. Role of food in hare population cycles. Kelley, K. W. 1980. Stress and immune function: review. Ann. Rech. Vet. 11:445-478. Oikos a bibliographic Lanciani, C. A. 1975. Parasite induced alterations in host reproduction and survival. Ecology 56:689-695. L€vin, B. R. 1982. Evolution of parasites and hosts (group report). Pages 213-243 in population biology of infectious diseases. R. M. Andersoniand R. M. May, eds. Springer-Verlag. 3l4pp. Lyon, L. J. 1979. Habitat effectiveness for elk as influenced by roads and cover. J. Forestry 77:658-660. MacKenzie, A. J., C. J. Thwaites, and T. N. Edey. 1975. Oestrus, ovarian and adrenal response of the ewe to fasting and cold stress. Aust. J. Agric. Res. 26:545-551. McNatty, K. P. and D. C. Thurley. 1973. The episodic nature of changes in ovine plasma cortisol levels and their response to adrenaline during adaptation to a new environment. J. Endocrin 59:171-180. 9 Moberg, G. P., C. D. Anderson, and T. R. Underwood. 1980. Ontogeny of the adrenal and behavioral responses of lambs to emotional stress. J. Anim. Sci. 51 :138-142. Myers, K., C. S. Hale, R. r~ykyto"Jycz.and R. L. Hughes. 1971. The effects of varying density and space on sociality and health in animals. Pages 148-187 j~_ behaviour and environment. A. H. Essner, ed. Plenum Press, New York. 411pp. �Colorado Division of Wildlife Wildlife Research Report July, 1984 167 JOB PROGRESS REPORT State of Colorado Project No. Work Plan No. 3 .' ----~----------- Job. No. 1 Period covered: Author: Big Game Investigations 45-01-503-15050 - Noncervids Pronghorn Investigations Pronghorn Popul ati on Dynami cs Study 7/1/83-6/30/84 T. M. Pojar ABSTRACTThe random quadrat and random strip sampling systems for censusing pronghorn (Antilocapra americana) were executed for the third year in the Limon/Hugo area and for the second year in the Great Divide area. Population size and herd structure were estimated in each area using both sampling designs. The objective is to determine if either of these designs will adequately estimate both population size and herd structure during the same fly-over in late summel~. The tests are conducted in 2 vegetative types; short grass prairie (Limon/Hugo) and sagebrush steppe (Great Divide). The quadrat sampling design consistently results in higher density estimates with the difference being rno re pronounced in the sagebrush steppe type. (Please refer to the accompanying tables.) In general, the precision level of both designs is comparable for population size estimates. The herd structure estimates (bucks:100 does and fawns: 100 does) are comparable between the 2 sampling systems. However, the precision level is lower from the quadrat system because of smaller sample size. High variability in the buck to doe ratio inhibits detection of a statistically significant difference between the 2 systems for this parameter. This ratio is consistently higher as estimated by the quadrat system, indicating that detection of solitary bucks may be more likely during the quadrat search. The low variability in the fawn to doe ratios results in relatively precise estimates. With the exception of the 1982 Great Divide data, there was no difference (P < 0.10) in the fawn to doe ra ti os between the 2 samp 1i ng me thods. - ��169 PRONGHORN POPULATION DYNAMICS STUDY Thomas M. Pojar P. N. OBJECTIVES 1. Test the efficiency and precision of a quadrat and strip census design on rolling sagebrush steppe and short grass prairie. 2. Test the efficiency and precision of herd structure estimates when done concurrently with a quadrat and strip census on both rolling sagebrush steppe and short grass prairie. 3. Test the reliability of doing strip and quadrat censusing during late summer when it is possible to do herd structure counts. 4. Test the predictability of the ONEPOP and the POP50 population simulators. 5. Measure and model the effects of reducing a pronghorn population density by 50 percent or greater. SEGMENT OBJECTIVES 1. Test efficiency and precision of quadrat and strip censuses on short grass prairie to estimate density and population structure during one fly-over in late summer. 2. Test efficiency and precision of quadrat census on sagebrush steppe pronghorn habitat to estimate density and population structure during one fly-over in late summer. 3. Simulate pronghorn population changes to evaluate results for each area. ACKNOWLEDGMENTS Appreciation is extended to the following for their cooperation and assistance: J. Ellenberger, J. Gray, G. Bryne, and M. Bauman of the NW Region and M. Elkins and D. Lengel of the SE Region. B. Gill and B. Hernbrode provided essential administrative support for this project. The consulting of D. C. Bowden, CSU Statistical Department, on statistical matters is appreciated. INTRODUCTION The purpose of this investigation is to evaluate the efficiency and effectiveness of 2 census methods in estimating population size and herd structure. This publication reports on the 3rd year of data collection for the short grass prairie area (Limon/Hugo) and the 2nd year for the �170 sagebrush steppe area (Great Divide). The 2 areas of study and the methods used in this experiment are described by Pojar (1983). RESULTS Both the stratified random quadrat and random strip experimental censuses were conducted during 15-18 August 1983 in the Limon/Hugo area. A Hughs 500 C helicopter was used during both counts. The 90% confidence limits for the quadrat census were much wider than in the past (Table 1). This was due to having a few quadrats with a high density of pronghorn and many quadrats with no animals. It may not be uncommon to encounter this situation because of either range conditions that affect pronghorn distribution or simply chance distribution of the pronghorn. Both censuses were completed in the Great Divide area during 6-9 September 1983. A Bell-Soloy helicopter was used for both counts. Once again, the quadrat census resulted in a much higher density estimate than the strip census (Table 2). LITERATURE CITED Pojar, T. M. 1983. Pronghorn investigations -- pronghorn population dynamics study. Colo. Div. Wildl. Res. Rep. July, Part 4:379-385. -. ,1L----1zL /1 (f/'iJay ( Prepared by ~t_/ ~'. Thomas M. POJar Wildlife Researcher �Table 1. Pronghorn census results for the Hugo Data Analysis Unit (Game Management Units A36, A37, A381) using a stratified random quadrat (mile2 quadrats) and a random strip (mile-wide strips) sampling design. 1981 1983 1982 Pop. Est. 90% C. L. Strip Quadrat Strip Quadrat Strip Quadrat 3,570 6,366 2,402 2,602 1 ,837 2,834 .±22% ±26% ±26% .±27% ±27% -±49% lower 2,773 4,710 1,774 1,904 1,342 1,436 Upper 4,367 7,976 3,030 3,301 2,331 4,232 2.14 3.82 1.42 1.54 1.09 1.68 47 67 44 45 44 67 ±19% ±48% ±31% ±51% ±23% ±58% lower 38 35 30 22 34 28 Upper 56 99 58 68 55 105 Fawns:100 Does 64 67 50 42 52 60 ±10% ±17% ±12% ±28% ±17% ±21% Lower 58 56 44 31 43 48 Upper 70 78 57 54 61 73 1,203 381 805 257 592 252 Mean Dens ity (Antmal s/mt , sq.) Bucks:100 Does 90% C. l. 90% C. L. No. C1ass ified I---' •••..J I---' �172 Table 2. Pronghorn census results for the Great Divide Data Analysis Unit (Ga.meManage Units A3, A301) using a random quadrat (mile2 quadrats) and a random strip (mile-wide strips) sampling design. Total area is 1,229 square miles 1982 Strip Pop. Est. 90% C. L. Lower Upper Mean Density (Animal/mi. sq.) Bucks:100 Does 90% C. L Lower Upper Fawns: 100 Does 90% C. L Lower Upper 1983 Quad Strip Quad -------------------------------------49347 16,650 8,868 ±31.3% 2,985 ±17.3% 13,770 ±16.9% 7.373 5~709 3.54 19,530 13.55 10,363 7.22 45.5 52.2 42.9 56.3 ±18.6% ±19.3% 42.2 62.3 83.2 ±14.4% ±17.2% 36.7 49.1 46.6 66.0 65.6 69.1 ±9.4% ± 11.2% 75.4 ±6.7% 61.2 91.1 70.0 61.4 76.8 37.1 54.0 66.8 ±9.4% 60.5 73.1 18,438 ±30.1% 12,883 23,993 16.39 No. Cl ass ified Bucks Does Fawns Total 336 324 603 337 738 493 620 1.404 599 516 921 414 1~567 1,460 1~350 �173 Colorado Division of Wildlife Wildlife Research Report July 1984 JOB PROGRESS REPORT State of Colorado Project No. Work Plan No. 45-0"1-503·-15050 4 ----------------- Job. No. Seasonal Habitat Selection and Act i vi ty of Sympatric Mountain Goat and Bighorn Sheep Populations Period Covered: Author: Rocky Mountain Goat Investigations 7/1/83-6/30/84 D. F. Reed .LSSTRA.CT Nine mountain goats wer'e marked with radio telemetry collars, neck bands , or eartags. Four radio te lerne try collars were rep laced on mountain goats whose telemetry co llars had exp'ired or were due to expire. One mountain sheep Vias marked with a radto telemetry collar. Of the 25 alpine habitats used in the study, mountain boats and sheep occupied 16 and 14 of the habitats, respectively. during September through May. The number of mountain goat groups or individuals occupying the 16 habitats was not greater (P > 0.40) than the same for mountain sheep groups of individuals. -Based on tests of preliminary data, the null hypothesis that mountain goats do not use alpine habitats disproportionately to their availability is rejected. The null hypothesis that mountain sheep do not use alpine habitats di spropor-t ionate ly to their availability is not rejected, and the null nypothesis that mountain goats and sheep do not use the same alpine habitats is rejected. The null hypothesis that mountain goats and sheep do not exhibit the same activity patterns (feeding and resting during the day, and feeding during the night) in alpine habitats is rejected, and the null hypothesis that mountain goats and sheep do not exhibit the same activity patterns (resting during the night) in alpine habitats is not rejected. Thirteen percent of adult female mountain goats had twins, 65 perce~t had 1 neonate or kid. and 22 percent had no kid(s). ��175 SEASONAL HABITAT SELECTION AND ACTIVITY OF SYMPATRIC MOUNTAIN GOAT AND MOUNTAIN SHEEP POPULATIONS Dale F. Reed P. N. OBJECTIVES Evaluate the extent to which mountain goat populations limit seasonal habitat utilization of mountain sheep in alpine environments and to describe patterns and rates of dispersal of mountain goats from colonization sites. SEGMENT OBJECTIVES 1. Test the hypothesis that mountain goats use alpine habitats disproportionately to their availability. 2. Test the hypothesis that mountain sheep use alpine habitats disproportionately to their availability. 3. Test the hypothesis that mountain goats and sheep use alpine habitats that are spatially and/or temporally discrete - their distributions are independent. 4. Test the hypothesis that mountain goats and sheep exhibit the same activity patterns in alpine habitats. ACKNOWLEDGMENTS I thank R. Reiner and L. Reiner of the Mount Evans Research Station, University of Denver, for providing unbounded support and a place for a snowmobile and other field equipment. E. Hashimoto and J. Stone provided field assistance. C. E. Braun provided aerial photography and vegetation maps that included the study area. R. Angel and others reported locations of marked animals during the summer. S. Bassow provided both field and office assistance - her review and discussions of competition models were most helpful. L. Lovett provided office assistance and moral support. DESCRIPTION OF AREA The study area has been described by Reed (1982). METHODS AND MATERIALS Methods and materials have been described by Reed (1981, 1982). �176 RESULTS AND DISCUSSION Capture and Marking Nine mountain goats were marked with radio telemetry collars, neck bands, or eartags. This brings the total of mountain goats marked in the study to 66 (Table 1). One mountain sheep was marked with a radio telemetry collar. Six mountain goat radio collars (4,5,7,17,20, and 55 - Table 2) were still active toward the end of the segment. Of these 6 radio collars, 4 of them were activity (tip switch) collars. Mountain goats 7 and 55 primarily occupied habitats located out of the study area. Hence, little or no data were collected on them for this reporting period. No signal has been received from or sighting made of mountain goat 35 since she was collared. The collar outfitted on mountain sheep 1 was an activity collar. This maintained the number of activity collars on bighorn sheep at 4 (1,7,9, and 10 - Table 2). Habitat Use Of the 25 habitats used in this study, mountain goats and sheep occupied 16 and 14 of the habitats, respectively, during September through May. Conversely, 9 of the habitats (3,4,5,6,10,11,19,23 and 24 - Reed 1982, Appendix A) were not occupied during this period by either species, and 11 were not occupied by mountain sheep. The number of habitats occupied by both species converged from fall to spring (Fig. 1). The number of habitats occupied by mountain goats was greater than the number of habitats occupied by bighorn sheep for the fall months and January (Fig. 2). During previous segments,mountain goats have generally used considerably more of the habitats than those used by the mountain sheep. The number of mountain goat gorups of individuals occupying the 16 habitats was not greater (P > 0.40) than the number of bighorn sheep groups or individuals occupying the 16 habitats during this segment. However, differences were apparent in habitats 2a(E) and 2a(S). These habitats are located above Lincoln Lake and were used frequently by mountain goats throughout the fall and winter seasons. During the past segments, mountain goats had also used these habitats frequently during the spring. However, heavy snow fall in March and April, 1984, may have resulted in different habitat occupancy patterns during the spring season. Also, habitat 2a(E) included a mineral lick and trap site. Similar features of 2a(E) and 2a(S) may account for a relatively high amount of mountain goat use. In theory, the same resources were available to mountain sheep. These habitats may not have been used frequently by mountain sheep because (1) mountain sheep had different requirements, (2) mountain sheep were substantially fewer in number, and/or (3) mountain sheep were excluded by the presence of mountain goats. The largest groups observed for mountain goats and sheep during the seg~: ment were 25 and 35, respectively. The mean size of groups for the 2 species was not different (f > 0.50) �177 Test of Hypothesis 1. The first hypothesis and its null were stated (Reed 1982:100) as follows: Hl = Mountain goats use alpine habitats disproportionately availability. H = Mountain goats do not use alpine habitats disproportionately their availability. o to their to Preliminary data on mountain goat habitat use (Sep-May of this segment) were analyzed by comparing observed with expected frequency distributions. Habitat use was analyzed for number of mountain goat groups (> 1) instead of number of individuals to avoid group behavior bias. Pooled data from other segments will be needed to adequately test such distributions across months and seasons. The number of mountain goat groups per hectare (ha) for each habitat were the parameters used in this X2 test (Table 3). The observed frequency distribution deviated significantly (X2 = 147.9, P < 0.005) from the expected. Hence, based on data from this segment, the null hypothesis that mountain goats do not use alpine habitat disproportionately to their availability is rejected. Test of Hypothesis 2. The second hypothesis and its null were stated (Reed 1982:105) as follows: H2 = Mountain sheep use alpine habitats disproportionately availability. to their Ho = Mountain sheep do not use alpine habitats disproportionately their availability. to Similar to the test of Hypothesis 1, preliminary data on mountain sheep habitat use (Sep-May of this segment) were analyzed by comparing observed with expected frequency distributions. Habitat use was analyzed for number of sheep groups (> 1) instead of number of individuals to avoid group behavior bias. Pooled data from other segments will be needed to adequately test such distributions across months and seasons. The number of mountain sh2ep groups per ha for each habitat were the parameters used in this X test (Table 3). The observed frequency distribution did not deviate significantly (X2 = 17.5, P > 0.10) from the expected. Hence, based on data from this segment, the null hypothesis that mountain sheep do not use alpine habitats disproportionately to their availability is not rejected. Test of Hypothesis 3. The third hypothesis and its null were stated (Reed 1982:106) as follows: H3 = Mountain goats and mountain sheep use the same alpine habitats. Ho = Mountain goats and mountain sheep do not use the same alpine habitats. Preliminary data on both mountain goats and sheep habitat use (Sep-May of this segment) were analyzed by measuring the difference between 2 �178 variables. The 2 variables were the number of mountain goat groups per ha and the number of mountain sheep groups per ha across 15 habitats (Table 3). For continuity with previous segments, habitat 14 (i.e. 16 vs. 15 habitats occupied by mountain goats) was excluded from this analysis. Pooled data from other segments will be needed to adequately test such correlations across months and seasons. There was no significant difference (P > 0.20). Hence, based on data from this segment, the null hypothesis that mountain goats and sheep do not use the same alpine habitats is rejected. Activity Telemetry tip-switch activity collars were maintained on 7 mountain goats and 5 mountain sheep during the segment (Table 2). Generally, both species separated into small groups and dispersed into areas located near the boundary of the study area during the winter. Habitats occupied during the recent winter were often left vacant. Of the 7 mountain goats with activity collars, mountain goat 17 (Blue diagonal - Ch 4) and mountain goat 20 (Black telemetry - Ch 6) provided the largest samples of activity periods. Their feeding and resting periods during the day and during the night (Table 4) were not different (P > 0.40). Variation between the pattern of activity over 24 hours may be compared by examining specific 24-hour periods of selected mountain goats and sheep (Figs. 3-9). In these cases, none of the animals compared had significantly different duration of day and night feeding periods (Table 5). Responses to sunrise and sunset appear to vary between animals and seasons. No responses to moonlight were detected. Generally, it is expected that such feeding and resting periods will vary between individuals, between species, and possibly between seasons, habitats, and environmental conditions. Test of Hypothesis 4. The fourth hypothesis and its null were stated (Reed 1982:108) as follows: H4 = Mountain goats and mountain sheep exhibit the same activity patterns in alpine habitats. Ho = Mountain goats and mountain sheep do not exhibit the same activity patterns in alpine habitats. The mean duration of feeding and resting periods during the day and the mean duration of feeding periods during the night of the 6 mountain goats were not different (P > 0.20, P > 0.20, and P > 0.50) than the corresponding values of 3 mountain sheep (Table 4)~ Conversely, the mean duration of resting periods during the night of 6 mountain goats was different (£ > 0.01) than the corresponding values of 3 mountain sheep. This difference may be due to longer resting periods exhibited by mountain sheep as a strategy to conserve energy during winter nights. The null hypothesis that mountain goats and sheep do not exhibit the same activity patterns (feeding and resting during the day, and feeding during the night) in alpine habitats is rejected. The null hypothesis that mountain goats and sheep do not exhibit the same activity patterns (resting during the night) in alpine habitats is not rejected. Partitioning of the overall hypothesis may be necessary to reveal more definitive variables. �179 Reproduction Of the female mountain goats collared, reproductive status was estimated on 8, 16, 21, 21, and 16 in 1980, 1981, 1982, 1983 and 1983, respectively (Table 6). The mean,age of mountain goats (n = 12) having their first kid or kids was 3.2 ± 0.9 (SO) years. The mean age of mountain goats (n = 8) having twins was 7.5 ± 2.5 (SO) years. Mountain goat 27 (Black collar 1) has had 2 sets of twins, 1 at age 9 and the other at age ll. She alternated twins and singletons during the 4-year period (Table 6). Thirteen percent of the adult (> 4 years old) females had twins, 65 percent had 1 kid, and 22 percent had no kid or kids (Table 7). This percent of twinning is relatively high compared to others reported (Brandborg 1955:94). The rate of twinning, as the rate of having singletons, is likely a minimum since some kids may die soon after birth. An example of this was reported previously (Reed 1983). No reproductive data obtained so far on mountain goats (Table 6) support Heimer's hypothesis of alternate-year production (Heimer and Watson 1982). Furthermore, applicability of this hypothesis to mountain goats in the Mt. Evans study area may be in doubt because dominant male behavior in mountain goats differs largely from that in wild sheep. Also, it would have to be assumed that success in harvesting dominant male mountain goats is comparable to success in harvesting dominant male mountain sheep. This mayor may not be the case. Of the marked or identifiable female mountain sheep, the reproductive status was estimated on 4 (Table 8). Data from mountain sheep 1 (Table 8) may be in support of the alternate-year production hypothesis. Conceptual Models of Competition Although it may be tempting to articulate simple ideas of competition between mountain goats and sheep in this study, in so doing we construct mental preconceptions (Pielou 1981). Despite such preconceptions, 1 simple idea of competition between mountain goats and sheep entails a graph of the 2 species estimated population curves (Fig. 10). If we are confident that these curvesl represent real populations, it is possible that mountain goats will increase to a point at which they will displace mountain sheep through interference competition, exploitation competition, or both. The results from these 2 types of competition are shown in Fig. 11. In the case of interference competition, data already collected on species interactions (N = 88) provide an approximate point on a negative effects continuum (A, Fig. 11; where SP 1 represents mountain goats and SP 2 mountain sheep). This point represents approximately 30 percent of the interactions between mountain goats and sheep where mountain sheep yielded space or other resources as a result of agonistic interspecific behavior. It has been stated that interference competition is unlikely to evolve lBased on Mt. Evans annual ground survey. �180 unless there is a potential for overlap in use of limited resources (i.e. exploitation competition){Pianka 1981). Hence, it is likely that in this case of mountain goats and sheep, resource or exploitation competition is occurring. Of course, it is one thing to assert exploitation competition, and quite another to test that assertion. Since no field data has been collected on exploitation competition so far in this study, a theoretical discussion may be useful. Assuming that both species interact only through use of limited resources, that the resources of forage and space are non-interactive (change in supply of 1 resource has no effect on rate of supply of another){Ti1man 1982), and that mountain sheep require more space and higher quality forage (based on estimates of greater selectivity in diet and habitat ), zero net growth isoclines for the 2 species can be drawn"{Fig. 12). Species A will be able to reduce resource levels to a point below that required for survival of Species B (area between curves A and B, Fig. 12). Species A will be able to competitively displace Species B in all habitats in which Species A is able to survive (total area to upper right of curve A, Fig. 12). Hence, if the system is allowed to reach equilibrium, mountain goats will be able to competitively displace mountain sheep. LITERATURE CITED Brandborg, S. M. 1955. Life history and ecology of the mountain goat in Idaho and Montana. Idaho Dept. Fish and Game Wildl. Bull. 2. 142pp. Craig, E. H. 1981. A relief-adjusted method for determining home range in mountainous areas. J. Mammal. 62(4):837-839. Heimer, W. E., and S. Watson. 1982. Differing reproductive patterns in dall sheep: population strategy or management artifact? Pages 330336 in J. A. Bailey and G. G. Schoonveld, eds. Proc. Bienn. Symp. Nort~Wildl. Sheep and Goat Counc. 3. Fort Collins, CO. 405pp. Pianka, E. R. 1981. Competition and niche theory. Pages 167-196 ~ R. M. May, ed. Theoretical ecology principles and applications. 2nd ed. Sinauer Associates, Inc., Sunderland, MA. 489pp. Pielou, E. C. J981 .. The usefulness of ecological models: taking. Qtrly. Rev. Biol. 56(1):17-31. Reed, D. F. 1981. Rocky mountain goat ecology study. Wildl. Game Res. Rep. July, Part 2:209-222. a stock- Colo. Div. of . 1982. Rocky mountain goat-bighorn sheep competitiu~ study. ---~Co10. Div. of Wi1d1. Game Res. Rep. Ju1y:79-128. . 1983. Seasonal habitat selection and activity of sympatric ------mountain goat and bighorn sheep populations. Colo. Div. of Wi1d1. Game Res. Rep. Ju1y:403-422. �181 Tilman, D. 1982. Resource competition and community structure. Univ. Press, Princeton, NJ. 296pp. Prepared Wildlife Researcher Princeton �•..... Table 1. Date~ age, sex, collar or tag, and selected measurements of mountain goats trapped in the Mt. Evans area durin se ment. Body Hind Horn length (cm) Trap Girth length Wt. foot (cm) (cm) Age Sex Collar/Tag location (cm) Left Right (kg) No. Date 58 6 Jul 83 2 F Orange eartag 6 DACa 59 7 Jul 83 9 F Black collar K DAC 60 7 Jul 83 1 M Orange eartag 7 DAC 61 7 Jul 83 6 F Black collar M 62 7 Jul 83 3 M 63 11 Jul 83 5 64 10 Aug 83 65 66 aOAC = 120 162 24. 1 24.4 - 31.0 DAC 111 175 24.7 22.4 - 31. 5 Orange eartag 8 DAC 107 158 21.3 21.7 - 31.0 F Black collar N DAC 110 166 24.7 24.7 - 30.5 3 F Black collar 0 DAC 112 157 17.3 16.2 - 29.5 27 Jun 84 1 F Blue eartag 1 DAC 88 113 12.0 11.9 - 26.5 28 Jun 84 3 F Black collar P DAC 114 151 23.2 23.1 - 33.0 Data acquisition center site at mile post 7. 00 N �183 Table 2. Number assigned to animal, telemetry collar, channel, frequency, pulse, and activation date for radios outfitted on mountain goats and sheeQ in the ~1t. Evans area. Pulse per Activation Frequency No. Collar Channel min. (ppm) date (mhz) Mountain goat 3 4 5 7 13 14 17 20 29 35 40 55 White Blue Fluorescent green Green 2 Green 3 Red-white-b1ue Blue diagC Black Green diag Blue diag wide Red Green diag 0 148.130 172.487 78 90-65a 22 Aug 80 26 Jul 83 1 2 3 2 4 6 5 4 7 5 172.237 172.262 172.287 172.262 172.312 172.387 172.362 172.312 172.412 172.362 11 48 82 65-90b 65-90 65-90 65-90 60 90-65 65-90 11 29 7 9 10 11 20 24 25 29 Jul Sep Oct Jun Aug Aug Oct Jun Jun Jun 83 80 80 81 83 83 81 82 82 83 Red 3 Yell ow 1 Black and white 9 Blue 8 Black 5 172.287 172.237 172.462 172.437 172.362 65-90 90-65 90-65 90-65 65-90 17 24 4 10 28 Jun Nov Dec Dec Dec 82 82 82 82 83 Mountain sheep 7 8 9 10 1 aActivity (tiP switch) co 11 ars ; 90 ppm = head up, 65 ppm = head down bActi vity (tip switch) collars; 65 ppm = head up, 90 ppm = head down cdiag denotes diagonal �184 Table 3. Habitats, area of habitats, estimated areas of drifted snow pack, estimated areas of habitat available, and mountain goat and mountain shee~ grou~s ~er hectare (ha), 1983-84. Groups per ha Area Habitat designation Habi tata Estimated snow packb Estimated available la lb 2a(W) 2a(E) 2a(S) 2b(E) 2b(W) 8a 8b(N) 8b(S) 9 18 20 21 22 174.1 42.4 14.0 50.6 69.5 17.7 94.2 31.3 5.8 22.6 16.5 32.9 9.9 11. 5 64.6 3.0 5.0 1.0 1.0 6.0 1.0 6.0 3.0 0.5 5.0 1.0 1.0 3.5 1.0 6.0 171 .1 37.4 13.0 49.6 63.5 16.7 88.2 28.3 5.3 17.6 15.5 31.9 6.4 10.5 58.6 TOTAL 657.6 44.0 613.6 Mountain goats 0.099 0.241 0.308 0.524 1.323 0.479 0.045 0.071 0.321 0.170 0.065 0.031 2.344 0.095 0.085 Mountain sheep 0.158 0.107 0.846 0.060 0.378 0.060 0.045 0.141 0.189 0.057 0.258 0 0.312 0 0 aBased on planimeter of habitats overlayed on Geological Survey Topographic Quadrangle Scale 1:24,000; not adjusted for vertical relief (Craig 1981). bOrifted snow sufficiently deep and/or compacted to preclude pawing for forage. Estimates derived from field inspections Sep-May. �185 Table 4. Mean duration of feeding, resting, and moving activity periods of 6 mountain goats and 3 sheep during day (sunrise-sunset) and night (sunset-sunrise). Mean activity periods (min.) Day (sunrise-sunset) No. Collar Feeding (n) Night (sunset-sunrise) Resting (n) Moving (n) Feeding (n) Resting (n) Moving (n) 73.6 (14) 81.2 (13) 10.0 (2) 14.9 (13) 148.4 (14 ) 24.1 (34) 38.2 (47) 37.5 (37) 12.4 (5) 48.0 (4) 67.1 (38) 11.7 (3) 128.5 (58) 207.4 (41 ) 86.5 (4) 117.5 (4) 17.6 (5) 13.7 (3) 26.7 (23) 171.9 (29 ) 9.3 (3) 34.4 (7) 27.8 (6) 222.6 (10) 321.7 Mountain goat 4 Blue 14 Red-white-blue 137. 1 (8) 48.1 (9) 17 Blue diagonal 20 Black 29 Green diagonal Red 40.3 (40 ) 51.3 (38) 101.0 (5) 48.2 (4) 19.3 (6) 8.0 (3) 5.0 (1) 40 88.7 (40) 67.4 (42) 62.0 (3) 49.0 (3) 64.6 (23) 71 .3 (24) 56.3 (19) 10.0 (3) 30.3 (4) Mountain sheep 1 Black 8 Blue 9 Black and white 59.3 (19) 87.3 (6) 68.3 (6) (9 ) �Table 5. Summary of feeding period tests represented in Figs. 3 through 9. Animal/Activity Means N periods y X Y X t SE SE Mountain goat 4 12-13 Oct 83 vs 24.6 9.2 30.7 1- 2 Nov 83 8 6 54.8 -1.066 Mountain goat 20 26-27 Oct 83 vs 15-16 Nov 83 7 4 61.4 22.1 169.0 128.3 -1.103 Mountain goat 17 28-29 Mar 84 vs Mountain goat 20 3- 4 Apr 84 106.6 63.8 19.6 5 5 8.0 1.265 Mountain sheep 8- 9 Feb 84 vs Mountain sheep 10 20-21 Mar 84 6 5 57.2 24.6 32.2 70.4 -0.333 Mountain sheep 10-11 May 84 vs Mountain sheep 10 30-31 May 84 10 9 42.8 10.2 51.3 16.7 -0.448 Mountain goat 17 25-26 Jan 84 vs Mountain sheep 1 2- 3 Feb 84 7 6 74.3 42.9 0.533 47.2 22.3 •...... co Q) df P - Fig. 12 > .20 3 9 > .20 4 8 > .20 5 9 > .50 6 17 > .50 7 12 > .50 8 �Table 5. (continued - page 2) Mountain goat 17 21-22 Sep 83 vs Mountain sheep 10 20-21 Mar 84 9 6 67.8 16.6 57.2 24.6 0.373 13 > .50 9 I--' co '-J �188 Table 6. Estimated reproductive status of collared female mountain goats in the Mt. Evans studt area. No. kids Date Est. age No. Collar banded Jun 84 1980 1981 1982 1983 1984 1b 3 White te1e (813) 22 Aug 80 Unk 8 1a 1 1 4 B1ue te1e (Ch 0) 18 Sep 80 1 2c 0 7 1 0 5 Fluorescent green 22 Sep 80 (Ch 1) 2 8 1 1 1 1 7 Green 2 (Ch 2) 29 Sep 80 10 1 Unk 1 ~d 8 Yellow diag (Ch 1) 30 Sep 80 6 0 0 Unk 10 Black collar 2 30 Sep 80 12 0 2 Unk 0 12 Black collar 3 7 Oct 80 6 0 0 0 1 0 13 Green 3 (Ch 3) 7 Oct 80 11 1 Unk Unk 1 d 14 Red-white-blue (Ch 2) 9 Jun 81 6 1 0 0 17 Blue diag (Ch 4) 16 Jun 81 6 1 1 2 1 17 Jun 81 0 19 Black collar 4 4 0 1 1 Unk 20 Black te1e (Ch 6) 29 Jun 81 9 le 1 1 28 Aug 81 22 Black collar 5 1 1 1 1 6 23 Black collar 6 30 Aug 81 1 1 0 7 24 Black collar 7 4 Sep 81 Unk Unk 8 1 26 Black collar 8 15 Sep 81 9 0 1f 2 2 27 Black collar 1 6 Oct 81 11 1 ld 19 29 Green diag (Ch 5) 20 Oct 81 9 1 Unk 35 Blue diag wide (Ch 4) 24 Jun 82 1 4 1 1 25 Jun 82 0 40 Red te1e (Ch 7) 8 29 Sep 82 1 2 1 Unk 47 Black collar A Unk 29 Sep 82 0 48 Black collar 8 3 30 Sep 82 0 0 50 Black collar E 5 16 Jun 83 0 53 Black collar F 3 29 Jun 83 0 55 Green diag (Ch 5) 6 30 Jun 83 0 5 57 Black collar H 1 30 Jun 83 1 1 6 42 Black collar Jh 2 7 Ju1 83 10 0 59 Black collar K 0 7 Ju1 83 1 61 Black collar M 7 2 1 11 Ju1 83 63 Black collar N 6 1 10 Aug 83 0 64 Black collar 0 4 1 28 Jun 84 66 Black collar P 3 aWh ite eartag 1. bConfirmed with 1 kid 2 Jun; without kid 29 Jun and thereafter. CParturition 1 Jun yielded 2 kids; with only 1 kid 4 Jun and thereafter. dMorta1ity fall 82 - summer 83. eB1ue "_"; after eartag pulled out known as "injured ear. fB1ack eartag. 9B1ue eartag. hprevious1y White eartag 4. II �189 Table 7. Number of adult (> 4 years old) marked female mountain goats associated with 0,1, or 2 neonates or kids across years in the Mt. Evans stud. Total Years No. of neonates 1980 0 1981 1982 1 2 1 5 0 0 9 2 4 14a TOTAL 6 11 1983 7 1984 No. Percent 2 10 2 4 8 3 16 46 9 22 65 13 20 19 15 71 100 Table 8. Estimated reproductive status of marked or identifiable female mountain sheep in the Mt. Evans study area. Collar or Date Est. Age No. Identification Banded Jun 84 1980 1981 1982 1983 1984 1 2 3 4 5 6 9 10 Black te1 a Yellow 195 Plain ye llowEartag 18 Eartag 4 Partial albino Black and white tele (Ch 9) Blue tele (Ch 8) 1 Feb 77 1 Feb 77 77 1 Feb 77 25 Jan 77 4 Dec 82 10 Dec 82 12 12 Unk 7 7 Unk 10101 1 0 Unk Unk o 0 o Unk 1 0 Unk 1 Unk 1 6 5 1 Unk Unk Unk Unk 1 2 o ,1 1 1 apreviously Yellow 17; Black tele collar added 28 Dec 83. b~1ost numbers have come off. Only stitchi ng and faded areas make identification possible. cNo stitching or faded areas apparent; collar frayed on bottom-posterior. �190 15 10 "d Q) ....• Po ::s t) t) 0 III ~ CIS ~ ....• ~ ::c: ~ 0 . 0 :z; 5 o Fall Fig. 1. Winter Spring Number of habitats occupied by mountain goats (dashed line) and sheep (solid line) by season (fall = Sep-Nov, winter = Dec-Feb, spring = Mar-May) 1983-84. �191 15 q -0 10 \ \ CI) '"::I P. \ CJ CJ o \ ? ~ I '1-1 o . o Z / 5 / / / \ b----d o Sep Fig. 2. Oct Nov Dec Jan Feb Mar Apr May Number of habitats occupied by mountain goats (dashed line) and sheep (solid line) by month 1983-84. �•..... lD N 2400 2400 er:~0600 1200 1200 A B Fig. 3. Mountain goat 4 (Blue telemetry - Ch 0) feeding (horizontal lines), moving (vertical lines) and resting activities during 24-hour periods 12-13 October 1983 (A) and 1-2 November 1983 (B). �2400 2400 1800ttL.~~t1= ~ 'fJ0600 1200 1200 A B Fig. 4. Mountain goat 20 (Black telemetry - Ch 6) feeding (horizontal lines) and resting activities during 24-hour periods 26-27 October 1983 (A) and 15-16 November 1983 (B). f-' 1.0 W �•..... 1..0 +=- 2400 b 1800 ?\Vz~:g 2400 r=*=~~0600 1200 1200 A B Fig. 5. Mountain goat 17 (Blue'diagonal - Ch 4) (A) and mountain goat 20 (Black telemetry - CH 6){B) feeding (horizontal lines), moving (vertical lines) and resting activities during 24-hour periods 28-29 March 1984 and 3-4 April 1984, respectively. �_j 2400 1800R' 5jJt>- ,,;:tr 2400 0600 . 1200 1200 A B Fig. 6; Mountain sheep 1 (Black telemetry - Ch 5)· (A) and mountain sheep 10. (Blue telemetry ::-._Ch 8) (B) _ feeding (horizontal lines), moving (vertical lines) and resting activities during 24-hour periods 8-9 February 1984 and 20-21 March 1984, respectively. t-' \.0 U1 �•..... 1.0 0) 2400 1800 F - - S?¥ 2400 >$ , t 7] 0600 === ~ 1200 1200 A B Fig. 7. Mountain sheep 1 (Black telemetry - Ch 5) (A) and mountain sheep 10 (Blue telemetry - Ch 8} (B) feeding (horizontal lines), moving (vertical lines) and resting activities during 24-hour periods 10-11 May 1984 and 30-31 May 1984, respectively. �, 2400 2400 0600 1800 1200 1200 A B Fig. 8. Mountain goat 17 (Blue,diagonal-Ch 4) (A) and mountain sheep 1 (Black telemetry - Ch 5) (B) feeding (horizontal lines), moving (vertical lines) and resting activities during 24-hour periods 25-26 January 1984 and 2-3 February 1984, respectively. I-' I.D -.....J �I-' 1800 t ':;;)jft' \.0 2400 2400 1\ J co 0600 1200 1200 A B Fig. 9. Mountain goat 17 (Blue diagonal - Ch 4) (A) and mountain sheep 10 (Blue telemetry - Ch 8)' (B) feeding (horizontal lines) and resting activities during 24-hour periods 21-22 September 1983 and 20-21 March 1984, respectively. �200 _......0 150 _,../ o \\ / ~/ »: -; " '\ \,\ 100 /f,O-::: /' '.. ./ / @- -<, -'- / 0'-' ' / <, ,'-..... . / -: ./ ------.___ ./ -.--0------ .-"" ~ // -'0 . /A 50 if -----·-·----- ..• ·----·----------r-----·-·---·--.,----·· 78 79 80 ----------.-,--------.. 81. ~-------~ 82 83 I-' 1.0 1.0 Fig. 10. Ground census estimates of mountain goats (dashed line) and mountain sheep (solid line) in the Mt. Evans area 1978-83. ---_--_--_ - --_-------- - �200 SP 1 SF 2 DISPLACEMENT U) Eo< u ~ INTERFEHENCE A ~§Z g;H E-' HZ E-'O <U C,!) ~ SP 2 SP 1 DISPU_CEMENT COMPETITION SP 1 SP 2 LOCALLY EXTINCT U) Eo< U ~..,.. ~S "-EXPLOITATION ~::::> Z --g;~ HZ jeOEXISTENCE @ EQUILIBRItTM _ _ RESOURCE PARTITIONING Eo< C' <U C,!) ~ SP 2 SP 1 LOCALLY EXTINCT Fig. 11. Results of competition (interference, exploitation, or both) as expressed on a continuum where species 1 (SP 1) and species 2 (SP 2) dominate at opposite ends of the continuum. Neither species dominates at a point represented by the middle of the continuum where negative effects are equal. Field data on interference between mountain goats and sheep are represented at point A on the continuum where SP 1 represents mountain goats and SP 2 mountain sheep. �201 B I j lUi~SO TReE 2 I \ \ \. ,,-.-- ...--- ....---.----.---~.---.-------.---.----------. - RESOURCE 1 Fig. 12. Zer-o nee 0r'o\JJ~:h t socl i nas for 2 species (A := mountain goats, mount.ai n ::..sheeo :~E'\ imount of resource ..J • '., e' .. ) "'('''rr' .c .. ::! resource '''>. ..L 'J l -: r,p-f'll'(" .. c.~.'.~ .:» ',., )1 <::Il-C'" ~~,d \.. and •• 2 represents emour.t of fora.:::e (modtf \"!d f\"'I)!rl Tt Iman 1982 :73). ~.'j .:\ lII. B = ��Colorado Wildlife JUlY Division Research 203 of Wildlife Reoort 1984 JOB PROGRESS REPORT State of Colorado Project No. 45-01- 503- -----'--15050 \~ork Plan No. 5 Job. No. Personnel: - Noncervids Black Bear Investigations Biack Bear Population Period covered: Author: Big Game Investigations Ecology 7/1/83-6/30/84 T. D. I. Beck J. T. Broderick ABSTRJ·\CT Twenty-seven bears were captured during the 1983 season, 17 initial captures and 10 recaptures. Most new captured bears were immigrant/ migrants but one old resident female was caught. Eight marked bears were known to die during the period; 2 illegal kills, 4 legal kills, 1 handling death, 1 natural death. Mean age of first reproduction for females is 5.7 years, litter frequency is 2.3 years, and mean litter size is 2.07. ~1a.lesappear to be denning about 2 weeks later than females. Denning occurs from early October to November. Males use rock caverns extensively for dens while females use more types of excavated dens as well as rock caverns. ��205 BLACK BEAR INVESTIGATIONS Thomas D. I. Beck P. N. OBJECTIVES 1. Develop techniques tion Ieve l s , 2. Determine 3. Describe population harvest and habitat habitat to accurately prefeY'ences and precisely of selected dynamics sufficiently manipulations. estimate bear popula- bear populations. to ailow analysis of various SEGMENT OBJECTIVES 1. Estimate age when female of litters. black bears have first litters, litter size, and frequency 2. Calculate home range size and seasonal home range dynamics for black bears 3. within the Black ~1esa study area. Document survival of radio-collared black bears with emphasis in age classes 2. 3, 4, and 5. on bears 4. Determine habitat preference of 8 female black bears in the study area. ACKNO~JLEDGtv1ENTS Assistance with field work was provided by the following Division of Wildlife personnel: G. Bock, J. Broderick, M. Dege, J. Ellenberger, J. Freedman, M. Grode , M. Haroldson, C. Haynes, R. Hoffman, D. Homan, M. Jandreau, G. Ti schbei n. Addi tf onat ty, H. Har-low from U. of Wyoming assi sted with den work . Thei r assistance vias necessary and much apprec ia ted. fvlETHODS AND fvlATERIAL.S .E.QQu1at i o_:~.Oeseri pti on Capture efforts lasted from 8 June to 10 July and 20 August to 12 September 1983, in the Black r"'esa study area. June and July efforts were concentrated in the Saddl e ~10unta'in area and the Dyer Creeks. Two specific females wi th inoperable collars were being sought in the Saddle Mtn. area whtIe the Dyer Creeks have not been adequately trapped because the snare line takes 5-6 hours to waIk , ~1uch of the roaded portion of the area was poorly accessible in June because of severe flooding and mudslides. All captures ~'Jen~made using Aldrich spring-activated snares. All snare sets \'/ere out of s iqht from roads and trails and were baited with scraps and honey. Snares were checked daily. �206 Snared bear's ~'Jere imnob t l ized with a combina ti on of ke tamtne hydrochloride and xyl az i ne hydroch 1ori dt? -j n a rr!; xture of "180 kg ketami ne and 45 mg xylazine per mi l l i l i ter. Drug vias adnrint s tered by use of a 1.8 m jab pole. Denned bears were drugged by use of a jab pole 0.5. 1.2, or 1.8 m in length. Dt'ug dosage was 6.6 mg ketamine per kg of body we iqht , Bears were ear-tagged in both ears with numbered, plastic Rota-tags. All female bears age c l ass (f-\C) :3 or older' and maIe Dears AC 5 and older were instrumented with a TeJonics radi o tr-ansmi t.ter collar. Yearling bears tha t were hand" ed 'i n dens with the ir mother, and thus known to have been born in the study area, were instrumented with small radio-transmitter collars designed to operate for 16 months, These small collars were removed from AC 3 b_ars during denning and replaced with larger radioThe small collars have a short section of surgical rubber tubing which will deteriorate ~nd allow the collar to drop free should we not be able to remove the collar in tie next. den. collars. Habitat Use Data on qenera l habitat use we re collected nr+mar i ly from ground tracking radio-collared bears supplemented by weekly aerial tracking. Den sites were located in October-Decembe~ 198~ by both ground and aerial tracking. Den entrances were marked wi th riu-stop nylon nagging to facilitate location in r~arch. Dens were v is ited in January, March, and April, 1984. and numerous phys ical measurements 01' the den were taken as well as a general site description. habitat se le .tion studies on g f'emal e black bears were conducted the year as part of M. H~roldson's maste~s degree program. Analyses are still in prDgre~s anci detailed reporting of this portion of the project will aw~it completion uf the thesis (target date December, 1984) . Deta-iled throughout All bear locations were recorded by UTM coor·dinates. The spatial description of the ranges employs the minimwn polygon procedure and utilizes the HOME computer program described by Harestad (1981). No significant changes in home range size occurred this year so reporting wi ll emphas i ze the migrant portion ot out' study population. r1igration distances are rep1rted -s straight line distances although the topography traversed dictates much longer trips. lhe migrant bears are seldom located more fre;iuent!.v than each TO days unt 1"1 they move into the a rea) thus times of travel are py-e:,~cntt;:, i::l~: mul t i;Ie tay slots. RESULTS AND DISCU3_ION Po pu 1.9-_t i_~_Il__Q_~?!~_tJL!:j_2_!_ The 1983 trapping season was quit~ successful with 17 initial captures 10 recaptures of bears caught in previous years. The 2 target females (F-13, F-..20) important collar with tnoperabl e co l l ars ItJp.l"e both caught in June , and Other recaptures included a migrant female who had slipped an earlier (F-·27), a resi dent f'ema l e v/'ithher' first 'l it ter that had been �207 caught but not collared as AC 3 (F-2·j L and a very 1arqe , mature rna le (M-16) who had slipped an earlier collar in October, 1981. We were unable to collar M-16 again as his neck has equal circumference as his head and our collars are not long enough. Of the 17 new captures, 4 were cubs (AC 1).2 were caught while snaring outside the study area, 11 were cauqht at. a time and place to suggest they are miqrarrts or' inmJigrallts~ and 1 adul t fema-le has probably been eluding us for the past 5 years (F-41). Since we do not rol1ar all captured bears, it will take future recaptures and current radio-tracking to determine the status of the l l imgrant/imll1igrant bears" Three of this group (t1-4·8, r~-43:, and F-47) are known to be migl~ants. Summar-ies of all bear handlings throuqh the 1984 den season are provided in Tables 1 and 2. Male black bear's continued to recei ve the brunt of man-re l ated mortality in 1983. However, illegal kills were much lower than previous years. M-5 was shot and abandoned during elk/deer season in an area open to hunting north of the study area .. F-7 WaS fatally shot in the study area on 5 November by an elk hunter WhD al"leged the bear was attacking him. She had 2 cubs with her on 4 November but hunter did not see them. No charges were filed against the hunter. Four male black bears were legally taken outside the study area (M-ll, M-17. r'1-25, n-so). M·30 WCi.S co llared 9···2···8"1 but the signa"i quit on 9-17-·8"1 while in the study area. ~rh:~n the co llar was returned in June, 1983, Vie discovered an "old wound" in the transmi t ter - a Iarqe caliber bullet lodged a~ainst tile tra.nsmitter housing. An AC 3 male, M-34, was found dead in his den in the summer of 1983. Deep snow and ava l anche danger prohib i ted examination of the densite in late May. thus the carcass was rompletely decomposed. The den had been completely flooded at some time 'in the spring, as evidenced by debris from the bed being stuck to the den roof and walls. The body of the bear was in a significantly different position than what he was left in after handl inq in January. Breath inq was normal dur-ing the winter handling. Although it is possible the bear died from handling, I believe the bear drowned in his den. The snowpack was >4 m deep of hardpacked snow; cold, wet weather i n April and May delayed snowme l t until the hot days of late May sent mas~ive floods off the slopes. There was a 14-cm di ameter water entry hole on the uphi l l side of his den and a downed log angled for 20 m which woul d divert water into the entry hole. I think these bad combinations caused rapid den nooding before the bear could dig out. This den was dug with >40 em of snow on ground because M-34 was disturbed from his first den by hunters. An AC 4 rna 1e , 1'+··33, di ed from an adverse reac tt on to druggi n9 duri ng 1984 den work. It is probable that the drug was injected into a blood vessel rather than muscle. About 2 minutes after injection, the bear gave one large exhalation and slumped. We artificially respirated (mouth-to-nose) the bear for 2.5 hours with no response. Fecundit.y of the study population remains Iow , ,1\11 litters born in 1984 were to females who had previously had cubs so mean age of first litter remains at 5.1 years. Of 15 newborn litters. 3 were singletons, 8 were twi ns , and 4 wen2 triplets; thus mean lttter size was 2.07. Of 31 �208 newborn cubs checked. 15 ~I/er'e females, "14ma es , and 2 unexami ned. Mortality of cubs in the first 6 months has been hi qh wi th 10 of the 31 cubs not denning with the mother in their first year. I am making the assumption that all cubs alive will den with their mother if she is alive during their first winter. Although this assumption is not strongly supported by difficult to ob tain data, the converse event (cubs den solitary when mother is alive) -is also poorly documented. Two periods of cub disappearance stand out - Jun~ and October-November. I suspect the losses in June are to non-man related causes primarily while the fall losses are to hunters. We have documented instances of the latter. The sex rat i o of the l ost cubs was 5 male, 3 f'emal e and 2 unknown. F··,29 had 2 cubs in 1983, apparently lost them in late June of 1983, then had another litter of 2 in 1984. She has apparently lost this litter also in early June. 1984. Of the 21 cubs surviving the first 6 months~ 5 died of starvation whi Ie denned with thei r mothers fonowing the catastrophic berry f'a i lure of -1981. Interval between litters was 2.3 years , based on 9 females involving 17 litters. Six females are lHtering every 2nd year while 3 are on irregular patterns. Generalizations from current population models are unwarranted because of the hiqh J\C 'I mortel ity as compared to an assumed AC -I mor-ta l t ty of 5% in the models and the s iqm+icant i'Jle~Jdl kill. Fecundi ty rates appear to be simi lar to other Rocky fVlountain areas although much less than eastern USA black bear populations (Bunnell and Tait 1981). Hopefully. within the next year more flexible population models will be available. It is wise to note the small sample sizes in all bear data and how rapidly the mean values can ch2nge. Although sounding trite. the longer the run of data. the Iess chance for mak i ng crl tical errors . Prel imi nary ana lyses of population modeling indicate d population with the characteristics of our study population coul d , at best, SUPPOt~t a mean annual mortality rate of 9%. As our population i s bel ieved to be at a dens i ty of approximately one-half the model popul ation, the sustainable mor te l ity is actua lly less than 9%. Documented morte llty on our- resident study population is approximately 10% annually. Habitat Use The period of August 15-20 once again marked the movement of all radioco 11ared bear's (N=31) into the Gambel oak-servi ceber-ry brush communiti es. Fourteen collared bears have been identified as migrants; 10 female and 4 male. Since we collar few male bears, it is likely that others of the males we tag and release are also migrants. A listing of seasonal ranges is provided in Table 3. The fall berry supply was excellent in 1983 and the Doug Creek area supported an especi a I ly abundant crop of servi ceberry (Amel anch i er a -j ni f'olia }. The importance of the Cow-Cl ear Fork-Doug Creek regioncallrloCb'e-overelnphasized. The s tudy area as originally outlined in 1979 did not include thes~ drainages on the north end. In August, 1979, all 8 collared bears moved into Cow Creek and our study area grew. Each year sinc~we have watched this concentration of bears in late August and tried to exp lo it i - f or better' trapping success. In September, -1983, all 31 collared bears spent at least part of 6 days in Cow Creek, an area of 18-20 km2. The entire Cow-Clear Fork-Doug Creek complex of Gambel oakserviceberry shrub lands 'i s approximately 42 km2 and prov+des critical fall habitat for an the col l ared resident bears of the 595 kmt study area and �209 a very significant number of migrant bears from as far as 37.0 km to the east. It would be useful to know what percentage of the bears in the areas to the east of the study area do.migrate to the Gambel oakserviceberry shrub type. This vegetation type does not extend east of the study area boundary in this region of the state. Management considerations abound. This particular area is obviously of vital importance to the 60-90 estimated resident bears in the Black Mesa study area for fall berr-ies and acorns. The area is also of great importance to some portion of the bear population residing in the region from Curecanti Creek east to Ohio Creek - an east-west linear distance of 32 km. Of the 42 km2 tha~ is considered of great importance, only the Cow Creek area (18-20 km ) is public land (USDA Forest Service). 2 Management for black bear habitat should concentrate on the entire 42 km area with the realization that half the area is being managed for multiple use and half for single use (cattle). Management for bear habitat will maintain the old-age structure of the Gambel oak (Quercus gambelii) stands and the chokecherry (Primus virginianus) and serviceberry shrub thickets. Our ignorance of hard and soft mast production in these 3 species is large. In reference to such ignorance it would be unwise to set about to alter the current situation until we could predict the response of these necessary berry producers. I am keenly aware of the ability of black bears to find every suitable microsite of habitat but here lies an area of 42 km2 that can truly be considered critical on a macro-basis and should be maintained. Enhancement would be nice but I am concerned the needed ecological understandings are lacking. These areas may only support 6-10 resident bears but are vital to the survival and reproductive output of 40-100 bears. Home Range Ana lyse_?_ All findings in 1983 that differed from 1982 were covered in the migration discussion under habitat. Basically, home range size and relative sizes between age and sex classes has not changed from last year. More detailed examination of family group home ranges through time and reproduction cycles are planned for the next 2 years. LITERATURE CITED Bunnell, F. L., and D. E. N. Tait. 1981. Population dynamics of bears implications. Pages 75-98 in C. W. Fowler and T. D. Smith, eds. Dynamics of large mammal populations. J. Wiley & Sons, N. Y. 477pp. Harestad. A. S. 1981. Computer analysis of home range data. Fish & Wildl. Branch, Fish & Wildl. Bull. B-11. 25pp. Prepared by __ 1 _1kL__ Beck Wildlife Researcher ThO~ Brit. Col. �210 Table 1. Estimated age and weight of captured male black bears, Bl ack ' Mesa study" __area L_COo . _ ---~.------Capture Estimated 1. D. No. Date Age Weight (kg) Remarks M-l 5-29-·79 6-·29,·,79 j~-2 6-B-79 6-27-- 79 7 7 141 132 9-15-81 M-3 7-25-79 -12 M-4 M-5 7-30· ..79 'lO7 5-2'7-·81 6 2 3 4 4 '1·-24-82 5 n8 M-6 9-n -79 :3 TI M-7 8··'11-79 5 84 M-8 8-27-79 ~~-9 9-20-,79 9-'J 2-80 2 1 2 52 48 M-10 5··-28,-80 5 t -114 M--dead 4-- 2·-82 6-- 3-80 9 159 4 6 52 70 95 80 5 5 118 6- 8,·,80 6 130 6-·16 .. 80 .J .~ 48 9-18-80 3 71 1"1--15 6--18-80 3 68 tJl--16 6--19-80 9 '139 155 6- 2-80 2-25-81 M-ll 6- 3--80 2-·23·-81 9-·29-81 1--23-·82 M-12 6- 4-80 "10- 7-,80 fVj- 13 ~1--'J4 7-· M-17 I:' o ") ,)-0,,) 6-25-80 3-2'7-8'1 9- g·-S-I 3--13-82 h ,) 5 12 3 4 4 5 15 km S of 68 4 4 6 8·- 2-79 Killed 7-12-79, study area 66 138 12 J Illegally killed in study a rea, 'lO-82 Illegally killed in study area, 11-80 41 76 89 86 61 Illegally killed 11-5-83, 2 km N of study area Killed 7-14-80 approximately 155 km SW of study area Suspected illegal kill but UNCONFIRMED Killed 9-81 by ADC9 S of study area 15 km 127 Illegally snare killed while in Killed 5-22-83 study area 0.3 km N of 91 Died from drug reaction in den. 3-81 Killed 6-8190.3 km N of study area Illegally killed 10-80 in study area Suspected illegal kill but UNCONFIRMED 55 89 95 Killed 6-7-83, study area 2 km S of �211 Table 1. M-18 M-19 M-20 (continued 3 7- 3-80 4- 3-82 1-28·-84 3 7- '12···80 3 6-24-81 4 5 2 2 M-22 ~1-23 9,·,-14-80 M-24 M·-25 9-25-80 9-25~,80 4- 1-8"1 10-20-83 M-26 M-27 M-28 2) 7_. 1-80 2-'16-82 2-15-82 8-·13-80 8-20-·80 M-21 -~ 10- 6-80 6-10-81 4,·· 1-8'J 9-· 1,·81 2-19~·82 57 5 Illegally killed in study area, 10-16-82 59 5 125 64 66 7 46 34 Killed 6-26-82 25 km N of study area 82 4 3 1 2 4 80 32 25 Killed 6.5 km N of study area 4 1 2 2 3 80 11 20 52 34 Illegally killed 11-10-82, 19 km N of study area 6-22-81 6-22-82 3 4 r"-30 9- 2·-81 o M-3l 9- 5-81 9-11-81 3 3 3 5 2 2 3 3 3 M-29 9-14-81 7- 9-·83 M-32 M-33 3-17 ·-82 2- 6-,82 1-11-83 6-17-83 6-19-83 c 2-16-84 3-17-82 3- 3-83 2- 6-82 L'f M-36 3·- 5-83 M-38 M-39 M-40 8-29-82 M-41 M-43 6-·21-83 8-21-83 M-44 8-27-83 M-45 8-29-83 M-48 2--24..·84 9- 2--83 2 3 2 2 6 5 4 5 6 '1 M-34 M-35 8-·29-82 8--25-83 2 3 2 61 59 . 102 Died in 7-82, believed to be natural causes Killed 6-22-83 19 km E of study area 56 56 56 82 16 23 Starved to death in den, 1982 48 Died of drug reaction in den 15 11 20 Drowned in den, Spring 1983 Starved to death in den, 1982 �212 Table 1. M-49 M-49-2 M-Sl M-52 (continued -~,-,,-e_3:c..<) 7- 6-83 9·· 2-83 9-· 4-83 9··'0···82 4 10 4 3 9- 4-83 4 2··15-84 2 2 _ 70 Was tagged as M-40 in 1982, changed to M-52 in 1983 M-59 M-62 3- 7-84 __ . 39 25 . Table 2. Estimated age and weight of captured female black bears. Black Mesa stuc!y ar~~2..CO: _ Capture Estimated 1.0. No. Date Age Weight (kg) Remarks F-l F··2 6-'13-79 6 70 6- 8···8"1 '"'? Uc. 3··14-82 8 9 107 6-'17-79 1 n LlIeqa l Iy killed in study area, 9- 12-82 Llleqa l ly killed in study area, 11-79; believed to be cub of F···I F-3 ].- ]-/9 k il Ied in study Illegally area, 11-79; believed to be cub of F-4 7-18·-79 8-'16-79 F·-5 F-6 F-7 ].·'-19··79 7-·20-79 7--2"1-79 7·· 9-80 3- 9-·81 3·-22-82 3·-19-83 2 2 2 4 6 7 8 9 10 F-l 27 33 27 D.·8 75 64 59 64 82 Il"Iegally killed in study 11-5-83 I il led 10-79, 1 m N of study area, F-8 8·· 1-19 5 43 F-9 8- 1-79 7 9 68 area 4- 1-81 6-"10···81 9 10 1'1 F-12 3- 15-82 3- 5-83 7- 5··83 2-29·-849··10-79 9··14-- 79 9-17-·]9 F-13 5-25-80 4 5 7 8 F-10 F-ll 3··19··81 6-·11·-83 3- 6-84 70 61 11 12 3 12 3 59 107 50 57 50 Killed 11-81 0.4 km E of study area �213 Table 2. F-P, F-15 F-16 F-17 F--18 (cont~nued - pa_9.e2) 6-- 7-80 3-24~81 3-28-82 3-27-83 3- 2-84 t~ 57 5 6 t 6- 8--80 8 10 3-28--81 9-22-81 3-21-83 3- 9-84 11 13 14 105 6--19-80 4 45 9- 5-80 4- 10- 3-81 5 54 71 6 68 3-20-82 4-17-83 3- 7-84 6-21-80 3-15--81 2- 6-82 3-12-83 2-15-846-24-80 6-20-81 n 98 95 86 Had open bullet wound 93 7 8 6 1 75 76 84 8 9 10 77 1 2 27 15 Had open bullet wound on 6- 20-81 . Ki 11 ed 11-6-81 0.8 km N of study area F-19 F-20 F-21 F-22 F-23 6-25-80 3 2-22--81 4 3-21-82 3-16-83 6-27--80 3-10-81 6- 9-83 3-· 8-84 6-30-80 9- 3-83 '1-22-84 7- 1-80 3-16-81 3-17-82 3-26-83 7- 9--80 5 6 5 3- 9-81 F-24 F-25 F-26 F-27 F-28 F--29 9-· 3-80 3-28·-83 10- 4-80 6-19-81 9-14--81 8-31-83 9-·16-81 3..24--82 3",15-83 9- 19-8'1 3--"19-82 6 8 9 3 6 7 9 10 11 12 1 2 5 8 2 43 57 61 61 66 30 74 64 91 13 20 52 2 36 31 39 4 15 82 2 16 17 7 8 84 �214 Tab'le 2. F-29 F-30 F-3i (conti nued - ~age 3) 3-17-83 3-29-84 9-20-81 1-·22·-82 3-19-84 3-19-82 3-21-83 1-2'1-84 F-32 3··24-82 ..)7-83 'l 9 10 2 3 5 2 3 4 2 3 4 2 2·· 5··84 F-34 2-· 6·,82 32 23 14 27 48 22 48 20 Starved to death in den. 1982 F-36 F-37 6- 4··82 2-- 7··83 6-19-83 3 34 4 61 4 5 4 3-23-84 8-29-82 F-·39 8·-31-82 '1·-27··84· 9-·15--82 F-40 6-·20·-83 F-41 8···25-83 10 3·· 6-··84 1"1 F-38 F-46 F-47 F-48 F-53 F-54 F-55 F-56 F-57 F-60 Illegally ki 11ed in study area, 9-82 2 4 4 3 86 59 93 '19 8-30···83 1 1··22-84 2 9- '···83 7- 6··83 9- 6-83 6 o" 1 66 16 9···11 ..·83 'j '15 9·-11-83 1-22-84 3·· 2·-84- ? 2 30 2-15···84 2 25 84 3- 2.. 2 , �215 Table 3. Seasonal ranges of migrant in the fall season. bears found in Black tlJesastudy area ---------"<"".-----.,-~----- Winter-Spring-Summer Straight-line 1. D. No. range Fall range distance (km) ._-----------------------_._----------E. Soap Creek E .. Red Creek Coal Creek Beaver Creek Curecanti Creek F-"Il Soap Cr-eek F-12 Soap Creek F-24 w. Elk Creek F-27 E. Soap Creek F-28 Soap Creek F-30 E. Soap Creek F-32 Curecanti Creek F-40 Coal Creek F-47 Beaver Creek --_----_._---_ _---.- •... _. M-6 M-30 M-48 M-43 F-7 ... Table 4. C~ronology 1980 1981 1982 1983 % Cumulative Females 1980 1981 1982 1983 % 24.0 30.5 27.2 37.0 14.5 17.7 16.0 24.1 20.8 18.8 20.9 17.7 19.2 37.0 of ,den en!El.._by radio-collared black bears. Den entry time October November Week 1 Males __ ._- Clear Fork Doug Creek Doug Creek Doug Creek Muddy Creek Cow Creek Vi rgi ni a Creek Cow Creek Cow Creek Doug Creek Doug Creek Doug Creek Doug Creek Doug Creek o 1 o o 3.8 3.8 o ° °1.5 1 1. 51 Cumulative ::=:::::::::::=========== Week Week 234 o o o o o 0 1 0 Week Week 1 Week Week 234 020 2 Week 1 320 1 o 1 .-~-o 1 1 7.7 7.7 3.8 11.5 19.2 o 3 420 1 2 5 2 'I 344 1 2 9 6.2 12.5 32 .8 7.7 20.2 53.0 Week December 1 3 3 ° °1 ° ° 11.5 11.5 0 Week 2 2 0 1 0 ° ° 7.7 0 19.2 26.9 3.8 38.4 65.3 76.8 88.3 92.1 99.8 1 020 2 3 1 300 2 0 ° °1 __ 0 0 0 3 3 0 1 7 • 2 1.:...:5~ • .::__6 _1__ O:::..:.~9_~O_.-:3'--". O::___ 70.2 85.8 96.7 96.7 99.8 99.8 �Ta.ble 5. Site characteristics of dens used by male black bears~ 1980-84. Overs tory Elevation Aspect Slope Relative VOlu~el Den entrance Den type (m) vegetation (degrees) estimator (m ) (%) (width x height; cm) a? Rock cavern Quga3 2583 360 33 1. 54 146 x 34a Rock cavern Potr 2730 20 1.28 66 x 53 60 Rock cavernb Psme 2867 30 2.49 137 x 82 69 b Rock cavern Psme 2882 67 65 3.91 83 x 54· c 2(\ Excavated-shrub Prvi 2556 22 1.47 61 x 31 rRock cavern~ Psme 2928 205 x 36 130 45 2.70 R' LOC!, cavern'd Psme 2867 112 53 1.67 35 x 74 d Rock cavern Ta.lus 2913 224 67 9.49 65 x 120 e ock cavern Psme 2577 1100 x 33 352 57.20 86 e Rock cavern Psme 2554 2.81 100 75 61 x 61 Rock cavern Psme 2837 297 41 x 40 77 nd Rock cavern Pien/Abla 3241 27 10.55 165 x 63 48 f Rock cavern Psme 2638 53 x 61 60 120 3.99 f Rock cavern' Quga 2715 270 34 x 45 54 115.09 Rock cavern f Pied/Jusc 2379 185 63 2.14 67 x 42 f Rock cavern QugajPied 2570 160 67 1. 17 48 x 44 Rock cavern PsmejPi fl 3,06 2806 250 105 280 x 50 Rock cavern Potr 2745 240 54 0.63 81 x 37 Excavated-sh rllb9 Prvi 2577 275 0.45 51 51 x 33 g Rock cavern PsmejAbla 3'11 2623 68 2.17 85 x 89 Excavated - tree Pien 3447 115 53 0.53 38 x 36 Rock cavern Potr/Feth 3325 118 42 x 37 60 13.74 Relative Volume Estimator = width x height x length of all tunnels and chambers in the den, measured in cubic meters. This is not an index, merely an estimator. ,) N I-' m �Table 5. (continued - page 2) 20ens denoted by common letter wer-e used by the same bear. 3Quga Pien PHl Table = Gambel oak = Engelmann spruce = Limber pine 6. Den type Potr = Aspen Abla = Subalpine fir Feth = Thurber fescue Psme = Douglas fir Pied = Pinyon pine Prvi = chokecherry Jusc = Juniper Site characteristics of dens used by female black bears, 1980-84. , Overs tory Aspect Slope Elevation Relative vOlu~e' vegetati on (degrees) estimator (m ) (m) (%) - Rock cavern Excavated-blowdown y5 Rock caverna2 Rock caven1d N Rock caverna Y Rock cavernb N Excavated-treeb Y Rock cavernb N Rock cavernb Rock cavern Rock cavernc ock cavernc Rock cavernc Excavated-treed Excavated-treed Pien~Abco/ Abla Pien/Abla open cliff Feth Psme/Abla open cliff Potr open c1iff Feth open cliff Psme Quga Quga Psme Pien/Ab1a - 3065 7 78 20.2 3172 250 210 130 350 180 102 250 34 232 260 280 80 160 20 54 98 nd 66 66 2.12 2.52 1.42 1.08 2.91 1.61 2.09 7.5 32.8 0.95 0.31 1.53 3035 2959 2867 3279 3126 3065 3203 3279 2852 2501 2394 3056 3294 Den entrance (width x height; cm) 68 60 56 55 69 72 63 62 74 87 1.99 99 x 40 nd 33 104 123 83 47 93 72 41 120 54 87 50 23 x x x x x x x x x x x x x 63 30 33 31 26 31 33.5 33 27 42 30.5 29 57 N I-' --J �Table 6. (continued - page 2) N Rock cavernd Feth/shrubs 2989 220 65 2.86 57 x 27 Y Rock cavernc! Feth/Se ra 3111 10 67 10.13 168 x 29 N Excavated-shrub e4 Quga 2638 112 40 0.46 56 x 40 Amal 2593 350 12 1. 18 g'! x 40 Quga 2638 112 40 0.46 56 x 40 Pien/Abla 3447 230 63 0.86 42 x 28 f Co. vern' Psme/Pifl 3325 70 80 2.65 48 Y R.ockcavernf open c1 iff 3172 259 79 2.4i 110 x 35 N Excavated-shrub9 Quga/Amal 25'16 330 53 0.57 81 x 41 Y Excavated-shrub9 Quga 2364 355 63 1,69 56 x 38 N Excavated-shrub9 N Excavated-shrubg Prvi/Quga 244,0 310 26 0.83 76 x 61 Quga 2410 280 39 0.71 61 x 41 Y Excavated-shrub9 Rock cavernh Quga.Amal 2684 303 55 1.81 65 x 37 Quga 2943 130 105 14.73 81 x 43 Excavated-shrubh Prvi/Quga 2623 272 66 0.64 61 x 38 N Excavated-shrubh Quga/Prv; 2516 235 56 0.71 65 x 58 Rock caverni Talus 2730 112 75 2.31 58 x 35 N Excavated-shrub; Ama1/Quga 2315 352 50 0.78 56 x 34 . p N Excavated-shrubExcavated-shrube4 Excavated-tree N R'aCK f N I--' x 57 co �Table 6. (continued - page 3) Y Excavated-shrub Arnal/Quga 2440 355 66 0.81 70 x 61 N Excavated-treej Pien/Abla 3424 240 80 1.52 77 x 44 Y Rock cavernj Potr/Pi en 3325 160 51 1.60 79 x 41 N Excavated-treej Pi en/Ab 1a 3172 290 77 0.88 42 x 72 Y Rock caven,.] Pien/~\b1a 3203 360 60 5.34· 89 x 57 Exca vated-b 1owdown Potr/Psme 2745 20 69 1.46 35 x 40 Rock cavern Quga/Potr 2928 169 65 0,77 98 x 24 Y Rock cavern open 2806 140 78 8,70 50 x 29 cliff Y Rock cavern'I' N Rock cavernk Quga 2501 280 81 nd nd Quga 2410 240 78 1.64 34 x 30 N Rock cavernk Quga 2379 122 67 0.33 112 x 40 Excavated-b1owdown1 Pien 2760 60 75 0.97 70 x 48 Excavated-tree' Psme 3050 271 39 0.29 46 x 38 Excavated-b1owdown' Rock cavernm Ab1a/Psme 2989 286 32 0.71 67 x 40 Quga 2715 206 84 2.42 23 x 55 Excavat;on-blowdownm Psme 2493 220 51 0.84 48 x 23 Above surface bedn Rock cavernn Abco 2861 96 60 NA NA Psme 2867 240 80 2.17 85 x 30 N •...... \.0 �Table 6. (continued - page 4) N N 0 Rock caverno Quga/Talu.s 2364 175 68 1.86 55 x 30 Excavation-shrubo Quga 250'j 136 53 2.17 72 x ,~W Excavation-shrub Potr 2951 67 .- ,I .J 1 69 62 x 35 Excavation-tree Pien 3386 96 63 0.18 61 x 36 Rock cavern Quga 2371 183 6r-o 1. 25 84 x 35 ---_-- ,. " e 1 iRelative Volume Estimator = width x height x length of all tunnels and chambers in the den. measured in cubic meters. This is not an index. merely an estimator. 2Dens denoted 3p. ' 1 en Fatr Sara =: commor. letter were used by Engelmann Thurber spi"uce fescue = Red e1derber-ry Py'vi by ::: chokecherry Abco = whHe fi r Psme = Douglas fir f; ~Same den used 5y = by same bear. 1981 and 1984. yearlings in den, N the same bear. = natal den Amal = Serviceberry Abla = Subalpine fir Potr = Aspen Pifl = Quga = Limber pine Gambel oak �Colorado Division Research Wildlife 221 of Wi1dlife Report Ju1y 1984 JOB PROGRESS State of Colorado Project No. 45-01-503-15050 Work Plan No. 6 ----- Job. No. 8io Game Investiqations _____,,' Mountain - Noncervids Lion Investigations f~oL1ntain Lion Period Covered: Author: REPORT Population Dynamics 7/1/83-6/30/84 A. E. Anderson {!'BSTRt,CT Five puma we re captured and ra.diocol1ared in 111 days of hunting effort November 19, 1983. to May 17, 1984. Two of the 5 puma died a violent death; one was cannabalized and the other died from unknown causes but a predacide was suspected. As of June 29. 1984. 8 of 16 puma captured and radiocollared since the study began were being radiotracked on an approximate \lJeek1y basis. Twelve puma. V{el~e tracked with aerial telemetry during this fiscal year and te lenetri c locations subjectiveiy rated as 'good" totaled 347. Since the study began, 431 "good" te lemet.r i c locations have been obtained from 7 puma yearlong wh t ch ranged from 39 to 99 "good" locations for individual puma. The elevation of puma locations averaged highest r~ay throuch October as did distances between successive locations. Three adult female puma appeared to occupy adjacent. discrete home ranges yearlong while the home ranqes of t'I~IO other females were superimposed. The yearlong home range of a mature male extended over a 37 x 16 km rectangle and encompassed the home ranges of 4 radi oco llared females. ��223 ~10UNTAIN LION Allen POPULp.TION DYNAMICS E. Anderson P. N. OBJECTIVES 1. To assess the effects of sports hunting on mounta.in lion populations. SEGMENT OBJECTIVES 1. Capture and mark up to 12 mountain 2. Test accuracy 3. Monitor mountain of aerial telemetry lions. Iocat.i ons . lion movements. ACKNOWLEDGf,1ENTS I thank C. Ander's, B. Bellqardt , D. C. Bowden, G. Cheney, A. Cunningham, D. Coven, C. Dunn, M. Hershcop', D. Kattner, D. Kattner, Jr., J. Kattner, B. Kattner, D. Masden, N. McEwen, J. Olterman, S. Steinert. L. Stevens, and M. Stone for their assistance. I am also grateful to Doug Miller, Director. Institute for Wildlife Research, National Wildlife Federation for faci li tet inq fl nanc i al assistance from N ..W. r. METHODS AND MATERIALS Methods and materials are described in Anderson (1982) and Anderson (1983). One hundred eleven days were spent hunting from November 19, 1983, to f'1ay 17, 1984. Estimated aer-i a I telemetry location sites of each puma vlere plotted on U. S. G. S. Topographic County Maps (Scale 1 :50,000; contour interval 80 ft) or on U. S. G. S. Topographic Quadrangles (Scale 1 :24,000; contour tnterva l 20 ft ) "if l ocati on s ites were densely aggregated. Airline distances between successive 10cation sites were measured to the nearest 0.1 km. Contour intervals were read to the nearest 100 ft. (County Maps) or 20 ft (Duadranqtes}. RESULTS AND DISCUSSION Allocation of Hunting Effort To help insure that each puma within the study area had a similar probabi 1 i ty of capture , hunt; n9 effo -t was allocated proporti ona 1 to area among 4 strata (rab!::: 1 and Fig. 1). The discrepancy between the actual effort (Table 2) and the planned effort (Table 3) was due to; (1) relative acces sibi l ity among strata duri nq the preva i l i nq severe winter, (2) stratum 3 was hunted less because of the risk of recapturing the several radiocollared plm resident thereon. and (3) adverse weather precluded hunting more often than an~icipated. �224 Puma Capture a!!_c!_~~l~~t"C.Y_ Five puma were captured, radt oco l I ared i nd released (Table 4) for a capture ra te of one puma per 22 days of hunt i ng. Aeri a 1 and ground telemetry locations of 12 tndtvi dua l puma are listed in Appendix Tables A through L. Excluding t~ose locations of puma subjectively rated as "poor", 347 locations were judged us abl e for computing home range areas and p l ot tinq puma di s trl buti ons (Teb Ie 5). As of June 29, 1984s 7 puma tracked yearlong have atout 40 to 100 locations (Table 6); the minimal number foY' tests of normality of home range data (Gustafson and Fox 1983). Puma number 4 was inadvertently treed on December 14. 1983. and recaptured April 14, 1984, to rep lace its 27-month--old radiocol1ar. During that episode a hound bit her slightly on the head and one canine penetrated her upper, posterior left thi qh about 1 em while she was ar-tt a l ly immobilized. Because we were also attempting to capture and radiocol1ar her 2 juveniles, circumstances precIuded obtaining her body weight and measurements. Subadul t pumas 8 and B were recaptured on Augus t 4, 1983. and October 11, 1983, respectively. to adjust their radiocol1ars for growth. Puma MortB:_litr Of 8 male and B female puma captured, radiocollared, and released since April 15, 1981, only 8 were definitely alive on June 29. 1984 (Table 7). Pertinent de tai l s are summar-ized 'in Tab l e 8. The c i rcums tances of the deaths of pumas numbers 8. 14, and 16 are elaborated as follows: Puma number 8 died fol1owing recapture to adjust "its COnal" for growth. A 3 1/2,1'11" chase on a very hot day, the high dosage (about 6 cc of Ketamine: Rompun 6:1) requi red to immobilize, and poss ibl e heal th problems were all likely factors in "its death. D. Schweitzer, D.V.~1., at the ~Jestern Slope Animal Diagnostic Laboratory, Co-lorado State Un-iversity. Grand Junction, examined the skinned carcass within 24 hours of death. He found 18 adult tapeworms in the small intestine and diagnosed the ultimate cause of death as inhalation pneumonia and esophigitis. The badly decomposed carcass of puma number 14 was found wi th ground telemetry within 300 m of a sheep camp. The position of the carcass (hind legs crossed and extended posteriorly and front legs straight and perpendicular to the body) and dried blood on the canines suggested a violent death, perhaps from a predacide. Maggots precluded external examination but no lesions were found on the cleaned skull. The cannabalized remains (only the head, tail. stomach and legs were uneaten) of puma number 16 were found along Big Dominquez Creek wi th qround telemetry. Examination of the cleaned skull revealed 2 canine punctures; through the left eye orb i t wall to the top of the skull and through the muzzle and proximal LO the eye orbit. Very l a rqe puma tracks and what were presumably the victim's tracks were abundant on game trails in the v i c ini tv. Since capture on January 15, 1984, this young male puma had crossed the formidable Unaweep Canyon at least 6 times. ~_Conce~ua l_~.QPu 1ati on !!i29_~1. Keith (1983) presented a model for wolves which - have modified slightly for puma (Fig. 2). The model should be useful in guiding future population research on puma. �225 Activity and Movement Elevations of the locations of 5 individual puma overlapped between the November-April and May-October periods but their means were from 153.3 m to 295.7 m greater during the May-October period (Table 9). Mean airline distances traveled between successive locations were greater during the May-October period for all puma except number 4. That animal had a litter of cubs during July or August, 1983, which probably explains her apparent limited travel during that period. Distances traveled by male number 5 averaged somewhat greater than those of females yearlong. Distances between extreme locations of the same 5 puma provide indices of home range (Table 10). Preliminary work toward a comparative analysis of the various methods of calculating home range size included obtaining a computer program for the harmonic mean procedure. These exploratory analyses will begin once numbers of locations approximate at least 50 for most of the pumas listed in Table 6. When plotted on large scale maps, location sites of those puma with yearlong location, suggest that female numbers 6, 7, and 12 occupied adjacent and discrete home ranges whereas the home ranges of female numbers 3 and 4 were almost superimposed. The inferred home ranges of male puma number 5 overlapped those of female puma numbers 4, 6, 7. and 12 and appeared to be in a state of flux since capture, Accuracy of Aerial Telemetry Locations Because of insufficient funds, no work was accomplished on a test of the accuracy of telemetry locations. Approximate ly 4 temporary employees working over 2 summers would be necessary to complete a statistically adequate test. Growth in Radiocollared Puma Recapture of juvenile puma to refit the radiocollar affords an opportunity to document growth in weight and body measurements in the wild. So far, only males have been re-measured (Tab1e 11). Mule Deer Killed by Puma The location and age structure of mule deer and elk killed by puma (Table 12) had a relatively large proportion of mule deer fawns compared to other winters (Table 13). Almost half of the 33 mule deer found killed by puma 1980-84. were less than 18 months of age (Table 13). Seven mule deer carcasses were not covered but snow may have been used initially in some cases. Three carcasses were almost completely covered by oak Quercus gambelli; leaves. Needles, branches and litter of pinyon pine Pinus edulis covered a large portion of the remaining covered mule deer carcasses. One elk carcass was not covered but records were incomplete on the second. Discovery of a mule deer carcass led to the capture and radiocollaring of male puma number 18. �226 LITERATURE CITED Anderson, A. E. 1982. Mountain lion investigations--·mountain lion population dynamics. Colo. Div. VJildl. Res. Rep. July, Part 2: 143-159. 1983. Program narrative. Project No. 45-01-503-15050, Work Plan 6, Job. No.1. Noncervid section, Research Center, Colo. Div. Wildl. Fort Collins. l8pp + 2 Figs + 2 Appendices. Erickson, J. A., and W. G. Seliger. 1969. Efficient sectioning of incisors for estimating ages of mule deer. J. Wildl. Manage. 33: 384-388. Gustafson, K. A., and L. N. Fox. 1983. A comprehensive interactive program for calculating and plotting home range and distribution. Pages 297-317 in D. G. Pincock, ed. Proc. 4th Intl. Conf. Wildl. Btote l emetry. -- Keith. L. B. 1983. Pages 66-77 lr:!. L. N. Carbyn, ed. Wolves "in Canada and Alaska: their status, biology and ~anagement. Can. Wildl. Servo Rep. Series No. 45. Ottawa. 135pp. .l, H. Olterman , and D. c. Bowden. 1980. A helicopter quadrat census for mule deer on Uncompahgre Plateau. J. Wildl. Kufe ld, R. C., Manage. 44:632-639. Masden, O. 1982. Units 61 and 62 (DAU) deer quadrat census (Memorandum to J. Olterman of June 1). C00l4 Southwest Regional Office Files, Montrose, CO. (unpaged). Reeves, R. C., A. Anson, and o. Lander, eds. 1975. Manual of remote sensing. Amer. Soc. Photogrammetry. Falls Church, VA. 214pp. Prepared by OL6,Jf)::a,-A~ A 11 en E. Anderson ze. Wildlife Researcher �227 Table 1. Allocation of hunting effort for puma, 1983-84. Uncompahgre Plateau (GMU 62). Strata Strata2 Percent of Total number numbera area (km ) hunting days tota 1 area 1,2 418.3 24.8 31 3,4 578.9 34.3 44 596 356.8 21.1 27 7,8 334.1 19.8 1688. 1 100.0 25 127b aKufeld et al. (1980) and Masden (1982), see Fig. 1. bEstimated days likely to be actually hunted from Nov. 17, 1983, through May 17, 1984. Number of hunting days projected for each stratum obtained by multiplying 127 by percent of total area for each stratum. �228 <, "':<'I ~.WHITE'''lATER o--- 5 10 MILES 2 Fig. 1. Unit 62 midwinter deer census area showing 8 strata and O.6475-km quadrat allocation within strata. From Masden (1982). �Table 2. Chronology of puma hunting effort by strata, 1983-84, Uncompahgre Plateau (GMU 65). re~resents one da,l hunting effort. Strata and dates Month Nov. 1 19,20,21,23, 27,28,29 7 Dec. 2,3,4,5,6 5 Jan. 3,4,5,6,9,10, 11,12,15,16,17, 18,30,31 Feb. 14 5,6,7,8,9, 12,17 May 26,27,28,29 4 19,23 2 7 0 1,8,12,13,14, 15 s 19 , 20 , 21 0 0 3 Total 3,8,9,10,11,12; 13,17,18,24,26. 30 1,2,7,8,9,14,16 11,12,13,15 Grand total 0 7 14,19,21,22 4 17 0 28,29 2 18 1 22 8 19 5 18 13 3 11 Total 4 4,2,29 2,7 2 9 5 5,6,23,26,28, 29,30,31 4.7.14.16, 19 12 7 4 Total 0 0 1,2,15,16,18, 21 ,22,23,24~26, 28 flJar. Apr; 1 2 Total Each date 1 1 10 3,4 2 10 - - - - - 33 45 11 22 111 N N 1.0 �230 Table 3. Planned and actual allocation of hunting effort for puma, 1983-,84~ Uncom~ahgre Plateau (GMU 62). P'lanned Actual Strata Number days Percent of total Number days Percent of total 31 24-.'8 33 29.7 2 44 34.3 45 40.5 3 27 21.1 11 9.9 4 25 19.8 22 19.9 127 100.0 111 100.0 �Table 4. Detai1s on the capture and body measurements of 5 puma radioco 11ared, 1983-84, Uncompahgre Plateau (GMU 62). Ear tatoo number 14 Sex Data of capture Estimated age (months) Legal descr. capture site 1/4 5-rI R U.T.M. capture site X y M 12-12-83 84+ NW 9 48N llW 753 4257 2042 15 16 17 18 F M 1-15-84 30 M M 4-14-84 8 4-16-84 36 N~~ 12-15-83 60-72 SW SE 5W 8 13 48N 11W 26 145 lOOW 752 4256 2073 4297 2103 Elevation (m) capture site Drug (2:1 Ketamine-Rompum) Dosage (cc injected) 3+ 5 Induction time (min.) 6 128 Radiocollar serial no. 12906 12902 Transmitter frequency (mhz) 149.9310 149.8900 Measurements (em) Body wt (kg) 48.0 Total body length 218 196 Taill ength 82 73 Head-body length 136 123 Chest girth 83 70 Neck circumference 48 37 Height at shoulder 75 69 Head length 22.7a 20.1 Zyomatic breadth 16.5a 13.9 Ear length 8.8 8.7 Hind foot 30.0 28.0 a Measurements on cleaned skull were 22.0 em and 15.0 cm b . Measurements on cleaned skull were 20.0 em and 14.3 cm 71<1 47N lOW 18 47N 9W 244 4246 2245 245 4246 2147 3 4 20 12897 149.8310 44 12910 149.9710 55.0 190 71 119 70 38 71 b 23.0b 15.5 9.0 28.0 36.2 185 69 116 57 32 60 21.6 13.3 9.0 28.0 5.5 18 12903 149.9010 64.3 214 86 128 80 46 75 23.4 17.5 10.0 29.0 N w �232 Table 5. Numbers of telemetric locations of 12 puma subjectively rated as either "fa i or good 1983-84, Uncompah9I:e Plateau (GMU 61.62.40). I~II II II, �Table 6. Number of aerial radiolocations of mature puma where N > 40a• Uncompahgre Plateau. Number of radiolocations Period of surveillance Sex F F f~ F Ear tatoo No. 3 4b 5 6 F F -12 ~1 13 7 Date From Radiocol1ared 1- 8-82 1-2-1-82 2- 7-82 2-10-83 3- 4-83 3-25-83 4-13-84 1-15-82 2-15-82 2-10-82 2-17-83 3-10-83 4·-1-83 4-·15-83 To 4-15-83c 6-29-84 6-29-84 6-29-84 6-29-84 6-29-84 2-20-84 c Tota,d Usa b'Ied 44 111 69 69 70 67 48 40 99 63 65 67 58 39 j _. - 478 431 aN ~ 40 recommended by Gustafson and Fox (1983:303) for normality tests. bRadiocollar replaced on 4-14-84. cLast signal received on that date. dllTotal" includes locations subjectively rated: locations rated "fair" and "good." poor, fair, good and "Usable" only those N W W �234 Table 7. Sex and age class or 16 individual puma at capture, 4-15-81 to 4-16-84. Uncom~ahgre Plateau (GMU 62)a. 24+ months of age Less than 24 months of age Males Females 'Males Females 5 2 8 1 14 3 10 16 18 _A l3a 17 6 7 12 15 ~----", Total 4 7 4 aUnderlined ear tatoo numbers are those puma al i ve as of 6-29-84. blast signal heard 2-20-84. �Table 8. Sixteen ~uma ca~tured and ra?iocol1ared 4-16-81 to 4-16-84, Uncom~ahgre Plateau (GMU 62). Estimated age Radioco 11ar Date (months) Ear tatoo Number Frequency Statusb Sex at capture Number Collared Breeding status 8389 149.5500 4-16-81 1 8772 149.7010 1- 5-82 2 8775 149.8005 1- 8-82 3 12898 149.8400 1-21-82 4a 8774 i 2904 12909 149.7815 149.9090 149.9605 2- 7-83 2-10-83 3- 4-83 5 6 12901 149.8810 2-20-83 8 12900 149.8705 3-23-83 10 12911 149.9800 3-25-83 12 7 Disappeared after 8-12-82 Ki1'1ed by hunter 1-10-82 Disappeared after 4-15-83 Alive F 20-21 Immature F 60 Unknown F 26 Unknown F 24 Alive Alive Alive M F F 60+ 30 48+ Died during recapture (for collar adjustment) 8-4-83 Found dead of unknown cause(s} 6-2-83 Alive M 6 M 6 F 60+ With 2 young 40-55 1bs at recapture ~ , 2-14-83, mother of l7c Unknown With 2, 15-30 1b cubs at capture, mother of 8. On 3-14-84 and 5-9-84, tracks indicated another litter of 2 Appeared pregnant at recapture 2-26-84, mother of 13 and 1 other cub N W U1 �Table 8. (con_tinued) _ N W 0'1 12896 149.8200 4-13-83 13 Unknown. last signal on 2-20-84 M 10+ 12906 149.9310 12-12-83 14 Found dead 6-22-84. Possible predacide victim (see text) M 84+ 12902 149.8900 12-15-83 15 Alive F 60+ 12897 149.8310 i6 Found cannabalized M 36+ 12910 149.9710 M M 8 149.9010 17 18 Alive 12903 -'-15-84 4-14-84 4-16-84 aAnimal recaptured and radiocollar Alive 2 kittens 5-12 days of age on capture 36 replaced on 4-14-84. bSased on aerial telemetric locations as of 6-29-84. Radlocul l ars have mortal ty siqnal . "Df sappeared'' or "unknown" have 4- possibilities; (1) transmt t ter failure and al ive , (2) tr ansm t ter failure and dead, (3) animal moved out of area of extensive search. and (4) animal alive but occupying topography where radio signal had very limited s trenqth and r-ange. CRecaptures were inadvertent in attempts to capture their young. �237 Table 9. Seasonal changes in elevation and airline distances between successive locations among 5 adult puma radiotracked at approximate weekly intervals yearlong. 1983-84. Uncoml2ahgre Plateau (GMU 62). Ear tatoo number Season, a 12 1983-84 5 6 7a Statistic 4 F Sex Nov-Aprilb (snow) Elev N (m) x SO Min ~lax Distance (km) Elev (m) 25 2147.1 22 2059.2 107.7 170.1 1829 2499 157.8 1951 2591 157.1 1768 2469 151. 1 1768 2438 22 22 25 22 1920 2377 20 x 5.19 5.21 0.8 Min Max 22.5 N 24 SO Min Max Distance (km) F 22 2164.2 N x F 22 2093.6 - -- F 20 2130.6 SO May-Octobel~C M 6.25 4.95 0.2 13.6 2283.9 88.3 2073 24 2353.3 197.9 2012 2450 24 N - x SO Min Max 2.97 3.21 0.1 10.7 3.95 2.85 0.8 10.4 24 2.89 1.86 0.5 6.5 cSeason without appreciable snow. 8.7 2379.9 140.0 2012 25 2332.2 109.7 2103 24 2274.5 105.1 1951 2682 2560 2560 2530 24 24 7.61 4.83 0.5 18.5 5.16 3.55 0.0 13.5 25 5.32 3.28 1.8 13.3 aWith 2 or more dependent young during a portion of sampling period. bSeason with snow. 2.99 2.29 0.4 24 4.18 2.31 0.1 8.2 �238 Table 10. Extreme limits (km) of locations of radiocol1ared puma tracked yearlong2 ___"l983-84. Uncompahgre Plateau ._,_{_G~_1U_6_2-,-)_. _ Ear tatoo number and sex 4,F --- 5,M 6.F 7,F 12,F ..-----.---~.---------.---- No. locations 45 46 46 50 45 East-West (km) 20 37 14 21 18 North-South (km) Total at~eaa (km2) 20 16 20 17 13 400 592 280 357 234 alndex only - computed by treating each area as a rectangle; multiply by 0.386 to convert to square miles. �239 Table 11. Growth in 2 young male puma over approximate 6-month periods, 1983-84, Uncompahgre Plat~au (GMU 62). Ear tatoo number 13 Dates captured. Body Wt. (kg) Total body length (em) Tail length (em) Body length (em) Chest girth (em) Neek circumference (em) Height at shoulder (em) Head length (em) Zygomatic breadth (em) Ears length (em) Hind foot (em) Hind paw length (em) Estimated age (mos)a 8 2-20-83 26 4-13-83 10-11-83 36 52 188 206 145 79 109 79 127 53 92 178 69 109 61 76 51 61 36 39 28 34 71 41 64 H.8 21. 1 14.9 20.1 "'2.1 15.3 9.5 8.2 8.3 10.2 12.7 9.0 25.5 29 9.3 15 8-4-83 32 8.0 3.0 6 12 aBased on dental development criteria summarized in Anderson (1983). blower right temporary and permanent canines both present. �Table 12. Discovery Date 8- 4-83 12-12-83 12-15-83 Twelve mule deer and 2 elk killed by puma, 1983-B4,_Unco_m_2ahgre Plateau Sex Est. Age (months) Unk adult M 6 F 17 Legal description U.T.M. (GMU 62). elev. 1/4 NW NE NvJ 5 T R X (east) Y (north) Habitat (m) 16 49N 13vJ 732 legs, skeleton 49N llW 756 riparian riparian 2347 26 4266 4263 1951 head, hi de ~ 1 egs • 2073 skeleton head. hide. spine, 17 48N 1H~ 752 4255 riparian 1/3 (elk) 2- 9-84 F Body Parts Remaining Approx. 57-69a NW 4 15S 99W 720 Ll.')Qt:; .f-oJ..., r+par+an 1829 (elk) a 44·-56 NW 34 49N 13\~ 734 4270 P-J 2195 9 NE 26 50N 12t~ 745 4272 ri pari an riparia.n 1707 hindquarters head, hide, skeleton head, hide, skeleton head, skeleton, hide .head, hide~ skeleton 2-16-84 F 3- 1-84 3-21··84 1'1 F 9-10 SE 8 47N 9vJ 248 4247 3-21-84 F 9-10 NE 8 47N 9'1II, 248 sagebrush 2042 head, skeleton 3-31-84 r~ 9-10 NW 4 47N 9W 249 4248 4249 P-J 2012 All 4-16-84 F iO NW 7 47N 9\~ 246 4247 P-J chaining 2164 4-17-84 Unk Unk SW 1 49N 13W 737 4267 sagebrush 2225 2042 except viscera, 2/3 thigh muscle 1/5 thigh, head, hl de , skeleton lower 1egs, portion of spine9 stomach 4-2L'~-84 F adult NE 7 49N 12~~ 740 4267 P-J 2225 5- 3-84 F 17 NW 7 47N 9W 246 4247 P-J-oak 2103 F 5- 8-84 9 Tatum Ridge ali extracted for age estimates by counting P-J dental cementum annuli (Erickson and Seliger. Intact except for left shoulder and thigh Intact except for viscera, all ribs left side, most of left thigh head, hide, skeleton 1969). N ..j::> o �Table 13. Age class and sex of cervids killed by puma, 1980-84, Uncompahgre Plateau (GMU 62). 18+ mos < 12 mas 13-17 mos Fiscal tlj Sex unknown ~1 F F Sex unknown yeara Species F r'i 1980-81 1981-82 1982-83 1983-84 Total Total - --. mule deer elk. 0 0 1 0 0 0 2 0 0 3 0 0 0 0 0 0 0 0 mule deer elk 1 0 0 0 2 6 0 0 a -' 0 0 0 0 0 0 a 0 0 mule deer elk 0 1 'J 2 0 5 0 0 9 0 0 0 0 0 0 0 0 0 mule deer 3 4 0 1 0 2 1 1 12 elk 0 0 0 1 0 1 0 a 2 mule deer elk 4 6 1 2 15 1 1 33 0 0 0 3 1 0 1 0 0 2 aNear1y all kills found from December through May. N ~ •....• �242 PUfv1A DENSITIES '\ / Puma numerical resp nse Il\ Puma social behavior / r1 \ EXPLOITATION I Puma predation I~ a.Lter-nat e=" prey Puma dietary ~--response DEER-ELK DENSITIES AND VULNERABILITY ~--- ..... 7 snow conditions "\... other predators ~. ~and disease forage production Fig. 2. Conceptual model of puma population dynamics (modified from Fig. 4 in Keith [1983:73J). �AEEendix Table A. Aerial telemetr,l locations of adult female l2uma number 4. U.T.M.a Legal descr. Distance (km) Approx. between Date S 1/4 T R X Y e1ev. (m) locations 6-30-83 7- 9-83 7-16-83 7-25-83 7-26-83 8- 3-83 SE SE NE NW NW SW 32 32 33 26 8 8- 9-83 10 17 48 48 48 48 47 47 4250 4250 4251 4252 4247 4245 2256 2256 2103 2316 2256 1.7 0.5 10 239 762 240 242 762 762 SE 18 47 10 762 4245 2316 0.7 8-16-83 NW 17 47 10 762 4245 2256 0.7 8-24-83 18 47 iO 4246 2450 L5 8 47 47 10 760 762 762 4247 4247 2225 2256 5.2 0.1 47 10 4246 2256 3.8 5 47 10 762 762 4249 2316 2.8 5 10 239 761 2286 2377 17 47 10 10 4249 4246 0.5 7 47 47 17 47 10 762 4246 4246 2316 2316 1.6 10-21-83 NW NW NW NW NE NE SE NW NW 0.3 10-28-83 NE 18 47 10 761 4246 2316 1.3 11- 4-83 NW 8 47 10 762 4247 2316 1.8 8-30-83 9- 4-83 9-12-83 9-19-83 9-28-83 10- 8-83 10-14-83 8 17 to 10 10 10 10 762 1.9 3.9 1.7 3.2 RatingC Major drainageb Spring Spring Spring Spring Spring E. Fork Spring E. Fork Spring E. Fork Spring Spring Spring Spring Spring Spring Spring Spring Spring E. Fork Spring Ck. W. Fork Spring Ck. Spring S L G G G G G G d G G G G G G G ,.. I.:! G G G G G G G G G G F F F G G G G G G G G G N ~ W �Appendix Table A. (continued - ~age 2) 11-11-83 SW 23 48 10 242 4253 11-24-83 NW 20 47 10 762 4243 1920 2377 10.0 7.3 33 48 10 240 4251 2103 7.0 12- 9-83 NE NE 33 48 10 240 425i 2073 0.3 12-14-83 NE 32 12-16-83 12-24-83 12-30-83 SW 33 48 48 10 10 239 240 4252 4250 NE 10 10 240 239 4251 4253 2103 3.0 1- 6-84 NE 9 48 48 48 1.7 1.1 0.8 SltJ 33 21 2134 2225· 2·!34 11 754 4257 2073 10.0 1-13-84 NE 9 48 11 754 4257 - - 1-20-84 SE 8 48 11 752 4256 - - 1-25-84 2- 3-84 S~J 4 11 11 4258 4256 - 18 753 751 - SW 48 [f8 2225 4·.3 2-13-84 SE 16 48 11 754 4255 2195 3.4 2-18-84 SW 16 48 1"1 753 4255 2073 1.0 2-26-84 SW 9 48 11 753 4257 2103 2.6 3- 2-84 3- 9-84 SW NW 2 16 48 48 11 11 758 753 4259 3.9 4255 2042 2042 8.6 3-17-84 NE NW 9 48 11 754 4258 2012 5.2 9 48 11 754 4258 2073 0.8 11-30-83 3-23-84 Spring E. Fork Spring Creek Spring Spring Lindsay Spring Spring Linscott E. Fork Dry Creek E. Fork Dry Creek t~. Fork Dry Creek Dry Creek ltJ. Fork Dry Creek E. Fork Dry Creek E. Fork Dry Creek W. Fork Dry Creek Coal Creek E. Fork Dry Creek Dry Creek W. Fork Dry Creek G F G F G F G G N -P> -P> e G r G F G F G F F P P P G P r> \:l F G F G F G F G c I G G G G G F r �Appendix Table A. 4- 1-84 NE (continued - page 3) 21 48 11 754 E. Fork 4254 48 47 47 48 10 10 10 11 761 4254 244 245 754 4245 4246 2225 N\lJ 20 14 13 21 4254 2195 1.5 22,5 5- 4-84 NW 21 48 11 754 4254 2195 0.7 5-11-84 5-18-84 5-26-84 SW 48 47 10 4250 4250 4244 2256 2408 1,7 SE 240 238 238 10.7 47 10 10 2225 NE 33 5 20 5,8 6- 1-84 NW 29 47 10 238 4243 2438 O,g 4- 6-84 4-13-84 4-20-84 4-29-84 6- 8-84 6-15-84 6-22-84 6-29-84 SW SE NE 2103 8.7 NVJ 20 47 10 762 4245 2316 1.6 SE N~J NE 8 8 1 48 47 46 9 248 239 244 4247 4248 4239 2073 2286 10.7 10 10 9.5 Dry Creel< Linscott Happy Happy E. Fork Dry Creek E. Fork Dry Creek Spring Spring E. Fork Spring Creek G p G F F p F F F F G F G F G F G G p F F F F F G G G p E. Fork Spring Creek E. Fork Spring Creek Dolores Spring Dolores = Universal Transverse Mercator coordinates (Reeves et a1. 1975) to locate puma on individual km2. b E = East, Fk = Fork, W = West aU.T.M. CSubjective rating of signal strength (S) and accuracy of puma locations (L): G = good, F = fair~ P When locations are rated "poor", elevations and distances between successive locations are deleted. = Poor, dGround telemetry, poor location. eThe initial ground observation was of tracks in snow; animal subsequently treed. N -lOa c...n �A~~endix Table B. Aerial telemetr~ locations of male adult puma number 5. Legal descr. Distance (km) U.T.M. a Approx. between T c y Date 1/4 R X e1ev. (m) locations '"' I 6-30-83 7- 9-83 NW SE 7-16-83 13 19 48 50 NE 25 7-26-83 NW 8- 3-83 8- 9-83 8-16-83 8-24-83 8-30-83 9- 4-83 9-12-83 9-19-83 9-28-83 10- 8-83 10-14-83 13 736 729 4259 4273 2621 2134 15.0 14.8 50 13 728 4272 2377 3.0 20 50 13 730 4273 2103 2.8 SE 11 49 13 SE 11 49 13 SI>J 34 NW 19 NI·J 7 NE 30 50 49 48 50 13 "136 736 734 739 741 730 4266 4266 4·270 4264 4257 4272 2347 2256 2164 2225 2499 2012 8.0 0.5 4.1 734 4262 11.4- 729 740 736 723 ll261 2438 2621 2316 2438 2560 4265 4264 4266 2073 16.3 2195 2499 13.6 4264 2225 11.9 3 S~J 27 NW NE SE SE 31 SE 12 12 13 19 26 49 13 4·9 13 49 12 49 13 16 49 i4 49 49 12 12 10-21-83 10-28-83 11- 4-83 SE 18 20 NW 13 49 13 741 742 728 11-11-83 NE 19 49 12 740 4264 4261 4264 ---- -------------- RatingC Major drainageb Criswe'll Dry Fork Escalante Dry Fork Escalante Dry Fork Escalante Monitor Potter Cottonwood S L G F G G G G G G G G G r» G \:l 7 , Cr i swe l l G G c 7.4 Terrib1e Dry Fork Escalante Potter Monitor Moore Criswell E. Fork Escalante Criswe11 Traver Dry Fork Escalante Criswell G F G l:" G F G F G G G ,'\;1" G F G G G G G F G F •• I 18.5 5.4 11.4 4·,3 13.5 2.6 \) , N ..j:>. O'l �Appendix Table B. (continued - ~age 2) 11-24-83 SE 27 50 13 735 4271 11-30-83 NE 49 14 728 1 4269 12- 9-83 12-16-83 12-24-83 12-30-83 1- 6-84 1-20-84 1-25-84 SE NW SE NW SE NE NE 27 25 28 23 19 17 2- 3-84 6 50 49 49 49 50 49 49 13 13 13 13 12 12 11 735 737 734 735 739 741 749 4271 4262 4261 4264 4273 4266 4269 SW 31 50 11 748 2-13-84 2-18-84 rm 35 11 SE 9 49 48 2-26-84 NW 15 3- 2-84 3- 9-84 3-17-84 3-23-84 4- 1-84 4- 6-84 4-13-84 4-20-84 4-28-84 5- 4-84 NW NW NE NW 2 -3 11 3 2 1 6 3 15 24 m-J NE SW SE SE NW 2256 2195 8.5 6.3 2134 2347 0.2 9.4 - - 2316 1981 3.7 9.7 - - 1890 W.O 4270 1951 1.6 4262 4257 1890 2073 907 11 755 755 48 11 755 4256 2134 0.6 48 48 48 48 48 48 48 49 50 50 11 11 11 11 11 11 10 11 13 14 755 755 757 755 756 759 760 754 735 727 4262 4259 4258 4258 4259 4259 4258 4269 4274 4274 2103 1890 2134 2073 2103 1951 1951 1829 2134 2195 3.6 1.6 3.2 2.6 2.0 2.5 1.3 12.3 17.8 8.0 5.4 Cottonwood Dry Fork Escalante Cottonwood Cri swe 11 Potter Potter Monitor ~1oore Roatcap Gulch Roatcap Gulch Dry Creek E. Fork Dry Creek E. Fork Dry Creek Dry Creek Dry Creek Coal Creek Dry Creek Dry Creek Coal Creek Coal Creek Coal Bank Cottonwood Escalante G F G F G F G F P P F F G F P P F F G F G G G F G F G G F F F F F G F G F F F G F G G G G G N ~ "'"-J �N co """ Appendix Table B. (continued - eage 3) SW 35 50 14 726 5-11-84 4270 ----~----~ 2195 4 , 'I 5-18-84- NE 5 48 14 724 4258 2682 12.0 5-26-84 6- 1-84 6- 8-84 6-15-84 6-22-84 SW SE SE NE 18 49 13 8.7 49 13 - - 26 4 14 SE 10 49 48 49 14 14 4265 4268 4261 4259 2408 1 729 738 727 725 725 4266 2560 2682 2469 4.2 2.8 7.8 6-29-84 Nt~ 27 49 14 725 4262 2530 4.0 aU.T.M. E. Fork Escalante Dry Fork Escalante G G G G Cot tonwood F F Potter P P Cot tonwood F F Cottonwood E. Fork Escalante Dry Fork Escalante G G F F G I.:! i" = Universal Transverse Mercator coordinates (Reeves et al. 1975) to locate puma on individual km2. bE. Fork = East Fork CSubjective rating of signal strength and accuracy (L) of puma locations: G = good. F = fair. P = poor. ~Jhen locations are rated "poor", elevations and distances between successive locations are deleted, �A~~endix Table.C. Aerial telemetr~ locations of female adult ~uma number 6. Legal descr. Distance (km) U.T J!1..a Approx. between Date S T 1/4· R X Y elev. (m) locations 6-30-83 7- 9-83 7-16-83 7-26-83 8- 3-83 8- 9-83 8-16-83 8-24-83 8-30-83 9- 4-83 9-12-83 9-19-83 9-28-83 10- 8-83 10-14-83 10-21-83 10-28-83 11- 4-83 11-11-83 11-24-83 11-30-83 12- 9-83 12-16-83 NE NE SW NE SE SE NW SE SE SE NE SE SE SW NE NE SW SE NW SW SE NE NW 20 2 28 5 7 35 29 3 11 4 29 27 30 48 48 49 48 48 48 48 12 13 12 12 12 12 12 13 12 12 12 12 49 49 49 13 13 12 1 30 48 49 13 12 740 739 4258 4262 2408 2377 3.8 3.4 6 48 48 49 48 49 49 12 12 13 12 13 12 742 4258 4257 4263 4258 4261 4266 2316 4.9 2256 2377 2103 2591 2316 3.8 10.4 1.3 9.8 6.2 6 5 10 24 4 34 18 48 48 49 48 48 744 739 742 742 4254 4259 4262 4259 2652 2530 2012 2408 741 743 741 736 743 746 748 745 732 4258 4257 4257 4260 4252 4258 4257 4258 2377 2377 2499 2499 2469 2256 2256 4261 426·1 13.5 2.2 4262 2530 2438 2347 734 740 745 737 744 734 739 10.2 7.2 3.6 3.3 2.7 0.5 2.6 5.2 10.7 6.7 1.3 5.6 Ratingb Major drainageb Roubideau Traver Traver ~Jright Bull Long Terrible c-: swe 1"1 Roubideau Roubideau Cushman Bull Monitor Potter Moore Wri ght Criswe 11 Terrible Roubideau Criswell Bull Potter Potter S L G G G G G G G G G G G G G G P F P F G F P P G F G F P F F F G F F F G F G F G F P F G F G F r-, +: ....: �Appendix Table C. 12-24-83 12-30-83 1- 6-84 1-20-84 1-25-84 2- 3-84 2-13-84 2-18-84 2-26-84 3- 2-84 3- 9-84 3-17-84 3-23-84 4- 1-84 4- 6-84 4-13-84 4-20-8'l u,-28-84 5- 4-84 5-11-84 5-18-84 5-26-84 6- 1-84 6- 8-84 6-15-84 6-22-84 {continued - ~age 2) NE 11 49 NE 17 49 SE SW NW NE SW NE SE SW SW NW SE SW NW NE SE SE SE SE SW SE NW SW SE NW 31 13 12 736 741 4267 4266 2316 2073 2.8 5,3 50 49 12 740 4270 2073 4.5 12 742 4266 1951 4.5 49 50 49 12 12 'J 2 13 49 12 2195 2042 2134 2164 2134 0.8 1.4 6 49 12 4267 4271 4268 4267 4269 4268 2.0 3.2 49 - - 6 49 49 12 12 12 740 740 739 738 740 739 739 4268 2042 l.i 9 8 31 6 12 6 6 5 12 739 741 739 12 13 738 739 740 738 24 49 50 50 49 18 49 13 1 13 1 26 49 49 49 13 6 49 12 36 3 22 8 25 49 48 13 13 49 48 49 13 12 13 30 30 13 738 738 739 738 736 734 743 738 2.8 2073 2073 1.3 2.5 4.6 4264 1951 1951 2256 4265 - 4268 4268 2225 2225 4262 2438 4268 4260 4258 4263 2042 4269 4268 4272 4272 4256 4263 2469 2560 2438 2469 2316 0.5 9. 1 .4.0 0.0 6.8 6.8 8,2 3.1 5.4 11.8 8.2 Monitor Moore Potter Moore Potter Potter Potter Potter Potter Potter Potter Potter Criswell Monitor Monitor Cri swe 11 Cr; swe 11 Potter Potter Cri swe 11 Potter ~loore Criswell Potter G F G F G F P F G F G F G F G F G F P P P F r» u F G F G F G F G F F P G G G F G G F F F F F F P F Bull F F Criswell P F N U1 0 �Appendix Table C. 6-29-84 aU.T.M. = NW (continued - page 3) 24 49 13 738 4264 Universal Transverse Mercator coordinates 2377 (Reeves 1.3 et al. 1975) to locate p F puma on individual km2. Criswell bSubjective rating of signal strength and accuracy (L) of puma locations: G = good, F = fair, P = poor. Vlhen locations are rated "poor"; elevations and distances between successive locations are de l e ted , N ,_. tTl �~endix Date Table D. Aerial telemetr~ locations of adult female ~uma number 7. a Legal deser. Distance (km) U.T.M. Approx. between 1/4 S X Y T R elev. (m) locations 6-30-83 NE 32 50 14 722 4270 234·7 2.0 7- 9-83 NE 24 50 13 728 4474- 2347 8.0 7-16-83 SW 36 50 14 728 4269 - - 7-19-83 NE 34 50 14 725 4270 2286 13.0 7-20-83 NE 3 49 14 725 4268 - - 7-26-83 8- 3-83 NE 32 13 3 13 731 734 4270 4268 2377 S\,J 50 49 2256 7.3 3.5 8- 9-83 5 49 13 8-16-83 NW SW NE 50 49 13 8-24-83 34 9 14 731 734 723 4269 4269 4·267 2377 2103 2347 3.2 3.2 11.3 8-30-83 SW 11 49 14 726 4266 2lD8 2.8 9- 4-83 NE 20 49 14 722 4263 2560 4.6 9-12-83 SE 1 49 15 719 4268 2438 3.9 9-19-83 9-28-83 S~~ 50 50 13 14 732 727 4269 4269 2377 2377 13.3 SE 33 35 10- 8-83 SW 18 49 14 729 4265 2377 3.7 5.1 Major drainageb RatingC N (.J1 S L Middle Fork Escalante Dry Fork Escalante Dry Fork Escalante E. Fork Escalante E. Fort Escalante Cottonwood Little Monitor G F G G G P G G a G G G G Cot tonwood G G Cottonwood E. Fort Escalante Dry Fork Escalante E. Fork Escalante Middle Fork Escalante Cottonwood Dry Fork Escalante Cottonwood G G G F G G G F G G G F G G G G N �A~~endix Table.D. (continued - eage 2) 10-14-83 SE 9 49 13 733 4266 7 8 49 49 13 11- 4-83 SE SE SE 7 49 13 11-11-83 11-24-83 11-30-83 NW SW NE 17 7 17 49 10-21-83 10-28-83 NE 33 SW 27 NW 26 12-30-83 NE 13 1- 6-84 SW 14 1-20-84 NW 23 1-25-84 SW 7 2- 3-84 NW 25 2-10-84 SE 27 2-13-84 SW 26 2-18-84 NE 35 2-26-84 SE 24 3- 2-84 NW 26 3- 9-84 SW 26 3-14-84 SW 21 (with 2 kittens) 3-17-84 NE 17 12- 9-83 12-16-83 2-24-83 2316 3.7 Little Monitor F F Cot tonwood G G Cottonwood Dry Fork Escalante Cottonwood G F G F G F Cot tonwood G F Little Monitor Cottonwood Cottonwood Cottonwood Cottonwood Cottonwood Cottonwood Cottonwood Monitor Cottonwood Cottonwood Monitor Monitor Cottonwood Cottonwood Dry Fork Escalante Dry Fork Escalante G F G F G F G F G F G I::" , F F G F G G G F G G G F G F G F 730 730 729 4266 4266 2377 2408 3.2 1.8 4266 2469 1.9 49 49 13 13 13 731 729 732 4266 4266 4265 2408 2438 2408 1.5 1.4 2.5 50 13 733 4270 2316 50 13 13 734· 735 4271 2195 5.2 1.5 4272 2073 1.6 13 13 737 735 4276 4274 2073 2195 4.4 2.8 735 4274 2134 0.6 50 13 12 739 4276 2042 5.6 50 13 4272 2103 4.4 50 50 13 737 735 4271 2134 2.7 13 735 4271 2073 0.5 50 13 735 4271 2073 50 50 13 13 738 735 4273 4272 50 50 13 13 735 732 4271 4273 2164 2103 2073 - 1.6 2.2 2.6 0.9 - 50 13 731 4275 2042 5.5 50 50 50 50 13 G F a G F N (.J1 w �AQ~endix Table D. (continued - ~age 3) NE 3-23-84 17 50 13 731 4275 -- -- - 4- 1-84 SW 35 5'1 13 735 -------_---- 4280 --- - - -_- -_-- - -- -- ---- --- ----- 2103 0.5 1768 5.5 4- 6-84 NW 16 50 13 732 4275 1981 5.4 4-13-84 SE 19 50 13 730 4273 2073 3.2 4-20-84 SE 17 50 13 731 4274 2134 1.8 4-28-84- NE 22 50 14 725 4273 2103 6.5 5- 4-84 SW 35 50 14 726 4270 2225 3.8 5-11-84 SE 25 50 14 5-18-84 SW 35 50 14 5-26-84 NE 32 50 14 6- 1-84 SE 36 50 15 6- 8-84 SW 26 50 14 6-15-84 NW 14 49 14- 6-22-84 NE 2 49 14 6-29-84 NE 29 50 13 aU.T.M. = Universal Transverse bE Dry Fork Escalante Dry Fork Escalante Dry Fork Escalante Dry Fork Escalante Dry Fork Escalante E. Fork Escalante G F G F E. Fork N (Jl .p. G F P F G F G F G G Escalante u F Dry Fork 728 2.3 4271 2286 Escalante G G E. Fork 726 4270 2195 2.2 Escalante F 722 Middle Fork G 4270 2195 42 Escalante r~idd'lePark F P 718 4269 2438 3.5 Escalante F F 725 4271 E. Fork 2134 7.4 Escalante 724 Dry Fork 4265 2499 8.7 F F Escalante 726 4269 2256 E. Fork G G 3.4 Escalante F 731 4272 G 2316 5.8 Cottonwood Mercator Coordinates (Reeves et al. 1975) to locate puma on individual kmi. I" = East CSubjective rating of signal strength and accuracy (L) of puma locations: G = good, F = fair, P = poor. When locations are rated "poor", elevations and distances between successive locations are deleted. dGround telem~try - poor location. �Appendix Table E. Aerial telemetry locations of juvenile male puma number 8a, Legal descr. U.T.M.a Distance (km) Approx. between Date 1/4 S T R X Y elev. (m) locations 6-30-83 SW 28 50 14 722 4271 2103 7.7 7- 9-83 SW 35 50 14 726 4269 2286 4.3 7-16-83 SW 35 50 14 726 4269 2286 0.2 7-16-83 SW 36 50 14 727 4270 2286 1.1 7-19-83 SE 34 50 14 725 4270 2286 1.2 7-20-83 NE 3 49 14 725 4268 2195 1.8 7-26-83 8- 3-83 NE SW 6 3 49 49 13 13 730 734 4269 4268 2408 2256 5.7 4.5 8- 4-83* NW 16 49 13 734 4266 2377 3.2 Ratingd Major inaqe ? dra Middle Fork Escalante E. Fork Escalante E. Fork Escalante E. Fork Escalante E. Fork Escalante E. Fork Escalante Cottonwood Little Monitor Little Monitor S L G F G G G G G G G G e G G G G G G aDied following recapture and drug immobilization to refit rad'iocollar 8-4-83, see text. bU.T.M. cFk = Universal Transverse Mercator Coordinates (Reeves et al. 1975) to locate puma on individual km2. = Fork, E = East dSubjective rating of signal (S) strength and accuracy of puma locations (L): G = good, F = fair. P = poor. When locations are rated "poor", elevations and distances between successive locations are deleted. eGround telemetry, poor location. * Location by ground telemetry, verified by tracks. N U'1 U'1 �Appendix Table F. Aerial telemetr~ locations of adult female ~uma number 12. Lega 1 descr. U.T.~1.a Distance (km) Approx. between Date S 1/4 T R X Y elev. (m) locations 6-30-83 NW 22 50 7- 9-83 7-16-83 NE 14 50 SE 4 7-16-83 NW 7-26-83 8- 3-83 8- 9-83 8-16-83 SE 14 724 4273 2256 4.0 726 724 4275 4268 1951 6.0 49 14 14 - - 3 49 14 724 4269 - - 4 14 4277 4273 4·278 4271 2103 2438 2134 2134 3.3 14 723 722 725 722 5.6 8.2 N~~ 21 NE SE 3 29 50 50 50 50 8-24-83 SE 6 50 14 720 4277 2225 7.0 8-30-83 9- 4-83 NE 25 15 21 718 723 4272 4273 2438 2438 . 5.7 N~j 50 50 9-12-83 9-19-83 9-28-83 10- 8-83 10-14-83 10-21-83 NE 35 SE 16 50 50 15 14 NE 17 50 14 17 15 50 NW 21 50 50 14 14 14 4270 4274 4275 4275 4275 4273 2530 2377 2164 2195 2377 2438 6.4 7.3 2.0 NE NW 717 723 722 721 724 722 10-28-83 NE 28 50 14 723 4272 2286 1.2 11- 4-83 NW 19 50 14 719 4274 2286 4.6 14 14 14 4.1 4. ~ 0.1 2.2 2.1 RatingC Major dra i nageb S L G G G F G P G P G G G G G r> (' ~ G G F G G G G Haley Draw G G Kelso G G Ke1so Ke1so Escalante Middle Fork Escalante Middle Fork Escalante Kelso G G F F G F G G G F G F Eo Fork Escalante Escalante E. Fork Escalante E. Fork Escalante ICe1so Ke1so Escalante Middle Fork Escalante North Fork Escalante Ke1so Middle Fork Escalante \;l N (J1 Cf'\ �A~Qendix Table F._ (continued - eage 2) 29 50 14 721 11-11-83 NW 4272 -~ 2438 2.9 11-24-83 11-30-83 SW NW 13 8 50 50 14 13 727 731 4274 4277 2286 6.3 - - 12- 9-83 SW 28 51 14 722 4281 - - Middle Fork Escalante Escalante Dry Fork Escalante Palmer G G G G F P P P G F G F G F G F G F F F G P G F G F G F G F G G G F G 12-16-83 12-24-83 12-30-83 NvJ 28 6 NE 1- 6-84 SE 4 51 50 50 13 13 13 733 729 733 4281 4279 4278 1829 2195 2062 8.7 4.8 4.4 SE 4 50 13 733 4278 1951 0.9 1-20-84 SW 26 51 13 735 4281 2012 4.2 1-25-84 NE 4 50 13 733 4279 2012 4.6 2- 3-84 SW 35 51 13 735 4280 - - 2-10-84 SE 34 51 13 735 4279 2012 0.5 2-13-84 SW 35 51 13 735 4279 1768 0.4 2-18-84 SE 27 51 13 734 4281 1951 1.5 2-26-84 NW 3 50 13 733 4278 2042 2.9 3- 2-84 SE 33 51 13 733 4279 2042 1.0 3- 9-84 SE 27 51 13 735 4281 1951 2.3 Tatum Escalante Dry Fork Escalante Dry FOI~k Escalante Dry Fork Escalante Dry Fork Escalante Dry Fork Escalante Dry Fork Escalante Dry Fork Escalante Dry Fork Escalante Dry Fork Escalante Dry Fork Escalante Dry Fork Escalante N (J1 '--J �A~~endix Table F. (continued - ~age 3) 3-17-84 NE 4 50 13 733 4278 2073 3 .<-? 3-23-84 NW 4" 50 13 732 4279 2103 Ll 4- 1-84 S\rJ 4 50 13 733 4278 2042 1 4- 6-84 NE 5 50 13 731 4278 2164 1.6 4-13-84 S~i 4 50 ' !" 732 4278 2103 1.2 4-20-84 N~l 4 50 13 732 4279 2056 0,9 .., i , 1 j Dry Fork Escalante Dry Fork Escalante Dry Fork Escalante Dry Fork Escalante Dry Fork Escaiante Di~Y Fork G F G F G F G F G G G F N Escalante 4-28-84 SE 11 50 14 726 4276 1920 5.? E. Fork F ,... r G F G G G F G F P F r::: , F P P G F G G 5- 4-84 NE 29 50 14 721 4272 2377 504 Escalante ~~iddie Foy'k Escalante 5-11-84 S~~ 32 50 14 720 4269 2256 3,4 Mi ddle Fork Escalante 5-18-84 NvJ 33 50 14 722 4270 2134 2 t! "j 5-26-84 NL~ 29 50 "14 721 4272 2i~38 1,8 6- 1-84 NE 12 50 15 719 4276 2316 5.2 IVli ddle Park Esca1ante Middle Fork Escalante Nor-th Fork Esca l ante 6- 8-84 6-15-84 6-22-84 SE NE SE 6-29-84 SE 9 50 14 723 4276 2073 4.6 23 50 14 727 50 H "720 2377 - 31 4274 4270 7.5 31 50 14 720 4269 2134 0.7 aU.T.M. : Universal Transverse Mercator Coordinates Kelso Escalante Middle Fork Escalante r~iddle Fork Escalante (Reeves et al. 1975) to locate puma on individual km2. ()1 co �4) Append; _x__T?_tD~__F ._______illntil}__u~d__-_j)_a_ge. bE = East CSubjective rating of signal (S) strength and accuracy of puma locations (L): G = good, F = fair, P When locations are rated "poor", elevations and distances between successive locations are deleted. = poor. N U"I 1.0 �App_endix Table G. Aerial telemetry locations U.T ,~1.a Legal descr. Date 1/4 6-30-83 NE 7- 9·-83 7-11-83 7-16-83 d Si~ 7-16-83d SW 7-26-83 8- 3-83 8- 9-83 8-16-83 8-24-83 8-30-83 NE 9- 2-83 9- 4-83 9-12-83 9-19-83 9-28-83 10- 8-83 10-10-83 10-14-83 10-21-83 10-28-83 11- 4-83 NE SE SE Nt~ NltJ SE NE SE SE SE SE SE SI-J SW NE SW S T 27 50 R X Y of subadult male puma number 13. Distance (km) Approx. between elev. (m) locations RatingC Major b drainage S L G G Kelso 4272 2347 4.5 715 No signal - thorough coverage of Escalante Canyon complex At about 2200 intermittent signa1s from vicinHy Short Points Upper North Fork~ Escalante North Fork G 26 50 16 706 4271 2621 8.8 Escalante rv II G North Fork 26 50 16 4271 2621 0.0 706 Escalante Q 7 B'j ue G v. , G 12 50 4276 2499 17 700 7 L1;275 \J G Blue 50 16 2652 1.7 701 G G 6 50 Blue 16 701 4275 2652 1.2 r» \,2 G 8 50 Blue 16 701 4276 2530 100 rl~ G 8 50 Blue 16 701 4276 2621 0.6 G Blue G 8 50 702 16 4276 2806 1.0 Fe ? ,-,IBlue F 12 50 17 700 4276 2499 G F 1 50 Biue 17 700 4·278 2652 1.3 F G 18 51 691 Calamity 17 4283 2286 10.3 F 23 Indian G 51 17 698 4281 2682 6.9 r= , 33 15 102 Calamity G 693 4285 2438 6.8 \;j 51 Calamity G 17 17 693 4283 2469 1.8 G G 17 51 Calamity 17 693 4283 2469 <0. 1 P 23 Indian 51 696 4281 F 17 F 12 F 50 17 699 4275 2438 9.8 Blue 24 G F 51 689 Calamity 18 4282 2286 11.4 F 33 G 51 Indian 694 17 4279 2225 5.5 15 f' i~ f' ! f' N 0) 0 �A~~endix Table G. (continued - ~age 2) NW 26 51 18 687 4280 11-11-83 SE 28 51 17 694 11-24-83 4280 11··30-83 SE 4 50 17 695 4276 12- 9-83 SW 11 50 17 697 4275 SW 15 50 18 686 12-16-83 4274 12-24-83 NW 2 49 18 687 12-30-83 NE 14 50 19 678 1- 6-84 SE 14 50 19 678 1-"13-84 1-20-84 1-25-84 2- 3-84 2-20-84 NE SW aU.T.M. bJ = G F r~averick c G Cow Creek Blue G F Blue F F P Little G Maverick 4268 G F 1720 12.3 Ca 1amityBlue J. 4274 G F 1798 11.1 Cottonwood DoloresR.. J. 4273 1.2 Cottonwood . G G 1768 South Side 4289 Dolores River G F 1890 6. 1 4280 2164 3.2 Maverick G G 4279 2042 2.6 Maverick G F 4277 Maverick G P Normal but faint signal heard (1/2-1 mi) vicinity upper Maverick about 2134 2347 2499 2499 6.9 6.9 3.3 2.4 I 36 51 19 680 29 51 18 683 NE 33 51 18 685 SE 5 50 18 683 Extensive night search. 1900 Universal Transverse Mercator coordinates (Reeves et al. 1975) to locate puma on individual km2. = Junction CSubjective rating of signal (S) strength and accuracy of puma locations (L): G = good~ F = fair, P = poor. When locations are rated IIpoorll,elevations and distances between successive locations are deleted. dSignals at 0800 and 2000, respectively. eGround telemetry, triangulated location. N ,_. m �ApRendi_x_J_ab1e H. Aerial telemetrl: locations Legal descr. U.T.M.a Date 1/4 S 12-12-83 12-16-83 Nt~ 9 NW 33 12-2L}-83 S~-J 19 12-30-83 1- 13-84· 1-20-84 1-25-84 N~J 20 2- 3-84 2-13-84 2-18-84 2-26-843- 2-84 3- 9-84 3-17-843-23-84 SE T 48 48 R 11 10 9 X Y of adult male j2uma number 14. Approx. e1ev. (m) 753 240 4257 4251 1951 2042 4243 4253 4268 2073 1951 4270 4255 2195 Distance (km) between locations 11.9 - RatingC Major b drainage West-Fork Dry Creek Spring P F F G F Cottonwood E. Fork Dry Creek P P F F Moore Potter Potter Potter Potter Potter Potter E. Fork Dry Creek G F G G G F G F G F G G F F G F F F G F G G G F F F P F 34 16 48 11 8 32 49 31 SE NW 31 31 32 50 49 50 NE 9 4-8 741 740 740 740 740 740 740 755 4266 4271 4270 4268 4270 4270 4271 4257 2134 2012 2103 2225 2073 2073 1951 2073 17.5 SE SW 12 12 12 12 12 12 12 11 4- 1-84 4·- 6-84 4-13-84 NW 30 4 16 50 48 48 12 11 11 739 754 755 4273 4258 4255 1951 1890 2225 22.5 20.9 3.2 Monitor Dry Creek E. Fork Dry Creek 4-20-84 4-29-84 5- 4-84 SE SE SW 22 31 16 50 50 48 13 12 11 734 739 754 4273 4270 4255 2196 2164 2073 26.5 Cottonwood Potter E. Fork Dry Creek SE SE SE NW SE SE SE 4 5 50 50 50 28.7 4.4. 0.5 1.9 2.3 0.2 0.5 20.0 5.7 20.7 (captured) P Dolores Linscott Roubideau N (j) N G 246 238 743 734 755 3.5 24.6 L G 47 48 49 50 10 12 13 S �Ap~endix Table H. (continued - eage 2) SE 2 5-11-84 48 12 748 4258 5-18-84- S~J 30 49 12 739 4261 24 48 12 749 6- 1-84 SE 4253 6- 8-84 6-15-84 NE SE 6-22-84 NW 11 20 48 48 20 48 11 9 11 = Universal Transverse bW = West, E = East aU.T.M. 2316 2499 2377 Cushman G 11. 1 Moore P F 12.8 W. Fork P F Dry Creek 754 4258 1951 6.4 Dry Creek P F 753 4253 2377 P 4.9 E. Fork F Dry Creek d 752 4254 2316 1.2 E. Fork Dr,:tCreek Mercator Coordinates (Reeves et al. 1975) to locate puma on individual km2. 6.3 CSubjective rating of signal (S) strength and accuracy of puma locations (L): G = good. F = fair. P ~Jhen locations are rated "poor" elevations and distances between successive locations are deleted. (' \) = poor. dGround telemetry location of carcass after receiving mortality signal from the air. N C1'I W �A~~endix Table I. Aerial te1emetrt locations of adult female ~uma number 15. Legal descr. Distance (km) Approx. between Date S T X Y locations 1/4 R elev. (m) 12-15-83 SW 8 48 11 752 4256 2073 - 12-16-83 SW 8 48 11 752 4257 - - 12-24-83 12-30-83 1- 6-84 N'l .v, 48 11 11 1829 8.2 NE 17 48 11 4260 4264 4256 - 49 752 755 753 - SE 5 23 2164 8.1 1-20-84 SvJ 9 48 11 754 4257 - - 1-25-84 SE 21 48 11 755 4254 2195 2.8 2- 3-84 2-13-84 2-18-84 SE 3 11 4 11 SE 9 11 755 754 754 4259 4258 4256 2012 1890 2012 6.5 SE 48 48 48 2-26-84 SE 9 48 11 755 4·257 2073 0.5 3- 2-84 SE 9 48 11 755 4256 2073 0.3 3- 9-84 NW 9 48 11 753 4257 2073 1.3 3-17-84 3-23-84 SE 10 11 NW 9 48 48 756 754 4256 4257 2225 2042 2.6 2.3 4- 1-84 4- 6-84 SW 4 11 NE 21 48 48 754 754 4258 4254 2073 2134 1.3 3.0 11 11 1.6 1.1 RatingC Major drainage b W. Fork Dry Creek W. Fork Dry Creek Piney Dry Piney J. E. Fork Dry Creek E. Fork Dry Creek E. Fork Dry Creek Dry Creek Dry Creek E. Fork Dry Creek E. Fork Dry Creek E. Fork Dry Creek tv, Fork Dry Creek Coal Creek E. Fork Dry Creek Dry Creek E. Fork Dry Creek S L (captured) P P F P G G G F P P F ,... r G F G G F F G F G F G F G F G F G F G F N O'l .+::> �A~~endix Table I. (continued - ~age 2) SE 2 48 12 4258 4-20-84 748 SW 15 48 11 4255 4-29-84 755 2316 2225 7.5 5.8 5- 4-84- NE 34 48 11 756 4251 2377 4.4 5-11-84 SW 22 48 11 755 4254 2195 4.3 5-18-84 SE 20 48 11 753 4253 2256 3.5 5-26-84 NE 19 48 11 750 4254 2347 1.7 6- 1-84 SW 28 48 11 754 4251 2438 3.6 6- 8-84 NW 27 47 11 755 4252 2316 2.3 6-15-84 6-22-84 NW SE 23 20 48 48 11 11 757 753 4254 4253 2256 2286 2.5 4.2 6-29-84 NE 21 48 11 755 4254 2073 2.0 Cushman E. Fork Dry Fork E. Fork Dry Creek E. Fork Dry Creek E. Fork Dry Creek W. Fork Dry Creek E. Fork Dry Creek E. Fork Dry Creek Coal Creek E. Fork Dry Creek E. Fork Dry Creek G G G G F F G F G F G F F F F F P F G F G F aU.T.M. - Universal Transverse Mercator Coordinates (Reeves et al. 1975) to locate puma on individual km2. bW = West, J = Junction, E = East CSubjective rating of signal (S) strength and accuracy of puma locations (L): G = good, F = fair, P When locations are rated "poor", elevations and distances between successive locations are deleted. = poor. N o(J1 �Ap_P~I1~ti_)(__T_abJE:_;J. __ P._eY'ialtelemetry locations of adult male puma number 16. Legal descr. U.T.M.a Distance (km) Approx. between Date 1/4 S T R X Y elev. (m) locations l4S 10m~ 714 15S 99t~ 718 . b Ratlng Major drainage S L Gtbbler G. (captured) o 1-20-84 N~~ Big Dominquez u 5.8 G F 1-25-84 NE 33 14S 99W 720 Big Dominquez G F 1920 4296 4.0 2- 3-84 SE 16 135 lOOW 710 2012 Bangs G 4310 F 17. 1 Canyon 2-10-84 SE 36 14S 100W 715 16,2 2256 Gibbler G 4296 F 2- 13-84 S~! 24 14S 100W 715 Gibbler G 4299 G 2134 3.3 2·- 18-84 St~ 5 13S lOOW 708 15,5 4313 Rough G G 2103 Canyon 2-26-84 SltJ 32 14·S 99t~ 718 15,5 4296 Big Dominquez G 2225 F 3- 2,-84 NE 5 15S 99W 719 4295 F 1981 1.0 Big DominQ4ez G 3- 9-84 N~~ 21 13S lOOW 710 4309 2012 Bangs G F 16.8 Canyon 3-17-84 N~J 26 12S 10HJ 703 6,1 F 4317 2073 Rough G Canyon 3-23-84 NE 5 15S 99W 719 4295 1890 27.0 Big Dominquez G F 4- 1-84 NE 6 15S 99W 717 4295 2164 1.6 Big Dominquez G FC d 4- 2-8~ SE 5 155 99W 718 4294 1903 _ 1.6 Bi9 Dom; nguez G aU.T.M. = Universal Transverse Mercator Coordinates (Reeves et a1. 1975) to locate puma on individual km2. bSubjective rating of signal (S) strength and accuracy of puma locations (L): G = good~ F = fair~ P = poor. ltJhenlocations are rated poor s elevations and distances between successive locations are deleted. 1-15-84 SE 26 II 4297 4293 2103 2073 I! CMortality signal late PM. dGround telemetry location of cannaba1ized carcass, see text. N O'l Q"l �A~eendix Table K. Aerial telemetr~ locations of juvenile male Quma number 17. a Legal descr. U.T.M. Distance (km) Approx. between Date S T R X Y elev. (m) 1/4 locations 4-14-84 4-20-84 4-29-84 SW 13 17 47N lOW 47 10 48 11 SE 12 SW 5- 4-84 NE 28 48 5-11-84 5-18-84 5-26-84 SE NE SE 33 6- 1-84 244 - 245 752 4246 4247 4255 2245 2196 1 .7 - - 11 755 4252 2316 15.4 5 20 48 47 47 10 10 10 240 238 238 4250 4250 4244 2103 2256 2408 10,8 3.8 6.0 SW 29 47 10 238 4242 2469 1.8 6- 8-84 NW 20 47 10 762 4244 2347 2.6 6-15-84 6-22-84 6-29-84 NE NW NW 8 48 47 47 9 10 10 249 243 242 4247 4245 4245 2042 2347 2377 23 23 aU.T.M. = Universal Transverse Mercator Coordinates bE RatingC Major b drainage Happy Happy E. Fork Dry Creek E. Fork Dry Creek Spring Spring E. Fork Spri ng E. Fork Spring E. Fork Spring Dolores Happy Happy S ! L. (captured) G F F P G F G F G G F F F F G G F F 6.2 F 1.4 (Reeves et al. 1975) to locate puma on individual km2. 11. 6 P G G = East CSubjective rating of signal (S) strength and accuracy of puma locations (L): G = good F = Fair, P Hhen locations are rated "poor", elevations and distances between successive locations are deleted. 9 = poor. N 0'\ '-l �Appendi x _IAb_l_e_L. Aerial telemetrt locations of adult male euma number 18. Legal descr. U.T.~1.a Distance (km) Approx. between Date S T R X Y elev. (m) locations 1/4 9W 19 47N 47 N~'J 17 47 9 SE 13 47 10 SW 33 9 10 27 47 47 47 10 46 10 14- 47 9 245 246 247 245 249 241 250 240 252 4-16-84 4-20-84 4-29-84 5- 4-84 NW 18 Nl~ 5-11-84 5-18-84 mJ 5-26-84 N~~ 6- 1-8Ll- N~j 6- 8-84- SW 27 9 9 6-15-84 NE 8 48 9 248 6-22-84 NE NW i3 47 13 47 10 10 244 244 6-29-84 aU.T.M. = 4246 4244 4246 4245 4240 4242 4243 4238 4245 4247 4246 4246 2147 2256 2134 2195 2225 2438 2195 - Rati ngb Major drainage 2.3 2.4 2.9 Happy Canyon Dolores Dolores Happy 6.2 8.3 9,5 S E:' ~ L (captured) G G F F F F Hcrsef ly G G Happy Horsefly G F G F 2621 11.0 Happy P F 2073 2073 2225 2225 11 .4· 5.1 Horsefly Dolores Happy Happy G F P F G F G t:" 4.2 0.2 N I Universal Transverse Mercator Cordinates (Reeves et al. 1975) to locate puma on individual km2. bSubjective rating of signal (S) strength and accuracy of puma locations (L): G = good, F = fair, P When locations are rated "poor". elevations and distances between successive locations are deleted. = poor. N CJ) (X) � https://cpw.cvlcollections.org/files/original/3930acb34c51ca5ad34bfed1bdd422f9.pdf 7d5581341a3b90d1b824cf8f2128efc6 PDF Text Text Colorado Division Wildlife Research September 1984 uf Wildlife Report JOB FINAL REPORT State of Project Colorado ----------------------------- Work Plan: Job: Job Title: Evaluation Period Covered: Author: Personnel: Wildl ife Law Enforcement of Law Enforcement Research Activities 1 July 1982 through 30 April 1984 John F. Smeltzer Janet E. Black, Clait E. Braun, Dave Croonquist, Richard M. Hopper, Donald L. Horak, Bob Le~sure, Kris Moser, Area Wildl ife Managers, District Wildlife Managers, Colorado Division of Wildlife. ABSTRACT An evaluation of the daily activities of District Wildl ife Managers (DWM's) and Area Wildlife Managers (AWM's) of the Colorado Division of Wildlife was conducted from 1 July 1982 through 30 June 1983 as the initial phase in a newly created and ongoing wildlife law enforcement research program. The 2nd segment of the study (1 July 1983 - 30 April 1984) focused on wi ldlife law enforcement information exchange, reporting forms, and wildl ife law enforcement Line/Staff relationships within the Division. Daily activity reports (DAR's) were collected and quantifiable data were totaled for each DWM. Four activities: wildl ife inventory, wildl ife law enforcement, public relations, and administrative and clerical services accounted for as much as 88% of a DWM's work time per month. Mean calculated values were: 201.5 hours worked per month, 23.48 days worked per month, 5.6 days off per month. AWM's spent most of their time in budget preparations, meetings, distribution of equipment and suppl ies, employee evaluation, and report preparation. There is no line authority between the Chief of Law Enforcement and the DWM-AWM-RLEC (Regional Law Enforcement Coordinator) positions which serve as the core of wildlife law enforcement in Colorado. The RLECis are responsible for planning, implementation, monitoring, and coordln~tion of a wide range of special ized wildl ife law enforcement tasks at the Regional level. A Iist of wildlife law enforcement activities conducted by the Division of Wildlife was developed. AI I Regions recognized each activity but variation existed between and within Regions in regard to wildlife law enforcement priorities and time expended on wi Idlife law enforcement. Data collection and reporting forms used for wildlife law enforcement purposes were collected from each �2 Reg~ons and compared for standardization. Only 25% of the forms currently used (15 of 60) were considered to be non-standard. Routes of information exchange were identified and classified as formal or informal. Line/Staff relationships within the Division were diagrammed. A review of wildlife law enforcement literature demonstrated that wildlife forensics is an accepted research area, non-forensic research is slowly evolving as a science, and current literature is predominantly descriptive or philosophical and not truly of a scientific nature. Weaknesses are identified and recommendations included. �3 RECOMMENDATIONS 1. Continued use of the present Daily Activity Reporting system (DAR) is recommended with reservation. The current DAR is inadequate as a tool to collect data for evaluation purposes. A new system should be developed but it should be entered into slowly and planned specifically to target job classifications and not generically developed. Wildlife law enforcement should be an integral part of the reporting process. A revised DAR is needed before substantive statewide evaluations of the wildlife law enforcement effort can begin. 2. Quantifiable wildlife law enforcement objectives must be a priority effort involving Denver Law Enforcement staff, Regional Law Enforcement Coordinators, and Wildlife Law Enforcement Research. 3. Standardized reporting should be required for all wildl ife law enforcement information. Forms should be recommended and developed by Denver Wildl ife Law Enforcement staff. 4. An interactive computer system should be available in each Region for entry, analysis, and print-out of wildlife law enforcement data. This system would enhance officer's safety, streamline record storage, simplify data retrieval, and allow cross-comparison of files between and within Regions. 5. District Wildlife Managers should become one of the primary target groups for feedback of information. DWM's serve as the primary nonresearch data collectors within the Division and may become disenchanted when information they provide '~isappears" for all practical purposes. Reports should be uncluttered and summaries should be concise. 6. Regional Law Enforcement Coordinators should be given Line authority over 1-4 new enforcement specialists per Region. These individuals would do primarily wildlife law enforcement work. ��5 EVALUATION OF LAW ENFORCEMENT ACTIVITIES John F. Smeltzer Enforcement of wildlife laws is an integral part of successful wildlife management programs. Increased demands on wildlife resources from increasing numbers of recreationists (hunters, fishers, animal watchers, etc.) and decreasing amounts of quality habitat make it imperative that wildlife populations be properly managed for the benefit of all people~ With increasing numbers of people interested in wildl ife, more leisure time, and sophisticated equipment, wildlife managers find it difficult to efficiently enforce wildlife regulations over large areas. Wildlife law enforcement research is relatively new although substantial research has been done on wildlife forensics,particularly the identification of cause and time of death of a variety of animals, and species identification. Additional research on wildlife law enforcement problems is needed to measure and improve the efficiency and effectiveness of wildlife managers and to provide tools for use in the field. However, before wildlife law enforcement research needs can be clearly identified, it is necessary to evaluate present wildlife law enforcement activities so that research efforts can be properly directed. P.N. OBJECTIVES 1. Examine and evaluate available wildlife managers in Colorado. data on law enforcement 2. Prepare a study plan on an appropriate research topic. wildlife activities of law enforcement SEGMENT OBJECTIVES 1. Examine available data on law enforcement Area Wildlife Managers. 2. Evaluate available data on law enforcement Area Wildlife Managers. 3. Interview selected personnel of the Colorado concerning law enforcement research needs. 4. Review available literature on wildlife past research and management efforts. 5. Prepare a study plan on an appropriate ment research topic. activities of District activities Division of District and of Wildlife law enforcement approved and wildlife problems and law enforce- �6 6a. Develop a comprehensive list of all wil~life of the Colorado Division of Wildlife. 6b. Organize 6c. Analyze the written reporting area, regional, and statewide 7a. Identify present routes and mechanisms for information the district, area, regional, and statewide levels. 7b. Analyze the present mechanisms and routes for information exchange at the district, area, regional, and statewide levels. 7c. Develop flow charts depicting the mechanisms and routes for information exchange at the district, area, regional, and statewide levels. 8a. Identify apparent strengths and weaknesses enforcement systems. 8b. Suggest possible alternatives system. 9. Describe the control mechanism for major wildlife law enforcement activities, including line authority, reporting responsibilities, and staff support responsibilities. and list those activities 10.. Assist in the development and formats statewide. 11. Compile data, analyze law enforcement activities by region. tools for each activity at the district, levels. in the wildl ife law to strengthen of standardized exchange at weaknesses information within the reporting forms results, and prepare progress or completion reports. �METHODS Daily Activity Reports Daily activity reports (DAR's) were collected from 12 of the 16 Division of Wildlife administrative areas. All 4 Regions were represented. The sample consisted of 514 months of activities by 32 District Wildlife Managers (DWM's, 30% of total) from January 1979 through June 1982. Total months evaluated for any individual DWM ranged from 4 to 28. Quantitative data, including days and hours worked per month, days leave. taken per month, and days off per month were totaled for each DWM. The information was used to calculate ranges and means by month for the following: days worked, hours worked, days leave taken, and days off. Weighted grand means were calculated for each parameter for both annual and monthly totals. Coded Daily Activity Reports Coded daily activity reports were re~eived from 10 DWM's. Their daily activities had been coded using a series of 18 activity classifications developed for the Division of Wildlife "EARS" program (Employee Activity Reporting System, Appendix A). This reporting system is no longer in general use. Work time per activity was recorded to the nearest 0.5 hour. A total of 133 months was evaluated. This subsample represented approximately 25% of the total months examined. Quantitative data were totaled by coded activity for each DWM. Calculations were made to determine mean hours and mean percent time spent on a coded activity by each DWM during a l-year period, January through December 1981 (106 months of coded work) and a 1.5-year period, January 1981 through June 1982 (133 months of coded work). Weighted monthly and annual means were calculated by activity-type for both periods. Annual leave hours (Code 78) were excluded in 2 series of calculations (Tables 1, 2). The percentages shown reflect time worked per activity divided by total hours actually worked (x 100). Code 78 data (annual leave) were included in Table 3. These data represent time spent on an activity divided by the total hours the DWM was paid salary by the 5tate (x 100). The category "uncoded" was included (Tables 1, 2, 3) to show an approximate coding error rate expressed as a percent of total hours worked. All hours included in this category had not been coded or were obviously miscoded. Field Observations Twenty-nine days were spent on field observations of the law enforcement effort. Personal contact was made with 32 DWM's, E Area Wildlife Managers (AWM's) and 4 Regional Law Enforcement Coordinators (RLEC's) in addition to the Chief of Law Enforcement and his 2 assistants. Subjective interviews were conducted with these personnel regarding wildlife law enforcement activities in Colorado and research needs. �8 Literature Review and Current Research Available wildlife law enforcement research information and wildlife law enforcement articles of a descriptive or philosophical nature, publ ished and unpublished, were collected and reviewed. A short, informal questionnaire was sent to each of the 50 states, 4 Canadian provinces, and Puerto Rico. The questionnaire was intended to identify current or recent law enforcement research/evaluation efforts and perceived wildlife law enforcement related problems. Study Plans Two study plans were drafted, a 3rd outlined, and a 4th proposed. The first 2 study plans have been implemented under Federal Aid Project FW-26-R, they are: Illegal Purchase of Resident Licenses by Non-residents, Work Plan 1, Job 2; and Preparation or-an Annotated Bibl iography for Wildlife Law Enforcement Personnel, Wor~Plan 1, Job 3. A 5-year-program plan has been completed which outlines future wildlife law enforcement research projects including: the development of measurable wildl ife law enforcement objectives and the evaluation of publ ic attitudes towards wildl ife laws and wildlife law enforcement. Wildlife Law Enforcement Activities A questionnaire and several informal interviews were used to develop a list of wildlife law enforcement activities related directly to Division of Wildlife operating procedures and responsibilities. This Iist was examined for differences between Regions. All written methods for collecting or reporting wildlife law enforcement information were examined. Forms were reviewed for uniformity, and non-standard reporting formats were identified. Suggestions were made to correct deficiencies. The Line-Staff organizational structure and responsibil ities specific to wildlife law enforcement within the Division of Wildlife were identified and flow charts were constructed that demonstrate how information is exchanged, i.e., formal or informal. Apparent strengths and weaknesses in the system were identified and discussed. RESULTS AND DISCUSSION Coded Data District Wildlife Managers of the Colorado Division of Wildlife operate under the "multi-purpose" concept. Their job is, by description, multifaceted; a composite of many activities (Appendix B). Law Enforcement is only one component of this multifaceted position. Any analysis of the law enforcement activities of the DWM must be done in relationship to other activities. Reported time relationships between these activities are illustrated in Tables 1, 2, and 3. �Table 1. Percent time spent on an activity by month. from 1 region).a (Based on reported coded data from Dally Activity Reports of 10 District Wildl ife Managers b Code 10 Month Jan 111 18-· - 20 26 30 34 38 42 46 50 17.750.1914.021.202.634.150.5527.380.030.134.05 54 58 1.78 2.16 3.140.1318.981.46 62 66 70 711 Uncoded 0.26 Feb 7.00 2.28 13.02 2.34 0.42 3.38 1.84 24.83 1.10 0.63 5.49 3.76 0.89 5.68 0.08 18.16 8.67 0.44 Mar 10.10 1.30 17.67 0.72 0.69 3.97 0.50 27.68 3.62 1.26 4.80 1.68 1.43 2.80 0.08 14.83 5.83 0.34 Apr 12.18 4.54 15.87 1.24 0.22 3.95 0.18 29.24 0.33 0.37 5.85 1.65 0.95 2.30 1.10 17.32 2.38 0.33 May 11.580.6620.540.740.393.370.0433.110.661.475.301.05 Jun 11.00 0.96 17.63 0.05 1.42 4.53 0.14 32.31 0.00 1.63 7.88 0.96 0.91 0.29 0.05 15.90 5.19 0.24 Jul 12.94 0.50 19.05 0.00 0.00 2.68 0.00 34.27 0.20 1.07 7.38 0.13 1.34 0.80 0.27 16.53 2.62 0.20 Aug 11.98 0.81 17.14 0.27 0.07 4.76 1.00 29.35 1.01 0.94 6.41' 0.13 1.14 1.27 0.40 15.33 5.84 2.15 Sep 11.760.0017.601.801.113.560.5240.531.561.243.980.00 Oct 11.66 0.21 20.24 0.16 0.52 1.66 0.26 45.31 0.23 0.00 4.69 o.oe 2.07 0.31 0.52 10.81 0.31 1.04 Nov 12.'18 1.19 18.40 0.17 0.00 2.71 0.07 42.44 0.07 0.20 3.81 1.59 1.26 0.60 0.20 14.13 0.93 0.07 Dec 13.34 0.69 19.35 0.07 0.55 2.80 0.00 38.91 0.14 0.73 3.90 0.35 2.14 0.76 0.00 15.34 0.73 0.21 ighted annual mean 12.03 1.24 17.26 0.75 0.76 3.43 0.46 32.67 0.79 0.79 5.28 1.26 1.49 1.88 0.33 15.59 3.48 0.50 1.161.320.0013.984.59 0.64 2.970.000.0012.100.83 0.45 \.Je apercentages derived from 133 months of coded activities from January 1981 through June 1982. All leave hours were excluded. bSee Appendix A for description of activity. 1.0 �Table from 2. Percent 1 reglon).a time spent on an activity by month. {Based on reported coded data from Dally Activity Reports of 10,District ~1I1dllfe Managers o b Code 10 14 18 22 26 30 34 38 42 46 50 54 58 62 66 70 7~ Jan 16.66 0.26 14.96 1.94 1.94 1.89 0.84 29.31 0.00 0.10 4.61 2.88 3.04 0.31 0.00 18.96 1.99 0.31 Feb 6.85 2.72 13.03 2.55 0.65 2.46 2.26 27.90 1.10 0.65 6.30 5.69 1.36 1.26 0.00 18.17 6.63 0.55 Mar 10.20 0.05 20.60 0.76 0.98 3.96 0.71 31.59 1.47 1.79 6.62 1.68 1.66 0.95 0.11 14.33 2.28 0.27 Apr 11.654.1116.181.460.313.850.2628.640.470.527.341.72 Month 1.350.731.3016.623.02 Uncoded 0.47 May 9.87 0.95 20.49 0.22 0.56 4.21 0.06 33.31 0.00 0.62 6.78 0.90 1.46 0.28 0.00 13.18 6.42 0.59 Jun 9.62 1.40 18.48 0.07 0.88 2.92 0.28 33.26 0.00 0.49 10.82 0.56 1.07 0.42 0.00 15.60 3.65 0.21 Jul 12.94 0.50 19.05 0.00 0.00 2.68 0.00 34.27 0.20 1.07 7.38 0.13 1.34 0.80 0.27 16.53 2.62 0.20 Aug 11.98 0.81 17.14 0.27 0.07 4.76 1.00 29.35 1.01 0.94 6.41 0.13 1.14 1.27 0.40 15.33 5.84 2.15 Sep 11.76 0.00 17.60 1.80 1.11 3.56 0.52 40.53 1.56 1.24 3.98 0.00 2.97 0.00 0.00 12.10 0.83 0.45 Oct 11.66 0.21 20.24 0.16 0.52 1.66 0.26 45.31 0.23 0.00 4.69 0.00 2.07 0.31 0.52 10.81 0.31 1.04 Nov 12.18 1.19 18.40 0.17 0.00 2.71 0.07 42.44 0.07 0.20 3.81 1.59 1.26 0.60 0.20 14.13 0.93 0.07 Dec 13.34 0.69 19.35 0.07 0.55 2.80 0.00 38.91 0.14 0.73 3.90 0.35 2.14 0.76 0.00 15.34 0.73 0.21 14eighted annual mean 11.61 1.09 17.98 0.81 0.65 3.11 0.52 34.46 0.51 0.68 6.03 1.34 1.79 0.63 0.25 15.08 2.92 0.54 apercentages bSee Appendix derived A for from 106 months description of of coded activity. activities from January tl]!c:l_ugh December 1981. All leave hours were excluded, �Table 3. Percent from 1 region).a time on an activity bv month. {Based on reported coded data from Dally Activity Reports of 10 District Wildlife Managers Codeb 10 Honth Jan 1~-Tg-n 17.19.· 0.19 22 26 30 34 38 42 46 50 54 58 62 66 70 74 ---'71J 13.58 1.16 2.54 4.02 0.53 26.52 0.03 0.13 3.93 1.73 2.09 3.05 0.13 18.39 1.41 3.14 0.25 Uncoded Feb ~.86 2.23 12.75 2.29 0.41 3.31 1.80 24.32 1.07 0.62 5.37 3.68 0.87 5.56 0.08 17.79 8.49 2.07 0.43 Mar 9.72 1.25 17.00 0.70 0.66 3.81 0.48 26.63 3.48 1.21 4.62 1.61 1.38 2.70 0.07 14.26 5.61 3.81 0.33 Apr 11.944.4415.561.220.223.870.1828.660.320.365.73 May 11.40 0.65 20.23 0.72 0.38 3.32 0.04 32.60 0.65 1.45 5.22 1.03 1.14 1.30 0.00 13.77 4.52 1.53 0.63 Jun 9.08 0.79 14.56 0.04 1.17 2.68 0.12 26.67 0.00 1.35 6.50 0.79 0.75 0.24 0.04 13.13 4.28 17.45 0.20 Jut 11.69 0.45 17.20 0.00 0.00 2.42 0.00 30.95 0.18 0.97 6.67 0.12 1.21 0.73 0.24 14.93 2.36 9.96 0.18 Aug 10.76 0.72 15.41 0.24 0.06 4.28 0.91 26.38 0.90 0.84 5.76 0.12 1.03 1.15 0.36 13.78 5.25 10.13 1.93 Sep 10.13 0.00 15.17 1.55 0.95 3.07 0.45 34.92 1.34 1.07 3.43 0.00 2.56 0.00 0.00 10.43 0.72 13.83 0.39 Oct 11.66 0.21 20.24 0.16 0.52 1.66 0.26 45.31 0.23 0.00 4.69 0.00 2.07 0.31 0.52 10.81 0.31 0.00 1. Nov 12.18 1.19 18.40 0.17 0.00 2.71 0.07 42.44 0.07 0.20 3.81 1.59 1.26 0.60 0.20 14.13 0.93 0.00 0.07 Oec 12.32 0.64 17.87 0.06 0.51 2.58 0.00 35.93 0.13 0.67 3.61 0.32 1.98 0.70 0.00 14.17 0.67 7.66 0.19 0.72 3.240.4430.840.750.744.991.191.41 1.77 0.30 14.72 3.29 5.60 0.47 1.610.932.261.0816.972.332.01 0.32 04 ~Ieighted annual mean 11.361.1716.290.71 8Percentages bSee Appendix derived from 133 months A for description of coded activities of activity. from January throuqh June 1982. All leave hours were included. �Four activities: Code 10, wildlife inventory; Code 18, public relations; Code 38, law enforcement; and Code 70, administrative and clerical services accounted for as much as 88% of the total time worked during a given month (~~ 77.5%, range 63.0-88.0%, Tables 1, 2, 3). Percent time spent on wi ldTife inventory (Code 10) was highest in January (~ 17.5%) and lowest in February (~7.0%). This was probably attributable to early winter survey counts of big game and waterfowl. Percent time spent on public relations (Code 18) remained relatively stable throughout the period but showed a slight increase during both May (early fishing) and October (big game season) reflecting increased public contact by DWM's. Law enforcement (Code 38) represented on 1y 25% of the workt ime in Februar-y but peaked at 45% in October during the big game seasons. Administrative and clerical services (Code 70, "paperwork") required 19% of the total worktime in January but only 11% in October. These numbers are virtually meaningless when considered alone in the absence of objective statewide standards for each activity. The information they provide relates to the relative distribution of worktime as described by this subset of 10 DWM's from 1 Region when those DWM's were restricted to 18 coded activities and definitions. They cannot be used to project specific statewide values with any degree of confidence. The percentages may sugsest the relative ranked priority of an activity during a month as described by the DWH's. These percentages would not be the best indicator of total workload during that month. Workload must be considered on a combination of characteristics and not just hours worked. Workload Beattie (1976b) described the workload of wildlife law enforcement officers as: liThe term workload, as it applies to wildlife law enforcement sections and their employees, will vary with the role of the officer. The workload of an officer represents the sum of all activities he (she) performs or is expected to perform. Actual (based on objectives) and perceived workload may be viewed differently by individual officers and the wildlife law enforcement agency. Some officers may, in actuality, have a lighter workload (based on agency criteria) than other officers but may perceive themselves as having a heavier workload. The important point is that agencies must establish and articulate meaningful criteria for determining individual officer workload and wildlife agency organizational workload." Wildlife officer workload would include the hours spent by an individual officer in the performance of his/her duties. Hours worked or percent time spent per activity would be one indication of workload. However, hours worked or percent time does not address the quality of those hours or the efficient and effective use of the time coded into a given activity category. District Wildlife Managers throughout the State consistently reported work hours in excess of those required of State employees by personnel regulations. It is the feeling of most field personnel that extra hours must be expended to properly do the job required of them. This assumption mayor may not be �13 true. McCormick (1970) documented the contact rate between wildl ife agents and the public prior to and following the enactment of a mandatory 40-hour work week for all wildl ife officers. McCormick's results suggested that, with improved planning of efforts and better definition of priorities, field contacts per agent actually increased after the 40-hour week was enacted. A 40-hour work week may not be practical in Colorado under the present multipurpose concept. A clear definition of priorities for the DWM position would be necessary along with average completion times by activity and average number of activities per day before a 40-hour week could be considered. What McCormick (1970) does suggest is that with an increased planning effort, efficiency may be increased. Major Work Period October was the peak work period for law enforcement activity and for total hours worked during January - December 1981. Approximately Mean 45% of all hours worked during October were assigned to law enforcement. DWM time expenditure during October was 275 hours and 27 days worked statewide. This is approximately 107 hours more than required by state personnel regulations (DWM's are exempt). 0" an annual basis, based on the information provided, the "average" DWM worked 417 hours in excess of those required (Table 4). This would suggest that statewide, DWM's reported approximately 47,000 "excess" or "overtime" hours during January - December 1981. The qual ity of those hours cannot be determined. Table 5 lists calculated values for individual DWM's. These values include: months evaluated, mean days worked per month, mean days off per month, mean hours worked per month, range in hours worked per month, and mean leave days (of all types) taken per month. A partial profile of an "average" DWM would read something like this: Every month he/she worked an average of 23.48 days; took off 5.6 days; worked approximately 201.5 hours, of which 34 hours could be considered "overtime"; and used 1.36 days leave. Hours worked per month ranged from 45 (excluding leave hours) to 420. During 1981 he/she issued approximately 40 Penalty Assessments (PAis) and Court Citations combined. Figure 1 graphically illustrates the average hours worked per month on each activity during January - December 1981. Wildlife inventory, publ ic relations, law enforcement, and administrative and clerical services are the predominant activities described by this subset of 10 DWM's. Division equipment and facilities maintenance (Code 50) is the only additional activity that on average required over 10 hours per month to complete. These figures represent an accurate picture of relative hourly work loads as identified by the individual DWM's. If these are to accurately represent the true distribution of DWH activities during January-December 1981, several assumptions must be made: 1. All activities have been defined in a similar manner by every DWH. 2. Time reported for each activity accurately represents the actual time spent on that activity. The quality standard between DWM's is the same. 3. All activities actually undertaken are represented act ivi ty report. in the daily �Table 4 . Weighted mean days and hours of reporteG work per montha by 32 DWMls during January 1979 - June 1982. (514 months of DARls evaluated.) .j::- Month Days/ month Hours/ month Mean hours/ actual days worked Required work days/month Required hours/ month Mean overtime hours/month Jan 24.86 200.14 8.05 21 i68 32. 14 Feb 20.68 172.29 8.33 18 144 28.29 Mar 24.63 196.02 7.96 22 176 20.02 Apr 23.38 200.52 8.58 22 176 24.52 May 24.72 208.48 8.43 21 168 40.48 Jun 21.96 187.78 8.55 22 176 11.78 Jul 24.05 210.99 8.77 21 168 42.99 Aug 22.32 192.30 8.62 21 168 24.30 Sep 21 .73 199.20 9. 17 21 16e Oct 26.95 274.70 10. 19 7.1 168 45.54 Nov 24.08 213.54 8.87 21 168 53.54 Dec 21.56 176.82 8.20 21 168 R.B3 aAnnually calculated mean hours/month = 201.52. AnnuaJ ly calculated 106.7 mean days worked/month = 23.48. �15 Table 5. Calculated values derived from daily activity reports (DAR's) of 32 District Wildlife Managers of the Colorado Division of Wildlife from January 1979 through June 1982. OWM Months evaluated 2 18 18 3 4 13 18 5 6 13 4 12 6 18 18 18 18 7 8 9 10 1I 12 13 14 IS 16 17 18 19 20 21 22 23 24 25 26 27 28 Mean days worked/me 29 30 31 25 12 32 _ll 13 Mean (weighted) Mean hours worked/rna Range hours worked/rna 24.50 23.00 22.08 21.56 24. 15 22.00 21.75 22.33 21.911 21.22 25.30 22.54 24.00 24.92 23.87 21.45 23.73 22.67 25.92 21.08 24.08 24.96 25.68 23.17 24.54 ~ 23.48 5.10 212.44 136-319 0.86 198.66 207.85 184.11 5.77 8.00 6.67 6.17 6.78 235.88 175.00 192.83 168.17 153-232 45-261 56-254 196-305 144-193 114-264 101-221 108-228 107-208 95-252 150-248 192-328 76-224 0.94 1.54 1.22 0.46 0.00 2.08 5.38 4.08 5.21 7.08 5.83 5.52 6.82 4.64 23.79 20.42 23.26 Mean days leave/rna (al~l. 6.50 6.70 7.66 7.39 3.94 5.72 3.83 7.15 6.00 23.72 26.50 21.54 12 13 24 24 24 24 12 23 23 II II 12 12 12 12 28 Mean days off/me 6.50 2.67 7.25 6.08 4.18 3.96 6.33 4.08 ~ 5.60 173.90 167.19 187.50 202.80 260.67 172.62 199.00 226.73 242.0B 191.42 172.83 207.39 205.52 168.27 182.18 1.33 1.67 2.28 1.60 1.06 0.00 1.77 1.91 143-313 129-420 152-351 95-262 0.96 1.44 103-259 87-31 I 114-278 2.83 1.35 1.04 89-207 116-224 2.09 2.00 142-293 162-328 115-240 140-238 71-241 114-296 134-306 1.25 1.83 2.17 0.25 1.32 0.76 1.29 204.96 225.67 192.96 188.75 198.04 241.52 216.67 176.38 164.08 114-329 91-226 0·92 1.85 1.46 201.52 118-271 1.36 �16 ACTIVITY 10: WILDLIFE ACTIVITY INVENTORY 14: REGIII.ATION RE':OMMEtlDATIONS 10 30 8 6 20 4 10 o J FHA ACTIVITY A H 18: PUBLIC 50 SON D M RELATIONS ACTIVITY A H 22: HUNTER M A A o N 0 o N D N 0 SAFETY 5 4 40 30 20 o F ACTIVITY 26: RESEARCH ACTIVITY 30: M GANE A DAMAGE 5 4 6 x SE o = 5.79 = 1.74 o M A t~ A o Fig. 1. Mean hours spent per month on a defined activity by 10 District Wildl ife Managers, N.E. Region, January-December, 1981, (note variable scale for hours). �17 ACTIVITY 34: OISTRIBuTE WILDLIFE ACTIVITY 30: LAW OIFORCEHENT 110 4 = 0.97 SE = 0.99 x 100 90 2 x = 64. I ~ o SE J F M ACTIVITY 42: A M HABITAT o A N = 16.46 o DEVELOPMENT ACTIVITY 3 46: PUBLIC FACILITIES MAINTENANCE 3 ' 2 o o SE J M ACTIVITY 50: A A M DIVISION 1.01 FACILITIES SE = 0.96 o D M MAINTEI~ANCE ACTIVITY 5~: A M ENVIRONMENTAL A o N D PROTECTION 19 10 16 6 13 4 10 7 1~ o M A M F F" i g. 1. (continued) A o N D F M A M A o N 0 �18 ACTIVITY 58: RESOURCE ACTIVITY RECONNAISSMICE 62: PLANNING/8UDGETING 5 4 x = 3.33 SE = 1.29 a o F ACTIVITY 66: REAL ESTATE ACTIVITY H A H A S 0 ~I o AOMINISTRATIVE/CLERICAL 70: x = 2~.O9 35 SE = 3.55 2 30 25 o o A ACTIVITY 711: IN-SERVICE N 0 LEAVE ACT IV ITY 78: TRAI N ItJr. 40 15 x = 3.85 30 12 9 20 6 10 0 0 M Fig. 1. (conti nued) A M A 0 N D �19 If these assumptions are not true, then the val idity of the results can be questioned. If these assumptions cannot be satisfied, the use of this technique to gather information must be questioned. Definition Problems Poor definitions can be a constraing to categorization. The problem Careful and thoughtbeing that "what is" to one person "is not" to another. ful definitions must be a part of any attempts to code and categorize activities. The system must be thoroughly explained through training so that all participants will respond in similar fashion to the question "What category should I place this activity in?" and all participants must report all activities correctly. The coded information examined in this study must be treated carefully with the understanding that it represents the best information available concerning daily activities of DWM's, but that it may contain serious reporting errors based on differential judgment regarding definitions of different activities and different reporting practices. Area Wildlife Managers The Colorado Division of Wildlife (CDOW) has 20 Area Wildlife Managers (AWM's or Area Supervisors), with 1 assigned to each of the 16 CDOW Administrative Areas and 4 serving as Regional Law Enforcement Coordinators (RLEC's). One RLEC is assigned to each Region. Like the DHM, the AWM position is multi-faceted (Appendix C). Generally speaking, AWM's are selected following an examination from the DWM ranks, although in theory it is possible to move into the AWM position after 4 years of experience at the DWM level or above. This experience must have included enforcement and administrative responsibilities. AWM's are directly responsible to the Assistant Regional Manager in their Region. The majority of the A\.JM'stime (exclusive of those AWM's serving as RLEC's) is spent in budget preparation, attendance at meetings, distribution of equipment and supplies, report preparation, and public contacts. AWM's typically work with DWM's as a supervisory field enforcement officer during peak law enforcement periods and on special enforcement problems. There is no line authority from the Chief of Law Enforcement or his assistants to any field law enforcement position, including the AWM and DWM as well as other CDOW commissioned officers. This lack of line authority can complicate the implementation of new law enforcement procedures. Likewise, it has hindered the standardization of law enforcement practices on a statewide basis. With the creation of the RLEC position some of these difficulties may be surmounted. Regional Law Enforcement Coordinators The 4 RLEC's are responsible for planning, implementation, monitoring, and coordination of all Regional wildlife law enforcement activities. The ~LEC conducts or supervises special Regional investigations; plans, conducts, evaluates, and coordinates Regional law enforcement training; conducts public education and information programs on a special needs basis; serves as Regional officer for license suspension hearings; and serves as staff advisor to the Regional Manager on law enforcement issues. The RLEC position is important to the standardization and coordination of wildlife law enforcement activities statewide as well as within the Re0ions. �20 Wildlife Law Enforcement Activities A Iist of wildl ife law enforcement activities conducted by or involving employees of the Division of Wildlife was formulated (Appendix D). These activities are not grouped according to job description because of overlap and multiple levels of involvement. The list was based on current job descriptions, Statutory and Regulatory requirements, Division Policies and Directives, standard operating procedures for wildl ife law enforcement, and specific Regional needs. It was determined from questionnaires completed by the RLEC's that these activities are recognized by all Regions of the state and can be considered standard statewide. It proved unnecessary to group activities by Region. The following differences between Regions (and within Regions) should be noted: 1. Variable wildlife 2. Different wildlife Regions. law enforcement effort between and within Regions. law enforcement priorities between and within No effort was made to quantify these differences. Factors that may influence these differences are: wildlife populations, density and species composition, public access, Area and Regional personnel attitudes, human population levels, socioeconomic conditions, and topographic features. Wildl ife Law Enforcement Data Collection and Reporting The assumption made at the beginning of this study was that many nonstandard reporting methods were used in each of the 4 Regions. Non-standard was defined as: any method of data collection or reporting of wildlife law enforcement data that varied between at least 2 Regions. A common example would be the reporting of similar information by different Regions on different reporting forms. Non-standard Forms Collection and examination of the different reporting forms supported this assumption. However, the problem was not as serious as initially estimated. Only 15 of 60 forms currently used were non-standard. These non-standard forms (Appendix E) were developed primarily on the Regional level and generally reflected information of Regional concern under the present reporting system. This procedure is acceptable if the information collected is irrelevant to the statewide wildlife law enforcement program or if it is used to evaluate a program or portion of a program of Regional concern only, If the information collected has statewide value it should be collected and reported in a standard way on standard forms that are approved by the Denver office. �21 The benefits of standard reporting include: uniformity in data collection statewide, an increased level of professionalism related to an increased abil ity to understand and communicate with others, the ability to analyze collected data more easily, ease of computer storage and retrieval of collected data, simpl ification of the communications process across Regional boundaries, and simpl ification of the report generating process. Standard Forms The 45 standardized data collection and reporting forms (Appendix F) were specifically developed to collect statewide data and out of the necessity for standardization due to legal concerns. An example of the latter would be the Penalty Assessment/Summons form. The ultimate use of the data collected was not determined for all of the documents. Likewise, standardization between Regions does not necessarily mean that all Regions use the form in a similar manner. Some forms may only be partly used while others may be completed entirely. Data currently collected for wildlife law enforcement are used primarily to meet mandatory reporting requiremenis establ ished at a variety of levels and to document work-related requirements of individual employees. An example of this would be a requirement made of DWM "Sm i th" that he/she make 200 field contacts with big game hunters between 1 October and 30 November 1984. DWM IISmithl1could record these contacts on a standard "Fi e ld Con t ac t " form. These data are not currently used to evaluate the overall efficiency or effectiveness of any given program; they only examine the performance of the individual. Its use could be expanded to evaluate the overall program efficiency and effectiveness. Individual performance may be summarized into monthly, quarterly, or annual reports. No uniform efforts are currently known to be in use that relate previous job efforts to current efforts as a tool of evaluation. Antiquated data storage and retrieval systems currently in place within the Division tend to perpetuate this problem. Current manual storage and retrieval methods cannot meet the ever-increasing volume of informa·tion. Manual techniques lack the flexibil ity of cross-reference to other valuable files. In wildlife law enforcement,this may cost an officer a case or in a worst case situation, someone1s life. Under the current system data filed are data lost. This is a serious weakness and should be corrected as soon as possible. Increased use of automated data storage, data retrieval, and standard data collection within and between all Regions is highly recommended. Considerable effort should be directed to the evaluation of the data collected and a system developed to feed back the results to the data col lectors. District Wildl ife Managers are the primary data col lectors for wildl ife law enforcement and should be one of the primary target groups in the reporting and feedback process. The reports should be uncluttered and concise. This evaluation effort should be a joint effort among Denver law enforcement staff, wildlife law enforcement research, and the Regional law enforcement coordinators. �22 Wildl ife Law Enforcement Information Exchange District Wildl ife Managers. - Wildlife law enforcement information exchange occurs at many different levels but primarily occurs at the DWM-AWMRLEC level. The DWM-AWM-RLEC link is at the center of wildl ife law enforcement in Colorado (Fig. 2). The DWM is the backbone of basic wildlife law enforcement in the state while the AWM and RLEC provide professional guidance, supervision, and specialized support. District officers gather routine intelligence information from field contacts, field patrol, Operation Game Thief calls (OGT), other officers and staff personnel, and from social contacts. This information may result in direct action by the DWM, no action, or an exchange of information with other DWM's and/or supervisors. Data collection, informal as it may be, results in an incremental growth in overall knowledge of violators and violations occurring within the state. This incremental growth does serve to significantly increase the knowledge of an individual officer. However, once an officer leaves the organization, moves to a new district, or changes jobs within the organization most of that knowledge is lost. Standard data collection, better documentation, and efficient data retrieval systems would minimize this information loss. Area Wildl ife Manager.- The AWM is a 2nd pivotal link in the exchange of wildl ife law enforcement information (Fig. 2). The attitude of any individual AWM towards any wildlife management activity, including wildlife law enforcement, has an immediate impact on the priority placed on that activity within an Area. The flow or collection of wildlife law enforcement information (or other wildlife management information) is hindered if an AWM does not strongly support wildl ife law enforcement conceptually and in ~ractice as a tool of wildlife management. This attitude may be reflected in the attitudes of the personnel he/she supervises and thus further compl icates information flow through appropriate channels in appropriate form. There appears to be no easy or immediate solution to this dilemma. Human behavior cannot be changed easily. However, with prope~ documentation of wildlife law enforcement problems through the collection of reliable data I believe that responsible people should respond in responsible ways when presented with facts. This is the appropriate way to attempt to change human behavior at all levels of authority. Regional Law Enforcement Coordinator. - The RLEC position has added a new look to wildlife law enforcement in Colorado. This position is a staff position. The RLEC serves as a consultant to upper level Regional administrators on wi ldlife law enforcement issues and as a coordinator for special wildlife law enforcement activities that occur within or between Regions. The objectives of the RLEC position are still being solidified but it is clear that the position should provide a more professional image to wildl ife law enforcement and will aid in the standardization process for many of the enforcement activities that must be performed with consistency statewide. Examples would include: handling of critical incidents, license suspension hearings, standard operating procedures, data collection, and evaluations. �Pub Iic Area ~IiId life Other DWM's Field Contact I ~ OGT LEGEND: Agencies ••• - Fig. -... 2. Formal Exchange I nforma I Exchange Wildl ife law enforcement information flow relationships. N \..N �24 The RLEC serves as the clearinghouse for Regional law enforcement information. Most of the information exchange routes are informal (Fig. 2). This lack of formal routes of information exchange is probably the most serious weakness of the position as there is no line authority over any field enforcement personnel. This could be solved by assigning wildlife law enforcement specialists to each RLEC. This move would enhance the statewide law enforcement effort by providing teams who could: (1) respond to special assignments on short notice, (2) spend more time on investigations, (3) conduct special investigations, and (4) serve as a data collection team to help evaluate program effectiveness. These positions would not eliminate the role of the DWM in wildlife law enforcement but would eliminate much of the extended investigative work, eliminate some routine enforcement, and would provide coordinated assistance to existing DWM-AWM law enforcement efforts. Wildlife Technicians. Wildlife technicians are frequently commissioned and assist in many wildlife law enforcement situations. They are generally assigned other primary responsibi Iities and are used on an "as needed" basis. Figure 3 summarizes the wildlife law enforcement Line/Staff relationships within the Division of Wildlife, including the wildlife technician. Denver Law Enforcement Staff. Another level of information exchange occurs at the Denver staff positions. Information flows between agencies, internal policies are developed and revised, and liaison with Regional law enforcement is maintained at this level. Contacts are maintained with other State and Federal wildlife agencies and with a wide variety of other law enforcement agencies. Intelligence information received through the Operation Game Thief (OGT) program and license violation information is passed on by the Denver staff to DWM's for field investigations. Any covert actions are coordinated through the Denver staff. Weaknesses in Information Exchange The primary weakness in wildlife law enforcement exchange within the Division of Wildlife is the lack of a centralized, automated repository for much of the data collected and the associated inadequate syste m for data retrieval. This could be solved by the acquisition of on-line data storage on any of a series of avai Jable computer systems. Data storage could be centralized in the Denver office or localized at the Regional level and made accessible through a tele-networking system. A secondary weakness is the lack of reliable information. Standard data collection needs must be identified with standard forms used as tools to measure program efficiency, effectiveness, and productivity. The RLEC's and the Denver law enforcement staff should recommend and provide necessary standard format ideas. r--' �r- WILDLIFE I COMlllSSION , • S' a ,.,------_.r-c;:s.C2t~--__ t.S) I • rooGDll--ceo-~C) 8 I ~ <S> ~ ~ e Fig. District \/i ld life Managers 3. "ECI Regional Managers Chief Law Enforcement 8. Line/Staff Wildlife • • G I Staff Line Area Wildlife Managers Regional Law Enforcement D 1+ I ~'8188 HI J I J I Ji"e ~'8£'8A~8 • Assistant Assistant fi 'i i G LEGEND: ••• Coordinators RcEC "EC AiM RLEC A~M Technicians relationships for wildlife law enforcement in the Colorado Division of Wildl ife. N V1 �26 Overview of the Literature Wi Idlife law enforcement research does not have a well developed literature base. ~uch of the information that is available is fragmented, poorly reported, incomplete, or the results are based on unfounded assumptions and poor research design. A significant portion of all published wildlife law enforcement information is philosophical or descriptive in nature but does generally contribute to the overall understanding of problems in wildlife law enforcement. Wildlife forensic research and much of the administrative-type research conducted at Virginia Polytechnic Institute and State University (VPI & SU), under the guidance of R. H. Giles, Jr. are notable exceptions to the descriptive or philosophical approach. In both cases, diligent attempts are being made, or have been made to maintain the standards of the scientific method. Wildlife law enforcement needs well designed research to help it evolve as a science. Forensic research must be carefully conducted so that results and applications can withstand rigorous testing in the courts. Work by Brohn and Korschgen (1950) helped establish the use of the precipitih Test as a tool of wildlife law enforcement following earlier work by Gay (1908) and Clark (1914). This test is useful in the identification of many animal species using only meat or blood samples. Oates et a1. (1974) and Glover and Korschgen (1980) have recently experimented with this particular technique. Johnson et al. (1980) used potassium levels in the vitreous humor of eyes from mule deer (Odocoileus hemionus) to calculate time of death. This work has recently been replicated in Missouri (R. Glover, pers. commun.) and Illinois (Woolf and Roseberry, no date). Much additional forensic research has been conducted and interested readers are referred to the publication by Wilson (1977) for additional information. Virginia Polytechnic Institute Studies Significant wildlife law enforcement research was undertaken by R. Giles and associates at VPI & SU during the 1970's as part of a joint law enforcement research effort. Funding came from a variety of sources including: the Wildlife Management Institute, National Wildlife Federation, National Rifle Association, American Petroleum Institute, Georgia State Game and Fish Division, South Carolina Wildlife and Marine Resources Agency, Tennessee Wildlife Resources Agency, and the Virginia Commission of Game and Inland Fisheries (Giles 1976b). These studies were some of the first attempts at an objective examination-of the wildlife law enforcement activity as an integral part of the overall job of wildlife management. Giles has been a primary motivating force behind wildlife law enforcement research since the 1960's when he served as advisor to J. R. Vilkitis at the University of Idaho. Vilkitis' research on the illegal take of big game and characteristics of big game violators in Idaho (Vilkitis 1968) was the first attempt to quantify poaching losses of big game. At VPI & SU, Giles continued law enforcement research projects into the late 1970's (Giles 1971; 1976~, ~,_;1978; Giles et al. 1971). �27 Other examples of the work and extent of the projects supervised by Gi les at VPI & SU include: Kaminsky (1974) examined the problem of spotl ighting of deer (Odocoileus virgianus) in Virginia. His analysis helped demonstrate that research could be conducted in wildlife law enforcement. Ritter (1975) developed measurable law enforcement objectives and performance criteria for state wildl ife agencies. Ritter's work could serve as the foundation in the development of comprehensive evaluation programs by other state wildlife agencies. Beattie (1975; 1976a, b ; 1978a, b; 1979; 1981a, b) and associates (Beattie et al. 1977a, b , c ; 1978a, b, c; 1980) has been-a -prolific writer and an advocate of well d;signed-law enforcement research projects. C. Cowles, a contemporary of Beattie's at VPI & SU, and also a student of Giles was instrumental in the development of optimum deployment techniques, based on computer models, for wildlife law enforcement agents (Cowles 1977; Cowles et al. 1977, 1978, 1979). The collective efforts at VPI & SU demonstrate the diversity possible for research in wildlife law enforcement. This work was by no means definitive, but should serve as the foundation for more comprehensive, integrated wildlife law enforcement research. Integration is the key word in successful wildlife law enforcement research. The problems facing wildlife law enforcement today are not strictly biological. The disciplines of Biology, Chemistry, Ecology, Sociology, Psychology, Politics, Economics, Computer Science, Statistics, Engineering, and Police Science must be fully integrated to help solve these problems. Other Research Efforts Other individuals and agencies have had active roles in the evolution of wildl ife law enforcement research. W. B. Morse, now with the Wildlife Management Institute, gathered wildlife law enforcement information for many years on a regional and later on a national basis. His reports to the Western Association of State Fish and Wildlife Agencies and other published works have stimulated thoughts on needed law enforcement research (Morse 1957, 1958, 1963, 1968, 1972, 1973, 1976, 1980, 1982). The law enforcement divisions of Michigan, California, and New Mexico have each conducted research projects to measure the effectiveness of their law enforcement efforts (Hussain 1977; Purol 1982a,b; Purol and Fournier 1979; Purol and Zambetis 1982' McCormick 1968, 1970; Mikel 1981a,b; Pursley 1977a,b; Vaught 1975; Vaught ~nd Turner 1975). These projects have met with varying success but have been successfully used to impact their law enforcement operations. Field oriented wildlife law enforcement research efforts have been reported from most states (Beattie and Giles 1979: Smeltzer 1983, Anne nd lx r,). These projects have generally been problem specific. The tendency to pursue only problems of immediate concern may be a weakness of many present wildlife law enforcement research projects. Wildlife law enforcement research appears to be slowly evolving into an accepted scientific discipline. The process has been and will remain slow. Researchers and other individuals involved in law enforcement can aid this process of evolution by lending support and constructive criticism to the overall process. �28 CONCLUSIONS Coded Daily Activity Reports Coded daily activities can be valuable in evaluating the efficiency and effectiveness of a program or an individual. When total hours expended per activity are weighted by agency objectives, the direction of the efforts can be evaluated. This would not be a total measure of effectiveness, only an indicator. Before a complicated reporting system is constructed,difficult questions must be answered: 1. Of what do we want to measure the efficiency and effectiveness? 2. How would these efforts 3. Can we make such a system work? 4. How will the information obtained be used? impact the total program? 5. How are the results to be communicated to the individuals who provided the information? 6. Who will be responsible for implementing, monitoring, and evaluating these efforts? Identification of the answer to question #1 is extremely important. Levels of efficiency and effectiveness can be measured only if we can clearly rlefine what we want to measure and then are given a standard to use for comparison. Lacking a standard, one must first be developed. This must be done before any changes are rn.rde to a program if we expect to measure the results of those modifications. Standards can be developed in 2 ways: known outputs such as miles driven, contacts made and recorded, programs given, articles written, reports completed, and arrests made can be used to set a "standard". The 2nd way involves establishing "desired" levels for difficult to quantify areas, such as tolerable levels of violation for specific laws. These standards should be developed based on discussions, establishing and implementing methods to measure current levels, and finally adjusting the standard as more information becomes available or as collection techniques improve. Objective evaluation is not possible without standards. The impact of any evaluation on the overall program must be considered. Just because we have the ability to evaluate a specific activity or function does not mean that we should evaluate it. A comprehensive reporting system will only be as good as the data received and the support system developed to evaluate them. It will be necessary to clearly define all activities or functions that wil I be monitored and evaluated. To encourage accurate reporting, there should be flexibil ity in some activity definitions and specific definitions in other activity areas. Above all, the information obtained must be used. �2':' USc of the information obtained is vital to the continuation of the system. If we don't use it, we shouldn't collect it. Use of the information to positively impact the overall program will encourage accurate and consistent reporting of data. Any information collected, analyzed, and reported should be made available to those individuals who spent time in the collection and analysis of the data. This is true of all data collection functions. Failure to complete the communication process by not supplying feedback to those who originally supply the data will result in rapid deterioration of the reporting system. Responsibility for the implementation, monitoring, and evaluation of any data collection efforts must be clearly defined. Failure to do so will result in another system in which lack of accountability causes the system to fa i I. Current Daily Activity Reports The current daily activity reporting form (DAR's uncoded) provides the information necessary to evaluate days and hours worked per month, days leave taken per month, days off per month, and gives a brief narrative description of the daily activities of the DWM or AWM. In its present form and as it is typically used, the DAR does not provide a convenient way to list daily activities by category or to quickly analyze them. It is, however, an adequate method to providt the information currently used by the CDOW to monitor the work of its personnel. With the addition of clearly defined, coded activities, it could be used as a coding form and could provide additional information on specific daily activities of all personnel. A support system would be needed to collect and analyze any information gathered. Training would be required to promote consistent reporting of activities. Any changes in the system must be evaluated and directed to impact the overall Division functions. Standardized Reporting Lack of standardized reporting of law enforcement information is due largely to the decentralization of law enforcement authority. Regional law enforcement programs have evolved based on the needs of a particular Region. The creation of the RLEC position should aid in the standardization of statewide law enforcement efforts. Standardized reporting and a system to collect and analyze these data must be in place before substantive efforts can be made to evaluate present levels of enforcement efficiency and effectiveness. Concurrent with these efforts must be the development of measurable wild] ife law enforcement objectives. Beattie et al. (1977a) stated: "A clear and precise formulation of wildl ife law enforcement objectives must precede an enforcement research program; they must be operationally defined so progress can be measured." Simply stated, we must know where we are, where we want to go, and how we wi II know when we get there. It should not be difficult to complete the standardization process. With only 15 non-standard documents currently in use out of ~O, we should be able to move rapidly towards standard information output. �30 LITERATlJRE C ITED Beattie, K. H. 1975. Anti-poaching campaigns: a tool of wildl ife law enforcement? Proc. Southeast.Assoc. Fish and Wild1. Agencies 29: 728-743. tors. 1976a. A descriptive assessment of Mississippi game law cooperaM.S. Thesis, Miss. State Univ., Starkville. 96pp. 1976b. A review of crimeload, workload, and manpower standards in wildlife law enforcement. VPI & SU, Southeast. Reg. Wildl. Law Enforcement Res. Proj. 53pp. 1978a. A comparison of hunting satisfaction of Virginia Wildlife and Colorado Outdoors hunter - subscribers. Proc. Southeast Assoc. Fish and Wildl. Agencies 32:738-744. 1978b. An initial bibliography of wildlife VPI & SU,-Southeast. Reg. Wildl. Law Enforcement law enforcement. Res. Proj. 21pp. 1979. A social psychological investigation Virginia sportsmen towards laws and regulations. VPI & SU, Blacksburg. 220pp. of attitudes of Ph.D. Diss., 1981a. The influence of game laws and regulations on hunting satisfaction. Wildl. Soc. Bull. 9:229-231. 1981b. Warnings versus citations Wildl. Soc. Bull. 9:323-325. , and R. H. Giles, Jr. ---research needs and current in wildlife law enforcement. 1979. A survey of wildlife law enforcement research. Wildl. Soc. Bull. 7:185-188. , , and C. J. Cowles. ---w--:-"ildlifelaw enforcement data. Agencies 31:698-708. 1977a. An analysis of nationwide Proc~ Southeast.Assoc. Fish and Wildl. , and 1977b. Fines in wildlife law enforcement. Proc-.-S-o-u-t'-heast. Assoc. Fish and-Wi rar , Agencies 31 :690-697. enforcement. and 1977c. Lack of research in wildl ife law Wildl. Soc. Bull.-5:170-174. 1978a. Quasi-experiments, multiple indica--_,--- , and tors, and enforcement effectiveness. Proc. West. Assoc. Fish and Wildl. Agencies 58:76-91. 1978b. Relative importance of enforcement ---- , and objectives and seriousness of violations in relation to objectives. Proc. Southeast.Assoc. Fish and Wildl. Agencies 32:808-815. �31 _..,.__ ' and 1980. Estimating illegal kill of deer. Pe+e s 65-71 in R. L. Hine and S. Heh ls , eds. White-taiJ.;d dec r popula t i on managc;;ent in the north central states. Proc. 1979 Symp., North Cent. Sect., The Wildl. Soc. _______ , T. A. Pierson, and H. l. Gilliam. 1978c. A readership preference survey of Virginia Wildlife subscribers. proc. Southeast.Assoc. Fish and Wildl. Agencies 32:732-737. Brohn, A., and l. J. Korschgen. 1950. Precipitin test -- A useful tool in game-law enforcement. Trans. North Am. Wildl. Conf. 15:467-478. Clark, F. C. 1914. Forensic value of the precipitin test in the enforcement of game laws in California. Univ. Calif. Pub!. Patho!. 2:131-138. Cowles, C. J. 1977. Optimum deployment of wildlife law enforcement agents: a problem analysis. VPI & SU, Southeast. Reg. Wildl. law Enforcement Res. Proj. 31pp. ___ ..,..' R. H. Giles, Jr., arid K. H. Beattie. 1977. Dyanmic deployment of wildl ife law enforcement manpower -- A decision aid. Proc. Southeast. Assoc. Fish and Wildl. Agencies 31 :679-689 . • K. H. Beattie, and R. H. Giles, Jr. 1978. A survey of methods of ---r-ecording reports of fish and wildlife violations. Fisheries 3:8-11. _____ , and pliance estimators. 1979. limitations of wildlife Wildl. Soc. Bull. 7:188-191. .,..,..,..-=- Gay, F. P. 1908. A contribution to the forensic precipitin test. J. Med. Res. 19:219-224. law com- value of the musculo- Giles, R. H., Jr. 1971. Wildlife law enforcement and research needs. Pages 131-133 in R. D. Teague, ed. Manual of wildlife conservation. The Wildl. Soc~ Washington, D.C. 1976a. Alpha-man: a theory of wildlife law violation. Southeast~ Reg. Wildl. Law Enforcement Res. Proj. 12pp. VPI & Sl, 1976b. The Southeastern wildlife law enforcement research project: -progress and perspectives. Proc. Southeast. Assoc. Fish and Wildl. Agencies 30:680-684. 1978. \~ildlife law enforcement. Pages 343-377 in R. H. Giles. Wildlife management. W. H. Freeman and Co., San Francisco, Calif. , M. Kaminsky, ---research -- The and J. McLaughlin. 1971. Wildlife law enforcement context and the needs. Proc. Southeast. Assoc. Fish and Game Comm. 25:677-687. �32 Glover, R. L., and L. J. Korschgen. 1980. Evaluation of two methods of preparing antisera for precipitin tests. Wildl. Soc. Bull. R:118-122. Hussain, N. G. 1977. An evaluation of the Michigan wildlife law enforcement effort. Proc. West. Assoc. State Game and Fish Comm. 57:35-66. Johnson, B. C., L. Maguire, and D. Anderson. 1980. Determining time of death in mule deer by using potassium levels in the vitreous humor. Wildl. Soc. Bull. 8:249-252. Kaminsky, M. A. 1974. Analysis of the spatial and temporal occurrence of deer spotlighting violations in Virginia. M.S. Thesis, VPI & SU, Blacksburg. 170pp. McCormick, J. B. 1968. A procedure for evaluating the effectiveness of wildlife law enforcement. Proc. West. Assoc. State Game and Fish Comm. 48:626~639 . . 1970. An evaluation of wildlife law enforcement -----W~est. Assoc. State Game and Fish Comm. 50:517-540. effort. Proc. Mikel, H. C. 1981a. Application of deterrents to reduce fishing violations. Proc. West. A~soc. State Game and Fish Comm. 62:246-249. 1981b. Illegal harvest of warmwater game fish. Game and Fish Fed. Aid Proj. F-51-R. 14pp. New Mexico Dep. Morse, W. B. 1957. The conservation officer, a personnel challenge. Proc. West. Assoc. State Game and Fish Comm. 37:63-69. 1958. Western wildlife law violators. Game and Fish Comm. 38:292-295. Proc. West. Assoc. State 1963. The new look in wildlife law enforcement. Assoc. State Game and Fish Comm. 43:287-294. Proc. West. 1968. Wildlife law enforcement Game and Fish Comm. 48:683-694. - 1968. Proc. West. Assoc. State 1972. Wildlife law enforcement Game and Fish Comm. 52:118-137. - 1972. Proc. West. Assoc. State 1973. Law enforcement Bull. 1:39-44. - one-third of the triangle. 1976. Wildlife law enforcement Game and Fish Comm. 60~162-180. Wildl. Soc. - 1976. Proc. West. Assoc. State 1980. Wildlife law enforcement - 1980. Fish and Wildlife Comm. 60:162-180. Proc. West. Assoc. State 1982. Sidearms policy and assaults on wildlife law enforcement officers. Ann. Meeting, Midwest Fish and Game Law Enf. Off. Assoc. Steamboat Springs, Colo. 6pp. �3. Oates, D. W., C. W. Brown, and D. L. Weigel. fication of selected birds and mammals. Comm., Fed. Aid Proj. W-38-R. 91pp. 1974. Blood and tissue identiPart I. Nebr. Game and Parks Purol, D. A. 1982a. Field estimating the legality of harvested Mich. Dep. Nat. Resour., Law Enf. Div. Rep. 4. 43pp. deer. 1982b. Recreational trespass enforcement in southern Michigan. Mich. Dep~ Nat. Resour., Law Enf. Div. Rep. 2. 43pp. , and T. J. Fournier. ---as a tool of enforcement 1979. A review of the arrest: contact ratio using prosecution data from the 1977 Michigan Mich. Dep. Nat. Resour;, Law Enf. Div. Rep. 1. 25pp. deer hunting seasons. --~":"" , and M. M. Zambetis. 1982. Enforcement of Michigan's hunting license revocations. Mich. Dep. Nat. Resour., Law Enf. Div. Rep. 3. 19pp. Pursley, D. 1977a. Illegal big game harvest during closed season. Mexico Dep. Game and Fish, Santa Fe. 5pp. New 1977b. Reducing illegal harvest of trout in streams. New Mexico Dep. Game and Fish, Santa Fe, Fed. Aid Proj. F-22-R-18. 27pp. Ritter, A. F. 1975. Objectives and performance criteria for state wildlife law enforcement agencies. M.S. Thesis} VPI & SU, Blacksburg. 198pp. Vilkitis, J. R. 1968. Characteristics of big game violators and extent of their activity in Idaho. M.S. Thesis. Univ. Idaho, Moscow. 202pp. Vaught, J. R. 1975. Wildlife law enforcement research in New Mexico. Proc. West. Assoc. State Game and Fish Comm. 55:152-162. , and F. L. Turner. ---and wildlife officer 1975. Wildlife officer district evaluation district law enforcement survey. New Mexico Dep. Game and Fish, Fed. Aid Proj. FW-15-R. 37pp. Wilson, M. (Editor). 1977. Bibliography of forensic science in wildlife enforcement. Alberta Recreation, Parks and Wildl., Fish and Wildl. Div. 171pp. Woolf, A., and J. L. Roseberry. No date. Forensic science studies: application of techniques to estimate the post-mortem interval of white-tailed deer in Illinois. Final rep , , 111. Dep. Conse rv. , Div. Law Enf. 10pp. " ,~ / /"!.~ (l 9 _''-., ».( ('I Ue~/J/ Prepared by,' i ' / John F. Smeltzer ~~ildlife Researcher ~lv'::::cr, /~ ' law �APPENDIX A 34 ACTIVITY Activity Code 10 14 18 22 26 42 46 50 54 58 62 66 70 74 7 10. DEFINITIONS Activity AND CODES Name Wildlife Inventory Wildlife Regulation Recommendation Public Relations Hunter Safety Research Game Damage and Animal Control Propagating and Distributing Wildlife law Enforcement Habitat Development and Maintenance Public Facilities Development and Maintenance Division Equipment and Facilities Maintenance Environmental Protection Resource Reconnaissance Planning and Budgeting Real Estate Administratio~ and Clerical Services In-Service Training leave Wildl ife Inventory Any work intended to provide information about the number, composition or distribution of wildlife, wildlife habitat, or publ ic use of wildlife. 14. Wildlife Regulation Recommendation Any part of the process of translating inventory recommendations for season regulations. 18. information into Publ ic Relations Any work done to inform, educate or assist the publ ic with the intent of increasing publ ic understanding or use of a wildlife resource. 22. Hunter Safety Any work performed with the intention of increasing fied hunter safety students. 26. Research Any work necessary project. 30. the number of certi- to accomplish the objectives of an approved research Game Damage and Animal Control Any work intended to control animals which are a nuisance or a danger to people or to properties and any investigation or processing of wildlife damage claims. �35 Appendix A (continued) 34. Propagating and Distributing Wildlife Any work intended to supplement or introduce wildlife by releasing captive animals into the wild. This category includes the capturing and caring for animals which are to be released for this purpose. 38. Law Enforcement Any work intended to detect violations action. 42. Habitat Development Any work 46. to legal and Maintenance intended to enhance Publ ic Facilities and bring the violators Development the ability of habitat to support wildl ife. and Maintenance Any work intended to result in the construction or upkeep of facil ities designed to provide comfort or convenience for wildlife users. 50. Division Equipment and Facilities Maintenance Any work intended to result in the construction or upkeep of facil ities or equipment used by Division personnel in their daily activities. 54. Environmental Protection Any work intended to minimize the adverse effects of land or water development on wildlife or their habitats. 58. Resource Reconnaissance Any work intended to famil iarize an individual wHh of responsibility. 62. his geographic area Planning and Budgeting Any work intended to influence the final form of a Division Strategic Plan, Operation Plan or budget document. 66. Real Estate Any work intended to result in the transfer of some degree of surface control of land to the Division. 70. Administration and Clerical Services Any work where the primary intention is to increase the efficiency or effectiveness of other Division personnel. Work in this category will result in some form of written or oral communication. Task force assignments are included in this category. 74. In-Service Training Any formally scheduled effort to improve the knowledge Division employees. or skills of �36 Ap endix A (continued) 78. Leave Any regularly scheduled working day when an employee is not on duty, except for holidays, weekends and compensatory time, should be recorded in this category. This includes all paid leave (annual, sick, funeral, etc.). Hours should be calculated at the rate of ~ hours per working day. �37 APPENDIX B DISTRICT WILDLIFE (Statutory Title: Wildlife MANAGER Conservation Officer) NATURE OF WORK This is a multiple range class with a broad range of duties and responsibil ities from the entry trainee through the fully operating journeyman level District Wildl ife Manager. This position requires the exercise of independent judgment, professional skills and abilities and technical expertise while engaged in wildlife management, including law enforcement, wildl ife inventories, population analysis, habitat maintenance and improvement, environments impact assessments, information and education, and a variety of additional wildlife-oriented responsibilities. RANGE A This is the beginning professional trainee level with formalized on-the-job classroom and field training under continuous supervision and control. The employee becomes knowledgeable of the Division's operation, appl ication of the State's laws, rules, and regulations, and techniques of wildlife resource management. RANGE B This range is a continuation of the developmental concept. This range requires less supervision and the individual is assigned a district or area of responsibil ity and is given more latitude to gain confidence and experience necessary to perform at the journeyman level. Although assignments given at this level are broader and more in-depth, the employee is still in a developmental stage. RANGE C This range identifies the fully functioning journeyman, full scope of assignment, usually non-supervisory level. A person of this level has full responsibility for all wildlife and related management practices and enforcement of wildlife laws in an assigned district. This level receives general supervision from an area supervisor but the district programs and how well they are managed and how effective they are is essentially the employee responsibil ity. SOME EXAMPLES OF WORK Initiates and conducts census and interprets other data. Initiates and performs complex wildlife management wildlife research studies. studies; assists in complex Initiates and assists in development of game management areas related to share crop, grazing leases,and contract farming arrangements, in addition to controlling and developing state-owned lands. �~u Appendix G (continued) DISTRICT WILDLIFE Page 2. MA~AGER (Con't) Develops district objectives and plans to achieve state goals and prepares budget estimates to achieve set goals. Develops technical wildlife material and writes articles for departmental and public use. Attends and participates in professional conferences and meetings and reviews literature to improve knowledge in the field of wildlife management. Performs liaison, coordinative dictions and other entitites. and contact work with various agencies, juris- Works with radio, television and press media, sportsmen, public schools, recreational ists,and rancher organizations and other groups in developing and promoting understanding and support of department policies and wildl ife management objectives. . Develops and supervises hunter safety programs within an assigned district. Conducts aerial and ground census trends, trappings, tagging, and banding studies; introduces and transplants game, nongame, furbearers, birds, and fish for experimental or management purposes and writes summary and conclusions. Conducts age-growth, length-weight, migration, distribution, food and life history studies; measure depth and surface areas of water, water flow, and the basic chemical composition of water. Conducts ecological surveys; investigates and assists in determining feasibil ity of planting of warm and cold water fish in streams, lakes, and rivers in the state. Participates in experiments with various feeds and rations of feeds, either in a laboratory or in the field. Responsible for initiating and planning of exclosures ranges to determine use of forage. and enclosures on game Performs field reconnaissance and planning for wildl ife habitat improvements; gathers information for mapping seasonal ranges and routes of migration of various wildlife. Assists in the planning, preparation program. Assists in check station operation of various game species. and planting of the annual fish planting for tabulation of wildlife harvest Investigates wildlife damage claims, plans and initiates actions to alleviate the damage,and makes recommendations on possible restitution; coordinates distribution on preventative materials within an assigned district. Investigates non-damage wildlife harassment and initiates actions on how such harassment can be alleviated. �Appendix 39 B (continued) DISTRICT WILDLIFE Page 3. MANAGER (Conlt) Coordinates with staff specialists district programs and projects. Participates Conservation professional and other conservation officers in the training of and supervIsion and evaluation Officer Als. Supervises subordinate professional employees. Acts for and represents the Area Wildlife Supervisor in of Wildlife and non- upon assignment. Initiates and conducts investigation of violations involving Division of Wildlife and Division of Parks and Outdoor Recreation laws and regulations, including federal statutes. Prosecutes said violator in courts of law. Initiate recommendation for state statute sion" regulation improvement. improvement and "Wildlife Commis- Responsible for the implementation and coordination within a given county or counties of the land and water use law. (Examples, HB 1041 and SB 97). Assists or initiates acquisition district. of wildlife habitat within an assigned Initiates and performs environmental analysis, participates in writing completion of environmental impact statements and Environmental Assessment Reports, i.e., highway projects, water development, coal and oi I shale, land reclamation, etc. Must review, assign and investigate all land use on private lands, responding in the preparation of technically written reports and effectively express orally, the interpretation of these technical reports to County Planning and Zoning Commissioners, County Commissioners, and Developers. Selects rivers, tributaries, and natural lakes to determine scientifically, minimum stream flows and lake levels to legally avoid dewatering by other users and to protect the natural environment and aquatic ecology. Generates public support and is contributing witness to various water boards and water courts. Performs related duties as assigned or required. KNOWLEDGES, SKILLS AND ABILITIES Knowledge of state, federal, wildlife,and outdoor modern law enforcement procedures and techniques. Knowledge of theories, principles,and of land development practices. Knowledge of stream and lake biological procedures. Knowledge techniques recreational of wildlife laws and management and surveying and sampl ing methods and of aquatic ecology and invertebrate zoology. �Appendix B (continued) DISTRICT WILDLIFE MANAGER (Con't) Page 4. Knowledge of microbiology, to wildlife management. and organic and inorganic chemistry Knowledge species. of the habits and ecology of wildlife Knowledge of related federal, including game and nongame state, and local cooperative Knowledge of the relationships of wildlife resource conservation concepts. conservation Knowledge all outdoor Skill of firearms, outdoorsmanship,and in the use of laboratory Skill and ability and happenings. in relating as they relate programs. to total natural recreation functions. and field equipment. in-depth observations of wildl ife incidents Abil ity to observe and classify wildlife from aircraft, fixed-wing hel icopters and also from the ground under varying conditions. Abil ity to complete complex wildlife management and biology assignments. Abil ity to analyze data and apply relevant wildlife the solution of problems. Abil ity to express oneself clearly and concisely, Abil ity to establ ish and maintain effective courts, district attorneys, law enforcement Ability to enforce wildlife rights of others. biologic principles to both orally and in writing. working relationships with officials, and the publ ic. laws firmly and tactfully, Ability to make independent decisions determined course of action. and with respect for the and act quickly and decisively on the Ability to create public awareness of and involvement in wildlife commission's programs and objectives and to provide leadership to the publ ic in this activity. Abil ity to work under stress and strain for prolonged Abil ity to use and maintain MINIMUM Education PREPARATION and extended periods. issued equipment. FOR WORK and Experience Graduation from an accredited college or university with a Bachelor's degree in Wildlife Biology, Biology or related field. �Appendix B (continued) DISTRICT WILDLIFE MANAGER Page 5. 41 (Con't) Necessary Special Requirement Valid Colorado Driver's License at time of appointment. NOTE: Employees are not adjusted to the "B" range until they have successfully completed a one-year training program and to the "C" range until they have completed one year at the "B" range with satisfactory performance or above, unless otherwise approved by the Division of Wildlife Director. �42 APPENDIX AREA WILDLIFE C SUPERVISOR NATURE OF WORK This is professional supervisory and administrative life management and lands program. work in the state wild- Under general direction of an assistant regional manager, plans, organizes, supervises, coordinates, and participates in wildlife management and enforcement activities, environmental impact programs and budget preparation for an assigned regional geographic area, or performs a wide variety of staff coordination functions in a regional office such as regional law enforcement coordination or similar staff positions within a wildlife region. Area wildlife supervisors assigned to an area within a wildl ife region must supervise three or more District Wildlife Managers and may also supervise Wildlife Technicians. Staff coordinator positions in a region must have regionwide responsibility for planning, implementing, evaluating, and coordinating a major wildl ife activity such as law enforcement or a comparable wildlife activity. SOME EXAMPLES OF WORK Plans, organizes, supervises, coordinates and participates in a wide variety of wildl ife management and enforcement activities, within an assigned geographic area. Consol idates and evaluates findings prepares management and enforcement supervised. and reports of subordinate personnel, and and other recommendations for the area Compiles a variety of reports on wildlife population and game land and environment conditions; develops program changes and management plans for present and future needs. Plans and inspects construction and maintenance projects, wildl ife improvement projects, and use of state game lands and private lands under state agreement or contract. Reviews reports and performs investigations in especially situations or litigation re9ulting from arrests, appeals, complex or unusual or hunting accidents. Promotes publ ic relations by preparing new releases, giving sl ide lectures, appearing on radio and television, preparing exhibits, and developing information for District Wildl ife Managers. Establ ishes and maintains working relationships with federal and local governmental agencies and other state agencies and provides assistance to other divisional programs on request. �Appendix C (continued) AREA WILDLIFE Page 2. Coordinates 43 SUPERVISOR and supervises (Con It) the areals hunter safety programs and schools. Consolidates area reports and recommendations and assists the regional manager in making evaluations, correlations,and regional consol idations for consideration and action at the Division and higher level. Coordinates the area's wildlife management and enforcement activities, publ ic use of Division properties, and area environmental issuei and projects. Assists with budget preparation, performs business management prepares a variety of reports and correspondence. duties and Coordinates the assignment of regional staff personnel, both professional and support, within and between areas and on special assignments. May serve as a regional staff coordinator for specialized programs such as regional law enforcement coordinator: Plans, implements, monitors, and coordinates regional law enforcement" activities; conducts and/or supervises special investigations of major and complex cases of suspected violation of wildl ife statutes; plans, conducts, evaluates, and coordinates law enforcement training programs for regional staff; conducts publ ic education and information programs; serves as regional hearings officer for license suspension hearings; performs related staff coordination activities. Performs related work as assigned or required. KNOWLEDGES, SKILLS AND ABILITIES Thorough knowledge of state wildlife laws and modern law enforcement procedures; the impact on the environment of natural resources development projects and the incidence of and problems inherent in publ ic use of Division properties. Thorough knowledge of wildlife management, control principles and practices. Considerable knowledge of federal, state,and related to wildlife management. land development,and local cooperative Considerable knowledge of the practices and objectives relations program. environmental programs of an effective Knowledge of modern administrative practices and procedures, including budgeting, supervision, management, planning,and organization. Skil I in communicating effectively, both orally and in writing. publ ic �44 Appendix C (continued) AREA WILDLIFE Page 3. SUPERVISOR (Con't) Abil ity to prepare a variety of reports and to keep personnel changes and additions to procedures and programs. informed of Skill and ability in establishing and maintaining effective working relationships with federal, state, and local officials, fellow employees and the publ ic. MINIMUM Education PREPARATION FOR WORK and Experience Graduation from an accredited college or university with an appropriate Bachelor's degree and four years of experience as or at the level of a District Wildlife Manager or above. The required experience may be in wildlife management or wildlife research and must have included enforcement and administrative responsibilities. Enforcement being defined as enforcing appl icable wildl ife statutes. Administrative responsibilities must have included developing and implementing work plans, preparing budget requests, providing input for operating and policy decisions. �WILDLIFE LAW ENFORCEMENT ACTIVITIES » APPENDIX D CONDUCTED BY THE COLORADO -0 -0 DIVISION (l) OF WILDLIFE ::J 0.. 1. Enforcement of smal I game regulations/statutes 2. Enforcement of upland game regulations/statutes 3. Enforcement of waterfowl regulations/statutes 4. Enforcement of big game regulations/statutes 5. Enforcement of fishing regulations/statutes 6. Enforcement of nongame regulations/statutes 7. 8. Enforcement of trespass Enforcement regulations/statutes of trapping regulations/statutes 29. Operation Game Thief implementation (OGT) coordination stake-outs/non-uniformed 30. Enforcement 31. Assist other Law Enforcement Agencies 32. Installation and maintenance of strategic 33. Distribution of regulations, brochures 34. Inspect and seal beaver, 35. Issue replacement safety cards 36. Hunter safety education programs bobcat, licenses, 9. Enforcement of road or area closures 10. Enforcement of safety 11. Enforcement of raptor regulations/statutes 37. Donate wildl ife 12. Enforcement regulations of other state/federal as required 38. Regulations/Statute 39. Planning 40. Maintain 41. Design and distribute 42. Develop 43. Organize 44. Serve warrants, 45. Deliver 46. Plan and conduct district, interstate meetings 13. Investigation 14. Conduct 15. Random enforcement 16. Non-random 17. Report writing .18. Meetings regulations/statutes laws or of wi Idl ife violations Iicense agent audits patrol or directed patrol with DA or County Attorney review and rewrites records forms and review Standard and conduct Operating check station assist in warrant 21. Respond to nuisance animal complaints 47. Night 22. Respond to citizens complaints 48. Phone call response 23. Area/Regional/Statewide 49. Suspension 24. Publ ic relations: 50. Participate 25. Law Enforcement training (participation) 51. Enforce game damage 26. Law Enforcement training (organization) 52. Law enforcement 27. Covert operations 53. Monitor law enforcement plan 28. Radio communications 54. Special law enforcement assignments 55. Coordinate & implementation Procedures (SOP) operations service tickets for docketing Case report reviews coordination tags, hunter and other publ ic education Court preparation day to day and scheduled LE signs bear permits, 19. meetings o patrol 20. and appearance X & law enforcement hearings, area, regional, patrol to publ ic inquiries review, in regional recommendations law enforcement associations statutes data collection pick-up of road kills .r:- \_;., �46 APPENDIX NON-STANDARD E DATA COLLECTION 1. Seizure Tag and Donation Certificate 2. License Agent Audit Summary - Regional 3. Warrant FORMS Record Records 4. Monthly Activity Reports 5. Regional L.E. Summary Reports 6. Monthly Report of Confiscated Game and Fish 7. Undercover Vehicle Use Form 8. License Agent Audit Form, Checklist 9. Residency 10. Training 11. Fisherman 12. Chain of Evidence/Evidence 13. Stolen License List 14. License 15. License Agent Investigation Form Report Count Form Cards Issuance Quality Report Inspection Summary �47 APPENDIX F STANDARD DATA COLLECTION 1. 2. 3. 4. 5. 6. 7. B. 9. 10. 11. 12. 13. 14. 15. 16. 17. lB. 19. 20. 21. 22. 23. 24. 25. 26. 27. 2B. 29. 30. 31. 32. 33. 34. 35. 36. 37. 3B. 39. 40. FORMS Lake/Wildlife/Park/Shooting Area/Commercial Hunting Appl ication Aerial/Motor Vehicle/Snowmobile Hunting Permit Application Compulsory Bighorn Sheep Check Incident Report, "Use of Force" Uniform Hunting Accident Report (NRA Form) Warrant Service Request (DOW-M-F-17) Special Wildlife Commission Application OGT Informant Interview Evidence Seizure Tage CDOW Donation Certificate (M-F-23~Bl) Disposition Form Compulsory Mountain Lion Check Duplicate License Affidavit Replacement Appl ication - Lost Hunter Safety Certificates Investigations Log CDOW Request for Wildlife Laboratory Examination Animal-Vehicle Kill Report Voluntary Statement (DOW-R-F-15) Court Transcript Record (DOW-M-F-21-B1) Intelligence Report (CDOW) Report of Investigation (CDOW) Continuation Sheet (CDOW) CBI Request for Laboratory Examination (CBI L 3-BOX) Check Station Report Form Appearance, Written Plea of Guilty and Waiver Case Report Daily Activity Reports OGT Log and Disposition Cold water fishing Contact Form Warm water fishing Contact Form Rec. Vehicle/Boat/Snowmobile Contact Form Hunting Contact Form Snowmobile Accident Report Boating Accident Report Penalty Assessment/Summons Form Dupl icate Carcass Tag License Copies Gift Affidavit PA Cash Receipt Chain of Evidence Envelopes �48 APPENDIX 1982-83 SUMMARY OF IN G STATE/PROVINCIAL RESEARCH/EVALUATION WILDLIFE LAW EFFORTS ENFORCEMENT by John Colorado F. Smeltzer Division of \>/ildlife Law Enforcement Research Wildlife Research Center 317 West Prospect Road Fort ColI ins, Colorado 80526 March 1983 �Appendix G (continued) TABLE PART I OF CONTENTS State/Provincial Summary of Research/ Evaluation Efforts - a narrative summary PART II ..... Responses to survey question: \.Jhatare the most serious problems facing wildlife law enforcement today? • . . . . . . . . . . . . 7 PART III .... Responses to survey question: What are the needs for wildlife law enforcement research? 9 PART IV ..... Names and addresses of contact persons throughout the U.S. and Canad~ who are most famil iar with their attempts to evaluate or research wildlife law enforcement problems. . . . . . . . . . .. 10 �..JU Appendix PART I G (continued) STATE/PROVINCIAL OF RESEARCH/EVALUAT SUMMARY Iou EFFORTS The fol lowing is a narrative summary of the information provided in response to the Colorado Wildlife Law Enforcement Research/Evaluation Survey. is not intended as a comprehensive summary of al I but simply as a source of information to responding agencies. It is based only on the information suppl ied. Alabama: a.) Special Task b.) Investigating Force development possibilities of covert a.) Computerization Arizona: a.} 10-year research plan being developed which will include wildl ife law enforcement research 1 person assigned to coordinate Wildl ife L.E. research plan Arkansas: Alberta: Cal ifornia: Colorado: No research/evaluation efforts data, operations Alaska: b.) of violation It violations/contact reported a.) Hunter attitudes toward wildlife laws and wildl ife officers in Alberta b.) Factors associated with wildl ife law violations in Alberta c.) The use of aircraft in wildl ife law enforcement d.) Compendium of Alberta Fish and Wildlife District W9rk Analysis Status 1980-81 e.) 1982 -- Helicopter patrol program llov. 1-30, 1982 f.) A selected annotated bibl iography of literature on wildl ife enforcement g.) Permanent personnel are assigned to Wildl ife L.E. research/evaluation No report a.)Computerized monthly and annual violation, court case, dismissed, and fines report b.) Wildlife Law Enfor~ement Plan -- 1981-84. Developed and being implemented. c.) Has wildlife law enforcement researcher d.) Draft study plans for the follow~ng: 11 legal license purchase evaluation -- Development of measurable enforcement objectives -- Public attitude survey e.} Developing a liS-yr. Operations Plan forWildlife Law Enforcement Research" F.) Using select co-l leqe students to quantify historic law enforcement info rma t ion. �Appendix G (continued) 51 Connecticut: No research/evaluation activities reported Delaware: No research/evaluation activities reported Florida: a.) A practical field method for blood and tissue identification b.) Intermediate non-lethal weapons for Florida wildlife officers c.} Decriminalization and recodification of Florida's wildl ife coCe d.) A three-year plan to curtail the commercialization of certain rare Florida wildl ife e.) What is enough wildlife law enforcement f.) The scope of the wildl ife trade in the United States g.) Development of a statewide citizen crime patrol reporting project (\.Jildlife alert) and impact h.) New approaches toward wildlife crime patrol i.) General orders manual for wildl ife law enforcement j.) Wildl ife officer productivity computer printouts k.) Spec ia I ized so I ut ions to a numbe r of wi I d life law enforcemen t prob Iems I.) Undercover investigations program m.) Accountabil ity-wildl ife enforcement goal for the 1980s Georg ia: No report Haws i i: No research/evaluation review measures which Idaho: a.) b.) c.) d.) III i no is: a.) b.) c.) d.) e.) f.) g.) h.) i.} Indiana: activities reported. Do have were not elaborated on. in-house Vilkitis poaching study Evaluation of purchase of resident I icenses by non-residents Computerized violation, I icense, and suspension' files Currently adding one person to staff to be responsible for wildlife law enforcement research projects Time of death studies - deer Blood identification test kits for deer Sonar research Lead and copper test kits Identification of illegal furs using scanning electron microscope (recommend discontinue) Long distance hearing devices Signal transmitters on illegal devices and commercial wildl ife Muscle tissue/parasite determination Post-Mortem vitreous humor potassium levels a.) Developed an extensive Standard b.} Coded violations c.) Coded activities by category Operating Procedures Manual �Appendix G (continued) Iowa: a.) Evaluation of wildl ife enforcement officer b.) Use of aircraft to detect spotl ighters Kansas: a.) Evaluate effectiveness of uniformed vs. non-uniformed personnel b.) Evaluate effectiveness of aircraft in aiding detection of spotlighters c.) Computerized boating registration files d.) Computerized activity analysis Kentucky: No report Louisiana: No research/evaluation Maine: a.) b.) c.) d.) reported Computerized officer daily activity reports Computerized prosecution reports Computerized complaint file Have looked at illegal purchase of resident by non-residents Manitoba: Recently Maryland: No research/evaluation Massachusetts: efforts replicated Vilkitis productivity poaching activities study licenses -- no result released reported No report Michigan: a.) An evaluation of the Michigan effort b.) Work on compliance estimation c.) Estimation of poaching levels wildlife law enforcement Minnesota: No report Mississippi: a.) Officer performance and operational cost analysis b.) Computerized citation files -- coded violation forms species, county, violation by: yet �Appendix G (continued) ~lissouri: a.) b.) c.) d.) e.) Montana: No report Nebraska: No research activities reported, doing substantial forensic work Nev ada : No research/evaluation activities reported Nev Jersey: No research/evaluation activities reported New Mexico: a.) b.) c.) d.) e.) f.) g.) h.) New York: Has wi Idlife law enforcement researcher Deer poaching and poacher study Lead detection analysis Meats and blood identification Computerization of arrest reports and agents monthly activity reports f.) Potassium levels in vitreous humor for time of death analysis - deer g.) Developed operation game thief program similar to New Mexico h.) Covert operations however, Oates has been Illegal harvest evaluation of warmwater game fish Illegal harvest of big game during closed season Reducing illegal harvest of trout in streams Wildlife officer district evaluation and w Ild l ife lawenforcement Wildlife Law Enforcement research in New Mexico Implication of illegal harvest on deer management Remote sensing equipment and radio telemetry testing Evaluation of illegal purchase of resident I icenses by non-residents a.) Computerization of violation information b.) Development of activity codes c.) Establ ishing objectives North Carol ina: No research/evaluation efforts reported North Dakota: No research/evaluation efforts reported Ohio: No research/evaluation activities Oklahoma: No report Ontario: Dave reported a.) Development of regional and district law enforcement plans b.) Development of a uniform officer reporting system and standard incident reporting system c.) Development of data base information on contacts, warnings, charges, t i me analysis, mi les driven, etc. d.) Draft proposal -- ~Improving Enforcement Efficiency" -- by Outdoor Recrea t ion Group. 1981-0}:"06. e.) Report on the "Role and Responsibi I ities of the Conservation Officer" s u rve �54 Appendix G (continued) Oregon: ~Jo report Pennsylvania: No research/evaluation Rhode No report Island: activities reported Quebec: a.) lden t lf i ca t i on o f v••r i ld bird blood, feather ana bone b.) Identification of deer and moose meat c.) Computer analysisofwildlife Ia ",I enforcement effort -- possible report to be given to mid-west L.E. meeting, Springfield, Illinois in June 1983 Saskatchewan: Canadian Saskatchewan: Tourism Wildl ife Service and Renewable a.) Computerized b.) Computerized analysis South Carol ina: South Dakota: Tennessee: Texas: Vermont activities reported Resources prosecution report work management report -- time/activity No current research activities report. Have been involved with research activities conducted through Virginia Polytechnic Institute. a.) Considering electronic surveil lance devices for back road use b.) Documented violations to establ ish projected/actual losses, and periods of highest illegal activity c.) Use of news media to· influence public attitudes towards poaching d.) Covert operations a.) Study completed on "High visibility versus vehicles in wildlife law enforcement" b.) Participated in VPI studies for 3 years c.) No additional research activities currently a.) b.) c.) d.) e.) Utah: - No research a.) b.) c.) d.) e.) low visibil ity planned Development of Statewide Manning Standards and Assiqnments Program. Deployment based on population density, type and area of patrol (land and we t e r ) , local hunting/fishing activity De~elopment of vi6lation trend charts and tables Evaluation of reorganization in 1975 Development of field contact survey program (FY-78) Development of "Wildl ife Law Enforcement Division Operational Plan" FY 1983. This is a 58-page plan including: work load summaries, 1982 accompl ishments and 1983 objectives for numerous wildl ife law enforcement activities. K-9 use in ~Ildlife law enforcement Effectiveness of t-wo-man vs. one-man patrol Effectiveness of various patrol techn i nue s Effectiveness of preventative law enforcement Time of death studies No report methods �Appendi~ G (continued) Virginia: Virginia: a.) Have b.) Some been involved I imited covert with research operations activities at VPI Virginia Polytechnic Institute and State University S.U.) Studies -- supervised by Dr. Robert H. Giles, (V.P. I. Jr. a.) Objectives and performance criteria Lor state w i ld l i f e law enforcement agencies b.) Analysis of the spatial and temporal occurrence of deer spotlighting violations in Virginia c.) Optimum deployment of wildl ife law enforcement agents: a problem analysis d.) A review and appraisal of crime load, workload, and manpower standards in wildl ife law enforcement e.) An initial bibliography of wildl ife law enforcement f.) Estimating illegal kill of deer g.) The influence of game laws and regulations on hunting satisfaction h.) Warnings vs. citations in wildl ife law enforcement i.) A survey of wildl ife law enforcement research needs and current research j.) Anti-poaching campaigns -- 3 tool of wildl ife law enforce~2nt? k.) Quasi - experiments, multiole indicators, and enforcement effectiveness 1.) Fines in wildl ife law enforcement ~.) An analysis of nationwide wi ldl ife law enforcement data n.) Relative importance of enforcement objectives. and seriousness of violations in relation to objectives 0.) Wildl ife law enforcement research - the context and the needs p.) Alpha - man: A theory of wildl ife law violation q.) Wildlife law enforcement r.) Limitations of wi ldl ife law enforcement compliance estimators s.) Dynamic deployment of wildl ife la0 enforcement manpower a decision aid Washington: West Virginia: Wisconsin: \Iyomi ng: a.) Regional workload assessment program b.) Development of data base information a.) No research activities reported b.) Do have standardized complaint forms No report a.) Computerized arrest records by violations reported by citizens, observed by officers, and by arrests from reports or observations . .b.} Developed case file referral system to keep track of current investigations and time spent per case c.} Have a stop-poaching program d.) District violation summaries (1978) e.} A study of user attitudes and violations f.) Has personnel assigned to wildl ife law enforcement research/ eva Iua t ion �Appendix PART II G (continued) Table 1. Survey serious problems a What are the most responses to the question : facinq wildl ife law enforcement today? ResDonse 1. 2. 3. 6. 7. 8. ..I • (26) aTwenty-six Other responses each) . Some of reSDonses 12 11 Funding problems - lack of funds Poaching, commercial poaching, marketing Lack of personnel Publ ic attitudes More people - increasing number of users Inabil ity to evaluate enforcement effort Program administration problems Civil suits - lawsuits against officers Spotl ighting 4. 5. a !l States/Provinces 2 responded to this problems. Habitat loss Inadequate detection rate Need to educate publ ic Low sa Ia r ies Lack of equipment Native hunting and fishing privileges Trespass problems Increasing wildl ife law enforcement responsibil of complete these 2 2 additional examples suggested 9 5 4 3 responses "We are seeing more his/her duties." against "We need to educate the hunting and responsibil ity." finding more a t tit u de " lack of criteria once ities and our officers fishing in the performance publ ic in ethical of conduct public attitude and the inability to prove activities in the eyes of the public and sophisticated law violators." the pub I ic - - the s ilen t ma j 0 r itY .' " 0f only s s !" "Our biggest problem is poor the worth of law enforcement the legislature." "We are (Mentioned are: lawsuits "H mv to do mo re, wit hIe question. I for officer evaluation." "Commercialization of w i ld l i f e resources, is our most serious problem." fish, game and non-game �Append~x ~ lcontlnued) "1 ack of money and "More people, "Program gaining "We are drugs, personne that's administration, issues, cornp seeing theft, "Trespassing relationships." " ... a serious for enforcing our , increased recreational on posted 1 to handle increasingly sophisticated vialiJtor problem." including time accrual, enforcement vehicles lands misunderstanding those laws." ... budgeting problems, collective barenforcement coverage, and producti'/ity." involvement in peripheral to name a fe\·/." resulting of the in bad laws and areas; landmmer/hunter the agency responsible ;." �_"V Append i x 11 (con t i nued) PART II I Table 2. Survey responses wildl iFe law enforcement to the questiona: research? What are the needs of res::;onses Response 1. 2. for 5 Evaluate manpower needs - allocation How much funding is needed? Undercover - Covert teams Develop comoliance estimates Measure wildlife commercial ization - poaching Study ways to educate the publ ic 3. 4. 5. 6. (17) aSeventeen Other responses once each) states/provinces suggested these responded additional are some examples of complete " ... to be effect ive research "\-/e must equipment question. ideas: (mentioned of enforcemen: have an undercover develop better, specialized w i t h limited budgets." equipment program." or obtain " .. , how we are going to establ ish an enforcement ooDit evaluate the effectiveness of an enforcement program." "Study of job ." required evaluate " to answer " " this funding poaching levels levels the question only responses: must ~'!e 2 to this Development and testing of new equipment Uniform, national records keeping system Forensic studies Develop a system to measure cost effectiveness Here 3 3 3 and the number by species "HOI'!much badly needed system to of personnel In a statistically lav. enforcement develop information to allow us to predict information wou ld save time and mileage." to do t he val i c manner." is enough?" patterns of violations, �APPENDIX PART IV 59 G NAMES AtJD ADDRESSES OF IND IV IDUALS TO CONTACT LAW ENFORCEMENT RESEARCH/EVALUATION PROGRAMS IN REFERENCE This list is not intended to be comprehensive. as a source of initial information. Follow-up some cases. TO STATE!PROV INC IAL Its purpose is only to serve contacts will be necessary in ALABAMA: Tim D. Cosby, Captain Law Enforcement Section Dept. Cons. & Nat. Resources 64 North Union Street Montgomery, Alabama 36130 FLORIDA: Lt. Colonel Brantley Goodson 620 South Meridian Room 201 - !:ryant.Bui Iding Tallahassee, Florida 32301 (904) 488-6251 ALASKA: Jim Nutgrass Fish & Game Enforcement Alaska State Patrol Anchorage, Alaska (907) 269-5535 GEORGIA: ALBERTA: Mike Melnyk, Research Officer Pro9r~m Evaluation & Res. Economics Fish & Wi Idlife Division 8th Floor - South Towe r 9915 - 108th Street Edmonton, Alberta CANADA T5K 2C9 (1;03) 427-6735 Colonel Drew ~hittaker Law Enforcement Chief Game & Fish Division Dept. of Natural Resources 270 ~ashington Street, S.~. Atlanta, Georgia 30334 (404) 656-3530 HAWAII: Maurice Matsuzaki, Enforcement Chief Dept. of Land & Nat. Resources Div. of Cons. & Research Enforcement 1151 Punchbowl Street Honolulu, Hawai i 96B13 (B08) 548-591,) IDAHO: Donald A. Nicholson Enforcement Operations Offic~r Idaho Dept. of Fish & Game 600 South Walnut, Box 25 Boise, Idaho 83707 (208) 334-3736 ILLINOIS: Tom ~akolbinger, Sergeant Division of Law Enforcement IIIinois Dept. of Conservation Lincoln Tower Plaza 524 South Second Street Springfield, III inois 62706 (217) 732-f,431 INDIANA: Major Phill ip Ohmit, Executive Law Enforcement Division 608 State Office Building Ind ianapo lis, Ind iana 46204 (317) 232-4010 I O\~A: Rick McGeough, Superintendent Law Enforcement Section Iowa Conservation Commission ~al lace State Office Building Des Moines, Iowa 50319 (515) 281-5918 KANSAS: Frank NeSmith, Chief La\1 Enforcement Kansas Fish & Game Box 54A, Rural Route 2 Pratt, Kansas 67124 (316) 672-5911 ARIZONA: ARKANSAS: CALI FORN IA: ·CONNECTICUT: COLORADO: DELA~ARE: AI LeCount, Research Star Route Tonto Basin, Arizona (602) 479-2335 Biologist 85553 W. F. Hailey, Ass't. Chief Enforcement Division 2 tlatural Resources Drive Little Rock, Arkansas 72205 (SOl) 223-6300 Ned Dollahite, Chief ~ildl ife Protection Division Cal ifornia Dept. of Fish & Game 1416 Ninth Street Sacra~ento, Cal ifornia 95814 (916) 445-3531 Frederick J. Pogmore Director of Law Enforcement State Office Building Hartford, Connecticut 06115 (203) 566-3978 John F. Smeltzer ~ildl ife Law Enforcement Researcher Wildl ife Research Center 317 West Prospect Road Fort Col Iins, Colorado BD526 (303) 484-2836, ext. 49 George ~. Stewart, Jr. Enforcement Administrator Division of Fish & Wildlife P.O. Box 1401 Dover, Delaware 19901 (302) 7%-4431 Officer �60 APPENDIX G PART IV (cont.) KENTUCKY: Steve Yontz, Director Division of Law Enforcement Dept. of Fish & Wildl. Res. #1 Game Farm Road Frankfort, Kentucky 40601 (502) 564-3400 MINNESOTA: Fredean Hammer, Director Division of Enforcement Dept. of Natural Resources 300 Centennial Building 658 Cedar Street St. Paul, Minnesota 55155 LOUISIANA: Susan K. Brittain Staff Development Special ist Enforcement Division Louisiana Dept. of Wildlife & Fisheries P. O. Box 15570 Baton Rouge, Louisiana 70895 (504) 568-5667 MISSISSIPPI: Phil ip J. Strong, Chief Law Enforcement Mississippi Dept. of Wildl ife Cons. Southport Mall, P. O. Box 451 Jackson, Mississippi 39205 (601) 961-5300 MAINE: John F. Marsh, Chief Warden Dept. Inland Fisheries & Wildl. 284 State Street State House Station 41 Augusta, Maine 04333 (207) 289-2766 Detective Richard Hennessey Augusta Regional Headquarters Dept. Inland Fisheries & Wi ldl. 8 Federal Street Augusta, Maine 04330 (207) 289-2175 MANITOBA: MARYLAND: MASSACHUSETTS: S. A. "Bud" Mcivor, Chief Field Services & Enforcement Manitoba Dept. of Natural Res. 1495 St. James Street Winnipeg, Manitoba CANADA R3H OW9 (.204) 786-9132 Ann Will iams (Computer Program) (601) 961-5313 John Given (Computer Program) (601) 961-5331 M ISSOUR I: Ron L. Glover Protection Research Special ist Missouri Dept. of Conservation Fish & Wildlife Research Center 1110 College Avenue Columbia, Missour i 65201 (314) 449-3761 MONTANA: Erwin J. Kent, Administrator L<l"'En f o rc ernen t Dept. of Fish, Wildl ife & Parks 11,20 East Sixth Helena, Mont<lna 59601 (406) 449-2452 NEBRASKA: Donald C. Schaepler, Chief t.av Enforce .ent Division Nebraska Game & Parks Commission 2200 North 33rd Street P.O. Box 30370 Lincoln, Nebr<lska 68503 (402) 464-0641 Jack T. Taylor, Deputy Supt. Dept. of Natural Resources Natural Resources Police Tawes State Office Building Annapol is, Maryland 21401 (301) 269-2248 James Jesseau Supervisor of Enforcement Dept. of Fisheries, Wi ld life & Recreational Vehicles 100 Cambridge Street ~oston, Massachusetts 02202 (617) 727-3900 Dave O<ltes, Forensic Research Nebraska Game & Parks Commission 2200 North 33rd Street P. O. Box 30370 Lincoln, Nebra~ka 68503 (204) 464-0641 NEVADA: Darrel D. Harold, Acting Chief Division of Law Enforcement Nevada Department of Wildlife 1100 Val ley Road, P. O. Box 10678 Reno, Nevada 89520 (702) 784-6214 �61 APPENDIX G PART IV (cont.) NE~I HAMPSHIRE: Major Mason S. Butterfield Chief of Law Enforcement New Hampshire Fish & Game Dept. Box 2003, 34 Bridge Street Concord, New Hampshire 03301 (603) 271-3421 NEW JERSEY: Hudson G. Amory District Conservation Officer Bureau of Law Enforcement CN 400 Trenton, New Jersey 08625 (609) 292-2965 NE\-j Harry Mikels Wildlife Law Enforcement Researcher N.M. Game & Fish Department Vi llagra Bui Iding Santa Fe, New Mexico 87503 (505) 827-2550 MEXICO: NE~I YORK: George Firth, Asst. Director Division of Law Enforcement N.Y. State Dept. of Environmental 50 Wolf Road Albany, New York 12333 (518) 457-5680 NORTH CAROLINA: Gene H. Abernethy, Chief Division of Enforcement N.C. Wildlife Resources Commission Archdale Building 512 N. Sal isbury Street Raleigh, North Carol ina 27611 (919) 733-7191 nORTH D:\KOTA: Harold H. Spitzer, Chief Law Enforcement Division State Game 0 Fish Department 2121 Lovett Avenu~ Bismarck, ~orth Dakota 58505 (701) 221,-2180 OHIO: Richard Francis Game Protection Manager Ohio Department of Nat. Resources Division of Wi ldl ife Fountain Square Columbus, Ohio 43224 (614) 466-5854 OKLAHOMA: Kenneth Van Hoozer, Chief La"1 Enforcement Dept. of Wildl ife Conservation 1801 ~orth Lincoln P. O. Box 531:65 lklahoma City, Oklahoma 73152 (405) 521-371<) ONTARIO: W. Dale Gartley Provincial Enforcement Spec. Room 2342, Whitney Block Queen sPark Toronto, Ontario CANADA M7A lW3 (416) 965-5661 I OREGON: L. R. Hyder Oregon State Police Fish & Game Enforcement Salem, Oregon 97301 PENNSYLVANIA: Edward W. Manhart, Chief Law Enforcement Division Pennsylvania Fish Commission P.O. Box 1673 Harrisburg, Pennsylvania 17120 (717) 787-2350 G. D. Kirkpatrick, Chief Division of Law Enforcement Pennsylvania Game Commission P. O. Box 1567 Harrisburg, Pennsylvania 17120 (717) 787-5743 Cons. QUEBEC: Jos-A. Saint-Pierre Canadian Wildl ife Service P. O. Box 10100 Tour Champlain, 4th Floor Ste-Foy, Quebec CANADA GIV 4H5 (418) 694-3914 RHODE ISLArID: Stephen P. Fongere, Chief Division of Law Enforcement Dept. of Environmental Mgt. 83 Park Street Providence, Rhode Island 02903 (401) 277-2284 SASKATCHEWAN: Garry Bogdon Enforcement Coordinator Western & Northern Region Canadian Wi Idl ife Service 115 Perimeter Road Saskatoon, Saskatchewan CANADA S7N ox4 (306) 665-4087 Barry F. Tether, Director Field Services Saskatchewan Tourism & Renewable Resources 3211 Albert Street Regina, Saskatchewan CANADA S4S 5W6 (306) 565-2323 �62 APPENDIX G PART IV (cont.) SOUTH CAROLINA: SOUTH DAKOTA: TENNESSEE: Pat Ryan, Director Division of Law Enforcement & Boating South Carol ina Wildlife & Marine Resources Department Rembert C. Dennis Bui Iding P. O. Box 167 Columbia, South Carol ina 29202 (803) 758-00~2 Ronald P. Catlin Law Enforcement Staff Specialist S.D. Game, Fish & Parks Sigurd Anderson Building 445 East Capitol Pierre, South Dakota 57501 (605) 773-3381 A. J. Gulley, Jr. Tennessee Wildlife Resources Agency 216 E. Penfield Street Crossville, Tennessee 38555 (800) 262-6704 TEXAS: Chester l. Burdett law Enforcement Director Texas Parks & Wildl ife Department ~200 ~mith School Road Austin, Texas 787~4 (~15) ~79-480::l, ext. 4845 UTAH: Bruce Johnson Enforcement Special ist Utah Division of Wildlife Resources )596 West North Temple Salt Lake City, Ut ah 84116 (801) 533-9333, ext. 217 VERI10NT: Roger Whitcomb Chief Gilli'e ~!arden Fish & Gilme Department Montpel ier, Vermont 05602 (802) 823-3371 VIRGINIA: Col. John H. Mclaughlin, Chief Law Enforcement Division Virginia G3me & Inland Fisheries Box 11104 Richmond, Virginia 23230 (804) 257-1000 Dr. Robert H. Gi l e s , Professor Wildl ife Manilgement V.P.I. & S.U. BI<Jcksburg, Virginia 24061 (703) 961-5910 WASH INGTOtl: Mike Shockman Wi Idlife Enforcement Department of Game 600 N. Capitol W"y I) Iymp ia, Iiashi ng ton (206) 753-5740 98504 WEST VIRGINIA: Nelson B. Shaw Law Enforcement Division Department of Natural Resources 1800 Washington Street East Charleston, West Virginia 25305 (304) 348-2783 InSCONS IN: Dona Id L. Begh in Bureau of law Enforcement Department of Natural Resources Box 7921 Madison, Wisconsin 53707 (608) 266-1115 IJYOHING: Tom Moore, Research Biologist Game & Fish Research lab Biological Science Building Box 3312 University Station Laramie, Wyoming 82701 (307) 766-63 I 3 Steve Smi th Law Enforcement Section Uyoming Game and Fish Department Cheyenne, Wyoming 82002 �Colorado Division Wildlife Research September 1984 63 (if Wildl ife Report JOB PROGRESS State of Colorado ProjectL FW-26-R-2 Work Plan Job: Job Title: Personnel: Wildl ife Law Enforcement Research 2 Illegal Purchase of Resident Period Covered: Author: REPORT 01 January - 31 December Licenses by Non-residents 1983 John F. Smeltzer J. Black, C. Braun, D. Croonquist, D. Mueller, H. Riffle, J. Sinley B. Leasure, K. Moser, ABSTRACT The illegal purchase of resident hunting and fishing licenses in Colorado is considered to be a serious source of revenue loss within the Colorado Division of Wildlife. Previous estimates placed the loss at $2,000,000 annually. This project is the first systematic one in Colorado and one of the first in the western states to address this problem using standardized sampling procedures, research design, and statistical analysis. Preliminary results from 1982 and 1983 sampling suggest an investigation rate of 8.0% per year for all deer and elk licenses sold over the counter. Pre-screening procedures being developed appear useful in identifying most potential investigations. Pre-screened licenses (17% of total) produced 72% of the investigations identified. Potential violations were initially investigated using existing criminal justice computer systems (CCIC - Colorado Cr.ime Information Computer; and NCIC - National Crime Information Computer) .. Use of these systems may streaml ine the investigative process and improve the Division's abil ity to detect and ultimately deter these violations. ��65 EVALUATION OF ILLEGAL PURCHASES OF RESIDENT LICENSES BY NON-RESIDENTS John F. Smeltzer P. N. OBJECTIVE Determine violation rates and potential Iicense revenue losses associated with the illegal purchase of resident licenses by non-qualified persons. SEGMENT OBJECTIVES 1. Develop and test procedures to evaluate the level of illegal purchase of resident licenses by non-residents in 6 license sales categories. a. Estimate the average investigative time necessary to pursue 1 investigation to the decision stage of: Wil I a citation be issued, yes or no? b. Encourage voluntary participation by field personnel in the investigative phase of the project during the pilot phase. 2. Randomly sample and computer screen sufficient licenses and/or license appl ications in each of the 6 categories to identify 100 investigations/ violations per category. 3. Assign possible Iicense purchase violations to field law enforcement personnel for follow-up investigation. All investigations will be routed through the proper chain of command. 4. Compile and analyze field investigation 5. Prepare job progress reports as they are returned. reports. MATERIALS AND METHODS Selection of License Sample Licenses were drawn from the remittance copy files of the Colorado Division of Wildlife for the 1982 and 1983 seasons. The 1982 sample consisted of 1,100 deer and 1,100 elk I icenses (Antlered only - Separate and Combined Seasons). The 1983 sample consisted of: 300 300 325 265 500 Antlered Antlered Antlered Antlered Resident deer - Combined season deer - Separate season elk - Combined season elk - Separate season Fishing �66 The 1982 I icenses were drawn randomly by groups of 100 Iicenses from the 1982 hunter survey sample. The 1983 sample was drawn systematically from the total population of 1983 licenses within that category after determining a random starting point. Pre-screening Procedures All licenses drawn were examined and divided into 2 categories, "pre-sorts" and "all others", based on a series of characteristics suggestive of possible violation. Characteristics used were: 1. No drivers license Iisted, subject 16 years old or older. 2. Out-of-State 3. License purchased drivers I icense listed. as a "Gift" license. 4. No signature. 5. Residency claimed of 1 year, 6 months or less. 6. Identification 7. Expired Colorado drivers 8. No residency 9. Mi Iitary status. card given for 1.0. purposes. license given. listed. 10. Signature by different 11. No hunter safety number given when required. These characteristics sorted". person. were coded and used to classify Post-screening licenses as "pre- Procedures All I icense purchasers were computer-screened through the use of their name and date-of-birth (DOB) and/or driver's Iicense number. Driver's Iicense files of the Colorado Department of Motor Vehicles (DMV) were checked using the CCIC system to verify proper drivers license information, to correct improper information, or to verify that an individual did not have a val id Colorado I icense. Information provided by this screening process was used to "Post-screen" the remainder of the licenses previously placed into the "all-others" category. The following characteristics were used to classify "Post-screened" licenses: �67 1. No record found through computer 2. No prior (previous) drivers 3. Out-of-State 4. Improper Iicense prefix sequence. prior drivers inquiry. license shown. license shown. 5. Date of last license update over 5 years old. 6. Expired license on file. These characteristics were coded and used to categorize a license as "Post-screened". The remaining licenses, neither pre-screened or postscreened, were set aside and were not examined further. Intensive Screening Procedure Pre-screened and post-screened licenses were re-examined and additional information on each subject was obtained, as available, using the CCICI NCIC system, metro-residential Iistings by address, directory assistance or phone directories, College or university student directories, and Department of Revenue tax information. Vehicle registrations, phone numbers, out-of-state drivers licenses, and wants or warrants were routinely checked for each subject in the pre-screen/post-screen group. The decision to investigate or not was made on the basis of the total information available. Rates of investigation, expressed as a percent per year, were determined for the pre-screen, post-screen, and total groups. Investigative Phase Intelligence reports were completed for each investigation initiated and a cover letter asking for voluntary assistance was attached to each report. Copies of each intelligence report were provided to Area Wildlife Managers (AWM's) and Regional Law Enforcement Coordinators (RLEC's) as required. RESULTS Work completed on the 1982 test sample of deer and elk licenses suggested an overall investigation rate of about 8.0% per year using the screening procedures described. Approximately 40% (163/400) of pre-screened licenses resulted in potential investigations while only 28% (61/219) of postscreened licenses resulted in potential investigations. Pre-screening in the trial study identified 73% (163/224) of all identified investigations. Post-screening identified only 61 additional investigations from the Iicenses not pre-screened (61/1,880). �68 Similar work on the 1983 sample has not been compiled. The investigation rate, based on 865 licenses examined in depth, remained at approximately 8.0% (74/865). Distributional data has not been developed due to data processing difficulties. DISCUSSION This project will continue into the next segment following approval by the Regional Staff. The 8.0% investigation rate cannot be used to general ize across al I license sales categories. Additional work is necessary to carry through with these investigations to help determine a violation rate. This project appears to have the potential to improve the Divisionis ability to detect and ultimately deter many of these violations. Wildlife Researcher v �Colorado Division of Wildlife Wildl ife Research Report September 1984 69 JOB PROGRESS State of REPORT Colorado ------~~~~--------~ FW-26-R-2 Project Work Plan Preparation Job Title: Enforcement Job: _...:...__ Wildlife Law Enforcement Research 3 __:;."..__ of an Annotated Bibliography for Wild] ife Law Personnel Period Covered: January - 31 December 1983 Author: John F. Smeltzer Personnel: J. Black, C. Braun, J. Dennis, M. Hershcopf, N. McEwen ABSTRACT The development and distribution of an annotated bib! iography for wildlife law enforcement is viewed as one way to familiarize field personnel and wildlife law enforcement administrators with the literature available on wildl ife law enforcement. Completion and distribution of the bibliography is projected for 30 June 1985. ��71 PREPARATION OF AN ANNOTATED BIBLIOGRAPHY FOR WILDLIFE LAW ENFORCEMENT PERSONNEL John F. Smeltzer P. N. OBJECTIVE Develop and distribute a comprehensive annotated bibliography on wildlife law enforcement for use by field personnel, administrators, and researchers. SEGMENT OBJECTIVES 1. Del ineate the scope of annotated 2. Review I iterature, law enforcement. 3. Prepare a draft annotated collect, bibliography. and annotate bibliography MATERIALS articles related to wildl ife on wildl ife law enforcement. AND METHODS Wildlife law enforcement articles, both popular and technical, were collected using manual and computer-assisted searches of journals, magazines, federal aid reports, dissertations, theses, books, and other miscellaneous sources. Computer-assisted searches were conducted through the Dialog Information Retrieval Service, California and through the Fish and Wildlife Reference Service. Articles were computer-selected based on the following descriptors: wildlife law enforcement, wildlife laws, poaching, illegal harvest, fish, fishing, and regulations. Descriptors were combined in different ways using Boolean logic commands where possible to maximize search capabilities and minimize unnecessary output. Manual searches were conducted using the resources of the Colorado State University Library, the Colorado Division of Wildl ife Research Library, and extensive use of Inter-l ibrary requests. RESULTS AND DISCUSSION Approximately 575 wildlife law enforcement related publications have been collected. Over 150 have been selected for inclusion in the final report. These have been read, summarized, and annotated. Each annotation is approximately 2 paragraphs long. Annotations include the highlights of each paper and any concluding remarks. The anticipated completion date is 31 December 1984 with publ ication and distribution to be completed by 30 June 1985. � https://cpw.cvlcollections.org/files/original/fbeb717f7a5b6ce9670d101d5c7a82f2.pdf 3a9897d7ddd34378053df2e442ea8b29 PDF Text Text Colorado Division of 0ildlife Wildl ;Fe Research Report October 1984 JOB PROGRESS REPORT State Colorado Project No. 45-01-506 - 15050 Work Plan Job Title: Period Covered: Job Migratory Bird Investigations --- Waterfowl Production Surveys 01 May through 30 June 1983 Author: Gerald M. Lorentzson Personnel: M. Bauman, G. Byrne, J. Creasy, J. Corey, J. Dennis, J. Frothingham, H. Funk, J. Hicks, G. Hinshaw, J. Kauffeld, D. Kenvin, G. Lorentzson, D. Masden, M. Nail, C. Nolde, T. Rauch, J. Ringelman, W. Russell, G. Saville, S. Steinert, M. Szymczak, R. Weldon and B. Widhalm. ABSTRACT vlater conditions were good for waterfowl production throughout the state early in the spring. Heavy snowfall caused a heavy runoff late in the sprin with severe flooding of most major streams. The total number of duck breeding pairs increased 86.1% over 1982 and 7.8% over the long-term average. The numbers of Canada geese increased throughout the state. Production was up 22.1% over 1982 in the northcentral but down 2.7% from the longterm average. ��3 WATERFOWL PRODUCTION SURVEYS Gerald Lorentzson P. N. OBJECTIVES 1. To estimate the number of duck breeding pairs, by species, on selected major waterfowl nesting areas in Colorado. 2. To estimate the number of goose breeding pairs, and in some cases, obtain production data on selected goose nesting areas in Colorado. 3. Compile data and submit reports to appropriate state personnel and the Fish and Wildlife Service for use in monitoring status of the various species and establishing hunting season recommendations. SEGMENT OBJECTIVES 1. To estimate the number of duck breeding pairs, by species, in the San Luis, Cache la Poudre, South Platte and Yampa Valleys, and in North Park and Brown's Park, using procedures presented in the Program Narrative. 2. To estimate the number of goose breeding pairs in the San Luis Valley, on the Yampa, Little Snake and Green rivers in northwest Colorado and in northcentral and westcentral Colorado using procedures presented in the Program Narrative. 3. Evaluate the present boundaries of the area sampled for duck breeding pairs in the San Luis Valley. 4. Alter the duck breeding pair survey in the Cache la Poudre and South Platte areas to compensate for sample sections which can no longer be counted because of changes in land use patterns. METHODS AND MATERIALS The 1983 duck breeding pair surveys were conducted during the period of May 1 through June 30. Surveys in North Park, the Cache la Poudre and South Platte Valleys were conducted exclusively from the air. Brown's Park and the Yampa River Valley were made on the ground. Aerial counts in the San Luis Valley and North Park were adjusted for visibility by air ground comparison studies. Pairs estimates for the Monte Vista National Wildlife Refuge in the San Luis Valley were obtained from nesting transects. The San Luis Valley has lost a great deal of habitat north of the Rio Grande River over the past ten years due to the installation of sprinkler-type irrigation systems. The method of doing the duck breeding pair survey was examined in great detail and the groundwork was laid for revision of the method in 1984. It was decided that we continue to do the 1983 survey as in the past so all survey methods in 1983, remained the same as in previous years. �4 Canada Goose surveys were conducted within the May 1 through june 30th period. Estimates of the Colorado, White River, Yampa and Little Snake River populations were obtained by direct counts from a fixed wing airplane. Aerial counts were not done on the North Fork of the Gunnison River again this year. Brown's Park continues to be a ground count conducted by the Brown's Park NWR personnel. All flying was done with Cessna 185 aircraft. Two observers were used when flying transects, while one observer was used for sampling sections. RESULTS AND DISCUSSION Water conditions in Colorado were better in 1983 than in recent years. Mountain snowpack was higher than normal throughout the state. During the sur"eys the rivers were high but within their banks. Later in the spring there was a good deal of flooding, especially on the Colorado River drainage, due to a late snow melt. The estimated duck breeding pairs in Colorado increased 86.1% over 1982 and 7.8% over the long-term average. The San Luis Valley continues to show a decrease in numbers of breeding pairs probably due to the decrease in habitat (Table 1). North Park, South Platte Valley and the Cache la Poudre Valley showed increases of 100-200% over 1982. Table 1. Summary of Colorado's duck breeding pair population estimates in selected areas , 1983. Total estimated Area San Luis Valley North Park South Platte Valley Cache la Poudre Valley Yampa Va IIey Brown's Park Totals breeding pairs Long- term average Percent 1982 change Long-term average 1983 1982 12,621 15,676 19,979 12,218 3,302 1,491 12,975 5,545 5,905 5,800 3,587 1,265 26,170 17,844 8,111 4,702 2,627 1,135 - 2.7% +182.7 +238.3 +110.7 - 7.9 + 17.9 - 51.8 13.8 +146.3 +259.8 20.4 "+ 23.9 65,287 35,077 60,589 + 86.1 + - 7.2 Mallards continue to be the dominant species breeding in Colorado comprising 33.8% of the total population (Table 2). Gadwalls showed the largest increase in percentage (17.9%) as opposed to 8.4% in 1982 and 10.7% in the long-term average. Total estimated breeding pairs by species and areas are shown in Table 3. �5 Table 2. tion. Species composition of Colorado's 1983 duck breeding pair popula- Number of breedin~ Percent species composition 1951Hl2 Species Mallard Blue w i nqed and cinnamon teal Gadwa II Pi n t a i I Shoveler Green-wi nged tea I Redhead Amer i can wigeon Other divers Common mergansers Wood ducks Red breas ted mergansers Es i 15,029 9,662 11,654 5,128 5,8116 1,156 3,187 1,480 4,6118 297 122 14 5,781 2,957 2,093 2,033 2,147 2,194 447 2,192 204 una t ed Species Mallard Blue winged & cinnamon teal Gadwa II Pintail Shoveler Green-wi nged teal Red head American wigeon Scaup & other divers Common mergansers Wood ducks Red bre as ted mergansers Totals 22,093 65,287 Tota Is Tdbie 3. 1982 ~~.- 35,077 popu l a t Sdn Luis _Valley .in North Park 1954-82 average 1983 1982 average 25,980 33.8 4218 47.2 14.8 17.9 7.9 8.9 1.8 4.9 2.3 7.1 0.5 0.2 16.5 8.4 6.0 5.8 6.1 6.2 1.3 6.2 0.6 11.5 10.7 6.4 6.5 5.9 5.8 2.1 3.9 100.1 99.9 100.0 6,323 5,876 3,515 3,582 3,2:62 3,218 1,168 2,156 55,080 of duck breed i fig pa irs in Colorado, Poudre River S.Platte River Yampa Va Iley Brown's Park 1983. Totals 5,578 4,680 5,809 4,245 1,499 282 22,093 3,198 1,431 1,051 591 135 177 402 54 4 0 0 1,754 2,127 3,111 417 444 580 519 2,044 0 0 0 1,995 1,099 177 1,271 268 498 62 905 34 100 0 1,687 6,470 561 3,374 3 1,781 360 1,409 67 22 0 701 358 96 69 206 28 83 69 179 0 14 327 169 132 124 100 123 54 167 13 0 0 9,662 11,654 5,128 5,846 1,156 3,187 1,480 4,648 297 122 14 12,621 15,676 12,218 19,979 3,302 1,491 65,287 The number of Canada geese seen on the Yampa and Little Snake Rivers show a sl ight decrease in nesting pairs and an increase in non-nesting pairs over 1982 (Table 4). Total birds observed showed an increase in 1983 over 1982 but were significantly below 1981 levels. �6 Table 4. Number of Canada geese observed on aer iaI surveys in Moffat and Routt counties. Nesting ~airs Non-nesting pairs Total adults 1983 1982 1981 1983 1982 1981 1983 1982 15 28 12 12 16 50 5 25 22 80 97 38 0 30 16 46 1981 Yampa River Steamboat Springs-Craig Craig-Juniper Springs Juniper Canyon Juniper Canyon-Lily 7 15 4 19 29 19 2 64 23 1 18 11 27 6 40 4 65 59 93 10 12 2 11 11 46 22 10 12 26 21 8 32 38 29 26 25 51 22 24 46 0 Park Li ly Park Totals 26 38 65 18 126 3 70 41 75 66 276 94 341 190 Little Snake River Cross Mt.-Powderwash Powderwash-Baggs Totals 17 68 142 48 55 70 125 85 190 Nesting pairs of Canada geese in Brown's Park N.W.R. showed an increase of 84.2% over 1982 (Table 5). Total birds observed showed an increase of 33.4%. Table 5. N.W. R. Numher of Canada geese observed on ground counts in Brown's Park Nesting pairs Area Brown's Park Estimated Total adults 1983 1982 1981 140 76 72 407 305 277 no. of goslings 1983 1982 1981 305 294 245 Breeding pairs in west central Colorado decreased 44.9% from 1982. Observed singles decreased 51% from 1982 (Tables 6 and 7). This was the lowest number observed for both singles and pairs since 1978. �1 Table 6. West central Colorado Canada goose breeding Singles Area pair survey, 1983. Pairs Groups Whi te River Meeker-Rio Blanco Lake Rio Blanco Lake-Rangely Rangely-Utah Line Subtotal Colorado 2 7 1 8 8 5 18 97 7 10 21 122 5 11 13 3 4 0 3 0 0 10 21 18 24 4 4 3 3 8 28 51 36 27 3 34 9 7 6 39 95 201 3 0 32 0 3 32 River Canyon Creek-Silt Silt-Rifle Rifle-Parachute Parachute-DeBeque DeBeque-Pal isade Pal isade-Grande Avenue Bridge Grand Avenue Bridge-Fruite Fruita-Horsethief Canyon Horsethief Canyon-Utah Subtotal Roaring Fork River E I Jebel-Carbondale Carbondale-Glenwood Subtotal Gunnison Springs 2 River N 0 T GRAND TOTAL 51 C 0 U N T E D 119 I N 1 983 355 �Table 7. Comparison of the number of Canada goose singles and pairs observed west central Colorado, spring 1978-83. in aerial surveys in 00 Number observed Area Singles 1981 1980 10 39 39 2 2 36 1979 1978 39 50 26 7 1 7 0 133 105 74 77 35 11 14 11 12 4 not done 4 5 1 0 not done not done 10 8 5 5 119 216 214 138 152 70 1978 1983 1982 30 46 31 21 60 74 2 6 3 2 3 12 56 31 47 41 30 81 3 7 9 6 4 4 14 Hotchkiss-Delta not done not done 0 2 0 0 not done Delta-Gr.Jctn. not done not done 6 3 0 4 87 94 94 71 Roaring Fork River 1982 Pairs 1981 1980 1979 White River 1983 Colorado River - Glenwood Springs5th St. Bridge Bridge-Utah Line Gunnison River TOTALS 51 104 �9 The total number of Canada geese observed in the San Lui~ Va Iley increased 51 .6%. The number of nesting pairs was down sl ightly but the number of non-nesting pairs increased dramatically (Table 8). Table 8. Results of San Luis Valley Canada goose breeding pair survey. Year Projected total number Non-nesting Pairs Grouped Birds 59 59 110 92 77 63 69 64 30 100 100 101 91 130 97 99 96 58 141 226 154 90 159 189 244 376 169 227 811 60 Nesting Pairs 1975 He Iicopter 1976 He Iicopter Fixed wing 1977 Hel icopter Fixed wing ) 1978 No counts ~ 1979 Fixed wing 1980 Fixed wing 1981 Fixed wi ng 1982 Fixed wi ng 1983 Fixed wing The Northcentral Colorado goose census was done on June 16 & 17, 1983. There was an increase of 7.7% on the total number of birds present over 1982. The greatest increase was in the number of gosl ings (22%) produced in 1983. Adults showed an increase of 4.5% (Tables 9, 10, 11). The only areas which showed a decrease in numbers of gosl ings produced were in the Fort Collins and Boulder areas. Boulder and Denver were the only areas showing a decrease in the number of adults. �10 TaLle 9. Results of Northcentral Colorado goose census, Total no. goslings Produc t ion Area Water Area Well ington Terry Lake Water Supply & Storage #4 Deines Pond Launer Pond Douglas Lake Stewart Pond N. Poudre #1 Dry Creek Reservoir N. Poudre #3 N. Poudre #2 N. Poudre #5 Bureau of Standards Reservoir #8 Elder Reservo ir #8 Annex Van Sant Pond Cobb Lake Dale Pond Watson Lake Curtis Lake Beghtoi Lake Subtotal Fort ColI ins Peterson Ponds Reservo~tMiller Ponds College Lake Dean Acres Claymore Sterling Gravel Pits Larimer Co. Ponds Lindenme ier Lake Grey Lake Nova k 's Pond Flatiron Gravel Pits Dixon Andersonls Pond Parkwood Kitchel Timnath Romily Gravel Pits Foss i1 Creek Horseshoe Wolaver Pond Subtotal Loveland Flatiron Reservoir Boedecker Flatiron Gravel Pits Kauffman Gravel Pits Big Thompson River McNeil Reservoir Welch Reservoir Reservoi r No. 12 Subtotal Boulder Ish Lake Crystal Lake Terry Lake Faivre Ponds Rest Home Ponds, Sawhill Ponds & Walden Ponds Valmont Reservoir Boulder Valley Farm Eddy Pond Subtota I Denver Kettring Centennial Columbine C.C. Chatfield Golf Course Bowles Lake King's Pond Tule Lakes Marston Reservoir Pinehurst C.C. Clarefield Reservoir Kendrick Lake Sloanls Lake Denver City Park Colo. Blvd. @ Quincy Blackmer Reservoir Subtotal June 16 & 17, 1983. 77 9 3 4 22 4 o 24 16 7 12 3 34 14 o Total no. adu Its and yearl ings 129 8 12 3 81 10 18 31 28 7 22 2 55 46 14 Total birds 206 17 15 7 103 14 18 55 44 14 34 5 89 60 14 13 37 29 15 24 252 26 o 40 4 40 2 325 945 1270 133 37 289 55 148 6 o 13 13 6 4 36 84 34 10 26 4 78 30 326 18 30 212 2 o 35 17 76 21 12 26 72 o 13 o 362 18 40 238 6 35 93 34 11 51 46 22 21 4 6 196 60 26 8 6 12 42 75 62 242 82 47 12 12 245 1298 1543 3 4 171 22 10 276 37 7 206 58 22 20 309 77 ..ll _2Q .!1l 198 624 822 4 16 29 8 47 28 38 33 16 3 35 36 8 10 33 40 19 2 44 57 14 24 66 30 82 152 5 95 4 _5 2 152 251 43 13 o 85 93 41 171 240 106 57 191 240 6 103 109 o 39 6 41 39 47 16 20 9 _7 403 128 o o 36 18 54 o 93 325 385 631 19 16 93 19 22 60 30 o 6 236 1347 o 93 1583 �11 Table 10. Number of adult Canada geese observed produc t ion trend areas, 1983. in northcentral Colorado Percent change From No. of adults Average 1982 1970-82 1983 1982 1970-82 Well ington For t Co 11ins Love land Boulder Denver 945 1,298 624 251 1,347 852 1 ,204 386 423 1,407 743 747 275 521 1 ,246 +10.9 + 7.8 +61.7 -40.7 - 4.3 +27.2 +73.8 +126.9 -51.8 + 8.1 Totals 4,465 4,272 3,532 + 4.5 +26.4 Area Table 11. production Number of Canada goose goslings produced trend areas, 1983. in northcentral No. of gos lings Average Area Well ington Fort Co llins Loveland Boulder Denver Totals / Prepared by Percent change From 1983 1982 325 245 198 152 236 186 299 132 176 154 257 326 117 204 284 +74.7 -18.1 +50.0 -13.6 +53.2 +26.5 -24.8 69.2 -25.5 -16.9 1 ,156 947 1 ,188 +22.1 - 2.7 ':\ ~7~ 1970-82 Colorado (1// ,t N~'I. ./.) . ~--Ge-r-a~l~d~M-.~L~o~r-en"t~z~s~o~n~~uA.--~~Jy-,(&_ Senior Wildlife Biologist 1982 1970-82 ��13 C010rado Division of Wildl ife Wildl ife Research Report October 1984 JOB PROGRESS State of Coloardo Project 45-01-506 Work Plan Job Title: Job: Ecological REPORT Migratory - 15050 Bird Investigations 14 Studies of the Flightless Period of Ducks in Colorado Period Covered: 1 April 1983 - 31 March 1984 Au thor: M. Szymczak and J. Ringelman Per sonne 1: J. Corey, S. Porter, S. Steinert, J. Wagner, J. Ringelman and M. Szymczak, Colorado Division of Wildlife ABSTRACT for 29 of the 34 photographed wetlands. Cover maps were'reconstructed Forty-seven percent and 40% of sample wetlands contained from 1 to 10 fl ightless adults during early season (26 July - 8 August) and late season (29 August - 15 September), respectively. Lake John Annex, Walden Reservoir, Boettcher Lake and Sneed Reservoir had more than 100 molting adults during the first count while Walden and Sneed reservoirs had more than 100 during the second count. Bi-weekly counts on selected ponds indicated peak numbers as follows: pintail - 24 July; mallard, blue-winged/ cinnamon teal, green-winged teal - 24 July-8 August; gadwall, American wigeon - 8 August. Weekly counts at Walden Reservoir indicated: a decided reduction in the number of adult ducks in late July corresponding to rising water levels; a molting population composed predominantly of gadwall; concentration of gadwall during the flightless period as opposed to a widespread distribution during the pre- and post-flightless period. . Gadwall fl ight feathers grow at the rate of about 5 mm/day resulting in a feather growth period for primary 10 of 31-34 days. Gadwall are capable of voluntary fl ight when primary 10 reaches 88% of total length. The chronology of molt of the male gadwall population in 1983 was remarkably similar to 1982, beginning in early August and lasting about 45 days. The female gadwall population's molt period began earl ier and was estimated to have lasted longer in 1983 than in 1982. The gadwall weight data by stage of feather growth in 1983 paralleled 1982 male and female data with a significant decl ine during the midportion of the flightless period. Female gadwall weights in 1983 did not indicate the mid-period decline, however females captured more than once during the period exhibited weight loss. A regression equation for estimating fat of flightless gadwall was developed using a combination of girth and length measurements. �14 Flightless gadwall collected for food habits had ingested primarily Elodea canadensis, Potamogeton fil iformes and filamentous green algae. Sulfur amino-acid composition for 7 aquatic plants common to North Park is presented. �15 ECOLOGICAL STUDIES OF THE FLIGHTLESS OF DUCKS IN COLORADO PERIOD Michael R. Szymczak James K. Ringelman P. N. OBJECTIVES 1. Document the species and sex composition and seasonal abundance ducks molting on selected wetlands in North Park. 2. Identify the physical and biological used by molting ducks. 3. Investigate spatial and temporal differences by molting ducks. 4. Determine the behavioral molting ducks. 5. Investigate differences in duration of the flightless period of selected species of ducks in relation to sex, body condition, habitat quality, and time of molt. 6. Determine the survival rate of molting ducks in relation to sex, body condition, habitat quality, and time of molt. 7. Identify and quantify duck molting wetlands characteristics of of wetlands in use of wetlands time budgets and net energy balance of in Colorado. SEGMENT OBJECTIVES 1. Document the species and sex composition and seasonal abundance ducks molting on selected wetlands in North Park. 2. Identify the physical and biological used by molting ducks. 3. Investigate spatial and temporal differences molting ducks. characteristics of wetlands in use of wetlands 4. Determine the behavioral time budgets and net energy balance of molting ducks. 5. of Investigate differences in duration of the flightless period of selected species of ducks in relation to sex, body condition, habitat qual ity, and time of molt. by �16 METHODS Wetland Characteristics and Waterfowl Use The vegetative composition and coverage of study wetlands was examined to determine the validity of interpretation of aerial photos taken in August of 1982. Considerable discrepancies were found in both composition and coverage and therefore wetland cover maps were reconstructed for 29 of the 34 photographed wetlands. Maps were also constructed for the 3 wetlands that were not photographed: Home Pd. (17), Clayton Res. (32) and Addison Res. (33). Cover maps will be completed for the additional 5 wetlands in 1984. Alkal inity, pH and specific conductivity measurements were recorded for each study wetland. Number of patches and area of each aquatic type and changes in physical features, where applicable were determined from the reconstructed maps. Updated basin area changes are presented in Table 1 for the 29 photographed areas. Physical, chemical and vegetative measurements for those 29 wetlands were stored in a computer file. Table 1. Number 1 2 3 5 6 7 8 9 10 11 12 13 14 16 18 20 23 24 25 26 27 28 29 30 31 34 35 37 40 Some characteristics of some study wetlands Name L. john L. john Annex Walden Res. Damfino Res. Pole Mountain Res. S. Delaney Butte L. E. Delaney Butte L. N. Delaney Butte L. Boettcher L. Sneed Res. Tricks Pd. #1 Ca r1strom Res. Tricks Pd. #2 Holzingers Antelope Pd. Case #3 Res. Goose Pd. Marsh Pd. Eagle Pd. S.School Section Pd. Will ford Pd. Spring Creek Pd. S. Allard Pd. Geisses Pd. Hebron Rich Pd. Hudspeth Richard Pd. E. Boettcher in North Park. UTM grid locationa 4515 4515 4509 4486 4490 4506 4507 4508 4520 4524 4522 4521 4523 4509 4503 4502 4502 4501 4500 4501 4502 4501 4595 4490 4490 4486 4499 4516 4523 x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 375 376 388 381 374 376 377 376 372 371 389 389 389 395 390 385 388 388 391 393 394 395 391 386 385 380 379 376 372 aGrid location near the center of the wetland. Area (ha) 201.99 69.37 1894. 16 8.95 49.16 59.08 25.92 44.55 52.80 68. 11 3.03 37.46 5.72 16.37 7.49 5.42 4.45 3.91 2.23 2. 11 3.45 10.28 5.54 3.44 2.68 2.31 4.90 5.71 3. 15 �17 Counts of flightless adults on all study wetlands except MacFarlane Reservoir were conducted at least twice during the summer period. The first count period began on 26 July 1983 and extended through 8 August 1983 while the second period began on 29 August 1983 and ran through 15 September 1983. Most counts were conducted in the early morning or late evening hours and involved 2 people; one to drive the birds into an observable position and another to count. Additional weekly counts were conducted during early morning hours on Walden Reservoir from 22 July through 2 September. A total count was made on 22 and 29 July, while other counts included only duck concentration areas. Biweekly counts were made on Boettcher Lake, Pole Mountain Reservoir, Elk and 76 ponds from mid-July through the 2nd week in September. MacFarlane Reservoir was not counted in 1983 because many ducks on the reservoir could not be classified at the long distance observation required. Data were coded and stored on a computer file, sorted and combined with the wetland characteristic file to facilitate analysis. Habitat Utilization of Wetlands by Molting Adults Marker buoys at 1,000 ft. intervals were again placed on Walden Reservoir in the same manner described by Szymczak and Ringelman (1983). Adult waterfowl concentrations were mapped on Walden at weekly intervals during early morning hours from late July through early September. Counts were discontinued after 2 September because the increasing size of young duckl ings made correct classification of ducks difficult. The species and possibly sex, and wing feather condition (fl ighted or flightless) were recorded for all adult birds observed. Composite maps of the distribution of flighted and flightless ducks were prepared. Back mounted radio transmitters (Dwyer 1972) were appl ied to 6 flightless gadwall captured on Walden Reservoir. Radios were placed on 2 males and 1 female on 12 August and on 2 females and 1 male on 29 August. The locations of these birds on Walden were recorded periodically from 2 days after release until they attained fl ight and left Walden Reservoir. Additional transmitters were placed on 5 gadwall nesting hens and 1 hen mallard in an attempt to document molt wetland habitat selection of females. Two gadwall were captured on nests in the Case Flats area of the Arapaho National Wildlife Refuge and 3 gadwall and 1 mallard were captured on nests on islands in Walden Reservoir. All hens were captured using a Coulter nest trap (Coulter 1958). The activity, movements and locations of these hens were monitored periodically until 11 October or when radio contact was lost. Energy Balance and the Effects of Wetland Variability on Molt Dynamics Because of time constraints no time budget data were collected during the summer of 1983. Some information on activities of gadwall was obtained through recording activity of radio-marked birds when contacted visually. In addition, a Russ-track recorder monitored 24 hour activity periodically for 2 flightless male gadwall and 1 flightless female mallard. Because of problems in circuit stability of the transmitters, environmental interference and attenuation of signal because of movement, consistent data were not obtained. �18 Adult gadwall trapped in conjunction with a banding operation were weighed and measured to classify stage of molt and relate stage to physiological condition using an index developed from fl ightless gadwall collected in 1983. In this case condition is analogous to percent body fat and therefore the objective is to estimate fat. Step-wise forward multiple regression analysis was used to formulate the equation. Use of the formula required a measurement of girth and U-B Length as described in Szymczak and Ringelman (1983). The lengths of primary 1, 5 and 10 were measured but in analysis only primary 10 was utilized to establish molt stage. If a bird was subsequently recaptured only weight girth and primary length were recorded. Birds were captured at MacFarlane Reservoir (8, 22 August; 8, 13 September), Walden Reservoir (9, 12, 29 August; 7, 12, 15 September), Lake John Annex (10, 31 August; 14 September) and Pole Mountain Reservoir (3 August). Nine adult gadwall (8 females, 1 male) in a pre-molt or flightless condition were collected in an attempt to document food utilized by flightless gadwall. Birds were shot with a .223 caliber rifle after they were observed to be actively feeding for at least 10 min. The contents of the G.I. were removed immediately and stored individually in 5% Formalin solution or AFA (30% Ethyl Alcohol, 10% Formalin, 10% Glacial Acetic Acid). Thirteen gadwall have been collected over a 2-year period. The dominant vegetative type by sample was recorded. Samples of 6 of the most common rooted submergent aquatic plants in North Park wetlands and green algaewerecollected to determine the relative amount of sulfur amino acids, crystcine and methionine by species (Table 2). Green algae was included because it was commonly found in the G. I. tract of f light less gadwa II. Both of these am ino ac ids have been found to be important components of feathers in domestic fowl (Mitchell 1959, Krimm 1960). Samples were dried to a constant weight at 50°C, ground in Wiley mill and submitted to Dr. David Clarkson, Department of Biochemistry, Colorado State University, for analysis. RESULTS AND DISCUSSION Duck Censuses on Sample Wetlands Counts of molting waterfowl were considered to be more reliable in 1983 than in 1982. As in 1982 flightless ducks were observed on nearly all wetlands. During the early count no flightless adults were observed on South or North Delaney Butte Lakes or on Goose Pd. During the late count no molting adults were observed on South or East Delaney Butte lakes, Turk's Pd. No.1, Carlstrom Reservoir, Home Pond, South School Section Pond, Hudspeth's or Wattenbury Pd. Nearly one-half (47%) of the ponds contained from 1-10 fl ightless ducks during the early count while 4 wetlands, Lake John Annex, Walden Res., Boettcher Lake and Sneed Reservoir had more than 100 molting adults. During the late count 40% of the ponds had from 1-10 fl ightless adults while only 2 wetlands, Walden and Sneel reservoirs, had more than 100 ducks. Generally diving ducks were more prominent during the later count period. �19 Table 2. Species Walden Reservoir. counts of duck concentrations compositions of adult ducks recorded during weekly Percent estimated flightless in parentheses. on Date of count Species 7/22 7/29 8/05 8/12 8/19 8/27 9/02 954 1191 (18·9) 392(67.3) 451 (92.5) 544(86.9) 582(34.5) 454(18.3) A. wigeon 518 501 (6.6) 603 (0.7) L. scaup 130 Gadwall Redhead 16 BWT/CT 34(26.5) Ringneck 5 Ruddy 118(32.2) 78(62.8) 183 (31.1) 332 (3.6) 51 32(28.1) 56(53.6) 118(80.5) 59( 10.2) 34 33 (3.0) 20(50.0) 32(37.5) 45(71.1 ) 17 25 10(50.0) 16 16(12.5) 20 43 26(57.7) 4 24 12 (8.3) Ma lIard 3 6(16.7) Pinta iI 8 39(17.9) GWT 4(25.0) 1 (100) 28(25.0) 49(22.4) 6 15 3(100) 132(65.9) 79 7(14.3) 27 35 32 181 22 3(33.3) 2 4(100) 8 0 0 3(66.7) 15 Buff Iehead 0 0 0 2(100) 0 N. Shoveler 0 0 0 2 3 6 0 0 0 0 262 0 1864 614 702 1132 1361 1405 Unknown Total ~ 1871 According to biweekly counts on 5 selected wetlands, peak numbers of fl ightless pintail occurred on or before 24 July, 1983. Peak numbers of mallards, blue-winged/cinnamon teal, and green-winged teal occurred around 8 August, however, those numbers were only sl ightly higher than those recorded on 24 July, 1983. Gadwa lI and American wigeon numbers peaked on 8 August, 1984. Flightless divers of all species present, lesser scaup, ringnecked duck, ruddy, redhead and canvasback were most numerous on 23 August 1983. Walden Reservoir Census During the census period peak numbers of birds were recorded on Walden Reservoir in late July (Table 2). A substantial reduction in duck numbers occurred between 29 July and 5 August, about the same time as water levels in the reservoir were being raised. Census techniques were changed between the 2 periods from a total count to counts of concentrations only, however, these changes were not responsible for reduced number. For example, of the 1191 gadwall recorded on 29 July, 1021 were located in 5 concentrations in 6 different 1,000 ft. quadrats and would have been included with either technique. The molting population was composed predominantly of gadwall, American wigeon, lesser scaup, ruddy ducks and redheads. Earlier molting species such as blue-winged/cinnamon teal, mallard, pintail and green-winged teal were represented by only a few birds with blue-winged/cinnamon teal the most prevalent. �20 Gadwall were the most numerous molting ducks with over 400 flightless birds present during the middle parts of August when most of the gadwall were fl ightless (Figure 1). At that time the male molting population had attained 40-50% of primary feather growth while females were just beginning the feather growth process (Figure 1). The location of gadwall on Walden Reservoir was somewhat widespread when birds were capable of flight with only 1 area accounting for more than 20% of the birds observed (Figure 2). Flightless birds restricted their distribution with most birds concentrated in the central portion of the reservoir or in the north bay (Figure 3). Preliminary analysis of data from radioed and other marked birds substantiated the distribution information obtained from counts, indicated that most birds have a limited home range when fl ightless, and suggested that the home range increases late in the feather growth period but prior to attaining flight. Wetland Selection - Nesting Hens The attempt to document molting habitat selection through applying transmitters to nesting hens was less than successful, probably in part because of what was considered aberrant behavior in response to the radio-transmitters. Of the 6 birds to which transmitters were attached only one went through a normal nesting brood-rearing cycle. Three of the birds, 2 gadwall and the mallard, abandoned broods shortly after hatch (Table 3). One bird did not return to the nest after capture and another abandoned the nest after 12 days of incubation. Table 3. Nesting and brood rearing history of hens to which radio transmitters were applied. Transm itter wt. as % of body wt. Nesting activity post-transmitter attachment Species We ight (gr) Gadwa 1I 570 4.7 10 Normal Gadwa II 590 4.5 12 Abandoned within 4 days Gadwa II 590 3.7 17 Abandoned wi thi n 4 days Gadwa 11 610 4.3 o (abandoned) Gadwa II 650 4.0 12(abandoned) Ma Ilard 880 2.5 7 Brood rear ing notes Abandoned wi th in 3 days �21 Only 2 of the birds were observed during the fl ightless period. The mallard, after abandoning its brood about 8 August spent time on the southeast corner of the reservoir and on the adjacent Illinois River bottom. About 6 September the mallard moved to the emergent cattail patch adjacent to the large middle island and molted its fl ight feathers. The bird was subsequently shot by a hunter on 2 October on Walden Reservoir with some primaries still in the blood quill stage. A gadwal I, which nested on Case Flats and abandoned its brood about 1 August, molted on Case Reservoir #3. It was observed as flightless on 14 September and moved to Case Reservoir #2 on 2 October. One gadwall which abandoned its nest after 12 days of post-marking incubation also spent time on the Illinois River and on Walden until 31 August. It was not found again until 21 September, when it was observed in the north bay at Walden. It seemed to have new fl ight feathers. Signals were lost for the other 3 gadwall on 6 August, 26 August and 12 September. One bird was shot near Laramie on 8 October. The hunter indicated that the bird had faded primaries when harvested. Feather Growth and Duration of Molt of Gadwall Recapturing a bird and measuring feather growth status provides data to estimate feather growth rates and the duration of the fl ightless period. Measurements of feather length of post-flightless gadwall provided an estimate of feather lengths at ful I growth (Table 4). Unfortunately sample sizes for females are very small, but, comparatively, sex variations for gadwall are quite similar to those found by Owen (1979) for the mallard. Table 4. Primary feather lengths of post-fl ightless gadwall as determined from measurements of birds captured 1977-83. Sex Pr imary No. Statistic Males Females Number Mean S.D. Variance 53 110.2 3.2 10.3 6 108.5 4.5 16.9 5 Number Mean S.D. Variance 35 148.7 6.8 45.5 5 145.2 5.4 22.2 10 Number Mean S.D. Variance 21 170.4 9.3 81. 6 3 157.7 6.4 26.9 �22 Number 10 primary feathers grew at the rate of about 5 mm/day for both sexes in both years (Table 5). Small but significant differences were noted in growth rates by sex in 1983, however these rates were for growth throughout the fl ightless period regardless of stage. Some preliminary analysis indicates reduced growth in later stages of the period and therefore growth rates may have to be prepared by period to obtain an accurate compa rison. Table 5. Number 10 primary growth rates (mm) of adult flightless captured more than once during the season in North Park 1982-83. gadwa 11 Year Sex No. Mean growth/day S.D. 1982 Male Female 44 11 4.97 5.06 0.51 0.83 F 0.73 1983 Male Female 23 16 5.29 4.98 0.44 0.37 F 0.02 At the estimated rates of growth it would take a male and female gadwall 32-34 days and 31-32 days, respectively, to complete the growth of a number 10 primary (Table 6). Ducks can fly prior to primary 10 reaching ful I growth. By throwing birds in the air Owen (1979) found mal lards could involuntarily fly when primary 10 reached 78% of full growth. Telemetry location data indicated that a female gadwall made a voluntary fl ight to an adjacent wetland when primary 10 length was an estimated 88% of tota 1. Table 6. Characteristics gadwall and mallards. of the duration of the fl ightless period in Species Sex Mean estimated total length, No. 10 primary(mm) Gadwa 11 Male Female 170 158 Ma 11a rd (Owen 1979) Male Female 178 167 Days required to atta in total length 32-34 31-32 34 32 Days until capable of fl ight 28-30 27-28 27 25 �23 Timing of Flightless Period Establishing the stage of feather growth of each gadwall captured provides information on the chronology of molt by sex. The male molt period in 1983 was remarkably similar to 1982 (Figure 4). The molt period began in early August and lasted for about 45 days in both years. In 1982 the female molt period paralleled the male molt, only approximately 20 days later. In 1983, female molt began earlier and was projected to end quite a bit later than in 1982. Data from 1982 were primarily from Walden Reservoir (97% males, 95% females) while in 1983 Walden birds did not completely dominate the sample (54% males, 55% females). Weight Dynamics of Flightless Gadwall In 1982 weight of gadwall as plotted by stage of molt showed a significant decl ine during the mid-portion of the flightless period for both sexes (Figure 5). In 1983. weights for males indicated a similar trend as in 1982 with weights for birds in the 5th stage (91-120 mm) being significantly lower than weights of birds in the 3rd stage (31-60 mm). Females in 1983, however, did not follow the same trend as birds showed no change in weight by stage (Figure 5). Since the 1982 sample was predominantly Walden birds, an analysis of Walden birds only was conducted, but the results were the same; no change in weight over time. Of greater value in examining changes in weight during the flightless period are data from gadwall captured more than once during the period. Weight trends of birds recaptured were plotted on graphs according to primary 10 development stage during the interval between captures. Birds were categorized according to 3 intervals which related to the trends observed for both sexes in 1982 (Figure 5). The results in Table 7 indicate trends in weight loss or gain similar to those hypothesized from weight-feather growth measurements of the general population in 1982. Measurements which occurred across periods showed definite weight declines. About 50% of the across period category measurements for males were taken before primary 10 reached 120 mm and only 3 were taken after primary 10 had reached 140 mm. For females, 10 of the 13 measurements in the across period category were recorded before primary 10 reached 120 mm. Table 7. Weight dynamics of adult gadwall captured more than once du ring the 1983 trapping (molting) season. Stage #10 primary (mm) pre-mo It - 60 60-120 > 120 Across stages Males Weight gain No. 9/dy 4.12 2 1 4.32 1.92 Weight loss No. s Idy 1 2 3.33 6.67 22 3.27 Females \~eight gain Weight loss No. No. q/dy g /dy 5 2.00 4.35 0.59 12 4.31 �24 Two males were recaptured 3 times. The first male gained 70 grams from pre-molt to 28 mm; then lost 20 grams from 28 mm to 81 mm. The second bird lost 140 grams from 56 mm to 147 mm but gained 40 grams from 147 mm to 166 mm. Both of these reversals reflect what would be expected considering trends in Figure 5. In summary these weight data continue to indicate the use of endogenous reserves during the flightless period. Female recaptures in 1983 (Table 7) contradict the weight dynamics information obtained for the general female population and indicates agreemment with 1982 data (Figure 5). These data will be analyzed using the condition index to adjust for structural size. Condition Index A regression equation to estimate fat condition of flightless gadwall wad developed using carcass composition data and physical measurements of adult males collected in 1982. Regression analysis resulted in the fol lowing formula. Es t imated Fat 2.14(girth)+17.09(girth/U-B Length)-438.71 R = 0.822 R2 = 0.636 F = 16.7, df 2/16, P < .000 Both variables are significant at P < .005 with girth being most significant. Biologically one would hypothesize that weight would be an important variable in developing an equation to estimate fat. However, for these data weight and girth were strongly positively correlated and in the regression analysis girth was the better predictor. Foob Habits Thriteen gadwall were collected over a 2-year period to examine food habits of the birds during the pre-flightless - flightless period. Statistics of the birds collected are presented in Table 8. Three species were commonly found in the esophagus of the fl ightless birds: Elodea canadensis, Potamogeton filiformes and a filamentous green algae. In relations to the food habit collection a ~ample of the common submergent plants found on most wetlands in North Park were collected and analyzed for sulfur bearing amimo acids. The results are presented in Table 9. Fi lamentous algae, which was hypothesized to be the possible key to fl ightless gadwall nutrition and was the exclusive item ingested by some flightless gadwall collected ranked lowest in combined sulfur amino acids. Myriophyllum which is ubiquitous in North Park wetlands and was not present in quantity in any gadwall collected contained the largest relative concentration. �25 Characteristics Table 8. Locat ion Walden of gadwall Date Time Sex collected for food hab its in North Park 1982-83. Vegetation ingested (major spp.) Molt stage Notes Res. 27 Jul 82 0710 Female Pre-fl ightless Zannichell ia palustris Potamogeton filiforis Prob. nes t ing L .John Annex 28 Jul 82 1410 Female Pre-fl ight less (flying) Z. palustris Small samp Ie Case #3 12 Aug 82 Male Pre-fl ightless Z. palustris 24 Aug 82 Male Post-molt (flying) Elodea canadensis Female FI ight less E. canadensis Walden Res. Case Flats 22 Aug 83 Hebron Pond 23 Aug 82 1600 Female Flightless Filamentous Hampton 24 Aug 83 1600 Female FI ightless E. canadensis 25 Aug 83 1900 Male FI ight less Filamentous 29 Aug 83 1600 Female Flightless P. fil iformes Hebron Pd. Pd. 1800 algae t ~ MacFarlane Res. 30 Aug 83 1915 Female FI ightless E. canadensis Eagle Pd. 31 Aug 83 1930 Female FI ightless Filamentous Giesses I Case d Pd. Small sample algae Pd. west of Walden Res. ~ ~ Small sample; Band #886-85299 Band #886-88008 algae 13 Sep 83 1800 Female FI ightless P. f iI if orrne s Filamentous algae 13 Sep 83 1900 Female FI ight less E. canadensis Table 9. Results of sulfur of North Park wetlands. amino acid analysis of com~on Band #886-88153 Small sample aquatic Nanno-moles/mg Carboxymethylated Methionine cysteine_ Species Myriophyllum Potomogeton Ceratophyllum Potamogeton Total filaformis 10.7 11. 1 25.8 25.2 36.5 36.3 demersum 8.2 25.4 19.0 18.3 14.8 33.6 26.9 24.6 exalbenscens richardson Ruppia mar it ima Elodea canadensis Filamentous plants algea ii 7.9 6.3 8.8 not detectable 18.3 23.6 18.3 �26 LITERATURE CITED w. Coulter, M. 1958. Dwyer, T. J. 1972. 43:282-284. Krimm, S. 1960. A new waterfowl An adjustable nest trap. radio-package Bird Banding 29:236-241. for ducks. Bird Banding Structure of frizzle mutant feather keratin. Mitchell, H. H. 1959. Some species and age differences in amino acid requirements. Pp. 11-43 in Albanese, AA. Protein and amino acid nutrition. Academic Press, New York. 604pp. Owen, M. 1979. The duration of the flightless mallards. Bird Study 26(4):267-269. period in free-living Szymczak, M. R. and J. K. Ringelman. 1983. Ecological studies of the flightless period of ducks in Colorado. Colo. Div. of Wildl. Fed. Aid Wi ld l, Res. Rept., Oct. Pp. 13-31. Prepared by: ~i1f~ Michael Wildlife R. Szymcz Researcher C �27 WALDEN RES. - ADULT GADWALL POPULATION 1983 100 , en en 80 w ,_J t- 60 :c (!) ,_J LL ,_J ,_J ..... 40 -c ..•• . ...... ..... ... ~ .••• ' FEMALES _ •• 0 « ..••.• .• •• o t- ,_J ::> 0 -c ..-.... ...... •• •• • •• 20 . •• •• • • LL 0 •• •• •• ::R 0 '..•• ••. ....: o~--------~--------~~----------~--------~AUG 1 AUG 15 SEPT 1 SEPT 15 Fig. 1. Number of gadwall present on Walden Reservoir and the percent estimated as flightless according to weekly counts 22 July through 2 September 1984. Male and female plots indicate mean stage of primary feather growth, in percent of anticipated total length of the number 10 primary, of flightless gadwall captured on Walden Reservoir. �28 WALDEN RESERVOIR DISTRIBUTION ADULT OF FLYING GADWALL .• <1% I3J 1-5% El 5-10% 1110-20% > 20% ,~ \U7 Fig. 2. General distribution of flying adult gadwall observed on Walden Reservoir from 29 July through 2 September 1984 according to weekly counts. , �29 WALDEN DISTRIBUTION ADULT RESERVOIR OF FLIGHTLESS GADWALL .•. < 1 % r Cd 1-5% rm 5-10% 1110-20% > 20% •I Fig. 3. Reservoir General distribution of flightless from 29 July through 2 September adult gadwall 1984 according observed on \.JaJden to weekly counts. �30 FULL GROWTH MALES - FEMALES MEAN LENGTHS 170 mm - 158 mm >- 0:: -c 1 ~ 0:: a.. MALES 0 T"" ~ I J: l- e) W ~ ...J <t , FEMALES ;, Z 40 .f ~ ~ ,f I I- I 0 o l- I I I I LL 0 cfi. 1982 ---- 1983 ----- 20 AUG 1 AUG 15 SEPT 1 SEPT 15 OCT 1 Fig. 4. Chronology of primary growth according to mean primary 10 feather lengths at date of capture of flightless gadwall captured in North Park 1982-83. �31 825 68 800 __ C/) _.... /~O 775 // ~ -c / 9 0:: / _..... ••••74 ••••.••••. •..•.• ••••.97 ••.• ~ / MALES 164 I / M' , ' / 'I 'I / "I 1 ~ 54 C!) 750 I-- I 1982 <!J w ~ 44 1983 ---- 725 >- Q o CO 700 FEMALES 675 27 pre-molt 0-30 31-60 =If: 10 PRIMARY Fig. 5. Relationship North Park 1982-83. 61-90 91-120 >120 LENGTH (mm) of body weight of gadwall to fl ightless stage in ��Colorado Division OF Wildl ife Wildlife Research Report October 1984 33 JOB PROGRESS State of Colorado --------------------------- Project 45-01-506 Work Plan Monitoring - 15050 Job: Development Job Title: Period REPORT Wintering Covered: Migratory Bird Investigations 15 and Use of a Physiological Mal lard Nutrient Condition Index for Reserves 1 April 1983 - 31 March 1984 Authors: James K. Ringelman and Michael R. Szymczak Per sonne I: G. Berl in, J. F. Corey, H. D. Funk, G. M. lorentzson, Pinkham, J. K. Ringelman, G. Shoener, M. R. Szymczak, Colorado Division of Wildlife; D. Bowden, D. Delong, A. Mendez, Colorado State University. M. K. ABSTRACT Mal lards (Anas platyrhynchos) were collected during winter 1982-83 in eastern Colorado, and subjected to whole-carcass analysis to determine carcass composition and develop a physiological condition index. Condition index was defined as body fat/fat-free body weight. Multiple regression equations using body weight and wing length were highly correlated with total body fat in male (R = 0.802) and female (R = 0.824) mallards. When tested on an independent data set, fat predictor equations accurately estimated measured body fat (r = 0.892). Weight and wing measurements were easily obtained on live mallards under field conditions with high repeatabil ity among observers (C.V. < 1%). By accounting for variability in structural size among birds, models were able to detect differences in body condition that were not detectable using body weight alone. Mallards were captured in bait traps during December, January, and February 1983-84 at Bonny Reservoir and Kodak Ponds. Additional birds were trapped at Segelke Slough and Valmont Reservoir during January, 1984. Condition index models were used to assess the physiological status of all sample groups. Significant differences in condition were noted according to a bird's age and sex, the month of trapping, and trap site location. During January, mallards at Kodak had higher energy reserves than at the other sites, although this relationship was not consistent for all age/sex groups. Overall mallard condition was lower during winter 1983-84 than 1982-83, but first-year males once again possessed the lowest, and adult females the highest, relative energy reserves. �34 Analyses of condition index values obtained from hunter-killed mallards at Bonny Reservoir indicate that condition changes rapidly in response to weather. Daily high temperatures during the previous two-week period corresponded with changes in mean condition levels. Examination of January weight data collected at Bonny from 1977-84 further supports the hypothesis that weather impacts condition and results in large annual differences in mean body weights. �35 DEVELOPMENT AND USE OF A PHYSIOLOGICAL CONDITION INDEX FOR MONITORING WINTERING MALLARD NUTRIENT RESERVES James Ringelman Michael Szymczak P. N. OBJECTIVES 1. Develop a physiological condition index to accurately nutrient reserves of wintering mallards. 2. Determine if differences exist in the physiological mallards wintering in several areas of northeastern 3. Document temporal changes in mallard physiological major Colorado wintering area. assess the condition Colorado. condition of on a 4. Relate spatial and temporal differences differences grain. in mallard condition, if such exist, to weather and the availability of waste cereal SEGMENT OBJECTIVES 1. Capture, weigh, and measure 160 mallards (40 of each age and sex) during winter at four northeast Colorado wintering areas. 2. Apply the physiological condition sex, and age-specific differences 3. Relate nutrient reserves to weather and the abundance cereal grains in the vicinity of capture sites. index to determine regional, in the level of nutrient reserves. of waste INTRODUCTION Interest in the nutritional status of waterfowl was spurred nearly 15 years ago, when Ryder (1970) hypothesized that the clutch size of arctic-nesting geese had evolved in relation to the level of endogenous reserves carried to the breeding grounds. Subsequent field studies supported Ryder's hypothesis (Ankney and Macinnes 1978, Davies and Cooke 1983) and documented the role of nutrient reserves in reproduction of Canada geese (Branta canadensis, MacLandress and Raveling 1981), maccoa ducks (Oxyura maccoa, Siegfried et al. 1976), common eiders (Somateria mollissima, Korschgen 1977), wood ducks (Aix sponsa, Drobney 1980), and mallards (Krapu 1981). Early studies of the-endogenous reserves of live waterfowl used body weight as an indicator of nutritional status (Hanson 1962, Folk et al. 1966, Street 1975, Owen and Cook 1977). Implicit in this approach were the assumptions that (1) lean body weight was constant within a species (Connell et al. 1960), (2) fat was most important in determining condition, and (3) structural size differences among individuals of a species did not affect �36 conclusions about nutritional status. Later researchers recognized that structural size biased indices based only on body weight, and "condition indices" were developed to eliminate such biases. Most indices have used the simple quotient of body weight divided by a measure of structural size such as wing, keel, tarsus, bill, or total body length, or a combination thereof. Whereas some researchers have used body measurements that correlate with lean body weight (Bailey 1979, Chappell and Titman 1983) or skeletal weight (Wishart 1979), others have adopted arbitrary structural measurements (Harris 1970, Bennett and Bolen 1978). Many authors have neglected to define condition (Harris 1970, Street 1975, Bennett and Bolen 1978, Wishart 1979, Campbell and Leatherland 1980, Reinecke et al. 1982), or have provided vague definitions relating condition to IIfitness of a bird to cope with its present and future needs" (Owen and Cook 1977, Owen 1981). We concur with Evans and Smith's (1975: 64) definition that "condition is a measure of the chances of survival of an individual at a particular time of the year and/or of its potential for breeding successfully". During winter, fat is the component of condition which is the most labile and potentially limiting to the mallard. In addition to serving as a supplemental energy source necessary for survival during winter periods of high energy demand and food scarcity (Jordan 1953), winter fat reserves may also provide a source of energy and nutrients used during reproduction (Krapu 1981). Waterfowl researchers are striving to understand the interrelationships among the breeding, post-breeding, and wintering periods. Energetics, with emphasis on the dynamics of nutrient reserves, is the common denominator linking these periods. A physiological condition index for the mallard, a species that serves as the focus for much duck research and management, would aid winter energetics studies by providing managers with a method to evaluate the biological affects of hunting regimes and habitat manipulations. METHODS Derivation of the Condition Index Mal lards were trapped, measured and collected at Bonny Reservoir during 2-7 December 1982, 15-21 January 1983 and 27 February-1 March 1983. Additional samples of birds were trapped and measured at three locations in northeast Colorado: Kodak Ponds, Valmont Reservoir, and Segelke's Slough (Fig. 1). Trapping on the latter three areas began on 11 January 1983 and was completed by 26 January 1983. Birds trapped in December and January were winter residents, but those sampled in late February may have included early spring migrants. For each location and time period, an attempt was made to measure 40 birds each of the four identifiable age and sex classes: adult females (AF), first-year females (FYF), adult males (AM), and first-year males (FYM). Birds were aged using the criteria described by Carney (1964) and Krapu et al. (1979). At Bonny, we measured and collected an additional eight birds of each age and sex class during the three periods, except nine AF were collected during the first period. All birds were captured in salt plains traps (Szymczak and Corey 1976) baited with dried, whole kernel corn. Birds were measured after being held for approximately 24 h to allow ingested corn to be voided. �37 Five physical measurements were taken on all birds (Table 1). Collected birds were euthanized immediately using C02 in an air-tight box. Birds were plucked and frozen in plastic bags for transport and storage. Later, carcasses were thawed and the bill, the feet below the proximal end of the tarsus, and the gut contents removed. The partially refrozen carcasses were weighted and ground three times through the 9 mm plate of a commercial meat grinder, then the homogenate partially refrozen and passed three more times through a 3.5 mm grinding plate. During the final grinding, two subsamples of about 20 g were selected and placed into pre-weighed 25 mm x 100 mm cellulose extraction thimbles. An additional three subsamples were taken from one bird. Between carcasses, the grinder was cleaned thoroughly in hot water. Samples in thimbles were weighed (0.0001 g), dried in a convection oven at 1000 C to a constant weight (~ 48 h) to determine % carcass water, then fat extracted for 6-8 hours using petroleum ether in a Goldfisch extraction apparatus. Grams fat for each bird was calculated as mean % fat in the samples x mean dry carcass weight. Our model ing approach followed Johnson et al. (in prep.) who modeled condition as the quotient of total body fat/fat-free dry weight, an assumed indicator of structural size. Dry weight is usually the preferred measure since water content as a percentage of fat-free weight (FFW) may vary within a species (e.g., Campbell and Leatherland 1980). However, since we could not detect differences in percent carcass water on a FFW basis among our sample birds, we adopted fat-free (wet) weight as our structural size indicator. Since FFW = field weight - fat derivation of the condition index was reduced to estimating our final index being modeled as: condition index total body fat, fat field weight - fat We derived an estimate of body fat by evaluating 12 variables (Table 2) in a stepwise, forward, multiple regression equation using extracted body fat as the dependent variable. Variables were eliminated that did not contribute significantly (p > 0.05) to an improvement in the multiple correlation coefficient, then a second multiple regression was performed using only significant variables to derive the final regression model. Models were compared among periods, sex, and age classes for statistical differences in intercept and slope of the regression coefficients (Zar 1974), and data pooled if differences were insignificant (p > 0.05). This procedure continued uncil statistically distinct regression models were derived for the highest order combination of periods, age, and sex classes. Mallard Trapping and Analysis of 1983-84 Data Forty mal lards of each age/sex class were captured in salt plains bait traps as follows: during December 1983, and January and February 1984 at Bonny Reservoir and Kodak Ponds; during January 1984 at Valmont Reservoir and Segelke Slough. All trapping was conducted between 1-9 December, 12-21 January, and 22-28 February. Only 35 adult females were captured at Kodak during December 1983. Body weight and wing length measurements �38 Table 1. Independent variables used in simple and multiple correlation analyses of mallard body fat. Length measurements are in mi 11 imeters. Variable Body weight to the nearest 10 g. Total body length from the longest retrix to the tip of the nail. Wing length of the straightened and flattened wing from the proximal of the carpo-metacarpus to the tip of the longest primary. end Bill length from the gape to the tip of the nail. Girth at midsection (behind wing) taken with a modified cinched to a spring-scale tension of 1000 g. seamstress Estimate of whole body volume [(girth) x (total/2.75)/12,566] Ringelman, unpublished data). Estimate of whole body density Weight/wing Weight/totai tape (j. K. (weight/volume). length. length. Weight/bi II length. Girth/wing Girth/total length. length. Table 2. ~1ean values of condition index of mallards during ear ly, mid, and late winter 1983-84 at four eastern Colorado locations. Age/sex class Kodak Bonny Valmont Segelke Unweighted ear It mid late early mid late mid mid AF .156 .141 .145 .172 .165 .159 .137 .130 .151 IF .123 .106 .148 .136 .151 .164 .124 .131 .135 AM .128 .124 .136 .158 .154 .156 .126 .130 .139 111 .115 .122 .135 .119 .154 .153 .121 .143 .133 mean �39 taken on all of the 1,260 mallards captured. and entered on computer for analysis. Check Station All measurements were coded Collections Division of Wildlife personnel at the South Republican State Wildlife Area were trained in techniques to age, weight, and measure mallard ducks. Weiqht and wing length data were obtained on all mallards checked through the experimental hunting area check station during the three days per week the region bel ow Bonny Dam wa s open to hunting. Data were obtained on 29 adult female, 21 immature female, 412 adult male, and 69 immature male mallards during 10 November-15 January. Mallard weight data, collected by DOW personnel during January trapping at Bonny Reservoir, we re obtained and entered on computer. Weight data co] Jeeted du r i nq 1982·-83 and 198}-84 under this project were reformatted and added to the 1977-81 information. Analysis of variance was used to examine the null hypothesis of no difference in weights within age/sex classes amOGg years. Climatological data we re obtained from summaries published by the National Oceanic and Atmospheric Adm i n i s t ra t i on , from the weather station operator at Bonny Reservoir, and from Colorado State University. Correlation analyO ses were used to explore the relationship between weather and mallard condition, particularly for 1977-84 January weight data and 1983-84 check stat ion informat ion. Wea t her data were encoded onto computer for those periods and locations where ma lla rd weight and condition data are available. Information includes location, day, month, year, precipitation, maximum and minimum temperature, snowfall, snow on ground, and average daily windspeed. Weather for 1,498 days is now accessible for analyses. RESULTS The Condition AND DISCUSSION Index Wing length was the morphological measure that best correlated with measured FFW (r = 0.838, ~ < 0.01). Models using wing length and body weight were best able to estimate total body fat. Slopes and intercepts of these fat predictor models did not differ among time periods within sex and age cla~)ses or between age classes within sex (p > 0.10), but did differ between sexes (p < 0.05). Therefore, a separat; equation was deemed appropriate for---;nalesand females, but within each sex, equations could be appl led regardless of age class or time of winter. The fat predictor' equation for' f ernaIe s was: �40 fat = (0.571 x body weight) - (1 .. 695 x wing length) + 59.0 .- (1.598 x wing length) + 31.5. and for males: A fat"" (0.539 x body weight) Correlations of estimated fat with measured carcass fat were highly significant (p < 0.001) for both males (r = 0.802, N = 47) and females (r = 0.824, N = 49),-and offered an improvement (p < 0.05) over the use of body weight alone, which accounted for" only 45.7%-of the variation in carcass fat for the entire data set (r = 0.676). Thus, an additional 18-20% of the variation in total carcass fat was attributable to differences in structural size as indexed by wing length. Performance of the Index The reliability of the condition index depends in part on the precision of body weight and wing measurements. We took body weight and wing length measurements three times on each of six mallards (total of 36 weight and wing measures) to determine observer error rates. Coefficients of variabi 1 ity within observers averaged 0.55 and 0.54% for wing length and 0.18 and 0.43% for body weight. Variability between observers was slightly higher, with C.V. = 0.62% for wing and 0.39% for weight. Because observer error rates were usually < 1%, we believe data collected by two or more persons could be pooled without introducing serious observer biases. A second source of error, the methodology used to determine carcass water and fat, was evaluated to measure the precision of our technique and the potential error in our fat predictor models. Five replications of one specimen resulted in Y water content = 56.8% (SO = 0.378, range = 56.3 57.2%). Fat content estimates, expressed as a percent of dry weight, were also precise (i = 46.7%, SD = 0.563, 46.1-57.4%). We conclude that our technique provided good estimates of water and (petroleum ether) extractable lipid content, and recommend this procedure to other researchers. A posteriori comparisons between a multiple regression model and the data Trom which that model was derived typically result in high correlations. Therefore, a valid test of a model's performance is best achieved through analysis of a different data set. Carcass fat extractions of 17 male and 13 female mallards obtained during winter 1981-82 provided an independent data set used to verify performance of the fat estimator models. The use of sex-specific equations resulted in a high correlation (r = 0.892, ~ < 000001) between measured and estimated total body fat (Fig. 2). Although the high correlation observed for the independent data set underscores the utility of the models, care must be taken when applying these findings to other populations. Considerable geographic variation in body size exists in the mallard (Palmer 1976:283), which could result in differences in structural size relationships among wintering populations. During periods of extreme physiological stress, waterfowl will catabolize protein as a major source of energy and thus alter the constancy of the FFltJcomponent. Equations should be applied only to birds within the weight range observed in this study (males: 850-1300 g, females: 740-1230 g). Waterfowl undergo seasonal changes in carcases protein content in response to increased demands for protein (Milne 1976, Korschgen 1977, Ankney and �41 Macinnes 1978) or possibly as an adaptation to increase the effective size of fat reserves (Reinecke et al. 1982). Growth of the reproductive organs contributes up to 100 g additional body weight in mallards (Krapu 1981). These relative changes in either protein or fat invalidate our fat predictor models. For example, a female mallard with an assumed wing length of 270 mm and weighing 1200 g would have a predicted fat content of 285 g. Krapu (1981) found that the total lipid content of a prelaying, 1200 g mallard hen averaged 110 g. The serious error evident in this example emphasizes the inappropriateness of using our condition indices during t lne s other than winter. Application of the Condition Index The use of condition indices as investigative tools will involve the detection of spatial or temporal differences in relative nutrient reserves of duck populations. By compensating for variability introduced by structural size differences, condition indices reduce the chances of fail ing to detect a true difference in condition when one actually exists (Type II error). Body weights and condition indices calculated for mallards captured at four locations in eastern Co Ior ado du r i nq January, 1983 (Fig. 3), exempl ify the value of condition indices. Analysis of field weight alone failed to detect mean weight differences among location' for FYF or AM (p > 0.05), impJying no difference in condition based on body we iqh t . However,-separation of mean condition levels by location was possible for each sex and age class using our condition index (p < 0.005). Comparison of mean condition indices for FYF indicates that birds at Bonny Reservoir were in poorer condition than at the other three sites (p = 0.002), and AM were in better condition at Segelke's than at Kodak (p = 0.(05). Because wing length differed by location within FYF and AM (~< 0.0005), the index was better able to separate condition levels by accounting for differences in structural size. Mean wing lengths (± SE) for FYF ranged from 267 ± 1.22 to 273 ± 0.97, and AM varied from 292 ± 1.19 to 299 ± 0.82. Therefore, a condition index is most effectively used when study objectives involve detecting slight differences in condition or when large sample sizes are unattainable. Physiological Condition of Mallards During 1983-84 Winter 1983-84 brought higher snowfall and colder temperatures than in 198283. Not only did fewer waterfowl overwinter in Colorado, but those that remained were generally in poorer physiological condition. A four-way analysis of variance, comparing differences in condition index by bird age and sex, month, and trapping location, revealed that differences in condition could be attributed to each effect (p < 0.001). Two-way interactions of main effects were also significant (p < 0.03), except for the sex x location interaction (p = 0.37). - �42 Large differences in condition index were apparent among time periods within locations (Table 2), but winter lows in condition varied from early winter in first-year males to late winter in adult females. In general, first-year males possessed the lowest (unweighted mean) reserve levels, and adult females the highest reserves. Differences in January condition among locations were apparent within each age/sex class (p < 0.0001). Adult males and adult females were in better condition at Kodak than at the other three sites, which did not differ among one another (p > 0.05). First-year females were in better condition at Kodak and poorer condition at Bonny than at the remaining two locations. First-year males had lower levels of reserves at Valmont and Bonny than at Segelke and Kodak. Check Station Data Because of small sample sizes of all but adult males, other age/sex groups were excluded from prel iminary analyses of check station data. Even with the larger sample size (N = 412), mean condition index values for adult males varied greatly from week to week and even among consecutive hunting days (Fig. 4). Nevertheless, it is apparent that during late November early December, adult male mallards at Bonny were in intermediate condition. In the third week of December, a relatively higher plane of nutrition was achieved. Finally, heavy utilization of endogenous reserves during late December - early January resulted in a drastic decl ine in mid-winter condition. Weather data, while falling short of demonstrating a causal relationship, suggests that temperature may playa role in regulating energy reserves. The low index level that existed during late December early January was preceeded two weeks earlier by record low, daily high temperatures. The high condition level the third week in December was preceeded by a two-week high temperature period. It has been recognized (Owen and Reinecke 1979) that the cost of thermoregulation in ducks can be high at temperatures below thermoneutrality (about 300 F). Historic Weight Data Data on Bonny Reservoir maliard weights, collected during January 1977-81 and 1983-84, were examined for yearly differences in mean weights as well as relationships between weights and weather. Body weight, rather than condition index, was used because no wing length measurements were taken during 1977-81. Annual differences in weight within each age/sex class were evident, particularly during the high in 1981 and the lows of 1977 and 1984 (Fig. 5). Patterns in weight change year-to-year were the same among each class. The daily high temperatures 10 and 20 days prior to trapping each year were positively correlated (P < 0.05) with body weights of each age/sex class (Table 3). Mean and daily low temperatures were less highly correlated with body weights. These results strengthen the hypothesis that temperatures impact energy reserves of wintering mallards. �Table 3. Simple correlation coefficients between temperature and body we iqh t s of mallards captured at Bonny Reservoir during January, 1977-84. All coefficients greater than 0.70 differ significantly from zero (P<0.05). Temperature (days before Dai ly high Da i 1'1 low Da i ly high Da i ly low Dai ly high Da i ly low Da i 1y high r r • temp. temp. temp. temp. temp. temp. temp. (5) (5) (10) ( 10) (20) (20) (30) Mean da i ly temp. Mean da i Iy temp. Mean da i ly temp. (30) (5) (10) (20) Mean da i ly temp. (30) Da; Iy r Variable trappi ng) low temp. We i ghts by age/sex class AF FYF AM FYM .79 .49 .92 .61 .63 .27 .85 .50 .43 .41 .47 .24 .74 .68 .76 .51 .85 .88 .53 .65 .47 .48 .80 .85 .62 .60 .71 .71 .74 .86 .57 .67 .77 .51 .71 .54 .68 .79 .67 .6Lf .69 .64 .44 .47 .38 .66 .76 .54 �44 LITERATURE CITED Ankney, C. D., and C. D. Macinnes. 1978. Nutrient reserves and reproductive performance of female lesser snow geese. Auk 95:459-471. Bailey, R. o. 1979. Methods of estimating total lipid content in the redhead duck and an evaluation of condition indices. Can. J. Zool. 57:1830-1833. Bennett, J. R., and E. G. Bolen. 1978. Stress response green-winged teal. J. Wildl. Manage. 42:81-86. in wintering Campbell, R. R., and J. F. Leatherland. 1980. Estimating body protein and fat from water content in lesser snow geese. J. Wildl. Manage. 44:438-446. Carney, S. M. 1964. Prel iminary keys to waterfowl age and sex identification by means of wing plumage. U.S. Fish and Wildl. Servo Spec. Sc i. Rep. Wi Id 1. 82. i!7r'f). Chappell, W. A., and R. D. Titman. 1983. Estimating reserve lipids in greater scaup and lesser scaup. Can. J. Zool. 61:35-38. Connell, C. L, E. P. Odum , and H. Kale. birds. Auk 77:1-9. Davies, J. C., and F. Cooke. geese: prairie droughts 291-296. Drobney, R. D. 430-490. 1980. 1960. Fat-free weights of t 1983. Annual nesting productivity in snow and arctic springs. J. Wildl. Manage. 47: Reproductive bioenergetics 1 of wood ducks. Auk 97: 4 Evans, P. R., and P. C. Smith. 1975. Studies of shorebirds at Lindisfarne, Northumberland. 2. Fat and pectoral muscle as indicators of body condition in the bar-tailed godwit. Wildfowl 26:64-76. Folk, C., K. Hudec, and J. Toufar. 1966. The weight of the mallard and its changes in the course of the year. Zool. Listy 15:249-260. Handon, H. C. 1962. The dynamics of condition factors in Canada geese and their relation to seasonal stresses. Arc. Inst. North Am. Tech. Publ. 12:1-68. Harris, H. J., Jr. 1970. Evidence of stress response winged tea I. J. Wildl. Manage. 34:747-755. 1 in breeding blue- Johnson, D. H., G. L. Krapu, K. J. Reinecke and D. G. Jorde. 198. Condition indices for the sandhill crane and greater white-fronted goose. J. Wild]. Manage. (In prep.). t J �45 Jordan, J. S. 1953. Effects of s t a rva t i on on wild ma lla rd s . MilnacJc.17:30~-311. Korschgen, C. E. 1977. Breeding stress of female eiders Wildl. Manage. 41:360-373. Krapu, G. L. 1981. Auk 98:29-38. The role of nutrient ---:-- , D. H. Johnson, and C. W. Dane. J. Wildl. Manage. 43:384-393. reserves 1979. J. Wild!. In Maine. in mallard J. reproduction. Age determination of mallards. McLandress, M. R., and D. G. Ravel ing. 1981. Changes position of Canada geese before spring migration. in diet and body comAuk 98:65-79. Milne, H. 1976. Body weights and carcass composition W iId f ow I 27 :115- 122 . of the common eider. Owe n , M. 1981. lhe field. Abdominal plofile -- a condition J. Wildl. Manage. 45:227-230. index for wild geese In and W. A. Cook. 1977. Variations in body weight, wing length and condition of mallards and their relationship to environmental changes. J. Zoo 1., Lond. 183: 377-395. Owen, R. B., Jr., and K. J. Reinecke. 1979. Bioenergetics of breeding dabbJ ing ducks. Pages 71-93 ~ T. A. Bookhout (ed.). WAterfowl and wetlands -- an integrated review. La Cross Printing Co., Wisconsin. Palmer, R. S. Waterfowl (ed.). 1976. Handbook of North American birds, Vol. 2. (Part 1). Yale Univ. Press, New Haven. 521pp. Reinecke, K. J., T. L. Stone, and R. B. Owen, Jr. 1982. Seasonal carcass composition and energy balance of female black ducks in Maine. Condor 84:420-426. Ryder, J. P. 1970. A possible factor in the evolution Rossi goose. Wilson Bul I. 82:5-13. of clutch size in Siegfried, W. R., A. E. Burger, and P. G. H. Frost. 1976. Energy requirements for breeding in the Maccoa Duck. Ardea 64:171-191. Street, M. 1975. Seasonal changes in the diet, body weight and condition of fledged mallard in eastern England. Congr. Int. Union Game BioI. 7: 339-347. Szymczak, M. R., and J. C. Corey. Plains duck trap in Colorado. 1976. Construction and use of the Salt Colo. Div. \.Ji ld l . Div. Rep. 6. 13pp. Wishart, R. A. 1979. Indices of structural wigeon. Can. J. Zool. 57:2369-2374. size and condition of American �46 Zar, J. H. 1974. Cliffs, N.J. Prepared Biostatistical 620pp. analysis. by: ~K.~n~ Wildlife Researcher B Prentice-Hall Inc., Englewood �47 .. :: WYOMING I NEBRASKA .---------------~-------------------- VALMONT 1-_1 1 1 1 I . . . : C/) <t: C/) . . Z <t: ~ ... . .: :. ~: :' . .. . : 40 TRAPPING KM SITES ---------------,------------1 Fig. Ioca t 1. Map of eastern ions. Colorado, indicating the four winter OKLAHOMA trapping �48 400 • 300 -- • Ol • l=- e « u, 0 ur • 200 I=- 0 0 W 0: 0.. r 100 = 0.892 P < 0.0001 o 100 200 MEASURED Fig. 2. The relationship estimated using multiple collected during winter, 400 300 FAT (g) between body fat measured by extraction regression models, for Colorado mallards 1981-82. and �Ai 44 A 29 ADUL T FEMALE A AI JB IB 38 IB 57 AI AI ADULT IA 36 AI AI JC 40 I B,C A 40 A 40 sl I IA IB,C IB 40 I 125011501050950 WEIGHT JA 40 40 AI FIRST-YEAR IA,B 40 AI BODY JC AI40 AI 40 FEMALE I IB 40 FIRST-YEAR MALE I A,B 42 BI MALE IA (g) .' JC , I .16 .18 CONDITION I .20 I .22 INDEX Fig. 3. Mean body weight and condition index values for four age-sex classes of mallards measured in January, 1983. Within each class, bars represent (top to bottom) Bonny Reservoir, Kodak Ponds, Valmont Reservoir, and Segelke1s Slough; sample sizes are indicated in body weight bars. Means with the same letter within each class do not differ (l-way ANOVA and Duncan1s Test, p < 0.05). �50 0.. ~ W l- 60 LL 40 I o I 0 -20 8 X W 0 .20 2 43 Z Z .18 0 18 I- 0 32 .16 Z 22 0 o .14 12" snowfall 2 1 Jan. 1 Dec. DATE 1984 Fig. 4. Daily high temperatures (top) and mean condition index values (bottom) for adult male mallards measured at the Bonny Reservoir, Experimental Hunting Area Check Station, 1983-84. Dashed Iines indicate mean values; numbers by daily index means indicate sample sizes. �1400 e 1300 ~ 1200 0) .,_.... t- ::c: C) 1100 W I S b,c _/\ . b,c a 1M 1000 AF~ 900 I ,\., / ~ b IF '77 '78 '79 '80 '81 '83 '84 ·YEAR Fig. each 5. Mean body weights of mal lards captured age/sex group, means with the same letters at Bonny Reservoir during January, 1977-84. do not differ significantly (p > 0.05). \J1 With i n ��Colorado Division of Wildl ife Wildl iie Research Report October 1984 53 JOB PROGRESS State of REPORT Colorado ---------------------------- Project 45-01-506 Work Plan Job: Job Title: Field-feeding Period Covered: Migratory - 15050 Bird Investigations 16 Ecology of Mallard Ducks 1 April 1983 - 31 March 1984 Author: James K. Ringelman Personnel: J. F. Corey, R. C. Pabst, J. K. Ringelman, S. F. Steinert, M. R. Szymczak, Colorado Division of Wildlife; J. Barnett, A. Mendez, Colorado State University. ABSTRACT Thirty-four mallard duckl ings were hatched from 47 eggs collected in 4 Colorado locations: Wellington Wildlife Management Area, Monte Vista National Wildlife Refuge, the Fort Collins Area, and North Park. Ducklings were imprinted and pinioned shortly after hatching. Persistent snow cover on cornfields in the Fort Collins area precluded preliminary foraging rate experiments during winter 1983-84. Ducks were exposed to simulated field-feeding conditions within the holding pen, with favorable results. Equipment for next winter's trials was located, and seed corn was purchased for planting at the Wellington Management Area, the site for next winter's foraging experiments. ��55 FIELD-FEEDING ECOLOGY OF MALLARD DUCKS James K. Ringelman P. N. OBJECTIVES 1. Determine the relationships between feeding rates of mallards. and waste corn density, foraging group size, sex, and age of duck. 2. Relate post-harvest 3. Document 4. Develop a model for post-harvest mallards. cornfield the foraging treatments to foraging habitat requirements management efficiency. of wintering of cornfields mallards. for wintering SEGMENT OBJECTIVES 1. Review optimal foraging literature cable to the experimental design. for findings and techniques appli- 2. Obtain eggs from wild nests, then rear and imprint ducklings as experimental birds. for use 3. Conduct preliminary foraging rate trials to test methodology birds to field-feeding situations. and expose ~ETHODS AND MATERIALS Nest searches were conducted in the Fort ColI ins area, North Park, and the San Luis Valley to obtain mallard eggs for incubation. Thirty-four mallard ducklings (18 females, 16 males) were hatched and reared from the 47 eggs collected from 7 nests. Of these, 1P (representing 3 nests) were collected on the Colorado Division of Wildlife's Wellington Wildl ife Management Area, 8 were obtained from a nest found near Rocky Ridge Reservoir, 3 originated from a nest on the Monte Vista National Wildlife Refuge in the San Luis Valley, and the remaining 5 came from 2 other nests in the Fort Collins area. The diverse origins of these experimental birds was deemed desirable to aid in evaluating the heritability of certain foraging traits. All duckl ings were reared successfully to adult size and acquired alternate plumage by mid-September. Numbered leg bandettes were affixed to all birds to enable histories to be kept on all individuals. Within 4 hours after hatching, ducklings were imprinted to the principal investigator through intensive bouts of following and brooding behavior. The right wing of all duckl ings was pinioned at 5-7 days to render them permanently flightless. �56 Young ducklings were fed a ration of Turkey Starter crumbles (23% protein) until 5 weeks old, at which time a pelletized laying ration was gradually added to the food. By 2 months of age, all birds were eating the pelletized ration exclusively (16% protein). In late August, whole kernel corn was added to the pelletized ration (approximately 10% by volume) and broadcast in the pen enclosure to encourage birds to pick up and consume corn as a food supp Iement. RESULTS AND DISCUSSION Record setting snowfall and cold, which began in November 1983, continued into February 1984. Cornfields selected as trial areas to test data collection methodology were completely blanketed with snow until late January, and held residual snow in furrows between rows until 8 February. The unusually persistent snow cover and subsequent work schedule confl icts made it impossible to conduct preliminary foraging rate trials as described in approach 4. However, progress was made in preparing for the foraging rate experiments to be conducted next winter. Penned birds were exposed to corn kernels under simulated field-feeding conditions, and responded as expected to changes in corn density and ground litter. A seed spreader was located and tested, and found to be satisfactory for dosing experimental plots. Seed corn was purchased, and arrangements were maJe with Northeast Region personnel to plant corn on the Wellington management area and to apply the desired post-harvest treatments. Nesting areas within the pen enclosure were establ ished for purposes of obtaining offspring from experimental birds to serve as first-year birds in next winter's foraging trials. Prepared b~1(~ ~ James K. In Wild] ife Researcher B �Colorado Division Wildlife Research October 1984 57 of Wildl ife Report JOB PROGRESS State or REPORT Colorado --------------------------- Project Work Plan 45-01-506 2 ---- Job: Monitor Job Title: Bird Investigations 9 ---- Banding of the Shortgrass Population Period Covered: Migratory - 15050 January in Southeastern and February, Prairie Canada Goose Colorado 1984 Au thor: Gerald M. Lorentzson Personnel: D. Lewis, J. Lorentzson, J. Marble, J. Russell, J. Slater, P. Tucker, B. Will, Colorado Division of Wildlife ABSTRACT Trapping and banding efforts in southeast of 20 large and 18 small Canada geese. Coloraoo resulted in the banding ��59 MONITOR BANDING OF THE SHORTGRASS PRAIRIE CANADA GOOSE POPULATIONS IN SOUTHEASTERN COLORADO Gerald M. Lorentzson P. N. OBJECTIVES The major objective of this job is to continually document, through monitor banding and analysis of recovery data, the annual and long-term status of the southeastern Colorado (Arkansas Valley) segment of the Shortgrass Prairie Canada Goose Population to provide a basis for annual hunting season recommendations and to contribute data to the Central Flyway Management plan for this population. SEGMENT OBJECTIVES 1. Band a minimum of 1,000 Canada geese in southeastern the post season period. Colorado during 2. Prepare and submit banding schedules, band recovery and return reports. 3. Prepare progress report. METHODS AND MATERIALS Geese banded in southeastern Colorado in 1984 were captured using cannon nets baited with wheat, milo and corn. The age and sex of the captured geese was determined through cloacal and tail feather examination. All banding results were forwarded to the U.S. Fish and Wildlife Service's Bird Banding Laboratory in Patuxent, Maryland. RESULTS AND DISCUSSION Efforts to band small Canada geese in southeastern Colorado met with little success in 1984. Winter storms and sub zero temperatures forced the small geese out early in December and they were unavailable for banding. Sites along the Arkansas River and at all the lakes in southeastern Colorado were checked for wintering birds. Lake Meredith and the Arkansas River from Pueblo to Manzanola held about 15,000 large geese and about 5,000 small geese. Two sites were baited but we could not attract any birds. Queens Lake held about 1,000 small geese but the area was covered by about 1~ feet of snow making the baiting and setting of nets impossible. One site at Turks Pond was baited and 38 geese were banded. of these birds may be found in Table 1. Ages and sex �60 Table 1. The number and location of Canada geese banded in southeastern Colorado, 1984. Males Location Adu 1ts Immature Adults Females Immature Turk's Pond Small geese Large geese 2 5 4 5 8 6 4 4 TOTAL 7 9 14 8 / / \ Prepared by: i . .' ... i i /'{i 4-'.I:'>ccci/.'" r. ' ., l/) Gerald M. Lore' zson & Senior Wildlife Biologist �Colorado Division Jiidiife Research October 1984 61 of Wild1 ife Report JOB PROGRESS State of REPORT Colorado Prcjec:::. ______~5-01-506 - 15050 2 J(::J Tit Geese le: J·::Jb: Period Covered: Bird Investigations 10 Distributional Inhabiting Migratory Characteristics of Some Populations of Canada Colorado 1 May 1983 - 29 February 1984 Author: Gerald M. Lorentzson Per sonne I: G. Bishop, G. Byrne, J. Corey, G. Claasen, D. Crawford, L. Crooks, 1<. Dillinger, J. Ellenberger, M. Etl, J. Frothingham, M. Gardner, J. Gray, W. Haggerty, T. Lines, G. Lorentzson, S. Porter, F. Pusateri, C. Reichert, J. Ringelman, B. Sigler, S. Steinert, M. Szymczak, J. Wagner, K. Wagner, P. Will, Colorado Division of Wildlife; G. Patten and staff, Arapaho National Wildlife Refuge. ABSTRACT The number of large Canada geese banded on production/brood rearing areas exceeded quotas in west central Colorado (312), North Park (176), Northeast Colorado (193), and South Park (27/{). There were no birds banded on production areas in northwest Colorado or on wintering birds in west central or northeast Colorado. ��63 DISTRIBUTIONAL CHARACTERISTICS OF SOME POPULATIONS OF CANADA GEESE INHABITING COLORADO Gerald Lorentzson P. N. OBJECTIVES 1. To document the wintering range and harvest distribution of Canada geese nesting in (1) northwest Colorado, (2) west central Colorado, (3) North Park, (4) northeast Colorado and (5) South Park. 2. To ascertain the breeding range of Canada geese wintering central and northeast Colorado. 3. To contribute data to the various Central and Pacific Flyway management plans for specific populations of Canada geese. in west SEGMENT OBJECTIVES la. Trap and band at least 150 Canada geese on production central Colorado. areas in west lb. Trap and band at least 150 Canada geese on production west Colorado. areas in north- le. Trap and band at least 100 Canada geese on production areas in North Park. 1d . Trap and band at least 150 Canada geese on production areas in northeast Colorado. 1e. Trap and band at least 75 Canada geese on production areas in South Park. 2. Trap and band at least 250 Canada geese on wintering west central Colorado. 3. Trap and band at least 250 Canada geese on wintering areas in northeast Colorado and take the measurements mentioned in the Program Narrative. 4. Submit banding schedules and recovery reports to the Bird Banding Laboratory. 5. Prepare progress report. areas in �64 METHODS AND MATERIALS Canada geese were trapped on production/brood rearing areas in westcentral Colorado, northeast Colorado, North Park and South Park during their fl ightless period using standard drive trapping methods. All captured geese were banded and released. RESULTS AND DISCUSSION Su~ner Populations Banding quotas were exceeded in all areas except in northwestern Colorado (Table 1). The Yampa and Little Snake rivers were flown to locate concentrations of molting geese, as in years past we were unable to find any concentrations of geese with broods even though nesting birds are observed during spring breeding pair counts. Winter Populations. Severe weather conditions forced most of the northeastern Colorado geese out of the area so I've were unable to trap or band after the 1983-84 season. No attempt was made to band geese in wintering areas in west central Colorado. Prepared by ~I" / ""V -.L- ,;), ,·(t l..(: , /VI' V~~~ ,\- rcz,/. -I,I cJ ~ ~I "Gerald M. Lor'entz6n Senior Wildlife Biologist J '. " ,_. 4 f �65 Table 1. The number of Canada geese banded on production/brood areas in Colorado, summer, 1983. Area/ Ioca t ion _.West-central.- Subtotal • , I LM 12 18 88 38 26 _7 159 12 11 4 - 1 28 33 8 13 7 61 2 24 10 _3 20 9 15 11 39 55 Toti)l 6 22 60 27 21 5 113 175 65 60 12 312 19 15 3 1 24 8 9 8 38 49 88 42 29 17 176 3 32 12 1 48 22 8 12 _9 51 15 7 North Park Waiden Reservoir Po 1e t-Iountai n Elk Pond McCannon Pond Subtotal Northeast Colorado Johnsons Pond Haxton Julesburg Reservoir Guenzi Project t It • r Number banded AF LF Colorado -------- S i It Radium Rifle Fingerock I M rearing Subtotal South Park Antero Reservoir Eleven Mil e Reservoir Subtotal TOTAL 47 73 49 24 193 unk 2 73 75 ____!1 160 Total -- __li 51 2 70 72 75 72 202 274 326 180 288 955 32 36 ��67 Colorido Division of Wildlife Wildl ,fe Research Report October 1984 JOB PROGRESS REPORT State Colorado ------------------------------ Project No. Work Plan 45-01-506 3 --~- Job Title: Period Covered: Job: Migratory - 15050 Bird Investigations 8 Monitor Banding of Eastern Colorado Mal lard Populations 30 April 1983 - 31 March 1984 Author: Gerald M. Lorentzson Personnel: C. Leonard and area personnel, D. Benson and area personnel, J. Young and area personnel, M. DePra and area personnel, J. Corey, G. Lorentzson, J. Ringelman, and M. Szymczak, Colorado Division of Wildlife. ABSTRACT A total of 4,059 mallards were banded postseason in thirteen locations in eastern Colorado in 1984. Banding records were entered on the computer and banding schedules were submitted to the Bird Banding Laboratory. ��MONITOR BANDING OF EASTERN COLORADO WINTERING MALLARD POPULATIONS Gerald M. Lorentzson P. N. OBJECTIVES 1. To establish mor.ltor banding of wintering mallard populations ee s t ern Co lcr ado as an annual management function. in 2. To continually document, through monitor banding and analysis of recovery d2ta, the annual and long-term status of eastern Colorado wintering mallards to provide a basis for annual hunting season recommendaTions. SEGMENT OBJECTIVES 1. Band a minimum of 4,000 mallards during t hc post-season period including a minimum of 500 birds in each of the following general areas of the South Platte Valley and Arkansas Valley: (1) Denver-Greeley, (2) Fort Col 1 i ns+Love land-w i nd sor , (3) Greeley-Fort Morgan, (4) Fort Morgan-Sterling, (5) Sterling-Julesburg, (6) Bonny Reservoir, (7) Manzanola-Lamar, and (8) Two Buttes Reservoir area. Divide the banded samples in each area equally among the four age and sex classes. 2. Submit banding schedules and recapture data to Bird Banding Laboratory. 3. Conduct an updated analysis of band recovery data, including the following major d e t errnlna t i on s for important population segments of mallards wintering in eastern Colorado: (1) distribution of harvest, (2) recovery rates, and (3) survival rates. 4. Prepare progress report and publish pertinent findings. METHODS AND MATERIALS Salt plains duck trap or modifications thereof were used to banded in this segment. Thirty-seven were banded on Turk's to goose trapping, using a cannon net. The research section on Bonny Reservoir and the rest were banded by the Southeast Regional personnel. capture the ducks Pond, incidental banded the ducks and Northeast RESULTS AND DISCUSSION A total of 4,059 mallards we re banded postseason in 1984 (Table 1). Quotas were almost met in all categories with the exceptions of adult females (-25) and immature females (-70). The weather was exceptionally cold during the period of banding resulting in concentrating the ducks on areas of warm wa ter sloughs. �7e Table 1. Mallards banded postseason banding areas, January 1984. AM Number of ducks banded Age and sex AF SF SM 81 0 153 137 137 50 145 76 9 157 132 123 50 130 703 677 150 92 125 0 32 124 1 94 123 5 28 399 1,102 Banding area Northeastern Subtotal 80 16 112 88 95 50 106 300 53 500 500 447 200 514 2,514 63 28 78 143 92 50 133 587 547 375 198 26 55 77 13 19 388 93 10 78 146 23 33 383 1 ,545 1 ,052 975 930 4,059 Colorado Bonny Reservo i1Turk's Pond Verhoeff Reservoir Las Animas Ponds CJay Pond Ma 1 on e S 1 oug h Subtotal GRAND TOTAL () Prepared by: Total Colorado Valmont Reservoir Propst Slough West Tamarack Shaefer Ditch Sege 1 ke Slough Ches tnu t Slough Kodak Ponds Southeastern by age and sex in eastern Colorado v. 1/// !,}J~ t( ~ AJA/cJd'J /Gerald M. Lorentz on Senior Wildlife Biologist 565 37 319 471 41 112 �Colorado Division Wildl ife Research October 1984 71 .f Wildlife Report JOB PROGRESS State REPORT Colorado ----------------------.-------- Project No. Work Plan 45-01-506 3 Job Title: Job: Migratory - 15050 Bird Investigations 10 Some Population Characteristics of Adult Gadwall Molting in North Park ---------------------------------------------------- Period Covered: 18 August 1983 to 31 March 1984 Author: Michael R. Szymczak Personnel: J. Corey, J. Ringelman, Division of Wildlife. S. Steinert, and M. Szymczak, Colorado ABSTRACT Trapping of gadwall (Anas strepera) in North Park resulted in the following number of birds being banded: 266 adult males, 212 adult females, 93 local males, 81 local females. The use of recaptures and returns in combination with recovery data improved the precision of survival rate estimates substantially. ��73 SOME POPULATION CHARACTERISTICS OF ADULT GADWALL MOLTING IN NORTH PARK Michael R. Szymczak P. N. OBJECTIVES 1. To document the distribution of harvest of gadwall YOl;ng or mo lt i nq adults in North Park, Colorado. 2. To estimate recovery and survival North Park, Colorddo. banded as flightless rates of adult gadwall molting in t SEGMENT OBJECTIVES • 1. Trap and band 400 adult males, 350 adult females and, if avai lable, up to 100 local gadwall in North Park on molting and brood rearing areas in North Park . 2. Submit banding schedules and recapture reports to the U.S. Fish and Wildl ife Service's Bird Banding Laboratory. File return information at the Colorado Division of Wildlife Research Center. 3. Recovery, recapture and return data will be analyzed to determine the adequacy of the banded samples. Recommend changes in sample sizes or duration of banding program if needed. 4. Prepare progress r report. METHODS AND MATERIALS Al I birds were captured from an air thrust boat at night using a hand-held 12-volt landing light and a long-handled net. Nearly all flightless adults were captured, weighed, and measured prior to being banded and released i.n conjunction with a study of the flightless period in ducks (Szymczak and Ringelman 1984, in press). The band numbers of birds captured that had been previously banded were recorded. Banding schedules and recapture reports were prepared and submitted to the U.S. Fish and Wildlife Servicels Bird Banding Laboratory. Information on returning birds recaptured in the same la-minute grid of banding were filed at the Colorado Division of Wildlife Research Center. Recovery and return and recapture matrices were constructed for adult males and females banded since 1975 and recovered through the 1982-83 hunting season or recaptured through the 1983 banding seasons (Mardekian and McDonald 1981). Banding and recovery matrices, banding and recapture and return matrices and the 2 matrices combined were subjected to program ESTIMATE (Brownie et al. 1978) to evaluate the degree of precision of the survival estimate using the specific types of band encounters. �RESULTS Band ing Over 470 adults and 170 ducklings were banded in North Park in 1983. Birds were captured on 4 areas but more than one-half the birds were trapped at Walden Reservoir (Table 1). Table 1. 1983. Number of gadv~a11 banded in North Park, August through September Adul t males Locat i on Walden Reservoir MacFa,1ane Reservoir Lake John Annex Pole Mountain Reservoir TOTALS __ --_--_--------_._ Age and sex Adult Local females males Local females Total 144 115 50 37 346 54 55 20 15 144 59 25 8 11 103 9 17 15 18 59 266 212 93 81 652 Quotas of 400 adult males and 350 adult females which were established in 1980 have not been met since that time and are probably unrealistic considering trapping results the last 3 years. The number of adults trapped annually since 1981 has been 338,320 and 260 males and 210,191 and 212 females. Survival - Use of Returns and Recaptures Mardekian and McDonald (1981) described the combined use of recoveries and recaptures encounters to improve survival estimates by using the models described by Brownie et aJ. (1978). Since adult gadwall apparently consistently util ize molting marshes from year-to-yec,r and are commonly captured in subsequent years after banding, recaptures were combined with recoveries to determine the level of increased precision which might be expected. Using the data from model 1 (Brownie's et al. 1978) as an example indicates use of the combined data increases precision substantially (Table 2). Male mean estimates are approaching a coefficient of variation of .05 at which survival can be estimated + 5%; the objective precision levels for the study. Female estimates, which are useless if recoveries only are utilized in this case, are much better using the recapture data. The inability to reach banding quotas for adults will hamper achieving precise survival estimates for this study, however, obviously the use of return and recapture data will enhance those estimates. ,• �75 Table 2. Example estimates of mean annual survival of adult gadwall banded in North Park since 1975 using recoveries, recaptures, and recoveries and recaptures combined. Recovery rate Parameter Arithmetic mean survival (± SE) Recoveries 2.28 55.09 (± 4.23) Recaptures 2.36 69.51 (± 5.40) 4.58 63.96 (± 3. 15) Recoveries 1.81 81.98 (±26.86) Recaptures 3 . 11 67.45 (± 8. 18) 4 .. 80 67.77 (± 6.25) Encounter type Males Both Females Both LITERATURE CITED Brownie, C., D. R. Anderson, K. P. Burnham, and D. S. Robson. 1978. Statistical inference from band recovery data - a handbook. U.S. Dep. Interior, Fish Wild!. Servo Resource Publ. No. 131. 212pp. Mardekian, S. Z., and L. McDonald. 1981. Simultaneous analysis of bandrecovery and live-recapture data. J. Wildl. Manage. 45(2):484-488. Szymczak, M. R., and J. K. Ringelman. 1984. fl ightless period of ducks in Colorado. Rep. Oct. Prepared by ]?ULft~J Michael R. Sz~~ \.JildlifeResearcher C Ecological studies of the Co 1O. D iv. Wi I d I. Wi I d I. Res. ��Colorado Division Wildl ife Research October 1984 77 of Wildlife Report JOB PROGRESS Colorado State of Project REPORT 45-01-506 - 15050 ------~----~----~~------ Work Plan Job Title: 3 ---- Job: Banding Migratory Bird Investigations 11 of Preseason Waterfowl Populations in the San Luis Va lley Period Covered: 1 July - 15 November, 1984 Author: Gerald M. Lorentzson Per sonne 1: M. Nail and staff, Monte Vista and Alamosa National Wildlife Refuges; J. Corey and G. Lorentzson, Colorado Division of Wi ld I if e . ABSTRACT A total of 1,439 mallards and 300 other ducks were banded in the San Luis Val ley during August and September 1983. We were 17 birds short of making our quota on adult female mallards, but were over our quotas on the other mallards. There was a marked decrease in other species banded this year, especially on pintails, blue-winged - cinnamon teal and green-winged tea 1. ��79 BANDING OF PRESEASON WATERFOWL POPULATIONS IN THE SAN LUIS VALLEY G. M. Lorentzson P. N. OBJECTIVES The objective of this job is to continually document, through monitor banding and analysis of recovery data, the status of the San Luis Valley preseason mallard population. SEGMENT OBJECTIVES 1. Trap and band at least 300 mallards of each sex and age group, adult males, adult females, immature males and immature females. 2. Band all other waterfowl mal lard trapping. 3. Submit banding schedules and recapture reports to the U.S. Fish and Banding Laboratory, file resulting recovery WildlifeService'sBirj cards at the Fort Collins Research Center. species captured during the period of 4. Prepare Progress Report. METHODS AND MATERIALS Ducks we re trapped in the San Lu is Valley from early August through mid-September, 1983. Mallards were the target species but all other ducks captured were banded. Salt Plains types of duck traps were used in the banding p roq rarn for capturing the birds. Barley was used for bait. Ducks were banded and recorded according to age and sex. All banding reports were sent to the U.S. Fish and Wildlife Service's Bird Banding Laboratory. RESULTS AND DISCUSSION Ducks were banded at Russell Lakes, White Mallard Club and at the Monte Vista Refuge during August and September, 1983. There was more water available in 1983 than in the three previous years. The Blue-winged Teal Club and parts of the White Mallard Club have such high numbers of carp churning the water that ducks were not using the ponds. The San Luis Lakes area had a lot of construction work, such as road building and well drilling, so we did not set traps in that area this year. Banding quotas for adult and immature mallard males and immature mallard females were easily obtained but the quota for adult mallard females was not reached. In most cases the adult mallard females either hadn't molted by September 15th or they had not regrown their fl ight feathers enough to fly. There was a sharp decline in the numbers of other ducks in 1983, especially in the number of pintails and teal on the areas. �80 Table 1. Number of ducks banded, by species, du ring the pre-season period, 1983. 1 in the San Luis Val ley Species Mal lard B Iue-w inged and/or Cinnamon teal Pintail Redheads Green-winged teal Total 1 Includes ducks banded National Wildlife Refuge" If Prepared by / / ::,/',1 .',.\ 'I' Age and sex IF LM AM 1M AF 337 336 283 474 25 25 3 5 82 21 6 5 8 395 450 by personnel 7i"" H\ ,~,= ~,h~ /1 <c )--- ','.'.:'J..Ii' ",' Gerald M. LorenYzson I" Senior Wildlife Biologiijst LF Total 7 2 1,439 1 71 16 11 5 0 0 2 0 0 0 0 0 186 73 23 18 306 577 9 2 1,739 11 _3 of the Alamosa '--'" .' and Monte Vista �Colorado Division Wildlife Research October 1984 81 I;Jildlife Report .f JOB PROGRESS State of Colorado Project 45-01-506 6 Work Plan Job Job Title: Migratory - 15050 Bird Investigations --- Migratory Period Covered: REPORT Bird Publ ications 01 Apr! I 1983 - 31 March 1984 Author: Howard D. Funk Personnel: C. E. Braun, J. K. Ringelman, Colorado Division of Wildlife; P. D. Curtis, T. E. Olson, R. A. Ryder, Colorado State Univers i ty. ABSTRACT Work was continued and publ ications accompl ished or submitted job during the segment were as fol lows: under this Curtis, P. D., and C. E. Braun. 1983. Radiotelemetry location of nesting band-tailed pigeons in Colorado. Wilson Bull. 95:464-466. ----;:- , and 1983. P.ecommendat ions for es tab Iishment and placement of bait sites for counting band-tailed pigeons. I;Jildl.Soc. Bull. 11: 364-366. --:=-:=--- and effects 54:381-386. , and R. A. Ryder. 1983. Wing markers: visibility, wear, on survival of band-tai led pigeons. J. Field Ornithol. Olson, T. E., C. E. Br aun , and R. A.. Ryder. 1983. Cooing activity and nesting of mourning doves in northeastern Colorado. Southwestern Nat. 28:335-340. Ringelman, J. K. 1984. Prepared 1983. Waterfowl Why band birds? mysteries. Colorado by Howard D. Funk Wildlife Research Leader Colorado Outdoors Outdoors 33(2):6-8. 32(6):4-6. ��Colorado Division of Wildlife Wildlife Research Report October 1984 JOB FINAL REPORT State of Colorado Project 45-01-506 Work Plan Job Title: Recovery, 8 Migratory - 15050 Bird Investigations Job: Computerized and Recapture Per i od Covered: System for Storing and Retrieval of Banding, Information 01 Apr i1 1983 - 31 March 1984 Au thor: James K. Ringelman Personnel: J. F. Corey, J. K. Ringelman, Division of \.Jildlife. M. R. Szymczak, Colorado ABSTRACT A computerized system for entering, storing, and retrieving banding data has been implemented and is fully operational; Three sub-systems provide flexibility in data entry and retrieval: archive files, data entry routines, and data processing programs. Archive files consist of over 260,000 records of Colorado-banded waterfowl, 20,000 records of hunterrecovered or foreign re-trapped waterfowl, and nearly 5,500 records of banded waterfowl recaptured in the same 10-minute grid in which they were originally banded. Eight wa t e rfow l species, sorted by year of banding or recovery, are represented in archive files. Data entry routines include programs to enter and re-format new banding data for entry into the archive records and to generate reports and sum8aries of banding information. Ease and accuracy of data input is facilitated by prog rams that a IIow for "short-hand" en t ry of data, check data for keypunching errors, then expand short-hand data into a format suitable for entry into archive fi les. Other programs generate banding schedules and tapes suitable for submission to the Bird Banding Laboratory. Data processing programs tally banding records to produce reports of bandings by species, location, and sex/age class, and search archive records to match newly-recaptured birds with banding records to derive banding histories. ��85 COMPUTERIZED SYSTEM FOR STORING AND RETRIEVAL OF BANDiNG, RECOVERY, AND RECAPTURE INFORMATION James K. Ringelman Colorado Division of Wildlife personnel have banded about one-quarter of a mill ion waterfowl in the past 20 years. Banding data continues to provide a useful tool for measuring survival rates and ascertaining harvest distributions. It became apparent that a computerized system for storing and retrieving banding data could increase efficiency, reduce errors, and provide additional information to researchers not readily available from the U.S. Fish and Wildlife Service's Bird Banding Laboratory. This report describes the structure and use of data files and computer programs developed specifically for the banding data storage and retrieval systems. P. N. OBJECTIVES 1. To establ ish a computer system for banding and recovery data. 2. To enter all necessary data from past and future banding programs into the system for accurate and efficient record keeping and analysis and reporting programs. METHODS AND MATERIALS The Gold Computer at Colorado State University was used to read and write computer tapes and for disc storage of data. System software procedures Sort and COPYT were used to manipulate banding information. Data reside on 5,2400' magnetic tapes stored at the CSU Computer Center tape library and accessed remotely via a terminal located in the Research Center headquarters. All computer programs were written by the principal investigator in Fortran 77 (Control Data Corporation NOS 2.1, FTN5). Listings of all programs are presented in Appendix A. RESULTS AND DISCUSSION Archive Files Archive files were initially produced from data tapes provided by the Bird Banding Laboratory. Banding tapes were read on the Gold Machine at Colorado State University, stored on disc, sorted by species, latitude-longitude, and date, and written back onto computer tape. This resulted in three tape files which contain all records of bandings from 1967-81 (Table 1). As birds are banded, additional records are added to Tape 2 (mallards) and Tape 3 (all other waterfowl). All data are written in EBCDIC on 9-track tape at 1600 BPI. The labeled, unblock tape is formatted as described in Table 2. �86 Recovery data were also obtained from the Bird Banding Lab then read, sorted, and re-written back onto tape in a manner similar to banding tub files. The result was two tapes (Table 3) containing all mallard recoveries (Tape 4) and recoveries of other waterfowl species (Tape 5). These data are kept current using annual tape updates provided by the Bird Banding Lab. Data are sorted by species, then date of recovery, in ascending order. Line format is identical to that used by the U.S. Fish and Wildl ife Service (Table 4). Nearly 5,500 returns (birds recaptured in the same 10-minute grid within which they were originally banded) were coded from field data forms, submitted for keypunching, and read onto tape. New techniques allow these data to be pooled with standard recovery and recapture data to provide more precise estimates of annual survival rates. Storage on tape assures ready access for analyses of this type. Line format of return data (Table 5) is simlar to recovery information. A data file of active trapping sites, which presently includes 72 locations, wa s built for Lise as a "look-up" file accessed by subroutines in the banding schedule generator and banding summary programs. Each entry consists of a unique, 4 digit wetland code, latitude and longitude, description of the trapsite in reference to towns or landmarks, and the name of the trap site (Fig. 1). Using only the wetland code, both the schedule and summary programs access the other data fields used in generating reports. This file is updated as new trapping sites come into use. Data Entry Routines Entry of new banding data remains the most time-consuming part of this largely automated system. To minimize time spent in this activity, provisions were made for "short-hand" data entry format. This format takes advantage of the fact that a banding crew often bands birds of the same species, sex, and age with consecutively numbered bands. When differences are encountered among consecutively numbered birds, they often involve changes in age, sex, or both. The first two columns (contro] field) foilowing the line number contain codes instructing the decoding program what to change between successive birds. For example (Fig. 2, top), "NLiI is used to indicate the start of data or any change which involves more than age, sex, or both. In line 1, bird banded with number 1377-04501 (AOU = 132.0, status = 300) is an ASY male banded on wetland #4215 on 20 January, 1982. Two blanks ( ) in the control field (as in lines 2 and 3) instruct the decode program-that birds 1377-04502 and 04503 have the same characteristics as the previous bird 04501 (i.e., "no change"). A IIA" in the control field instructs the program to change only the bird age, leaving everything else the same. The new age is listed immediately following the IIA". Thus in line 4, �87 the age is changed to "SY" from "ASyll, so bird 1377-0lr504 is an SY male banded on wetland 4215 on 20 January 1982. To change sex, a S" is entered in the control field, followed by 3 spaces, then the new sex (e.g., 1 ine 14). An "AS" in the control field, followed by the new age and sex, changes both characters. New lines (NL) must be entered in cases of a new trapping wetland (e.g., line 21) or date change, and to indicate replaced bands as shown (e.g., line 7). The expanded data file, after the short-hand file is processed by the decode program, is shown at the bottom of Figure 2. When entering 4,000-5,000 bandings typical of a winter banding program, the savings in time realized with short-hand entry is appreciable. II An error check program checks for obvious or typographical errors in the data file. If an error is detected, a diagnostic message of the problem (along wi th the 1 ine number on wh ich the error was detected) is pr inted. Nine parameters are checked with the present program (Table S). In addition, hand verification of data is performed prior to submitting records to the Bird Banding Laboratory. Banding schedules, which are required of all banders by the Bird Banding Laboratory, are produced by a computer program using the decoded data file (Fig. 2, bottom) as input. In addition to listing the characteristics of all banded birds, it also indicates replaced bands, latitude-longitude, and descriptions of trap sites (Fig. 3). The latter two variables are determined by use of the trap site data file (Fig. 1). The overall format of the schedule conforms to rigid standards set by the Bird Banding Laboratory. Cost of computer time and paper for schedules describing 1,000 bandings is $3.12. A computer tape of banding records is submitted, together with computergenerated schedules, to the Bird Banding Lab. In doing so, the data can then be entered directly into BBL data files without the need for keypunching and verification. The decoded, raw data file must first be re-formatted to conform to specifications (Table 2) before being written to tape, and a special util ity program was written to accompl ish this change. Data are also written onto archive files prior to being purged from disc storage. Data Processing Programs Summary tables describing the number of birds banded by location are needed for internal reports and record keeping, and for evaluating the success of banding operations. These reports were previously produced by timeconsuming hand tallying methods. The new system uses a Fortran computer program that accesses raw data files and the trap site look-up file to produce a computer-generated summary (Fig. 4). The summary 1ists the number of birds banded by age/sex class and location for each species. �88 A second data processing program derives the history of recaptured birds by matching a list of "recaptured" band numbers with a banding archive file likely to contain data on the original banding. Before building the archive file, the user should consider the species recaptured as well as the survival rate expected for banded birds. As an example, a winter banding program in eastern Colorado would result in mallards comprising over 95% of all recaptures. Moreover, with an average annual survival rate of about 60%, less than 8% of a banded cohort would be expected to be al ive 5 years after banding. The desired archive file should therefore consist of all mallards banded during the previous 5 years. This increases the efficiency of the search by restricting the archive to 20,000-30,000 birds. Though somewhat expensive in computer time, the search program reduces from days to minutes the time needed to determine banding histories. The user first builds a file listing the band numbers and wetland code for the recapture site (Fig. 5, top). The archive file is built from data files on tape s toraqe (e.g., Fig. 5, bottom). The search program then I ists not only the date and place of the original banding (Fig. 6), but also whether the bird was a return (recaptured in the same 10-minute grid as banded) or recapture. This last distinction is important, since returns, unl ike recaptures, are not reported to the Bird Banding Laboratory. The program also I ists bands for which no history was found. These presumably represent birds banded out-of-state or birds whose history predates the archive file. Manual determinations of history may be required for these cases. Prepared bY,(/C_'_ \~~K. Wildlife r::t:~ Rin lma Researcher B �o OOO-O(('(l ?002 400-1 (,< 1. ~."1I. :1401 4~4-11)"" ~lJ ~41(') 3407 340~ 3409 0,,,2 ?403 3404 3405 041)6 3702 270< 3705 405-11'", 40~-g~2 ,Qf 401-10,,7 40 '1-1 C'''? 404-1:,1., 4(l?-Hfl 404-1(11'1 404-10": 4C4-1 ""1 40~-lnz4 40<-1(.7' en 405-1 '''nG 404-~');'~ 37]0 ., f'(\ ~ I ,'11' l.?Oi; I, 4204 4'14 , , '1 ? t •• r • l ELLl[l~' rCheN uL,". t'(I\[,l!\r COILMC~r. t.1~ 11 ~~ Lr LF L l(~C~cy, ••L~'·y, ~AccARLAN~ r.~: :,,' \J 0\ , !~ AL r " r, , J /10 ( RES •• 12 S ~F 5 C t-. l L 0 9 \. l K J t CKS '~~ I ": : ~;• J s, L ~ ~ c'" c L t-- 'OAST TAf'lHACI<, J' I ~,)oJ K,;,.. <, J I.'" 'I:j ~,~So, " (8 t.F L 1 1u·" ", 00 l.'10 4211 4~(" l.~j~ "~l' 1',1 C? ( !.o(. ~ 1"- 0 -' l.Q?-10'·l. l.c\~-l "'l. 'l 4 (I ?-l "l.) 4C1-1P4~ l.I'1-1(,40, 401-1(,41 4 -,':-::-1 ,'"4~. 4,'" -~ "l.' ~z \oI_LC ',. •. l~: c :. Y J <;cV'~~?r;C~, ~-:\::Kt,i,i_:- k' -: f ·..J!"P)L;'., r ,. 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'; II T:, i L ,1 I 1" ,., " h ~ u F ,..,1-;, T ~ ~ r i~ 4'1':> 4CI'-1rf··1 4 r S -; .: ? ., .J I! '.' 'J c: /.: j ., '1 ~ 1 t " i· ~ L I- (..R L':'r-., C:' 3 <;~_1 .. ':': O,\:T.\"", 0"1 (1] b;".t:,i., l;_., CL ronl 4nO-1 rl,~ c ! '"r :. r L C K, J :, \'..r '( r-' .' t, '"L C U f .•.•,~ l ("~L , i t l t !'-" J. F L l' r: ,.... r?04 3G3-1r74 1~1? !t (' '#_ 1..1'. I Ii '. i., J •..:_r< ~,:~", C L o! C (_ ',)06 4<)"-1 "1'1 H'YT~~. P~lLLl~~ Cu., CO ~"0~ 'l ~ ,~,- J ~ .,., l·', eTC' :, l d 0" •. h _'f ;,; I:' Tc.t, c:~q(]7 3~n-1(?' L \ ~ : L ~ ) L .~L U', 4 iii '_ f (, ~ 1 ~ 1 U, " r. ~7')7 4r'-100? v - ;: r . i. , L f.;. f'. t l • ILL ~NIM~~, Et~l CO., CD 450" 31'("-10'l1 l~S ,,~~o, ;"", C,. ~1 I' 1 ;., " .) ~ l_ Figure 1. Trap sites (. data III file. " C j\ RANt" ~ :":1-,,= f-ei< Gl I c+ :d,.:LI', SL£.:uCI\ .. r"::'i,'S ::'lCL(;H JCI-i',:iLN r: "" L ~ 1 'I r (ril.JTto.LT FCr.C k f. ::. • :LULlr SLuLllS ::LCt.;C,1- k~'MLrZ~LL \..'-~. ~-" ~LLL(t- rt~. I\~'tll- c' •. 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L \,.tl[ t ~ ., ::, .: !-- , 3 PlJl\L !-LATS f-'_iKI:~' ~Ct.(~ F;:L~ r r s , H:~. I t. ~.: J,j;-'~ ~ALf"'LANc Id:S. f~~c en ~ S L L L (_H , p S. DL,ll. 1,.' '·'ILee( ~~1'11 3e5-J055 '"'"Of. 3f5-1"5' ",.)"4 1 (' .:) 7 K"t':.cy, 39'1-1c?n ,C,-1C7' ~71 T, ? 01 -1 '''r ') ~6(W, (40'5 390-10°4 l- j (,'1. 40n-1(\'1) ,.401 "1'7'-:,1" 4503 4~04 5701 f'? L L t_.lr~~~~;::, 0', t. '! Y r' 4Q2-104~ t...~4 ,C"-' . "7 ~4i)2 K A I" '" ; • Z ~ L:;; \'"L~. ,_ F ~ l LDr' ~,c 1 ~T itS. (.Hd, LA~': (1-/" Fl~,;~t ~ ~~ Lf l~GLK. on ~.5 ~~ OF STLkll~G, GUf~Zl kANC~. SCI" ~ t f "K (.J ~ALn-~, (.F C"O[K. L;; l ~K, l r L ~ L C k, ([ u- r V,\L·~ F'_:tT ("::10 \. l SE ~'l '" C~ > C[ t, ., (.:.:., 1 TA~A~t(k. r C~., C JACKSCN r: 1 I L r ,_~ -I, v t.CK S L N C L ., l C"ll"'jl\il, JAlK$[1\ Ll:., L L -\ I-'C J L H:"i, : Lt·. 'r, [., F ~ Co l L ;...I ~~'T 401,-IO'l1 40l-lt"40'-10'1' (U., L~K~, <:" t: C L ~ t ~ L L i.. {- ~'. 4 r, ~ '- r h'( L L>: :' :; c, r:r 401-;(,i F l' T ~ L' G ~ 1\. ,', L ~ (~1. t!o., C C 4301 405-1,)~~ J'lh N S 0, i' u t; (' , 3 ~ t '"I- LV lOp CO 402-10'P p;, r- w I 1 1 :-. '. ~ L t!'" ~ r·.l r" (. , C r. 4101 .L!jLd-, 3 ;,L lr ~llLLr;(O~',(r, 401-10l.4 cL.!f';r:,Lf 3p·04 3 ~r'5 ~ ,J,t, 1L :. r L '- L G b I". r T L'-K~ I-llrlLl:n ~'HL". i,1TCr, ~ALKt~ ~lL~l. tKcA ~.H l L T ,~ f t. r, C I- ,_ ;oU,[, DLTTt:: I<i..:'. JChN ~ARllr; ns. VtrfJd-f ~LS. l I'Fc~ QL~~N'S ~~~. ~ l c N[;,hL ~.E~. toNIlKlJ RE:!i. i . Mil" hS. lC~~A ULtlN'~ RlS. Tu~K r «. l L •. 0''1" ~ i:. r, ••,.. p-", TtAL lLUIo l U :..r, i>ChL c~r L"t<.c:' h:p:) l:l,K 5~1, Ll;lS L LAKt: SMlir-: :...cs. rU~Llt"Ht'S ~"lrLt~ LAn leller. ns. ~Lt--T~ VI'::T t: r ~ ...:r- r: ~TS~'_LL • 1 ~ I A ~ ~ r< JALK;,C:', l"K t ~LCL(;h ;'L.,lT~ Li:.K(~ lioF~(r 5CLrri r<. .! L t t: \..' .-<- ~ ~ • ~AL~.l!r jlb~l. H"':~.\ i If,,; ".j(~ (H.!~ fC~.t: <. l;.H ..U; f·tJl,L, c"l~ V I-A X T ~ I.~,1 r L •• ;.. 1 , 'L" us ~ 1 r--l L[l.(1- LGt.;(;f, l LL (; r, "S HATCh. �90 = -:-ggggiNl 00003 00004 .\ SY 00005 AASI 00006 131704507 00007 NL 00003 0000'1 A SY 00010 AASI 132J 300 ASI M 4215 012082 REPLACES 1077-562~5 REPLACoS 717-93756 88gB 00013 OJJl ~ 0001~ co o i e F A SY HSY A 51 AA5V - 88gt~ 00019 4 51 A AS'( 00)20 . 00021 ~L 137704521 S .. '1 00022 00023 A SY OU024 AASY oa025 OO')Z 6 gC027 0028 1320 300 ASY M 4102 0120e2 A 5Y AASY 00:>29 137704530-1320· 00030 NL 001)31 001)32 A SY AASY 00033 00034 NL ·137704534 1370 AASI 00035 00036 1)0037 00038 ';---1'. 00039 00040 Nl 137704540 1320 M 00041 S 00042 00043 AASI 00044 A 5Y 00045 AASY 00046 00047 Nl 1377C4547 1320 (lOa48 00-.,49 00050 300 ASY 4102 012082 300 ASY M 4102 012082 300 ASY F 3601 0120a2 4 • 300 .SI F 2802 012e62 4 f • 1 g8gg~~~BH8f~8~T~E8 ~88iH ~ ~~B 8!:~8:R~ 00003 137704503 132.0 300 ASY M 4215 01-20-ez 8ggg;--l~t+g~~g; i~~:8j58 4~1~ 8i:~2:~~ 8gg89--1IJJg:;g9-1~I:g ~gg :~, ~ ~~l~ gl:18:~~REPLACES 00008 13770450B 132.0 300 ~SY M 4215 Ol-2G-82 A~~ C0009 0001(1 00011 00012 13770450'1 1371U4510 137704511 137704512 132,0 132.u 132.0 132.0 ~ 300 5Y M 4215 30~ AS' M 4Z15 300 AS' M 4215 30u ~sv M 4215 1071-56295 JI-20-~2 ~1-20-e2 01-20-82 vl-2u-c2 8gg1~ ij~t8~§l~ i~~:g~3g :g~ ~ 4~!5 gr:~g:~~ 00015 137704515 132.0 300 SV 4215 01-~O-b2 888t9 ·1~Jt8~~I~ 1 ~I:g~gg A~'~ ~~i~ 8t:~8:~1 00g18 1377(4516 132.0 300 ISY F 4215 81-Z0-e2 f 00 19 00020 00J21 ~00022 00023 00024 00J25 0026 0027 g 131704519 13770452~ 137704521 13710~522 137704523 1377C4524 1377C4525 137704526 137704521 132.1) 132.0 132.0 132.0 132.0 132.0 132.0 132.0 132.0 300 300 300 300 300 300 300 300 300 SV F 4215 1-20-62 ASY F 4215 JI-2C-e2 ~Sf ASY ASY SY AS! ~Y A~Y M ~ M M M M M 41ez 4102 4102 4102 4102 41a2 ~102 01-20-62 01-2C-B2 01-20-b2 0l->U-E, 0 -2u-a, 01-20-82 01-20-E2 883~~ 137104530 l~fjg~5~S i1~:g ~38 ~~~ ~ ~18~~1:28:~~ ·00030 132.0 300 ASY F 4102 01-20-E2 REPLACES 00031 137704531 132.0 00032 1?77045;2 132.0 00033 137704533 132.0 00034 131104534 137.0 00035 137704535 137.0 00J36 137704536 137.0 00037 - 131704537·137.? OC~3~ - 1377C4538 137.0 00039 137704539 137.0 00040 137704540 132.0 00041 137704541 132.0 00042 137704542 132.0 00043 137704543 132.0 0044 137704544 13Z.V 0045 131704545 132.0 00046 137704546 132.1) OuJ47 1377&4S47 132.0 00J48 137704548 132.3 oe049 137704549 132.0 00050 137704550 132.0 8 3eo 300 3)0 300 300 300 300 300 300 300 30G 30) 300 300 30) 300 300 30a 30J 300 ASY Sf ASY 6S' AS' AS' ASY AS' ASY ~SV ASY ASY ~SY Sf AS' I.SY $Y SY SY SY F F F M M M M F F F ~ M M M M M F F F F 4102 4102 410Z 4102 41C2 41~2 41D2 4102 410Z 36Jl 3601 3601 3601 360~ 360. 3601 2b02 28J2 2~J2 2802 777-q315~ 01-20-t2 01-20-t2 01-20-t2 01-20-e2 01-20-e2 01-20~t2 01-2u-E2 Ol-2G-BZ 01-20-02 ~1-20-fZ 01-20-e2 01-20-62 01-20-62 01-20·02 01-20-22 01-20-b2 01-20-62 01-20-62 01-2~-b2 01-20-B2 .. Figure 2. Short-hand banding data file (top) and completed banding record (bottom) produced by computer program BANDIN (data files reduced in scale). �91 PE~~1T NO PERMl TT EE lUSTER IIASTER I I S C H E --- B A N 0 I ~ G 06529 COLui1.ADO OII/ISION OF WiLDLIFE SANDING A WINDSQR, 6~LD cn •• CO RE eRVOIR • • ASHINGT8N CO.,CO 8 PREwITT LAR IMER CO. , CO C FORT COLLINS, r l • BAND NUM8~R Col1Mu:~ NA:1c 1377<----04501 IIAl~ARD C2 03 010 05 Ob 1077-56295 07 HPLACES IIALhAl<D 08 l~ 19 20 .. .... .... 00 H ~~ 26 27 2B • ---------43 -----_. .~.;-. _- - ~ .. Sy ASY SY AH ASY .... .... .... AH SY A?'y .... .. AMHICAN IIIG<.ON IIALhARD ., .. R ~g -- --- ------.-- --- BOULDER HGIJN CO., CO LAT-LJNG LoC .. ..A .... 317 IoiJ2-1045 .. .... co •• .. .... .... .. .... 8A TE 110- AY-YR 01-2~-82 .... .... •• F ~ - .. BAND NOS INCLUSIVE 1377-04 ~D 1 04550 THRU 1002:1032 .. •• ".. ..~ .... .... .... .... .. •• F .. 137.0 I.p .." .... 13~ .0 .... .... II .. F . 04550 -R8MARK~ I 7 RE LA~ES 00 REPL A ~~ II SY 602· ""105 106 47 - SEX ..SY .... ASY .... .... SY ASY .. REPLACES 777-93756 lIt.lhARD 32 33 34 35 36 37 38 39 40 41 -- Ap IV .... 23 B 31 AGE 300 .... .. .... .. 00 it 15 H Id r " SUTUS 00 12 t 13200 LOCATIONS 0 "OULDER, E F .. C9 ~ AoJ U L E i) .... .. .. ~ " .. SY ASY .. - .... .." ·SY F .... .. .... .... 1003:1050 ~ .... 0 101)0-1051 .... 1077-56295 777_Q'l7"" Figure 3. Computer-produced banding schedule of data in Figure 2. Schedule produced by program BSGEN (schedule reduced in scale). �92 ~~U.ARO , ., ~INTER BAt-lCING SUMMARY NuMBER BANDED BY AGE AND ~~~~----~~-~-------------------AM AF 1M IF ,. ~--=-~ ----~------~~~-~-------~ RfSo 81 eo ~ LOCATION - .. :SEX TOTAL " VAlf"lONl 16 63 () 9 28 !~3 157 PROPST SlCUGH TA~t.RACK ldESY 300 78 16 112 500 53 SC ~AEFER DITCH 131 132 143 8a 500 SEGEll<E SLOUGH 131 123 92 95 It41 50 5C 5(; 50 200 13C 133 106 514 124 198 93 565 a .l. 26 j,O 31 92 9~ 55 18 319 125 123 77 a 5 13 23 It1 32 26 19 33 .1.12 1 ----- 4 CHESTNUT SLOUGH KOCAI< PONDS BONt-.Y RES., rURKoS ?OND vERHJEFF LAS 1~5 !~y RESo AN I ~AS tJATCI-- •• CLAY PON!) SLOUG~ MAl(lI.£ ---~--~~-----~----TOTAL BAND:D 1.102 CANADA LOCATION GeOSE WINTER B~NDING N~MBER BANDED 1M Ml TURK~S POND TOTAL BANDED Figure 4. j.Y5.2 975 j,,46 930 1t11 ItO~" I • , f t • SUMMARY BY AGE AND SEX AF , IF . TOTAL 4 5 Printout produced by banding summary program. 20 20 �93 , Figure 5. Recaptures requiring banding histories (top) and recent archive file to be searched for matching banding records (bottom). �94 ------------------------------------------------- L;~,~'A~~" NO.":. SUTUS ;~. 133794702 ~F.TURN 133794719 fETUKN 337947b9 FcTUFN .~. 19788670 ~iCAPTUR~ 1197B88~3 ~Ll.PTURe NO 'ANDING ~lST(RY WAS FOUND i / ,""'~. ,'.~ •. :.,~ ..J.':'::'~:::"':: _:..•..., ~ . ~~::." :...~. ;'i)T;~"T'U8 FI~t>~AI\Di~GS ". , ..•., .' l:;.·· ...:\;?:!:.:....:~:·~·f.i~~~:{1·.:··: .. ·:r:~:.'··;~·~~,:·..·S!2\·' }.A.T-l[;~G.~. LtC Ar~p". .. ~-::".,.,;:.:r. r: <..:' <.,.-: .••. ~, .. ., .._u: •.•••.. " ~~~~~;;~~~;A~c~~5;cg~{;:·--~~ :.._ ce., 3H-1060 393-1\)20 C'lTA. DelT' 3t4-1080 393-1020 '. BON,.y Rf.S •• i)ON,.y RFS.. 393-1020 FOR THE FOllCwlN~ !'.,: ':~!k:;'~'i';' ;~.~.~~ ..::~:.:: .•., .." ",'" :~~..' 1Z9787b91 987b5432 '0,1 ,:::: CO· .IIIII .OF Of HAU.i:CD'?l'.,. HAll:.I_CO ~IROSI :J.~:~;» ··>."t:,:r;,~.:;~:t.~; .c.~:.;;~~·Jl_l~t~~$~l.~~1it .-. ;{ftt~'~i~~;~';'~"~~" ;'~?>'~: 7 IiE~·APTu~is:\'Is·TtD'·'·:'\~~;2'1i! ,~.::~~~:: > :::.:t.",r;·; ·:<···~!)f5t'~,:", :~~;:.;:~:;·:~ •.:;...~.~~;·~':'?,:_t_~J~,~Xt-,:.~ , :~·~;.·:'{:;:!(~;:"1.;': -: ~~ . .? .. • Figure 6. Output of search program depicting of birds used in the Figure 5 data set. banding histories �95 Table 1. Contents of banding Fi 1e # Tape 1. 1 2 3 4 5 6 7 8 9 10 11 12 13 Tape 2. 1 2 Tape 3. 2 3 4 5 6 7 8 tub tapes 1, 2, and 3. # Records File contents Mallard bandings, 1967-79. 1967 1968 1969 1970 1971 1972 1973 1974 1975 1976 1977 1973 1979 11,656 11 ,417 10,030 8,751 9,377 9,984 9,753 10,815 12,622 11,653 10,921 12,815 10,956 1980 1981 (part) 10,945 5,473 Combined blue-winged teal and cinnamon teal bandings, 1967-81 Green-winged teal bandings, 1967-81 Gadwall bandings, 1967-81 Pintail bandings, 1967-81 Small Canada goose bandings, 1967-81 large Canage goose bandings, 1967-81 Redhead bandings, 1967-81 Wigeon bandings, 1967-81 11,115 19,853 5,393 40,350 9,349 13,627 2,253 1,337 Mallard Mallard Mallard Mallard Mallard Mallard Mallard Mallard Mallard Mallard Mallard Mallard t1allard Mal lard handings. bandings, bandings, bandings, bandings, bandings, bandings, band lnqs , band ngs, band ngs, band ngs, band ngs, band ngs, band ngs, 1980-81. Mal lard bandings, Mallard bandings, Other duck and goose bandings. �96 Table 2. Column(s) 1 10 - 9 18 19 - 23 24 - 27 28 - 31 32 - 33 34 35 - 37 38 - 40 41 - 44 45 46 - 51 Card-image format of banding tub file records. Data Band number Replaced band number Permit number A.O.U. number Status Age Sex Banding region Banding latitude Banding longitude Direct ion Date (mo-da-yr) �97 3. Table Contents of :-ecovery tapes Fi le # Tape 4. Recovery Mallard recoveries, 1 3 4 5 6 7 8 9 10 11 12 13 14 Tape 2 3 4 5 6 7 8 9 5. Other duck and goose # Recoveric5 year(s) 1963-81. 1963-67 1967-69 1969-70 1970-71 1971-72 1972-73 1973-74 1974-75 1975-76 1976-77 1977-78 1978-79 1979-80 1980-81 2 4 and 5. 3,224 3,041 1,657 1,769 1,523 1,68? 1,488 1,146 1,541 1,356 1,457 1,547 1,398 1,166 ( in pa rt) recoveries. Blue-winged teal recoveries, 1963-1981 Green-winged teal recoveries, 1963-1981 Gadwa I I recover ies, 1963-1981 Pintail recoveries, 1963-1981 Small Canada Goose recoveries, 1963-1981 Large Canada Goose recoveries, 1963-1981 Blue-winged and cinnamon teal recoveries, Redhead recoveries, 1963-1981 Wigeon recoveries, 1963-1981 1963-1981 304 896 484 3,037 r , 503 3,409 332 275 162 �98 Table 4. Column(s) 9 1 10 11 - 17 19 21 23 25 16 - 18 - 20 - 22 - 24 - 29 30 31 33 41 46 - 32 - 40 - 45 - 49 50 51 - 53 54 - 55 56 57 - 67 68 - 72 74 75 76 - 79 80 - 81 Table 5. Column(s) 1 10 11 - t 9 6 17 - 18 19 - 25 26 - 29 30 31 - 32 33 - 35 36 - 39 40 Card-image format of waterfowl recovery data. Data Band number Replaced band code Date recovered (mo-da-yr) How obtained code Who reported code Present condition code Why reported code Batch number Flyway code State code Where recovered (lat.-long.-dir.) Permit number A.O.U. number Status code Additional status data Age Sex , Where banded (flyway-state-lat.-long.-dir.) Date banded (mo-da-yr) Band type code Date processed Hunting seasons survived Card-image format of waterfowl return records. Data Band number Recapture code Date recaptured How obtained code Blank field - for future use Wetland identifier number Flyway State Latitude Longitude Blank column �99 Table 6. t t f , Parameters checked by the banding data error check program. Parameter Logic Band number Num~er N( ) must equal N(a_l)+l or if N(a) = 99 (end of string) t~en N(a+l) = O. AOU number Number must equal an AOU number of a species likely to be captured (132, 135,137, 139, 140, 140.1, 141, 143, 146, 172, 172.9). Status Status must equal a likely status code (300, 370, 639, 647, 689). Age Age must equal an age class likely to be banded (AHY, HY, L, U). Sex Sex must equal a sex likely to be banded (M, F, U). Trap site First 2 digits of code must be less than 61 but greater than 16. Month Month must equal a month in which banding programs are conducted (1,2,6,7,8,9). Day Day must be less than 32. Year Year must equal designated year of banding. ��101 APPENDICES COMPUTER PROGRAMS DEVELOPED FOR STORING RECOVERY, • FOR THE COMPUTERIZED AND RETRIEVAL AND RECAPTURE OF BANDING, INFORMATION SYSTEM �102 100100 00110 DOI?O 00130 00140 00150 Ml60 0(1170 ' 00 80 00190 00200 O(l?lO 00220 00230 00240 00250 00260 00270 00290 007.90 00300,:':, , I I oo n o 00320 C 00330 C 88~~g C OO::lbO 00 laC. J • 1.110e Qf6D(1,10,~~u~300l DUP,~EWAGE,NEwSlX,PREFIX,NU~,j(U,~TAT~S, + ~rE,s.x,SIlt,~(.CA,y~,REP 00370 ,10 cORI'CAT( I.2.A 3,A1. 17 biZ. f 5 .1.1lI.13. lX,A 3, IlI.H, lX.lIt,. lX,3I 00330 + 41<;) 00390 00400 C --- CHfCK I. ~fW LINE [R ClFLICATE Cf ~RfVlnus LINF 00410 C IF IriJP .'EQo 'NL,) GC TO HL:-':, ',: 00420,::";:, PRFF;x 0 OlCP~E 00430 0044() NU~ • "lt~UN • 1 er u • JLDJ,Jt.: 00450 '$HTU~ 0 OlDSTA ~ ,. C ,', A(.E a ,1LOAG' s~x ~ 'llD~tx OOVO )pr u OlDSH 004~0 ;<10 c I1l0l'O 00500 r.~ " 'ill"rA 00')'0 OO<;?O YR • nLDYR 1< ~ P " , 00530 ';:,":"; 00540 C O(l5~() C ~nrIFY 'G~ !ND/DR ~EX 'CL'SS IF Nltl~~APY' OO}bO C 00570 IF (~UP .EC. I A') AlE • hl~'GE OOStlO IF (OUP .ECo ' 5') SiX a Nt~SfX 00590 IF IOUP .EQ. 'A~'l T~~N AGE c "IEoHE 00600 $ ~ ~ = ~H IiS!- X 00~10 HID IF 00620 00630 C P~INT lI~E 1F ~Aw (~lPLl DATA --00640 C 00650 C 00660 00670 006RO 00690 C SAVF OLD VALUES FOR USE IN DUPLICA1ING NEXT LI~E_--00700 C 00710 C rU,PRr 0 'RFFIX 00720 (jlCtiV~ • NUM 00730 01. Q}"U " },QU 00740 Ol [ISlA· ,TATuS 00750 CL(lAH HI:' 00760 r'.DSCY ,00770 007'10' CLOSH • SITE 00790 CLOl"O a "'3 "L~rlA • DA. OOROO ['LOn = y~ OO~lO 100 CJtn!NL'~ OOE20 00"""0 300 JU"IK = 1 00Q40 STOP PlO 00P58 006t'> IfOR 00870 I~FAD,COR? 00880 IH'F 2 ,IX. ':., gg~~g ':r:':' :~'-";,,: '. ,q Program to Decode Shorthand Banding Data Format '!,-., to Full Data Set. �103 i , « .. 00100 ".::''t,-.: 88H8 I 00130 . ~:..' . 001~0 OOl~O 00160 00170 00180 00100 00200 00210 00220 , 00230 , 00240 00250 002bO _ '" ,', 00270 007~ 0 002'10 .. ~::~;. 00300 00310 C e~NCING D~TA FILfS FOR r~~c~s ••• 003?0 C $*. DR~GRA~ FO~ C~ECKINE 6.* ~R'TTE~ ~V Ji~ klNGllM~N, rcw ••• 003'10 C 00340 C IIRITt14,51 ,"'':'~' ""';} 00350 00360 5 FOF~br," --- l~ROR OlA~[STICS FOR BANDING DATA FIle ---"III 00370 rpIOO',I a,1.110C, , ,V' " t: "-":4:', ,<h':' 00360, C ',., RE~D(1.lC,ENC·~99) LINl,F~EFl),NU~,.rU,ST.TUS,~(E,Sl),LC(1,LCCZ, 00390 ~ MD,DA,VR 004(10 10 F1R~ATIT~,2X,I7,I2,H.f~.1,H,13,1).,1.3,1X,A1,1~,212,lX, 00410 OO~20' '.'<1-', ..;"" r2.1X.I2.1~PI2) '" ,,:i'e .. ", .., 00430 C 9** BEGIN llGICAL TEST QLESTICNS 00440 C I 004~0 ~o. C 0041>0 ',~,~',:,,',: IF r r •E o , OD470 IF IClDNL" 11 004~0 0041)0 00500 oNE ••• IF 00570 005"0: 00500 00600 00610 00620 006,0 006~O 00(51) 00660 .~c. + (",,', .~,. ,HMO oNEe I~ (DA IF IYF C 00700 00710 oorzn 00730 00740 14 16 00750 17 00760 00770 007RO ,~ lq 20 007'10 OMOO 22 '.3 e .~t. .~i. llNIU".]1 :.'.. llN~ \i~!TE{4,131 LINE: 7) 0)1 .CT. .N~. .~NDn (n .~E. !:) liRlIU4,2CI 311 .'''iJ. 'MD .NE. LINt '" ,', LINE r. oNE. CI.RIll(4,231 9) .~I\(i. " ,',,,:',;,; ~RITI(4,22) e3 .A~C. lINF 11 FOP"!T( 12 F8PMA'( 13 00"10 C " .~E. CI W~TT~(4,1~1 .~C. (IA~U .NE. 132.0) .t~C. {leL 135.01 .A~r. 137.01 .~~C. (AGt: 139.01 .UiO. OOt; .IiE. o.er :;';:." :>i' (AOU .Nf. 140.CI .~NC. (~CL .~~. 14C.1) "~l. 141.C~ .,he. (AOU .NE. 143.0) .AND. 'Jel .~E. 146.C) ItOU .Nt. 172.C) .~~L. (tCL .~~. 172.9) ~k.T~(4,141 Ll~E: IF ({nATU~ .N~. CI .t.H. (SHHS .NF. 300) .AI'C., "./:':(:',,'1';;, +(ST!TUS .NF. 370) .~~~. (STAllS .Nt. 630) .ANC. (~lA'U! .NE. ~ 6~7) oHID. (~lAlv~ .Nto tB~I) w~ III '4,16) lI~f ye ((Ar,r .~E. ',l.HY·) .~NC. ("n .!.t.' HY" .Ar!'. + (AGE .NE. 'ASY') .A~I). (AGE .NE. • SY'I .A"'D. !AGE .Hf., }:,~':"<';:~',,:,y~,,. + I l'l oAt-:O. (AGt .N~. f U') .A"L. !!~f .NF.' 'II + \J~ITt(4.17111N;: IF «(SC:x .1'':. 'M'l .AN[.. (St) .Nt. IF!) .AND. (SE~ .liE. f I) ~:;., ~ oANO. (SEX .Nt. ,uo I I wRlT~ (4,1(' I LINE ,'''-:','"..;:'' 1F 'IL[lCl .GT. 60) .cs , u oc z .GT. 16)) Io'RIH(4,HI lli<E TF (I~O .NF. 1) .ANe. (~G .~•• 2) .~ND. (M( .~F. t) .AliC. 005~0 00%0 00820 NIJM .I<"t. +!AOU '.N~. '.AND. +(ACU +.ANP. 00540" OM10 ~ 9D IF ':" 00510 00520 00530 OOh70 006QO '0061)0 {('U)"U'" (;0 TO 90 o~~. 9~ .,NC. NLM u,o:; C "LINE ",15.~z "lI~E ·,15,·' ",!~" "l f 0 Q ~ ,\ T ( "lIN E ••, 1 5, " I FO~~tT( "LTNE ",15."1 F 0 ~,., t T ( "l t ~ E ",! 59" I Fnp~tT( "L'NE ~,I5,~: FOQ~~T( "LI~E ",15,"1 FO~~AT( "lINE ".15.": f",!P~'T( "Ll~E ",15,01: FOR~AT( "lINf ",i5.": FDRfoIATf ttl IN£. INITIAlIZ" OLD VAL~ES fER SUBSEQUENT cr~p.PtSo~~ ••• -~ .. 00~40 0llo50 OO~60 00"70 OO~'O C r: gg:~g C.lna 00010 OO~?O 00Q10 00Q40 00Q50 Program CrNTINUE w~ITr(4,301 COL~T 30 Cnp~AT(II!4,1X,"dANDIN( <TOP END ~~lRltS for Checking Data for Errors. 9CQ WtRL CHECKED") IE'OF Banding �ggi~~ USER, AOcS, P•.30. ROUTE,CUTfLT.IL·40. 88H8 (lOliot GETITAPEloe~ND$3) 0015~ GETITAPE2 iRPLO() FiNS.lOnO. i ggn~ lGC. OOlSC GAYFILbDFl I JGB Or ~.AA.OEF. o o 00190 REPlAC •• Ofl. 200 ExIT • o08 210 DAYF lL b rF L. REPLACbDHo HO~ PROGRAM SCHEOIINpuT.OUTPUT.TAPE1~TAPF.2.TAPE4.~UTPUT) 00250 C PRCGRAM FOR GENiRATING A BANDING SCH~OULf FROM PAW BANDING DATA 00Z6C C INP~TTEO FROM DATA FlLE "BANDS" --- wkITTEN BY JIM PINGELMAN. (OLO~AD1 DIVISION OF WILDLIFE IMPLICIT I 'Ti:GE~ IA~ZI· 0031 CH.RACTER.S AOU CH~~ACTE~.37 PlAC~.lOC (HARACTER.19 NA~E.~EPl 0340 CHAhACT[~.a LATlCN,l.LC,OATE 035C CHARACTER$3 AGE,REGION.STATUS 003bO CHtPACTER.l SEX,CC CIMENSlON PRE1I~O).PRE215Q),NUMI50),AOUI5~).STATUSI50).AGEI50)._ + SEXI50).SITtI5C).DATEt5Q),REPLI~O),N.~EISOI.LATL(NI~Q).((1501, + PlACEl61 c lOTCL ••.O. 88H8 OOHO a._ 88~~f E 8g~~8°~ 88H2 8 8g~~8 88~ZB 88H8 88 ~~O c 00lt5!) C H(I N 1.20 H!~l~I~IAL ULLE5 FOR VARIA8LI:S Its REGICN" '311' It9 -... DO 510. t:tlcl.t 500.__. PLACE IN,.,'c' ... si~-·; oCgNTINUE g~0!~80.C o DC 500. 0 0 __ 8 S8iiS 8S~~g~. 88H8 c C SET LINE PRINTER WRITE(4.371 ,. FOR t LINES/INCH 37 FORt:ATI"S") READ iN DATA IN BLOCKS (iF 50: llNt:S~·. 00 ~2C, ! c 1,5C READ(1,10pEhDo~~9) PRE1(1),PRE2(I).NU~III.AOU(II.STATUSIII.AGEIII +,SEXIII,$ITElll.DAJEII).REPLII) ... __ .~O FORMAT(7X,1,.13.12,11.A5.1X,A3.1X.A3.1X.Al.lX.I4,lX,A8,1X,AI91_ TOTcY ~ TOleT.l 00630 IF IAOUIXI oNc. 0 0.00 oAHO. FLAG .NE. II T~EN FHST I<UMIlL _ Ctb!5 _0(;66(1 flAG" 1 LL c ! WD IF 0068 If IAGEIlI .NE. '.. -') THEN .. . _ LAST· I'\UMll'--_ 0070 .. K~ ~ 1 00710 lENt IF oone 0073e C (HECK TO SEE If TRAP SITE SAME AS PR~V!OVS ENTRYl IF NOT. CAll OOHC c SUBROUTINt TO CERIVe LAT=lONG ANG D"SCRIPTION INFORMATION --00750 C 007bO IF SHEIHkST_) .01< •. ISITEII) .NE~ SITFII-l~)~ THE~_ . rEWIND 2 ... L2~to~'~il!Et~tG5IT.LAlO.LOCI CO 15, 11'\ 1,!-1 IF I~UMIII .EO. FIRST) GO TO 1& IF (LAlG .E~. LAllONII~11 GO TO ~5· 15 CONTINUE HSI:. 0085C .LAnOWlll. o.lATlONll=ll.·_. GO TO 35 00880 END If 16MoM+1 00890 PLACE I!") = lUC 35 JUr.~ 1 MAll ARD IF 1A0UI I) .EQ. '132.0·) NAMEIII , NAM~ 111 GADliAll '13500') 4'1FOIt A"I II IGEON '137.0') NAII~II m' TEAL IF IACUI II "EO' '139.0') NAMElll •• • G~~F~-WINGfD '140.01) HAMEl I 0' IF lAoue II 0_0, 8lUE-WING£D TEAL !F I~Ou( I) ,~Q. 'lltO.l' ) "'AME IXI 00910 ' U"IID~"ITIFlfD T~AL 0141.0' ) NAME 11) CINNAMI'1N TEAL IF I"OUI I) .~Qo CCJ6Q 099(, PINTAIL IF l~lJUI!l ocJ. f 1" 3 o· ) tiAMEIII m IF (AOUI!) oEQo '14b.0· ) HAME II I REDHF.lD 01000 , CANADA GOOSE H (ADUII) .EO. '172.0'1 NAME lIla IF I"Gut I) .jOe. '172.9'1 NAI1[II), a 'SMALL CANAD~ GOOSE 01030 C 10lte 520 CONTINue 00588 88~Z~ 88g~8 OOb"8 COb7C OOb9g i~Hlg&nt Q g~~iF! 0 88U8 &gg~~. .8~~~8 ~8U88g~~~. 0 H gBbl U :£g: g 8ig~g 5( C 8100b 1 0 C 0 818818° 09 8ilu~ Ollle O 1 ,0 01130 Ol14C 115C 1160 01170 C 8 8HS8 01l0C) Program 0 • m , 0 , ,. .--, 0 0 PRINT TABLE HEACrNG - 3t ~RITlI4,21) ~RlT~I~.30) ~PITEI~,4u) PKEl(Ll),PRE2Ill),FI'ST wRITE(4,5U) PRE2(KK),lAST ~RIIEI4.6°1 ~wl E 4.70 PlACEIl),PtACEI4) ~~lfi:(4,80) PlA(~121.PlACE(5) ~FITE(4.QO) PLAC~(3).PlACE(6) .RITEI4.100) ~RITl(4.110) PRE1ILL). 20 FC~r.ATI"1",T22."· - B ,. N 0 I N G S c H E 0 u i. E --") 3? FOkl1ATI " MtSTE~ ~[KMIl NOI 'b52~"6Tb~'''INClUSIVE BAND NOS") 40 FO~MATI Q MASTEl PERMITTEE I COLORA 0 OIVISION OF WIlDLIFE".T67, to Generate a Banding Schedule. �105 8 1210 1220 14,"-", Ij o3~12(2) + 50 FGR~ATI Tbb,"TH"U",2X,I3.3,I2.2) 01Z30 bO Fu.roATI T34,"6A~OING LOCATIONS"I 012'0 7~ FOR~ATI· A",lX,A37,T42,·0",lX,A31) 01250 eO" f(j~~ATI" 9",D.~37,r42."E",lX,A311 GlZbO. _ 90 FOR~ATI· C·,IX,,37.T42,"F·,IX,A371 °112287'0 1~u fORrATI"OB~~O Nl~~ER",4X,"COMMON h~MF",5X~"t~U .",lX,·STATUS·,lX, + "AGE·.l ••·StXR.lX,"REGIOh".lX,·LlT-L~~G·,1X, LOC·,3X. °01Z9t ~ ~DATE~I 1300 110.fO~~ATIIX,14,"<·--~",T73."MO-DAY-YR"1 1310 C I 01320 COUNT 1 1 ~A! 1.50 ~1358... IF IA<;EIJI .EQ.' 'I THEN 36 ' Iil<lTEI4,SINUMIJI.REPLIJI 31 _ J. tDRMArI9.,12.211 ••Al~1 ····£NgcI~o 530 g I ~H2~·-.~~ 55 Ho; 8i 8HS8-··-- D 01400 C 0141(; C ~o~ 0142C C 01"3C 01440 01450._ .. _ 014bO._. _ l't7(; 1'080 149C __.... _ 150IL. __ . 01510 8 g SiHt . IF BEGIN COI1PA~ISO~S FO~ LUPliCATE DATAl DUPLICATE, ENTER DITTOI If NOT u~PLICAT". ASS~ME CHARACTER V LU~ BY DfFAULT --IF (~UMIJI .E~. FIRSTI GO TO 205 IF (N,A~HJI .EO. I,AI'I[lJ-1)1 "A"E(JI • , • l.F IAOUIJI .1cQ. AQUIJ III AOU(JI m , .• • -. ... . .. _- .• -..•• IF ISTATUSIJ .EQ. STATl-S (J.l) STATlISIJI.tl...t. W. L ._ -.-_ ....IF IAGkIJ) .EQ. AbEIJD111 AGclJI • , " , IF ISE.IJ' .EQ. SEXIJal SEXIJI. ,., IF .!J .GT. 11 k£G1CN IF .. IlATlONIJI .EQ. LATlONIJ"'UI THEN - .... - .... -.LATLONIJI •• • , EL~~IJ.~ a .• .,o__ D , ".' . 8g~8-·--·-·' IF ~rAttg~df :LA:JliTLONiIII) 015bO CCIJI ~ CCI1I1 01570 - .. 10 180-. 01SBCi _ ._.._ E,,(j f . _ . . THEN - _ .. GO 0159C 1bO CLNTINUE 01bOO CO~NT COUNT + 1 _Olb1CL_._. IF ICCIUNT cEQ. 2) CCIJ): '8' __ .. 01b20 ... If "CUhT cEQ. 3) CClJ I 'C' .. ---. 01b30 IF ICOUNT o~Qc 4) CCIJI '0' 01b~a IF (COUNT .EG. 51 CCIJI m 'E' 01b50. IF ((OU,.,1oHIo t) CClJ I a '.F' Glbbel C 180 lND If. - .... .. - .-- .-. IF (DATEIJI .EQ. DATEIJ~111 DATEIJI • t w t 01090 __.. IF IREPLIJI .N~o ' ') HAMFIJI • REPL(JL-.--01100_.__205 IF IOiUlIIJI .I::Q.FIRST) .OR. ItWIIIJ) .EG. LAST)) THEN ..-.--.~~1~14'1~CI 017jC .. __ LA jAOU ~J I.S T ~ T~S.I.~I.' AGE IJ I, SEX ~ J!.~RE~I~H'_ ... 01740.__ GO TO 210 ..._ .._ -- .... --..- .... -..... -.-- ...-.--._ ... -- .. ---. --.----01750 l~O If 017bO .RI1EI4.2~OI HU"IJI,NAIIEIJI,AOUIJI,STlTUSfJI,lGEIJ),SeX(J),REGION, 0117.0__ .. +. _. LATLONIJI,CC(J).OA1EIJI .. .. .. _. _ 0178 (I 20u FDR I'A T19 X. 12.2, IX,A19. lX,A5. lX. Al. 3X,A 3, 2X, Al, 3 X. A3, 3X. AB, 2X•. 01790 + Al,2X,A6l ··C·~1~Q~<~~~AT:~( ~~; H;.~i. ~X~A~~..~.~l~'_l~.A_5.2~'~.~:~:~ X'~.3 :.2)(, A_~~3 X, ~.~~~~'-=:-_ .. :~~ 01830 C CHA"Gl DITTOS Of SUBSCRIPTI::OVARIABLES BACK TO CRIGIHAL VALUES --210 '1 AO~(JI •• AOU;~ ••~~~EIJI • ~Ar1~(~-l1._. __ ._._ .. '«)1Bbu_._ If ISTATUSIJI oEQ •. ·-" 'I STATUSIJ I • STATUSIJ-l) _.. - ..._ .---.--.__ 01B7C IF (AG~(JI .EQ. ' ~ 'I AGilJI • AGEIJ-l1 01880 IF ISfXIJI .EQ ••••• ) SEXIJI • ScXIJ-lI 018'10-IF (LATllJl'dJI LQ. _'.,!' ') LATlONIJI a LATLONIJ-l) .._.. _. __._..__. 01900 __ ._ lLIDiHIJI .EO. ' ._ 'I OATUJ) • OATEfJ-lI .._.- --.- --- .. ._ 019l() IF ICCIJI .!:Q. '''') CCIJ) • (C(J-l) 01920 530 CC~TI~UE .01930 C ENTEk REPlAkKS AT THE BCTTOI1 OF THE PAGE -_ ••.. OH~O_._.. loIRlTEf4.2901 ..-... -.... 01'150 290 FO~MATI~ R~NARKS") 019bO SPACE = 0 .01'170._ .. _.. tCl 5"0, K • l,5e «:198(;____ IF IREPL(KI .NE. ' 'I THEN 01990 ~PACE = ~PAC~ + 1 02000 IF ISPACE 1) WRITEI4.300) "U~fK"REPLlKI 02010._ .IF ISPACe 21 wRlTEI4,301l "UNI K I,REPLI K I 02020 __ . Ii' ISPACE .EC e . 31 IIRITEIIt.3021 NIJr1IKI.REPUkl 02030 IF ISPACE 41 wRITE14.3031 hU"I(i( I,~EPLIk I 284~ IF ISPACE .EC. 51 W~ITEI4.3041 NllM(K)'"fPUI(, 2 5 iF ISPAC~ .EC. 01 ~RIT!:14,3u51 NUMIK"~EPlIK I 02 b· IF ISPACE .Glo 61 bRITEI4.30bl NUl'lKI ,REPlI K I 02070 3C~ FCR~AT(·+·.T~,".".X2.2.1X.A191 02080 301 FOR~ATI".".T34,"U",I2.2,IX.A191 02090 302 FOkI1AT("+·,T5q,·~",12.2.1x,AI9) 0210~ 3{3 FORMATI" ·,T~,·.·.I2.2.1X.A191 0211' 304 FOkMATI"+",T34."#",I2.2,lX,AI91 02120 305 fGR~AT("+",T~9,"#",12.~,lX.A19111 213C 3Ub FGRHATIIX,",·,12.2,IX,AI'1l . Zl'tt. END If .. 0215C 540 CCNTINU~ 021bO C 0217C 500 CONTINUE 0218C 0219C C '~9' 9 ~~ilil~,2201 TOTtT' 02200 20 FOR~AT("lA,I~,A BANDING RECORDS ~ERE PROCESSED FOR SCHEDLlES·I STCP 02210 ~NO 02220 SUB~D~TINE TRPSITE (NSITE,LALO.DES) CHARACTER*37 OES.D C~A~ACTcR¢6 lALl,lOCA OZZ~O 00 100. 1 • l.b4 022bO ~cA012,lOI LlC.LDCA,O 02270 fORM~TlbX'I4.1KfAe.1XtA371 IF (lCC ocQ. ~s TEl GIJ TO 200 02300 . 100 CONTINUE LALC • 'CvC=OOOC' C23l1. O~S c •••• tRAP SITE UNLISTED •••• 02320 GO TO 300 ao LALIJ = LOCA DUaD (.1235 Ii 300 ~ETLRN 023bO E.ND {EOF 0 D 0 81~~~ 8iJ1g ! Hb~U ~~~U ~s~~~Ht ~ 8U~~ O~_ 2U;8 __ H l~B~~~1).E~:·_.''' .D __ .,Q. .H. .E~. 8 8 8~~~8 gH~8 gB~~ 8B~g Program to Generate a Banding Schedule (continued). �106 0,,010" c~ll 0 ('CPO ~.'" (\."" o-uvo 'l'I' , :'"'\("11...) ,':;" 1 7 ) O"l~O "rl~,) (\nZ')(l () ~l ('I" on:'l?"l ,)') "I 31 )'l?4 ) o C' - t: ~ Aw ~"r r ~ f ( i 1< I L T, (J L 1 P LT. 1 A F L 1 , 1 t I~~l !~JT !""'r en (.I.-Z) (:~'~t~Tr;~~ LL~'~LL 00")t;t') oo~!>') ~,i?7"" J:'?" r ,')~.,~""; r 1'10300 r 0"~t'") O"J',\ ot'" ( 1)~3" \ rd"~" ') ~ ~I 11. -: • : • \.: • 5 C r A~ WQ1TI~~ AN >A=>:TT:~,1 Jl~~ c '''n, nl 411 ..;y r:!." ~ !T;., i o gg~jg ~r u :- .• ~ '; ( ~ 0 I")~~AT( .• B.~CIN' r t:t T ~ I. R #01.:'; l. 1 I,d.; L~ 6 T6'~ l~ ~l!~L TC Th~ :l.l~[':' hIlh ft:fLACcL.: r~MPUT~~ 01'\ ~ ,,') flr4"J,) (Ii TJ QrfO~~.T ''i:LI'~::~ '. TAO c 4· r l Tf ~II G," 6(T;""1 ~QnrRl· r f ~ '),:) O'l~41 17 ~(l'i.J ~"H D' = I "A" VNDE~ ~'TA Fk~M C~LORtnn CO. F(k~,J I[ L. P\:-: t.1, t r- ~ t.: ~:- !.~ c: tNt. to Ai ..•.,'\ L ~, ul~~ t,~·DI~.~ l'R. r~.~ ~~~f~Ah iJ/..lll.,~ bAND "lr~('::l~t.r~, TYP~ CLLur..t.~u F~"""'! T~c l~ C~Lr~~ V";"; lr~'lt,u, M': \o.:"'Ll ~? 1~/(1/f3. 1,~OOO ' 1. ~, J ~J L e .4 J ~ f'4L , t. G U 1, t. w IJ c I ~ I /..T , .l L r , G;':"', 7X, ,'), iX, j_:;, j), 11,11<, 13,lx,;.",1 x,~ 1,lX, ~"J , ,_'f. A -:.., 5 : T = , l~ L, l..,. , 't !4, 311x,IZ I j(, J.. C L. r,44'l Tu ('1'"\4'\;) !:VI.; cu 0046U 0r.47~ C"4";) Aut TC 500 0 "li: :")'4"~ 005,1~ • ~ /..Gl £. :.g~ 3 (')('. e ~ ') 005'0 AC, i; I> AGt A('E 7 0(54) Qf) 0('· c:::. '\ r:,~. ~. Tr.: ",.:;. 71 : I' 0(050):) n'\~o') r. OI)~O() C YI~\) C -- L '1" K " " {,..~. Cl)~'(\ ('I0t., Go;) (\~,t..,;,:! ~ I t tTl It.;) .re. ,:: " • 5 0 X sb = _ t t ,,0 l [ " G 1 TL l) CLU~ITI l F k U, «< TL At< r r r i' ,n if .I ~ A T H .•" -- T~cN ., ol = n~t-c:~ ~ C (}) 7'l·J i f r 0"74" (. Jr, 1': ) -; "'\ (\"7":) 'Y: ) 0""00 on""'! () 01'0'1 ,,,,,,, ".:-\ f I) .: ,A, :.; ") 0)71') 0~710 ('0770 .,,)'1 T S':') I ~ I) ~ILl 'T·r~IT~(~IT~'Ltl,lL~EI •...• r "I -r ~ L i LU"'~ :~, nl.OLJN c ': i IT = !JI ')LA] t '-",:,": -= lvLL~i~ r: I.,J ~ - I;: i)0~~' Jr' ')7'"': 0;'171(;r',"''' o ~: :'; , II' (STTe 00"20 ~)"\ ..=0.~'.'u') ( GP' I r.c~ ( (7, ~-J IF 00 5~O ~" r: ""'l t.: ~ ~ T:: r (•• 1 :;.) I T f I "J • q , !:' ~' : ,~• :t t II • '" I ,t • G.i:'" •••..; ~' j 1 t- e ~ II ~ GL· , .i. 11 ~ Lv;'" f~, \;._. ,t ~ tt J •••• L t... '- 11 t: :..t. ~ It S 1 i~1 11 J \; c: , "'? Y , L t: T, l r ~(:, r L' ~ /-Ii, 1 K 1 ;" i 1 • 1 • I ~ , 0" • I 2 • 2, 11 II ,I 3 1 7 It , i 3 • 3,1 II It 'It''q-:- = "'!T~ J: ,- •• ': "'11:,: 1:":qqC; 5TOo -:"''"'1 ) 00'·0 :)(.:'t '\ 00 '''0 00~'" 0f\~':)' .,.j "::J,) !)r,Q,,) n"'ol r; :"~"'; ""0'" 0"(4) 0"')1: 0 !~TE~·~ [~,LJC,LAT1,LGN(1 f'; .I _:~, -', :: .•., J.v0 ==!rr7,~,~~'~;2L'J) ~~~,ltTi'L[NGl r.,~ •.. · ',T ( ~ 'I , ~"1I 1 i, .i 3, 1) 11 14 I rr IL1: .cC. ID) (,u TO 200 r'-p'T r·qJ:"'!) ?v ) l ~= I !:' t ~'=l (,"Ir! ,'=rUr'l .. , i) r>"':)4,') Program to Reformat Data to Conform to Bird Banding Lab Specifications. �107 , l t t Program to Summarize Banding Data by Species, Age/Sex Class, and Location. �108 Program to Summarize (cont inued). Banding Data by Species, Age/Sex Class, and Location �109 '·..·t -v-.=:-::..'~. otOo LA .A~C. LeNt • HAC E oCO. LDI THFN ..•...;.;, .... Program to Search Banding Archives to Determine History of Recaptures. ��Colorado Division of Wildl ife Wildl ife Research Report October 1984 111 JOB PROGRESS REPORT State of Project Colorado 45-01-506 - 15050 Migratory --------~--~~----~~------ Work Plan 10 Job Title: Period Covered: Bird Investigations Job: Cooperative Management Programs 01 April 1983 - 31 March 1984 Author: Gerald M. Lorentzson Personnel: John Corey, Howard Funk, Gerald Lorentzson, Jim Ringelman, Steve Steinert, Mike Szymczak, Colorado Division of Wildlife. ABSTRACT Work accomplished during tive management programs in the Central Flyway. Service consistent with this segment consisted with the four regions Help was also given to the objectives of this of assistance in cooperaof Colorado and the states the u.S. Fish and Wildl ife work plan. ��113 COOPERATIVE MANAGEMENT PROGRAMS Gerald M. Lorentzson P. N. OBJECTIVES The major objective of this job is to provide liaison with the various DOW sections in order to assist and work with them in migratory game bird matters, particularly those relating to management responsibil ities placed upon the various sections. SEGMENT OBJECTIVES The procedures for this job wil I be to provide liaison and expertise to the Regions and other DOW entities in assisting them with management activities for which they have been assigned main or partial responsibilities. This assistance will be given in the way of training and/or help in design of cooperative management programs such as monitor banding and various migratory game bird surveys. Assistance will be given in design of goose transplant programs and monitoring of results. Procedures for federal hunting season requirements and limited permit hunts will be explained and assistance given in setting up seasons and monitoring results. As necessary, assistance will be given in sampling schemes, hunter surveys and statistical analyses of results. Cooperative effort will be suppl ied in preparation of strategic plans for migratory game birds. Project personnel will take part in various technical committees, status and regulations meetings and management plan workshops both in-state and out of state as necessary for cooperative research and/or management-oriented purposes. METHODS AND MATERIALS Due to the diverse nature of this program, various procedures and material were used to accompl ish the objectives. The methods and materials used for trapping, banding, inventory studies, and the compilation of harvest data have been well documented in other segments of this program and will not be repeated in this report. RESULTS During this segment, assistance was given to the regions in the areas of breeding pair counts in the San Luis Valley and in North Park. Assistance was also given for the goose production surveys and the December and January inventories taken on waterfowl. Many contacts were handled regarding goose nesting structures and other day-to-day problems concerned with waterfowl. �114 Several training sessions and other informative meetings were held for regional personnel and other community-oriented organizations. Assistance was given to the Fish and Wildlife Service in collecting and speciating the duck wings for their waterfowl parts collection program. Help was given setting up material for the Wing Bee and for actual assistance during the Wing Bee. Cooperative efforts with other states and the Central and Pacific Flyway included administration of Colorado's dove call-count routes and submission of data, the Rocky Mountain Canada Goose Sub-committee, the Central Management Unit Technical Committee Meeting, the spring and summer meetings of the Central Flyway Waterfowl Technical Committee and Central Flyway Council meetings, and the Subcommittee on Nesting Studies in the Dakotas and eastern Montana as well as the Research Needs Subcommittee. Bands and banding materials were distributed to the regional personnel and banding records forwarded to the Bird Banding Laboratory. Band recoveries and recaptures were forwarded to the Bird Banding Laboratory. Assistance was given to the Regional personnel for picking up dead ducks which died of Aspiringillosis. Help was also given in the removal of nuisance geese from the Denver and Fort Collins area. (~ Prepared bY~ -._ _'--~A~1_-~~~~~~ __~ ~_'~~~~-~~(i~~_':~· M. Lorentzson Senior Wildlife Biologis �Colorado Division of Wildl ife Wildl ife Research Report October 1984 115 JOB PROGRESS State of Project REPORT Colorado 45-01-506 - 15050 ------~----~----~~------ Work Plan Job Title: 11 Job Bird Investigations ----- Migratory Period Covered: Migratory 1 April Game Bird Project Administration 1983 - 31 March 1984 Author: Howard D. Funk Personne 1: Janet E. Black and Howard D. Funk. ABSTRACT Program Plans for migratory game birds were completed as part of the Strategic Plan in the previous segment. However, the specific plan for ducks was highlighted and utilized with some others as part of the final approval of the Comprehensive Plan option for the Division of Wildlife. Planning for and approval of a new job (Work Plan 1, Job 16, Field Feeding Ecology of Mallard Ducks) was completed and gained and the study was initiated this segment. All other jobs continued as scheduled. ��117 MIGRATORY GAME BIRD PROJECT ADMINISTRATION Howard D. Funk P. N. OBJECTIVES To supervise and administer game birds in the project. research and management-related jobs on migratory SEGMENT OBJECTIVES Supervise and administer all research and management-related necessary on a day-to-day basis. jobs as METHODS AND MATERIALS Planning for new jobs and documentation of jobs already on line was guided by the Strategic Plan and the Program Plan of the Division of Wild1 ife for a 5-year period (1983-84 through 1987-88). The Program Plan for ducks was one utilized as a sample in the final approval for the Comprehensive Plan contract. RESULTS Program Plans for migratory game birds were completed in the previous segment. However, as part of the final approval for the Comprehensive Plan option for the DOW, sample programs were described as to methods and materials used in specific portions of the Program Plan of the Strategic Plan. The Program Plan for ducks was one of the examples. Descriptions of the processes utilized to obtain data pertinent to current status (included averages of number of hunters, number of birds harvested, average season bag, recreation days, numbers of birds present in Colorado in mid-winter), and projected objectives and/or trends for the above items for the near future. Narratives on these subjects were submitted to Denver for inclusion in early July, 1983 with information utilized in gaining final approval of the Comprehensive Plan option. Planning was completed and approval gained for initiation of Work Plan 1, Job 16, Field Feeding Ecology of Mallard Ducks, in late June and early July, 1983. The Program Narrative was submitted and work was initiated on this job in this segment (see progress report elsewhere in this set of reports) . �118 Work was continued on all other jobs as planned and as per progress final reports found elsewhere in this set of reports. Prepared by Howard D. Funk Wildlife Research Leader and � https://cpw.cvlcollections.org/files/original/e5e5f4d2575e6134d3d2de136ccae598.pdf 7f5061e35d73398d8066887cafe6409f PDF Text Text Colorado Division of Wildlife Wildlife Research Report July 1985 1 JOB PROGRESS REPORT State of Colorado Project No. 01-03-047 Mammals 1 Research Work Plan No. Multispecies Investigations ------------------ Job. No. 1 ---------------- 7 -------------------- Period Covered: Author: July 1,1984 Publishina Mammals Research Results - June 30,1985 L. H. Carpenter Personnel: R. B. Gill., L. Lovett, N. McEwen, and all Mammals Researchers ABSTRACT During the 1984-85 segment, the Mammals Research Sections had 17 manuscripts published, and 3 others accepted for publication. ��3 MAMHALS RESEARCH PUBLICATIONS Len H. Carpenter P. N. OBJECTIVE To publish results of research conducted under the auspices of federal aid projects 01-03-047-15085 and 01-03-048-15080 in a variety of professional journals and other indexed publishing media to insure widespread dissemination and availability of this information to natural resource managers and ecological scientists. SEGMENT OBJECTIVES Following is a list of publication working titles and Progress Reports that will be prepared and/or published under this job during this segment. The publication outlet is tentative depending upon acceptance by the particular periodical. 1. Job Progress Reports 2. Anderson, A. E., and O. C. Wallmo. Mammalian Species. 3. Baker, D. L., and D. R. Hansen. in mule deer and elk. 1984. Mule and black-tailed deer. 1984. Comparative digestion of grass, 4. Baker, D. L., N. T. Hobbs, and W. Vanhoven. 1985. In vitro rates of digestion of native forages. J. Anim. Sci. (In prep.). 5. Bartmann, R. M., and D. C. Bowden. 1984. Predicting mule deer winter mortality from weather data. Wi1dl. Soc. Bull. 6. Bear, G. D. 1984. Expanding telemetry collar for elk calves. Div. Game Info. Leafl. Colo. 7. Bear, G. D. 1985. Mark-recapture method applied to elk population estimate. J. Wildl. Manage. 8. Bowden, D. C., A. E. Anderson, and D. E. Medin. 1984. Sampling plans for sex and age ratios of mule deer. J. Wildl. Manage. 9. Dailey, T. A. N. T. Hobbs, and T. N. Woodard. 1984. Experimental comparisons of diet selection by mountain sheep and mountain goats. J. Wildl. Manage. 10. Freddy, D. J. 1984. Heart rates for activities of mule deer at pasture. J. Wildl. Manage. 11. Freddy, D. J., W. M. Bronaugh, and M. C. Fowler. 1985. Reactions of mule deer to human harassment during winter. Wi1d1. Soc. Bull. �4 12. Hobbs, N. T., B. C. Welch, T. E. Remington, and R. Sayre. 1984. Effects of sagebrush on in vitro digestion of grass cell wall. Wildland Shrub Symposium. The Biology of Artemisia and Chrysothomnus. 13. Hobbs, N. T., and R. A. Spowart. 1985. Effects of prescribed fire on nutrition of mountain sheep and mule deer during winter and spring. J. Wildl. Manage. 14. Kufeld, R. C., M. L. Stevens, and D. C. Bowden. 1985. Winter variation in nutrient and fiber content and in vitro digestibility of mountain mahogany (Cercocarpus montanus) and serviceberry (Amelanchier alnifolia) from diversified sites in Colorado. J. Range Manage. 15. Lance, W. R., and T. M. Pojar. 1984. Diseases and parasites of pronghorn: a review. Colo. Div. Wildl. Spec. Rep. 16. Pojar, T. M., and L. L. W. Miller. 1984. Recurrent estrus and cycle length in pronghorn. J. Wildl. Manage. 17. Torbit, S. C., L. H. Carpenter, A. W. Alldredge, and D. M. Swift. 1985. Mule deer body composition--a comparison of methods. J. Wildl. Manage. 18. Torbit, S. C., L. H. Carpenter, D. M. Swift, and A. W. Alldredge. 1985. Differential loss of fat and protein by mule deer during winter. J. Wildl. Manage. PUBLICATION PROGRESS 1. Job Progress Reports These reports have been printed in: Colo. Div. of Wild1. 1985. Res. Rep. Wildl. 2. Anderson, A. E., and O. C. Wallmo. 1984. Mule and black-tailed deer. Mammalian Species. This manuscript was published as: Mammalian Species - 1984. Odocoileus hemionus. Amer. Soc. r·1amm.No. 219:1-9. 3. Baker, D. L., and D. R. Hansen. 1984. Comparative digestion of grass in mule deer and elk. This manuscript as published in J. Wildl. Manage. 1985. 49(1):77-79. 4. Baker, D. L., N. T. Hobbs, and W. Vanhoven. 1985. In vitro rates of digestion of native forages. J. Anim. Sci. (In prep.). No progress was made on this manuscript. 5. Bartmann, R. M., and D. C. Bowden. 1984. Predicting mule deer winter mortality from weather data. Wild1. Soc. Bull. This manuscript was published in Wildl. Soc. Bull. 1984. 12:246-248. 6. Bear, G. D. 1984. Expanding telemetry collar for elk calves. Div. Game Info. Leafl. No progress was made on this manuscript. Colo. �5 7. Bear, G. D. 1985. Mark-recapture method applied to elk population estimate. J. Wi1d1. Manage. The second draft of this manuscript is being prepared. 8. Bowden, D. C., A. E. Anderson, and D. E. Medin. 1984. Sampling plans for sex and age ratios of mule deer. J. Wi1d1. Manage. This manuscript was published as: Bowden, D. C., A. E. Anderson, and D. E. t4edin. 1984. Sampling plans for mule deer sex and age ratios. J. Wi1d1. Manage. 48(2):500-509. 9. Dailey, T. A. N. T. Hobbs, and T. N. Woodard. 1984. Experimental comparisons of diet selection by mountain sheep and mountain goats. J. Wi1d1. Manage. This manuscript was published as: Dailey, T. V., N. T. Hobbs, and T. N. Woodard. 1984. Experimental comparisons of diet selection by mountain goats and mountain sheep in Colorado. J. Wi1d1. Manage. 48(3) :799-806. 10. Freddy, D. J. 1984. Heart rates for activities of mule deer at pasture. J. Wi1d1. Manage. This manuscript was published in J. Wi1d1. Manage. 1984. 48(3):962-969. 11. Freddy, D. J., W. M. Bronaugh, and M. C. Fowler. 1985. Reactions of mule deer to Human harassment during winter. Wi1d1. Soc. Bull. This manuscript has been accepted for publication by the Wildl. Soc. Bull. 12. Hobbs, N. T., B. C. Welch, T. E. Remington, and R. Sayre. 1984. Effects of sagebrush on in vitro digestion of grass cell wall. Wildland Shrub Symposium. The Biology of Artemisia and Chrysothomnus. This manuscript is in press. i 13. Hobbs, N. T., and R. A. Spowart. 1985. Effects of prescribed fire on nutrition of mountain sheep and mule deer during winter and spring. J. Wild1. Manage. This manuscript was published in J. Wi1d1. Manage. 1984. 48(2):551-560 .. 14. Kufe1d, R. C., t,1. L. Stevens, and D. C. Bowden. 1985. Winter variation in nutrient and fiber content and in vitro digestibility of mountain mahogany (Cercocarpus montanus) and serviceberry (Ame1anchier a1nifo1ia) from diversified sites in Colorado. J. Range Manage. This manuscript has been accepted by J. Range Manage. as Kufe1d, R. C., ~1. L. Stevens, and D. C. Bowden. 1985. Winter variation in nutrient and fiber content and in vitro digestibility of mountain mahogany (Cercocarpus montanus) and serviceberry (Amelancier a1nifo1ia). 15. Lance, W. R., and T. M. Pojar. 1984. Diseases and parasites of pronghorn: a review. Colo. Div. Wi1dl. Spec. Rep. This manuscript was published as Colo. Div. Wi1dl. Spec. Rep. No. 57. 14pp. 16. Pojar, T. M., and L. L. W. Miller. 1984. Recurrent estrus and cycle length in pronghorn. J. Wi1d1. Manage. This manuscript was published by J. ~lildl. Manage. 1984. 48(3):973-979. �6 17. Torbit, S. C., L. H. Carpenter, A. W. Alldredge, and D. r~. Swift. 1985. Mule deer body composition--a comparison of methods. J. Wildl. Manage. This manuscript was published by J. Wildl. Manage. 1985. 49(1):86-91. 18. Torbit, S. C., L. H. Carpenter, D. M. Swift, and A. W. Alldredge. 1985. Differential loss of fat and protein by mule deer during winter. J. Wildl. Manage. This manuscript was published by J. Wildl. Mange. 1985. 49(1):80-85. Additional publications which were not scheduled but which were published or accepted for publication during the 1984-85 Segment, are listed below: Baker~ D. L., and N. T. Hobbs. uutdoors 33(5):11-15. 1984. A diet for deer. Colo. Garrott, R. A., R. t~. Bartmann, and G. C. Whi teo 1985. Compari son of radio-transmitter packages relative to deer fawn mortality. J. Wildl. Manage. 49(3):758-759. Hobbs, N. T., and D. r~. Swift. 1985. Estimates of carrying capacity incorporating explicit nutritional constraints. J. Wildl. Manage. 49(3):814-822. Lee, J. E., G. C. White, R. A. Garrott, R. ~1. Bartmann, and A. W. Alldredge. 1985. Accessing (sic) accuracy of a radiotelemetry system for estimating animal locatons. J. Wildl. Manage. 49(3):658-663. Strong, L. L., R. W. Dana, and L. H. Carpenter. 1985. Estimating phytomass of sagebrush habitat types from micro densitometer data. J. Photog. Engin. and Remote Sensing. 51(4):467-474. White, G. C., L. H. Carpenter, and D. R. Anderson. 1985. Application of expert systems in wildlife management. Trans. N. Amer. Wildl. Conf. 50 (in press). A~~ Prepared By ~ Len H. Carpenter Wildlife Research Leader �Colorado Division of Wildlife Wildlife Research Report July 1984 7 JOB PROGRESS REPORT State of Colorado Project No. 01-03-047 Mammals 1 Research Work Plan Multispecies Investigations Job. No. ------------------No. 1 --~------------ 8 -------------------- Period Covered: Authors: Personnel: July 1,1984 Mammals Publication Editing and Library Services - June 30,1985 M. W. Hershcopf, L. H. Carpenter R. B. Gill, N. McEwen, L. Lovett, D. Phillips ABSTRACT During the 1984-85 Segment, 11 books were purchased for permanent reference by DOW personnel. Twenty-five additional publications were located, ordered, and obtained free of charge for use. Twenty-four theses were purchased, obtained on interlibrary loan, or given to the library. An additional 600 individual references requested by Mammals Researchers were located by library staff and made available for use. About 35 of these requests were not available locally and were obtained through interlibrary loans. ��9 MAMMALS PUBLICATION EDITING AND LIBRARY SERVICES Marian W. Hershcopf and Len H. Carpenter P. N. OBJECTIVE To provide a centralized support program for mammals research manuscript editing and library services so that Mammals Research scientists can allocate additional time to research. SEGMENT OBJECTIVES To provide coordinated, efficient, and economic editing and library services to all Colorado Mammals Research programs (Federal Aid Projects). SUt1MARYOF SERVICES Publications Purchased with Mammals 1 and 2 Funds and Placed in the Research Center Library Bunnell, F. L., D. S. Eastman, andJ. M. Peck, eds. 1983. Symposiumon natural regulation of wildlife populations. Proc. NW Sec., The Wildl. Soc., March 10,1978, Vancouver, B.C. 225pp. Caughley, G. 1983. The deer wars: the story of deer in New Zealand. Heinemann Publ., Exeter, NH. 187pp. Halls, L. K., ed. 1984. The white-tailed deer: ecology and management. Stackpole Books, Harrisburg, PA. 870pp. Hubner, K. 1983. Critique of scientific reason. Chicago, IL. 283pp. Univ. of Chicago Press, Krausman, P. R., and tJ. S. Smith, eds. 1984. Deer in the Southwest: a workshop, New Mexico State Univ., Las Cruces, April 16 and 17,1984. Univ. Ariz. Coop. Wildl. Res. Unit., Tucson. 131pp. Medawar, P. B. 1984. The limits of science. New York. 108pp. Harper and Row, Publ., National Res. Coun., Comm. Animal Nutrition. Nutrient requirement of beef cattle. 6th Rev. Ed. Subcommittee on Beef Cattle Nutrition, Committee on Animal Nutrition, Board on Agricul., Natl. Res. Counc. Natl. Acad. Press. Washington, DC. 90pp. Romesburg, H. C. 1984. Cluster analysis for researchers. Learning Publ., Belmont, CA. 334pp. Russell, F. 1983. The hunting animal. Life-time Harper and Row, New York. 211pp. �10 Seber, G. A. F. 1982. The estimation of animal abundance and related parameters. 2nd Ed. MacMillan Pub1. Co., New York. 654pp. Wolf, R. R. 1984. Tracers in metabolic research: radioisotope and stable isotope - mass spectrometry methods. Alan R. Liss, Inc., New York. 287pp. Publications Obtained Free or at Low Cost In addition to books purchased with Federal Aid Funds, about 25 free reports and short publications from state or federal agencies or from private sources, were located, ordered and obtained for use by Mammals Research personnel. Theses Purchased, Obtained on Interlibrary Loan or as Gifts for Use by Researchers Amidon, P. H. 1968. New York deer hunters: a comparison of deer law violators and non-violators. M.S. Thesis, State College of Forestry at Syracuse Univ. Syracuse, NY. 143pp. Aune, K. 1981. Impacts of winter recreationists on wildlife in a portion of Yellowstone National Park. M.S. Thesis, Montana State Univ. Bozeman. 111 pp. Barrows, P. L. 1981. Studies on Sarcocystis infections in domestic and wild swine. Ph.D. Dissertation, Univ. of Georgia, Athens. 95pp. Beidleman, R. G. 1954. vertebrate habitat. The Cottonwood river-bottom community as a Ph.D. Thesis, Univ. of Colo., Boulder. 357pp. Bessey, K. M. 1983. Analysis of the illegal harvest of white-tailed deer in Agro-Manitoba: implications for program planning and management. M.N.R.M. Thesis, Univ. of Manitoba, Winnipeg, CAN. 139pp. Bouckhout, L. W. 1972. The behavior of mule deer in winter in relation to the social and physical environment. M.S. Thesis, Univ. of Calgary, Alberta, CAN. Campbell, R. B. 1970. Pronghorn sheep and cattle range relationships in Carter County, MT. ~1.S. Thesis, Montana State Univ., Bozeman. 87pp. Crum, J. M. 1977. Some parasites of black bears (Ursus americanus) in the southeastern United States. M.S. Thesis, Univ. of Georgia, Athens. 76pp. Eiler, J. H. 1981. Reproductive biology of black bears in the Smokey Mountains of Tennessee. M.S. Thesis, Univ. of Tennessee, Knoxville. 127pp. E1owe , K. D. 1984. Home range, movements, and habitat preferences of black bear (~rsus americanus) in western Massachusetts. M.S. Thesis, Univ. of ~1assachusetts, Amherst. 112pp. �11 Hakonson, T. E. 1967. Tissue distribution and excretion of 134CS in the mule deer. M.S. Thesis, Colo. State Univ., Fort Collins. 121pp. Halfpenny, J. C. 1980. Reproductive strategies: intra and interspecific comparison within the genus Peromrscus. Ph.D. Thesis, Univ. of Colo., Boulder. 160pp. (Courtesy - C. oeffler).Helm, D. J. 1981. Vegetation Helm, D. J. 1981. Vegetation diversity indexes in several vegetation types of western Colorado. Ph.D. Thesis, Colo. State Univ., Fort Collins. 113pp. Hunt, C. 1984. Behavioral responses of bears to tests of repellents, deterrents, and aversive conditioning. M.S. Thesis, Univ. of Montana, Missoula. 137pp. Huot, J. 1984. Body condition and food resources of white-tailed deer on Anticost i Island, Quebec. Ph.D. Dissertation, Univ. of Alaska, Fairbanks. 258pp. Koch, D. B. 1983. Population, home range and denning characteristics of black bears in Placer County, Calif. M.S. Thesis, California State Univ., Sacramento. 71pp. Kopf, V. E. 1983. The relationship between food intake and thyroid hormone concentrations in white-tailed deer. Ph.D. Dissertation, Virginia Polytech. Inst. & State Univ., Blacksburg. 163pp. Lee, J. E. 1984. Mule deer habitat use and movements on Piceance Basin winter range as estimated by radiotelemetry. M.S. Thesis, Colorado State Univ., Fort Collins. 59pp. Nowlin, R. A. 1985. Distribution of moose during occupation of vacant habitat in northcentral Colorado. Ph.D. Thesis, Colorado State Univ., Fort Collins. 60pp. Certley, K. D. 1981. Studies on toxoplasmosis in white-tailed deer (Odocoileus virginianus). M.S. Thesis, Univ. of Georgia, Athens. 124pp. Parmenter, R. A. 1984. Food and energy intake of coyotes determined from turnover of tritiated water and sodium-22. M.S. Thesis, Colorado State Univ., Fort Collins. 55pp. Pulliam, D. E., Jr. 1978. Determination of digestibility coefficients for quantification of fecal analysis with elk. M.S. Thesis, Washington State Univ., Pullman. 34pp. Skiba, G. T. 1981. Ecological evaluation of the Dinosaur National Monument bighorn sheep herd. M.S. Thesis, Colorado State Univ., Fort Collins. 107pp. Smith, R. 1983. Mule deer reproduction and survival in the LaSal '''ountainsof Utah. M.S. Thesis, Utah State Univ., Logan. 104pp. �12 Reference Document Location and Delivery The Research Center Library staff also located and delivered about 600 individual articles on request for Mammals Researchers during this segment; about 35 were not available locally and were obtained through interlibrary loan procedures. Prepa red by ev-~"-~rf-- M ~.r;~ t-J. H Marlan w. Aershcopf Librarian Len H. Carpenter Wildlife Research Leader I �Colorado Division of Wildlife Wildlife Research Report July 1985 13 JOB PROGRESS REPORT State of Colorado Project No. 01-03-047 Mammals 1 Research Work Plan No. Deer Investigations -------------------- Job. No. 2 ----------------- 1 ---------------------- Period Covered: Author: July 1,1984 Quantifying Capacity of Winter Ranges to Support Deer--Evaluation of Thermal Cover Used by Deer - June 30,1985 D. J. Freddy Personnel: W. M. Bronaugh, D. C. Bowden, G. R. Winfield ABSTRACT Effects of the presence and absence of thermal cover on nutritionally stressed mule deer were monitored for 108 days from 11 December 1984 - 29 March 1985. Percent weight loss was not different between deer with and without thermal cover (P > 0.50). Deer with cover spent more time lying (P < 0.09) and less time Toraging (P < 0.5) than deer without cover, impTying that deer without cover attempted to compensate for an energetic disadvantage by seeking forage. Nutritionally stressed deer used thermal cover more than did well-fed deer and differences in behavior patterns between deer with and without cover increased when deer were nutritionally stressed. Interactions among nutritional status, use of thermal cover, and behavior are implicated. However, absence of thermal cover did not appear to reduce potential for deer to survive winter. ��15 NUTRITIONAL BASIS FOR QUANTIFYING CAPACITY OF WINTER RANGES TO SUPPORT DEER Davi d J. Freddy P. N. OBJECTIVE To determine if a system can be developed to estimate number of deer winter ranges are capable of supporting. SEGMENT OBJECTIVES 1. Conduct further experimentation on the interactions between nutritional level and use of thermal cover by deer. 2. Complete data summary and analysis of 1983-84 experiment. 3. Prepare a manuscript inclusive of results from experiments in 1982-83, 1983-84, and 1984-85 for submission to an appropriate journal. ACKNOWLEDGMENTS The U.S. Forest Service, Rocky Mountain Forest and Range Experiment Station, kindly loaned instrumentation for measuring weather conditions. Whitcomb M. Bronaugh provided diligent assistance in collecting and summarizing data, and Lynn L. Stevens provided laboratory analyses of rations fed to deer. G. R. Winfield provided customized software for summarizing data on an Apple lIe. ~1ETHODS AND t'ATERIALS See Program Narrative (Appendix A). RESULTS AND DISCUSSION Effects of presence and absence of thermal cover on deer behavior, weight performance, and intake of pe11eted ration were monitored for 108 days from 11 December 1984 - 29 March 1985. Behavior of 12 deer, of which 6 had access to thermal cover, was monitored during 6, 6-day, 24-hr trials conducted about every 2 weeks. Body weights of deer were measured every 18' days while intakes of restricted amounts of pe11eted ration were monitored daily. Deer adjusted well to prolonged confinement in isolation pens and exhibited minimal nervous behavior. Ten of 12 deer used in 1985 were common to experiments in 1984. Weather Conditions Weather conditions were relatively moderate throughout the winter and during behavior trials. Snow coverage was complete within cover and no-cover �16 isolation pens during 5 of 6 trials. did heavy fog (Table 1). Strong winds occurred sporadically as Comparisons of weather conditions between trials in 1984 and 1985 revealed that temperatures were generally warmer in 1985 and were significantly warmer (P ~ 0.10) primarily during trials 3 and 5, and solar radiation did not vary-between years (P ~ 0.20}(Tab1e 2, unpaired t-tests). Assuming a lower critical temperature of -23C (Mautz et a1. 1984), deer were not metabolically stressed in 1985 but were likely stressed in 4 of 6 trials in 1984 (Table 2). Deer Diet Beginning in October, 1984, all deer were fed ad libitum a pe11eted ration (special deer-elk wafers) having 70% in vitro digestible dry matter (IVDDM), 3.16 kcal DE/g, and 24% crude protein. During the experiment, 2 rations were fed: deer-elk wafers were fed 11-25 December 1984; and Low Energy II pellets having 60% IVDDt~, 2.59 kcal DE/g, and 13% crude protein were fed 26 December 1984 - 29 March 1985 (see Freddy 1984 for ingredients of rations). Additionally, 50g of chopped grass hay (61% IVDDM, 2.64 kca1 DE/g, 7% crude protein) were added to each of 2 allotments of pelleted ration given daily to all deer. The ration was fed to each deer on the basis of metabolic body weight so that all deer received 70-75% of their estimated digestible energy required for maintenance (Ullrey et al. 1969). To fulfill an apparent need by deer to forage on fibrous native vegetation (essentially not available in isolation pens), 15 x 45-cm lengths of freshly cut aspen (Populus tremuloides) logs were given to deer every 5-7 days. All deer actively ate the aspen bark and chewed on remaining wood fiber. Snow was available to deer as a source of water throughout the experiment. Three deer (157, 176, 177) chewed and injested their hair during the experiment. Deer 157 (cover deer) died during trial 5 from malnutrition which was possibly hastened by extensive hair coat deterioration. Both deer 176 (no cover) and deer 177 (cover) suffered from moderate loss of hair and were used as a pair in the experiment. Intake of Digestible Energy All deer ate their al10ted ration with few exceptions. Deer 177 reduced intake for 12 days due to apparent digestive problems while deer 191 voluntarily reduced intake for short periods of time on an irregular basis. Intake of digestible energy was, therefore, the same for deer with and without cover. During 1984, when deer were fed ad libitum throughout the experiment, intake of digestible energy was also not different between deer with and without cover (Freddy 1984). Changes in Body Weight There were no differences in percent weight loss between deer with and without cover over the 108-day experiment (P ~0.50, l-sample t-tests). This conclusion was based on 3 tests conducted at the end of trials 4, 5, and 6 to reflect the decrease in numbers of healthy deer pairs which declined from 6 to 4 (Table 3). Average percent weight loss for deer with �17 cover and deer without cover at the conclusions of trials 4, 5, and 6 were, respectively: -14.8 + 2.0(SE), -13.4 + 1.3; -16.1 + 1.1, -16.4 + 2.2; and -17.4 + 1.2, -18.3+1.0. Two pairs of deer were removed from the experiment because of excessive weight loss. Deer 157 lost 22% body weight and died of malnutrition during trial 5, thus effectively removing deer 150 from the experiment. After trial 5, deer 176 had lost 24% body weight and was obviously weak. This resulted in the removal of deer 176 and 177 (Table 3). Similar changes in body weight for deer with and without cover strongly suggested that thermal cover provided little, if any, energetic advantage, even when deer were nutritionally stressed. There was also no difference in weight loss between deer with and without cover 1984 when deer were fully fed. Percent weight loss in 1984 was 6.2 + 0.7(SE) for adult deer having cover and 7.1 ~ 3.4(SE) for adults not having cover. This experiment contrasted deer with and without cover; an assumed difference of 100% in potential treatment effect. Torbit et ale (1985, Fig. 1) demonstrated that body weights of mule deer declined an additional 12% when dietary energy inake was reduced only 5%. This small change in energy input caused a much greater change in body weight than the dichotomy of cover - no-cover. Use of Thermal Cover Avoidance or selection of cover treatments by deer was determined by comparing the average proportion of lying activity for all cover deer/treatment with the area that each treatment accounted for within pens. This assumed that if deer were not selecting for cover treatments, they would lie on treatments in proportion to treatment areas. In 1984, when deer were fully fed, deer with cover avoided lying on control areas (~~ 0.10, l-sample t-tests) in all trials except trial 5 (Fig. 1). In 1985, nutritionally stressed deer with cover avoided lying on control areas during all trials (P ~ 0.03, Fig. 1). The proportion of lying activity on control areas-was not related to body weight or percent body weight loss (linear, exponential, and logarithimic regressions, r < 0.63, ~~ 0.10) during either 1984 or 1985. During lying activity, deer spent 43 + l(SE) and 14 + l(SE) % of their time on control areas in 1984 and 1985, respectively. This large difference in use of control areas may reflect differences in thermal cover treatments (see Appendix A, Freddy 1984 and this report), but also suggests that nutritional status of deer affected their use of thermal cover. Greater use of thermal cover in 1985 occurred even though temperatures were warmer in 1985 (Table 2). During both years, relative use of thermal treatments changed between trials, suggesting deer altered their preference for cover. In 1984, use by deer of the dry, dark-soil treatment inceased through trial 4 (Fig. 1). In trial 5, this soil treatment was wet from rapid snow melt (likely responsible for the �18 insignificant use of cover in trial 5). During the warmer temperatures of trial 6, deer often sought shade afforded by the tree treatment, suggesting heat loss was minimally important to deer. In 1985, use of the west-facing, dark-soil wind panel treatment markedly inceased after trial 1 and remained high until trail 6 when deer utilized shade, provided primarily by the hut treatment (Fig. 1). During both years, deer strongly associated with dark soil treatments suggesting that conductive heat loss may be an important energy drain. Use of shade by mule deer during winter requires further analysis, but deer could be heat stressed in late winter. In 1985, deer with cover spent varying amounts of time sleeping or ruminating while standing. During all trials, deer avoided standing on control areas (P < 0.01, 1-samp1e t-tests) even though they were frequently on control areas (Fig. 2). Use of the hut treatment markedly increased between trials 1 and 6. Because the hut provided either warmer temperatures or shade, such increased use is difficult to interpret. I suspect the high use during trial 6, however, is related to deer seeking shade. Behavior Comparisons Behavior activities were compared between deer with and wouthout cover. In 1984, frequencies of lying and foraging activity were not different for all trials combined (P > 0.40, 1-samp1e t-tests) or within trials (P > 0.12, paired t-tests). -L5!ing activity inceased and foraging activity-decreased for all deer between trials 1 and 6 (Fig. 3). The decrease in foraging activity corresponded to a voluntary reduction in intake of pe11eted ration that was 35% for fawns and 38% for adults from trial 1 to trial 6 (Freddy 1984) • In 1985, deer with cover spent more time lying (P ~ 0.09) and less time fora~i ng (P ~ 0.05) than deer without cover across all tria1s (Fig. 3). Withln trials, differences in lying activity occurred in trials 1 and 2 (P < 0.10) and differences in foragfng occurred in trials 1, 2, 3, and 4 (~ ~ 0.08, Fig. 3). Deer without cover, therefore, spent more time seeking limited native forage or chewing on aspen logs because all deer quickly ate the limited amounts of pe11eted ration that were offered twice/day. The data imply that deer without cover attempted to cmpensate for an energetic disadvantage by seeking forage. Differences in time spent lying by deer with cover and without cover occurred on specific days within trials (P < 0.10, paired t-tests). On these days, deer without cover were more active during daylight hours and spent more time foraging during daylight hours (P ~0.10, paired t-tests). These comparisons suggested that deer without cover "waited" for temperatures to moderate before they attempted to "capture" energy by increased foraging. The implication of this behavior pattern may be that if forage is available, even low quality forage, deer without cover can compensate behaviorally and achieve the same rate of body weight loss as deer with cover. �19 CONCLUSIONS Degree of weight loss in well-fed and nutritionally stressed deer was not affected by the presence or absence of thermal cover. Thermal cover, therefore, appeared to minimally enhance the potential for deer to survive winter. Paradoxically, use of thermal cover by deer increased as nutritional status declined. Nutritional status also altered the relative behaviors of deer with cover and without cover which suggested that deer without cover were adjusting to or compensating for the lack of thermal cover. If low quality forage is available, deer without cover may be able to sufficiently alter their behavior so as to achieve similar rates of body weight loss as deer with cover. Further experimentation should monitor physiological parameters of individual deer as they are experimentally forced to adjust to the presence or absence of thermal cover. LITERATURE CITED Freddy, D. J. 1984. Evaluation of thermal cover used by deer. Wi1d1. Game Res. Rep. July (1):21-50. Colo. Div. r·1autz,W. \4., P. J. Pekins, and J. A. Warren. 1984. Cold temperature effects on metabolic rate of white-tailed, mule, and black-tailed deer in winter coat. Proc. Int. Conf. on Biology of Deer Production. Dunedin, New Zealand. Torbit, S. C., L. H. Capenter, D. M. Swift, and A. W. Alldredge. 1985. Differential loss of fat and protein by mule deer during winter. J. Wi1d1. Manage. 49:80-85. Ul1rey, D. E., W. G. Youatt, H. E. Johnson, L. D. Fay, B. L. Schoepke, and W. T. Magee. 1969. Digestible energy requirements for winter maintenance of Michigan white-tailed does. J. Wi1d1. ~'anage. 33:482-490. �20 Table 1. Experimental conditions for cover trials in 1984-85. Trial Dates Conditions 1 16-21 Dec 1984 100% snow coverage in all pens; strong wind. with blowing snow 2 3-8 Jan 1985 100% snow coverage in all pens; light wind. considerable overcast. some fog and light snow 3 22-27 Jan 1985 100% snow coverage in all pens; coldest temperatures; some heavy fog; no wind. heavy snowfall 4 11-16 Feb 1985 100% snow coverage in all pens; some strong wind; moderate temperatures 5 27 Feb-4 Mar 1985 100% snow coverage in all pens; moderate wind and temperatures 6 18-23 Mar 1985 Snow in pens rapidly melting; warm temperatures; exposed soil in pens wet; cover treatments needed constant maintenance to keep dry; strong wind �21 Table 2. Temperatures and solar radiation values for behavior trials during cover ex~eriments in 1984 and 1985. Temperature °c Total solar ambient radiation black 910be Trial 1 2 Dates cal/cm2/day 0 1 -1 -15 1 -14 2 17a 2 lla 2 184 17 171 29 -22 2 -18 3 _30d 14 3 13 1 208 18 196 21 1 _22d 2 lle 1 20e 2 258 20 19 2 22 3 338 6 303 29 16-21 Dec 1984 SE x SE x SE x SE -21 2 -18 3 -8 2 x SE -30b 1 -22b 2 -16c 1 _7c 2 x SE x SE -22 2 -5 1 -16 3 0 3 -24f 2 -18f 3 x SE x SE -209 2 -159 2 _3h 1 2h 2 -23 i 2 _16i 2 23 2 25 3 384 25 406 29 x SE x SE -10 1 -8 1 2 1 6 1 -llj 1 j -9 1 28 4 20 3 428 36 490 31 3-8 Jan 1984 16-21 Jan 1984 1-6 Feb 1984 11-16 Feb 1985 17-22 Feb 1985 27 Feb-4 Mar 1985 6 max -13 1 -11 1 x SE 5 min x 22-27 Jan 1985 4 max 13-18 Dec 1983 3-8 Jan 1985 3 min 7-12 Mar 1984 18-23 Mar 1985 1 -4 2 apairs of letters indicate means different (f _2_ O. 10) . 265 20 �22 Table 3. Percent cummulative change in body weight for cover (c) and no-cover (nc) deer at the conclusion of each of 6 behavior trials during 1985. Deer listed in order of exeerimental eairs. Behavior trial Deer 1 16Bc -4.9 162nc 2 3 ' 4 5 6a -17.0 -11. 3 -20.7 -17.6 -11.4 -4.6 -11.0 -10.4 -B.9 -15.6 -10.4 191c 225nc -2.6 -6.5 -6.7 -B.3 -13.5 -15.0 -3.4 -B.9 -6.3 -lO.B -13.9 -16.B 157c 150nc -7.2 -4.2 -10.4 -11.4 -13.5 -9.9 -22.2 -15.5 lnc 176nc -1.0 -4.0 -15.4 -9.6 -14.B -12.2 -17.7 -lB.O -19.9 -24.2 156c 194nc -1.9 -3.0 -6.4 -7.3 -6.3 -13.5 -10.4 -15.3 -14.3 -17.6 -17.5 154c 144nc -2.7 -2.0 -6.7 -7.2 -B.3 -11 .7 -15.1 -15.0 -16.4 -21.4 -7.0 -9.0 apercent weight change at end of lOB-day experiment. -lB.3 �23 60 II O 1985 HUT WEST WIND-SOIL 50 m 40 30 20 SOUTH WIND-501L HUT- Uld_• 10 ••• III LJ I n ID b 0 ........~...,......~ ::~TROL MAR ;:) ... Z ••• U tIC ••• A. 70 1984 a a 60 b b b 50 40 30 20 II TREE EJ D SOIL • r: r WIND CONTROL 10 0 DEC JAN 2 . f JAN 3 4 TRIA Fig. 1. E. 5 6 L Average percentages of lying activity on cover treatments by 6 mule deer during 6 trials conducted between December and March in 1984 and 1985. Only those trea~ments receiving >2% of the lying activity within a trial are shown for 1985. Avoidance of control areas significant at: a = P < 0.10, b = f ~0.05, c = P < 0.001. -- �24 ~ o HUT • HUTSOIL D 70 D HUTEAST WEST WIND-SOIL b b 50 b ;:) ... Z 30 I¥ 20 A- c 40 ••• U ••• CONTROL b b 60 ••• In • SOUTH WI NO-SOIL I~ 10 ~ 0 DEC JA N 2 118 JAN FEB 3 4 FEB-MAl 5 MAl 6 TR IA L Fig. 2. Average percentages of standing activity on cover treatments by 6 mule deer during 6 trials conducted between December and March in 1985. Only those treatments receiving >2% of the standing activity within a trial are shown. Avoidance of control areas significant at: b = f ~0.05, c = P < 0.01. �25 70 65 60 55 1985 25 20 ... -> -... u )0- c( 15 •...... b ~-~ •• 10 5 ... 0 Z 75 III b u III: 70 III A. 65 60 198'4 55 20 15 10 5 0 2 3 4 5 6 TRIAL Fig. 3. Average percentages of lying and foraging activities for deer with and without cover during 6 trials conducted between December and March in 1984 and 1985: deer with cover lyi ng (.) and foragi ng (.), deer wi thout cover lyi ng (0 ) and foraging (6.). Differences in activities between deer with and without cover significant at: a = P < 0.10, b = P < O. 05. - - �26 APPENDIX A .' ?;:WGR':\~1 NAR.I~ATIVE 11cS~ State: Colorado Project Title: Big Game Investi~ations Project No. 01-00-071, 15050 ~.;rork Plan Title: Work Plan No. 2 Job Title: Job No. 1 Prepared Submitted Deer Investi gati o_I]_?__--;----:=--_ Nutritional Basis for Quantifying Capacity of Winter Range -- Evaluation of Thermal Cover Used by Deer by: by: Dave Freddy Date: November 7. 1984 Dave Freddy Date: November Date: Date: Date: Date: Date: Approved by: Date: Date: 1984 �27 11/84 ammend. PROGRAM NARRATIVE State: Colorado Big Game Investigations Proj ec t No._-,0-,-1_-0,;;...0,---,-0...;..7...;..1 ':_clc...;.5_:_0..:;..50;;___ __ Work Plan 2 ---------=------------1 Job No. A. - Cervids Deer Investigations Nutritional Basis for Quantifying Capacity of Winter Range -Evaluation of Thermal Cover Used by Deer NEED This study, which addresses the effects of thermal cover on mule deer energetics during winter, is now entering the third, and likely, final year. During the first winter of work in 1982-83, tame mule deer were provided artificial structures that could be used by deer as thermal cover (Freddy 1983). Because deer used these structures for a significant amount of time, a procedure for determining the effects of thermal cover on deer energetics was established: namely, provide artificial cover in a controlled experiment. In 1983-84, an experimental test of the effects of thermal cover on deer was conducted (Freddy 1984). Twelve tame mule deer were involved in this paired-design experiment. Six deer had access to artificial cover structures while 6 deer had no thermal cover during the lOa-day period of 9 December 1983 to 17 March 1984. All deer were fed the same pelleted ration ad libitum. Body weight, intake of digestible energy, and behavior were monitored for both groups of deer throughout the lOa-day period. Results of the experiment were: l} there was no difference in the degree of body weight gain or loss between cover and no-cover deer, 2} there was no difference in intake of digestible cover and no-cover deer. energy between Thus, the net f low of energy in deer, as judged by changes in body weight and intake of energy, was not affected by the presence of thermal cover. These results supported conclusions of Swift et a1. (1980) and Gilbert and Bateman (1983) and opinions of Peek et al. (1982) and Moen (1966) that thermal cover plays a minor role in the survival of deer during winter. Is this conclusion warranted, or could further experimentation provide a more conclusive or even divergent answer? There are 2 possible weaknesses in the design experiment. First, deer had unlimited access and, thus, unlike wild deer, were not stressed although the cover structures provided thermal of the 1983-84 to a pe11eted ration nutritionally. Second, advantages to deer, �28 Freddy cover may not have been adequate enough to affect the physiological performance of deer. During this third winter of the study, the experiment will be modified to address these 2 possible inadequacies. First, deer will be fed a pe11eted -ration.but at-a rate below maintenance, and feed will be available only at certain times of the day. Deer will thus be stressed nutritionally both.in quantity and availability of feed. This feeding regime will more accurately reflect the status of wild deer during winter. -Second, cover structures will be modified so that deer have shelter far in excess of what would be available to wil d deer. To state that thermal cover is unimportant to mule deer during winter is contrary to many habitat improvement and mitigation programs. Thus, there is a need to continue this project through the third winter of experimentation in order to more completely clarify the role of thermal cover in the over-winter survival of mule deer. B. OBJECTIVES This research will compare the rate of body weight loss between nutritionally stressed deer having, and not having access to thermal cover. Additionally, food intake and activity patterns of cover and no-cover deer will be compared. This study is sequential to work completed during the 1983-84 winter. C. EXPECTED RESULTS OR BENEFITS Completion of this research will aid in developing a habitat evaluation system by establishing the role of cover in maintaining energy balance of deer during winter. The habitat evaluation system will provide a quantitative basis for the Division of Wildlife to assess and improve winter ranges used by deer. D. APPROACH The approach of this study will be to monitor the effects of the absence and presence of thermal cover on deer energetics during winter. There will be 2 groups of 6 tame deer. While being confined to individual isolation pens (Freddy 1984), deer in one group will have access to thermal cover while deer in the second group will not have access to thermal cover. Twelve isolation pens wi ll be divided into 6 pairs of pens so that pens within pairs are as identical as possible. A 3-sided and roofed artificial cover structure will be randomly placed within 1 pen of each pair (Fig. l). 7welve adult deer will be grouped into 6 pairs �29 Freddy based on similar body weights, age, and sex (Table 1). One deer from each pair will be randomly assigned to be hous~d in an isolation pen with cover and the other deer of each pair to a pen without cover. Deer will be housed in pens during a 108-day period from 11 December 1984 through 28 March 1985. Primary measurements on deer will be: 1) body weight at 18-day intervals, 2) frequency of orts (food offered but not eaten) within each 18-day interval, 3) percent lying and up activity during 6 24-hour behavior trials, and 4) frequency of use of thermal cover during 24-hour trials. Rationale: Compairing physiological and behavioral responses of nutritionally stressed deer having access to cover with deer not having access to cover should provide a strong experimental test of the role of thermal cover in deer energetics during winter. Hypotheses that will be tested: 1. Rate of body weight loss will not be different between "cover" and "no-cover" deer. 2. Frequency of arts (>25g/feeding) will not be different between "cover" and "no-cover" deer. 3. Activity patterns will not be different between "cover" and "no-cover" deer. Additionally, l. 2. 3. 4. for those deer having access to cover: Deer wi 11 Deer wi 11 Deer wi 11 Deer will March) . Maintenance exhibit no preference for thermal cover. use cover independent of time of day. use cover independent of ambient conditions. use cover independent of time of Year (December - of Experimental Deer Deer will be fed a pelleted ration having 58% in vitro digestible dry matter and 12.6% protein (Freddy 1984). Each deer will be fed at a rate approximately equal t8 60% of maintenance energy requirements (158-160 kcal DE/kg BW .75/day equals maintenance, Ullrey et al. 1969, 1970). This rate is conservatively predicted to cause weight losses of 10-17%, depending on initial body weights, during the l08-day period. Body weights will be determined every 18 days and if weight loss is inadequate, feed wi 11 be further res tri cted for a11 deer to a lower percentage of the maintenance requirement. Predicting weight loss is difficult because weight loss is dependent on intake and the time over which a given intake is �30 Freddy sustained. Carpenter (1979, 1980) achieved weight losses in adult deer of 17~ when deer were fed at 60% of maintenance for 70 days and losses of 16% when deer were fed at" 40% of maintenance for 56 days. Over a 140-day period, fawns fed at 75% of maintenance lost 16% of their weight and those fed at 60% of maintenance lost 24% (Carpenter 1980). I believe feeding initially at 60;; of maintenance is a reasonable approach to invoking nutritional stress on deer. A body weight loss of 20% over the 108-day period is desirable. Deer will be fed 1/2 their daily ration twice per day from 07300900 and from 1600-1730 hours. When behavior and cover-use trials are in progress, deer will be fed from 0600-0700 and from 17001800 hours to coincide with the ending of observation shifts (Freddy 1983). This twice/day instead of once/day feeding schedule will ameliorate the anticipated long time periods over which deer might have nearly empty rumens due to rapid passage of the pelleted ration (Robbins 1983:317). In past nutrition experiments where deer were fed only once/day at low percentages of maintenance, considerable hair chewing and deterioration of pelts occurred (Carpenter 1980). Th is prob 1em may be caused by boredom and/ or the feel i'ng of an "empty rumen." I must avoid this type of pelt deterioration and, therefore, have decided to invest in the extra labor needed to feed t\'1i ce/ day. Amount of ration fed to each deer will be based on body weight at the beginning of each lS-day period. Ration will be preweighed and bagged for each deer to facilitate the feeding process. Orts will be left in the feeders to allow each deer to consume their allocated amount of ration. Occurrence of orts (>25g/ feeding) will be recorded for each deer. Adequate snow or water will be provided to deer. Deer will be weighed using a platform scale. If deer reach a degree of body weight loss considered potentially lethal, deer will be removed from the experiment. Body weight loss of 25:; of the initial December body weight will be considered critical for adult deer (Carpenter 1979). Any deer having excessive hair loss due to ingestion (Carpenter 1979) will be removed from the experiment. Deer reaching critical body weights and removed will be refed using a pelleted ration and small amounts of alfalfa. Isolation Pens and Artificial Cover Isolation pens will be the same as in 1983-84 (Freddy 1984). However, artificial cover will be changed and will consist of a 3-sided-roofed structure having a windbreak on the open side (Fig. 2). Additionally, dark soil will be placed near the structure and windbreak on south and west exposures. A small �31 Freddy feeder located at the center of all pens will be the only structure within "no-cover" pens. Snow will be compacted within and around pens as in previous years and the dark soil will be manually kept free of snow. The 6 pairs of deer will be rotated through all 6 pairs of pens in a Latin Square design so that deer are in a different pen during each of the 18-day intervals (Table 2). Deer Behavior Observations of deer behavior will be made during 6 trials beginning in mid-December and ending in late March. Data collection periods will consist of 6 days each trial in which deer are observed during 4 3-hour observation periods: 03000600, 0900-1200, 1400-1700, and 2000-2300 hours (MST) (Freddy 1983) . At 10-minute intervals, behavior of all deer will be quickly determined using scan sampling (Freddy 1983). Behavior will be classified as bedded head-up, bedded head-down, standing, feeding, and other. Deer having cover will be assigned to a cover treatment and deer without cover will be assigned to a quadrant (Freddy 1983). Criteria for deciding whether deer are on cover treatments will be the same as in previous years (Freddy 1983). Ambient Conditions Ambient temperature, wind speed, wind direction, and solar radiation will be measured continuously using a self-recording thermograph, wind monitoring system, and pyranograph. Black globe temperature will be measured each one-half hour only during trial observation periods (Freddy, 1983, 1984). Data Analysis This experiment follows a paired comparisons design. Primary comparisons between pairs of "cover" and "no-cover" deer will be: 1) percent change in body weight; 2) frequency of orts; 3) frequency of activities, primarily time spent lying and active; and 4) frequency of head-up and head-down postures when lying. These variables can be compared within each 18-day interval and over the entire 108-day experimental period. Additi ona l ly , use of cover by "cover" deer wi 11 be determi ned to be significant or non-significant. Use of cover will also be assessed for relationships to ambient conditions. �32 Freddy Schedule Period Activity October 1984 Construct new cover structures; prepare isolation pens for experiment; deer weighed regularly at Fort Collins pen complex November 1984 Deer transported to Junction Butte facility; begin behavior adjustment to isolation pens 11 Decenber 1984 Deer weighed and placed in cover isolation pens; begin l08-day experimentation period; monitor intake 16 Decerrner 1984 Begin behavior trial 1 29 Deceraer 1984 Deer weighed and placed in new isolation pens; monitor intake 3 January 1985 Begin behavior trial 2 16 January 1985 Deer weighed and placed in new isolation pens; monitor intake 22 January 1985 Begin behavior trial 3 3 February 1985 11 February 1985 Deer weighed and placed in new isolation pens; monitor intake Begin behavior trial 4 21 February 1985 Deer weighed and placed in new isolation pens; monitor intake 27 February 1985 Begin behavior trial 5 11 March 1985 Deer weighed and placed in new isolation pens; monitor intake 18 j·la rch 1985 Begin behavior trial 6 28 ~'la rch 1985 END EXPERIMENT April - June 1985 Data summary; data analysis; prepare yearly reports �33 Freddy Personnel David J. Freddy Principal Investigator Uti1ity Worker I Field Assistance Clerk-Typist Secretarial Support Estimated Annual Costs Person-Days (01) Personal Services David J. Freddy 264 Utility Worker I 130 Clerk Typist 22 Retirement and Insurance (21) Operating Supplies and Services (28) Travel (31) Capital Expenses TOTAL Costs $36,876 7,020 1,455 5,378 $10,672 840 o $62,241 E. LOCATION Field work will be conducted at the Division of Wildlife Junction Butte Research Center located 6 km south of Kremmling, Colorado. F. RELATED FEDERAL AID PROJECTS 01-00-071,15050, WP2, Jobs 6 and 10 01-00-071,15051, Noncervid Research LITERATURE CITED Carpenter, L. H. 1979. Nutritional basis for quantifying capacity of winter ranges to support deer. Colo. Div. Wild1. Game Res. Rep., July Part II. 69-74pp. Carpenter, L. H. 1980. Nutritional basis for quantifying capacity of winter ranges to support deer. Colo. Div. Wildl. Game Res. Rep., July Part I. 84-98. Freddy, D. J. 1983. Evaluation of thermal cover used by deer. Colo. Div. Wildl. Game Res. Rep., July Part I. 31-68pp. �34 Freddy Freddy, D. J. 1984. Evaluation of thermal cover used by deer. Wild1. Game Res. Rep., July Part I. 21-50pp. Colo. Div. Gilbert, F. F., and M. C. Bateman. 1983. Some effects of winter shelter conditions on white-tailed deer, Odocoileus virginianus, fawns. Can. Field Nat. 97:391-400 Moen, A. N. 1966. Factors affecting the energy exchange and movements of white-tailed deer, western Minnesota. Ph.D. Thesis, Univ. of Mi nnesota. 121 pp , Peek, J. M., M. D. Scott, L. J. Nelson, and D. John Pierce, and L. L. Irwin. 1982. Role of cover in habitat management for big game in northwestern United States. Trans. North Am. Wildl. Nat. Res. Conf. 47:363-373. Robbins, C. T. 1983. York. 343pp. ~Hldlife feeding and nutrition. Academic Press, New Swift, D. t·!., J. E. Ellis, and N. T. Hobbs. 1980. Nitrogen and energy requirements of North American cervids in winter--a simulation study. Pages 24~-251 in E. Reimers, E. Gaare, and S. Skjenneberg, eds. Proc. 2nd Int. Reindeer/Caribou Symp., Roros, Norway, Ullrey, D. E., ~. G. Youatt, H. E. Johnson, L. D. Fay, B. L. Schoepke, and W. T. rlagee. 1969. Digestible energy requirements for winter maintenance of Michigan white-tailed does. J. Wildl. Manage. 33:482-490. Ullrey, D. E.. :L G. Youatt, H. E. Johnson, L. D. Fay, B. L. Schoepke, and W. T. Magee. 1970. Digestible and metabolizable energy requirements for winter ~aintenance of Michigan white-tailed does. J. Wildl. Manage. 34:863-869. �35 Freddy Table 1. Deer pair 1 Probable Qairings of deer for the 1984-85 cover eXQeriment. Age Weighta (yrs , ) Deer Sex 59.5 162 168 F F 2.5 2.5 61.0 2 225 191 F F 4.5 4.5 64.0 67.0 3 150 157 F 2.5 2.5 68.5 1.5 1.5 75.0 69.5 2.5 4.5 79.0 72.0 2.5 7.5 87.0 97.5 4 M 176 177 5 6 ~1 M 156 194 M 154 144 M F M 67.0 aBody weights (kg) as of 13 November 1984. Table 2. Rotation'of 6 pairs of deer (1-6) through 6 pairs of isolation eens (A-F)a during 6 18-da~ time Qeriods during 1984-85. Deer pairs (see Table 1) Time period 2 3 4 6 1 beginning 5 Dec Dec Jan Feb 21 Feb 11 Mar 11 29 16 3 aA E = = pens 2 & 11; B pens 6 & 10; F A B B C 0 E C D E C D E D E E F A F A B F A F A F = = F A B B C B C D C D E pens 5 & 12; C = pens 1 & 4; D pens 7 & 8 (see Fig. 1). = pens 3 & 9; �36 PEN 98 ISOLA710N / -/ ~------~---------------~ ,,----, FE.€OCR_ ,.•.••..••. '-l. WIt. ~--r--- (,0 eM .. Wltto aReA\{ r-~~ I ,,0, I i, ~------) A :. 3- SIOE'D B = mn e« ecceet: !4.l,{"r D~ R.l!...SOIl. ~ c.: Cu.TER. DRRk Co ,: Z. 2. m 1. 60 I L c. I. S I'f1 z. ~ I rn'L = 4.2. ml. e, z.: 2.0 rn" o= 4)IIUD - SIIA OE SHELTeJl. 0,; 1M)!) P1k"~'b SNoW =.s 1M2- Z"t./M'L 01.= 2..'''''1. rt. B/9f1 Fig. 2. :: ().IZO Location of artificial cover within cover isolation pens for 1984-85. �Colorado Division of Wildlife Wildlife Research Report July 1985 37 JOB PROGRESS REPORT State of Colorado Proj ect No. _0_1_-_0_3-_0_4_7 _ Mammals 1 Research Work Plan No. Deer Investigations Job. Winter Habitat Selection and Activity Patterns of Mule Deer in Front Range Shrubland and Forest Habitats 2 ---------------6 No. -------------------- Period Covered: Author: June 1,1984 - July 30,1985 R. C. Kufeld Personnel: L. Roberts, D. Schrupp ABSTRACT Triangulation data consisting of approximately 1,500 locations and 250 hrs of 24-hr activity data involving 13 radio-collared mule deer were collected during the November 12, 1984, through March, 1985, period. These data are currently undergoing analysis. Home ranges for 23 deer, which lived on !he study are for at least 1 and up to 3 yrs before their demise, averaged (x) 2.2 km2 in size and ranged from 1.2 to 3.2 km2• Two deer which migrated long distances during summer returned to the same home range during winter. The rest stayed in their home ranges year long. ��39 WINTER HABITAT SELECTION AND ACTIVITY PATTERN OF MULE DEER IN FRONT RANGE SHRUBLAND AND FOREST HABITATS Roland C. Kufeld P. N. OBJECTIVE 1. To test Telonics telemetry equipment to determine its accuracy at various distances in locating a transmitter on the Horsetooth Mountain study area and the ability of an observer using that equipment to correctly detect deer activity patterns by habitat type. 2. To determine habitat selection and activity pattern of mule deer within habitat types in the Horsetooth Mountain area during winter. SEGMENT OBJECTIVES 1. Honitor radio-collared mule deer to determine habitat selection and activity patterns throughout 24-hr periods. 2. Develop computer methodology for analysis of telemetry data on location and activities of mule deer. ACKNOWLEDGMENTS L. Roberts assisted in collection, tabulation and computerization of field data. D. Schrupp coordinated creation of a system for computerizing habitat and telemetry data. r~ETHODS AND MATERIALS Monitoring of Radio-Collared Deer Thirteen radio-collared adult female deer were periodically monitored from the same 2 triangulation points from November 12, 1984, through March, 1985, to determine habitat selection and activity. Equipment, procedures, and monitoring schedules were described by Kufeld (1981, 1982). The monitoring schedule included 3 sunrise, 3 daytime, 3 sunset, and 3 nighttime periods during November, January, February and t·1arch;and 2 of each period during December. Each period lasted 6 hrs. During the rest of the year, periodic checks using hand-held telemetry equipment were made at intervals of 1 week to 10 days to determine general loation of all radio-collared deer. �40 RESULTS AND DISCUSSION Telemetry data showing approximately 1,500 deer locations and 250 hrs of recorded deer activity involving 13 radio-collared deer were collected during the November 12, 1984, through March, 1985, period. These data, along with comparable data collected during the November-March period of 1982-83 and 1983-84, are currently undergoing analysis an will be presented in a future report. The following is a summary of information on movements of radio-collared deer at Lory State Park during the 3 yrs they were monitored. Only 4 of the 28 instrumented deer have left their home ranges since being intrumented. Two of these only went a short distance (less than 1.6 km). The rest appear to be year-around residents. The movement history of the 4 deer is as follows: 1. Deer #149.641 was instrumented in Lory State Park 1-19-82. It left its home range between 6-9-82 and 6-18-82. It was located 7-13-82 in a valley of about 32 ha, 3.2 km north of Pennock Pass, which is 29 km west of its home range. It spent the entire summer there, and was still there 10-25-82. On 11-1-82 it was back in its home range in Lory State Park. It remained in its home range until sometime between 8-28-83 and 9-21-83, when it disappeared. On 9-21-83 it was located in the valley described above. It was still in the valley on 10-26-83. On 11-14-83 it returned to its home range at Lory State Park and remained until its death by falling off a cliff on 4-5-84. 2. Deer #149.671 was instrumented in Lory State Park 1-27-82. It left its home range between 5-27-82 and 6-9-82. It was located in the valley 3.2 km north of Pennock Pass (see deer #149.641 above) on 7-13-82 where it spent the entire summer. It was still there 10-25-82. On 11-1-82 it was back in its home range in Lory State Park. It remained in its home range until sometime between 6-1- and 6-10-83 when it disappeared. It was located in the valley 3.2 km north of Pennock Pass 6-22-83 where it again spent the summer. It was still there on 10-26-83. This was 3 days before the 1983 separate deer season. It had not returned to Lory State Park as of 4-5-84. An aerial search on 1-10-84 of the entire Poudre River and Big Thompson drainages including the valley where it was located on 10-26-83 could find no trace of it. I suspect it was killed by a hunter during the 1983 season and the collar transported out of the area. *Deer #149.641 and 149.671 were almost constant companions when on the study area at Lory State Park. Although they left a week apart in the spring of 1982, they both migrated to the same valley, 3.2 km north of Pennock Pass where they spent the summer together. They returned together to Lory State Park between 10-25-82 and 11-1-82. The following year #149.671 migrated to the valley in June but #149.641 did not follow until September. Deer #149.641 subsequently returned to Lory State Park in November, 1983; but #149.671 may have been killed in the valley or enroute to Lory State Park. �41 3. Deer #149.921 was instrumented in Lory State Park 2-3-83. Between 6-10-83 and 6-20-83 it swam eastward across Horsetooth Reservoir and spent the rest of the winter in its regular home range in Lory State Park. 4. Deer #149.879 - At about 5:30 AM on November 13, 1984, 4 instrumented deer (#149.879, 149.732, 149.808, and 149.911) left their home ranges on the west side of the reservoir and, together, moved eastward across the north dam of Horsetooth Reservoir to an area immediately northeast of Horsetooth Reservoir. The next day, #149.732 and 149.808 had returned to their regular areas on the west side. Deer #149.879 stayed in the area northeast of the reservoir until late December, 1984, when it, too, returned to its regular area on the west side. Home ranges for 23 deer which lived on the study area for at least 1 and up to 3 yrs before their demise averaged (x) 2.2 km2 in size and ranged from 1.2 to 3.2 km 2. Those which migrated long distances during summer (#149.641 and 149.671), returned to the same home range during winter. The rest were nonmigratory and stayed in their home ranges year long. LITERATURE CITED Kufeld, R. C. 1981. Winter habitat selection and ativity patterns of mule deer in front range shrubland and forest habitats. Colo. Div. Wildl. Game Res. Rep. July (1):97-110. 1982. Winter habitat selection and activity patterns of mule deer in front range shrubland and forest habitats. Colo. Div. Wildl. Game Res. Rep. July (1):29-34. Prepared by ,,~:~(Ct/~vdC~ )d. l:lt{ Roland C. Kufe d Wildlife Researcher C ��Colorado Division of Wildlife Wildlife Research Report July 1985 43 JOB PROGRESS REPORT State of Colorado Project No. 01-03-047 ~'amma1s 1 Research Work Plan No. Deer Investigations -----------------2 ---------------- Job. No. 10 Period Covered: Author: July 1,1984 Mule Deer habitat Use in Piceance Basin - June 30,1985 R. M. Bartmann Personnel: A. W. Alldredge, L. H. Carpenter, J. Depperschmidt, D. A. Garrott, R. A. Garrott, J. Graham, D. Weybright, and G. C. White ABSTRACT Field work on the first phase of the cooperative mule deer study in Piceance Basin between Colorado State University and the Colorado Division of Wildlife was completed. Drafts of 2 manuscripts, 1 covering deer movements and the other deer mortality, are being prepared. The following is the abstract from the annual report prepared by CSU cooperators: In the fall of 1984, 60 fawns were radio collared on both the C-b Tract and Little Hills study areas, bringing the total instrumented population in Piceance Basin to 160 at the onset of winter. The 1arge data base collected during the previous 4 years of study coupled with budgetary constraints resulted in the termination of the movement studies in November, 1984. Deer show strong fide)ity to both their individual summer and winter ranges and appear to use the same migratory route to travel between the two. Timing of the fall migration remains consistent between years and is not stimulated by snow. storms. Spring migrations varied by as much as a month from year to year and appear to be correlated with winter severity. Adult doe winter survival was ~5% for the instrumented C-b Tract population and 100% for Little Hills animals. Fawn survival was 9% for the C-b Tract animals and 31% for Little Hills animals. The high fawn mortality for C-b Tract animals is attributed to an incease in predation rates from an average of 44% of the instrumented population during winters 1980-81 through 1983-84 to 77% this past winter. Both coyote and bobcats were responsible for the increased predation. For the third consecutive year at least 40-50% of the instrumented fawns in the Little Hills area succumbed to starvation. These relatively high losses even during Ifnorma11fwinters (1982-83 and 1984-85) suggest that adequate forage may not be available for the number of deer wintering on the Little Hills range. ��45 MULE DEER HABITAT USE IN PICEANCE BASIN Richard M. Bartmann P. N. OBJECTIVE See Segment Objective. SEGMENT OBJECTIVE Administer contract funding and cooperate in studies of mule deer seasonal movements, survival, and habitat use in proximity to oil shale development projects by means of radiotelemetry methods. RESULTS AND DISCUSSION Dr. Gary C. White, Principal Investigator for the Los Alamos National Laboratory, moved to Colorado State University (CSU). The contract with the Department of Energy for the cooperative mule deer study was then renewed for 1 year with CSU to allow completion of ongoing field studies. A report of work accomplished during 1984-85 was prepared by CSU personnel and is included as Appendix A. All data are presently being analyzed and manuscipts prepared for the seasonal movements and mortality aspects of the study. ;2:/Jlv,,p_ ~ Prepared by R1~harC1 r~. Bartmah~'::--b Wildlife Researcher C �• 46 APPENDIX A PICEANCE BASIN MULE DEER STUDY FISCAL YEAR 1985 REPORT Robert A. Garrott and Gary C. White Colorado State University Fort Collins, CO 80523 ABSTRACT In the fall of 1984 sixty fawns were radio collared on both the C-b Tract and Little Hills study areas, bringing the total instrumented population in Piceance Basin to 160 at the onset of winter. The large data base collected during the previous fo~r years of study coupled with budgetary constraints resulted in the termination of the movement dies in November 1984. stu- Deer show strong fidelity to both their indivi- dual summer and winter ranges and appear to use the same migratory route to travel between the two. Timing of the fall migration remains con- sistent between years and is not migrations stimulated by snow Spring varied by as much as a month from year to year and appear to be correlated with winter severity. instrumented animals. Fawn survival was 9% for the C-b Tract Hills animals. C-b Tract Adult doe winter survival for the Little storms. was 85% population and 100% for Little Hills animals and 31% for The high fawn mortality for C-b Tract animals is attributed to an increase in predation rates from an average of 44% of the instrumented population during winters 1980-81 through 1983-84 to 77% this past winter. Both coyote and bobcats were responsible for the increased For the third consecutive year at least 40-50% of predation. the instrumented fawns in the Little Hills area succumbed to starvation. These relatively high losses even during "normal" winters (1982-83 and �47 Mule Deer Study Progress Report 1985 1984-85} suggests that adequate forage may not be available for the number of deer wintering on the Little Hills range. This report summarizes the results of the fifth year of field on the Piceance Basin mule deer study. Administration of the project was transferred from Los Alamos National Laboratory University during the current to Colorado State 1984 and both R. A. Garrott and G. C. White are now affiliated with Colorado State University. ends work phase The past year of field work of studies and the results of this five year research effort will be compiled, analyzed, and interpreted in a report scheduled to be disseminated in January 1986. emphasize manipulative experiments on both Future research will the C-b Tract and Little Hills study areas. TRAPPING Mule deer were trapped on C-b Tract 26-30 November 1984 and on Little Hills area from 30 November to 5 December 1984. two deer were captured using (Table drop nets with Two hundred and fawns instrumented 1). Only fawns were instrumented this year as the movement stu- dies were concluded in November 1984. study 120 the During the five years of this a total of approximately 1100 deer have been captured with 528 of these animals instrumented (Table 2). Transmitters on many of the adult does collared last 12 months. of during the early years of this study expired during the This resulted in nearly a 50% reduction in the number instrumented adult does returning to the C-b Tract area in the fall. �· 48 Mule Deer Study Progress Report 1985 After the November-December mented population 1984 trapping operation Sex Age Female Male Female Male Fawn Fawn Adult Adult Tota 1 s Little Hills instru- and released on 2 study areas Trapped C-b Tract total in the Piceance Basin was 160. Table 1. Mule deer trapped, instrumented, in Piceance Basin during fall 1984. Area the Female Male Female Male Number of Deer Instrumented Recaptured 34 33 43 1 30 30 0 0 10 0 111 60 16 33 29 29 0 33 27 0 0 0 2 3 0 91 60 5 202 120 21 Fawn Fawn Adult Adult Totals Grand Totals 3 3 Table 2. Mule deer instrumented on 2 study areas in Piceance Basin as of 15 December of each year. Area Year Fawns C-b Tract 1980 1981 1982 1983 1984 Little Hills 1982 1983 1984 Totals Adult does Total 28 66 61 60 60 25 51 51 43 22 53 117 112 103 82 59 60 60 21 29 18 80 89 78 454 260 714 -- �49 Mule Deer Study Progress Report 1985 FALL MIGRATION Due to budgetary constraints no data were collected during the 1984 fall migration. A summary of the previous 3 years of data on the timing of the fall migration for C-b Tract animals is presented in Figure 1. Timing of the migration was relatively synchronous among individuals and remained consistent over the three years we monitored instrumented deer. Deer began migrating the last week of September with the peak of migra- tion occurring during mid-October. Little Hills deer were only followed during the fall 1983 migration and thus provided little additional data, however, the same general pattern was observed for these animals. deer biologist believe that fall migrations are stimulated by the arrival of the first winter storms in the high country. not Many These data do support this contention as deer from both study populations usually initiated migrating before winter storms had occurred. this difference The reason for in the movements of Piceance Basin mule deer is uncer- tain but we hypothesize that the instrumented deer may have developed tradition of a early fall migration in order to utilize the high quality forage available in the irrigated hay meadows present on the winter range. WINTER MOVEMENTS In late November 1984, after the fall migration completed, have been a flight was made to locate all instrumented deer. As in the p~st, all deer showed strong animal should returning to the fidelity same to wintering geographic winters. The winter movements of deer November flight, were area areas with each occupied during past not monitored after the however, incidental field observations indicated that �· 50 Mule Deer Study Progress Report 1985 PERCENT OF DEER MIGMTINC 1981 100 = N 80 27 1982 H = 52 60 H 40 1983 31 = 20 e- 0 I M I "" 'U0"' ~ (\l Q.. 101 ~ ""coI '"' U 0 ~ I I ""::> ""eoI (\l N ::> 0 :z: 'U0"' 'U0"' I/) t'" (\l eo "" N I In ..; 0 :z: WEEJ< Fig. 1. Timing Tract, 1981-83. of fall migration of instrumented deer from the C-b the movement patterns were similar to those Upon winters. observed during arrival on the winter range C-b Tract deer concentrated along the periphery of the irrigated hay meadows along These Piceance meadows were used extensively for two to four weeks. deer were scattered throughout the C-b Tract area where relatively previous sedentary until mid-winter. Creek. By December they remained In late January and February deer moved off the C-b Tract area onto predominately south facing slopes to the north and northwest. This movement is thought to be a response to accumulation of snow on the predominantly north facing slopes of C-b Tract area. the In late March and April these deer again moved back onto the C-b Tract area and concentrated along the irrigated meadows �51 Mule Deer Study Progress Report 1985 until they began spring migration. Some Little Hills deer showed simi- lar movements to the irrigated hay meadows along the White River and the Dry Fork of Piceance Creek in spring and fall. In addition, a few deer moved onto south facing slopes during the winter, however, most remained relatively sedentary throughout the winter. SPRING MIGRATION Deer were not monitored during the spring migration this year, however, data from 1981-84 have revealed that timing of spring migration may vary by as much as a month from one year to the next (Fig. 2). There appears to be a correlation between timing of migration and severity of winter with deer migrating later after more severe winters. these data we hypothesize migration, which places reverse the negative that further From in order for deer to initiate their energy demands on them, they must energy balance experienced during the winter and improve their physiological condition. This can only be accomplished by intake of high quality forage with deer in poorer condition needing a longer period of spring foraging then deer in better condition. Hence, after relatively mild winters deer reverse their negative energy balance quickly and migrate to summer range early, while spring foraging must winters This delay in During the winters of and 1983-84 when C-b Tract deer delayed their spring migration, snow pack on the summer range persisted after severe be extended, delaying migration. migration may also provide a secondary benefit. 1982-83 after longer into the the milder winters of 1980-81 and 1981-82. been delayed after the more severe winters, impeded movement. In addition, plant this spring than If migration had not snow pack may have phenology on the summer range �52 Mule Deer Study Progress Report 1985 would undoubtedly have been retarded at a time when energy demands on adult does were high due to the late stage of pregnancy. MORTALITY The number of previously of collared adult does surviving to the onset winter was 22 and 18 for the C-b Tract and Little Hills study areas, respectively. The collars of two C-b Tract does originally collared fawns dropped off the deer and three other does died (2 starvation predator kill) for a survival rate of 85%. Little placed on fawns in each area, radio and 1 does on the Hills area survived the winter for a 100% survival rate. 60 transmitters PERCENT All instrumented as signals Of the were lost OF DEER MIGRATING 80 1981 /\ N = 29 I \ I ' 60 / / \ / / . / '\ I . "'L /' "" .j -, __ - \ /;~':.... ..- ••• •••I co N I it') )0 ••• (t )0 )0 J: J: I it') I N. N )0 Ill: J: ••• =: ~ (t ~ -- ... _ _L~~_L 'If 0 I e (t J: ••• e '. " '-." \ e- ••• N ('I) \. \, , ..• """" 46 \ N 1984 34 = = ~_ ••• I N N ••• e 1983 39 " \ r- ('I) N \ \ \ /,i .,-_ .• _ _.:... ""." -----O~~--~~--~~--~~~~----~ ) '\ / ,../ '\ l ',' i .'/ \-""" -,0" = -7'"\ \ / . \ ,i-j'/--o-----'/./ :'~--~,'-..\ \ iI 20 \ \ J I 40 1982 N I Z ::::I "') Olf •••I co Z ::::I "') WEEJ< Fig. 2. Timing Tract, 1981-84. of the spring migration of instrumented deer on C-b �53 Mule Deer Study Progress Report 1985 for two animals from the C-b Tract area and 11 from area. the Little Hills We failed to contact these signals throughout the winter despite repeated aerial searches of the entire winter range. several of these The signals transmitters were abnormal just prior to permanently losing contact with them, indicating malfunctioning circuits. tion from In addi- to these transmitter malfunctions, information on a third instru- mented fawn on the C-b Tract was lost when the collar was entangled in a fence and broke off the animal. Of the remaining 57 fawns from the C-b Tract area, 52 died during the winter for a 9% survival rate and 34 the of remaining 49 instrumented fawns from the Little Hills area died for a 31% survival rate. Although these survival rates generally fell within those recorded the range since the inception of these studies in 1980 (Table 3), several interesting discrepancies with previous data are apparent. Little Hills study perturbations C-b Tract This from oil shale development. rates. time. perturbations two areas must track and fawn overwinter survival rates were similar. vival rates over two winters of dissimilar fawn areas this year. on each other Data from the previous two winters supported the conten- tion that Little Hills was an adequate control for C-b However, on In order for Little Hills to be a valid control for weather, survival rates on the through expected control was needed to separate the effects of winter severity and oil shale survival The area was incorporated into our studies during the winter of 1982-83 to serve as a control for deer of survival rates Tract as adult The agreement of sur- severity was encouraging. were markedly different between the two The reason for this dissimilarity is uncertain, but �, S4 Hule Deer Study Progress Report 1985 causes us to re-evaluate the applicability of the control-treatment experimental design. Causes of fawn mortality followed the same pattern documented ing previous years. dur- The major cause of mortality on the C-b Tract area was predation, while starvation (winter kill) was the prominent cause of mortality for Little Hills fawns (Table 4). There was, however, a change in the proportion of fawns killed by predators on the area (f<O.Ol). From C-b Tract 1980-81 to 1983-84 documented predator kills for the C-b Tract fawns ranged from 44-48% of the total instrumented population but during the winter of 1984-85 77% of the instrumented fawns were killed by predators. severe the Although increased this past winter was not abnormally mortality from predators resulted in a fawn sur- vival rate almost as low as that documented for the winter of 1983-84 when many animals starved to death. extremely harsh Although coyotes, the major predator, killed more fawns this past winter, bobcat predation also increased. Table 3. Survival rates of instrumented mule deer on 2 study Piceance Basin. Area C-b Tract Little Hills Fawns Adult does 0.22 0.35 0.38 0.05 0.09 0.92 0.90 0.84 0.85 0.85 1982-:-83 0.36 1983-84 0.05 1984-85 0.31 0.90 0.81 1.00 Year 1980-81 1981-82 1982-83 1983-:-84 1984-85 areas in �55 Mule Deer Study Progress Report 1985 Table 4. Causes of winter fawn mortality on 2 study areas in Piceance Basin. Numbers in parenthesis are percentages of instrumented population. Mortality category C-b Tract Little Hills Total Predator Winter kill Road kill Undetermined Totals 44(77) 5( 9) 2( 4) 1( 1) 52(91) 7(14) 24(49) 1( 2) 2( 4) 34(69) 51(48) 29(27) 3( 3) 3( 3) 86(81) Mortality rates and causes for Little Hills fawns during the winter of 1984-85 were similar to the winter of 1982-83 with starvation the major cause of mortality (Table 4). progressed during Mortality increased as the winter both winters with the largest number of animals suc- cumbing in March (Fig. 3). During the winter of 1983-84 most fawns died in January However, we feel such early winter mortality and February. is abnormal and was a result of the onset conditions in December. Although of extremely severe winter we have no quantitative measure of coyote populations on either study area, the lack of heavy predation pressure on Little Hills fawns when compared to C-b Tract fawns is probably a reflection of lower coyote densities in the The 40-50% starvation relatively normal winters rate Little Hills area. for fawns in the Little Hills area during may indicate that these animals are not obtaining adequate forage during the winter. ROAD REFLECTOR EXPERIMENT The experiment to test the effectiveness of the Swareflex reflector was continued Piceance Creek Road. nate weeks along four 1.6-km wildlife (1 mile) sections of the Two sections of reflectors were covered on alter- from October through Hay when deer were on the winter range �- 56 Mule Deer Study Progress Report 1985 NUMBER OF DEER 20 r01Tl'act e-x ~N = 52 P.77l Li ttle Hi lIs 34 tLLJN 16 = 12 8 4 Feh Jan Dec Mal' MONTH Fig. 3. Timing of instrumented fawn mortalities from 2 study areas in Piceance Basin, 1984-85. and the number of road-killed deer found on each section additional of 1984-85. tions, 24 ten An deer were killed on the test sections during the winter To date, 41 deer have been killed on the four test sec- (60%) when the reflectors were covered and 17 (40%) when the reflectors were operational. the recorded. effectiveness of These data are insufficient the reflectors to determine as 95 animals must be killed in order to achieve the power needed to test the hypothesis at a meaningful level. At the present rate of road kills the experiment would have to continue for 4-5 more years. HABITAT STUDIES �57 Mule Deer. Study Progress Report 1985 The third year of field work has been initiated on the summer habitat study which is being conducted on the Roan Plateau in the vicinity of the Colony Shale Oil Project. estimate summer-long habitat Objectives of the study are 1) to selection and activity patterns of adult female mule deer over 24-hour periods and 2) to test for the effects construction disturbance on activity of and movement patterns of adult female mule deer. Permanent 4-tower telemetry systems were two during the summer of 1983 and 18 adult does were cap- study areas tured and instrumented. extensive test of During the spring the accuracy From June through September 1984 tracked for tions. following year on an of the triangulation system was con- ducted. approximately of the erected the instrumented deer were 580 hours generating over 5000 deer loca- An additional two adult does from the study areas were also cap- tured in drive nets and instrumented. Efforts during the summer of 1985 will concentrate on collecting additional activities of the instrumented data on the movements and deer and exploring the possibility of modifying the investigation from a descriptive to an experimental study. Two papers dealing with biotelemetry accuracy and triangulation are currently being prepared for publication (see project publications). Accuracy data indicates that the telemetry towers are capable of obtai~ing extremely accurate and precise bearings fr.omtransmitters, however, when transmitters are not line-of-sight with the receiving tower signals may be reflected "bounce" signals causing are erroneous incorporated bearing estimates. When such into triangulation procedures, the resulting location estimates are inaccurate. Quantitative techniques to identify poor location estimates have been developed so that these estimates can be removed from the movement data before analysis of habitat �• 58 Mule Deer Study Progress Report 1985 selection. ACKNOWLEDGMENTS We gratefully acknowledge the encouragement, advice, tion of and coopera- personnel from the Bureau of Land Management, Cathedral Bluffs Shale Oil Co., Colorado Division of Wildlife, Colorado State University, Exxon Company USA, Los Alamos National Laboratory, Minerals Management Office, Rio Blanco Oil Shale Co., and many support was provided by local ranchers. Financial the U. S. Department of Energy, contract W- 4305-36 to Los Alamos National Laboratory and FG02-85ER60297 to Colorado State University. Supplementary financial support was provided by Cathedral Bluffs Shale Oil Co., Colorado Division of Wildlife, and Exxon Company USA. RECENT PROJECT PUBLICATIONS Garrott, R. A. and R. M. Bartmann. 1984. Eval uat ion of implants for mule deer. J. Wildl. Manage. 48:646-648. Garrott, R. A., R. M. Bartmann, and G. C. White. 1985. radio-transmitter packages relative to deer fawn Wildl. Manage. 49:758-759. vaginal Comparison of mortal ity. J. Garrott, R. A. and R. W. Hayes. 1984. A radio-controlled triggering traps. Wildl. Soc. Bull. 12:320-322. device for Garrott, R. A., G. C. White, R. M. Bartmann, and D. M. Weybright .. 1986. Bearing accuracy and location estimation with three-tower triangualtion systems. J. Wildl. Manage. 50:. Submitted. Lee, J. E., G. C. White, R. A. Garrott, R. M. Bartmann, and A. W. Alldredge. 1985. Assessing the accuracy of a radiotelemetry system for estimating mule deer locations. J. Wildl. Manage. 49:658-663. �59 Mule Deer Study Progress Report 1985 White, G. C. 1985. Optimal locations of towers for triangulation studies using biotelemetry. J. Wildl. Manage. 49:190-196. White, G. C. and R. A. Garrott. 1984. Portable computer system for field processing biotelemetry triangulation data. Colorado Div. of Wildl. Game Information Leaflet 110:1-4. i 1986. Effects of biotelemetry White, G. C. and R. A. Garrott. triangulation error on detecting habitat selection. J. Wildl. Manage.:. Submitted. ��Colorado Division of Wildlife Wildlife Research Report July 1985 61 JOB PROGRESS REPORT State of Colorado Project No. 01-03-047 -------------------- Work P1an No. 2 ----------------- Job. No. 11 Period Covered: Author: Mammals 1 Research Deer Investigations Testing of Mule Deer Census ~1ethodology July 1,1984 - June 30,1985 R. t,1. Bartmann Pesonnel: A. W. Alldredge, D. C. Bowden, L. H. Carpenter, D. A. Garrott, R. A. Garrott, J. Graham, D. Weybright, G. C. White, and numerous CSU student volunteers. ABSTRACT A manuscript entitled IIAccuracy of Helicopter Counts of Mule Deer in PinyonJuniper Woodland was submitted to the Wildlife Society Bulletin for review. The abstract is as follows: Accuracy of aerial mule deer (Odocoileus hemionus hemionus) counts from a helicopter was evaluated on plnyon-Juniper {Plnus edulis-Juniperus osteosterma} winter range in Piceance Basin, Colorado. Deer were radio-col ared and stocked in 4, 58-70-ha pastures at densities ranging from 9-46/km2. Aerial count accuracy ranged from 35-86% in individual pastures and from 59-68% for 5 evaluation periods over 2 winters. Differences in mean proportions of deer counted were not significant (P > O.05) between morning and afternoon counts, deer densities, quadrat sTzes, or oservers, but significance (P < 0.05) was found with search direction. There were indications of a conditioning response by deer to repeated helicopter pverflights, but any effects on results seemed small. Under optimal conditions, 2/3 seems like the maximum proportion of mule deer that can be seen from a helicopter on pinyon-juniper winter range. A second manuscript entitled IIAerial Mark-Recapture Estimates of Confined Mule Deer in Pinyon-Juniper Wood1and was nearly completed. The abstract of that manuscript is as follows: Four 58-70-ha pastures with known numbers of marked (radio-collared) and reasonably well known numbers of unmarked mule deer (Odocoi1eus hemionus hemionus) were used to calculate 114 Linco1nPetersen estlmates of population size, with resightings obtained from a helicopter. Three approaches to combining population estimates for a pasture (mean, median, and hypergeometric maximum likelihood) were explored. The median is least sensitive to outliers, but the hypergeometric maximum likelihood provides ~40% smaller confidence intervals. About the same percentage of confidence intervals for all 3 estimators (64-73%) overlapped at least part of the ranges of true population size. For all 3 estimators, a large proportion (>45%) of the population should be marked to obtain more reliable estimates and greatest confidence interval coverage. However, there is still high probability that mean population estimates will be low. ll ll ��63 TESTING OF MULE DEER CENSUS METHODOLOGY Richard M. Bartmann P. N. OBJECTIVES 1. Test for sex bias in aerial deer counts. 2. Test for effects of observers in numbers of deer counted. SEGMENT OBJECTIVES Same as P. N. Objectives. RESULTS AND DISCUSSION A manuscript entitled "Accuracy of Helicopter Counts of Mule deer in PinyonJuniper Woodland" was submitted to the Wildlife Society Bulletin for review. A second manuscript, entitled "Aerial Mark-Recapture Estimates of Confined Mule deer in Pinyon-Juniper Woodland" was ne~rly completed. /IJvy 4. Prepared by ~J ~~/ . / Rl card tt Bartmann-Wildlife Researcher C -=-=. ~J,.___~ ��Colorado Division of Wildlife Wildlife Research Report July 1985 65 JOB PROGRESS REPORT State of Colorado Project. No. 01-03-047 Mammals 1 Research Work Plan Elk Investigations -----------------No. 3 ---------------- Job. No. 2 Period Covered: Author: July 1,1984 Elk Population and Ecology Study - June 30,1985 G. D. Bear ABSTRACT A total of 139 elk (68 cows, 43 calves, and 28 bulls) was captured with Clover traps and marked on the North Park study area from December 18, 1984, to March 7, 1985. Mean population estimate for 3 aerial surveys using the mark-recapture technique was 3,543 + 231 elk. A suitable test of accuracy was not found for the method. A total of 3,763 + 1,015 elk was estimated using the variable-sited quadrat method. There was a problem recognizing and distinguishing collars in both methods. Migration patterns for 45 elk during December 1983, through February, 1984, are described. Twenty-thee percent (18) of ~O radlo-collared elk were harvested during the fall hunting season. ��67 ELK POPULATION AND ECOLOGY STUDY George D. Bear P. N. OBJECTIVE Develop techniques to more accurately and precisely estimate elk population levels. SEGMENT OBJECTIVES 1. Capture and mark approximately 250-300 elk in the North Park study area. 2. Conduct aerial surveys and estimate population densities. 3. ~10nitor movements of telemetered elk. METHODS AND MATERIALS For a complete description of methods, refer to the Program Narrative (Bear, 1983; DOW files). Following is a brief description of methodology used in the variable-quadrat counts. This information is provided since it was additional to previous work. Mark-Recapture Experiment The North Park elk winter range was divided into 28 sampling units (5 high, 3 moderate, and 20 low) based on expected elk densities. Boundaries of these units were based on observations during the past year and from talking to local District Wildlife Managers. All high and moderate density sampling units were surveyed but only 3 low density sample units in the northern portion of North Park and 3 low density sample units in the southern portion of North Park were flown. These 6 low density sample units were randomly selected before surveys were flown. Three separate counts were flown on all sample units. Seventy-five radio collared elk were used to obtain a sightability correction factor. This factor was used to project number of elk observed on each survey to total elk. Before each survey it was necessary to determine by radio telemetry, location of each collared elk and record the sample unit where it was located. Aerial flights were conducted from a helicopter during March, 1985. Flights were conducted in early morning (within 3 hrs of sunrise) when light conditions were acceptable for sighting elk and while elk were still active in open areas. The entire sampling unit was flown using general features of the terrain to divide the unit into counting strips. An effort was made to obtain "maximum" counts, yet avoid duplication. The population estimate and its variance were obtained by estimating the number of elk and variance for each stratum and then summing all strata estimates to arrive at the total for the survey area (Gasaway et ale 1981). �68 1. The following symbols were used in the calculation of each individual stratum population estunate and variance. = A total surface area (mi2 or kms) in a particular stratum Yi = number of elk in the ith sample unit xi = number of mi2 (kms) in the ith sample unit x = mean size of all sample units surveyed in a particular stratum = N total number of sample units in a particular stratum T = total population estimate for a particular stratum 2. The following calculations were performed for each stratum: a. The density of elk for each stratum (r) was the number of elk per mi2 (km). n total no. of elk observed in all L y. sample units that were surveyed = i = 1 1 t-o~ta~'~s~u~rf~a~c~e~ar~e~a~o~f~a~'~'~sa~m~p~'~e~u~n~i~t-s n (mF) (km2) that were surveyed L X· i = 1 1 b. The population estimate for each straum. T = density of elk x total surface area of the stratum or T" c. = r . A Variance [V(T)] for the stratum population estimate • V(T) = . [ A2 1 --:::-z . x 3. Total population estimate ~ ~ Tt = where h = ~ 5 n = strata population estimates A Th + Tm + T£ (rh • Ah) + (rm . Am) + (r£ . A£) high densi ty stratum, m = medium, and £ = low. = �69 4. Variance of the = L variance of the strata population estimate population estimates. ~ A V(Tt) = V(Th) A A + V(Tm) + V(T~) = + [A2~• V(r ~)] 5. Calculation of the confidence interval (CI) for the population estimate of the survey area. C1 = total population estimate ~ (t y) a 6. V variance of the total population estimate Correction of the estimate for sightabi1ity was calculated by dividing number of collared elk known to be in the survey area by number of collared elk observed. This number was then multiplied by the population estimate and confidence interval. RESULTS AND DISCUSSION A total of 139 elk (68 cows, 43 calves, and 28 bulls) was captured in Clover traps and marked at 7 different locations (Appendix A). This was below the predetennined goal of 250-300 elk to be marked because the 1984-85 winter was considerably milder than nonna1 with little snow cover to aid trapping. Elk were scattered throughout the winter range and showed minimal interest in the alfalfa hay used as bait.for the traps. Trapping was more successful on the southern half of the study area where the elk were more concentrated. Forty-five elk were fitted with telemetry collars during 1984-85. These elk combined with 30 functioning collars from 1983-84, gave a total of 75 radiocollared elk. Counting all collars, there are now 168 marked elk in the population. Three aerial surveys were conducted over the entire study area and separate population estimates derived from the ratio of marked to unmarked elk (Table 1). Population estimates for the 3 surveys were 3,810,3,412, and 3,407, respectively, with a mean of 3,543 + 231 elk, for a precision level of 6.5%. This small variance was surprising Tn that the desired number of marked animals was not achieved and considering the poor distribution of marked animals in the northern half of the study area. Perhaps the random observation of elk on the aerial surveys compensated for the poor distribution of marked elk in the population. Precision of the estimate was better for the 3 surveys in the south-half of the study area. Here approximately 9% of the estimated population was marked, and the estimates was 1,163 + 45 elk. In the north half where approximately 3% of the estimated population was marked, the estimate was 2,181 ! 216 elk (Tables 2 and 3). . �70 Efforts were made to determine accuracy of the mark-recapture estimate. This test was to be done by marking a subpopulation within the overall marked population by using nyanzol dye to make numbers on the backs of collared elk. This approach proved unsatisfactory. Numbers were easily read when elk were found in open areas; however, they most often could not be identified on elk found in timbered areas. A second approach to measure accuracy was to regard radio-collared elk as a subpopu1ation within the 168 marked elk. This yielded an estimate of 230 ~ 5 elk for the 3 surveys {Table 4}, a 37% error from the known population of 168 marked animals. However, this test is suspect since on several occasions the record for the dyed identification numbers indicated a radio-collared elk; yet at the time of the observation, the radio collar was not seen. This would bias the estimate upward. Since identification numbers could not be recorded for all marked elk, it is impossible to determine a correction factor for radio-collared elk observed but not recognized. It is important to continue this experiment another year to better assess accuracy and precision of the estimate. Precision appears to be good, however, an intensive effort should be made to mark at least 10-15% of the estimated population. In testing for accuracy the marked animals must be readily identified. It is hoped that this can be accomplished using brightly colored collars. Variable-sized Quadrats Three aerial surveys were conducted on the variable sized-quadrats. Population estimate derived from these flights were: 2,600, 4,219, and 4,469 with a mean of 3,763 + 1,015 elk for the North Park study area {Table 5}. Projections for the south half of the study area were: 842,1,667, and 1,100 with a mean of 1,203 + 422 elk. Estimates for the north half were: 1,767, 2,425, and 3,518 with-a mean of 2,570 ~ 884 elk {Tables 6 and 7}. Precision of the variable-sized quadrat method was large. This technique is, in essence, a variation of the mark-recapture technique but with fewer marked animals {78 radio collars or 2% of the estimated population}. Advantages of the variable-quadrat technique are less flight time and fewer animals to be trapped and marked. However, the disadvantages are the radio collars are expensive and precision is less acceptable at this low level of sampling. If the sample size was increased, the increased cost in radio collars and trapping expense would make it comparable in cost to the mark-recapture technique. It is undesirable at this time to continue evaluating the variable-sized quadrat technique on this study area. When and if the present radio collars are retrieved, reconstructed and color marked so they are more visible, perhaps additional tests of this method will be warranted. Trend Count Three population estimates were derived for the North Park study area using 3 different procedures. All 3 techniques yielded similar estimates. Personnel from the Northeast Region of the DOW conducted aerial trend-count in North Park and observed 1,828 elk. Using this information and the projected hunter harvest data in a repeated IItria1 and error IIprocess, they concluded there were 3,400 elk in North Park. The mark-recapture and variable-sized quadrat �71 techniques produced estimates of 3,543 + 231 and 3,763 + 1,015 elk, respectively. This similarity of these estimates is interestlng. The process needs to be repeated to provide a better evaluation of all methods. The mark-recapture and the variable-sized quadrat methods are more desirable in that they provide estimates of precision which can be used to compute confidence limits for the mean estimates. The trend count method provides no measure of variation. However, the mark-recapture and variable-sized quadrat approaches are also more costly. ~1igrati on Fifteen elk (13 cows, 2 bulls) were trapped on Owl Ridge and marked with radio collars. Trapping stations were located on upper Owl Ridge and Owl Creek (Fig. 1). As snow depths increased in late winter, many of the marked elk moved from Owl Ridge towards the Arapaho National Wildlife Refuge. As spring progressed, these elk reversed their migration back towards Owl Mountain. A few individuals moved up Owl Creek to the logged areas on the northern slopes of Owl Mountain for the summer. Most of the marked elk migrated around the south side of Owl Mountain to Taylor draw and the Elk Creek area. In late summer they moved to the alpine ranges at Bear Paws and Nount Cindy (Table 8). In late fall and early winter this migration was reversed and marked elk returned to the Owl Mountain area. Five cow elk were trapped and marked with radio collars on Buffalo Ridge in southern North Park. These 5 elk remained on the Buffalo Ridge area until spring, then migrated to higher elevations along Green Ridge (Table 9, Fig. 2). One cow moved eastward to the logged areas on the northern exposure of Parkview ~lountain for the summer. The other 4 elk migrated to the logged areas at Grassy Run for a brief time, then they moved to the alpine ranges along Arapaho Ridge. In the fall this migration pattern was reversed, with elk arriving back on the Buffalo Ridge area by early winter. Eight elk (5 cows, 3 bulls) were trapped and radio collared on the Sentinel Mountain in northeastern North Park. One of the bulls died of unknown causes shortly after release. As winter progressed, 2 cows migrated northward to winter on lower Camp Creek, then moved back to the Sand Hills area in the spring (Table 10, Fig. 3). The other 5 remained in the Sentinel Mountain area, then moved to the Sand Hills area as spring progressed. One bull moved westward across North Park into the ~·10untZirkel area then south to Rabbit Ears Pass, where he was shot in the fall hunting season. The 4 remaining elk used the higher elevations in the Rawah Range as far south as Montgomery Pass throughout the summer and fall. By late winter they had returned to the Sand Hills and Sentinel Mountain areas. Fifteen elk (12 cows, 3 bulls) and 3 elk (2 cows, 1 bull) were trapped and fitted with radio collars while on Independence Mountain and the Delaney Buttes area, respecti ve1y. E1k co11 ared on Independence ~~ountain generally moved to Fischer Peak with a few moving to the north side of Independence ~10untain into the sagebrush. As winter became more severe (Fig. 4, Tables 11 and 12), these elk returned to the southern exposures of Independence t~ountain and the sagebrush areas near Delaney Buttes as spring progressed. �72 Elk marked on the Swift Ranch near Delaney Buttes remained in the sagebrush areas of the ranch until spring. They then joined a group of elk from Independence Mountain and moved westerly to the foothills along the west side of North Park. These elk migrated westward during the sunvner to alpine ranges in and near the Mount Zirkel Wilderness area. Eleven of the 18 collared elk moved up the Newcomb Creek and Lone Pine Creek drainages. In late fall and early winter, most collared elk returned to the Delaney Buttes and Independence Mountain winter ranges. Two of these cows moved to the south end of North Park; one remained on Buffalo Peak, the other was shot on Indian Creek in the fall. Also, two cows moved over the Rabbit Ears Pass to winter range near Steamboat Springs. Mortal ity Twenty-three percent of the 80 elk trapped and marked with radio collars durin~ the 1983-84 winter were harvested during the 1984 hunting season. One cow dled from a gunshot wound suffered during the early muzzle-loading season, and 1 bull died from unknown causes while still on the winter range. LITERATURE CITED Gasaway, W. C., S. DuBois, and S. Harbo. 1981. Estimating moose abundance and composition. Annual Report. Dept. Fish and Wi1d1., Univ. Alaska. 62pp. Prepared by �73 Table 1. Total number of elk, number of marked elk, and projected population estimate for the North Park Study Area as determined from 3 aerial surveys during March, 1985. Population Elk Co11 ars estimate Survey observed observed 1 2 3 Mean Std. Oev. 1,202 914 1 ,866 53 45 92 3,810 3,412 3,407 3,543 231 168 collared elk in population Table 2. Total number of elk, number of marked elk, and projected population estimate for the south-half of the North Park Study Area as determined from 3 aerial surveys during March, 1985. Survey 2 3 Elk observed 299 270 631 Co 11ars observed Population estimate 26 22 1 ,139 1 ,215 55 1,135 1 ,163 Mean Std. Oev. 45 Table 3. Total number of elk, number of marked elk, and projected population estimate for the north-half of the North Park Study Area as determined from 3 aerial surveys during March, 1985. Population Co 11ars El k estimate Survey observed observed 2 3 Mean Std. Oev. 903 644 1,235 27 23 37 2,308 1,932 2,303 2,181 216 �74 Table 4. Results of test using number of radio collars observed to estimate total marked in the population. True number of radio collars 75 and total collars = 168. Radio collars observed Survey 2 3 Mean Stud. Dev. Co 11ared elk observed Number 53 45 92 17 15 30 Percent 23 20 40 Marked population estimate 234 225 230 230 5 = �Table 5. Population estimates for the entire North Park Study Area projected from the variable-sized quadrat surveys. Elk density Elk observations by strata (e1k/mF [km2 J) Population Projected Correction estimate Survey H-M Low tota 1 factora Low H-M 2 3 1 ,150 1 ,165 1,432 34 101 48 17.3 [6.7J 19.3 [7.5J 24.9 [9.6J 1.0 [0.4J 2.6 [1.0J 2.5 [1.0J 1 ,215 1,348 1,552 2.14 3.13 2.88 Mean Std. Oev. 2,600 4,219 4,469 3,763 1 ,015 aCorrection factor is sightabi1ity index as determined from number of known radio collars observed each flight. Table 6. Population estimates for the north-half of the North Park Study Area projected from variable-sized quadrat surveys. Elk density Elk observations by strata (e1k/mF[km2J) Population Projected Correction estima te Low Low Survey H-M H-M total f'ac tors ' 2 3 Mean Std. Oev. 654 904 926 32 78 31 15.0 [5.8J 24.4 [9.4J 29.5[11. 4) 1.7 [0.7J 2.4 [0.9J 2.4 [0.9J 716 1 ,041 1,005 2.63 2.33 3.50 1 ,767 2,425 3,518 2,570 884 aCorrection factor is sightabi1ity index as determined from number of known radio collars observed each flight. -..J Ln �Table 7. Population estimates for the south-half of the North Park Study Area projected from the variable-sized quadrat surveys. Elk observations Elk density by strata (elk/mi2[km2J) Population Projected Correction estirnate Low Low total Survey H-M factor" H-M 2 3 Mean Std. Dev. 484 261 430 0 14 17 22.0 [8.5J 11.1 [4.3J 18.4 [7.1J 0.0 [O.OJ 3.2 [1.2J 2.8 [1.1J 484 303 466 1.74 5.50 2.36 842 1 ,667 1,100 1,203 422 aCorrection factor is sightability index as determined from number of known collars observed each flight. -...J 0\ �Table 8. Summary of movement eatterns for elk radio-collared on Owl Ridge, North Park. Winter - Spring Summer - Fall Wi nter Telemetry collar Location Location Location Date number Sex Date Date 20 Cow Jan/May Owl Ridge 40 Bull Feb/Apr 60 Cow Jan/May Baller RanchBuffalo Ridge Owl Ridge 140 Cow 190 Cow Jan/Apr May Jan/Apr May Owl Ridge NW Owl Mtn. Owl Ridge NW Owl Mtn. 240 Bull Jan/May Owl Ridge 350 Cow Jan/Apr Owl Ridge 400 Cow Jan/May Owl Ridge 510 Cow Jan/May Owl Ridge Jun Aug Oct/Nov May/Oct S Owl Mtn. Bear Paws W Owl Mtn. S Owl Mtn. Jun Oct Nov Nov Jul/Sept Oct Jul Aug/Oct Nov Jun Jul Aug/Oct Oct Jun/Jul Aug Sept/Oct Oct Jun/Sept Oct Jun/ Jul Aug/Sept Oct/Nov S Owl Mtn. Parkview S Owl Mtn. Shot by hunter Bear Paws SW Owl Mtn. NE Owl Mtn. W Owl Mtn. Shot by hunter Owl Mtn. Bear Paws SW Owl Mtn. Shot by hunter SE Owl Mtn. Bear Paws Owl Mtn. Shot by hunter N Owl Mtn. NW Owl Mtn. SE Owl Mtn. Bear Paws SW Owl Mtn. Jan/Mar Owl Ridge Jan Owl Ridge Jan/Mar SW Owl Mtn. Jan/Mar Owl Ridge Jan/Mar SW Owl Mtn. '-J '-J �Table 8. (Continued) -...J 520 Cow Jan/Apr May Owl Ridge SW Owl Mtn. 530 540 Cow Cow Jan/May Jan/Apr May Owl Ridge Owl Ridge SW Owl Mtn. 560 Cow Jan/Apr Owl Ridge 570 Cow Jan/May Owl Ridge Jul Aug/Sept Oct/Nov Jun/Oct Jun/Jul Aug Sept/Oct Nov May/Oct Nov Jun Aug Oct Nov NW Owl Mtn. Bear Paws N Owl Mtn. Owl Mtn. S Owl Mtn. Bear Paws S Owl Mtn. Shot by hunter SE Owl Mtn. SW Owl Mtn. NW Owl Mtn. Bear Paws SW Owl Mtn. Shot by hunter Jan/Mar Owl Ridge Mar Owl Ridge Jan/Mar SW Owl Mtn. ex> �Table 9. Summary of movement ~atterns for elk radio-collared on Buffalo Ridge, North Park. Telemetry Winter - Spring Summer - Fall Winter co 11ar number Sex Date Location Date Location Location Date 50 Cow Jan/May Buffalo Ridge 210 Cow Jan/Apr Buffalo Ridge 450 Cow Jan/May Buffalo RidgeBuffalo Cr. 550 Cow Jan/May Buffalo Ridge 580 Cow Jan/May Buffalo Ridge Jul Aug Sept Nov Jun Aug Oct Nov Jun Jul/Aug Oct/Nov Jul/Sept Nov Jul/Aug Oct/Nov Green Ridge Grassy Run Buffalo Cr. Buffalo Pk. Lower Willow Cr. Parkview Upper Willow Cr. Green Ridge Green Ridge Sheep Mtn. Grassy Run Arapaho Ridge Buffalo Cr. Arapaho Ridge Buffalo Ridge Jan/Mar Buffalo Ridge Jan/Mar Buffalo Ridge Jan/Mar Buffalo Ridge Jan/Mar Buffalo Ridge Jan/Mar Buffalo Ridge --.I \,Q �Table 10. Summar~ of movement ~atterns for elk radio-collared in Sentinel Mountain, North Park. Telemetry Winter Wi nter - Spri ng Summer - Fall collar Date Location Location Location number Sex Date Date 100 Cow 200 Cow Jan/May Jun Jan/Apr Bull May/Jun Jan/Apr 220 May/Jun 260 Bull Jan/May 270 Bull Jan/Feb 280 Cow Jan/May 430 Cow Jan/Apr Cow May Jan/Apr 550 May Sentinel Mtn. E Sand Hills Sentinel Mtn. Kings Canyon N Sand Hills Sentinel Mtn.Camp Cr. N Sand Hi 11sE Sand Hills Sentinel Mtn.Kings CanyonCamp Cr. Sentinel Mtn. (died unknown causes) Sentinel Mtn.Camp Cr. Sentinel Mtn.Camp Cr. N Sand Hills Sentinel Mtn.Kings Canyon N Sand Hills Sept/Oct Oct/Nov Aug/Nov Montgomery Pass E Sand Hills Shipman Mtn. Sept Swamp Park (NE Steamboat Sprgs.) Shot by hunter Rabbit Hills Pass Sh ipman Park Oct Aug/Sept Jun/Nov Jun/Oct Jun/Nov E Sand Hill sShipman Park E Sand Hi11sShipman Park E Sand HillsShipman Park Jan/Mar Camp Cr. Jan/Mar N Sand Hills Jan N Sand Hills Jan N Sand Hills Jan/Mar N Sand Hi 11sSentinel Mtn. (X) 0 �Table 11. Summar~ of movement ~atterns for elk radio-collared on Inde~endence Mountain, North Park. Winter - Spring Summer - Fa11 Winter Telemetry coll ar Date Location Location Location Date Date number Sex 30 70 Bull Cow Jan/May Jan/Apr May/Jun 90 110 130 Bull Jan/May Cow Jan/Mar Cow Apr May Jan/Apr May 150 Cow Jan/Apr Independence Mtn~ Jun Sept/Nov Independence Mtn.- Jul/Oct Fischer Pk. Nov Lower Roaring Fk.Newcomb Cr. Independence Mtn. Jul Independence Mtn.Fischer Pk. Boettcher Ridge Lower Lone Pine Independence Mtn.Sagebrush Flats Boettcher Ridge Independence Mtn.Sagebrush Flats Aug/Oct Oct Jun/Sept Oct/Nov Jun/Oct Nov May/Aug Nov 180 Cow Jan/Apr May Independence Mtn.Fischer Pk. Indian Cr. Jun/Nov Lower Lone Pine W Big Cr. Lakes Upper Newcomb Cr. Lower Roaring Fk.Delaney Butte Illinois Cr. (E of Rand) Owl Mtn. Shot by hunter Upper Newcomb Cr. Lower Lone Pine Lower Newcomb Cr.Upper Newcomb Cr. Lower Newcomb Cr.Lower Lone Pine Lower to Upper Newcomb Cr. 10 mi. NE Steamboat Sprgs. Buffalo Pk.Buffalo Cr. Mar Jan/Mar NW Independence (Wyo. ) Delaney ButteFischer Pk. Jan Delaney ButteFischer Pk. Jan/Mar Delaney ButteFischer Pk. Jan/Apr Buffalo Cr. ,_. 00 �Table 11. (Continued) Cow Jan/Mar 300 Mar/May 00 Independence Mtn.Fischer Pk. Boettcher Ridge Jun/Aug Oct/Nov 320 Cow Jan/Apr Independent Mtn.Fischer Pk. Jul/Aug Oct 330 Bull Jan/Jun Independence Mtn. Jul/Nov 370 Cow Jan/Apr Independence Mtn.Fischer Pk. Jun Oct/Nov 390 Cow Jan/Jun 410 Cow Jan/May Independence Mtn.Fischer Pk. Independence Mtn.Fischer Pk. 420 Cow Jan/May 9.420 Cow Jan/Apr Jun/Jul Aug Jan Aug/Oct Nov Aug Nov Nov Independence Mtn.Fischer Pk. Independence Mtn.Fischer Pk. Lower to Upper Lone Pine Upper to Lower Lone Pine Mexican Ridge Wounding loss in hunting season Big Cr.Roaring Cr. Upper Newcomb Cr. Lower Lone PineLower Roaring Fk. Lower Forester Cr. Upper Lone Pine Lower Roaring Fk. Upper Lone Pine Lower Lone Pine Upper Big Cr. Independence Mtn. Shot by hunter Jan/Mar Fischer Pk.Delaney Butte Jan/Apr Delaney Fischer Delaney Fischer Jan/Mar Mar ButtePk. ButtePk. Fischer Pk N �Table 12. Summary of movement patterns for elk radio-collared on the Swift Ranch (Jct. North Platte River and Lake John Road), North Park. Telemetry Winter - Spring Summer - Fall Winter collar number Sex Date Location Date Location Date Location 340 Bull Feb/Apr Swift Ranch Jun Sept/Oct 440 Cow Feb/May Swift Ranch 460 Cow Feb/Apr May Swift Ranch Da 1aney Butte Oct Sept Oct/Nov Jun Sept/Oct Nov Lower Lone PineBig Ck. Lakes Divide W Big Ck. Lakes Shot by hunter Upper Lone Pine Lower Lone Pine Lower Red Canyon Upper Roaring Fk. 10 mi. NE Steamboat Sprgs. Jan/Mar Delaney Butte ex> VJ �To Walden AN R ..• /,,< _ ~ '\ -. .•, \ ;.,." ''''''''; ~ 1.1 (5,;;" '''''11" /', , .~"-""'""" •••••••••.• ) "". ~ ~ r-. ,·'0 ""'C~ 00 ;:!~Cu.t.r .__ +=- Mtn. "'•. .~ " \ I ?,,;t- ~,;. • 1 r . ..",...,..... ~. ( N 1 r;:.j .~'" ..:> • f(,~ o I Mil •• 2 . f(,~. ./cot' Fi g. 1. Migration routes of elk collared ~ -~. 'J .c: (_'---" ~t.~i ~t.~r· DIVIDE', - .,..- 't\~ on the Owl Ridge winter • \....J range. ~ �To Waldan\ I I I '--",".....• I I -- - ..•_,- e \ ,;".'t!j.tt I . '·'·'111Iit.-~0 ,,'" ~/ ,,-1> ''0 :;,~ ~<f' , / I, . \ . .I \~S'ia 'I Res. I '41,- I /. I r- ) . / . o 1 Mile. r \ _-- . -1>;~,> "'1-0\ '> .~ <: , -,," '- r. '", ~ I<, 2 ' '-. \ \ ./ ,/ l ) 'J- J---~•. (/' ../- -:.. ""'. ~ I ", "'-".; ',<- \\_\~- -'/ 1 "~ 1.\" 'I ~/ N 0 " { /' r\J , , Arapa, La k e "T'I~",. .~)._ GO~ . ~<~.'-. ", .~, .', I• lJ r>: __ ~''''..... .~~ ~.iT"'. MO., 0"IV, ,0 '\ /- . l $/l_ "'r>. / , •••.• I I I. '\, . ••.•.•..'~Haystack ", ..._,J Fig. 2. Migration routes of elk collared on the Buffalo Ridge Winter range. j' . M,., / .~ ...r __,.~" , Willo"" C •••• ~Parkvie"" M •• , •••• 00 Ul �'t4 ILDERNESS ", Me Inl yr. Burn .oJ f.p.'I" p.'" co o- 111 1 " ~II, "" 1 Jo",,"~ 7fl,,,,, "f, .: /. I·_ ~\ ~I~\ ~ iI~ ~ ; I~ ~\t_ ->: , •..•• ,. "tf/"tlt '1111 f'f f' ' 00 111",1,/ .i r· ... ~.. " tIIM\n;,.,' 'I' I'IIiU ~ /' f\\n~ I Miles winter Fig. 3. Migration routes of elk marked on the Sentinel Mountain range. �87 ':;~ I,..-"- .... ~ , I;" ,;.. ( I I I I I \ I III III 1&1 Z II: 1&1 Q ...• So.ttc her Lake ...• 1&1 ¥ II: N •.. Z :::I o ~ N 1 o Fi g. 4. Migration routes of elk marked on the Independence Butte winter ranqes. and Dalaney 2 �88 APPENDIX A ELK TRAPPING - NORTH PARK December 18. 1984 - March 7. 1985 Location of Traps Independence Mountain - along the loop road and halfway up Dead Horse Draw (7 traps) Sentinel Mountain - north side at start, later moved to east side (4 traps) Camp Creek - approximately 3 miles above junction with Platte River (4 traps) Delaney Butte - east side at base of cliff (3 traps) Baller Ranch - approximately 2 miles SW of junction of Highway 14 and Rand-Gould road (3 traps) Owl Ridge - Speck's Draw - approximately 1 mile SW of junction of Gould-Rand road wi th Owl Creek (4 traps) Owl Ridge - state property - on the east side of the road where the GouldRand road crosses Owl Ridge (4 traps) Buffalo Ranch - along side of the road in Buffalo Pass (7 traps) Marking of Elk Elk were marked with metal yellow eartags with the Fort Collins Division of Wildlife address and an identification number. Also, a green collar with white numerals (corresponding to the eartag 10 number) was placed on the animal. Regular Trapping Crew DOW George Bear Richard Byrd Al White Steve Porter Keith Kaler USFWS Gale Brewer Gary Sull ivan Dave Johnson Jackson County Sheriff Department Rick Rizor Other Contributors Vicki Anthony - DOW John Hobbs - DOW Chuck Cesar - BLM Gene Patten - USFWS Mike Hopper - Colorado Parks and Rec. Bob Miller - Colorado Parks and Rec. Al Purcell - USFS Kit Buell - USFS �89 Elk Trapping - North Park Injuries - Mortality -Elk - calf (RC 75) neck injury, recovered -Al Purcell - back injury, also recovered -Bull - (214) missing lower portion of right hind leg, apparently a hunting mishap, otherwise in good body condition; seen in mid-March - still doing very well. Summary of Elk Trapped Calves Yr. Ad. cows Sentinel Mt.Camp Creek cows Male Female 18 2 2 8 5 3 5 2 Buffalo Ranch 21 3 6 Owl Ridge-Baller 14 2 11 58 10 24 Independence Mt.Delaney Butte Tota 1 Recaptures from last year: Recaptures of this year: New Radio Collars: 45 2 7 Spikes Bulls 2 Total 32 18 3 7 6 44 8 7 3 45 19 17 11 139 �90 ELK TRAPPED - NORTH PARK 1984-85 Collar No. 1 2 3 4 5 6 9 10 11 12 13 14 15 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 Date 12/18 12/18 12/19 12/19 1/3 1/3 1/3 1/3 1/4 1/4 1/4 1/8 1/8 1/8 1/8 1/9 1/9 1/9 1/9 1/9 1/10 1/10 1/16 1/11 1/16 1/11 1/11 1/16 1/17 1/17 1/17 1/17 1/17 1/17 1/17 1/17 1/22 1/22 1/22 1/22 1/22 1/22 1/22 1/22 1/23 1/23 1/24 1/23 1/23 1/23 1/23 1/24 1/24 1/24 1/24 1/28 1/28 1/28 1/28 2/7 Sex Age F A F A F A F A F A F A F A M Ca F A F A F A F A F F Yr Ca Yr F A F A M M Ca Ca Ca Ca Ca F A r·, Ca F M F Ca F A F A F A F A F A F Yr F A F Yr F A M M Ca Ca F A M M Ca Ca Ca Ca F A M F Ca Ca F A F M Ca Ca F A M F M Ca F A F A F A F A F A Ca F l/r F A F Ca Ca Ca Location Buf Buf Buf Buf Buf Buf Buf Owl Buf Spk Spk Buf Buf Ba1 Ba1 Buf Ba1 Ba1 Ba1 Ba1 Owl Ba1 Buf Buf Ba1 Owl Owl Buf Buf Buf Owl Buf Buf Buf Ba1 Buf Buf Buf Buf Spk Spk Spk Spk Ba1 Buf Buf Spk Owl Buf Owl Owl Buf Buf Buf Ba1 Camp Camp Camp Camp Sent Remarks RC 590 RC 170 RC 290 RC 310 RC 71, Lg. RC 360 ET 16 Lg (230 1b) Last year =57 Lg Lg Sm Very 19 RC 79 RC 9.09 19 RC 190 Lg (220 1b) RC 9.42 RC 57 ET 36 ET 34 ET 35 Lg Sm ET Lg Lg Lg Lg RC Lg Lg (275 (150 216 (290 (230 (230 (255 320 (255 (215 lb) 1b) 1b) 1b) 1b), RC 9.08 1b), RC 62 1b) lb ) Very Lg Lg (235 1b) RC 11 RC 340 Very 19 Lg RC 90 Lg Lg (230 lb) Sm RC 82 �91 Collar No. Sex Age 1/29 1/29 1/31 F A F A F A s/5 F A F A F A F A F A F F Ca Ca F A 93 94 95 96 97 98 99 100 101 102 103 104 105 106 107 108 109 110 2/5 1/31 2/7 2/5 2/7 2/9 2/7 2/7 2/8 2/8 2/8 s/o 2/9 2/9 2/11 2/9 2/11 2/11 2/11 2/11 2/11 2/15 2/22 2/15 2/21 1/14 1/14 1/14 1/14 2/21 2/22 1/14 2/26 2/26 2/27 2/27 2/28 3/1 3/4 3/5 3/4 3/4 3/4 111 3/5 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 112 113 114 200 201 202 203 204 205 Date 3/7 3/7 3/7 1/3 1/3 1/4 1/8 1/8 1/8 Remarks Location F A F Ca Camp Camp Camp Camp Sent Camp Sent Sent Sent Owl Sent Ind Spk Owl Owl Owl Owl Sent Camp Ind Ind Camp Camp Camp Camp Camp Ind Camp Ind Buf Buf Spk Spk Sent Sent Owl Ind Ind Butte Ind Ind Ind Butte Sent Sent Sent Sent M Ca Sent Lg RC 9.15 Butte Butte Butte Buf Buf Owl Buf Bu1 Ba1 RC RC RC ET ET 9.17 Lg 9.18 Lg 68 7 RC 340 8 F Yr F A F A F A F A M Ca A F Yr Ca Ca Ca Ca F A F A M M Ca Ca F A F F Ca Ca Yr F Unk F M F A M Ca F A F A M Ca F A F A A F Yr Yr F A M Ca Ca F F A M Ca Ca M F A M A M A M Yr M A M Yr Yr M RC 220 RC 240 RC 270 Lg RC 9.10 Sm RC 9.20 ET 78 ET77 Lg RC 9.18 ET 83 Very Lg (210 1b) ET 82 Lg RC 9.19 Lg (275 lb) Lg (260 1b) Lg Lg RC RC Sm RC 9.12 (320 1b) 9.02 78 Sm (180 1b) Sm RC RC Lg (170 1b) 9.07 76 (250 1b) RC 9.10 RC RC RC RC RC RC 9.22 ET 103 9.03 ET 102 9.16 67 9.05 75 Sm ET 15 4x4 RC Y.13 �9/. Collar No. Date 206 207 208 209 210 211 212 213 214 1/9 1/10 1/10 1/16 1/11 1/11 1/22 1/23 1/16 215 216 217 218 219 220 221 222 223 224 225 226 227 1/23 1/23 1/24 1/24 2/6 2/5 2/6 2/8 2/8 3/4 2/20 3/4 3/7 Sex Age Yr A Yr Yr A A Yr Yr A Buf Spk Spk Buf Buf Buf Buf Buf Owl M Yr Yr A A Yr A Yr Yr Yr Yr M A M Yr Yr Buf Buf Buf Owl Owl Camp Spk Spk Owl Butte Sent Butte Butte M M M M M M M M M M M M M M M M M M M Remarks Location 3x4 Last Year #1 (84-1 ) RC 540 4x4 4x4 ET 209 4x4. Right Hind Leg ~li55 in9 4x5 5x5 RC 60 5x5 �Colorado Division of Wildlife Wildlife Research Report July 1984 93 JOB PROGRESS REPORT State of Colorado Project No. 01-03-047 Mammals 1 Research Work Plan Elk Investigations ------------------No. 3 ----------------- Job. No. 3 Period covered: June 1,1984 Author: Evaluation of Factors Influencing Elk Nutrition Status and Population Performance - July 30,1985 D. L. Baker ABSTRACT Comparative intake rates, digestion rates, and rates of passage were studied in mule deer, mountain sheep and elk consuming native diets. Mean voluntary intake in sheep and elk decreased with increasing amounts of browse in the diet while intake for deer remained relatively constant. Apparent digestibilities of dry matter and fiber decreased as browse content increased. Sheep and elk were more efficient digesters of these constituents than mule deer. Passage rates for all species increased with increasing percentage of browse in the diet. With the exception of mule deer, intake, digestibility and retention time were directly related to the amount of browse consumed. In accordance with theory, the larger bodied ruminants (sheep and elk) showed increased retention time and greater digestive efficiency. ��95 EVALUATION OF FACTORS INFLUENCING ELK NUTRITIONAL STATUS AND POPULATION PERFORMANCE Dan L. Baker P. N. OBJECTIVES .,..,.-. 1. To develop and test a system for evaluating the potential of habitats to support elk. 2. To improve the predictive capability of this system by identifying and quantifying variables influencing the range supply-animal demand model of nutritional carrying capacity. • SEGMENT OBJECTIVES 1. Complete data analysis and prepare manuscript on deer-elk comparative digestion, voluntary intake and rate of passage. 2. Complete data analysis and prepare manuscript on in vitro rate of digestion experiments. ACKNOWLEDGMENTS N. T. Hobbs, P. Neil, J. Ritchie, and L. Stevens assisted in various aspects of thi s study. METHODS AND MATERIALS Experimental methods pertinent to these studies have been previously described (Baker 1983:126-131). For 1984, 3 tame adult mountain sheep (x weight = 93 kg) were included in the comparative digestion studies. Using these sheep, we conducted 3 complete balance and rate of passage trials from October, 1984, to February, 1985, then compared these results to those previously collected from mule deer and elk fed the same diets. We followed the same conventional in vivo digestion protocol as for deer and elk in prior experiments. Diets fed were: 0% browse (all grass), 50% grass: 50% browse and 25% grass:75% browse (Table 1). Throughout this report I refer to these diets as O-diet, 50-diet, and 75-diet which corresponds to the % of browse in each diet. For passage measurements, we marked the diets and diet components with the same rare earth elements as in previous trials with deer and elk (Baker 1983:130). Analysis of feed and feces for chemical constituents and marker concentration was analogous to previous methods. �96 RESULTS AND DISCUSSION Voluntary Intake We observed different patterns of intake among species as we increased amount of browse in the diet (Fig. 1). Intake for sheep and elk decreased with increasing amounts of browse while intake for deer remained constant. All species consumed similar amounts of the O-diet (f > 0.05), but deer ate more of the 50- and 75-diet than either sheep or elk (P < 0.05). Intakes for mountain sheep and elk were not different (f < O.O~). Apparent Digestibility Apparent digestibilities of dry matter (OM) and neutral detergent fiber (NDF) decreased as browse content increased (Fig. 1B, Fig. lC). The decrease in NDF digestion was most apparent between the 0- and 50-diet (P < 0.05) and greatest for mule deer. Mule deer showed a 47% decrease; mountain sheep 34%; and elk 38%. The decrease in dry matter digestion was, again, most notable between the 0- and 50-diet but in contrast to NDF digestion, was greatest for mountain sheep. Sheep and elk were similar (P > 0.05) in their digestion of OM and NDF and higher than deer (V < 0.05). For the 50-diet, deer and elk showed similar and lower (P > 0.05) digestive efficiencies than sheep for OM and sheep and elk showed similar (P > 0.05) and higher efficiencies of NDF than mule deer. For the highest browse diet, all species were alike in their digestion of OM (~> 0.05), while sheep showed the most efficient utilization of NDF, followed by elk, then mule deer (P > 0.05). There was no significant interaction (P > 0.05) between animal and diets for either OM or NDF suggesting that the magnitude of the difference in OM and NDF is the same for each diet for all species. Turnover Time Passage rates appeared to be directly related to percentage of browse in the diet (Fig. 1C). Turnover times were generally more rapid for deer than mountain sheep and elk with the magnitude of the difference similar for all diets (no significant interaction). The higher passage rate in deer may be attributed to their smaller gut capacity and higher metabolic rate per unit of body wei ght. Rate of Digestion Experiments No further progress was made on thi s phase of the study. LITERATURE CITED Baker, O. L. 1983. Elk investigations--eva1uation of factors influencing elk nutritional status and population performance. Colo. Div. Wi1d1. Res. Rep. Ju1y:103-154. Prepared by ~-f---' /~D.' _-' _ . , _ oi?J. Baker ...,C,_,-r-, ...,..-:,··'l l1AA.=;\ Wildlife Researcher C �97 Table 1. Chemical composition (%) of experimental diets fed to deer, mountain sheeE, and elk. Test diets Item Crude protein NDF lignin 100% Grass 0% Browse 50% Grass 50% Browse 25% Grass 75% Browse 6.1 6.3 6.8 69.5 68.4 67.3 4.5 10.9 17.0 �98 A 16 §' CD a1~ at, 14 a1, C) ~ <, - ~ 12 C) Q) ":-b2 " ....•. b2 •••••..•••• c2 " 10 :::t:. es .E -a1 a1- 18 8 "" ", "c2 ••••••• Deer Sheep --- Elk Intake for mt. sheep and elk was similar and decreased for both; intake for deer did not change and was higher than for sheep or elk. 6 8 - 60 '#. 50 £ :.a en 40 Q) Ol (5 u... 75 50 0 a2 a2~~ ~ a1 ~~~ ~" ~~bc2 ~ b2 - •••••••• c2 ..- " 30 0 z b1~C3 20 -- 75 50 65 60 /C2 /"b12 •.... s: Q) b2," E t- b1 La 1/ 40 30 Deer "'••••••• Sheep ---Elk ~"", 50 J= •.... Q) > 0 c •.... ::J Digestion of NDF was more similar and higher for sheep and elk than for deer; NDF digestion decreased with increased browse. c1 0 C Deer ·······Sheep =----Elk ,/' a1;1' a1/ a1 at Retention time was more similar and longer for sheep and elk than for deer; retention time decreased with increased browse for deer, sheep and elk. 20~ I I I 0 50 75 % Browse In Diet Fig. 1 A-C. Comparative intake, digestion and passage of diets consumed by mule deer, mt. sheep and elk. Different numbers indicate significant differences (p 0.05) among species within diets. Different letters indicate differences among diets within species. = � https://cpw.cvlcollections.org/files/original/9a400fb773cfc10d5b9ec11e75e9cfc1.pdf ef5e3b7c77d49a09ecd97cdec9611b41 PDF Text Text Colorado Division of Wildlife Wildlife Research Report July 1985 99 JOB PROGRESS REPORT State of Colorado Project No. 01-03-048 Mammals 2 Research Work Plan No. Multispecies Investigations --------------------------1 ----------------------- Job. No. 1 Period Covered: JulY 1,1984 Author: Animal and Pen Support Facilities for Big Game Research - June 30, 1985 P. H. Neil ABSTRACT Seventeen pronghorn and 5 mule deer were hand reared and trained during the fiscal year. All were raised on undiluted canned evaporated milk. The pronghorn were used in a winter wheat game damage study in eastern Colorado. Twelve adult mule deer were involved in a winter range thermal cover study in Kremmling, Colorado, and returned to Fort Collins in good condition other than weight loss. Intake, digestion, and rate of passage trials were conducted on 3 bighorn sheep and 3 Rocky Mountain goats. A 4-phase Chronic Wasting Disease management program was established for the Fort Collins and Kremmling facilities with phase 1 being completed at the end of this fiscal year. ��101 ANI~mL AND PEN SUPPORT FACILITIES FOR BIG GAME RESEARCH Paul H. Neil P. N. OBJECTIVES To provide and maintain populations of captive big game animals and pen facilities to support big game research programs. SEGMENT OBJECTIVES 1. Continue to develop and maintain facilities at CSU Foothills Campus and Wildlife Research Center. 2. Coordinate rearing, training, and research activities with captive wild and tame big game animals for all big game cervid and noncervid research projects. 3. Integrate big game animals and physical plant support facilities, personnel, and fiscal resources into a single budget. METHODS AND MATERIALS Routine neonate rearing procedures were used to complete the hand rearing of 17 pronghorn and 5 orphaned mule deer fawns. These animals were reared and trained for various nutritional and game damage studies. Twelve adult mule deer were transported via stock trailer to the Junction Butte Research Facility near Kremmling, Colorado, in November in support of the winter range thermal cover study - Dave Freddy, Principal Investigator. One deer died, and the remaining 11 returned to the Fort Collins facility in April in good condition other than weight loss. Intake, dige,stion, and rate of passage trials were conducted on 3 bighorn sheep and 3 Rocky Mountain goats during the winter. Diet on the first trial consisted of 50% low quality grass hay and 50% vaccinium. Diet on the second trial consisted of 75% vaccinium and 25% low quality grass hay. Fecal samples were taken to the Denver Federal Center, U.S. Geological Survey Laboratory where they were bombarded with nutron activation and markers then counted for the rate of passage study - Tom Hobbs, Dan, Baker, Principal Investigators. Nine pronghorn were transported to eastern Colorado for use in a game damage study on winter wheat. All were returned to the Fort Collins facility in May and in good condition. A 4-phase chronic wasting disease management program was established for the Foothills and Kremmling facilities. The 4 phases consist of the following: (l) Elimination of the entire captive deer and elk population; (2) disinfection of the facilities; (3) construction of a perimeter barrier fence, and (4) reintroduction of captive deer and elk. Phase 1 of the program was completed in late June under the direction of Dr. Beth Williams of the Wyoming �102 State Veterinarian's staff who has been working on the slow virus for many years. Brain and spleen samples were taken from each animal for diagnosis. Phase 2 was begun and is at this time about half completed. Disinfection is being accomplished using a 65% active chlorine solution with an application rate of 1,000 ppm. Application is done by mobile sprayer and by helicopter. The holding pens will be unoccupied for 1 year before reintroduction of animal s. A 12' X 50' mobile home was placed at the Foothills facility to provide housing for temporary employees and to replace a trailer lost to fire in November. Construction of a waterfowl research area was begun at the Foothills facility. The pens and pond area will house approximately 75 ducks. RESULTS AND DISCUSSION Minimum diarrhea problems were encountered during the hand rearing of pronghorn and mule deer. The few problems that did occur cleared up within a few days. We anticipate staying with the undiluted, canned milk formula for future hand rearing of big game animals. The final results of the intake, digestion, and rate of passage studies will be reported by Dan Baker and Tom Hobbs when completed. An interesting sidenote to this study is that the bighorn sheep experienced very little weight loss after 2 trials in comparison to the Rocky Mountain goats. From beginning of intake trials to end of rate of passage trials, maximum weight for the sheep was 8 kg vs. 18 for the goats. The goats could not be used on the second trial due to stress and loss of weight. The results of the winter wheat game damage study are not available at present but should be reported in the near future by Steve Torbit and Len Carpenter, Principal Investigators. . The chronic wasting management program was established in an effort to break the cycle of the slow virus disease which has been affecting our captive herd for many years. Other facilities have executed the same or similar procedures and have thus far reported being successful in breaking the cycle. The application rate of 1,000 ppm Calcium Hypochlorite (65% available chlorine) is a more than adequate rate for disinfection. The holding pens will remain unoccupied for one full year before reintroduction of animals. At present, the captive big game herd consists of 4 Rocky Mountain goats, 17 bighorn sheep, 13 pronghorn antelope, and 1 domestic cow. Ten pronghorn were lost during the year as a result of fire and harassment by dogs. Construction of the waterfowl research facility began during the summer. The facility is being moved from the old Wildlife Research Station to the Foothills Research Center as a result of interstate highway construction. A concrete pond, pen framing and water lines were installed by the end of June. Other activities at the Foothills facility during the year consisted of educational tours for wildlife personnel from various state and federal agencies, Colorado State University personnel, summer science classes and local Boy/Girl Scout troops. Prepared by: Q~-~ Paui:Niil Wildlife Technician III �Wildlife Research Report July 1985 103 JOB PROGRESS REPORT State of Colorado Project No. 01-03-048 Mammals 2 Research Work Plan Multispecies Investigations --------------------No. 1 -------------------- Job. No. 4 Period Covered: July 1,1984 Authors: Personnel: Big Game Forage Selection Dynamics - June 30, 1985 S. C. Torbit, J. Liewer, A. W. Alldredge, R. B. Gill P. H. Neil, J. Stone, C. Chaffee, R. Sayre ABSTRACT Grazing exclosures which had been established on 2 January 1984 on a wheat field in northern Weld County were harvested on 16 July 1984. fvleangrain yield for grazed plots was 2840 kg/ha and for ungrazed plots was 3070 kg/ha. Mean total biomass was 7120 kg/ha for grazed plots and 7750 kg/ha for ungrazed plots. Statistical analysis revealed no significant differences existed between grazed and ungrazed plots in either total biomass or grain production. A 7-ha enclosure was erected on a wheat field in central Weld County and 12 pastures of 0.6 ha each were constructed within the enclosure. Hand-reared pronghorn, 5 months of age, were randomly assigned to 1 of 4 grazing treatments: control (no grazing); early grazing (16 November10 March); late grazing (10 March-6 May); and continuous grazing (16 November6 May). Wheat biomass was estimated monthly and on 18 July 1985 all wheat was harvested. Mean grain yields (kg/ha) for control, early, late and continuous treatments were 317.6, 295.8, 319.7 and 294.5, respectively. Statistical analysis showed no significant differences in grain yield among treatments. Biomass estimates revealed that standing biomass is an appropriate index for pronghorn grazing pressure but was not correlated to final grain yield. Aerial surveys were continued on both study areas in eastern Colorado. Pronghorn distribution shifted from wheat to grasslands in March and April, 1985. Landowners known to have pronghorn grazing on wheat were surveyed and asked to supply July, 1984, yields. Sixty-nine percent of landowners contacted supplied the necessary data, but no relationship could be established between days of pronghorn use and grain yield. Phenology plots were established on 4 grassland sites in Weld County and phenology of native plants tracked through 1984-85 winter. Time of green-up was not related to pronghorn distribution as determined from aerial surveys. �104 Twenty-four collars were recovered from pronghorn marked in the northern unit during January and February, 1984. Seventeen marked pronghorn were legally harvested during fall 1984, the remaining 7 collars were recovered from winter-killed or illegally harvested animals. Marked pronghorn foraged extensively in wheat fields north of Riverside Reservoir during 1984-85 winter and then retreated to the southwest portion of the study area or moved north onto the Pawnee National Grasslands to summer. Fifteen marked pronghorn were legally harvested in the southern study area during fall 1983. Eight of the legally harvested pronghorn were taken north of the study area. Pronghorn were again trapped in the southern area and 107 additional animals were marked. Observations of marked pronghorn demonstrated that animals continued to use wheat fields in the vicinity of the trapsites until March and April and then migrated to the northern and eastern portions of the study area. Fourteen additional pronghorn were instrumented with radio-telemetry equipment during January, 1985, in the northern study area. Telemetered pronghorn began abandoning wheat fields and moving into native prairie by 19 March 1985. All telemetered pronghorn had abondoned wheat fields and moved to grassland by mid-April, 1985. Attempts were made to monitor nighttime foraging activities of instrumented pronghorn during March, 1985. Two males foraged on wheat during darkness on 3 of 8 occasions during March. Feeding bouts were short (approximately 20 mins) and insufficient data was collected to quantitatively assess significance of night foraging. �105 INTRODUCTION Results from year 2 of our 3-year investigation of the effects of pronghorn grazing of winter wheat are presented herein. Results of each subproject are presented separately. Because this is the second progress report for the pronghorn-wheat project, frequent reference is made to our initial progress report. Details of methods employed and objectives will not be repeated here; instead the reader will be referred to our initial progress report (CSU - Pronghorn-Wheat Project, Progress Report #1), which reported work performed from the period 1 September 1983 to 1 July 1984. �106 P. N. OBJECTIVE Evaluate the impacts of pronghorn antelope on winter wheat yields. SEGMENT OBJECTIVES 1. Test accuracy of fecal analysis to estimate diets of big game ruminants. 2. Investigate impacts of pronghorn antelope grazing activities on winter wheat yields. 3. Develop a conceptual model of pronghorn habitat interactions and develop a reseach proposal to validate the conceptual model. FECAL ANALYSIS Hypothesis Biases of ruminant diet composition estimation will increase with increasing proportions of shrub stem in the diet. Methods and Materials A pilot trial was conducted with a fistulated elk at the Colorado Division of Wildlife's Foothills Research Center. That trial was used to perfect the techniques for introducing diets into the rumen, for collecting fecal samples, for preparing and mounting samples on fecal slides, and for training the graduate research assistant in the microhistological technique. Subsequent experiments will be conducted with fistulated goats. Experimental Protocol This experiment will be conducted by feeding diets of known composition directly into the rumen of up to 4 fistulated goats. Each feeding trial will last 7 days. Dates of the 1985 trials will be April 10-17, June 2-9, and June 17-24. The goats will be fed a low (25%) shrub diet during trial 1, a medium (50%) shrub diet during trial 2, and a high shrub (75%) diet during trial 3. Shrub components of all diets will be stem material; and remaining portions of each diet will consist of equal proportions of grass and forbs. Plant collections will be made in the Fort Collins, Poudre Canyon area during February and March, 1985. Native forages will be mountain mahogany (Cercocarpus montanus), skunkbrush (Rhus trilobata), crested wheatgrass (Agropyron cristatum), little bluest~Sch;zachyrum scoparium), and fireweed (Kochia scoparia). Alfalfa (Medicago sativa) will be obtained commercially. After sun mulcher. fragments through a from each drying, the forages will be chopped and rechopped in a shredder Fragment sizes will range from .5 cm to 10 cm. _To insure that <2 cm are not included in the diet, the forages will be filtered 1/2-cm screen. Dry matter will be determined by drying samples forage at 500 C for 48 hrs. �107 Daily rations will be weighed, each corrected for dry matter variation by multiplying percent 3rymatter X the percent allocated for the diet (Tables 2-4). Daily rations wlll be separated, labeled, and stored. In order to prevent digestive upsets caused by rapid diet changes, a 14-day adaptation program will be conducted. The goats will be isolated in a 6- x 8-m pen and will be fed only wafers and grass hay with the percentage of wafers decreasing and percentage of grass hay increasing. After 10 days, increasing amounts of chopped mountain mahogany and skunkbrush will be inserted into the rumen. At the end of 14 days, the animals should be sufficiently adapted to a low quality, high shrub diet. Each feeding trial will last 7 days, but the first 3 days will serve as an adjustment period because it will take at least 48-72 hrs for the test diet to begin passing. The entire diet will be fed directly into the rumen. After taking a grab sample, the forage will be soaked in water; this facilitates handling and the forage, once hydrated, will not expand in the rumen. The goat will be fed 3 times daily with about .5 to .7 kg each meal. Starting on day 4 and continuing through day 7, a rumen sample, in addition to a grab sample from the test diet, will be collected prior to feeding. Fecal samples will be collected immediately after each meal. In the evening, after the last meal of the day, the pen will be cleaned of all fecal matter and other debris. This will insure there is no mixing between daily fecal depositions. Each sample will be placed in a separate labeled and dated ziploc plastic bag. All samples will be freeze dried, ground through a micro-wiley mill with a l-mm screen, and soaked in domestic bleach as a preparation for mounting. After rinsing with water, standard templates (Hansen et al., unpubl.) of fragments will be mounted with Hoyers solution on the microscope slide. After spreading the fragments uniformly with a probe and placing a cover slip over the Hoyers-fragment mixtures, each slide will be held over a lit bunson burner for a few seconds to seal the cover slip. Five slides will be made from each sample, and 20 valid observations per slide at 100X magnification (light microscope) will be recorded. At least 2 identifiable fragments with cellular mateial will be the requirements for a valid field. Estimated percent composition of each sample will be calculated with the frequency addition procedure (Holechek and Gross 1982) where: Estimated Dry Weight = Freruency of Each Species Tota Number of Frequencies for All Species Analysis: The statistical design will be a 2-way analysis of variance, with the 3 diets and daily samples as treatments: �108 ANOVA Tabl e Source Degrees of Freedom Total 20 Diets 2 Days 6 Error 12 Differences in accuracy of estimation will be determined according to the above ANOVA design. Similarity between actual dry weight and estimates from undigested, rumen, and fecal samples will be calculated with Kulcyznki IS formula (Oosting 1956). Results Microhistological slides have been prepared from feces collected during the pilot study with the fistulated elk. Analysis of slides is now commencing. Four goats have been selected for fistulation and fistulas have been ordered. �109 PRONGHORN DA~JAGETO ~JINTER HHEAT CotJTROLLED GRAZING BY FREE-RANGING PRONGHORN ~·1ethods Thirty paired plots which had been established on a wheat field in northern Weld County on 2 January 1984 were harvested on 16 July 1984. Pronghorn had been allowed to graze 1 plot of each pair and both biomass and numbers of pronghorn uti1izing the field were estimated bimonthly from January through t~ay, 1984 (CSU Progress Report #1 1984). Each permanent sub-plot wi thin paired plots was hand clipped at ground level and vegetation placed in a paper sack and transported to Fort Collins. The grain was separated from the head by hand and total wei ght of both grain yi eld and total bi onass were compared for grazed and ungrazed plots by a paired t-test (f = 0.01) Thirty paired plots were reestablished on a neighboring 81-ha wheat field on 21 November 1984. Numbers of pronghorn and total biomass were estimated bimonthly from December, 1984, through Hay , 1984. Total biomass Has estimated by a procedure different from that employed the previous winter. Ocular double samp1ing (Laycock 1962, Range Resea rch ~·1ethodsSubcommi ttee 1962) was employed to estimate biomass from each plot from December, 1984, through nay, 1985. Results Significant numbers of pronghorn utilized the experimental field from January through May, 1984. Numbers of pronghorn grazing on this experimental field ranged from 117 in January to 38 in f'lay(CSU Progress Report #1 1984). No significant differences in total biomass could be detected between grazed and nongrazed plots within sample date (January - t·1ay,1984) (CSU Progress Report #1 1984). Mean wheat biomass for grazed plots at harvest was 7120 kg/ha, and mean biomass for nongrazed plots at harvest was 7750 kg/ha (Table 1). Mean total grain yield for grazed plots was 2840 kg/ha, and mean total yield for nongrazed plots was 3070 kg/ha (Table 1). No significant differences could be detected between grazed and nongrazed plots for either total biomass or grain yield. Mean grain yields were 17.2 bushels per ha for grazed plots, while nongrazed plots averaged 18.5 bushels per ha. The average gain yield for the entire 160 acre field was 18 bushels per ha (J. Fiscus pers. comm.). The unusually dry, windy 1984-85 winter prevented successful replication of this grazing experiment. Weather data collected from an atmospheric measurement station near the experimental field demonstrated that the 1983-84 winter was wetter than the 1984-85 winter (Table 2). Wheat plants successfully emerged during early winter 1984 but due to extreme winds and low precipitation many of the wheat plants suffered from w+nd burning and did not provide succulent forage for pronghorn. No pronghorn were observed to utilize the experimental field until 10 ~1arch 1985 when herd of 20 animals was observed grazing on the field. All pronghorn had abaondoned the experimental wheat field by 25 ~~arch 1985. Estimates of standing biomass �110 (wet weight) for all experimental plots never exceeded 500 kg/ha even after spring green-up. Accordingly, the owner of the wheat field plowed and reseeded the entire field to oats during late M~, 1985. Therefore, no wheat was harvested from this field during summer 1985. �Table 1. Wheat biomass and grain weight at harvest (16 July 1984) for paired wheat plots in northern Weld County, Colorado. Grain yield Grazed Total biomass Ungrazed Ungrazed Grazed kg/ha 1b/acre kg/ha 1b/acre kg/ha 1b/acre kg/ha 1b/acre Mean 2840 2534 3070 2739 7120 6352 7750 6915 S.D. 1290 1151 1380 1232 3280 2926 3530 3149 S. E. 240 214 260 232 610 544 660 589 •.....• •.....• •.....• �112 Table 2. Normal precipitation (cm) and 20 year average temperature (C) for the Kauffman site (northern Weld Co., Colo.); mean precipitation (cm) and mean temperature (C) collected from October, 1983, to March, 1985, for that same site. Date Mean precip. (em) Normal precip. (cm) Mean temp. (C) 20 Year ave. temp. (C) Oct, 1983 0.86 1.32 10.3 9.6 Nov 2.31 0.71 1.9 2.2 Dec 0.69 0.66 -9.5 -1.8 Jan, 1984 0.38 0.79 -3.4 -3.4 Feb 2.54 0.33 0.9 -0.8 Mar 2.69 1.63 1.8 1.8 Apr 6.50 3.02 3.6 7.5 May 2.62 6.02 18.59 14.48 Oct 3.43 1.32 7.1 9.6 Nov 0.00 0.71 2.3 2.2 Dec 0.41 0.66 -2.1 -1.8 Jan, 1985 0.58 0.79 -5.4 -3.8 Feb 0.00 0.33 -3.7 -0.8 Mar 0.00 1.63 4.42 5.44 TOTAL TOTAL �113 --~----~------~14~------------------~ ·BRIGGSDALE ~fr~ WELD CO MORGAN CO. 4S~w, 00 POINT OF D f:"1...4E.,.N ·ROCKS 0 S GREASEWOO~ LAKE I I I 18 ·II~ V OeD Uo:: MACKSON RES . 1 :31 VERSIDE RES. ~ ~ I ____ ~--~--~I~ ~~~~ ~UTH 2 mi Fig. 1. PL~\\"- 2km Location of experimental pronghorn grazing enclosure winter wheat field in central Weld County, Colorado. FT. MORGAN (EN) on �114 EXPERU1ENTAL GRAZING HITH HAND-REARED PRONGHORN r1ethods An enclosure was erected on a wheat field located in east-central Weld County (T6N, R61W, Sl) (Fig. 1). The fence enclosed 7 ha that had been planted on 5 September 1984 with Baca variety of hard red winter wheat and fertilized in June, 1984, with anhydrous ammonia and liquid phosphorus at 39 and 13 kg per ha, respectively. The enclosure was divided into 12 pastures of 0.6 ha each and each pasture was randomly assigned 1 of 4 treatments: early grazing, late grazing, continuous grazing, and no grazing (control). This design allowed each grazing treatment to be replicated 3 times. Six hand-reared pronghorn were used in the grazing experiment. These pronghorn had been reared and trained at the Division of Wildlife ungulate research facility during summer 1984. Experimental animals were at least 5 months old at the beginning of grazing trials. Three male and 3 female pronghorn were randomly assigned to graze 1 of the experimental pastures. Grazing was initiated on early and continuous pastures on 16 November 1984, and on late pastures on 10 t{larch1985. Early grazing was terminated on 10 r~arch 1984. All animals were randomly reassigned a treatment and moved to the appropriate pasture at the beginning of late grazing. Pronghorn were removed from all experimental pastures and returned to Fort Collins on 6 May 1985. Early grazing, representing use during winter dormancy, consisted of 115 days of animal use (16 November - 10 r1arch). Late grazing occurred during spring green-up and late jointing, lasting 57 days (10 r~arch - 6 t'lay). Continuous grazing occurred during the entire experimental period of 172 days (16 November - 6 May) applying grazing pressure during both vegetative and early reproductive stages of winter wheat. The experimental stocking rate of 1 pronghorn per 0.6 ha is greater than densities observed for wild pronghorn utilizing wheat fields in eastern Colorado. Animals were allowed to graze wheat ad libitum and without interference throughout the entire trial. Water was provided ad libitum daily and animals were supplied with a windbre~k in each pasture. Vegetative biomass was estimated monthly on 24 permanent plots (0.2 m2) randomly established in each pasture. Biomass was estimated by ocular double sampling (Laycock 1962), and all monthly biomass data was standardized to biomass on a dry matter basis. Each pasture was harvested separately with a wheat combine on 18 July 1985. Grain was unloaded from the combine into a scale wagon located on the pasture and total weight of harvested grain was determined. Five random samples were withdrawn before the grain was transferred to a truck, and moisture and protein content were determined for each grain sample. Total grain yield was converted to number of bushels per ha, standardized for 10% grain moisture content. Differences in grain yield among treatments were analyzed by a one-way analysis of variance (P = 0.01), both with and without use of November biomass as a covariate. �115 Results Mean November biomass for all pastures was 152 + 20 kg/ha (dry matter basis). By February, significant differences in biomass existed among treatments; early, continuous, late and control pastures measured 43, 34, 131 and 192 kg/ha, respectively. Mean February biomass of early and continuously grazed pastures was significantly different from mean biomass of both late and control pastures (neither had been grazed by pronghorn). Spearman's rank correlation coefficient (Snedecor and Cochran 1971) was used to order monthly biomass rankings and compare rankings of July grain harvest from individual pastures. Significant discordance was shown between pasture biomass and final grain yield, indicating that measurement of wheat biomass serves only as an index to grazing pressure, but not an accurate predictor or final grain yield. Grain yield from experimental pastures ranged from 9.5 to 14.1 bushels per ha. Mean grain yields for early, late, continuous and control grazing treatments were 12.0, 13.0, 11.9 and 12.8 bushels per ha, respectively (Table 3). No significant differences in grain yield were detected among treatments. Grain yield for that portion of the wheat field outside the enclosure averaged 13 bushels per ha (K. Harpring pers. comm.). Experimental grazing will be repeated during the 1985-86 winter. The enclosure will be reassembled on the same wheat farm stocked with handreared pronghorn and the same wheat grazing treatments applied. �•....• •....• o- Table 3. Mean grain yield from wheat pastures grazed by pronghorn, November, 1984 - May, 1985. Mean yield Standard deviation Range Treatment bu/acre bu/ha bu/acre Control (no grazing) 31.8 12.9 29.3 - 34.5 Early grazing (16 Nov - 10 Mar) 29.6 12.0 Late grazing (10 Mar - 6 May) 32.0 Continuous grazing (16 Nov - 6 May) OVERALL bu/acre bu/ha 11.9 -14.0 2.2 0.9 23.4 - 33.9 9.5-13.7 4.5 1.8 13.0 26.9 - 34.9 10.9-14.1 3.6 1.5 29.4 11.9 25.4-34.1 10.3 -13.8 3.6 1.5 30.7 12.4 23.4-34.9 9.5-14.1 3.9 1.6 bu/ha �117 WINTER WHEAT QUALITY VS. NATIVE FORAGES Methods Quality of native forage diets was estimated bite-count observations of dietary items selected by free-ranging, hand-reared pronghorn fawns (6-12 mos old) from February through June, 1985 (Hoover 1981). Samples were collected from pronghorn dietary items that exceeded 2% of the total bites for each month. These samples were analyzed for fiber constituents, cell contents, crude protein, and in vitro digestibility at the CDOW Research Laboratory. Estimates of diet quality were derived by weighting the individual species values proportionally to average bite size for each species (Hobbs et al 1981 ). Winter wheat samples were collected at one month intervals from November, 1984, through June, 1985, and analyzed at the Colorado Division of Wildlife's Research Laboratory for fiber constituents, cell contents, crude protein and in vitro digestibility (Goering and Van Soest 1971, Horwitz 1980, Tilley and Terry 1963). Results Gi11 (1984) specul ated that the key to changes in habi tat use by pronghorn from native forage pastures to winter wheat in late fall and from winter wheat back to native forage pastures in spring could be found in comparative forage quality dynamics of the 2 habitats. In fall following emergence after planting winter wheat, forage value was expected to dramatically exceed forage value of native forage diets because native forages were declining in quality prior to winter senescence. Gill (1985) speculated that pronghorn would switch from native forage pastures to winter wheat fields whenever the inclining curve of winter wheat quality intersected the declining curve of native pasture forage (Fig. 2). Likewise, pronghorn would evacuate winter wheat fields for native pastures when declining quality of winter wheat intersected inclining quality of native forage diets in the spring. Observations of radio-collared animals indicated that pronghorns had moved from native forage pastures to winter wheat fields by mid-November, 1984. By the time the first samples were collected for forage quality analysis in November, winter wheat forage quality had already surpassed quality of native forage diets in terms both of crude protein and cell contents (Figs. 3, 4). In spring, cell contents of native forage diets exceeded cell contents of winter wheat by mid-April, 1985, which coincided with the period when radio-collared pronghorn abandoned winter wheat fields for native forage pastures (Fig. 3). Crude protein content of native forage diets exceeded crude protein of winter wheat by the beginning of r~ay, 1985 (Fig. 4). �ll8 100 90 80 ,.- - / 70 I 60 " " \ Cell contents (%) 50 \ r 40 \~_" F' .% G: 30 20 / -;5" j en / 'pi ~ I '\ 10 \ 7 F M A til a Ju I '\ ., Jl ., I / 7 Au " 5 q If N 0 0 Month Fig, 2. Hypothetical chronology of winter wheat forage quality and native forage diet quality dynamics. �119 V') I- z: w I- z: <:) u .....J .....J W U • \ \ \ \ I \ I \ \ I 'J 30'~------~~--~--~--~~--~--NOV DEC JAN FEB MAR APR MAY JUN Fig. 3. Comparisons of cell contents of seasonal winter wheat (. .) and native forage (.--- ...• ) samples. �120 32 28 24 ~ :z •....• w 20 f- a , a:: CI... w / 16 / Cl ::J a:: u I 12 I •... ] - / ...• I ...........•. _-4 NOV DEC JAN Fig. 4. FEB MAR APR MAY JUN Comparisons of crude protein of seasonal winter wheat (. .) and native forage (e--- ....• ) samples. �121 AERIAL SURVEYS Methods The 2 study areas (Figs. 5 and 6) and methods used to survey winter pronghorn distribution have been described previously (CSU Progress Report #1 1984). Pronghorn distributions (wheat vs. grassland) were tallied after each flight and relative distributions tracked through time. Legal descriptions of wheat fields consistently used by pronghorn were recorded and owners of these fields were contacted and asked to supply grain harvest from their fields. Pronghorn occupancy was correlated with grain yield by simple regression techniques. Four sites were selected on native grassland along a north-south gradient in units A-17 and A-13l (Fig. 7), and 4 permanent plots (1 m2) were established at each site. Phenology of native vegetation was monitored bimonthly and described according to the scale described by Dickinson and Dodd (1976) (Appendix 1). An attempt was made to relate pronghorn distribution to phenology of native grasslands. Our objective was to determine if pronghorn movement to native ranges in spring was related to green-up, by comparing habitat use (determined from aerial surveys) to phenology at grassland sites. Results The majority of observed pronghorn occupied wheat or wheat stubble fields through early February (Table 4). However, by 19 February the majority of pronghorn in the southern study area were occupying grassland habitat. Pronghorn in the northern study area were utilizing wheat and grassland types in equal proportions through early April (Table 4). Total number of pronghorn observed in each study was reduced compared to 1983-84 winter, even though more surveys were conducted during 1984-85 winter (CSU Progress Report #1 1984). Eight hundred seventy-five pronghorn were counted in 4 aerial surveys during 1983-84 winter, and 815 pronghorn were counted in 7 aerial surveys during 1984-85 winter in the northern unit. In the southern unit, 2622 pronghorn were counted in 3 aerial surveys in 1983-84, and 2425 pronghorn were counted during 6 aerial surveys in 1984-85. One hundred twenty-four landowners were contacted and asked to supply wheat harvest figures from their fields. Sixty-nine percent of all landowners contacted supplied the necessary information; however, the response rate was much higher for landowners in the southern unit (79%) than the northern unit (62%). Only 2 northern unit landowners whose wheat had been grazed by pronghorn responded to the questionnaire. Accordingly, yield data was combined for both study areas and not analyzed separately. No significant relationship could be detected between grain yield and pronghorn occupation for wheat fields in eastern Colorado. Yield data (Table 5) were extremely variable and any effect of pronghorn grazing was masked by other factors (moisture, fertilization, seed bed preparation, etc.). Grain yields for fields which were never observed to be occupied by pronghorn ranged from 4.3 to 21.5 bushels per ha (Table 5). Yields for fields observed to be grazed by pronghorn were less variable than those which pronghorn did not use (Table 5) and averaged 10.0 and 10.4 bushels per ha for 1-20 and 21+ pronghorn days of occupancy, respectively. �122 Phenology plots were generally characterized by winter dormancy until sampled on 31 March 1985. Phenology rankings on that date showed that significant green-up was beginning: Agropyron smithii plants were in stages 3, 4 and 5; Artemisia frigida was in stages 5 and 6 and very green; an unidentified bunchgrass was in stage 5. Habitat distribution (as determined from aerial surveys) in the northern unit showed nearly equal pronghorn distribution between grassland and wheat (Table 4) by 1 April. A strong relationship between occupied habitat and grassland phenology was not demonstrated. The phenology project will be continued through the 1985-86 winter. �123 --~----~------~14~------------------_ -BRIGGSDALE GIY~ WELD ------ CO MORGAN CO. 4S~w. 00 POINT OF D 1='1..4'/" -ROCKS Q s. GREASEWOOQ LAKE I • 18 "I~ 0(9 u~ _II ~ 010 MACKSON RES V . VERSIDE RES, W 3. ~--~--~~~I~ ~~~~ SOUTH PL~"('\~ 2 mi H 2 km FT. MORGAN Fig. 5. Pronghorn-wheat study area in northeastern landmarks are indicated. Colorado, important �-, .i . ~: '_.:- •••• KIOWA I• . • .. • il i@ I • I >~> l- l- CALHAN z·z ~w ::lD::l .. • I A-35 0110 OgO 0·1(/)gc::: ~.~ ..JrI...,J W·UJ I ELBERT •111•_. • 1. LI NCOLN -§...-_.-----....---@ YODER I • I COUNTY • - • - • _ • _ tJ COLl NTY • #pUNKIN CENTER >1 1-- zil ::l. all o. O~ (/). A-42 ~g • • u1:1 I • ! ._._._.-._._.-.-._.- Fig. 6. LINCOLN COUNTY CROWLEY COUNTY Pronghorn-wheat study area in southeastern important landmarks are indicated. Colorado, �125 4;11 3 --~----~------~14~------------------~ -BRIGGSDALE G,l? WELD ------ CO MORGAN CO, ~4S~ WOOD POINT OF -ROCKS ;:-"4-'" Q 'S I GREASEWOOQ LAKE I I Is 2 MACKSON RES V . SOUTH 2.mi Fig. 7. H 2km Locations of 4 phenology plots (numerals) located on native grassland in northeastern Colorado. Plot number 4 is located 18 mis northeast of New Raymer. FT. MORGAN �>--' N (J\ Table 4. Pronghorn distribution (%) by habitat type determined by aerial surveys for 2 areas in eastern Colorado, November, 1984 - April, 1985. A-17 Flight date A-35/A-42 Wheat Stubble Grass Other Total observed Nov 13 Nov 15 78.5 16.9 4.6 0.0 130 Dec 18 Dec 19 30.8 Jan 21 Feb 6 Feb 7 Feb 18 Feb 19 0.0 Mar Mar Apr Apr 6 7 1 9 100.0 40.8 50.7 42.5 61. 2 0.0 0.0 3.3 0.0 15.9 8.2 0.0 0.0 44.1 49.2 41.6 0.0 100.0 0.0 11.8 0.0 0.0 Wheat Stubble Grass Total observed 22.3 25. 1 52.5 394 0.0 0.0 100.0 13 72.8 11. 3 15.9 353 33.5 4.4 62. 1 639 18.6 0.8 80.5 601 6.1 0.0 93.9 425 147 40 95 152 138 113 �Table 5. Wheat yield from fields in eastern Colorado that were observed to have been grazed by pronghorn during 1983-84 winter. Yields were supplied by owner/operators responding to a mailed questionnaire, 124 owner/operators were contacted. o Pronghorn days 1-20 Pronghorn days 21+ Pronghorn days bu/acre bu/ha bu/acre bu/ha bu/acre bu/ha Mean wheat yield (bu/acre) 29.8 12.1 24.7 10.0 25.6 10.4 S.D. 11.7 4.7 8.4 3.4 9.0 3.6 10-53 4-21 8-36 3-15 15-34 6-14 33 11 11 7 7 Range (bu/acre) Number of responses (n) 33 ,_. N -.._J �128 REOBSERVATI ON OF t·1ARKEDPRONGHORtJ Methods Pronghorn were again trapped in both study areas during January and February, 1985 (Figs. 8 and 9). Fourteen pronghorn were trapped in the northern study area and equipped with radio transmitters. One hunded seven pronghorn were trapped in the southern unit and banded with orange collars marked with black numbers. Pronghorn that were using or in the vicinity of wheat fields were trapped and marked. Data on marked pronghorn were collected by methods described previously (CSU Progress Report #1 1984) and from winter-killed or harvested pronghorn. Results In the northern study area, 17 collars were returned from pronghorn that were legally harvested during fall 1984. Seven additional collars were recovered during winter and spring 1984-85: 5 collars were recovered from winterkilled pronghorn, 1 from an illegally harvested animal, and 1 collar from a wounded pronghorn. The majority of harvested pronghorn were taken between Highway 14 and the South Platte River, west of Morgan County line (Fig. 10). Twenty-three percent of collars placed on pronghorn in February, 1984, had been recovered by July, 1985. A maximum of 92 collars remain in the northern study area as of 1 September 1985. Reobservations of marked pronghorn during winter 1984-85 showed extensive use of wheat fields between Greasewood Lake and Riverside Reservoir and west of the Weld County-Morgan County boundary (Fig. 11). Large tracts of grassland border these wheat fields, and marked pronghorn would travel out of grasslands to graze on wheat fields. As spring progressed, pronghorn were observed deeper into this large tract of grassland. After 1 May, no marked pronghorn were observed to utilize wheat fields in unit A-17. Marked pronghorn--occupied grassl ands in the southwest corner of the study area or moved north and west of Briggsdale onto Pawnee National Grasslands (Fig. 11). Fifteen marked pronghorn were legally harvested in the southern study area during fall 1984. No other mortality of marked pronghorn has been discovered in the southern unit. Eight of the legally harvested pronghorn were harvested outside of the study area north of Highway 24 (Fig. 9), and only 2 collars were recovered from legally harvested pronghorn in A-42 (Fig. 12). A maximum of 228 marked pronghorn remain in the vicinity of the southern study area; 11% of collars placed on marked pronghorn were recovered from hunters during fall 1984. Observations of marked pronghorn are much less extensive than in the northern unit. Wheat fields near grasslands sively in the central portion of the study area. During pronghorn were observed to move north to the vicinity of of Punkin Center (Fig. 13). in the southern unit were used extenspring 1985, marked Highway 24 and south �129 --~----~------~14~------------------~ ·BRIGGSDALE Gfi>~4S~WO POINT OF • ROCKS WELP CO . °D 21 f:'( MORGAN CO . 47"'S IJ • GREASEWOO~ LAKE I • 18 3 'I'~ MACKSON RES V O(!) Un:: VERSIDE RES. ~ r-~ Fig. 8. H 2km W 5:. I __ ~~ SOUTH 2 mi ~IIs . ~~~ ~\ PL~'""- Locations of trapsites (numerals) used to trap pronghorn during winter in northeastern Colorado. Trapsites 1 and 2 were used during January and February, 1984; and trapsite 3 was used in January, 1985. FT. MORGAN �130 ••• KIOWA • I • • ....,. CALHAN;j ·• D@ • I >"'> rill- z·z I 6:5 A-35 U;U 4g 0-1- 2 W·UJ • . LINCOLN •••_.___ @ ,_., •....• oa u. o! U)- • _ JI COUNTY • IpUNKIN I • ~ - I-111 _ - ELBERT COUNTY - • • >. ~iI ::::l- -- I ..,Ja..,l • • I (J)ao:: «.UJ a. co _<§...l~D._E_R , CENTER 1 A-42 ~n -• uj~ 3 I • I LINCOLN COUNTY .-.-.~--.-.-.-.-.-.CROWLEY COUNTY Fig. 9. Locations of trapsites (num era.l s) used to trap pronghorn during winter in southeastern Colorado. Trapsites 1 and 2 were used during January, 1984; trapsites 3 and 4 were used during February, 1985. �131 ---r----~------~14r-------------------~ x ·BRIGGSDALE x x x G x "f?~4S~w, TRAPSIT~ #;..1 POIN\ OF OOD f:'1.. • ROCKS 47'S .~ 'V 0 x N • GREASEWOOQ LAKE x xX WELP CO (88)MORGAN CO . x • 0 x I x. 18 TRAPSITE #1 U"1 . 'I'~ 0 X o 0 X~IVERSIDE RES, (.!) a::: ~IIs MACKSON RES V . W 5:. ~--~--~--~I~ ~~~~ SOUTH PL~_'(~"- 2 mi H 2 km If) Fig. 10. Locations of recovered neckbands from marked pronghorn in northeastern Colorado. Locations of legally harvested animals are indicated by an X; locations of other mortalities are indicated by O. FT. MORGAN �132 GIY~ 4s~ INT OF • ROCKS I WoO D .-------f:-( WELD CO MORGAN CO. 47's , ~ EASEWOOQ LAKE I , 18 N o- ~ di~ V MACKSON RES RES. . W 3:, I SOUTH I---i 2 mi H 2km r-_ .. Fig. 11. Movements of banded pronghorn March through June, 1985, in .northeastern Colorado; width of each arrow represents relative number of collars observed. A •• �133 x ••• KIOWA x SIM.~~ __ !I x X. x D@ I • • >~> l- I- -L CALHAN I x z·z A-35 6~g u~u -I 0-1- • II • CJ)go::: ~. w a. cc ..Jll_J w W· I x ! ELBERT COUNTY r:. __ • LI NCOLN COU NTY • I 'C'-!---------~ , :3 YODER -Q94~. • • _ 1:11 • 13 •••• Q • • 7 PUNKIN: CENTER I >1 _ • _ ~ x 1-- z~ =:>::1 O~ x u. H 1 mi H 1 km Oa CJ). c:a• A-42 ~:l • I • ._._._.-._.-.-.-._.I • Fig. 12. LINCOLN COUNTY CROWLEY COUNTY Locations of recovered neckbands from marked pronghorn southeastern Colorado. Locations of legally harvested animals are indicated by_an X. in �134 •• I • I • I •• I •• ... B@ • II : >9> ;- I- •• z·z A-35 :=l~:=l 0110 USU oal- (/)gc: ~.~ ..J1.1....J UJ·UJ I ~ YODER -'e;J __ •••• • ELBERT • :;a •• _ • - • • . LINCOLN II @l COUNTY • _ •• _ • _ • _ I « COUNTY • rpUNKIN I CENTER •• >1 1-- z!i :=l- og U. 01 (/). A-42~ ~a • ~:I ~ •• I .-.-.~.. -.-.- .. -.-.- .. • I LINCOLN COUNTY CROWLEY COUNTY Fig. 13. Movements of banded pronghorn February through June, 1985, southeastern Colorado; width of arrow represents relative numbers of collars observed. �135 RADIO TELE~lETRY Methods Five pronghorn (4 males, 1 female) were monitored in the northern area during summer and fall 1984. Our objective was to determine summer and fall habitat use by pronghorn in the northern unit. A total of 17 pronghorn (7 males, 10 females) were equipped with radio transmitters by February, 1985. Winter habitat use by pronghorn was monitored at least bimonthly during 1984-85 winter to determine grazing pressure on wheat by pronghorn and duration of grazing. Three radio collars were equipped with activity sensors that allowed us to differentiate foraging from other activities. During March, 1985, activity collars were monitored on 8 occasions for 18 hrs each to determine pronghorn foraging activities at night. Requirements for collection included: visual confirmation of "head-down", "head-up" signal with each radio; pronghorn occupying wheat continually during darkness; and clear, calm weather. Radio-equipped animals were monitored at 20-min intervals during 12 hrs of darkness. Results During summer and fall 1984, radio-equipped pronghorn occupied 2 different habitat types. Three of the 5 active radios occupied large expanses of grassland (13 km2 or larger) during summer and fall. Two adult males occupied small areas (5 km2) of grassland that were entirely surrounded by cropland. All radio-equipped pronghorn interacted with other pronghorn during the rut and had abandoned grassl and to forage on wheat by midNovember, 1984. radio-equipped pronghorn were killed during fall 1984. One female was legally harvested, and 1 adult male was killed illegally on a wheat field north of Riverside Reservoir. T\'IO Fourteen radio collars were placed on pronghorn in January, 1985, bringing the total of instrumented pronghorn to 17. Eleven of these radio-equipped pronghorn occupied a single wheat field from January through r~arch, 1985, along with 120 noninstrumented pronghorn (Fig. 14). The remaining 6 radioinstrumented animals utilized large wheat fields directly north of Riverside Reservoir, occasionally retreating westward to bed and forage on native pasture. The wheat field occupied by the 11 radioed animals is surrounded by grassland, and pronghorn moved easily between both habitats, foraging and bedding in both. An aerial search for radio-equipped pronghorn on 19 March 1985 showed that pronghorn were beginning to abandon winter concentration areas. Two adult males had returned to summer territories by that date. The large concentration of pronghorn on the isolated wheat field had abandoned the field by mid-April. Six radio-equipped pronghorn continued to use grasslands within 5 km of that isolated wheat field throughout summer 1985. The remainder of the radio-equipped pronghorn moved south an west to summer (Fig. 14). Data was successfully gathered during 3 of the 8 attempts to gather night foraging behavior by pronghorn. Radio-equipped pronghorn were observed to �136 --~----~------~14r------------------__ -BRIGGSDALE QIY~ 4S~w. POINT OF -ROCKS I OOD ;:::-(4 Q ts MORGAN CO. I -+ ~GREASEWOO~ .t--RIGH , / E WELP CO I LAKE I 18 -II~ 0(9 . . ~ 00:: ::1' ~ MACKSON RES V . VERSIDE RES, W ~I ~----~~--~I ~~~~ SOUTH PLr\\~ 2 mi H 2 km FT. MORGAN Fig. 14. Location of single wheat field (HIGH USE) on which 11 radio-collared pronghorn foraged from January through March, 1985. Approximately 120 additional pronghorn also utilized this wheat field from January through March, 1985. Movements to summer ranges of telemetered pronghorn are indicated by arrows. Width of each arrow represents relative number of pronghorn moving to summer range. �137 bed on wheat at sunset on these 3 occasions, and 2 adult males were monitored. Changes in impulse frequency indicated that both males foraged on wheat for short periods. Foraging bouts occurred after midnight and peaked at 1:00 am and at 3:00 am. Foraging episodes were short, lasting (x + s) 28 + 7 mins (n = 8). These data confirm that pronghorn forage on wheat after sunset but cannot quantify extent of nighttime foraging. An attempt will be made during the next segment to monitor nighttime pronghorn foraging behavior. SUMMARY Results presented to date in progress reports are for intenal use only within Colorado State University and Colorado Division of Wildlife. The third and final year of the pronghorn-wheat project is upcoming, and important replications of experiments will be performed. Therefore, until the final report is composed, care must be exercised in interpreting these preliminary data. All subprojects must be completed and integrated before results of the pronghorn-wheat project are released to the general public. To date, no effects (positive or negative) of pronghorn grazing of wheat during winter have been demonstrated. Total grain yield and total final biomass have not been significantly affected by pronghorn grazing. Experimental stocking rates of 1 pronghorn/0.6 ha for 172 days is heavier use of wheat than has been documented for wild pronghorn in eastern Colorado. Radio-telemetry studies have shown that pronghorn do not forage exclusively on wheat during winter but also move to grassland sites to forage. Aerial surveys and radio telemetry studies indicate that pronghorn occupy wheat fields from mid-November through April. Above-ground biomass is an appropriate index of pronghorn grazing pressure, but apparently is not indicative of final grain yield. LITERATURE CITED CSU Progress Report #1. July, 1984. Colo. State Univ., Fort Collins. Dep. Fishery and Wildlife Biology. 20pp. Dickinson, C. E., and J. L. Dodd. 1976. Phenological pattern in the shortgrass prairie. Am. Midl. Nat. 96:367-378. Gill, R. B. 1984. Big game forage selection dynamics. Game Res. Rep. July Part 2:133-137. Colo. Div. Wild1. Goering, H. K., and P. J. Van Soest. 1971. Forage fiber analysis. (Apparatus, reagents, procedures, and some applications). U.S.D.A. Agr. Res. Servo Ag. Handbook. No. 379. 20pp. Hobbs, N. T., D. L. Baker, J. E. Ellis, and D. M. Swift. 1981. Composition and quality of elk winter diets in Colorado. J. Wi1d1. Mange. 45:156-171. Horwitz, W. (ed.). 1980. Official methods of analysis of the Association of Official Analytical Chemists. Association of Official Analytical Chemists. Washington, D.C. 1018pp. �138 Laycock, W. A. 1962. Range research methods. Pub. No. 940. U.S.D.A. For. Servo r·1isc. Range Research r'lethodsSubcommittee. 1962. Basic problems and techniques in range research. Natl. Acad. Sci. Pub. No. 890. Snedecor, G. W., and W. G. Cochran. Univ. Press. Ames. 593pp. R. B. Gill 1971. Statistical methods. Iowa State �139 APPENDIX I Scale used to describe phenology of native grassland vegetation during winter and spring, 1985, in northeast Colorado after Dickinson and Dodd (1976). Description Stage Winter dormancy 1 First visible growth 2 First leaves fully expanded 3 Middle leaves fully visible 4 Middle leaves fully expanded 5 Late leaves fully expanded 6 First floral buds 7 Mature floral buds 8 Floral buds and open flowers 9 ��Colorado Division of Wildlife Wildlife Research Report July 1985 141 . JOB PROGRESS REPORT State of Colorado Project No. 01-03-048 Mammals 2 Research Work Plan No. Bighorn Sheep Investigations ------------------2 ----------------- Job. No. 4 Period covered~ July 1,1984 Authors: Personnel: Plane of Nutrition and Bighorn Sheep Population Performance - June 30,1985 N. T. Hobbs, M. W. Miller L. L. Miller, W. H. Rutherford, T. R. Spraker, G. G. Schoonve1d ABSTRACT Pilot studies revealed large interanima1 variation in physiological responses to acute stress as indicated by blood cortisol assay. Despite this variation, chronically stressed bighorn sheep showed higher maximum cortisol production in response to acute stress than we observed in unstressed bighorns (P < 0.0001). Maximum cortisol production increased with time in chronically stressed animals (P < 0.0001). Injectible ivermectin proved to be an efficacious treatment for lungworm infections in bighorn sheep. Larval output of treated animals was reduced to 0.0 within 1 month of dosing treated animals; larval output of control animals remained unchanged during that period. Study plans were prepared to investigate relationships between nutritional plane, stress, and disease resistance in bighorn sheep. ��143 PLANE OF NUTRITION AND BIGHORN SHEEP POPULATION PERFORMANCE P. N. OBJECTIVES Same as Segment Objectives. SEGMENT OBJECTIVES 1. Prepare a detailed study plan to investigate relationships between nutritional plane and parasite resistance in bighorn sheep. 2. Conduct pilot experiments to determine the most effective method to challenge bighorn sheep with Protostrongylus sp. larvae. 3. Rear and train bighorn sheep lambs for controlled nutrition-parasite experiments during 1985-86. RESULTS AND DISCUSSION We prepared a detailed study plan describing future experiments on identifying and managing environmental stress in bighorn sheep (Appendix A). We conducted experiments on methods for treating lungworm infections in bighorn sheep (Appendix B) and planned future work in this area (Appendix C). We reviewed literature and undertook preliminary investigations of the stress-disease complex in bighorn sheep (Appendix D). Prepared by u:»vl ~;1 TTL~I N. iiompson Aob6s Wildlife Researcher Ml&1.rr.~~ Graduate Student ��145 APPENDIX A Preliminary draft of study plan describing experiments on identifying and managing stress in bighorn sheep. STUDY PLAN State of Colorado Project No. 01-03-048-15080 Mammals 2 Research Work Plan No. Bighorn Sheep Investigations Job. No. A. 2 ----------------4 Experiments on Identifying and Managing Stress in Bighorn Sheep NEED A fundamental consideration in managing Rocky Mountain bighorn sheep is the role of infectious disease in their population dynamics. Throughout North America, mortality resulting from bacterial bronchopneumonia appears to limit the abundance of bighorns (Buechner 1960, Forrester 1971, Bear and Jones 1973, Feuerstein et al. 1980, Spraker and Hibler 1982, Proc. Bighorn Sheep Dieoff Workshop, unpubl. rep.). Parasitic lun~/orms may contribute to onset and severity of pneumonia in bighorns, but are not essential to pathogenesis of the pneumonia complex (Forrester 1971, Spraker and Hibler 1982, Spraker et al. 1984b). Pneumonia outbreaks in bighorn herds occur as acute epizootics, often during the fall or winter months. These infections affect all age groups, and may markedly reduce populations (March 1930, Buechner 1960, Spraker and Hibler 1982, Spraker et al. 1984b). The inability to predict or control (lungworm treatment program). These are among the most intensive management practices for any game species in the state. Colorado's bighorn populations have responded by more than doubling estimated numbers (2,000 to 5,000 head) in 6 years (Spraker and Schmidt 1983). Despite intensive management, Colorado's bighorn herds (like other herds throughout North America) continue to experience periodic, precipitous declines associated with pneumonia epizootics. Since 1979, acute outbreaks of pneumonia have been documented in 4 bighorn herds (Taylor River, Waterton Canyon, Little Hills, Ouray) (Feuerstein et al. 1980; Spraker et al. 1984b; Colorado Division of Wildlife Research Center, unpubl. data; T. R. Spraker, pers. comm.); mortalities were invariably attributed to bacterial bronchopneumonia with or without a verminous pneumonia outbreaks continually limits the success of bighorn management. Programs for managing bighorn sheep in Colorado currently use some combination of habitat preservation, population control (limited hunting, trapping), range extension (transplanting to historical ranges), habitat improvement (controlled burning), and winter feeding/parasite control (lungworm) component. Evidence exists for similar declines in at least 3 additional herds (Ford Creek, Upper Poudre Canyon, Lone Pine) (R. L. Schmidt, pers. comm.). Consequently, understanding the conditions leading to these pneumonia epizootics is the next step in developing a successful management program for bighorns. �146 Central to most contemporary hypotheses on susceptibility of bighorn populations to pneumonia outbreaks is the role of chronic environmental stress as "a dominant influence on the dynamics of bighorn sheep populations and on the initiation and continuation of disease outbreaks" (Proc. Bighorn Sheep Dieoff Workshop, unpubl. rep. 11:15). Experts agree that developing techniques for detecting and quantifying the effects of chronic stress on free-ranging populations is vital to future success of bighorn management (Proc. Bighorn Sheep Dieoff Workshop, unpubl. rep.). The hypothesized role of chronic stress in the pneumonia complex of bighorn sheep is based on concepts derived from research using several animal models. Stress (Selye 1950, 1975) is defined as a state requiring abnormal or extreme adjustments in physiology or behavior in order to cope with adversities of environment or management (stressors) (Fraser et al. 1975). This generalized response to an environmental stressor (reviewed by Stephens 1980:180-182) is initiated by impulses conducted from the cerebral cortex to the hypothalamus (integrator of emotion and physiological function), stimulating the autonomic nervous system. The adrenal gland responds to autonomic stimulation by incresing secretion of epinephrine and norepinephrine into the blood. These endocrine events mobilize body resources to facilitate heightened activity levels during "fight or flight." If exposure to the stressor persists beyond this immediate, emergency phase, the animal secretes corticotrophin-releasing factor from the hypothalamus, which stimulates release of adrenocorticotrophic hormone (ACTH) from the pituitary. Increased ACTH levels, in turn, cause the adrenal cortex to secrete corticosteroid hormones (cortisol), further mobilizing body resources needed for activity. The stress response evolved as a protective mechanism enhancing survival (Selye 1950, 1975; Fraser et al. 1975), but chronic stress leads to a variety of physiological dysfunctions that adversely affect survival. These include adrenal hyperfunction/hyperplasia and elevated blood glucocorticoids, decreased vigor, impaired reproductive performance, and immune system failure (Ganjam et al. 1972, Petropoulos et al. 1972, Weiss 1972, Stein et al. 1976, Trindle et al. 1978, Herrenkohl 1979, Kelley 1980, Stephens 1980, Riley, 1981, Dantzer and Mormede 1983, Stoskopf 1983)• Immunosuppression resulting from chronic environmental stress increases susceptibility to infectious agents, implicating stress as a common component of several disease syndromes of domestic animals (reviewed by Kelley 1980). Both corticosteroid-dependent (reviewed by Baxter and Harris 1975) and -independent (Keller et al. 1983) mechanisms for immunosuppression have been observed. Effects of corticosteroiddependent mechanisms appear more significant; inhibitory activities of 9lucocorticoids have been described for virtually every aspect of the lmmune response (Baxter and Harris 1975). One key immunosuppressive activity of corticosteroids is depression of lymphocyte function (Nd4aster and Franzi 1 1961; Hudson 1972, 1973; Zeman et al. 1972; Bach et al. 1975; Baxter and Harris 1975; Butler 1975; Fauci 1975; Kelley 1980; Riley 1981; Stoskopf 1983). Lymphocyte populations are largely responsible for both cellular and humoral (antibody) aspects of the immune response (Tizard 1982). Glucocorticoids preferentially depress function of T-lymphocyte precursors (cellular immunity) and activated �147 B-lymphocytes (humoral immunity) (McMaster and Franzil 1961, Bach et al. 1975, Baxter and Harris, 1975, Butler 1975, Fauci 1975, Kelley 1980, Riley 1981), cause reduction of circulating lymphocytes (Fauci 1975), and degeneration of lymph and thymic tissues (McMaster and Franzil 1961, Weis 1972, Baxter and Harris 1975, Hartman et al. 1976, Stein et al. 1976, Kelley 1980, Stephens 1980, Riley 1981). Indications of lymphocyte dysfunctions secondary to chronic stress have been reported for ungulate species (reviewed by Kelly 1980, Stephens 1980), including bighorn sheep (Hudson 1971,1973; Spraker and Hibler 1982; Proc. Bighorn Sheep Dieoff Workshop, unpubl. resp.). Investigations are underway in Colorado (Appendix D) and Wyoming (E. S. Williams, pers. comm.) to demonstrate the relationship between chronic stress and immune failure in bighorns. Thus, corticosteroid-induced immunosuppression and accompanying increased susceptibility to infectious agents provide a plausible mechanism supporting the hypothesis that chronic environmental stress predisposes bighorns to pneumonia, and that the cumulative effects of chronic stress are ultimately responsible for the mortality that limits abundance of bighorns. It follows that the ability to detect and quantify the effects of chronic stress in bighorns is essential in predicting pneumonia outbreaks in bighorn populations, and in identifying key environmental stressors that can be manipulated through management to reduce these stress levels. Here, we propose to develop techniques to detect and quantify chronic stress in bighorns, and to evaluate the sensitivity of bighorn sheep to commonly encountered environmental stressors. B. EXPECTED RESULTS AND BENEFITS The experiments we propose will provide a foundation for laboratory and field investigations of the stress-pneumonia complex of bighorn sheep. Initially, we will determine if physiological responses to chronic stress can be detected and quantified reliably in bighorn sheep. We will examine and compare urine, fecal, and serum cortisol concentrations under experimental conditions, and correlate these with indicators of immune competence. If these measures prove reliable, we will use them in additional experiments testing specific hypotheses on the role of individual stressors (plane of nutrition, population density) in contributing to stress-induced immune failure in bighorn sheep. Our view of the role of this research in providing future benefits to management of bighorn sheep is illustrated in Figure 1. C. OBJECTIVES The specific objectives of our work include: 1. Determine if adrenal responses to acute stress can be used as a reliable indicator of chronic stress. 2. Correlate the magnitude of adrenal responses to acute stress with measures of immune competence in bighorn sheep. �148 3. Experimentally compare the utility of blood, urine, and fecal cortisol concentrations as indicators of exposure to chronic stress in bighorn sheep. 4. Determine the magnitude of adrenal responses in bighorns to malnutrition and subsequent dietary improvement. 5. Determine magnitude of adrenal responses to chronic exposure to a high population density and subsequent reduction of density. D. APPROACH Detection and Quantification of Adrenal Responses to Chronic Stress in Bighorns Overview: The detrimental effects of chronic stress on mammalian systems are well-documented (Selye 1975, Kelley 1980, Stephens 1980, Dantzer and Mormede 1983, Stoskopf 1983); among these, immunosuppression links chronic stress to disease syndromes of domestic (reviewed by Kelley 1980) and wild (Fiennes 1982, Stoskopf 1983) animals, including the pneumonia complex of bighorn sheep (Proc. Bighorn Sheep Dieoff Workshop, unpul. rep.). Most previous research has concentrated on observations of acute stress (reviewed by Stephens 1980), with few experimenters attempting to objectively measure the physiological effects of chronic stress (Dantzer and Mormede 1983). Adrenocortical hormone concentrations in blood and excreta appear to be the most promising indicators of chronic adrenal stimulation and activity (Beisel et al. 1964, Perry 1973, Stephens 1980), although interpretation of these responses must be made with care (Perry 1973, Dantzer and Mormede 1983). Our initial investigations will determine reliability of serum, urine, and fecal cortisol concentrations in detecting adrenal responses to experimentally-imposed chronic stress in bighorn sheep. The methods developed in these studies are critical to any further field or laboratory investigations of environmental stressors affecting bighorn populations. Experiment 1.1: Comparison of serum cortisol concentrations in response to exogenous ACTH administration and restraint in bighorn sheep. Hypothesis: Adrenal responses of bighorn sheep to (restraint) and exogenous ACTH administration will as measured by slope and asymptote of the cortisol serum cortisol concentration (ng/m1) vs time (min) acute stress be equivalent response curve: (Fig. 2). Rationale: Evidence' exists for use of maximum cortisol concentrations ("cortisol plateau") in blood as an indicator of past exposure to chronic stress (Friend et a1. 1977,1979; Fulkerson and Jameson 1982; Dantzer and Mormede 1983; Sap10sky 1983; Spraker et al. 1984). In domestic sheep, maximum cortisol concentrations attained 10-30 mins. following exposure to an acute stressor (shearing, yearding) did not differ from those observed following exogenous ACTH administered as an intravenous bolus (Moberg et a1. 1980, Fulkerson and �149 Jamieson 1982); however, chronically stressed baboons responded to restraint with higher cortisol plateaus than to exogenous ACTH administration (Saplosky 1983). Manual restraint is currently used as an acute stress to elicit maximum cortisol responses in bighorns (Appendix 0, Spraker et ale 1984a), but variability in response to similar types of restraint, both within and between individuals, has been observed (Appendix D). Therefore, we need to determine whether manual restraint is superior or equivalent to exogenous ACTH in producing maximum cortisol responses in our tame bighorns; these results will establish the most consistent and reliable method for obtaining maximum serum cortisol concentrations to be used in comparison with urine and fecal cortisol concentrations as indicators of chronic stress in bighorns. Methods: We will compare maximum serum cortisol responses of bighorn sheep to restraint and exogenous ACTH administration using 5 sheep in a 2-way crossover design. On day one, 2 sheep will be caught and restrained in lateral recumbency with leg hobbles, the other 3 will be caught, administered 20 U ACTH intravenously (N), and placed in small (about 20-m ) isolation pens. Blood will be drawn from indwelling jugular catheters at 0, 7, 15, 30, 45, and 60 mins. post-treatment, collected in clot tubes, and serum harvested and stored at -700C. Seven days later, treatment groups will be reversed and the experiment repeated. Serum cortisol concentrations will be determined by radioimmunoassay (RIA) (Hasler et ale 1976). Analysis: We will use Marquardt's least squares regression to fit models to the relationship between serum cortisol concentration (ng/ml) and time (min) after initiation of acute stress (restraint) or exogenous ACTH administration. We hypothesize that a natural growth function Y = a(l-e-bx) will represent that relationship. We will use the parameter describing the asymptote (a) to estimate maximum cortisol concentration (cortisol plateau). We will analyze differences in maximum cortisol concentrations between restraint and ACTH groups with a paired t-test. We anticipate no significant differences between the 2 treatment methods. Experiment 1.2: Development of sampling techniques for determining urine and fecal cortisol concentrations in bighorn sheep. Hypothesis: Minimal daily variations in urine or fecal cortisol concentrations for individual bighorns will be observed over a 3-day period. Urine and fecal cortisol concentrations from grab samples will not differ from 24-hr composite samples. Rationale: Remote sampling methods for detecting chronic stress in bighorn sheep are needed for field and laboratory studies. t,1easuring cortisol concentrations in excreta offers the most promising approach to this problem. Free cortisol in urine is a sensitive indicator of adrenal hyperfunction (Beisel et ale 1964), and urine can be readily obtained from habituated research animals. Feces can also be collected easily from captive animals and can be obtained from wild animals in the field as well. There is limited evidence that fecal �150 cortisol concentrations may indicate chronic stress in domestic and bighorn sheep (E. Mock, pers. comm.). The first step in designing experiments to assess the reliability of these indicators is development of techniques for sampling and processing urine and feces for cortisol assays. Further, we need some indication of the variability encountered within these samples to assure adequate sampling in future studies. Methods: Urine and fecal samples will be collected on 3 consecutive days from 3 bighorns housed in individual metabolic cages (about 20 m ) after thorough habituation and a 24-hr acclimation period. Random voided urine and fecal samples will be collected hourly from 8 AM to 6 PM from individuals, as will 24-hr composite urine and fecal samples. All samples will be labeled and stored in appropriate containers at -700C. Cortisol concentrations will be determined by RIA following an extraction process currently under development (E. Mock, pers comm.). Urine creatinine concentrations will be determined by colorimetric methods (Caraway 1960) as an index of urine production per unit time, and urine cortisol will be expressed as cortisol:creatinine ratio (Berman et al. 1980). Fecal cortisol concentrations will be expressed as ng/g dry matter. Analysis: We will use netted analysis of variance to estimate the number of animals and number of fecal and urine grab samples per animal necessary to provide a reliable (+ 20% of mean, 90% of the time) estimate of the true mean cortisol-concentration in urine and feces. Experiment 1.3: Tests of serum, urine, and fecal cortisol concentrations as reliable indicators of chronic stress in bighorn sheep. Hypotheses: Concentrations of cortisol in blood, urine, and feces of chronically stressed bighorn sheep will be higher than those concentrations in "unstressed" bighorns. There will be no difference in coefficients of variation among cortisol concentrations in blood, urine, and feces. Rationale: In order to detect and alleviate cumulative environmental stress affecting bighorn populations, wildlife managers need a reliable measure of chronic stress. Behavioral parameters have been used to measure stress in domestic animals (reviewed by Stephens 1980), but behavior may not accurately reflect stress responses by wild ungulates, including bighorn sheep (Moen 1973, Freddy et al. 1979, Moen and Chevalier 1977, Stemp 1983, Stoskopf 1983). Physiological measures have included assays of adrenal response, as indicated by glucocorticoid (cortisol) levels in blood, to assessstress in domestic (reviewed by Stephens 1980) and wildlife species (Franzmann et al. 1975, Kirkpatrick et al. 1979, Wesson et al. 1979, Saplosky 1983, Stoskopf 1983, Spraker et al. 1984a). The relative stability of cortisol in plasma or serum (72h at 40C) (Reimers et al. 1983), and the development of a sensitive and specific radioimmunoassay for cortisol (Hasler et al. 1976), make cortisol assays adaptable to laboratory and field studies of stress in nondomestic animals. �151 Cortisol is secreted by the adrenal gland in response to noxious stimuli; stressful events increase blood corticoid levels, which return to normal after the stress is removed. Chronic exposure to a stressor causes adrenal hyperfunction and hyperplasia, thereby increasing the adrenal's capacity to secrete cortisol. The resulting elevations in serum cortisol concentrations at baseline (resting) levels and in response to persistent acute stress have been used as indicators of chronic stress (Jensen and Rasmussen 1970; Willett and Erb 1972; Halley et al. 1975; Johnson and Vanjonack 1976; Friend et al. 1977, 1979; Trenkle 1978; Burchfield et al. 1980; Stephens 1980; Fulkerson and Jamieson 1982; Dantzer and Mormede 1983; Saplosky 1983). Variations in baseline cortisol levels within and between individuals (Reid and Mills 1962, Jensen and Rasmussen 1970, Wagner and Oxenrider 1972, Willett and Erb 1972, McNatty and Thurley 1973, Holley et al. 1975, Arave et a1. 1977, Trenkle 1978, Kattesh et al. 1980), compounded with diurnal effects (McNatty and Thurley 1973, Perry 1973, Holley et al. 1975, Turner 1984) and influences of sampling procedures (Holub et al. 1959, Bassett and Hinks 1969, Garner et al. 1980, Fulkerson and Jamieson 1982, Saplosky 1983), limit use and interpretation of these values for studying stress in wild animals. Alternatively, maximum cortisol production in response to persistent acute stress has been used as an indicator of chronic stress (Friend et al. 1977, 1979; Fulkerson and Jamieson 1982; Dantzer and Mormede 1983; Saplosky 1983; Spraker et al. 1984a, Appendix D), with maximum cortisol concentrations (cortisol plateau) in chronically stressed animals reaching higher levels than those in controls. Using this method, maximum cortisol output overrides variations caused by sampling and temporal fluctuations. These are compelling arguments for using maximum cortisol concentrations to study chronic stress in wildlife species. Preliminary results of ongoing research indicate that maximum cortisol data may reflect exposure to chronic stress in bighorns (Spraker et al. 1984a, Hobbs et a1. Appendix D). Unfortunately, there are several limitations in using maximum serum cortisol values in bighorns. In the laboratory, the 15-30 mins. of restraint required to achieve maximum cortisol concentrations may harm experimental animals and, consequently, limits sampling intervals. In the field, the obvious requirements for obtaining blood from free-ranging animals limits sampling to winter months when sheep can be concentrated at bait stations and trapped; circumstances surrounding the baiting and trapping may contribute to environmental stress perceived by baited sheep, making them poor representatives of overall herd status. Logistically, it is difficult to sample several herds in a short enough timespan to eliminate temporal and climatic variations when comparing herd data. Therefore, we need an indicator of chronic stress than can be sampled remotely; this would allow frequent, unbiased sampling from individuals, without handling, in the field or laboratory. Measuring cortisol concentrations in excreta (urine, feces, saliva) offers an alternative to serum cortisol sampling. Metabolic clearance of cortisol into urine and feces should reflect accumulated cortisol secretion over time, thus making urine and fecal cortisol �152 concentrations less sensitive to temporal variation than blood sampling. Cortisol. is inactivated, mainly by the liver, through reduction and subsequent conjugation with glucuronic acid. Conjugated cortisol is rapidly excreted by the kidney (Brooks 1979). However, a fraction of total cortisol (0.5%) is passively excreted unchanged in the urine, and this free cortisol is directly related to free cortisol concentrations in plasma, and thus to adrenal function (Deise1 et a1. 1964, Brooks 1979). Interspecific variation in cortisol metabolism and excretion has been reported (Taylor and Scratcherd 1963, Kine 1975), but urinary cortisol excretion has been shown to increase in response to confinement stress in domestic sheep (Berman et a1. 1980), indicating potential for use of urine cortisol in detecting chronic stress in bighorns. Although urine sampling is no more useful than blood sampling in the field, laboratory experiments involving tame, approachable bighorns are ideal conditions for applying urine cortisol assays in studies of chronic stress. Our capability for 24-hr urine collections further enhances the integrating properties of urine cortisol data (Brooks 1979). Presence of cortisol in feces probably relates to the high lipid solubility of this molecule, which allows passive diffusion across cell membranes down concentration gradients. Biliary excretion of cortisol metabolites does occur and is highest in cats and rodents (Taylor and Scratcherd 1963). However, no data on this phenomenon have been reported for ruminants. If free cortisol concentrations in feces result from passive diffusion, then fecal cortisol may serve as an indicator of plasma cortisol concentrations. Preliminary evidence suggests that fecal cortisol may indicate exposure to chronic stress in domestic and bighorn sheep (E. Mock, pers. comm.), and the potential for applying this assay to monitoring stress in free-ranging bighorn populations warrants further investigation. Here we will evaluate blood, urine, and fecal cortisol concentrations as indicators of responses of bighorns to experimentally-imposed chronic stress. Low doses of repositol ACTH will be used to stimulate adrenal activity, simulating low-level chronic stress (Ganjam et ale 1972, Fulkerson and Jamieson 1982); in this way, we hope to uniformly impose a controlled, known level of stress across a treatment group, and compare it to an untreated control. Methods: We will examine the reliability of urine cortisol concentrations (UCC) and fecal cortisol concentrations (FCC) as indicators of adrenocortical function and serum cortisol concentrations using 10 bighorn sheep in a blocked, 2-way crossover design. The 10 sheep will be assigned to 5 pairs using sex, age, body size, and serum cortisol concentrations as criteria for pairing. One animal from each pair will be randomly assigned the chronic stress treatment. The other will be an untreated control. Treatment animals will receive ACTH gel (20 U subcutaneously) on alternate days to simulate chronic stress in stimulating adrenal activity; controls will remain untreated. All animals will be housed individually in bare-ground isolation pens (about 50 m ); alfalfa hay, pe11eted concentrate, and water will be provided ad libitum. On day 31, treatment and control assignments will be reversed, and the experiment repeated. �153 Samples for cortisol determination will be collected from all animals on day 0, 15, 30, 45, and 60. All sheep will be housed individually in metabolic cages (about 10 m ) beginning 24 hrs. prior to sample collection. After a 24-hr acclimation period, urine and feces will be collected for 24 hrs and composited for individual animals. Immediately following urine and fecal collections, blood will be obtained for serum cortisol determination via indwelling jugular catheters at 0, 7, 15, 30, and 45 mins; use of restraint or ACTH stimulation to obtain maximum cortisol response will be determined by results of Experiment 1.1. Blood (10 ml) will be collected into 10-ml clot tubes and serum harvested within 12 hrs. Urine, fecal, and serum samples will be stored at -700C until processed. Cortisol concentrations will be determined by RIA; creatinine concentrations will be determined for urine samples as previously described. Sampling and handling procedures may be altered where indicated by preceeding experiments (1.1,1.2); all animals will be closely monitored throughout the study for cortisol responses and signs of pneumonia or other diseases, and experimental design may be modified by adverse reactions to treatment. Analysis: We will use Marquardt's least squares regression to fit models to relationships between maximum serum, fecal, and urine cortisol concentrations for treatment and control groups. Differences in values of maximum serum, fecal, and urine cortisol concentrations between treatment and control groups will be analyzed using a 2-way factorial analysis of variance for a blocked crossover design with repeated measurements. We will replicate by individual animals and will repeat measurements over sample dates. We anticipate significant main effects for treatment (ACTH) and treatment x time interaction. Experiment 1.4: Correlation of measures of adrenal response with measures of immune function in bighorn sheep. Hypothesis: Immunocompetence of bighorn sheep, as indicated by ability to process complex antigens, will be reduced during prolonged exposure to chronic stress; no reduction in immunocompetence will be observed in control sheep. These effects will correlate with cortisol concentrations in serum, urine, and feces. Rationale: The immunosuppressive effects of chronic stress (reviewed by Kelley 1980) have been implicated in several disease syndromes, including the pneumonia complex of bighorn sheep (Proc. Bighorn Sheep Dieoff Workshop, unpubl. rep.). These effects are primarily glucocorticoid mediated (Baxter and Harris 1975). Evidence of lymphocyte dysfunction has been reported for captive bighorn sheep (Hudson 1972), but little experimental evidence exists to support these observations. Processing of complex antigens (e.g. red blood cells) requires participation of macrophages and both B- and T-lymphocytes (Baxter and Harris 1975, Tizard 1982), and stress-induced suppression of anyone of these cell lines will decrease the immune response to a complex antigen. Therefore, differences between stressed and unstressed �154 bighorn sheep in their production of antibodies in response to a complex antigen should serve as an indicator of stress-induced immune compromise. Methods: We will conduct this experiment in conjunction with Experiment 1.3. On days 15 and 45, all animals will be injected intramuscularly (1M) with 1.0 cc washed canine and feline erythrocytes, respectively. Serum will be retained on day 15, 30, 45, and 60 and submitted for antibody titer determinations. Analysis: We will use least squares regression to fit models to relationships between maximum serum cortisol concentrations and antibody titers for treatment and control groups. Differences in antibody titers between treatment and control groups will be analyzed as described in Experiment 1.3. Investigation 2: Adrenal Responses to Individual Stressors in Bighorn Sheep Overview: Environmental stressors include a variety of stimuli, usually categorized as physical (heat, cold, fasting, pain) or emotional (fear, anticipation) (Selye 1950, 1975; Kelley 1980, Stephens 1980; Dantzer and Mormede 1983, Stoskopf 1983). Emotional stressors have profound effects on domestic (Kelley 1980, Stephens 1980, Dantzer and Mormede 1983) and wild animals (Fiennes 1982, Stoskopf 1983), and appear to have the greater impact on bighorn sheep (Stemp 1983, Proc. Bighorn Sheep Dieoff Workshop, unpub1. rep.). Stressors regarded as particularly significant in predisposing bighorns to pneumonia outbreaks include density of sheep populations, disturbance and harassment by humans, malnutrition, trace mineral deficiencies, adverse weather, parasitism, predation, and interspecific competition (Buechner 1960, Hudson 1972, Feuerstein et a1. 1980, Risenhoover and Bailey 1980, Foreyt and Jessup 1982, Goodson 1982, Spraker and Hibler 1982, Stemp 1983, Spraker et a1. 1984b, Proc. Bighorn Sheep Dieoff Workshop, unpub1. rep.). Individually, these stressors appear surmountable, and selective pressures have favored those bighorns capable of adapting to or coping with them (Geist 1971). However, cumulative effects of multiple stressors have severe physiological consequences (especially immunosuppression) that may exceed the bighorn's ability to cope; the manifestation of these chronic effects appears to be widespread mortality from bronchopneumonia. Although the effects of chronic stress can be severe, they are reversible if the source of persistent stress is removed, or if acclimation is achieved (McNatty and Thurley 1973, Burchfield et a1. 1979, Berman et al. 1980, Dantzer and Mormede 1983, Appendix D). Observed reversal of steroid-induced immunosuppression (McMaster and Franzl 1961; Hudson 1972, 1973) is particularly significant to the concept of managing the stresspneumonia complex in bighorns. If critical sources of environmental stress can be identified and managed, it may be possible to prevent pneumonia outbreaks. To do this, we must first establish the relative significance of environmental stessors; management efforts may subsequently emphasize control of the identified key stressors. In this �155 investigation, we propose to begin evaluating adrenal responses of bighorns to individual stressors. Experiment 2.1: Adrenal Responses to Chronic Malnutrition in Bighorns Hypothesis: Chronic malnutrition in bighorn sheep will elevate cortlsol levels in serum, urine, and feces, as compared to controls fed a high quality diet. Cortisol levels in malnourished animals will decrease in response to improved diet quality. Rationale: Seasonal reduction in forage quality and/or intake represents a potential source of stress for many bighorn populations. Loss of winter ranges to development or plant succession may magnify these effects. The timing of "classical" pneumonia outbreaks implicates change in diet (fall) or malnutrition (winter) as contributing or precipitating factors. Ovine species appear relatively resistant to effects of malnutrition, and studies of stress associated with malnutrition in sheep present conflicting results. Malnutrition, in the form of short-term undernutrition (Thwaites and Edey 1970), chronic underfeeding (Keenan et ale 1968), or fasting (Belonje and VanNiereck 1968), had little or no effect on adrenal weights of domestic sheep. However, fasting or undernourishment did increase cortisol responses to physical (cold) (Panaretto and Ferguson 1969, MacKenzie et ale 1975) or psychological (Reid and Mills 1961) stress, and represents a predisposing factor to cold-induced mortality (Hutchinson 1968, Panaretto 1968, Hutchinson and MacRae 1969).We will examine adrenal reponses of malnourished bighorns compared to those maintained on a high plane of nutrition. Methods: We will investigate effects of plane of nutrition on adrenal responses using a blocked, 2-way crossover design with 5 replications per cell. Animals will be paired and assigned to treatment as described in Experiment 1.3. The control group will receive pelleted rations (Baker and Hobbs 1986) ad libitum. Treated animals will receive the same ration at 50% of the ad libitum rate. Animals will be housed individually in bare-ground isolation pens. They will be weighed weekly. Any animal losing more than 25% of its pre-treatment body weight will be removed from the experiment. After 60 days, diets of animals in the treatment and control groups will be switched. Samples for cortisol determination will be collected and processed as described in Experiment 1.3 on days 0,1,30,45,60, and 120. (Results of Experiment 1.3 may influence our choice of samples collected for cortisol assay.) In addition, we will record body weights weekly for all animals. Analysis: We will analyze differences in cortisol responses between treatment and control groups as described in Experiment 1.3. �156 Experiment 2.2: Adrenal responses to increased population density in bighorns. Hypothesis: Increased population density will elevate cortisol levels in bighorns over time, as compared to cortisol levels of bighorns at a low population density. Cortisol levels in a highdensity population will decline when density is reduced. Rationale: Stress associated with high population density ("crowding") regulates population dynamics of several rodent species (reviewed by Christian and Davis 1964, Andrews 1979). Regulation occurs primarily by glucocorticoid-mediated alterations in reproductive and immune system functions (Christian 1950, Christian and Davis 1964, Archer 1970, Brain 1971, Brayton and Brain 1974, Andrews 1979). Moreover, crowding affects individuals within a population differentially according to social rank (reviewed by Andrews 1979). Social organization, behavioral conditioning, and genetic background all appear to influence thresholds of response to population density in mammal species. Population growth is believed to increase the likelihood of pneumonia outbreaks in bighorns (Feuerstein et ale 1980, Spraker et ale 1984b) because increased density is viewed as a significant source of stress in bighorn populations (Proc. Bighorn Sheep Dieoff Workshop, unpubl. rep.). The gregarious social organization of bighorn is characterized by aggressive encounters (Geist 1971). These interactions, combined with restrictions of movement imposed by plant succession or human encroachment, and the bighorn's own reluctance to pioneer unfamiliar range, may collectively increase the "ecological density" (Bailey 1984:151) bighorns experience. Tolerable density levels appear to vary among populations and may be overestimated by management standards used for other species. Population control, achieved by limited hunting and/or trapping, is a widely accepted strategy for managing bighorn populations. In principle, this strategy differs from practices (range improvement, parasite control) that attempt to increase bighorn numbers on a given range. Wise management of bighorn populations and the habitats they occupy fundamentally depends on the ability to set appropriate goal densities in specific bighorn populations and to alter management strategies in accord with population responses. Here we will examine adrenal responses of bighorns to manipulations of population density over time in an attempt to evaluate the significance of high population density as a source of chronic stress in bighorns. Methods: Adrenal responses to population density will be measured uSlng a blocked, 2-way crossover design with 5 subsamples per cell. We will pair and assign animals to treatment as described in Experiment 1.3. A group of 5-8 additional bighorns will serve as the density treatment. These sheep will be introduced into the treatment pasture to double the treatment population density. �157 Treatment and control groups will be housed in adjacent pastures (about 1/2 hal. Alfalfa hay, pelleted ration, and water will be provided ad libitum for both groups. Animals will be monitored for signs of pneumonia or other disease problems and adverse reactions to treatment may modify experimental design. After 60 days, the density treatment sheep will be removed and introduced into the control pen, thereby switching treatment and control assignments. Samples for cortisol determination will be collected and processed as described in Experiment 1.3 on days 0, 15, 30, 45, 60, 75, 90, 105, and 120. Results of preceeding experiments may influence our choice of samples collected, as animals will not be removed from pastures to metabolic cages during the experiment. Analysis: We will analyze differences in cortisol responses between treatment and control groups as described in Experiment 1.3. We anticipate significant treatment (density) effects and a significant treatment x time interaction. Additional Experiments: Examining adrenal responses of bighorns to plane of nutritlon and population density are only initial steps in a series of potential investigations. Evaluation of other stressors (harassment, visibility, parasitism) and of stressor interactions will add to understanding of the stress-pneumonia complex of bighorns. Schedule October, 1985 - Experiment 1.1,1.2 October-December, 1985 - Experiment 1.3 January-April, 1986 - Analyze Data, Prepare Publication(s) MaY-August, 1986 - Experiment 2.1 September-October, 1986 - Analyze Data November, 1986 - February, 1987 - Experiment 2.2 March-April, 1987 - Analyze Data, Prepare Publication(s) E. LOCATION This study will be conducted in Fort Collins at facilities of the Colorado Division of Wildlife Research Center. F. RELATED FEDERAL PROJECTS This work will complement the ongoing studies of E. T. Thorne, E. S. Williams et al., "Heart rate response and physiologic changes �158 associated with stress in Rocky Mountain bighorn sheep" (Wyo. Wildl. Fed. Aid Proj. No. UJJ ). We are. in continual contact with their research team during our studies. G. LITERATURE CITED Andrews, R. V. 1979. The physiology of crowding. Compo Biochem. Physiol. 63A:1-6. Arave, C. W., C. H. Mickelsen, R. C. Lamb, A. J. Svejda, and R. V. Confield. 1977. Effects of dominance rank changes, age, and body weight on plasma corticoids of mature dairy cattle. J. Dairy Sci. 60:244-248. Archer, J. 1970. Effects of population density on behaviour in rodents. In J. H. Crook, ed. Social Behavior in Birds and Mammals. Academic Press, London and New York. 386pp. Bach, J. F., D. Duval, J. Dardeene, J. C. Salomon, T. 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Stress-induced suppression of immunity in adrenalectomized rats. Science 221:1301-1304. Kelley, K. W. 1980. Stress and immune function: a bibliographic review. Ann. Rec. Vet. 11:445-478. Kirkpatrick, J. F., C. W. Baker, J. W. Turner, R. M. Kenney, and V. K. Ganjam. 1979. Plasma corticosteroids as an index of stress in captive feral horses. J. Wi1d1. Manage. 43:801-804. MacKenzie, A. J., C. J. Thwaites, and T. N. Edey. 1975. Oestrous, ovarian and adrenal response of the ewe to fasting and cold stress. Aust. J. Agric. Res. 26:545-551. Marsh, H. 1938. Pneumoni a in Rocky Mountain bighorn sheep. J. r~amm. 19 :214-219. McMaster, P. D., and R. E. Franzi1. 1961. The effects of adrenocortical steroids upon antibody formation. Metab. 10:990-1005. McNatty, K. P., and D. C. Thurley. 1973. The episodic nature of changes in ovine plasma cortisol levels and their response to adrenaline during adapttion to a new environment. J. Endocr. 59:171-180. Moberg, G., C. O. Anderson, and T. R. Underwood. 1980. Ontogeny of the adrenal and behavioral responses of lambs to emotional stress. J. An. Sci. 51:138-142. . Mock, E. 1985. Personal communication, July. Department of Physiology and Biophysics, Colorado State Univ. Moen, A. N. 1973. Wildlife ecology: an analytical approach. W. H. Freeman and Co., San Francisco. __ , and S. Chevalier. 1977. Analyses of te1emetered ECG signals from white-tailed deer. Pages 118-125 in Proc. Biotelem. Conf., Univ. of Wyoming, Laramie. -- �161 Panaretto, B. A. 1968. Some metabolic effects of cold stress and undernourished non-pregnant ewes. Aust. J. Agric. Res. 19:273-282. __ , and K. A. Ferguson. 1969. Comparison of the effects of several stressing agents on the adrenal glands of normal and hypophysectomized sheep. Aust. J. Agric. Res. 20:115-124. Perry, G. 1973. Can the physiologist measure stress? New Sci. 60:175-177. Petropoulos, E. A., C. Lau, and C. L. Liao. 1972. Neurochemical changes in the offspring of rats subjected to stressful conditions during gestation. Exp. Neurobio1. 37:86-99. Proc. Bighorn Sheep Dieoff Workshop. 1983. Found. N. Amer. Wild Sheep, Cranbrook, B.C. (mimeo) Reid, R. L., and S. C. Mills. 1962. Studies on the carbohydrate metabolism of sheep: XIV. the adrenal response to psychological stress. Aust. J. Agric. Res. 13:282-295. Reimers, T. J., J. P. McCann, and R. G. Cowan. 1983. Effects of storage times and temperatures on T , T , LH, prolactin, insulin, cortisol, and progesterone in blood samples from cows. J. Anim. Sci. 57:683-691. Riley, V. 1981. Psychoneuroendocrine influences on imunocompetence and neoplasia. Science 212:110-1109. Risenhoover, K. L., and J. A. Bailey. 1980. Visibility: an important habitat factor for an indigenous, low-elevation bighorn herd in Colorado. Proc. Biennial Symp. No. Wild Sheep and Goat Counc. 3:18-28. Sapolsky, R. M. 1983. 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Biochem J. 86:114-119. Thwaites, C. J., and T. N. Edey. 1970. The adrenal glands of the ewe. Aust. Vet. J. 46:599-603. Tizard, I. R. 1982. An Introduction to Veterinary Immunology, 2nd ed. W. B. Saunders Co., Philadelphia, PA. 363pp. Trenkle, A. 1978. Relation of hormonal variations to nutritional studies and metabolism of ruminants. J. Dairy Sci. 61:281-293. Trind1e, B. D., L. D. Lewis, and L. H. Laverman. 1978. Evaluation of stress and its effects on the immune system of hand-reared mule deer fawns (Odocoileus hemionus). J. Wild1. Dis. 14:523-537. Turner, J. C. 1984. Diurnal periodicity of plasma cortisol and corticosterone in desert bighorn sheep demonstratd by radioimmunoassay. Can. J. Zool. 62:2659-2665. Wagner, W. C., and S. L. Oxenreider. 1972. Adrenal function in the cow. Diurnal changes and the effects of lactation and neurohyphophyseal hormones. J. An. Sci. 34:630-635. Weis, J. M. 1972. Psychological factors in stress and disease. Sci. Am. 226:104-113. Wesson, J. A., P. F. Scanlon, R. L. Kirkpatrick, H. S. Mosby, and R. L. Butcher. 1979. Influence of chemical immobilization and physical restraint on steroid hormone levels in blood of white-tailed deer. Can. J. Zool. 57:768-776. Willett, L. B., and R. E. Erb. 1972. Short-term changes in plasma corticoids in dairy cattle. J. An. Sci. 34:103-111. Williams, E. S. 1985. Personal communication, June. Wyoming State Veterinary Laboratory. Zeman, G. D., G. Cohen, and M. Budry. 1972. The effects of plasma cortisol levels on lymphocyte transformation tests. J. Allergy Clin, Immunol. 49:10-15. �163 Can physiological responses to stress be reliably measured in bighorn sheep? No Yes Are responses to stress related to immune ~funct;On? End Project Yes End Project No No Are blood indicators correlated with fecal indicators? Yes Develop fie d sampling procedures for estimating stress levels in bi ghorn Figure 1. Proposed research and its application. Does animal density or plane of nu tr t t t on Ll affect responses to stress? Yes Develop population and habitat guidelines for mitigating environmental stressors �164 ?-Cortisol " Plateau ,....... E <; O'l c:: ,....... o VlI or- ....., S- o U -0 o o co ,....... 10 20 30 Time After Initiation of Acute Stress (min) Figure 2. Relationship between blood cortisol levels and time following an acute stress. �165 APPENDIX B Experiments on methods for treating lungworm infections in bighorn sheep. Michael W. Miller Colorado Division of Wildlife Research Center 317 W. Prospect St. Ft. Collins, CO 80526 RH: INJECTABLE IVERMECTIN • Miller et ale EFFICACY OF It~ECTABLE IVERMECTIN FOR TREATING LUNGWORM INFECTIONS IN MOUNTAIN SHEEP MICHAEL W. MILLER. Colorado Division of Wildlife, 317 W. Prospect Street, Fort Collins, CO 80526 N. THOMPSON HOBBS, Colorado Division of Wildlife, 317 W. Prospect Street, Fort Collins, CO 80526 WILLIAM H. RUTHERFORD, Colorado Division of Wildlife, 317 W. Prospect Street, Fort Collins, CO 80526 LISA L. W. MILLER, Department of Animal Sciences, Colorado State University, Fort Collins, CO 80523 Mortality caused by bronchopneumonia limits the abundance of mountain sheep (Ovis canadensis) throughout their North American range (Buechner 1960, Thorne 1982). Infections of parasitic lungworm (Protostron~lus spp.) frequently contribute to development of bronchopneumonia in mounta1n sheep (Forrester 1971, Hibler et ale 1982), and have been associated with elevated mortality in young and adult animals (Spraker and Hibler 1982). Consequently, controlling lungworms with anthelmintic drugs offers an important tool for managing some mountain sheep populations (Hibler et ale 1976, Schmidt et ale 1979). A number of oral anthelmintics, most notably benzimidazoles (albendazole, cambendazole, fenbendazole, thiabendazole), can reduce lungworm infections in mountain sheep (Hibler et ale 1976, Schmidt et ale 1979, Foreyt and Johnson 1980). Fenbendazole drenching is recommended for treating mountain sheep prior to transplant (Hibler et ale 1982). Unfortunately, no single benzimidazole is uniformly effective against all developmental stages of protostron~lus. Fenbendazole is most effective against adult lungworms, cambendazQ(e against stored third-stage larvae (L3) (Hibler et ale 1982). Other disadvantages of these drugs include potential toxicity of cambendazole (Roberson 1982), as well as logistical problems caused by the oral delivery of large volumes of drugs necessary for effective treatment. An alternative treatment is offered by the avermectins, agents with a broad spectrum of activity against internal and external parasites. Although ivermectin (22.23-dihydroavermectin Bl), an injectable �166 avermectin, is safe and effective for treating external parasites in mountain sheep (Kinzer et ale 1983), as well as for treating many parasitic conditions in domestic animals (Egerton et ale 1980, Hotson 1982, Campbell et ale 1983), its value in reducing lungworm in mountain sheep remains undescribed. We evaluated the efficacy of ivermectin for treating lungworm in mountain sheep under field and laboratory conditions. MATERIALS AND METHODS Laboratory Test We observed changes in concentrations of first stage larvae (Ll) in feces of 6 hand-reared mountain sheep (Table 1) in a 2-way factorial experiment with pretreatment observations. We randomly allocated 4 sheep to the treatment group and 2 to the control. Animals were housed individually in bare-ground pens (about 200 m2). Three days prior to treatment, we weighed all animals to the nearest kg and examined them. On 27 April 1983, we injected treatment animals subcutaneously with 0.2 mg/kg ivermectin (10 mg/ml). Treatment and control animals were also injected intramuscularly with about 22,000 iu/kg procaine penicillin G to prevent secondary bacterial infections at injection sites. We observed animals for clinical signs of toxicity (ataxia, paralysis, death) as well as local and systemic side effects (swelling, inflammation, anaphalaxis). Fecal samples from early morning defecations were collected daily from all individuals beginning 14 April 1983 for 14 days prior to treatment and 10 thereafter. Post-treatment samples were then collected on alternate days for the next 6 days. Additional samples were collected on days 21 and 29 post-treatment. We dried all feces for 24 hours at room temperature. Immediately after drying, half of each fecal sample was stored in sealed plastic bags at -20 C for later determination of percent dry matter (Assoc. Off. Anal. Chern. 1980), while the other half was used for larval counts by a modified Baermann technique (Beane and Hobbs 1983). Results were expressed as Ll/g dry matter. We examined the post-treatment change in fecal output of L1 (= weekly average pretreatment - weekly average post-treatment) with a factorial analysis of variance for a fixed-effects model in an unbalanced, blocked design. We predicted significant main effects for treatment and time as well as a significant time x treatment interaction due to time lags in treatment effects. Field Test We observed concentrations of L1 in feces collected from a transplant of wild mountain sheep before and after their treatment with ivermectin. On 19 April 1984, animals were trapped under a drop-net near Baker Mountain in the Never Summer Mountains, Colorado, as part of a routine trapping and transplanting operation. We estimated body weights visually (adult females and yearling rams -- 60-65 kg, lambs -- 30-45 kg), and injected each of 19 animals subcutaneously with 0.5 mg/kg ivermectin (20 mg/ml) (Eqva1an, Merck and Company Inc., Rahway, NJ, 07065, USA.) and intramuscularly with 22,000 iu/kg procaine penicillin G. Sheep were ear tagged, and adult ewes were collared; these animals were subsequently transported to Echo Park in Dinosaur National Monument, Colorado, and released the next day. �167 Immediately after capture, 11 fresh fecal samples were collected from the surface of the snow adjacent to the trap site. Because snow had fallen the previous night, we inferred that all fecal samples were fresh. We counted larvae as described above. Post-treatment samples were collected from sheep in the Echo Park transplant herd during 10 July 1984 to 14 September 1984; samples were collected from transplant sheep either immediately after observing defecations, or from the immediate vicinity of bedding areas after observing bedded sheep thus assuring that field samples were collected from treated animals. Samples were individually bagged and identified by date and collection site and transported to our laboratory. Because all fecal samples were not collected from known individuals, they did not represent independent replications allowing statistical inference based on known degrees of freedom. Consequently, we performed no statistical analysis on field data. RESULTS Laboratory Test Ivermectin sharply reduced Ll output in treated mountain sheep (treatment effect P = 0.004, time effect P = 0.0001, Fig. 1). Ll output of control animals remained unchanged (treatment x time interaction P = 0.0001). Within 4 weeks of treatment, Ll were eliminated in all animals receiving ivermectin. We observed no adverse local or systemic effects of treatment. Field Test Concentrations of Ll in fecal samples collected from wild mountain sheep after treatment with ivermectin were markedly lower than pretreatment. Larvae were recovered from all pretreatment samples, and 7 of 11 samples had concentrations 100 Ll/g feces. In contract, no Ll were recovered from 17 of 23 post-treatment samples (up to 4 months posttreatment); another 5 of 23 post-treatment samples had concentrations 5 Ll/9 feces. DISCUSSION Ivermectin offers an effective means of treating lungworm infections in mountain sheep. Our results resemble findings on efficacy of ivermectin against Dictyocaulus infections of domestic sheep (Wescott and LeaMaster 1982, Yazwinski et al. 1983) and cattle (Alva-Valdes et al. 1984, Benz et al. 1984). It may be possible to reduce lungworm infections more rapidly than we observed. Higher dosages of ivermectin (0.4-1.0 mg/kg) have eliminated L1 output in mountain sheep 7-14 days post-treatment (M. W. Miller, Co. Div. Wild1. unpubl. data). Similarly, elimination of Q. viviparus larvae production in cattle occurred 7 days after ivermectin therapy (0.2 mg/kg) (Benz et a1. 1984). Protostrongylus adults may be somewhat more resistant to the effects of ivermectln than other species of lun~~orm. Others observed variability in efficacy of ivermectin against filarial nematodes �168 and intestinal parasites (Egerton et ala 1980, Campbell 1982, Hotson 1982, Campbell et ala 1983). Safety of ivermectin is well documented. Subcutaneous injection of 30 times the recommended dosage in cattle produced no signs of toxicity, but toxicity and death occurred at 40 times the recommended dose (Campbell et ala 1983). Ivermectin has been used safely in mountain sheep at 1.0 mg/kg (Kinzer et ala 1983, M. W. Miller, Co. Div. Wildl. unpubl. data). Although various adverse effects, mostly minor, have been reported with intramuscular administration of ivermectin in horses (Karns and Luther 1984), only transient injection site irritation has been observed in other domestic species. We have not observed any adverse effects of ivermectin on pregnancy or fetal development in ewes, either in the field or in captivity. Ivermectin offers several advantages for treatment of lungworm during trapping operations. It can be delivered more easily and quickly than drugs requiring drenching. A single dose is broadly effective against external parasites (Kinzer et ala 1983) as well as internal ones, and antiparasitic effects persist for 14 days after administration (Barth 1983). A novel mode of action appears to preclude the development of cross-resistance between ivermectin and other antiparasitic agents, including benzimidazoles, to which resistance has developed (Campbell et ala 1983). No data are available on efficacy of ivermectin against stored L3 Protostrongylus or on transplacental efficacy against in utero lungworm lnfectlons. Immature stages of a number of nematodes are susceptible to the effects of ivermectin, including infective L3 of several species of gastrointestinal parasites of cattle (Barth 1983, Bremner et ala 1983, Swan and Harvey 1983), infective second-stage larvae of Toxocara canis in mice (Abo-Sheda and Herbert 1984), fourth-stage larvae (L4) of D. viviparus in cattle (Alva-Valdes et ala 1984, Benz et ala 1984), arrested or hypobiotic L4 Ostertagia of sheep and cattle (Hotson 1982, Campbell et ala 1983), and migrating Stronfflus vulgaris larvae in horses (Slocombe and McGraw 1981). The high ef lcacy of ivermectin against adult Protostron~lus, in combination with its broad spectrum of activity against a varlety of internal and external paraSites and wide margin of safety, should make this drug a valuable tool in management efforts to transplant mountain sheep to new ranges and in other management and research situations that require parasite control. For these purposes, we recommend treating with 0.2-0.5 mg/kg delivered subcutaneously. ACKNOWLEDGMENTS We thank J. M. Cheney for providing experimental drugs; T. R. Spraker for help in experimental design; R. L. Schmidt and M. A. Bowser for help in the field; C. A. Weinland for counting larvae; S. J. Petersburg, J. F. Hogan, and J. E. Friedlander for collecting Echo Park samples; and L. E. Lovett for assisting with manuscript preparation. D. L. Baker, A. E. Anderson, G. D. Bear, and D. F. Reed offered particularly helpful comments on the manuscript. This research was supported by the Colo. Div. Wildl. Fed. Aid Wildl. Restoration Proj. 45-01-502-15050. �169 LITERATURE CITED Abo-Sheda, M. N., and I. V. Herbert. 1984. Anthelmintic effect of levamisole, ivermectin, albendazole, and fenbendazole on larval Toxocara canis infection in mice. Res. Vet. Sci. 36:87-91. Alva-Valdes, R., G. W. Benz, D. H. Wallace, J. R. Egerton, S. J. Cross, and J. W. Wooden. 1984. Efficacy of ivermectin in oral paste formulation against immature gastro-intestinal and pulmonary nematodes in cattle. Am. J. Vet. Res. 45:685-690. Association of Official Analytical Chemists. 1980. Official methods of analysis, 13th ed. Assoc. Off. Anal. Chern., Washington, D.C. 1018pp. Barth, D. 1983. Persistent anthelmintic effect of ivermectin in cattle Vet. Rec. 113:300. Beane, R. D., and N. T. Hobbs. 1983. The Baermann Technique for estimating Protostrongylus infection in bighorn sheep: effect of laboratory procedures. J. Wildl. Dis. 19:7-9. Benz, G. W., J. V. Ernst, and J. R. Egerton. 1984. Anthelmintic activities of ivermectin against immature and adult Dictyocaulus viviparus. Am. J. Vet. Res. 45:771-772. Bremner, K. C., D. A. Berry, and I. K. Hotson. 1983. Persistence of the anthelmintic activity of ivermectin in calves. Vet. Rec. 113:569. Buechner, H. K. 1960. The bighorn sheep in the United States, its past, present, and future. Wildl. Mono. 4. 174pp. Campbell, W. C. 1982. Efficacy of the avermectins against filarial parasites: a short review. Vet. Res. Comm. 5:251-262. ___~, M. H. Fisher, E. o. Stapley, G. Albers-Schonberg, and T. A. Jacob. 1983. Ivermectin: a potent new antiparasitic drug. Science 221:823-828. Egerton, J. R., et ale 1980. 22,23-dihydroavermectin Bl, a new broadspectrum antiparasitic agent. Br. Vet. J. 136:88-97. Foreyt. B., and R. Johnson. 1980. Treatment for lungworms (Protostrongylus sp.) in Rocky Mountain bighorn sheep (Ovis c. canadensis) with albendazole. Proc. Bienn. Symp. North~a Sheep and Goat Counc., 2:248-259. Forrester, D. J. 1971. Bighorn sheep lungworm-pneumonia complex. Pages 158-173 in J. W. Davis and R. C. Anderson, eds. Parasitic diseases of wild mammals. Iowa State Univ. Press, Ames. 364pp. Hibler, C. P., T. R. Spraker, and R. L. Schmidt. 1976. Treatment of bighorn sheep for lungworm. Proc. Bienn. Symp. North. Wild Sheep Counc., pp. 35-39. ___~' , and E. T. Thorne. 1982. Protostron~ylosis in bighorn sheep. Pages 208-213 in E. T. Thorne, et al., eds. 01seases of wildlife in Wyoming, 2nd ear. Wyo. Game and Fish Dep., Cheyenne. Hotson, I. K. 1982. The avermectins: a new family of antiparasitic agents. J. South Afr. Vet. Assoc. 53:87-90. Karms, P. A., and D. G. Luther. 1984. A survey of adverse effects associated with ivermectin use in Louisiana horses. J. Am. Vet. Med. Assoc. 185:782-783. Kinzer, H. G., W. P. Meleney, R. E. Lange, Jr., and W. E. Houghton. 1983. Preliminary evaluation of ivermectin for control of Psoroptes ovis in desert bighorn sheep. J. Wildl. Dis. 19:52-54. Roberson, E. L. 1982. Antinematodal drugs. Pages 808-818 in N. H. Booth and L. E. McDonald, eds. Veterinary pharmacology and therapeutics. 5th ed. Iowa State Univ. Press, Ames. �170 Schmidt, R. L., C. P. Hibler, T. R. Spraker, and W. H. Rutherford. 1979. An evaluation of drug treatment for lungworm in bighorn sheep. J. Wildl. Manage. 43:461-467. Slocombe, J. O. D., and B. M. McGraw. 1981. Controlled tests of ivermectin against migrating Strongylus vulgaris in ponies. Am. J. Vet. Res. 42:1050-1051. Spraker, T. R., and C. P. Hibler. 1982. An overview of the clinical signs, gross and histological lesions of the pneumonia complex of bighorn sheep. Proc. Bienn. Symp. North. Wild Sheep and Goat Counc. 3:163-172. Swan, G. E., and R. G. Harvey. 1983. Persistent anthelmintic effect of ivermectin in cattle. J. South Afr. Vet. Assoc. 54:249-250. Thorne, E. T. 1982. Pasteurellosis. Pages 72-77 in E. T. Thorne, et al., eds. Diseases of wildlife in Wyoming, 2nd ed. -Wyo. Game and Fish Dep., Cheyenne. Wescott, R. B., and B. R. Leamaster. 1982. Efficacy of ivermectin against naturally acquired and experimentally induced nematode infections in sheep. Am. J. Vet. Res. 43:531-533. Yazwinski, T. A., T. Greenway, B. L. Presson, L. M. Pote, H. Featherstone, and M. Williams. 1983. Antiparasitic efficacy of ivermectin in naturally parasitized sheep. Am. J. Vet. Res. 44:2186-2187. Received Accepted �171 Table 1. Characteristics of mountain sheep used in laboratory tests of the efficacy of ivermectin for treating lungworm infections. Animal no. no. Sex Age (years) Weight (kg) Assignment M 2 41 Treatment 6 42 Treatment 3 Ma Ma 6 80 Treatment 4 F 2 91 Treatment 5 F 6 61 Control 6 F 6 64 Control 1 2 aCastrated. �500 0----0 I-' TREATMENT CONTROL -....I N 400 ,P / ~ / 300 / 0> / <, W « > 0: « __J / / 200 ----0..... . ----0 100 .' 0' o I I I 1 2 3 I 4 r: 5 ~ 6 WEEKS Fig. 1. Counts of lungworm larvae in feces of mountain sheep before and after treatment with ivermectin. Animals treated after week 2. Control animals were not treated. Each point shows mean weekly counts for 1 experimental animal· �173 APPENDIX C Study plan describing experiments on efficacy of oral ivermectin for treating lungworm infections i.nbighorn sheep. STUDY PLAN A. NEED: Controlling lungworm (Protostrongylus sp.) levels in bighorn sheep with anthelmintic drugs is an important tool for managing bighorn populations in Colorado (Hibler et a1. 1976, Schmidt et a1. 1979); the lungworm treatment program has improved the status of several existing herds and allowed successful establishment of many transplant herds in the state (Schmidt et a1. 1979, Spraker and Schmidt 1984). Until 1983, oral benzimidazoles were the only anthelmintics with demonstrated efficacy in treating lungworm infections in bighorn sheep (Hibler et ale 1976, Schmidt et ale 1979, Foreyt and Johnson 1980), although no single benzimidazole was uniformly effective against all developmental stages of Protostrongylus (Hiber et ale 1982). Recently, we demonstrated efficacy of ivermectln (22.23-dihydroavermectin Bl) in its injectable form (Eqvalan R, Ivomec R, Merck and Co., Inc., Rahway, NJ 07065, . USA), against lungworm infections of bighorns (Miller et-al., in prep.). After completion of field and laboratory tests, ivermectin was adopted for use in transplants of bighorns in Colorado (Schmidt, pers. comm.). Ivermectin's broad spectrum of antiparasitic activity and wide margin of safety (Egerton et ale 1980, Hotson 1982, Campbell et ale 1983) make the paste form of this drug attractive as an adjunct to the existing fenbendazole regimen used in oral treatment of lungworm in free-ranging bighorns. Efficacy of iVermectin paste has been demonstrated against nematodes of several domestic species, including lungworms of cattle (Alva-Valdes et ale 1984). Advantages of this combined therapy include broadening the spectrum of antiparasitic activity, especially against external parasites, minimizing opportunity for development of resistant internal parasites (Campbell et ale 1983) and further decreasing lungworm burdens in free-ranging bighorns. - Before oral ivermectin is used in a large-scale management program, controlled studies should be performed in bighorns to demonstrate efficacy against Protostrongylus sp. We propose this study to experimentally determine efficacy and appropriate dosage of oral ivermectin (EqvalanR Paste, Merck and Co., Inc., Rahway, NJ 07065, USA) in treating lungworm infections in ,bighorn sheep. B. EXPECTED RESULTS AND BENEFITS: We anticipate efficacy of ivermectin paste against lungworm to be demonstrated at some dosage level by this experiment. Efficacy for injectable ivermectin has been demonstrated at 0.2 mg/kg, but reduction of firststage larval (Ll) output is more rapid at 0.5 mg/kg (Miller et al., in prep.). Once efficacy and an appropriate dosage are determined, �174 ivermectin paste can be adapted for use in the lungworm treatment program; potential benefits are as previously described. c. OBJECTIVES: Specific objectives of this study include: 1. Determine if ivermectin paste (Eqvalan), administered orally, is effective in reducing or eliminating lungworm infections in bighorn sheep. 2. Determine minimum dosage at which ivermectin paste is effective against lungworm in bighorn sheep. D. APPROACH: We will test the hypothesis that ivermectin paste (oral administration) will reduce lungworm infections in bighorn sheep with no adverse effects. Four sets of 2 tame bighorns will be paired randomly and assigned to 1 of 4 treatment groups: control (no treatment), 0.2 mg/kg oral ivermectin, 0.5 mg/kg oral ivermectin, and 1.0 mg/kg oral ivermectin; treatment pairs will be housed in separate bare-ground pens. Lungworm infections will be established naturally, or by oral administration of infective third-stage Protostrongylus larvae (L3) derived from naturally-infected bighorns and cultured in the laboratory (Lange 1973). Comparable lungworm burdens, represented by comparable Ll outputs, will be demonstrated in all experimental animals prior to beginning the study. Ll output will be determined prior to and during the study using fecal samples collected from observed defecations of individual-animals. Fecal pellets will be air-dried, and larvae will be recovered and counted using a modified Baermann procedure described by Beane and Hobbs (1983). Aliquots of each air-dried sample will be used for determination of drymatter according to AOAC (1980); Ll output will be expressed as Ll per 9 drymatter feces. The study will consist of a 2-week pretreatment period and a 4-week posttreatment period. Fecal samples will be collected from all animals and Ll output determined on alternate days throughout the study as previously described. After 2 weeks of pretreatment observations, all sheep will be weighed and animals from the 3 treatment groups will be administered invermectin paste (Eqvalan) per os at the respective dosage levels; the control group will remain untreated. In addition to fecal sampling and Ll output determination, daily observations of all sheep will be made for the first week posttreatment to record any adverse effects of oral ivermectin therapy. �175 Differences in Ll output between treatment and control animals and betw~en pr~- and posttreatment Ll output of treated animal s, will be examlned wlth a factorial analysis of variance for a fixed-effects model in an unbalanced, completely random design. Significant main effects are predicted for treatment, time, and treatment x time interaction; comparisons of dosage levels will be made in a similar manner. Ivermectin paste will be judged effective against lungworm if significant reduction in Ll output occurs and if no detrimental side effects or toxic reactions are observed; the most effective dosage level will be the minimum dosage with the maximum significant Ll output reduction which produces no toxic effects. Recommendations for field use of ivermectin paste in bighorn sheep will be based on these findings. E. SCHEDULE: June - July, 1985 - Determine lungworm burden by Ll output for selected tame bighorns, and administer infective L3 to noninfected animals, establish comparable lungworm infections in all experimental animals by late July. 1-14 August 1985 - Two-week pretreatment period. 15 August 1985 - Administer ivermectin paste to treatment animals. 15 August 12 September - Four-week posttreatment period. F. BUDGET: Drugs - $50.00 All other expenses will be covered by existing overhead. LITERATURE CITED: Alva-Valdes, R., G. W. Benz, D. H. Wallace, J. R. Egerton, S. J. Cross, and J. W. Wooden. 1984. Efficacy of ivermectin in oral paste formulation against gastro-intestinal and pulmonary nematodes in cattle. Am. J. Vet. Res. 45:685-690. A.O.A.C. 1980. Official Methods of Analysis, Thirteenth Ed. Assoc. of Official Analytical Chemists, Washington, D.C. l018pp. Beane, R. D., and N. T. Hobbs. 1983. The Baermann Technique for estimating Protostrongylus infection in bighorn sheep: effect of laboratory procedures. J. Wildl. Dis. 19:7-9. �176 Campbell, W. C., M. H. Fisher, E. O. Stapley, G. Albers-Schonberg, and T. A. Jacob. 1983. Ivermectin: a potent new antiparasitic drug. Science 221:823-828. Egerton, J. R., J. Birnbaum, L. S. Blair, J. C. Chabala, J. Conroy, M. H. Fisher, M. Mrozik, O. A. Ostlind, C. A. Wilkins, and W. C. Campbell. 1980. 22.23-0ihydroavermectin Bl, a new broadspectrum antiparasitic agent. Br. Vet. J. 136:88-97. Foreyt, B., and R. Johnson. 1980. sp.) in Rocky Mountain bighorn Pages 248-259 in W. o. Hickey, Sheep and Goat~ounc., Salmon, Treatment for lungworms (Protostrongylus sheep (Ovis c. canadensis) wlth albendazole. ed. Second Proc. Bienn. Symp. Northern Wild 10. 668pp. Hibler, C. P., T. R. Spraker, and R. L. Schmidt. 1976. Treatment of bighorn sheep for lungworm. Pages 35-39 in Proc. Bienn. Symp. Northrn Wild Sheep Counc., Jackson, WY. 165pp. _~, , and E. T. Thorne. 1982. Protostrongylus in bighorn sheep. Pages 208-213 in E. T. Thorne, ed. Diseases of Wildlife in Wymoing, Second Ed. Wyoming Game and Fish Dept., Cheyenne. 353pp. Hotson, I. K. 1982. The avermectins: J. So. Afr. Vet. Assn. 53:87-90. a new family of antiparasitic agents. Lange, R. L. 1973. Epidemiology of lungworms (Protostrongylus stilesi and rushi) in Rocky Mountain bighorn sheep (Ovis canadensis canadensis) M.S. Thesls, Colorado State Univ., Fort Colli~ 64pp. Miller, M. W., N. T. Hobbs, W. H. Rutherford, and L. L. W. Miller. in prep. Efficacy of injectable ivermectin for treating lungworm infections in mountain sheep. Schmidt, R. L. 1985. Colorado Division of Wildlife. --of Pers. Comm. April. , C. P. Hibler, T. R. Spraker, and W. H. Rutherford. 1979. An evaluation drug treatment for 1ungwonn in bighorn sheep. J. Wi1dl. Manage. 43 :461-467. Spraker, T. R., and R. L. Schmidt. 1983. Area experiences of dieoffs and disease: Colorado. Pages 6-7 in Proc. Bighorn Sheep Oieoff Workshop, Mimeo. Foundation N. Amer. Wild Sheep, Cranbrook, B.C. 35pp. �177 APPENDIX D Literature review and report of pilot experiments on the stress-disease complex in bighorn sheep. DEVELopr'1ENTOF TECHNIQUES FOR IDENTIFYING STRESS-INDUCED FAILURE OF IMMUNE REPONSES IN BIGHORN SHEEP N. T. Hobbs. Colorado Division of Wildlife, 317 W. Prospect St., Fort Collins, CO 80526. T. R. Spraker. College of Veterinary Medicine and Biomedical Sciences, Colorado State University, Fort Collins, CO 80523. G. G. Schoonve1d. Colorado Division of Wildlife, 317 W. Prospect St., Fort Collins, CO 80526. M. W. Miller. Colorado Division of Wildlife, 317 W. Prospect St., Fort Collins, CO 80526. The fundamental limit on the abundance of bighorn sheep in the Rocky Mountains appears to be high levels of mortality of young and adult animals resulting from bacterial pneumonia (March 1938, Buechner 1960, Bear and Jones 1973, Spraker 1979, Feuerstein et ale 1980, Foreyt and Jessup 1982, Spraker and Hibler 1982). These outbreaks of pneumonia may be associated with parasitism by lungworm (Protostron9¥lus spp.); however, they may also develop in the absence of significant paraslte loads (Forrester 1971, Spraker and Hibler 1982, Spraker et ale 1984). Thus, although lungworm may contribute to pneumonia infections, they apparently do not ultimately cause them (Spraker and Schmidt 1983). Alleviating this problem is made difficult by an inability to predict the likelihood of pneumonia outbreaks in specific populations, as well as an inability to intervene effectively once these outbreaks become acutely apparent. The ability to identify conditions leading to pneumonia-induced mortality and to effectively alter the course of events leading to that mortality depends on understanding the susceptibility of bighorns to pneumonia. It is generally agreed among experts on the ecology and physiology of bighorn sheep that failure of the immune system resulting from effects of excessive environmental stress offers a plausible hypothesis explaining the widely observed epidemics of pneumonia in bighorn populations (Proc. Bighorn Sheep Dieoff Workshop, Unpubl. Rep.). This hypothesis is based on the following observations (reviewed by Stephens 1980:180-182). In response to an environmental stressor, impulses are conducted from the cerebral cortex to the hypothalamus thereby stimulating the autonomic nervous system. Following that stimulation, the adrenal gland increases secretion of adrenaline and noradrenaline into the blood. These endocrine events mobilize body resources to facilitate heightened levels of activity during "fight or flight." If the stressor eersists beyond this immediate, emergency phase, the animal secretes cortlcotrophin-releasing factor from the hypothalamus, which stimulates release of adrenocorticotrophic hormone (ACTH) from the pituitary. Increased levels of ACTH, in turn, cause the adrenal gland to secrete corticosteroid hormones, further mobilizing the body's resources needed for activity. However, in addition to these mobilizing effects, corticosteroids decrease circulating lymphocytes �178 and cause degeneration of lymph and thymic tissue, thereby inhibiting the body's ability to resist disease (Weis 1972, Jensen and Rasmussen 1970, Paape et a1. 1973, Hartman et a1. 1976, Stein et a1. 1976). Kelley (1980:446) summarized these effects: "Therefore, the association between environmental stressors and lymphoid dysfunction may be the critical link between host resistance and environmental changes." It follows that when environmental stressors are excessively intense or prolonged, bighorn sheep will become increasingly susceptible to disease (Fig. 1). During February, 1983, 23 experts on disease outbreaks in bighorns concluded that developing techniques to quantify the influence of environmental stressors on populations of bighorn sheep was fundamentally important to future management of bighorn herds (Bighorn Dieoff Proc., unpub1. rep.). In particular, such techniques should allow prediction of disease outbreaks before their clinical signs appear; that is, they should offer "an early warnlng system" to facilitate intervention by wildlife managers before the effects of chronic stress become irreversible. Here, we propose to begin developing techniques for early detection and remedy of excessive stress in bighorn sheep. The classic causes of stress in bighorn sheep appear to involve rapid or gradual alteration of the environments bighorns occupy. These changes in environmental conditions include increased density of bighorn populations (Feuerstein et a1. 1980), disturbance by human activities (Spraker et a1. 1984), plant succession (Risenhoover and Bailey 1980) or competition with other herbivores (Goodson 1982). The impacts of such broad scale changes are difficult to duplicate with experimental treatments. However, these changes share a common feature; all lead to a shift in the collective environmental conditions that bighorn experience. Thus, from an experimental standpoint, the effect of artificially changing natural, familiar environments bighorn occupy can be simulated by moving natural populations to artificial, novel environments. We surmise that both situations have the same fundamental consequence: an increase in stress that results from changing the conditions bighorn experience from those faimi1ar and predictable to those foreign and uncertain. There is extensive evidence that movement of domestic animals from familiar to novel environments constitutes a strong environmental stressor. Dantzer and Mormede 1981:55 reviewed studies of effects of stressful stimuli in pigs and concluded that "•••psycho10gica1 stressors (exposure to novelty) are as effective as physical stressors (electric shock) in increasing plasma corticosteroids. They further suggested that such changes are particularly well suited as experimental treatments because "Exposure to a new environment is indeed a powerful stimulus that has the advantage over other stressors of not inducing physical pain while allowing qualitative and quantitative measurements •••" (Dantzer and Mormede 1983:8). Similarly, Stephens (1980:184) argued that "Nove1 stimuli which accompany environmental change or interference are probably the most potent short-term stressors for cattle of all ages. Reid and t~i11s (1962:292) found that movi ng domesti c sheep from "•••one place to another, whether it be merely walking from the paddock to enclosed yards or movement some distance by road transport" increased plasma cortisol by nearly an order of magnitude. II II �179 Many workers have observed that the average level of corticosteroids in the blood of mammals increases in response to chronic stress (Jensen and Rasmussen 1970, Willett and Erb 1972, Halley et a1. 1975, Johnson and Vanjonack 1976, Trenkle 1978, Burchfield et a1. 1980, Stephens 1980, Dantzer and Mormede 1983). However, measurement of IIbasalllor lIaveragelllevels of plasma corticosteroids in wild animals has serious drawbacks. The most formidable of these dilemmas is that blood corticosteroid values show extreme variation among different animals at any given time, and within the same individual at different times (Reid and Mills 1962, Jensen and Rasmussen 1970, Wagner and Oxenreider 1972, Willet and Erb 1972, McNutty and Thurley 1973, Holley et a1. 1975, Arave et a1. 1977, Trenkle 1978, Kattesh et a1. 1980). Consequently, repreated measures on many experimental animals are needed to provide precise estimates of average values. Such replication usually requires that blood be sampled from indwelling catheters (Trenkle 1978), a feasible procedure in domestics, but an extremely difficult one in wild species. In addition to these problems of precision, it is difficult to accurately measure IIbaselinellor lIaveragellconcentrations of plasma corticoids because the act of sampling itself causes sharp elevations in those concentrations (Bassett and Hinks 1969, Sap10sky 1983). What is needed, then, is a procedure that reveals the physiological response of bighorn to stress without requiring overly-frequent sampling. In addition, this procedure should not be biased by stress resulting from the sampling process itself. It is plausible that the magnitude of an animal's response to an acute stressor may serve as a reliable integrator of its past exposure to chronic stressors. When an animal experiences an acute stressor (electric shock, restraint, sudden loud noise), cortisol levels in the blood rise to a high point 10-30 mins. following the stressful event (Bassett and Hinks 1969), Venkateshu and Estergreen 1970, Wegner and Stott 1972, Wegner et al. 1973, ~1cNutty and Thurley 1973, Burchfield et ale 1980, Dantzer and Mormede 1980, Sapolsky 1983). If the stressor is removed, cortisol levels decline to levels similar to those prior to the event. If the stressor remains, cortisol levels tend to persist at high levels (Fig. 2). We will refer to these persistent high levels as the IIcortiso1 p1ateau.1I There is evidence that chronic exposure to environmental stressors elevates the magnitude of the cortisol plateau. Dantzer and Mormede (1981) found that the increase in plasma corticoids following acute stress (restraint or fighting) was greater in chronically stressed subdominant animals than in dominant ones. Friend et ale (1977, 1979) concluded that adrenal responsiveness to exogenous ACTH was a reliable indicator of exposure to past stressful conditions in cattle; exposure to increased stall density and social disruption increased adrenal sensitivity to ACTH stimulation. They surmised (Friend et a1. 1977:1960) that the adrenal response to acute stress offered a more reliable indicator of chronic stress than average glucocorticoid levels because: IIBasal glucorticoids varied greatly over time and were not affected by days at high density, time of sampling, age, days in 1actati on, or body wei ght. II However, there is also evidence that adrenal sensitivity to acute stress declines following prolonged exposure to chronic stress, as the animal adapts to stressful conditions (McNutty and Thurley 1973, Ader 1975, Burchfield et a1. 1980, Dantzer and Mormede 1981). If the peak or plateau in plasma corticoids following acute stress shows a consistent, increasing relationship �180 to chronic stress in bighorn sheep, an increase that eventually terminates in widespread disease and mortality, then the cortisol plateau will offer a reliable indicator of exposure to chronically stressful conditions. However, if that plateau rises and falls as the animal adapts to chronic stress, or if it varies unpredictably, then measures of adrenal responsiveness to acute stressors will not be predictive of exposure to chronic stressors. We tested the hypothesis that exposure to chronic stress elevates the cortisol plateau in bighorn sheep. We tested further that elevations in the cortisol plateau increase consistently with time after initiation of the chronic stress. METHODS Our experimental treatment was exposure to a novel environment; the control, exposure to a familiar environment. A treatment group of 10 wild bighorn sheep was captured in a drop net near Grant, Colorado, and transported to 1/2-ha holding pens in Fort Collins. The control group included 10 bottleraised, tame sheep thoroughly habituated to their surroundings and to human activity. During the weeks of March 11, May 20, and July 15, we drew blood from all animals by jugular venipuncture. Samples were collected every 5 mins. for 20-30 mins. Plasma cortisol cotent was analyzed by the radioimmunoassay method of Hasler et ale (1976). The cortisol plateau was estimated as the maximum blood cortisol concentration (ng/ml) observed after 20 mins. following the initiation of acute stress, restraint and venipuncture. Effects of month and treatment on the cortisol plateau were examined with a 2-way factoral analysis of variance for a fixed-effects model. RESULTS AND DISCUSSION Despite large interanimal variation in blood cortisol values, chronically stressed animals showed greater elevations (P 0.0001) in blood cortisol than control animals following an acute stress (Table 1). This effect of treatment was not significant until 5 mos. after initiation of the chronic stress. Both stressed and control sheep showed substantial temporal variation in maximum cortisol values; in particular, those values consistently increased with time after initiation of chronic stress in treatment animals (Table 1). However, the large increase in cortisol production we observed in control animals during March-May casts substantial doubt on their value as "unstressed" animals. This amplifies the need to control stress levels physiologically in future experiments. CONCLUSIONS We surmise that chronic stress causes physiological changes in bighorn sheep that may be detectable in trapped animals. However, to be useful, any physiological index must show lower variation than we observed. This �181 suggests that samples from urine or feces, which respond to the animal's physiological status over relatively long periods of time, may be superior to blood samples, which are obtained over relatively brief periods. LITERATURE CITED Ader, R. 1975. Early experience and hormones: emotional behavior and adrenocorticol formation. Pages 7-33 in B. E. Eleftheriou and R. L. Sprott. Hormonal correlates of behavior. Vol. 1: A lifespan view. Plenum Press. New York. Arave, C. W., C. H. Mickelsen, R. C. Lamb, A. J. Svejda, and R. V. Confield. 1977. Effects of dominance rank changes, age, and body weight on plasma corticoids of mature dairy cattle. J. Dairy Sci. 60:244-248. Bassett, J. M., and N. T. Hinks. 1969. Microdetermination of corticosteroids in ovine peripheral plasma: effects of venipuncture, corticotrophin, insulin, and glucose. J. Endocr. 44:387-403. Bear, G. E., and G. W. Jones. 1973. History and distribution of bighorn sheep in Colorado. Colo. Div. Wildl. Fed. Aid Proj. W-41-R, Job Final Rep. Jun: 1-231. Buechner, H. K. 1960. The bighorn sheep in the United States, its past, present, and future. Wildl. Mono. 4:74. Burchfield, S. R., S. C. Woods, and ~·1.S. Elich. 1980. Pituitaryadrenocortical response to chronic intermittent stress. Physiology and behavior 24:297-302. Foreyt, W. J., and D. A. Jessup. 1982. Fatal pneumonia of bighorn sheep following association with domestic sheep. J. Wildl. Disease 18:163-168. Forrester, D. J. 1971. Bighorn sheep lungworm-pneumonia complex. Pages 158-173 in J. W. Davis and R. C. Anderson, eds. Parasitic diseases of wildlife---.Iowa State Univ. Press. Ames. Friend, T. H., F. C. Gwazdauskas, and C. E. Polan. 1979. Changes in adrenal response from free stall competition. J. Dairy Sci. 62:768-771. , C. E. Polan, F. C. Gwazdauskas, and C. W. Heald. 1977. Adrenal -----g~lucocorticoidresponse to exogenous adrenocroticotropin mediated by density and social disruption in lactating cows. J. Dairy Sci. 60:1958-1963. Goetsch, D. D., L. E. McDonald, and G. Odell. 1959. The effects of four synthetic corticosteroids in leukocytes, blood glucose, and plasma sodium and potassium in the cow. Amer. J. Vet. Res. 20:697-701. Goodson, N. J. 1982. Effects of domestic sheep grazing on bighorn sheep populations: a review. Proc. Biennial Symp. No. Wild Sheep and Goat Counc. 3:287-313. Hartman, H., P. Hielman, H. r~eyer, and G. Steinbach. 1976. General adaptation syndrome in the calf (Selye). 5. Effect of increased glucocorticosteroid levels in phagocytosis actiity of leukocytes, RHS function, and morphology of lymphatic organs. Arch. Exp. Vet. Med. 30:59-73. Hasler, M. J., K. Painter, and G. D. Niswender. 1976. An125I-labelled cortisol radioimmunoassay in which serum binding proteins are enzymatically denatured. Clin. Chern. 22:1850-1854. Holley, D. C., D. A. Beckman, and J. W. Evans. 1975. Effect on confinement on the circadian rhythm of ovine cortisol. J. Endocr. 65:147-148. �182 Huber, H., and S. D. Douglas. 1971. Functional impairment of lymphocytes and monocytes: assessment in vitro. Seminar Hemotol. 8:192-215. Jensen, M. M., and A. F. Rasmussen. 1970. Audiogenic stress and susceptibility to infection. Pages 7-19 in B. L. Welch and A. S. Welch, eds. Physiological effects of noise. Phenum Press. New York. Johnson, H. 0., and W. J. Vanjonack. 1976. Symp: stress and health of the dairy cow. J. Dairy Sci. 59:1603-1617. Kattesh, H. G., E. T. Kornegay, J. W. Knight, F. G. Gwadauskas, H. R. Thomas, and D. R. Notter. 1980. Glucocorticoid concentrations, corticosteroid binding characteristics, and reproduction performance of sows and gilts subjected to applied stress during mid-gestation. J. An. Sci. 50:897-905. Kelly, K. W. 1980. Stress and immune function: a bibliographic review. Ann. Rech. Vet. 11:445-478. Kristensen, F., B. Kristensen, and S. Lazary. 1982. The lymphocyte stimulation test in veterinary immunology. V~t. Immunol. Immunopathol. 3:203-277. Marsh, H. 1938. Pneumonia in Rocky Mountain bighorn sheep. J. Mamm. 19:214-219. McNutty, K. P., and D. C. Thurley. 1973. The episodic nature of changes in ovine plasma cortisol levels and their response to adrenaline during adaptation to a new environment. J. Endocr. 59:171-180. Moberg, G., C. O. Anderson, and T. R. Underwood. 1980. Ontogeny of the adrenal and behavioral responses of lambs to emotional stress. J. An. Sci. 51:138-142. Paape, M. J., W. D. Schultze, and R. M. Miller. 1973. Leukocytic response to adrenocortiotrophic hormone as influenced by the infectious history of the mammary gland. J. Dairy Sci. 56:733-737. Risenhoover, K. L., and J. A. Bailey. 1980. Visibility: an important habitat factor for an indigenous, low-elevation bighorn herd in Colorado. Proc. Biennial Symp. No. Wild Sheep and Goat Counc. 3:18-28. Sapolsky, R. M. 1983. Individual differences in cortisol secretory patterns in the wild baboon: role of negative feedback sensitiity. Endocrin. 113:2263-2267. Soper, F. F., C. C. Muscoplat, and D. W. Johnson. 1977. In vitro stimulation of bovine peripheral blood lymphocytes: analysis of variation of lymphocyte blastogenic response in normal dairy cattle. Am.J. Vet. Res. 39:1039-1042. Spraker, T. R. 1979. The pathogenesis of pulmonary Protostrongylus in bighorn lambs. Ph.D. Thesis. Colorado State Univ., Ft. Col1ins. 233pp. ______,'C. P. Hibler, G. G. Schoonveld, and W. S. Adney. 1984. Pathological changes and microorganisms found in bighorn sheep during a stress-related die-off. J. Wildl. Dis. 20:319-327. , and C. P. Hibler. 1982. An overview of the clinical signs, gross ---~a'nd histological lesions of the pneumonia complex of bighorn sheep. Pages 163-172 in J. A. Bailey and G. G. Schoonveld, eds. Proc. of the Biennial Symp.lNo. Wild Sheep and Goat Counc. 3:163-172. , and R. L. Schmidt. 1983. Area experiences of die offs and ---~d'isease: Colorado. Proc. Bighorn Sheep Die-Off Workshop. Mimeo. Foundation N. Amer. Wild Sheep. Stein, M., R. C. Schiavi, and M. Camerino. 1976. Influence of brain and behavior on the immune system. Science 191:435-440. Stephens, D. B. 1980. Stress and its management in domestic animals: a review of behavioral and physiological studies under field and laboratory situations. Adv. Vet. Sci. Compo Med. 24:179-210. �183 Trenkle, A. 1978. Relation of hormonal variations to nutritional studies and metabolism of ruminants. J. Dairy Sci. 61:281-293. Venkatasseshu, G. K., and V. L. Estergreen, Jr. 1970. Cortisol and corticosterone in bovine plasma and the effect of adrenocorticotropin. J. Dairy Sci. 53:480-483. Wagner, W. C., and S. L. Oxenreider. 1972. Adrenal function in the cow. Diurnal changes and the effects of lactation and neurophyphophyseal hormones. J. An. Sci. 34:630-635. Wegner, T. N., and G. H. Stott. 1972. Serum minerals, leukocyte profiles, and plasma corticoids in dairy heifers after an injection of corticotropin. J. Dairy Sci. 55:1464-1469. __ , D. E. Ray, C. D. Lox, and G. H. Stott. 1973. Effect of stress on serum zinc and plasma corticoids in dairy cattle. J. Dairy Sci. 56:748-752. Weis, J. M. 1972. Psychological factors in stress and disease. Sci. Am. 226:104-113. Willett, L. B., and R. E. Erb. 1972. Short-term changes in plasma corticoids in dairy cattle. J. An. Sci. 34:103-111. Williams, E. S., and C. P. Hibler. 1982. Survey of Colorado and Wyoming bighorn sheep and mountain goats for paratuberculosis. Pages 173-187 in Biennial Symp. No. Wild Sheep and Goat Counc. 3:173-187. �184 Table 1. Averagea maximum cortisol concentrations (ng/ml) in blood of bighorn sheep 20 minutes after initiation of restraint. Treatment Month - March 49.1 a 37.3 - May 68.1a 57.2 - July 97.3a x Control - x 95% C1 61.0 28.7a 14.5 - 42.9 79.9 67.7a 53.6 - 81. 9 85.9 - 108.6 61 .1b 49.8 - 72.4 95% C1 aMeans within months that do not share the same superscript significantly different at P < 0.05. are �185 Human Activity .~ Poor Nutrition High Density Poor Visibility ••. Stress )II'"- Endocrine ---~-Response ~ Reduced Resistance MortalityFigure 1. Hypothesized relationship between environmental and mortality in populations of bighorn sheep. stressors ?-Cortisol '\ Plateau ~I ~I III I rl 81 -glo r- eo 10 20 30 Time After Initiation of Acute Stress (min) Figure 2. Relationship between blood cortisol levels and time following an acute stress. ��COloradO U1V1Slon ot Wlldllte Wildlife Research Report July 1985 187 JOB PROGRESS REPORT State of Colorado Project No. 01-03-048 Mammals 2 Research Work Plan No. Pronghorn Investigations ------------------------- Job. No. 1 ----------------------- Period Covered: Author: 3 --------------------July 1,1984 Pronghorn Population Dynamics Study - June 30,1985 T. M. Pojar ABSTRACT The random quadrat and random strip sampling systems for censusing pronghorn (Antiloca~ra americana) were executed for the fourth year in the limon/Hugo area and or the thlrd year in the Great Divide area. Population size and herd structure were estimated in each area using both sampling designs. The objective is to determine if either of these designs will adequately estimate both population size and herd structure during the same fly-over in late summer. The tests are conducted in 2 vegetative types; short grass prairie (limon/Hugo) and sagebrush steppe (Great Divide). The quadrat sampling design consistently results in higher density estimates with the difference being more pronounced in the sagebrush steppe type. In general, the precision level of both designs is comparable for estimates of population size. The herd structure estimates (bucks:100 does and fawns:100 does) are comparable between the 2 sampling systems. However, the precision level is lower from the quadrat system because of smaller sample size. High variability in the buck to doe ratio inhibits detection of a statistically significant difference between the 2 systems for this parameter. This ratio is consistently higher as estimated by the quadrat system, indication that detection of solitary bucks may be more likely during the quadrat search. The low variability in the fawn to doe ratios results in relatively precise estimates. ��189 PRONGHORN POPULATION DYNAMICS STUDY Thomas M. Pojar P. N. OBJECTIVES 1. Test the efficiency and precision of a quadrat and strip census design of rolling sagebrush steppe and short grass prairie. 2. Test the efficiency and precision of herd structure estimates when done concurrently with a quadrat and strip census on both rolling sagebrush steppe and short grass prairie. 3. Test the reliability of doing strip and quadrat censusing during late summer when it is possible to do herd structure counts. 4. Measure and model the effects of reducing a pronghorn population density by 50% or greater. SEGMENT OBJECTIVES 1. Test efficiency and precision of quadrat and strip censuses on short grass prairie to estimate density and population structure during 1 fly-over in late summer. 2. Test efficiency and precision of quadrat and strip censuses on sagebrush steppe to estimate density and population structure during 1 fly-over in late summer. 3. Simulate pronghorn population changes to evaluate results for each area. ACKNOWLEDGMENTS Special appreciation is extended to Kathy Cushman and Larry Dickerson for their dedication to the task of checking and re-marking quadrat corners in the Great Divide area. The following Colorado Division of Wildlife personnel from the NW Region also contributed to this study: J. Ellenberger, provided administrative and financial support; M. Bauman, K. Navo, and J. Haskins assisted as,navigators/observers during portions of the census flights. Mark Elkins of the SE Region provided administrative and financial support of this project. METHODS AND MATERIALS The winter of 1983-84 was extremely severe in the northwest portion of Colorado. Due to the severe winter conditions, the pronghorn population suffered high mortality, especially in the fawn segment of the population. There was also evidence of aberrant movements of groups of pronghorn, both into and out of the study area (Units A3 and A301). �190 A group of approximately 2,000 pronghorn were reported to have moved from the Red Rim area of Wyoming, along Fortification Creek to the vicinity of Ralph White Reservoir where most of them perished in the deep snow (Mike Bauman, Col. Div. of Wildl., pers. comm.). There also was some evidence of pronghorn movement from the nothwest portion of the study area to areas 30+ mi. to the south and west (John Ellenberger, Col. Div. of Wildl., pers. comm.). The census in the Great Divide area (Units A3 and A301) was of special interest because of the potential of measuring effects of the severe winter on the pronghorn population size and structure. An attempt was made to tally the number of yearling bucks as a measure of fawn survival from the previous winter (Table 1). The two areas of study and the methods used in this experiment are described by Pojar (1983). RESULTS Great Divide During the summer of 1984, an attempt was made to visit every marked quadrat corner in Units A3 and A301. There are a total of 109 quadrats with 436 potential corner markers. However, there are actually 409 marked corners because 27 of them are in hayfields, wheat fields, or etc., and were not marked. From the ground search and from notes taken during the quadrat census, it was determined that 139 of 409 markers (34%) were in need of repair. These markers were originally installed during the summer of 1982. A Bell Soloy helicopter was used in both the transect and quadrat census. The transect census was conducted 13-14 August and the quadrat census was completed during 14-17 August. As in the past, the quadrat census yielded a much higher pronghorn density estimate than the transect method (Table 1). Each method, however, reflected the dramatic reduction in the popUlation; the reduction from the previous year in the transect and quadrat estimates was 52.9% and 50.7%, respectively. Limon/Hugo The transect census was conducted on 27-28 August and the quadrat census was completed during 28-30 August 1984 in the Limon/Hugo experimental area. A Bell Soloy helicopter was used in each census. The east central area of the state (i.e. Limon/Hugo) did not experience the severe winter weather that affected the northwest portion of Colorado. The 1983-84 winter in the Limon/Hugo area would have to be considered "normal" to "mild" (Table 2). Summary During the next segment, a publication comparing the quadrat and the transect census methods will be prepared and submitted to either the Wildlife Society Bulletin or the Journal of Wildlife Management. The publication will include 4 years of data from both the short grass prairie and sagebrush steppe �191 habitat types. The analysis will focus on the efficiency and reliability of both census methods and will include management recommendations for conducting pronghorn censuses to obtain the most reliable information for population modeling input data. LITERATURE CITED Pojar, T. M. 1983. Pronghorn investigations--pronghorn population dynamics study. Colo. Div. Wildl. Res. Rep. July, part 4:379-385. �192 Table 1. Pronghorn census results for the Great Divide Data Analysis Unit (Game Management Units A3, A301) using a random quadrat (mile-square quadrats) and a random transect (mile-wide transects). Total area is 1,229 mi2• 1984 1982 1983 Strip Pop. est. 90% C.!. Lower C.L. Upper C. L. Mean density Quadrat 4,347 +31% 16,650 +17% 2,985 5,709 3.54 13,770 19,530 13.55 Buc ks :100 Does 90% C.!. Lower C.L. Upper C.L. 45.5 +19% 37.1 54.0 Fawns:100 Does 52.2 :!:_19% 42.2 62.3 66.8 +9.4% 60.5 73.1 90% C. I. Lower C.L. Upper C. L. No. Classified Strip Quadrat Strip Quadrat 8,868 +17% 18,438 +30% 12,883 23,993 16.39 4,689 +13% 9,345 +18% 7,373 10,363 7.22 4,052 5,327 3.82 7,644 11.046 7.60 42.9 +14% 36.7 49.1 56.3 +17% 34.8 +18% 46.6 66.0 28.5 41.0 55.9 +24% 42.4 69.4 65.6 69.1 +11.2% 61.4 76.8 47.7 +11 .5% 42.2 53.2 54.4 +20.4% 43.2 65.4 83.2 +9.4% 75.4 91.1 +6.7% 61.2 70.0 Bucks 336 324 337 603 1 Does 738 620 1,404 599 Fawns 921 414 _____lli_ _ill. 1,567 1,460 Total 2,928 1,350 *1n a sample of 108 bucks, there were 14 yearlings; i.e. 13% yearlings. **Of the total bucks observed (218), only 14 were yearlings; i.e. 6% yearlings. 323* 929 443 1,695 218** 390 212 820 Table 2. Pronghorn census results for the Hugo Data Analysis Unit (Game Management Units A36, A37, A38, A381) using a stratified random quadrat (mile-square quadrats) and a random transect (mile-wide transects). Total area is 1,688 mi2• 1984 1981 1982 1983 Quadrat Strip 2,402 +26% 1,774 3,030 1.42 47 6,109 +26% 4,507 4,507 3.62 67 +19% 38 56 Strip Pop. est. 90% C. I. Lower C.L. Upper C.L. Mean Density Bucks:100 Does 90% C1 Lower C.L. Upper C.L. Fawns: 100 Does 90% C.!. Lower C.L. Upper C. L. No. Classified 3,570 +22 2,773 4,367 2.14 64 +10% 58 70 1,203 Quadrat Quadrat Strip Quadrat :!:_27% 1,812 3,312 1.54 1,837 +27% 1,342 2,331 1.09 44 45 44 2,841 +49% 1,443 4,237 1.68 67 +48% +31% +51% +23% +58% +29% +50% 35 99 30 58 22 68 34 55 28 105 26 48 25 73 67 50 +12% 42 +28% 31 54 257 52 +17% 60 +21% 52 +11% 37 +32% 43 61 592 48 73 252 :!:_17% 56 78 381 44 57 805 2,602 Strip 3,046 +24% 2,312 3,779 1.80 3,555 +40% 2,128 4,983 2.11 38 49 46 57 1 ,023 25 49 308 �vVIV,QUV UIVI::>IVII 01 VlliGI1Te Wildlife Research Report July 1985 193 . JOB PROGRESS REPORT State of Colorado Project No. 01-03-048 Mammals 2 Research Work Plan No. Rocky Mountain Goat Investigations --------------------4 ------------------ Job. No. 1 Period Covered: July 1,1984 - June 30,1985 Author: Seasonal Habitat Selection and Activity of Sympatric Mountain Goat and Bighorn Sheep Populations D. F. Reed ABSTRACT Sixty-nine mountain goats were marked with radio telemetry collars, neck bands, or eartags. Ten mountain sheep were marked with radio telemetry collars or neck bands. Of the 27 alpine habitats used in the study, mountain goats and sheep occupied 16 and 19 of the habitats, respectively, during September through May, 1981-85. The number of mountain goat groups or individuals occupying 16 habitats was not greater (P > 0.50) than the same for mountain sheep groups 6~ individuals. Based upon tests of pooled observations, mountain goats selectively used alpine habitats; that is, they used alpine habitats disproportionately to their availability or abundance. Similarly, mountain sheep selected among alpine habitats such that habitat use was not random. Mountain goats tended to occupy more habitats within a year than mountain sheep, and the difference in numbers of occupied habitats was largest in the fall months (September-November). The null hypothesis that mountain goats and sheep exhibit the same activity patterns (feeding and resting during the day, and feeding and resting during the night) in alpine habitats is not rejected •. Ten percent of adult female mountain goats had twins, 67% haal neonate or kid, and 23% had no kid(s). ��195 SEASONAL HABITAT SELECTION AND ACTIVITY OF SYMPATRIC MOUNTAIN GOAT AND MOUNTAIN SHEEP POPULATIONS Dale F. Reed P. N. OBJECTIVES Evaluate the extent to which mountain goat populations limit seasonal habitat utilization of mountain sheep in alpine environments and to describe patterns and rates of dispersal of mountain goats from colonization sites. SEGMENT OBJECTIVES 1. Test the hypothesis that mountain goats use alpine habitats disproportionately to their availability. 2. Test the hypothesis that mountain sheep use alpine habitats disproportionately to their availability. 3. Test the hypothesis that mountain goats and sheep use alpine habitats that are spatially and/or temporally discrete - their distributions are independent. 4. Test the hypothesis that mountain goats and sheep exhibit the same activity patterns in alpine habitats. ACKNOWLEDG~1ENTS I thank R. Reiner and L. Reiner of the Mount Evans Research Station, University of Denver, for providing support and a place for a snowmobile and other field equipment. E. Hashimoto and J. Stone provided field assistance. C. E. Braun provided aerial photography and vegetation maps that included the study area. R. Angel and others reported locations of marked animals during the summer. S. Bassow provided both field and office assistance - her review and discussions of competition models were most helpful. L. Lovett provided office assistance and moral support. DESCRIPTION OF AREA The study area has been described by Reed (1982). METHODS AND MATERIALS Methods and materials have been described by Reed (1981, 1982). �196 ,RESULTS AND DISCUSSION Capture and Marking Sixty-nine mountain goats were marked with radio telemetry collars, neck bands, or eartags (Table 1). Of this 69,10 were harvested and 5 died during the study. Ten mountain sheep were marked with radio telemetry collars or neck bands. Three other sheep were individually identifiable from a previous marking program in 1977 (collars Yellow 19, Plain Yellow; eartag 4), and 1 was a readily distinguished, partial albino. Six mountain goat radio collars (4,5,7,17,27, and 55 - Table 2) were still active at the end of the study. Of these 6 radio collars, 4 were activity (tip switch) collars. Mountain goats 7 and 55 primarily occupied habitats located out of the study area. Hence, little or no data were collected on them. No signal has been received from or sighting made of mountain goat 35 since she was .collared. Collars outfitted on mountain sheep 1,4,9,10,11,12, and 13 were activity collars. This maintained the number of activity collars on mountain sheep at 4 (1,7,9, and 10 - Table 2). Habitat Use Of 27 habitats used in this study, mountain goats and sheep occupied 16 and 19 of the habitats, respectively, during September through May, 1981-85. Conversely, 6 of the habitats (3, 5, 6, 10, 23 and 24 - Reed 1982, Appendix A) were not occupied during this period by either species, and 11 and 8 were not occupied by mountain goats and sheep, respectively. Mountain goats generally used more habitats than those used by mountain sheep except for 1984-85 (Fig. 1). The mean number of habitats occupied by mountain goats was greater than the mean number of habitats occupied by mountain sheep for each season (f. < .05) and month (f. < .01) (Figs. 2 and 3). The number of mountain goat groups of individuals occupying 16 habitats was not greater (P > 0.50) than the number of bighorn sheep groups or individuals occupying the-same 16 habitats during the study. However, differences were apparent in habitat 2a(E) and 2a(S). These habitats are located above Lincoln Lake and were used frequently by mountain goats throughout the fall and winter seasons. During 1982 and 1983, mountain goats had also used these habitats frequently during the spring. However, heavy snow fall in March and April, 1984, may have resulted in different habitat occupancy patterns during the spring season. Also, habitat 2a(E) included a mineral lick and trap site. Similar features of 2a(E) and 2a(S) may account for a relatively high amount of mountain goat use. In theory, the same resouces were available to mountain sheep. These habitats may not have been used frequently by mountain sheep because (1) mountain sheep had different requirements, (2) mountain sheep were substantially fewer in number, and/or (3) moyntain sheep were excluded by the presence of mountain goats. The largest groups observed for mountain goats and sheep during the study were 25 and 35{ respectively. The mean size of groups for the 2 species was not different f. > 0.50). �197 Test of Hypothesis 1. The first hypothesis and its null were stated (Reed 1982:100) as follows: Hl = Mountain goats use alpine habitats disproportionately to their avai1abil ity. Ho = t,10untaingoats do not use alpine habitats disproportionately to their availability. Pooled data on mountain goat habitat use (September-May, 1981-85) were analyzed by comparing observed with expected frequency distributions. Habitat use was analyzed for number of mountain goat groups (> l) instead of number of individuals to avoid group behavior bias. The number of mountain goat groups per hectare (ha) for each habitat were the parameters used in this x2 tes~ (Table 3). The observed frequency distribution deviated significantly (x = lOl.5), P < O.005} from the expected. Hence, based on pooled data from the overall study, the null hypothesis that mountain goats do not use alpine habitats disproportionately to their availability is rejected. Test of Hypothesis 2. The second hypothesis and its null were stated (Reed 1982:105) as follows: H2 = Mountain sheep use alpine habitats disproportionately to their availability. Ho = Mountain sheep do not use alpine habitats disproportionately to their availability. Similar to the test of Hypothesis 1, pooled data on mountain sheep habitat use (September-Nay, 1981-85) were analyzed by comparing observed with expected frequency distributions. Habitat use was analyzed for number of sheep groups (> l) instead of number of individuals to avoid group behavior bias. The numoer of mount~in sheep groups per ha for each habitat were the parameters used for this x test2(Table 3}. The observed frequency distribution deviated significantly (X = 28.3, P < O.Ol) from the expected. Hence, based on data from the overall study, the null hypothesis that mountain sheep do not use alpine habitats disproportionately to their availability is rejected. Test of Hypothesis 3. The third hypothesis and its null are re-stated as follows: . H3 = Mountain goats and mountain sheep do not use the same alpine habitats. Ho = Mountain goat and mountain sheep use the same alpine habitats. Pooled data on both mountain goats and sheep habitat use (September-May, 1981-85) were analyzed by measuring the difference between 2 variables. The 2 variables were the number of mountain goat groups per ha and the number of mountain sheep groups per ha across 15 habitats (Table 3). For continuity within years, habitat 14 (i.e. 16 vs. 15 habitats occupied by mountain goats) was excluded from this analysis. There was a significant difference �198 (P < 0.05). Hence, based on pooled data from the overall study, the null hypothesis that mountain goats and sheep use the same alpine habitats is rejected. Activity Telemetry tip-switch activity collars were maintained on 8 mountain goats and 9 mountain sheep for varying periods during the study (Table 2). Generally, mountain goats were accessible for obtaining activity data. However, mountain sheep were more transitory and often dispersed into areas located near the boundary of the study area. Hence, substantially more activity data were collected on mountain goats than on mountain sheep during the study. Of the 8 mountain goats with activity collars, mountain goat 17 (Blue diagonal) and mountain goat 20 (Black te1e) provided the largest samples of act1vity periods (Table 4). Their feeding and resting periods during the day and during the night were not different (Table 5, Tests 1-4). Variation between the pattern of activity over 24 hours may be compared by examining specific 24-hour periods of selected mountain goats and sheep {Figs. 4-10}. In these cases, none of the animals compared had significantly different duration of day and night feeding periods (Table 6). Responses to sunrise and sunset appear to vary between animals and seasons. No responses to moonlight were detected. Generally, it is expected that such feeding and resting periods will vary between species, and possibly between seasons, habitats, and environmental conditions. Test of Hypothesis 4. The fourth hypothesis and its null are re-stated as fol lows: H4 = Mountain goats and mountain sheep do not exhibit the same activity patterns in alpine habitats. Ho = Mountain goats and mountain sheep exhibit the same activity patterns in alpine habitats. The mean duration of feeding and resting periods during the day and the mean duration of feeding and resting periods during the night of the 7 mountain goats were not different (P > 0.10, P > 0.50, P > 0.50, and P > 0.20) than the corresponding values of 4 mountaln sheep (Table 4). Longer resting periods exhibited by mountain sheep as a strategy to conserve energy during winter nights were not detected. The null hypothesis that mountain goats and sheep exhibit the same activity patterns (feeding and resting during the day, and feeding and resting during the night) in alpine habitats is not rejected. Partitioning of the overall hypothesis may not be necessary to reveal more definitive variables. Reproduction Of the female mountain goats collared, reproductive status was estimated on 8,17,21,23,19, and 17 in 1980,1981,1982,1983,1984, and 1985, respectively (Table 7). The mean age of mountain goats (n = 12) having their first kid or kids was 3.2 + 0.9 (SO) years. The mean age of mountain goats (n = 8) having twins was ~5 + 2.5 (SO) years. Mountain goat 27 (Red te1e II) has had 2 sets of twins, 1 at-age 9 and the other at age 11. She alternated �199 twins and singletons during the 5-year period (Table 7). Ten percent of the adult (> 4 years old) females had twins, 67% had 1 kid, and 23% had no kid or kids (Table 8). This percent of twinning is relatively high compared to others reported (Brandborg 1955:94). The rate of twinning, as the rate of having singletons, is likely a minimum since some kids may die soon after birth. An example of this was reported previously (Reed 1983). No reproductive data obtained so far on mountain goats (Table 7) support Heimer's hypothesis of alternate-year production (Heimer and Watson 1982). Furthermore, applicability of this hypothesis to mountain goats in the Mt. Evans study area may be in doubt because dominant male behavior in mountain goats differs largely from that in sheep. Also, it would have to be assumed that success in harvesting dominant male mountain goats is comparable to success in harvesting dominant male mountain sheep. This mayor may not be the case. Of the marked or identifiable female mountain sheep, the reproductive status was estimated on 8 (Table 9). Data from mountain sheep 1 (Table 9) may be in support of the alternate-year production hypothesis. Species Interaction One hundred instances of direct interaction have been noted between mountain goats and sheep throughout the study. The intensity of these interactions ranged from neutral or no overt response to moderate or intense flight reaction. Fifty-four interactions were neutral, 37 resulted in apparent deterrence of sheep from use of some resource; 1 resulted in a sheep following several mountain goats with no apparent response by the mountain goats (considered positive interaction), and 8 resulted in slight to moderate flight reaction in mountain goats from the presence of sheep. In all, agonistic interspecific behaviors observed, the mountain goats initiated the encounters and the sheep were the recipients. Although it could be suggested from these data that greater than 30% of mountain goat-mountain sheep interactions result in sheep yielding space or other resources, small sample size data should be used with caution. Conceptual Models of Competition Although it may be tempting to articulate simple ideas of competition between mountain goats and sheep in this study, in so doing we construct mental preconceptions (Pie10u 1981) •. Despite such preconceptions, 1 simple idea of competition between mountain goats and sheep entails a graph of the 2 species estimated population curves (Fig. 11). If we are confident that these curves represent real populations, it is possible that mountain goats will increase to a point at which they will displace mountain sheep through interference competition, exploitation competition or both. The results from these 2 types of competition are shown in Fig. 12. Based on Mount Evans annual ground survey. �200 In the case of interference competition, data already collected on species interactions (n = 100) provide an approximate point on a negative effects continuum (A, Fig. 12; where SP 1 represents mountain goats and SP 2, mountain sheep). This point represents approximately 30% of the interactions between mountain goats and sheep where mountain sheep yielded space or other resources as a result of agonistic interspecific behavior. It has been stated that interference competition is unlikely to evolve unless there is a potential for overlap in use of limited resources (i.e. exploitation competition) (Pianka 1981). Hence, it is likely that in this case of mountain goats and sheep, resource or exploitation competition is occurring. Of course, it is one thing to assert exploitation competition, and quite another to test that assertion. Since no field data was collected on exploitation competition in this study, a theoretical discussion may be useful. Assuming that both species interact only through use of limited resources, that the resources of forage and space are non-interactive (change in supply of 1 resource has no effect on rate of supply of another) (Tilman 1982), and that mountain sheep require more space and higher quality forage (based on estimates of greater selectivity in diet and habitat), zero net growth isoclines for the 2 species can be drawn (Fig. 13). Species A will be able to reduce resource levels to a point below that required for survival of Species B (area between curves A and B, Fig. 13). Species A will be able to competitively displace Species B in all habitats in which Species A is able to survive (total area to upper right of curve A, Fig. 13). Hence, if the system is allowed to reach equilibrium, mountain goats will be able to competitively displace mountain sheep. LITERATURE CITED Brandborg, S. M. 1955. Life history and ecology of the mountain goat in Idaho and Montana. Idaho Dept. Fish and Game Wi1d1. Bull. 2. 142pp. Craig, E. H. 1981. A relief-adjusted method for determining home range in mountainous areas. J. Mammal. 62(4):837-839. Heimer, W. E., and S. Watson. 1982. Differing reproductive patterns in da11 sheep: population strategy or management artifact? Pages 330-336 in J. A. Bailey and G. G. Schoonveld, eds. Proc. Bienn. Symp. North Wild Sheep and Goat Counc. 3, Fort Collins, CO. 405pp. Pianka, E. R. 1981. Competition and niche theory. Pages 167-196 in R. M. May, ed. Theoretical ecology principles and applications. 2nd ed. Sinauer Associates, Inc., Sunderland, MA. 489pp. Pie10u, E. C. 1981. The usefulness of ecological models: Qtrly. Rev. Biol. 56(1):17-31. Reed, D. F. 1981. Rocky mountain goat ecology study. Game Res. Rep. July, Part 2:209-222. a stock-taking. Colo. Div. of Wild1. ___~. 1982. Rocky mountain goat-bighorn sheep competition study. Div. of Wildl. Game Res. Rep. July:79-128. Colo. �201 Reed, D. F. 1983. Seasonal habitat selection and activity of sympatric mountain goat and bighorn sheep populations. Colo. Div. of Wi1d1. Game Res. Rep. Ju1y:403-422. Tilman, D. 1982. Resource competition and community structure. Univ. Press, Princeton, NJ. 296pp. Prepared ~ ;1C2~ Da e F. Reed Wildlife Researcher C Princeton �Table 1. Date, age, sex, collar or tag, and selected measurements I'-. C of mountain goats trapped in the Mt. Evans area. r-, Horn 1ength (cm) No. 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 Date 22 22 22 18 22 29 29 30 30 30 7 7 7 9 10 10 Aug Aug Aug 5ep 5ep 5ep 5ep 5ep 5ep 5ep Oct Oct Oct Jun Jun Jun 80 80 80 80 80 80 80 80 80 80 80 80 80 81 81 81 16 17 17 29 Jun Jun Jun Jun 81 81 81 81 29 28 30 4 7 15 Jun Aug Aug 5ep 5ep 5ep 81 81 81 81 81 81 6 Oct 81 14 Oct 81 Age 5ex Y F K F A F A F' A F K F A(6} 1 F 2 yr F K F A(8} F K M 2 yr A(7) 2 yr F F F Y M A M A(3} F A(3} M Y F A(6} F K M 3 F 4 F 5 F 2 M 6 F 8 F K t~ Co 11ar/tag Trap location Black collar White eartag White tele Blue tele R2 R1 Rl R2 Yellow tele Orange eartag 008 Green 2 tele (Ch 2) Yellow diag (Ch 1) Yellow eartag 79 Black collar 2 Yellow eartag (no no.) Black collar 3 Green 3 tele (Ch 3) Red-White-Blue te1e (Ch 2) Yellow w/Green "_" eartag Yellow w/Green "+" eartag Blue diag (Ch 4) 5 Yellow w/Blue "+" eartag Black collar 4 Black tele (Ch 6) Yellow w/Bl ue "_" eartag Black collar 5 Black collar 6 Black collar 7 Green eartag Black collar 8 Red tele (Ch 7) Black eartag Rl 53 53 52 TC 53 53 TC 52 Rl R2 R2 51 51 52 52 TC R2 Rl R2 R2 52 52 Girth (cm) Length (cm) Left Right 16.5 16.5 Wt. (kg) Horn to muzzle (cm) Front leg fm scapul a (cm) Hind foot (cm) ~ 100 89 119 102 114 90 111 159 165 102 152 118 99 142 113 128 147 143 125 183 149 147 126 157 51 96 101 111 92 105 133 97 81 146 145 159 138 147 140 114 114 116 77 116 94 69 116 72 95 121 21. 6 21. 3 2.8 22.9 4.4 13.2 21.3 3.3 23.1 4.3 12.7 20.3 27 74 41 21 25 41 77 21.0 21.5 14.0 14.1 25.0 24.8 21. 7 22.4 23.8 15.0 24.0 23.0 15.5 23.8 0.4 19.3 0.4 19.4 23.0 23.4 21.4 22.1 17.8 17.1 21.9 21.7 21.821.2 5.3 5.3 23.9 79.4 22.2 80.0 60.0 57.0 88.0 73.0 18.5 26.0 19.0 25.5 19.0 87.0 51. 5 25.0 12.0 20.5 73.0 42.0 76.0 74 45 58 22.8 23.5 19.5 23.0 79 33 23.8 15.2 82.0 81.0 73.0 76.0 78.0 9.9 61.0 30.0 27.0 34.0 30.5 31. 5 27.0 31.0 21.0 30.3 29.5 30.5 30.0 30.0 26.8 �Page 2 (continued) Table 1. 29 30 31 32 33 34 35 36 20 Oct 81 20 Oct 81 6 F K M 20 20 20 21 24 25 Jun Jun Jun Jun Jun Jun 82 82 82 82 82 82 Y Y Y F F F y M 2 Y M 37 38 39 25 25 25 25 26 26 26 26 Jun Jun Jun Jun Jun Jun Jun Jun 82 82 82 82 82 82 82 82 y Unk Y M y F F F F F 27 27 29 29 30 Jun Jun 5ep 5ep 5ep 82 82 82 82 82 52 53 54 55 30 16 16 16 16 29 5ep Jun Jun Jun Jun Jun 82 83 83 83 83 83 56 57 29 Jun 83 30 Jun 83 40 41 42 43 44 45 46 47 48 49 50 51 6 Y 5 Y 5 4 4 Unk F M F F y F F K M 3 Y F F F F Green diag (Ch 5) Blue eartag Grey eartag Black 1 dangle Black 2 dangle Black 3 dangle Blue diag wide (Ch 4) White eartag 1 White eartag 2 White eartag 3 White eartag 5 (right ear) Red tele (Ch 7) Black 4 dangle White 4 eartag (left ear) White 7 eartag White 6 eartag (right ear) White 9 eartag White 8 eartag (left ear) Black collar A Black collar 8 Orange eartag 1 (left ear) DAC R2 Rl DAC DAC DAC F M Orange eartag 5 (right ear) DAC DAC 4 F Black collar H DAC M 87 153 107 23.4 4.2 22.7 150 119 110 22.2 12.0 3.8 22.3 12.1 3.8 142 20.6 20.3 4.1 75 30 24.0 16.0 84.0 64.0 32.0 26.5 79 - - 32.0 32 - - 25.0 - - - - 32.0 - - - 32.0 DAC DAC DAC Rl DAC DAC DAC 3 5 3 2 118 DAC DAC DAC DAC BPP DAC Black Orange Orange Black Orange Green y collar E eartag 2 (left ear) eartag 3 (left ear) collar F eartag 4 (right ear) diag 112 (Ch 5) 51 51 DAC DAC DAC DAC DAC 115 90 78 120 118 182 23.1 23.6 (Harvested 10 5ep 83) 5E Rogers 119 171 23.7 23.7 30.0 N 0 W �N 0 .p.. Table 1. Page 3 (continued) 2 9 1 F F M Orange eartag 6 DAC 59 60 61 62 63 6 Jul 83 7 Jul 83 7 Jul 83 Black collar K Orange eartag 7 7 Jul 83 7 Jul 83 11 Jul 83 6 3 5 F Black collar M M F Orange eartag 8 Black collar N DAC DAC DAC DAC DAC 64 65 66 67 68 69 10 27 28 13 14 20 3 1 3 2 3 2 F F F F F M Black collar 0 Blue eartag 1 Black collar P Black collar V Black collar Y Orange eartag 2 (left ear) DAC DAC DAC DAC DAC DAC 58 Aug Jun Jun Jul Sep Sep 83 84 84 84 84 84 120 162 24.1 24.4 - - - 31.0 111 24.7 21.3 24.7 22.4 21.7 24.7 - - - 107 110 175 158 166 - - - 31. 5 31.0 - 30.5 112 88 114 110 111 157 113 151 137 141 17.3 12.0 23.2 16.2 11.9 23.1 - - - - - - - - - 18.2 20.1 20.2 - - 18.5 21.2 20.2 - - 29.5 26.5 aTC = Tumbling Creek at first switchback, Sl and S2 = Saddle below curve and mile post 6 (1 and 2, east and west trap, respectively). Rl and R2 = Rogers east above Lincoln Lake drainage and road between mile post 6 and 7 (1 and 2, north and south trap, respectively). DAC = Data acquisition center site at mile post 7. BPP = Below photo point on road shoulder between Rl - R2 and mile post 7. 33.0 • 31.0 30.0 �205 Table 2. Number assigned to animal, telemetry collar, channel, frequency, pulse, and activation date for radios outfitted on mountain goats and sheep in the Mt. Evans area. Activation Frequency Pulse per No. Coll ar Channel min. (ppm) date (mhz) Mountain goat 3 4 5 7 13 14 17 20 27 29 35 40 55 White Blue Fluorescent green Green 2 Green 3 Red-white-blue Blue diagC Black Red II Green diag I Blue diag wide Red I Green diag II 0 148.130 172.487 78 a 90-65 22 Aug 80 26 Jul 83 1 2 3 2 4 6 7 5 4 7 5 172.237 172.262 172.287 172.262 172.312 172.387 172.412 172.362 172.312 172.412 172.362 11 48 82 65-90b 65-90 65-90 75-120 65-90 60 90-65 65-90 11 29 7 9 10 11 . 14 20 24 25 29 5 1 3 1 9 8 2 3 6 172.362 172 .237 172.287 172.237 172 .462 172.437 172.262 172 .287 172.387 65-90 65-90 65-90 90-65 90-65 90-65 65-90 75-120 65-90 28 8 17 24 4 10 5 5 25 Jul~83 Sep 80 Oct 80 Jun 81 Aug 83 Aug 83 Sep 84 Oct 81 Jun 82 Jun 82 Jun 83 Mountain sheep 1 5 7 8 9 10 11 12 13 aActivity Black I Black & yellow Red I Yellow Black white Blue White Red II Black II Dec Feb Jun Nov Dec Dec Sep Sep Oct (tiP switch) collars; 90 ppm = head up, 65 ppm = head down. bActi vity (tiP switch) co llars; 65 ppm = head up, 90 ppm = head down. cDiag denotes diagonal. 83 85 82 82 82 82 84 84 84 �206 Table 3. Habitats, area of habitats, estimated areas of drifted snow pack, estimated areas of habitat available, and mountain goat and mountain shee~ grou~s ~er hectare (ha), 1981-85. Area Groups per ha Habitat designation la lb 2a(W) 2a (E) 2a(S) 2b(E) 2b(W) 8a 8b(N) 8b(S) 9 18 20 21 22 TOTAL Estimated snow packb Estimated available 174.1 42.4 14.0 50.6 69.5 17.7 94.2 31.3 5.8 22.6 16.5 32.9 9.9 11.5 64.6 3.0 5.0 1.0 1.0 6.0 1.0 6.0 3.0 0.5 5.0 1.0 1.0 3.5 1.0 6.0 171 .1 37.4 13.0 49.6 63.5 16.7 88.2 28.3 5.3 17.6 15.5 31.9 6.4 10.5 58.6 657.6 44.0 613.6 Habitata Mountain goats Mountain sheep 0.073 0.468 0.323 0.958 0.737 0.419 0.062 0.099 2.302 0.341 0.129 0.038 1.375 0.238 0.077 0.126 0.102 0.523 0.145 0.283 0.060 0.040 0.177 0.038 0.057 0.116 0.006 0.469 0 0.009 aBased on planimeter of habitats overlayed on Geological Survey Topographic Quadrangle Scale 1:24,000; not adjusted for vertical relief (Craig 1981). bOrifted snow sufficiently deep and/or compacted to preclude pawing for forage. Estimates derived from field inspections Sep-May. - �207 Table 4. Mean duration of feeding, resting, and moving activity periods of 7 mountain goats and 4 sheep during day (sunrise-sunset) and night (sunset-sunrise). Mean activity periods (min) Day (sunrise-sunset) No. Collar Night (sunset-sunrise) Feeding (n) Resting (n) Moving (n) 83.3 (32) 137.1 (8) 75.6 (71 ) 72.7 (28) 48.1 (9) 41.1 (72 ) 67.4 (42) 94.5 (19 ) 62.0 (3) 49.0 (3) 51.3 (38) 44.4 (20) 101 .0 (5) 48.2 (4) 8.0 (3) 24.0 (1) 5.0 (1) 68.8 (35) 59.3 (19 ) 58.4 (36) 56.3 (19) 68.3 (6) 18.8 (12 ) 10.0 (3) 30.3 (4) Feeding (n) Resting (n) 24.6 (29 ) 175.3 (30) 24.1 (34) 34.4 (55) 37.5 (37) 31.8 (13 ) 12.4 (5) 48.0 (4) 67. 1 (38) 172.4 (71) 207.4 (41) 386.6 (17) 86.5 (4) 29.1 (29) 34.4 (7) 27.8 (6) 29.5 (2) 168.6 (36) 222.6 (10) 321.7 (9) Moving (n) Mountain goat 4 Blue 14 Red-white-blue 17 Blue diagonal 20 Black 27 Red II 29 Green diagonal 40 Red I 8.0 (5) 19.3 (6) 11. 7 (3) 17.6 (5) 13.7 (3) 117.5 (4) r~ountain sheep 1 Black 8 Blue 9 Black and white 87.3 (6) White 43.8 (12 ) 11 190.7 (3) 9.3 (3) �208 Table 5. Independent t-tests of selected activity patterns (F = feeding, R = resting) of 4 activity-telemetered mountain goats. P Test Sample N Mean SO t 1 Blue diag day F Black day F 71 42 75.56 67..38 75.20 86.96 .527 = .599 2 Blue diag day R Black day R 72 38 41.08 51.26 38.91 44.40 -1. 242 = .215 3 Blue diag night F Black night F 55 37 34.38 37.46 36.02 36.25 -.401 = .664 4 Blue diag night R Black night R. 71 41 172.37 207.42 181. 15 160.74 -1.027 = .307 5 Blue day R Blue diag day R 28 72 72.68 41.08 66.09 38.91 2.958 = .004 6 Blue night F Blue diag night F 28 54 25.14 34.70 40.62 36.28 -1.086 = .280 7 Blue night R Blue diag night R 30 70 175.33 165.77 163.06 173.70 .256 = .686 8 Black day R Red II day R 38 20 51.26 44.45 44.40 33.71 .600 = .556 �Table 6. Summar~ of feeding ~eriod tests re~resented in Figs. 3 through 9. Means N Animal/Activity periods X Y X Y t SE SE Mountain goat 4 12-13 Oct 83 vs 1- 2 Nov 83 8 6 24.6 9.2 -1. 066 54.8 30.7 Mountain goat 20 26-27 Oct 83 vs 61.4 22,1 169.0 15-16 Nov 83 7 4 128.3 -1 .103 Mountain goat 17 28-29 Mar 84 vs Mountain goat 20 3- 4 Apr 84 5 5 106.6 63.8 19.6 8.0 1 .265 Mountain sheep 1 8- 9 Feb 84 vs Mountin sheep 10 20-21 Mar 84 6 5 57.2 24.6 70.4 32.2 -0.333 Mountain sheep 10-11 May 84 vs Mountain sheep 10 30-31 May 84 10 9 42.8 10.2 51.3 16.7 -0.448 Mountain goat 17 25-26 Jan 84 vs Mountain sheep 1 2- 3 Feb 84 7 6 74.3 42.9 47.2 22.3 0.533 Mountain goat 17 21-22 Sep 83 vs Mountain sheep 10 20-21 Mar 84 67.8 16.6 57.2 24.6 9 6 0.373 df P Fig. 12 > .20 9 > .20 5 8 > .20 6 9 > .50 7 17 > .50 8 12 > .50 9 4 N 0 1.0 13 > .50 10 �210 Table 7. No. 3 4 5 7 8 10 12 13 14 17 19 20 22 23 24 26 27 29 31 35 40 47 48 50 53 55 57 42 59 61 63 64 66 67 68 status of collared female mountain goats in the Mt. Evans stud,l area. No. kids Date Est. age Date replaced 1983 1984 1985 banded Jun 84 1980 1981 1982 Estimated re~roductive Coll ar 22 Aug 80 White tele (813) 18 Sep 80 Blue tele (Ch 0) Fluorescent green (Ch 1) 22 Sep 80 29 Sep 80 Green 2 (Ch 2) Yellow diag (Ch 1) 30 Sep 80 Black collar 2 30 Sep 80 Black collar 3 7 Oct 80 7 Oct 80 Green 3 (Ch 3) Red-white-blue (Ch 2) 9 Jun 81 Blue diag (Ch 4) 16 Jun 81 Black collar 4 17 Jun 81 Black tele (Ch 6) 29 Jun 81 Black collar 5 28 Aug 81 Black collar 6 30 Aug 81 4 Sep 81 Black collar 7 Black collar 8 I 15 Sep 81 Red tele II (Ch 7) 6 Oct 81 Green Diag (Ch 5) 20 Oct 81 Black collar zg 18 Sep 84 Blue diag wide (Ch 4) 24 Jun 82 Red tele I (Ch 7) 25 Jun 82 29 Sep 82 Black collar A 29 Sep 82 Black collar 8 II 30 Sep 82 Black collar E Black collar F 16 Jun 83 Green diag II (Ch 5) 29 Jun 83 30 Jun 83 ~i 30 Jun 83 Black collar K 7 Jul 83 Black collar M 7 Jul 83 11 Jul 83 Black collar N 10 Aug 83 Black collar 0 28 Jun 84 Black collar P B 1ack co 11ar V 13 Jul 84 Black collar Y 14 Sep 84 ~~:~~ ~~n:~ 26 Jul 83 8 7 8 10 Aug 83 14 Sep 84 10 6 12 6 11 6 6 4 9 6 7 8 9 11 9 3 4 8 Unk 3 5 3 6 5 6 10 7 6 4 3 2 3 la 1 1 1 0 0 0 1 lb 1 2 Unk 0 2 0 1 0 1 0 le 1 1 1 lf lh 1 aWhite eartag 1. bConfirmed with 1 kid 2 Jun; without kid 29 Jun and thereafter. cParturition dMortality 1 Jun yielded 2 kids; with only 1 kid 4 Jun and thereafter. fall 82 - summer 83. eBlue "-"; after eartag pulled out, known as "injured ear." fBlack eartag. gPreviously Grey eartag. hBlue eartag. ipreviously White eartag 4. 1 2c 1 1 Unk 0 0 Unk 1 1 1 1 1 1 Unk 0 2 1 1 0 1 0 0 Unk 0 1 0 1 1d 1 0 Unk Unk 1 Unk 0 2 1 1 1 0 Unk Unk 0 Un~ 1 _d 0 Unk 1 2 Unk 0 0 0 0 1 0 1 1 0 2 1 0 Unk 1 Unk Unk 1 Unk 1 1 Unk Unk Unk Unk Unk 1 2 0 2 1 1 0 0 1 0 1 Unk 0 1 0 �211 Table 8. Number of adult (> 4 years old) marked female mountain goats associated with 0, 1, or 2 neonates or kids across years in the Mt. Evans stud~. Years Total No. of Percent neonates 1980 1983 1984 1985 No. 1981 1982 0 0 4 7 4 4 20 23 5 9 14 10 8 12 58 67 2 0 2 2 2 3 0 9 10 TOTAL 6 11 20 19 15 16 87 100 Table 9. Estimated reproductive stud area. Collar or No. identification 1 2 3 4 5 6 9 10 11 12 13 status of marked or identifiable Black telg Ia (Ch 5) Yellow 19 Plain yellowC Eartag 4 Black and yellow tele (Ch l)d Partial albino Black and white tele (Ch 9) Blue tele (Ch 8) White tele (Ch 2) Red tele (Ch 3) Black tele II (Ch 6) apreviously Date banded Est. age Jun 84 1 Feb 77 1 Feb 77 77 25 Jan 77 1 Feb 77 4 10 5 5 25 Dec Dec Sep Sep Oct 82 82 84 84 84 12 12 Unk 7 7 Unk 6 5 3 Unk 3 female mountain sheep in the Mt. Evans 1980 1981 1982 1 1 Unk 0 0 0 0 Unk 0 0 1 1 0 Unk 1 1 1 0 Unk Unk Unk Unk 1 0 1 Yellow 17; Black tele collar added 28 Dec 83. bMost numbers have come off. Only stitching and faded areas make identification cNo stitching or faded areas apparent; collar frayed on bottom-posterior. dpreviously 1983 eartag 18; Black and yellow telemetry collar added 8 Feb 85. possible. 1984 1 1 2 1 1 1 1 1 1985 0 �212 0-----0- 15 -- ----~ \ \ \ \ \ \ \ \ 'o 10 "0 OJ "r'f p.. ::l t) t) 0 U) .j.J qj .j.J "r'f .0 qj ::t: 4-l 0 . 0 z 5 81-82 Fig. 1. 82-83 83-84 84-85 Number of habitats occupied by mountain goats (dashed line) and sheep (solid line) by year 1981-85. �213 15 10 · zo 5 'all Fig. 2. W:l.nt-er Spring Mean no. of habitats occupied by mountain goats (dashed line) and sheep (solid line) by season (fall = Sep-Nov, winter = Dec-Feb, spring = Mar-May) 1981-85. �214 15 10 ~ \ \ · o Z \ '0-_ 5 Sep Fi g. 3. Oct Ndv Dec Jan Feb Mar Apr May Mean no of habitats occupied by mountain goats (dashed line) and sheep (solid line) by month 1981-85. �2400 2400 1200 1200 A B Fig. 4. Mountain goat 4 (Blue telemetry - Ch 0) feeding (horizontal lines), moving (vertical lines) and resting activities during 24-hour periods 12-13 October 1983 (A) and 1-2 November 1983 (B). N •..... VI �N •..... ~ 240;0 2400 1800t=' Fig. 5. ~~=------ tel::z;4f' y 1200 1200 A B 'W Mountain goat 20 (Black telemetry - Ch 6) feeding (horizontal lines) and resting during 24-hour periods 26-27 October 1983 (A) and 15-16 November 1983 (B). . 0600 activities �2400 1800t 2400 -== - ?:j~~d 1200 1200 A B Fig. 6. Mountain goat 17 (Blue diagonal - Ch 4) (A) and mountain goat 20 (Black telemetry - Ch 6)(B) feeding (horizontal lines), moving (vertical lines) and resting activities during 24-hour periods 28-29 March 1984 and 3-4 April 1984, respectively. ,_. N -...J �N I-' co 2400. 1800t:=! t·f 2400 >k 0600 1200 1200 A B ~' , t",_ l' Fig. 7; Mountain sheep 1 (Black telemetry - Ch 5)' (A) and mountain sheep 10.jBlue,Jel~metry -=-_Ch8) (B) feeding (horizontal lines),'moving (vertical liries) and resting activities during 24-hour periods 8-9 February 1984 and 20.-21 March 1984, respectively. �2400 1800 F S?¥" 2400 '$ 1 ~ ~===-~. ~0600 1200 1200 A B ~ Fig. 8. Mountain sheep 1 (Black te lemetrv - Ch 5) (A) and mountain sheep 10 (Blue telemetry - Ch 8) (B) feeding (horizontal lines), moving (vertical lines) and resti~g activities duri~~ ~4-hour periods 10-11 May 1984 and 30-31 May 1984, respectively. N •..... 1.0 �N N o , 2400 2400 (+ 1800 } 1200 1200 A B Fig. 9. Mountain goat 17 (Blue,diagonal-Ch 4) (A) and mountain sheep 1 (Black telemetry - Ch 5) (B) feeding (horizontal lines), moving (vertical lines) and resting activities during 24-hour periods 25-26 January 1984 and 2-3 February 1984, respectively. �2400 1800~f Fig.10. 2400 §~~~d 0600 1200 1200 A 8 Mountain goat 17 (Blue diagonal - Ch 4) (A) and mountain sheep 10 (Blue telemetry - Ch 8)' (B) feeding (horizontal lines) and resting activities during 24-hour periods 21-22 September 1983 and 20-21 March 1984, respectively. N N ,_. "". ro �N N N 200 .o, /'" ,,/ 150 ,,/ JY' "- "" '0 ,,/ 100 / / / 50 c( 78 Fig. 11. 79 80 81 82 83 84 Ground census estimates of mountain goats (dashed line) and mountain sheep (solid line) in the Mt Evans area 1978-84. �223 ,- SP 1 SP 2 DISPLACEMENT CI.l E-I u IJ::I INTERFEHENCE A ~~ Z g:H E-' HZ E-'O <U C,!) ~ SP 2 SP 1 DISPU_CEMENT COMPETITION SP 1 SP 2 LOCALLY EXTINCT CI.l E-I U ~ ~""" ~S L.. EXPLOITATION ~:::> Z --g:~ ~OEXISTENCE @ EQUILIBRTlTM _ _ RESOURCE PARTITIONING HZ E-IO <:u C,!) ~ SP 2 SP 1 LOCALLY EXTINCT Fig. 12. Results of competition (interference, exploitation. or both) as expressed on a continuum where specics 1 (Sp 1) and species 2 (SP 2) dominate at o?pcsite ends of the continuum. Neither species dominate at a point re:;:"",sented by the middle of the con.tinuum where negative effects are equal. Field data on i~terference betHeen mountain goats and sheep are represented at point A on the continuum. where SP 1 represents mountain goats and SP 2 mountain she2p. :~":"":'.'"'' . �224 A B RESOURCE 2 RESOURCE 1 Fig. 13. Zero net growth isoclines for 2 species (A = mountain goats, B = mountain sheep) where resource 1 represents amount of space and resource 2 represents amount of forage (modified from Tilman 1982:73). �Colorado Division of Wildlife Wildlife Research Report July 1985 225 JOB PROGRESS REPORT State of Colorado Project No. 01-03-048 r,1ammal s 2 Research Hork Plan Black Bear Investigations ------------------No. 5 ------------------ Job. No. 1 Period Covered: July 1,1984 Author: Black Bear Population Ecology - June 30,1985 T. D. I. Beck ABSTRACT Sixteen black bears were snared in 1984; 9 recaptures and 7 new bears. Five of the 7 new bears were migrants, 1 a resident, and 1 unknown. Three yearlings were tagged in dens; bringing total individuals tagged in the study to 56 females and 53 males. Five marked bears were known to be killed in 198485; 4 legal and 1 illegal. Of the 109 tagged bears, 36 are known to be dead. Average age of first litters was AC 6; mean litter size was 2.09 with 50:50 sex ratio; and frequency between 1itters was 2.5 years. First year mortal ity in black bears may run as high as 56%. Den entry times were strongly peaked in the third week of October, an event I believe to be a direct response to unusually heavy snowfall in mid-October. Data were collected from 25 dens. ��227 BLACK BEAR INVESTIGATIONS Thomas D. I. Beck P. N. OBJECTIVES 1. Develop techniques to accurately and precisely estimate bear population levels. 2. Determine habitat preferences of selected bear populations. 3. Describe population dynamics sufficiently to allow analysis of various harvest and habitat manipulations. SEGMENT OBJECTIVES 1. Estimate age when female black bears have first litters, litter size, and frequency of litters. 2. Calculate home range size and seasonal home range dynamics for black bears within the Black Mesa study area. 3. Document survival of radio-collared black bears with emphasis on bears in age classes 2, 3, 4, and 5. 4. Determine habitat preference of 8 female black bears in the study area. ACKNOWLEDGMENTS Assistance with field work was provided by the following Division of Wildlife personnel: J. Broderick, M. Dege, M. Grode, D. Homan, M. Jandreau, G. Tischbein, and J. Toolen. B. McEwen volunteered during den season and his long hours were both appreciated and valuable to our efforts. METHODS AND ~~TERIALS Population Description Capture efforts lasted from 18 August to 20 September 1984 in the Black Mesa study area. No spring trapping was attempted because extensive mud slides and flooding restricted access to crucial areas. All captures were made with Aldrich snares; with the exception of M-49. Snare sets were out of sight from roads and trails and were baited with meat scraps and honey. Snares were checked daily. Snared bears were immobiled with a combination of ketamine hydrochloride and xylazine hydrochloride in a mixture of 180 kg ketamine and 45 mg xylazine per milliliter. Drug was administered by use of a 1.8-m jab pole. Denned bears were drugged by use of a jab pole 0.5, 1.2, or 1.8 m in length. Drug dosage was 6.6 mg ketamine per kg of body weight. �228 Bears were ear-tagged in both ears with numbered, plastic Roto-tags. All female bears age class (AG) 3 or older were instrumented with a Telonics radio transmitter collar. Yearling bears that were handled in dens with their mother, and thus known to have been born in the study area, were instrumented with small radio-transmitter collars designed to operate for 16 months. These small collars were removed from AC-3 bears during denning and replaced with larger radio-collars. The small collars have a short section of rubber surgical tubing which will deteriorate and allow the collar to drop free should we not be able to remove the collar in the next den. Data on litter size and survival of yearlings were obtained from visiting dens in the winter. Survival of older bears was monitored by radio telemetry and hunter contacts. Habitat Use Data on general habitat use were collected primarily from ground tracking radio-collared bears supplemented by weekly aerial tracking. Den sites were located in October-December, 1984, by both ground and aerial tracking. Den entrances were marked with rip-stop nylon flagging to facilitate location in March. Dens were visited in January, March, and April, 1985, and numerous physical measurements of the den were taken as well as a general site description. Analyses of detailed habitat selection by 8 female bears has not been completed. This portion of the study was contracted to a graduate student and while the student has been busy analyzing, he has lost sight of the realities of timeliness and obligation. Hopefully, correspondence with university officials can speed this project up. The major advisor has a long record of not pushing students and allowing them to hang on for a few extra years. Home Range Calculations All bear locations were recorded by UTM coordinates. The spatial description of the ranges employs the minimum polygon procedure and utilizes the HOME computer program described by Harestad (1981). Detailed reporting of home range sizes, spatial and temporal changes, and family relations will be deferred until completion of the final field season (March, 1986). Migration distances are reported as straight line distances although the topography traversed dictates much longer trips. Migrant bears are seldom located more than once each 10 days until they move into the study area in mid-August. Times of travel are estimated in multi-day periods. RESULTS AND DISCUSSION Population Description Twenty black bears were captured in 1984; 16 in snares, 3 in dens, and 1 surrounded on the ground (Table 1). Of the 16 snare captures, 9 were recaptures from previous years. Of 6 new females snared in 1984, 5 were classified as migrants based on movement out of the study area in late September. The adult female classified as a resident (F-61) has her home range centered in the headwaters of the S. Smith Fork, a roadless area of approximately 40 km~ This �229 area had only been trapped for 1 week in August, 1980. All male captures except M-73 were recaptures or den handlings. Little data was obtained on M-73 because I mistakenly injected him with dopram instead of ketamine. Dopram is a respiratory stimulant and even after correctly dosing him with ketamine, I was only able to get ear tags in and the snare off his foot before he began aggressive behavior. M-49 was surrounded on the ground in mid-November by 3 project personnel. He was seriously injured, and we were able to immobilize him with the jab stick. He had been shot with a high-powered rifle through one thigh and shattered the other ankle. The shooting occurred within the closed area. He died a few days after this capture. A major problem in any population study is missing animals. They are not known to be alive or dead. The group of missing animals of most concern are those radio-collared bears that go off the air during the fall big-game season. In 1984 radio contact with 4 females was lost during the big game season. All 4 were residents with well-documented seasonal ranges. It is probable that 1 collar malfunctioned from battery failure as it had been operating for 30 months. The other 3 females were wearing collars of 6-15 month operational life. We plan to concentrate trapping activities on fall ranges of these bears in 1985 in hopes of finding 1. Otherwise, we can never know their fate--only suspect. Four tagged bears were legally killed duing the 1985 spring season. F-19 was shot by 2 outfitters training hounds in r~ay. Apparently, she would not tree and was attacking their dogs. She was roughly 1.5 km north of the closed area. M-45 was shot on a bait site 0.1 km north of the closure. He was the third tagged bear killed at the bait site during the study. M-43 was a longdistance migrant who was killed over bait in Antelope Creek, a distance of 24 km east of the study area and 44 km east of his fall range within the study area. F-74 was shot over bait in r<1ay0.1 km east of the closed area and 24 krn east of her capture site last fall. Fifty-three males have been tagged so far, of which 25 are known to be dead. The area closed to bear hunting is roughly 700 km2• Of the 25 known deaths, 11 were legal kills, 8 were illegal kills, 4 were natural deaths, and 2 were handling losses in the den. Of the 56 females tagged, 11 are known to be dead; 4 legal and 6 illegal kills and 1 natural death. The fact that we have caught more females than males each year of the study and the documented losses suggest this population is suffering from heavy exploitation, even with a closed season. . Fecundity of the population remains low and is not changing as sample size slowly grows. Age of first litter remains at AC 6 (x = 6.4, median = 6). Based on 17 females, the frequency distribution for age of first litters was: 6% at AC 5, 53% at AC 6, 29% at AC 7, 6% at AC 8, 6% at AC 2 9. Frequency of litters was calculated 2 ways. The first, and less preferable, is to add the total years of study since first litter for 9 females and divide that number (44) by number of litters in the period (18). This gives a litter frequency of 2.44 years. I prefer the second method, or frequency distribution. Of the 9 females with 4 or more years of data, 5 littered every second year, 2 littered at 3-year intervals, 1 each at 2.5 and 4 year intervals. An average from the distribution was 2.5 years. �230 Litter size (N = 22) averaged 2.09 with an even sex ratio (50:50). Of 18 litters of 2 or more, 8 litters had same sex litter mates. More distressing than the low fecundity was 'the apparent low survival of offspring to the age of estrangement from their mother (16 months). Of the cubs checked in dens as cubs, we believe we know the fate of 30. The remainder were in dens that we missed or are currently cubs. I make 1 very important assumption: cubs not present in their mother1s den in the winter after birth are presumed dead. I know exceptions can occur. LeCount (1983) documented fall estrangement in 2 instances when the female gave birth to a litter than winter. I have not witnessed this in our study area. I know cubs orphaned in the fall can survive to their yearling summer (F-7 and young in our study) however, I can find no reports in the literature of voluntary fall separation and consequent survival of the cubs. LeCount (1983) did not know the fate of the early estranged cubs. Furthermore, I have not captured any yearlings that were marked as cubs except those that denned with their mothers. The disappearance of cubs occurred in 2 rather distinct periods; May-June and October-November. The earlier period is probably mostly natural mortality, although I have good reason to believe 2 cubs may have died from pulling M-44 coyote-getters. The latter period coincides with deer and elk season and our cubs weigh 34-48 kg. I believe that nearly all disappearance in this period is man induced--the cubs are shot. Not having captured any of these IImissingllcubs as yearlings and the documented shooting of F-2 and F-3 (sibling cubs) support my belief. I also assisted a neighboring officer investigate the shooting of 2 cubs from a tree by a hunter. So I know that such actions occur. Thirteen of 30 cubs have survived to estrangement (7 ~, 6 ~) in the presence of their respective mothers. Assuming all missing are dead, first year mortality was 56.7%. Assuming half of the missing were dead, first year mortality was 28.3%. Either figure is a far cry from the assumed 5% first year mortality used by Bunnell and Tait (1981) in their early modeling attempts. I believe high cub mortality, coupled with the documented adult mortality, has kept the study population from increasing. Sample sizes for population structure, mortality, and natality estimates are small in absolute terms because the total black bear population is small. Sample sizes as a proportion of the total population component are comparable to population dynamics studies of other large mammals. Mortality estimates have remained relatively constant for the past 3 years. Given low recruitment and high mortality even in this supposed unhunted population, a population crash seems inevitable. Habitat Use General trends of habitat use continue the same as in previous years. Migrant bears appear on the study area in mid-August then return to denning sites off the study area in late September and early October. All the migrations have been into the study area. Again, all the resident collared bears and the collared migrant bears used the Cow Creek-Doug Creek-Clear Fork area. The soft mast crop was poor and the bears moved more in response. A migrant female moved north about 14 km, thus extending her airline distance from berries to den to 50 km. Den entry times were difficult to obtain in 1984 because of extremely heavy snowfall in mid-October (over 1.3 m at 3,300 m). The most reliable entry times are presented in Table 2. Den characteristics are in Table 3. �231 Four instances of den reuse were documented this year. An AC-2 female denned in a rock cavern in an aval~nche chute on Bonfisk Peak; the same den used by F-ll in 1979-80. One is inclined to speculate the 2 bears may be related. Two AC-3 females (sisters) used dens that had been previously used by their mother. One young female reused their natal den while the other used her mother's den from the winter prior to their birth. The family ties and learning behavior get more interesting. An AC-2 female used a ground den in 1984-85 that was used by an AC-2 male in 1982-83. These young bears share a common mother but come from different litters. Evidence of prior use in dens continues to be common. Detailed analyses of habitat use have not been completed by the graduate student. Unless such analyses are completed by the end of the next segment, permission to use the material for a thesis will be rescinded, and I will complete the analyses. LITERATURE CITED Bunnell, F. L., and D. E. N. Tait. 1981. Population dynamics of bears implications. Pages 75-98 in C. W. Fowler and T. D. Smith, eds. Dynamics of large mammal populations. J. Wiley & Sons, N.Y. 477pp. Harestad, A. S. 1981. Computer analysis of home range data. Fish & Wildl. Branch, Fish & Wildl. Bull. 8-11. 25pp. Brit. Col. LeCount, A. 1983. Evidence of wild black bears breeding while raising cubs. J. Wildl. Manage. 47(1):264-268. Prepared by: ~~ Ihorns: ~ BeCk Wildlife Researcher C �232 Table 1. Black bears snared or newly tagged in dens in 1984-85, Black Mesa, CO. 10 no. Date Estimated Age Wt (kg) Remarks F-38 F-53 F-54 F-55 F-61 F-63 F-65 F-66 F-67 F-69 F-74 8-20-84 9-: 3-84 8-20-84 9- 2-84 8-27-84 8-30-84 8-30-84 8-31-84 9- 1-84 3-19-85 9-13-84 F-78 M-38 M-39 M-44 M-45 M-49 M-62 M-73 M-79 3- 7-85 9- 6-84 9- 7-84 9- 5-84 9-12-84 11-12-84 9- 5-84 8-29-84 3- 8-85 Table 2. Chronology of den entry by radio-collared (known) (known) (known) (known) 4 2 2 2 6 10 2 3 3 2 4 77 .3 23.6 nd 45.5 59.1 nd nd 36.4 36.4 nd nd (known) 2 (known) 5 5 5 6 5 2 (known) 4 2 (known) recapture recapture recapture recapture nursing cub(s) 3 cubs daughter of F-39 killed 5-5-85 0.1 km E of study area daughter of F-9 recapture recapture recapture recapture died from gunshot wounds recapture nd nd 113.6 94.1 nd 56.8 nd son of F-61 female black bears. Den entry time November October Week 1 Week 2 Week 3 1984 1 9 Week 4 2 Week 1 Week 2 Week 3 2 o Week 4 �233 Table 3. Site characteristics of dens used b~ black bears, 1984-85. Overstory Elevati on Aspect Slope Relative volume Den type vegetation estimator (m3)1 (m) (degrees) (%) Rock cavern V2 Excavated-tree Excavated-shrub N Excavated-shrub Na Excavated-treea Rock cavern-Na Rock cavern Rock caverna Rock cavern Rock cavern-N Rock cavern-Na Rock cavern-Va Excavated-shruba Rock cavern-Na Excavated-blow down Rock caverna Excavated-treea Excavated-shruba Rock caverna Excavated-tree Va Rock caverna Excavated-treea Rock caverna Rock cavern Rock caverna Portr/Psme3 Pien/Abla Quga Quga Pien/Abla Psme/Quga Psme/Quga Quga/Pied Quga/Pied Potr Psme/Quga Holodiscus Quga Pien/Abla Potr/Psme Potr Psme Prvi Potr Pien/Abla 3,050 3,111 2,562 2,440 3,294 2,593 2,623 2,501 2,577 2,623 2,410 2,776 2,333 3,172 2,867 3,325 2,928 2,577 2,989 3,233 Avalanche chute Pien/Abla Potr Psme/Potr Potr Den entrance (width x height; cm) 115 48 30 25 50 110 155 57 58 37 70 70 40 49 45 55 80 51 65 57 69 0.95 0.67 0.91 1.38 3,279 78 6 116 360 95 100 182 143 206 260 22 52 350 56 30 128 180 275 220 60 232 70 38 94 63 92 84 40 58 1.02 1.26 1.95 0.60 3.78 2.30 2.50 6.79 1.51 1.86 0.26 1.17 0.45 0.45 2.86 1.15 2.09 x 30 x 45 x 46 x 46 x 51 x 38 x 40 x 28 nd 80 x 33 103 x 33 65 xl19 78 x 42 54 x 31 43 x 35 76 x 50 45 x 45 51 x 33 57 x 27 30 x 27 41 x 33 3,203 2,547 2,684 138 50 254 60 70 100 0.08 2.74 nd 30 x 46 110 x 34 nd 2,928 30 35 4.03 42 x 27 lRelative volume estimator = width x height x length of all tunnels and chambers in the den, measured in cubic meters. This is not an index, merely an estimator. 2V = yearlings in den N = natal den 3Quga = Gambel oak Potr = Aspen Psme Pied = Pinyon pine Prvi - Chokecherry a = den used previously by bears = Douglas fir Pien ,. , Engelman spruce Abla = Subalpine fir ��Colorado Division of Wildlife Wildlife Research Report July 1985 235 JOB PROGRESS REPORT State of Colorado Project No. 01-03-048 t1larnmal s 2 Research Work Mountain Lion Investigations Job. ------------------Plan No. 6 -----------------No. 1 --------------------- Period Covered: Author: July 1,1984 Mountain Lion Population Dynamics - June 30,1985 A. E. Anderson ABSTRACT ; Eight puma were captured, weighed, measured, ear-tattooed, radiocollared and released. In addition, a 14-kg female puma caught in a commercial leg-hold trap was immobilized, weighed, measured, ear-tattooed and released. Fifteen puma radiotracked on an approximate weekly basis yielded 470 usable locations which ranged from 1 to 51 for individual puma. Since the study began, 729 usable telemetric locations have been obtained for 2 male and 5 female resident puma yearlong which ranged from 59 to 149 for individual puma. Of 13 male and 11 female puma captured, radiocollared and released since April 16, 1981, only 6 males and 7 females were alive and being radiotracked as of June 28, 1985. The elevation of male and female puma locations averaged higher May through October. However, mean distances traveled between successive locations were greater for male but not female puma May through October. The yearlong movements of 2 resident male puma were in sharp contrast; one exhibited extensive and unpredictable movements over a wide elevational range, the other generally limited and fairly predictable movements over a small elevational range. Birth intervals for 3 mothers were grossly estimated as 13, 16, and 22 months. Similarly, of 7 births from 4 mothers, 1 each occurred during May~ July, September; and 2 each·during August and December. ��237 MOUNTAIN LION POPULATION DYNAMICS Allen E. Anderson P. ~J. OBJECTIVES To assess the effects of sport hunting on mountain lion populations. SEGMENT OBJECTIVES 1. Capture and mark up to 12 mountain lions. 2. Monitor mountain lion movements. ACKNOWLEDGr~ENTS I thank D. Allen, M. Blymyer, G. Bock, C. Carroll, G. Cheney, A. Cunningham, A. Haley, R. Hicks, D. Coven, F. Fields, L. Green, G. Hanson, M. Hershcopf, J. Humphrey, J. Kane, B. Kattner, D. Kattner, Jr., D. f4asden, K. Miller, S. Music, J. Olterman, M. Potter, S. Steinert, M. Stevens, G. Stone, A. Theobald and F. Wild for their assistance. D. M. Kattner, project puma hunter, whose efforts were consistently above and beyond the call of duty, deserves a special note of thanks. I am also grateful to D. Miller, Director, Institute for Wildlife Research, National Wildlife Federation, for facilitating financial assistance from N.W.F. METHODS AND MATERIALS Methods are described in Anderson (1982) and Anderson (1983). One hundred three days were spent hunting from November 19,1984, to f4ay 17,1985. Puma were located with aerial telemetry at approximate weekly intervals and with ground telemetry opportunistically. Based on the results of estimated and actual locations of 3 radiocollared puma carcasses, 1 living puma, and 1 radiocollared hound, aerial puma locations were rated subjectively as being within a circle of l-km radius = good, 1.5-km radius = fair, and 2-km radius = poor. Estimated telemetry locations of each puma were plotted on U.S.G.S. topographic county maps (scale 1:50,000; contour interval 80 ft) or U.S.G.S. topographic quadrangles (scale 1 = 24,000; contour interval 20 ft) if location sites were densely aggregated. Linear distances between successive location sites were measured to the nearest 0.1 km. Contour intervals were read to the nearest 100 ft (county maps) or 20-ft (quadrangles). Hunting effort was allocated proportional to area among 4 previously established strata (Fig. 1 and Table 1) so that each puma within the study area had a similar probability of capture. �238 RESULTS AND DISCUSSION Allocation of Hunting Effort I failed to achieve proportional allocation (Table 2) because: (l) I underestimated the time required to recapture 4 radiocollared puma for radiocollar replacement, most of whom were within stratum 2; (2) abundant predacides employed within stratum 3 precluded use of hounds except when weather and track ing condi tions were optimum; and (3) about 7 days of hunti ng were lost because of illness and adverse weather conditions. Division of Hunting Effort Because the batteries of 4 transmitters would expire during summer, 1985, and another radiocollar required adjustment for growth, considerable effort was expanded to recapture and replace or refit the radiocollars of those 5 puma. In addition, temporary loss of hounds following some puma chases and subsequent searches required several days. Thus, of 103 days of hunting effort, 77 days were spent hunting uncollared puma, 20 days hunting radiocollared puma, 5 days searching for hounds, and 1 day attempting to capture and remove a radiocollared, transplanted puma from its capture (and presumably) sheep-killing locale (Table 2). However, during some of these activities, it was sometimes possible to capture and radiocollar puma; in fact, male puma #18 and female puma #21 were both captured during searches for hounds, and male puma #17 captured during the recapture and recollaring of his mother, puma #4. Capturing Puma - Measures of Success During 1984-85, we required 11.0 days to capture (with hounds) and radiocollar 1 puma once and 22.2 days during the relatively severe winter of 1983-84 (Table 2A). Capture rates reported in the literature mayor may not include radiocollaring or marking and 1 or more recaptures but are generally much less than ours; 4.3 days (Hornocker 1970), 8.7 days (Donaldson 1975), 6.0 days (Ashman 1975), 3.4 and 54 days (Currier 1976), and 3.6 days (Logan 1983). Another measure of success is the percentage of successful and unsuccessful pursuits by hounds (Table 2B). However, I know of no similar compilations for comparison. Puma Capture and Telmetry Eight puma were captured, radiocollared and released during 1984-85 (Table 3). In addition, a puma (#25) too small to radiocollar was immobilized, released from a leghold trap, and ear-tattooed (Table 4). On December 16, 1984, a young puma large enough to radiocollar was caught on the ground and injured by hounds to the extent that it was decided to forego the trauma of drug immobilization and handling. On December 20, 1984, this animal identified by one slightly bloody track, was apparently traveling normally through about 10 cm of snow wi th an adul t puma. Aerial and ground telemetry locations of 16 individual puma are listed in Appendix Tables A though P. Excluding those locations rated as "poor", 470 locations were judged usable for computing home range areas (Table 5). As of �239 June 28, 1985, 3 male and 6 female resident puma tracked yearlong or approximately yearlong, have 39 to 149 locations per animal (Table 6). Forty locations have been estimated as minimal for certain contemplated statistical tests (Gustafson and Fox 1983) and 100 to 200 locations for reliable estimates of the home range area of large carnivores (Bekoff et a1. 1984). Puma ~1orta1ity Of 13 male and 11 female puma captured, radioco11ared and released since April 15,1981, only 6 males and 7 females were alive and being radiotracked as of June 28, 1985 (Table 7). The tabulation of this mortality by sex and age class shows a preponderance of males dying among puma less than 24 months of age (Table 8). Thus, excluding the 2 instances of unconfirmed mortality and 1 of unknown status, sex and age class of definitely known mortality was 4 males and 1 female <24 months of age and 2 males and 1 female >24 months of age. Neither sex ratio, however, differs significantly (~ < 0~05) from equal ity. Puma Activity and t~ovement Elevations of approximate location sites for 7 adults overlapped between the November-Apri 1 and r~ay-October periods, but thei r means were 20.4 (#18) to 287.1 m (#5) greater during the May-October period (Table 9). Mean linear distances traveled between successive locations were greater during the MayOctober period for the 2 males (3.9 km - #5 and 4.2 km - #18) but slightly less for each of the 5 females. During each period, mean elevations of all puma locations were much more variable than the mean linear distance traveled between them (Table 9). Extremes of variation in mean distance traveled occurred during the May-October period among the 2 males; coefficients of variation ranged from 49.3% (#5) to 113.6% (#18). The latter value is largely owing to a 47.0 km journey recorded on June 21,1985. A similar distance traveled by #18 but not entered into these calculations because of a poor location, was recorded on November 30, 1984 (Appendix Table H). Except for these 2 movements, yearlong locations of #18 suggest he moved mainly between the Horsefly, Happy, and Dolores drainages and generally remained within each drainage for about 1 to 3 weeks with little elevationa1 change. This predictability is in contrast to_male #5 whose movements have been extensive and unpredictable within an area of about 37 km in length and 16 km in width. Although not readily converted to numerical analysis, inspection of the plotted locations of the 7 puma in Table 9, suggests subtle annual changes in their distribution within their respective home ranges. Interestingly, no radiocollared puma have been located in the considerable land area above 2,684 m but at least 3 have passed through that area. Subjectively, the winter of 1984-85 was much milder than in 1983-84. Possible weather-induced variation in the approximate mean elevations and mean linear distances moved in successive locations among the 5 adult puma present during both winters (November-April) is explored in Table 10. Based on 95% confidence intervals computed about the coefficients of variation (%), I conclude that weather appeared to have been a minor influence on this variation. �240 Puma Home Range No progress was made on a comparative analysis of the various methods of calculating home range area (Anderson 1984). This will be a priority activity during the next fiscal year. The inferred home range of male puma #5 overlapped female puma #s 4,6, 7, 12 and 15. Since the demise of male puma #14, no other resident male appeared to be present within the home range of #5. However, subadult male #20 has been a presumably temporary resident and mature male #18 a rare visitor to the inferred home range of #5. Reproduction A compilation of the reproductive history of 7 mature female puma (Table 11) suggests that 3 females gave birth at 13, 16, and 22 month intervals. Of 7 births from 4 mothers, 1 each occurred during May, July and September and 2 each during August and December. The foregoing can only be regarded as gross approximations. Information on litter size is too fragmentary to summarize. Cervids Killed by Puma The location and age structure of 7 mule deer found killed by puma (Table 12) included 4 heavily scavanged and uncovered carcasses whose age and sex could not be determined. Two of the 3 carcasses identified by age and sex were almost completely covered by needle litter of pinyon pine (Pinus edulis) while the third was uncovered. Since 1981,40 mule deer and 2 elk have been found which were killed by puma (Table 13). Of the 40 mule deer, 19 were less than 18 months of age. Their sex ratio (60:100) did not differ significantly (t < 0.05) from equality. However, the sex ratio (33:100) of the 18 deer more than 18 months of age did differ significantly (~ < 0.005) from equality. LITERATURE CITED Anderson, A. E. 1982. Mountain lion investigations--mountain lion population dynamics. Colo. Div. ~~ildl. Res. Rep. July, Part 2:143-159. • 1983. Program narrative. Project no. 45-01-503-15050, Work Plan 6, ---'-Jo'b.1. Noncervid Section, Research Center, Colo. Div. Wildl. Fort Collins. 18pp + 2 Figs. + 2 Appendices. 1984. dynamics. Mountain lion investigations--mountain lion population Colo. Div. Wildl. Res.•Rep. July, Part 2:221-268. Ashman, D. 1975. Mountain lion investigations. Job Performance Rep., P-R Project W-48-6, Study S & I, Job 5. Nev. Fish and Game Dep. 18pp. Bekoff, M., T. J. Daniels, and J. L. Gittleman. 1984. Life history patterns and the comparative social ecology of carnivores. Ann. Rev. Ecol. Syst. 15:191-232. �241 Currier, M. J. P. 1976. Characteristics of the mountain lion population near Canon City, Colorado. ~l.S. Thesis, Colorado State Univ., Fort Collins. 8lpp. Donaldson, B. 1975. Mountain lion research. Job. Prog. Rep., P-R Project W-93-R-l7, Work Plan 15, Job 1. New Mexico Game and Fish Dep. l8pp. Gustafson, K. A., and L. N. Fox. 1983. A comprehensive interactive program for calculating and plotting home range and distribution. Pages 297-317 ~ D. G. Pinock, ed. Proc. 4th Intl. Conf. Wildl. Biotelemetry. pp. Hornocker, M. G. 1970. An analysis of mountain lion predation upon mule deer and elk in the Idaho Primative Area. Wildl. Monogr. 21. 39pp. Kufeld, R. C., J. H. Olterman, and D. C. Bowden. census for mule deer on Uncompahgre Plateau. 1980. A helicopter quadrat J. Wildl. Manage. 44:632-639. Logan, K. A. 1983. Mountain lion population and habitat characteristics in the Big Horn Mountains of Wyoming. M.S. Thesis, Univ. Wyoming, Laramie. 101 pp, r~asden, D. 1982. Units 61 and 62 (DAU) deer quadrat census. (~1emorandum to J. Olterman of June 1.) CDOW SW Regional Office files, f4ontrose. Unpaged. Prepared by: LLLhz._~ e. ~ JfffenY.'Ande~sor;j$ • Wildlife Researcher C �242 o 10 20 KI LOMETERS Fig. 1. Unit 62 midwinter deer census area showing 8 strata and 0.6475-km2 quadrat allocation within strata. From Masden (1982) after Kufe1d et al. (1980) . �243 Table 1. A 11ocati on of hunting effort for puma, 1984-85, Uncompahgre Plateau (GMU 62). Strata area (km2) Percent of total area Total number hunting days 1 418.3 24.8 27 = 2 578.9 34.3 38 = 3 4 356.8 334.1 21. 1 19.8 23 22 1,688.1 100.0 Strata numbera 1, 2 3, 4 5, 6 7, 8 110b aKufe1d et al. (1980) and Masden (1982), see Fig. 1. bEstimated days likely to be actually hunted from November 19, 1984, through May 17, 1985. Number of hunting days projected for each stratum obtained by multiplying 110 by percent of total area for each stratum. �Table 2. Chronology of puma hunting effort by strata, 1984-85, Uncompahgre represents one day of hunting effort.a Each date Plateau (GMU 62). Strata and dates 1 Month Tota 1 2 Total 3 Nov. 29 1 l2_,_gQ_,n,27 4 Dec. 13,14,17 3 4,5,6,7,10 5 Jan. 7,8,9,N, 14, 15 ,IT, 30 8 3,28 2 Feb. 6,7,28 3 4,5,12,13, 14, 18 ,20, 22,25 9 t~arch 3,4,5,13, 30,31 6 11,12,18,19 20,21,25 7 7,10 Apri 1 1,g,,9,24 5 8,10,11,12, 8 15 0 1,2,3,6,7,8, 9,1'0=:1"'1,13-:14,15,16,17 14 allocation Total Grand total 1 28,30 2 8 0 1,2,3,9,11, 12,16,31 8 16 6 1 1,2,5,20, 22,23 6 17 19 1 11 ,21 ,23,24, 26,27 6 19 2 0 15 1 0 14 0 0 14 26 - 49 6 - 26 22 103 27 38 23 22 110 - Target _- 4 l§_, 17 ,~, 30 26 May Tota 1 aSing1e underlined dates indicate major effort was spent attempting to recapture radioco11ared puma to replace or refit radioco11ar; double underlined dates indicate major effort was made to find lost hounds; triple underlined indicates major effort was to recapture a transplanted puma (#19). See text. N .p. .p. �245 Table 2A. Rate of radi oco 11aring puma using hounds GMU 62, 4-9-81 to 6-28-85. Period No. days Radiocollaring No. From To radiocollared ratea hunteda 4- 9-81 12-14-81 1- 2-83 11-19-83 11-19-84 4-29-81 2-21-82 5- 6-83 5-17-84 6-28-85 Total 1 3 7 5 7 23 16.0 10.7 16 32 91 111 13.0 22.2 77 11.0 327 14.2 aNo. days to radiocollar 1 puma; does not include replacing or refitting 4 radiocollars. Table 2B. The history of hound l2ursuit of l2uma, G~1U 62 4-9-81 to 6-28-85. Hunter A N Successful pursui ts B % N % N % N % 0 0 0 0 18 10 29.5 16.4 23 10 28.8 12.5 0 4 32 6.6 52.5 5 38 6.2 47.5 Radi oco 11ared Recaptureda 5 27.8 0 0 Treed or b bayed only 1. Total Unsuccessful pursuits Total pursuits Total C 6 5.5 33.3 12 66.7 100.0 29 47.5 42 52.5 18 100.0 100.0 61 100.0 80 100.0 aAnimal radiocollared; recaptured refit radiocollar (2). 0 inadvertently (4) or to replace (4) or bAnimal not radiocollared; either escaping (3) or released because of probable injury which further handling might exacerbate (1). �N .j::- Table 3. C)'\ Details on the ca~ture and bod~ measurements of 8 ~uma radiocollared 1984-85, Uncom~ahgre Plateau (GMU 62). Ear tatoo number Sex Date of capture Estimated age (months) Legal descr. capture site 1/4 S T R U.T.M. capture site X Y Elevation (m) capture site Drug (2:1 ketamine-rompun) Dosage (cc injected) Induction time (min) Radiocollar serial no. Transmitter freq. (MHz) Body wt (kg) Measurements (cm) Total body length Tai 1 length Head-body length Chest girth Neck circumference Height at shoulder Head length Zygomatic breadth Ear length Hind foot 19a 20 21 22 23 24b 26 27c F M 11-26-84 10 F 12- 7-84 12 M 1- 5-85 15 M M 1-23-85 18 M 3-20-85 10 F 4-24-85 14 NW 2 47N 9W SE 27 48N 11W NW 12 50N 14W NE 30 46N 8W NE 32 14S 99W SW 17 46N 8W NE 14 50N 14W SW 29 51N 14W 253 4249 1890 756 4252 2286 727 4277 2134 256 4232 2347 718 4297 2195 257 4235 2108 726 4275 2074 721 4281 2377 3.3 8 8771 149.6710 56.0 3.1 12 8390 149.5020 44.0 6.0 50 8773 149.7210 7-18-84 7 3.0 5 12908 149.9510 28.0 190 72 118 55 32 67 20.0 12.7 9.5 27.5 5.0 - 12899 149.8600 44.5 206 81 125 70 39 69 3.5 10 12905 149.9200 36.8 193 77 116 63 35 - - 14.4 8.0 27.0 12.1 8.1 24.5 215 85 130 72 38 77 20.6 14.0 9.0 30.0 1- 9-85 12 195 79 116 69 39 74 22.0 14.0 9.0 29.0 194 74 120b -b 72 23~0 9.5 30.0 5.0 40 12906 149.9305 44.0 180 67 113 66 36 66 19.5 13.9 9.0 27.0 3.0 12 8769 149.6310 36.3 187 75 112 66 34 68 18.6 12.0 8.0 25.5 aAnimal captured in leghold trap at site of sheep killing by ADC, USFWS trapper. Moved 45 linear km from capture site to upper Criswell drainage SW 1/4, S18, T48N, R13W, 732-4254, 2806 m and released. Shot in GMU 70 as a sheep killer 1-15-85. bMany healing scratches on badly swollen shoulders, neck, and head; chest girth - 81 cm, neck circumference 54 cm, zygomatic breadth Died on date of capture. 15.5 cm. Left upper canine broken off within its socket but healing externally. cHeel pad from right front foot gone; wound completely healed. �247 Table 4. Physical characteristics of female puma (25) about 4 months in estimated age; ear-tatooed, and released from a leghold trap on 1-24-85. Characteristic Measurement Body wt (kg) Measurement (cm) Total body length Tail length Body 1ength Chest girth Neck circumference Height at shoulder Head length Zygomatic breadth Ear length Hind foot length 14 128 51 77 42 22 52 15.0 10.2 6.0 21.0 Table 5. Number of aerial telemetric locations of 15 puma radiotracked 7-6-84 through 6-28-85, GMU 40, 61, 62, 65, 70. GMU locations Ear tatoo Capture Est. Age No. locations number date Sex Majority Minor on 6-85 1984-85 4 5 6 7 12 15 17 18 19 20 21 22 23 26 27 1-21-82 2- 7-83 2-10-83 3- 4-83 3-25-83 12-15-83 4-14-84 4-16-84 7-18-84 11-26-84 12- 7-84 1- 5-85 1- 9-85 3-20-85 4-24-85 F M F F F F M r~ F M F M M M F 65 89 58 75 86 78 17 50 12 17 17 20 17 13 16 49a 51b 47c 49d 48 51e lf 48g 8 26 29 24 21 11 10 470 62 62 62 62 62 62 62 62 62 62 62 65 40 62 62 0 0 0 0 0 0 61,40 61 ,70 62 62 61 aIncludes 1 visual and 1 ground telemetry location. blncludes 2 ground telemetry locations. cIncludes 2 ground tel ernetry locations. dIncludes 1 visual location. eIncludes 1 ground telemetry location. fShot by a sport hunter 1-11-85 in GMU 42; 87 km distant from capture site, age is at death. gIncludes 3 ground telemetry locations. Animal shot as a sheep killer 1-15-85 in GMU 70; 54 km distant from capture site, 31 km distant from transplant site; age is at death. �Table 6. Number of radiolocations and vicinity; GMU 62,61. of resident puma where N approximated >40, Uncompahgre Plateau Period of surveillance Sex Ear tatoo no. Date radiocollared From To Number of radiolocationsb M 5 2- 7-83 2-10-83 6-28-85 112 M 13a 4- 7-83 4-15-83 1-25-84 39 M 18 4-16-84 4-20-84 6-28-85 59 F F 3 4 1- 8-82 1-21-82 1-15-82 2-15-82 4-15-83 6-28-85 40 149 F F 6 7 2-10-83 3- 4-83 2-17-83 3-10-83 6-28-85 6-28-85 112 F 12 3-25-83 4- 1-83 6-28-85 106 F 15 12-15-83 12- 16-83 6-28-85 75 aLocations 116 primarily in GMU 61; locations of all other puma in GMU 62. bLocations made for 1 year or longer total 729 for 2 males and 5 females. Comments Replaced radiocollar 5-17-85 A poor ground telemetry location 9-5-84 Located 40 km N of previous home range 6-21-85 Last signal on 4-15-83 Replaced radiocollar 4-14-84 Replaced radiocol1ar 5-14-85 Replaced radiocollar 4-30-85 N .j::-- <Xl �24<; Table 7. Twenty-five puma captured 4-16-81 to 4-24-85, Uncompahgre Plateau (G/·1U 62). Radioco11ar Estimated age Date Ear tatoo (months) e Frequency collared number Status Sex at capture Number 1-21-82 Disappeared after 8-12-82 Killed by hunter 1-10-82 Disappeared after 4-15-83 Alive 8389 149.5500 4-16-81 8772 149.7010 1- 5-82 2 8775 149.8005 1- 8-82 3 8773 149.7215 F 20-21 unknown 26 Unknown F 24 With litters of 2 & 1 on 12-14-83 and 1-24-85, respectively 149.7815 2- 7-83 Alive M 60+ 12904 12909 149.9090 149.9605 2-10-83 3- 4-83 Alive Alive F F 30 48+ 12901 149.8810 2-20-83 8 M 6 12900 149.8705 3-23-83 10 M 6 12911 149.9800 3-25-83 Died during recapture (for collar adjustment) 8-4-83 Found dead of unknown cause(s) 6-2-83 Alive F 60+ 12896 149.8200 4-13-83 13 Unknown, last Signal on 2-20-84 M 10+ 12906 149.9310 12-12-83 14 M 84+ 12902 149.8900 12-15-83 15 Found dead 6-22-84. Possible predacide victim (see text). Al ive F 60+ 12897 12910 149.8310 149.9710 1-15-84 4-14-84 16 17 M 36+ 1·1 8 12903 12908 149.9010 149.9510 4-16-84 7-18-84 M F 36 12899 12905 8771 8390 149.8600 149.9200 149.6710 149.5020 11-26-84 12- 7-84 1- 5-85 1- 9-85 20 21 22 23 Found cannabilized Ki11ed by hunter 87 km linear distance from capture site on 1-11-85. Al ive Killed 1-15-85 by USFWS ADC hunter as a sheep killer in Naturita Ck. A1 ive Al ive Alive Alive M F M M 10 12 15 12 8773 149.7210 1-23-85 24 M 18 1-24-85 25 3-20-85 4-24-85 26 27 12906 8769 dfi4 149.9305 149.6310 Immature 60 8774 Found dead 1-26-85; was severely wounded by another puma when captured (see text) Too small to radi 0collar; ear tatooed only. (See text; Table 11). Alive Alive Breed ing status and comments 4 M F 10 14 Unknown 2, 15-30 1b cubs at capture, mother of 8. On 3-14-84 and 5-9-84, tracks indicated another litter of 2 t·/ i th Appeared pregnant at recapture 2-26-84, mother of 13 and 1 other cub Offspring of #12 2 kittens 5-12 days of age on capture Offspring of #4 Extensive movements including 1 visit to capture site 11-28-85 Offspring of #15 Offspri ng of ."7 Resident within GMU 40 since 2-85 Caught in a commercial trapper's leghold trap. Offspring of #4 Offspring of #12 Unknown recaptured and radioco11ar replaced 4-14-84; 12898,149.8400. b'5 recaptured and radiocol1ar replaced 5-17-85; 8772-01,149.7010. c'7 recaptured and radioco11ar replaced 5-14-85; 12901-01,149.8810. d'12 recaptured and radiocollar replaced 4-30-85; 8773-01,149.7210. eBased on aerial telemetric locations and compiled information as of 6-28-85. fCaptured at ~ite of sheep kill with 1eghold trap by USFWS, ADC trapper; tranquilized, measured and transplanted by me to a site about 37 km distant and 196 m higher in elevation. �.t..JV Table 8. Sex and age class of 25 puma at capture, 4-15-81 to 6-28-85, Uncompahgre Plateau (GMU 62) and their subsequent fate. Numbers are ear tatoos of individuals. Year Less than 24 months of agea 24 months and older Total Sex M 1981-82 a Total a - F M F lb a 2c 3d 4 3 a 4 8e lOf 13g a 5 6 7 12 Total 3 a 1 3 1983-84 17h a 15 Total 1 a 14 ~ 16J 18 3 19k 21 25m 27 a a Total 20 22 231 24 26 5 4 a a 9 Total 9 5 4 7 25 1982-83 1984-85 1 apumas a~ la, 17, 19 were 6-8 months in estimated age. bl unconfirmed mortality. c2 illegally shot. d3 unknown. e8 died during recapture 8-4-83. flO found dead 6-2-83, cause(s) unknown. g13 unconfirmed mortality. h17 shot by sport hunter 1-11-85. i14 suspected predacide found 6-22-84. j16 probable cannabalism found 4-2-84. k19 shot as sheep killer 1-15-85. 124 died day of capture; severely injured a few days before capture presumably by another puma. m25 - 4 months of age but not radiocollared. 7 5 �251 Table 9. Seasonal chanqes in elevation and linear distances between successive locations amonq 7 adult ~ radiotrackp.d at approximate weekly intervals yearlonq, 1984-85. Uncompahgrg_?lateau (GMU 62~ Ear tatoo number Season ---------------------_._._-12 15 Statistic 5 18 7 1984-85 4 6 Nov-Apri 1 (snow) Sex E1ev. N (m) x SO Min Max Range Oistance (km) N x SO Min Max May-October (no snow) E1ev. N (m) SO Min Max Range Distance (km) aA11 females N X SO Min Max M M Fa Fa 27 2073.8 146.2 1829 2316 487 27 10.15 5.94 2.1 22.3 25 2232.4 75.5 2098 2347 249 25 4.19 2.51 0.6 7.6 26 2206.3 123.6 2042 2499 457 26 3.77 2.21 1.1 8.8 27 2166.4 152.4 1951 2499 548 27 3.50 2.57 0.3 13.4 24 2360.9 180.6 1920 2652 732 24 9.82 4.85 1.9 20.5 23 2252.8 153.8 1920 2499 579 23 8.37 9.48 0.9 47.0 23 2328.3 103.3 2103 2499 396 23 3.69 2.89 0.2 9.9 20 2439.7 98.1 2256 2560 304 20 3.08 I.91 0.6 6.7 Fa Fa 26 2068.1 140.2 1707 2316 609 26 5.52 3.22 1.4 13.0 25 2184.9 155.6 1859 2499 640 25 6.14 3.34 0.3 15.5 25 2191.1 137.4 2012 2560 548 25 3.36 2.22 0.6 7.9 23 2313.8 132.6 1951 2560 609 23 5.42 3.69 0.3 14.7 23 2361.5 133.3 2073 2560 487 23 4.96 3.50 0.6 13.9 26 2307.0 151.1 1951 2621 670 26 2.62 2.03 0.3 7.9 were mothers with young, Table 10. Coefficients of variation (+ 95% confidence interval) of seasonal and annual mean elevations (m) and mean linear distances (km) traveled between successive locations by 5 adult puma, Uncompahgre Plateau (GMU 62). Ear tatoo number Year Season Variable 1983-84 Nov-Apr e1ev 1984-85 Nov-Apr e1ev 1983-84 Nov-Apr distance 1984-85 Nov-Apr distance 1983-84 May-Oct e1ev 1984-85 May-Oct e1ev 1983-84 May-Oct distance 1984-85 May-Oct distance 4 8.12 5.6- 10.7 7.04 5.1- 9.0 79.2 54.4-104.0 58.5 42.1- 74.9 8.41 5.9- 10.9 7.64 5.4- 9.9 63.4 44.5- 82.3 49.3 34.6- 64.0 5.05 3.4- 6.7 5.60 4.0- 7.2 100.3 67.1-133.5 58.6 41.9- 75.3 3.86 2.7- 5.0 4.40 3.0- 5.8 108.0 75.8-140.2 78.3 54.4-102.2 12 6 7.29 5.0- 9.6 7.03 5.1- 9.0 72.2 49.6-94.8 73.4 52.9-93.9 5.88 4.1- 7.6 4.02 2.7- 5.4 68.8 48.3-89.3 62.0 41.5-82.5 7.32 5.2- 9.5 6.78 4.8- 8.7 64.4 45.6-83.2 58.3 41.6-75.0 4.70 3.4- 6.0 5.73 3.9- 7.5 67.7 47.9-87.5 68.1 47.3-88.9 7.34 5.0- 9.6 7.12 5.0- 9.2 76.6 52.6-100.6 54.4 38.5- 70.3 4.62 3.2- 6.0 5.64 3.9- 7.4 55.2 38.7- 71.7 70.5 48.9- 92.1 �Table 11. Known and inferred history of the reproductive Plateau (GMU 62), 1982-85. Ear tatoo number Date captured Est. age (mos) S tatus on 6-28-85 shot 1-10-82 disappeared 4-15-83 olive 3 1- 5-82 1- 8-82 40+ 26 4 1-21-82 24 2 Date status of radiocollared N 20 alive 7 3- 4-83 48 alive 12 3-25-83 60 0 2 2 2-20-83 2? 3-14-84 5- 9-84 12- 7-84 2 2 4-13-83 1 2-26-84 0 60 alive 12-15-83 6-22-84 11-26-84 aWt. visual estimate under poor conditions. young 18-25° June-Aug 1983 36.2 Aug 1983 No evidence of lactation at capture. #4 treed but no evidence of young. Large difference in size of treed young. 8 unk M 4 F 14.0 Sept 1984 6 M 25.9 Aug 1982 12 F 36.8 Dec 1983 9 M 36.3 July 1982 10 M 44.0 May 1984 unk unk Dec 1983 Young unobtainable 44 Dec 1983 Tracks of 1 young vicinity of #15. Radiocol1ared #20, left #15 in April 1985 at about 16 mos of age. 4-6 1? 2 N Comments and radiocollared No indication of lactation. No indication of lactation at capture. o alive 4-30-85 12-15-83 Est. mots) year of birth o 3-20-85 15 Postnatal young Est. age wt (mos) Sex (kg) o 1-24-85 2-10-83 24 months estimated age at capture, Uncompahgre - o 5- 2-83 12-14-83 4-14-84 6 puma> 5-12 days 11 M Radiocollared #17, tracks of another young; #17 left mother early July 1984, shot 1-11-85, GMU 42. Released from leghold trap, ear tatooed #25, with #4, current status unknown. Suspect young born during late summer, 1984, based on localized movements of #6. On 2-12-85 tracks of adult and 2 young within her home range but recei ved no te 1emetry signa 1. 1 of at least 2 young believed with mother prior to her radiocollaring. This radiocollared puma (#8) died on 8-4-83. When #7 was radiocol1ared, #8 and at least lather young was with her. Based on tracks and telemetry. Based on tracks and telemetry. Radiocollared #21 with another young. #21 may have left #7 in May 1985 at about 17 mos of age. No indication of lactation at capture. Radiocollared #13, left mother July 1983 at about 14 months of age, unconfirmed mortality late 1984 in GMU 61. Bayed, no indication of young but appeared pregnant. Radiocol1ared #26, left mother June 1985 at about 13 mos of age. Recaptured and recol1ared #12, no indication of lactation. Tracks and telemetry indicated absence of #26 and presence of 1 young. in crevice. V1 N �253 Table 12. Discovery date Seven mule deer found .killed by puma, 1984-85, Uncompahg_re Plateau (GMU 62). Es t. U. T.t1. Approx. Leg"l descr. age elev. Body pa rts Sex (mos) 1/4 S T R X Y Habitat remaining (m) 12-5-84a 6 SW 50N 13W 729 4276 P-J chaining 2256 all except viscera P-J chaining P-J 2225 only hair, puma tracks, drag path skeleton but no head P-J P-J chaining P-J chaining P-J sagebrush 1951 2256 1-15-85 unk unk SW 23 ·15S 99W 722 4290 1-15-85 unk unk Nvl 26 15S 99W 722 4288 3-12-85 4- 1-85 unk unk unk unk NW SE 36 17 51N 15S 13W 99W 737 721 4280 4291 4-18-85 M 108+ SE 8 49N 12W 741 4267 5- 1-85a M 11 NW 29 50N 13W 731 4272 aAlmost completely covered wi th Table 13. Fisca year J 1:.. 2134 stomach, hide skeleton and hide fragments skeleton, most of skull all except viscera 2103 2196 edulis needles. Age class and sex of cervids killed b~ puma, Uncompahgre Plateau (Gt·1U62); 1981-85. 18+ mos < 12 mos 13-17 mos Sex Age Sex and Speci es M F F F unknown unknown age unknown M M Total 1980-81 mule deer el k 0 0 1 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 3 0 1981-82 mule deer elk 0 0 .D 0 0 0 0 2 0 6 0 0 0 0 0 0 0 9 0 mule deer el k 0 0 0 0 2 0 0 0 5 0 0 0 0 0 0 0 9 0 mule deer elk 3 0 4 0 0 0 0 0 2 1 0 0 0 0 12 2 mule deer elk 0 0 0 0 0 0 4 0 0 Total mule deer 5 4 40 Total elk 0 0 2 1982-83 1983-84 1984-85 1 0 0 0 0 0 0 3 3 15 0 aNearly all ki 11s found from November th rough May. 0 0 0 0 0 �254 Appendix Table A. Aerial telemetry locations of adult female puma #4. Legal descr. U.T.M. Approx. Distance (km) elev. between Major Date 1/4 S T R X Y (m) locations drainage 2377 2225 2438 7- 6-84 7-13-84 7-20-84 NE SE SE 20 24 13 47 47 47 10 10 11 238 245 758 4244 4243 4245 7-27-84 8- 6-84 SE SE 7 47 47 10 10 760 760 4247 4247 8- 9-84 8-17-84 8-24-84 8-31-84 9- 7-84 9-14-84 NE SW NE SW NW NE 26 24 23 24 25 18 47 47 47 47 47 47 10 10 10 10 10 10 243 244 243 244 243 760 4442 4243 4244 4243 4243 4246 2347 2103 2316 2377 2286 9.5 2.4 2.4 1.2 0.2 9-21-84 9-28-84 10- 5-84 10-13-84 10-19-84 10-26-84 11- 2-84 11-10-84 11-16-84 11-23-84 SE SW NE NE SW SE NE SW SW SE 21 23 23 27 14 12 24 19 25 19 47 47 47 47 47 47 47 47 47 47 10 10 10 10 10 10 10 9 10 10 240 242 243 241 242 245 245 245 244 760 4243 4244 4244 4243 4245 4247 4244 4243 4242 4243 2438 2225 2347 2438 2316 2164 2195 2286 2134 2438 4.1 2.5 0.8 1.2 2.8 2.8 2.3 1.3 1.8 8.0 11-30-84 12- 7-84 NW SE 9 24 47 47 10 11 239 758 4248 4244 2195 2499 5.1 5.8 12-16-84 12-21-84 12-29-84 1- 4-85 1-11-85 1-18-85 1-25-85 1-24-85 2- 1-85 2- 8-85 2-15-85 2-22-85 3- 1-85 3- 8-85 3-14-85 3-22-85 3-31-85 4- 5-85 4-11-85 NW NW NE NW NW SE SE NW NE SW NW NE NW NE SW NW NE NW SE 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 11 241 241 240 241 239 240 239 238 240 241 762 762 239 240 238 239 240 239 755 4250 4250 4249 4253 4252 4251 4246 4248 4251 4250 4248 4247 4252 4250 4251 4252 4250 4252 4251 2103 8.8 28 33 29 28 33 28 27 48 48 47 48 48 48 47 47 48 48 47 47 48 48 48 48 48 48 48 2225 2073 2103 2134 2256 2286 2042 2103 2316 2225 2073 2103 2164 2134 2073 2164 2316 1.4 3.9 4.8 1.8 1.1 4.6 4.4 1.3 3.8 1.1 5.5 2.5 3.7 1.7 1.8 3.8 6.5 4-19-85 4-27-85 NW NE 29 47 47 10 10 761 761 4247 4243 2286 2438 6.7 4.5 5- 3-85 5-10-85 5-16-85 5-25-85 SW NW SW NW 30 47 47 47 47 10 10 10 10 239 244 761 759 4248 4246 4248 4242 2196 2225 2347 2499 5.5 5.6 7.9 6.2 5-31-85 SE 20 47 10 761 4244 2438 2.9 6-10-85 NE 29 47 10 239 4243 2469 0.5 6-14-85 SE 18 47 10 760 4245 2286 2.8 6-21-85 NE 18 47 10 760 4246 2347 0.4 6-28-85 NW 8 47 10 761 4247 2347 2.0 34 34 4 27 28 28 8 8 33 34 8 8 8 4 13 5 3.5 7.7 9.9 Rating Signal Location Spring Dolores Wes t Fk Spring Spring Spring G p p Happy Happy Happy Happy Happy West Fk Spring Spring Happy Happy Happy Happy Happy Happy Dolores Dolores East Fk Spring Spring Middle Fk Spring Spring Spring Spring Divinney Lindsay Spring Spring Spring Spring Spring Spring Spring Lindsay Spring Lindsay Lindsay Spring Lindsay East Fk Dry Ck Spring East Fk Spring Spring Happy Spring Middle Fk Spring East Fk Spring East Fk Spring Middle Fk Spring Middle Fk Spring Spring F F P F F F G G F F G P G F G F p p Ground Telemetry F F G F G F F F P F G G G F F F F F P F F F F F G F F P G F F F F F G G Visual F F F G G F F F G F F F F F F G G F F G G F F F F F F G G F F G F F F G F G G G F G F G F �:L55 Appendix Table B. Aerial telemetry locations of adult male puma #5. Legal descr. U.T.M. Approx. Distance (m) e1ev. between y (m) locations Date 1/4 S T R x 2347 2560 8.5 2.7 5.8 15.0 Potter Moni tor Potter East Fk. Escalante 4265 4258 4261 4261 4264 2408 2560 2012 2530 2530 12.3 7.0 7.0 9.1 13.5 741 744 738 742 748 743 744 742 751 4262 4257 4262 4264 4259 4268 4259 4262 4256 2316 2256 2499 2195 2286 1920 2286 2073 2286 20.5 5.7 6.8 5.0 8.2 10.7 9.7 3.4 10.6 12 12 11 740 738 754 4261 4272 4257 2316 1920 1920 12.0 12.0 21.6 50 49 49 49 12 13 12 11 738 735 739 750 4273 4269 4265 4268 1951 2103 2316 1951 22.3 4.8 4.5 11.4 49 49 48 11 11 11 752 756 754 4267 4265 4258 1920 1829 2073 2.1 4.9 8.0 49 49 48 49 50 49 11 11 11 12 12 11 49 49 49 50 49 49 11 13 12 13 11 13 752 755 754 744 742 751 754 736 742 738 751 737 4264 4266 4258 4266 4270 4263 4264 4269 4266 4270 4263 4266 2012 1859 2103 2103 1951 2042 1951 2012 2073 2225 2042 2164 6.2 4.9 8.1 13.7 4.5 11.4 2.7 18.8 6.5 5.2 14.9 14.5 Moni tor Criswe11 Wright Potter East Fk Escalante Traver Bull Criswe11 Roubideau Cushman Roubideau Terrible Traver West Fk Dry Ck Traver Monitor Eas t Fk Dry Ck Monitor Monitor Cri swe 11 Big Sandy Wash Coalbank Dry West Fk Dry Ck Cushman Dry Ck Dry Ck Roubideau Criswe 11 Cushman Dry Ck Moni tor Moore Monitor Cushman Potter 11 13 12 12 13 750 734 741 745 735 4258 4271 4259 4258 4269 2286 2225 15.1 20.8 35 48 50 48 48 50 2377 2286 2134 14.0 4.1 15.0 19 11 33 28 3 18 49 48 49 49 48 49 12 12 13 13 12 13 739 748 733 733 746 729 4263 4257 4260 4262 4259 4265 2286 2377 2652 2469 2256 2377 7.3 10.5 14.8 1.9 7- 6-84 7-13-84 7-20-84 7-27-84 NW NW SE SW 8- 2-84 8- 9-84 8-17-84 8-24-84 8-31-84 SW NW SW NE 9- 7-84 9-14-84 9-28-84 10-13-84 10-19-84 10-26-84 11- 2-84 11-10-84 11-16-84 SW NW 29 NE 36 21 2 11-23-84 11-30-84 12- 7-84 NW NW NE 12-16-84 12-21-84 12-29-84 1- 4-85 NW NW 30 N,/ 18 NW 8 1-11-85 1-18-85 1-25-85 SW SW SW 13 2- 1-85 2- 8-85 2-15-85 SW NW SE 21 14 2-22-85 3- 1-85 3- 8-85 3-14-85 3-22-85 3-31-85 4- 5-85 4-11-85 4-16-85 NE 15 33 28 22 4-19-85 4-27-85 5- 3-85 5-10-85 5-16-85 SW SW NW SW SW 5-25-85 5-31-85 6-10-85 6-14-85 6-21-85 6-28-85 SW NE NW NE SE NW SW SE SW NW SE NW SE SW SW NW NE SE SE SW NW Rating Major drainage 28 11 8 15 3 28 33 19 9 4 4 28 7 32 30 9 2 9 4 4 2 8 36 21 13 6 27 6 3 48 49 49 49 13 13 13 14 733 732 736 721 4259 4262 4266 4266 2591 2438 49 48 49 49 49 13 13 12 13 14 734 736 742 733 720 49 48 49 49 48 49 48 49 48 12 12 13 12 12 12 12 12 11 49 50 48 12.7 17.7 Piney Ck Cottonwood Wright Bull Little Monitor Criswell Cushman Potter Potter Roubideau Cottonwood Signal G Location F F F G G F F Ground Telemetry G F G F G G F F G F G F G F G F F F F G G G F G F G G G F G F F G F F F G F P F G F G F F F G F G F G F G G G F G G G G G F P F G F G G Ground Telemetry F F F F G F G F F F G F F F F F G F G F G F G F �256 Date Aerial telemetry loations of adult female puma #6. Legal descr. U.T.M. Approx. Distance (km) Major elev. between drainage S T R X Y (m) locations 1/4 7- 6-84 7-13-84 7-20-84 7-23-84 SW SW SW NW 7-27-84 8- 9-84 8-18-84 8-24-84 9- 3-84 NE 26 49 13 Criswell 738 4263 2377 1.3 NW 24 49 13 738 4264 2377 1.3 Criswe11 NW 34 49 13 Potter 734 4260 2591 4.6 SE 35 49 13 Criswell 737 4260 2560 2.5 No specific location but within Dry Fk Escalante; probably vicinity Iron Spring Mesa SW 5 48 14 Dry Fork Escalante 34 49 13 NW Potter 2.5 2286 734 4261 31 49 12 SE Traver 5.6 2377 740 4260 30 49 12 NE Moore 2.2 2377 740 4263 NW 48 12 Terrible 6.1 2499 741 4257 NW 4 49 12 Criswe11 742 4269 13.4 1951 NW 20 49 12 Moore 1981 741 4265 4.6 16 49 12 NE Roubideau 2073 742 4266 2.2 29 49 12 NW Traver 3.7 2347 740 4263 NW 29 49 12 Moore 0.3 2377 740 4262 Criswell 3.2 2377 738 4264 24 49 13 NW NW 24 49 13 738 4264 2316 0.4 Criswe11 SW 15 49 13 734 4265 2438 3.1 Potter SE 17 49 12 741 4265 2012 6.7 Moore NW 25 49 13 738 4263 2377 2.0 Criswe 11 NW 19 49 12 739 4264 2073 2.0 Criswe11 NW 17 49 12 741 4265 2073 2.8 Moore NW 32 50 12 740 4270 2012 5.0 Potter NW 6 49 12 739 4269 2164 1.5 Potter NW 16 49 12 742 4266 2073 4.5 Moore NW 6 49 12 740 4269 2134 4.4 Potter SE 17 49 12 741 4265 2073 4.8 Moore SW 33 50 12 742 4270 2012 5.1 Criswe11 NE 8 49 12 741 4267 2103 2.7 Criswell SE 12 49 13 738 4267 2134 3.3 Potter NE 6 49 12 739 4269 2103 2.6 Potter NE 1 49 13 738 4269 2164 1.0 Potter NW 8 49 12 740 4268 2134 2.4 Criswell NW 17 49 12 740 4266 2073 1.7 Criswell Appendix Table C. 9- 6-84 9-28-84 10-13-84 10-26-84 11- 2-84 11-10-84 11-16-84 11-23-84 11-30-84 12- 7-84 12-16-84 12-21-84 12-29-84 1- 4-85 1-11-85 1-18-85 1-25-85 2- 1-85 2- 8-85 2-15-85 2-22-85 3- 1-85 3- 8-85 3-14-85 3-22-85 3-31-85 4- 5-85 4-11-85 4-18-85 4-19-85 4-27-85 5- 3-85 5-10-85 5-16-85 5-25-85 6- 1-85 6-10-85 6-14-85 6-21-85 6-28-85 SE NW NW NW NE SE SE SE NW SE NE 18 36 25 24 18 30 31 31 9 8 2 26 35 12 3 49 49 49 49 49 49 49 49 48 48 48 49 49 48 48 12 13 13 13 12 12 12 12 12 12 13 13 13 13 13 740 737 737 738 742 739 739 740 744 743 738 738 736 740 737 4265 4260 4262 4264 4265 4263 4261 4261 4257 4256 4258 4262 4261 4256 4258 2256 2438 2408 2377 2134 2286 2408 2438 2377 2499 2560 2408 2560 2560 2560 2.0 5.1 1.5 1.8 l.5 3.5 1.8 0.6 6.7 1.8 5.0 4.1 l.1 6.5 3.7 Criswe11 Criswell Criswe11 Criswell Moore Criswell Moore Traver Bull Bull Traver Criswe 11 Criswell Terrible Moore Rating Signal G Location G F F G G Ground telemetry G G F F G F P F F F Ground telemetry Ground telemetry P p G F F F F F G F p F F F F F P F P p F p F p F F P F F F G F G F G G G F G F G F F F G G G F G G G G G F G F Ground telemetry G F F F G P G F P F G F F F G F G F G F G F G F �257 Date Aerial telemetry locations of adult female puma #7. Legal descr. U.T.M. Approx. Distance (km) elev. between S T R X Y (m) locations 1/4 7- 6-84 SE 7-13-84 NW 7-20-84 Appendix Table D. 49 14 728 4268 2316 4.3 34 50 14 724 4270 2377 4.6 NE 2 49 14 727 4268 2377 3.4 7-27-84 SE 32 50 14 722 4270 2408 6.0 8- 9-84 SW 32 50 14 721 4269 2316 0.9 8-18-84 SW 36 50 15 718 4269 2256 3.2 8-24-84 NE 9 49 14 723 4267 2377 6.7 8-31-84 NE 16 49 13 733 4266 2377 10.0 9-14-84 9-28-84 SE NE 7 6 49 49 13 14 729 720 4266 4268 2408 2347 3.4 10.1 10- 5-84 NE 33 50 14 723 4270 2256 5.1 10-13-84 10-19-84 10-26-84 SW SE SE 24 12 7 49 50 50 14 13 13 728 738 730 4263 4276 4276 2530 1951 2256 8.0 14.7 8.0 11- 2-84 SE 13 51 13 737 4284 1707 11.1 11-10-84 11-16-84 NE NW 25 35 50 50 13 14 738 726 4272 4270 1890 2042 12.2 4.8 11-23-84 NW 4 50 13 732 4279 2042 4.2 11-30-84 12- 7-84 12-16-84 12-21-84 SW NE SE NE 31 31 31 4 51 51 51 50 13 13 12 13 729 730 739 733 4279 4280 4279 4278 2012 1951 1829 2042 3.2 1.4 9.4 6.2 12-29-84 NW 31 50 13 729 4270 2073 9.3 1- 4-85 NW 21 50 13 732 4274 2042 4.5 1-11-85 NE 18 50 13 730 4275 2286 2.5 1-18-85 SW 9 50 13 732 4276 2164 2.4 1-25-85 SW 20 50 13 731 4272 2164 3.9 2- 1-85 2- 8-85 2-15-85 2-22-85 SW SW NW SW 11 25 15 9 50 50 50 50 13 13 14 13 735 737 724 732 4276 4271 4275 4276 2073 2073 2316 2073 5.5 5.1 13.0 7.5 3- 1-85 3- 8-85 NE SE 22 31 51 51 13 13 734 729 4283 4279 1890 2103 7.0 6.1 Ra ting Major drainage Dry Fk Escalante Middle Fk Escalante Dry Fk Escalante Middle Fk Escalante Middle Fk Escalante Middle Fk Escalante East Fk Escalante Little Moni tor Cottonwood Middle Fk Escalante Middle Fk Escalante Cottonwood Cottonwood Dry Fk Escalante Dry Fk Escalante Monitor East Fk Escalante Dry Fk Escalante Escalante Escalante Co t tonwocd Dry Fk Escalante Dry Fk Esca 1 ante Dry Fk Escalante Dry Fk Escalante Dry Fk Esca 1 ante Dry Fk Escalante Grade Gulch Monitor Escalante Dry Fk Escalante Escalante Escalante Signal Location G F F F G F G F G F G G F F G F G G G F F F G G G F F F G F P F G F G G F F p F F F F F G F p F G F G G G F G F G F G F F F G F G F �258 Appendix Table D. 3-14-85 NE (continued) 17 50 13 731 4275 2164 4.6 3-22-85 3-31-85 SW SW 11 9 50 50 13 13 735 732 4276 4276 2042 2073 4.4 3.4 4- 5-85 NW 15 50 13 733 4275 2134 1.4 4-11-85 4-19-85 SW NW 12 31 50 50 14 13 727 729 4276 4271 2316 2196 2.4 6.0 4-27-85 SE 19 50 13 730 4273 2073 2.1 5- 3-85 SW 20 50 13 730 4273 2103 0.7 5-10-85 NE 19 50 13 730 4273 2196 0.3 5-16-85 5-25-85 SE 32 32 50 50 13 14 732 722 4269 4270 2256 2134 4.3 10.0 6- 1-85 6-10-85 NW NE 24 10 49 49 14 14 729 725 4264 4267 2560 2377 9.3 5.0 6-14-85 NW 12 49 14 727 4267 2316 2.7 6-21-85 6-28-85 SE SW 7 8 49 49 13 13 730 731 4266 4266 2377 2347 3.0 0.9 NE Dry Fk Escalante Grade Gulch Dry Fk Esca 1 ante Dry Fk Escalante Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Esca 1 ante Dry Fk Escalante Cottonwood Middle Fk Escalante Cottonwood East Fk Escalante Dry Fk Escalante Cottonwood Cottonwocd Appendix Table E. Aerial telemetry locations of adult female puma #12. Legal descr. U.T.M. Approx. Distance (km) elev. between Major Date 1/4 S T R X Y (m) locations drainage 29 15 15 33 50 50 50 50 14 14 14 14 721 724 725 723 4272 4274 4275 4269 2438 2195 2103 2469 2.8 3.9 0.7 5.6 24 32 50 50 15 14 717 -721 4273 4270 2347 2195 7.3 5.3 NE 30 36 50 50 14 15 720 718 4271 4270 2499 2499 2.1 1.9 9- 7-84 NE 29 50 14 722 4272 2377 3.7 9-14-84 NW 21 50 14 723 4273 2408 1.8 9-21-84 SE 24 50 13 728 4273 2377 5.4 9-28-84 NE 31 50 14 720 4270 2316 8.3 10- 5-84 SE 20 50 14 722 4273 2408 3.3 10-13-84 10-26-84 11- 2-84 11-10-84 SE 20 16 24 23 50 50 50 50 14 14 15 14 720 723 718 727 4272 4275 4273 4274 2408 2195 2438 2.1 4.1 5.7 Kelso Esca 1 ante Escalante Middle Fk Escalante Kelso Middle Fk Esca 1 ante Kelso Middle Fk Escalante Middle Fk Escalante Middle Fk Escalante Dry Fk Escalante Middle Fk Escalante Middle Fk Escalante Kelso Kelso Kelso 2256 9.0 Escalante 7- 6-84 7-11-:84 7-20-84 7-27-84 NW SW 8- 9-84 8-17-84 NW 8-24-84 8-31-84 SE NE SE NE NE SW NE F G F F G G G F G F G F G F G F G F G F F F G F G F G F G F G F Rating Signal Location G F F F G F G F G F G G F F G F G F G G F F G F G F G G G F G F p F �259 Appendix Table E. (continued) 23 50 14 11-16-84 NE 24 50 15 11-23-84 NW 26 SO 14 12- 7-84 NE 727 718 727 4274 4273 4272 2225 2164 22S6 0.3 8.9 9.0 12-16-84 NE 26 50 14 727 4270 2195 1.2 12-21-84 sw 3 50 13 733 4278 2042 9.S 12-29-84 SW 8 50 13 730 4276 2103 3.3 1- 4-85 1-11-85 1-18-85 1-25-85 NW NW NW NE 27 30 7 13 13 13 14 733 729 728 72S 4282 4281 4277 4278 2012 1859 2256 S.9 4.2 4.6 3 51 Sl 50 50 2- 1-85 SE 15 50 14 725 4274 2164 4.7 2- 8-85 2-15-85 2-22-85 SE SW SW 26 lS 4 Sl SO SO 14 13 13 726 733 732 4281 4274 4278 2164 222S 2073 6.9 9.8 4.1 3- 1-85 3- 8-85 3-14-85 3-22-85 SE SE SW SW 36 10 17 3S Sl SO 50 50 14 14 14 14 728 725 721 726 4279 4276 4274 4269 18S9 2134 2103 22S6 4.S 4.4 3.9 7.2 3-31-85 SW 26 Sl 13 735 4281 2073 15.5 4- 5-85 SW 13 SO 14 727 427S 2286 10.0 4-11-85 4-19-85 NE NE 28 36 SO SO 15 15 714 718 4271 4270 2438 2499 S.7 4.7 4-27-85 NE 26 SO 14 726 4272 22S6 8.4 4-30-85 NW 24 SO 14 728 4274 2286 2.0 5- 3-85 5-10-85 SE SW 14 11 SO 49 14 lS 727 716 4274 4266 22S6 2499 0.6 13.9 5-16-85 SE 36 SO lS 719 4269 2347 4.S 5-25-85 NW 36 SO 15 718 4270 2S60 1.6 6- 1-85 SE 33 SO 14 723 4269 2469 6.0 6-10-85 SE 32 SO 14 722 4269 6-14-85 6-21-85 NW SW 28 6 SO 49 is 14 713 719 4271 4267 2439 2438 10.8 7.5 6-28-85 NE 14 50 14 727 427S 2073 10.9 Escalante Kelso East Fk Escalante East Fk Esca1ante Dry Fk Escalante Dry Fk Esca1ante Tatum Draw Escalante Tatum Draw North Fk Escalante East Fk Esca 1ante Esca 1ante Grade Gulch Dry Fk Escalante Escalante Kelso Kelso East Fk Escalante Dry Fk Escalante Escalante G F G F G F G F F F G F G F F F Kelso Middle Fk Escalante East Fk Esca 1ante Escalante G F G F G G Esca 1ante Middle Fk Escalante Middle Fk Escalante Middle Fk Escalante Middle Fk Escalante Middle Fk Escalante Kelso Middle Fk Escalante Esca 1ante Appendix Table F. Aerial telemetry locations of subadu1t male #13. Legal descr. U.T.M. Approx. Distance (km) Major e1ev. between Date 1/4 S T R X Y (m) locations drainage 9- 5-84 NE 7 48 14 722 4256 North Fk Tabeguache F p p p F F G F G F G F G F F F F F G F G F Sighted while ground tracking G F G F G F F F G G G p G F G F G F Rating Signal Location Ground telemetry G P �260 Appendix Table G. Aerial telemetry locations of adult female puma #15. Legal descr. U.T.M. Approx. Distance (km) Major between elev. y (m) locations drainage R 1/4 S T x Date 7- 6-84 NW 21 48 11 754 4254 2256 1.2 7-13-84 SW 19 48 11 750 4253 2377 3.3 7-20-84 NW 28 48 11 754 4252 2377 3.6 7-26-84 NE 28 48 11 754 4253 2286 0.4 7-27-84 NW 28 48 11 754 4252 2377 0.4 8- 9-84 ' SE 20 48 11 753 4253 2256 1.2 8-17-84 NW 17 48 11 752 4256 2042 2.8 8-24-84 SE 7 48 11 751 4256 2225 0.8 8-30-84 NW 17 48 11 752 4255 2195 1.1 8-31-84 SW 20 48 11 752 4253 2316 1.8 9- 7-84 SW 21 48 11 754 4253 2286 3.0 9-14-84 NW 27 48 11 755 4252 2408 1.5 9-21-84 SW 21 48 11 754 4253 2286 1.5 9-28-84 NW 27 48 11 755 4252 2316 1.7 10- 5-84 SW 8 48 11 753 4256 2195 6.3 10-13-84 SW 18 48 11 750 4255 2347 2.5 10-19-84 NE 9 48 11 754 4257 1951 4.9 10-26-84 NW 9 48 11 754 4257 2042 0.7 11- 2-84 11-10-84 SE NE 4 17 48 48 11 11 754 753 4260 4256 2073 2134 1.6 3.2 11-16-84 SW 20 48 11 750 4253 2438 6.5 11-23-84 NE 5 47 11 753 4249 2560 5.0 11-30-84 SW 34 48 11 755 4250 2408 2.5 12- 7-84 NE 8 48 11 753 4257 2012 7.8 12-16-84 NE 20 48 11 753 4254 2256 2.8 NE 12 48 48 48 48 48 11 11 11 11 11 759 759 759 758 758 4257 4258 4258 4252 4254 2073 2073 2042 2316 2256 7.2 0.8 0.6 6.2 1.8 12-21-84 12-29-84 1- 4-85 Hl-85 1-18-85 SE NE SW SW 12 26 24 Rating Signal Location East Fk Dry Ck West Fk Dry Ck East Fk Dry Ck East Fk Dry Ck East Fk Dry Ck East Fk Dry Ck West Fk Dry Ck West Fk Dry Ck West Fk Dry Ck West Fk Dry Ck East Fk Dry Ck East Fk Dry Ck East Fk Dry Ck G F F F G G East Fk Dry Ck West Fk Dry Ck West Fk Dry Ck East Fk Dry Ck West Fk Dry Ck Dry Ck West Fk Dry Ck East Fk Dry Ck East Fk Dry Ck East Fk Dry Ck West Fk Dry Ck East Fk Dry Ck Coa 1 Ck Coal Ck Coal Ck Coa 1 Ck Coak Ck Ground telemetry G G G F F F F F G F G F G G G G G F G G F F F F G F F G F F F G G G G G G G F G F G F G F G F G F G F �i 261 Aeeendix Table G. SE 1-25-85 (conti nued) 11 7 48 751 4256 2347 7.9 2- 1-85 SE 16 48 11 755 4255 2195 4.2 2- 8-85 NW 22 48 11 755 4254 2196 1.2 2-15-85 2-22-85 SW NW 10 16 48 48 11 756 753 4256 4256 2225 2.0 11 3- 1-85 3- 8-85 3-14-85 3-22-85 3-31-85 NW SW NW NW NW 12 18 12 12 10 48 48 48 58 48 11 10 11 11 11 758 760 758 758 755 4257 4255 4257 4258 4257 2103 2134 2103 2042 2134 2.6 3.7 3.2 0.6 3.3 4- 5-85 4-11-85 SE NE 11 16 48 48 11 11 758 755 4257 4255 2103 2164 2.7 3.3 4-19-85 4-27-85 NE NE 22 21 48 48 11 11 756 755 4254 4254 2256 2134 1.7 1.6 5- 3-85 NE 19 48 11 752 4254 2316 2.7 5-10-85 NE 28 48 11 754 4252 2316 3.0 5-25-85 6- 1-85 6-10-85 SE SE SE 35 35 34 48 48 48 12 12 11 748 748 756 4250 4250 4250 2621 2591 2377 6.4 0.3 7.9 6-14-85 SE 28 48 11 754 4252 2438 2.2 6-21-85 SW 33 48 11 754 4251 2499 1.6 6-28-85 NW 25 48 11 758 4253 2286 5.2 West Fk Dry Ck East Fk Dry Ck East Fk Dry Ck Dry Ck East Fk Dry Ck Coal Ck Coal Ck Coal Ck Coal Ck East Fk Dry Ck Coal Ck East Fk Dry Ck Coa 1 Ck East Fk Dry Ck West Fk Dry Ck East Fk Dry Ck Gray's Ck Gray's Ck Cottonwood East Fk Dry Ck East Fk Dry Ck Coal Ck G F G F P F G F P P P P F F F F F G G G F G F F F G F G F F F G F F F G F G F G F F F F �262 Date Aerial telemetry locations of male adult puma #18. Legal descr. U.T.M. Approx. Distance (km) elev. between y (m) locations S T R x 1/4 7- 6-84 7-13-84 7-20-84 NW SE NW 25 12 4 47 47 46 10 10 9 243 245 249 4243 4246 4240 2316 2134 2316 2.1 0.9 7.9 7-27-84 8- 9-84 8-18-84 8-24-84 8-31-84 9-14-84 9-21-84 SE NE SW NW SE NW SW 26 35 33 23 23 13 9 47 47 47 48 47 47 46 10 10 243 243 249 243 243 244 249 4242 4241 4240 4254 4243 4245 4237 2377 2438 2286 1920 2377 2256 2377 8.1 1.2 6.2 15.0 9.8 2.3 10.7 9-28-84 10- 5-84 10-13-84 NE NW NW 27 27 5 47 47 45 251 241 247 4242 4243 4239 2134 2438 2377 6.2 10.1 7.4 10-19-84 10-26-84 11- 2-84 11-10-84 11-16-84 11-23-84 11-30-84 12- 7-84 SE NW NW SE SW SE NW NE 8 47 47 47 47 47 47 49 46 248 245 247 246 245 251 739 248 4246 4248 4246 4243 4243 4245 4262 4239 2073 2134 ·2134 7.4 2.7 2.7 2.8 1.0 6.0 2347 6.8 12-16-84 12-21-84 12-29-84 1- 4-85 1-11-85 1-18-85 1-25-85 2- 1-85 2- 8-85 2-15-85 2-22-85 3- 1-85 3- 8-85 3-14-85 3-22-85 3-31-85 4- 5-85 4-11-85 4-19-85 NE SW SW NE SW SE NE NE SW NW NW SW NW SW SW NW 251 246 246 245 251 251 244 245 249 245 249 249 244 249 249 251 245 244 249 4242 4245 4244 4246 4241 4245 4243 4244 4240 4243 4239 4240 4246 4240 4240 4241 4245 4245 4240 2134 2195 2256 2195 2134 2098 2347 2286 2256 2286 2316 2256 2196 2256 2256 2134 2286 2196 2316 4.8 5.6 1.0 2.6 7.5 3.4 7.1 1.1 5.7 4.4 4.8 0.6 7.5 7.6 0.9 1.7 5.7 0.7 7.0 246 244 245 247 250 251 4243 4244 4246 4246 4233 4243 247 251 741 739 4245 4232 4264 4262 2256 2286 2134 2134 2469 2103 2103 2499 2134 4.8 1.7 2.0 1.5 13.4 10.2 4.8 13.8 47.0 Appendix Table H. 7 17 19 19 15 25 5 10 10 10 9 9 10 9 9 9 9 9 9 9 13 9 NW 33 13 33 33 34 13 13 4 47 47 47 47 47 47 47 47 47 47 46 47 47 47 47 47 47 47 46 10 10 9 4-27-85 5- 3-85 5-10-85 5-16-85 5-25-85 SW NW NW NW NW 19 24 18 17 27 47 47 47 47 46 9 10 9 9 9 5-31-85 6-10-85 6-14-85 6-21-85 6-28-85 SE NE NW NE NW 27 17 26 20 25 47 47 46 49 49 9 SE SE 27 18 9 19 13 27 15 25 24 33 30 4 9 9 9 10 9 9 10 10 9 9 9 9 10 9 9 9 9 9 12 13 2286 2286 2103 Rating Major drainage Signal Location Happy Happy West Fk Horsefly Happy Dolores Horsefly Spring Happy Happy Eas t Fk Horsefly Horsefly Happy West Fk Horsefly Dolores Happy G p F G F P F G F G F G F Dolores Dolores Dolores Horsefly Cri swe 11 West Fk Horsefly Horsefly Dolores Do 1ores Happy Horsefly Horsefly Happy Happy Horsefly Dolores Horsefly Horsefly Happy Horsefly Horsefly Horsefly Happy Happy West Fk Horsefly Do 1ores Happy Happy Dolores McKenzie Ck Horsefly G F F F G F G F P P F F G F F F Dolores Fisher Moore Criswell G G G F G G G F G F G F F F F F G F G F G G F F F F G G G F G G G G G G G F F F F F G F G G G G G G G G G G F G G F G G G F F F G F G F G p �263 Appendix Table I. Aerial telemetry locations of subadu1t female puma #20. Lega 1 descr. U.T.M. Approx. Distance (km) elev. between X y Date S T (m) 1/4 R locations Rating Major drainage Signal Location 11-26-84 SE 27 48 11 756 4252 2286 capture 11-30-84 12- 7-84 SE NW 35 49 48 11 11 756 754 4261 4257 1920 2103 8.9 4.3 12-16-84 12-21-84 12-29-84 1- 4-85 1-11-85 SE SE SW SE NW 25 35 31 11 10 49 49 49 48 48 11 11 11 756 755 758 758 755 4264 4261 4261 4259 4257 1951 2012 1981 2073 2103 6.9 4.0 2.5 2.5 3.1 1-18-85 NE 17 48 11 753 4256 2164 2.3 1-25-85 NW 17 48 11 752 4256 2164 1.4 2- 1-85 NE 16 48 11 755 4255 2042 3.0 2-15-85 NW 10 48 11 755 4257 2134 1.9 2-22-85 SW 16 48 11 754 4255 2196 3.1 3- 1-85 3- 8-85 3-11-85 NW 14 49 11 755 4266 1859 11.5 NW 14 49 11 755 4266 1859 0 Two signals received: (1) hill W of Smith Cabin Escalante Sawmill Mesa 3-18-85 Escalante Ground telemetry P p 3-19-85 One signal received hill W of Smith Cabin Escalante C. at 0830 One signal received lower Escalante C. at 0809 Escalante Ground telemetry 3-20-85 One signal received lower Escalante C. at 0809 Escalante Ground telemetry 3-21-85 One signal received lower Escalante C. at 0843 Escalante 3-22-85 3-31-85 4- 5-85 4-11-85 4-19-85 5- 3-85 5-10-85 5-16-85 NW 14 49 11 NW 27 49 11 SE 26 49 11 NW 23 49 11 No specific location, SE 23 49 11 NW 33 49 12 SW 23 50 12 5-31-85 NE 6-10-85 6-14-85 6-21-85 6-28-85 SW SW SE NW 9 2 11 East Fk Dry Ck Dry Ck West Fk Dry Ck Dry Ck Dry Ck Dry Ck Coal Ck East Fk Dry Ck East Fk Dry Ck vJes t Fk Dry Ck East Fk Dry Ck East Fk Dry Ck East Fk Dry Ck Dry Ck Dry Ck C; (2) lower G F G F G G G F G F G F G F G F G F G F P F P F Ground Telemetry p P P p 48 6 34 49 49 29 49 16 49 11 11 12 12 12 p p Ground telemetry p 28 F G 1890 0.4 755 4266 Dry Ck F 753 4263 2042 3.1 Dry Ck G 1859 2.8 756 4263 Dry Ck P 1829 2.5 755 4265 Dry Ck G very faint signal while above E rim of Roubideau near Traver Ck. 1.3 1829 Dry Ck 755 4264 G 13.4 Wright 742 4261 2225 G 11.6 745 4273 1707 CriswellG Roubideau 22.8 755 4252 East Fk 2377 G Dry Ck 7.9 Piney Ck 750 4259 2286 G 6.6 Roubideau 744 4260 1981 G 2.9 741 4262 Traver 2196 G 742 4266 4.3 Moore 2012 G p F F F F F F F F F F F F �264 Appendix Table J. Date Aerial telemetry locations of subadult female puma #21. Legal descr. U.T.M. Approx. Distance (km) Major elev. between y (m) locations drainage S R x 1/4 T 12- 7-84 12-16-84 NW NW 12 15 50 50 14 13 727 733 4277 4275 2134 2073 Capture 6.2 12-21-84 SE 27 51 13 735 4281 1920 5.6 12-29-84 SW 30 50 13 729 4271 2073 11.2 1- 4-85 NW 17 50 13 731 4275 2225 4.3 1-11-85 SE 50 13 729 4276 2256 1.8 1-18-85 SW 17 50 13 730 4274 2195 2.2 1-25-85 SW 19 50 13 730 4273 2195 1.3 2- 1-85 SW 31 50 13 729 4269 2225 4.2 2- 8-85 SW 9 50 13 732 4276 2196 7.8 2-15-85 2-22-85 SW NE 36 17 50 50 13 13 737 732 4270 4275 2196 2103 8.5 7.4 3- 1-85 SW 16 50 13 732 4274 2134 1.1 3- 8-85 SE 17 50 13 731 4275 2103 1.0 3-14-85 NW 16 50 13 732 4275 2103 0.7 3-22-85 SE 8 50 13 731 4276 2164 1.1 3-31-85 SW 9 50 13 732 4276 2073 0.7 4- 5-85 SE 36 50 14 728 4269 2286 8.5 4011-85 SW 10 50 13 733 4276 2073 8.3 4-19-85 NE 31 50 13 730 4270 2377 6.5 4-27-85 SE 19 50 13 730 4273 2286 2.5 5- 3-85 5-10-85 5-16-85 SE NW SE 14 24 25 50 50 50 14 14 14 727 727 728 4274 4274 4271 2225 2286 2316 3.5 0.6 2.2 5-25-85 NW 31 50 14 720 4270 2438 8.3 6- 1-85 NW 49 15 718 4268 2347 2.7 6-10-85 NW 10 49 14 724 4267 2103 6.1 6-14-85 SE 2 49 15 717 4267 2530 6.9 6-21-85 NW 9 49 15 713 4266 2745 4.1 6-28-85 SW 21 49 13 732 4263 2469 19.5 Escalante Dry Fk Escalante Dry Fk Esca 1ante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Monitor Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Esca 1 ante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Escalante Escalante Dry Fk Escalante Middle Fk Escalante Middle Fk Escalante East Fk Escalante Middle Fk Escalante Middle Fk Escalante Moni tor Rating Signal Location G F F F G F G F G F G F G F G F G F G G G F F F F F F F G F G F G F G F G F G F G G G G G F F F G F G F G F G F G F �265 Appendix Table K. Aerial telemetry locations of subadult male puma #22. Legal descr. U.T.M. Approx. Distance (km) elev. between Major Date 1/4 S T R X Y (m) locations drainage 1- 5-85 NE 30 46 8 256 4232 2347 Capture 9 4238 4243 4231 2408 2134 2256 9.8 6.1 14.1 1-11-85 NE 8 1-18-85 2- 1-85 SE SE 22 28 46 47 46 9 8 248 251 259 2- 8-85 NE 29 46 8 258 4232 2196 1.5 2-15-85 NE 20 46 8 258 4234 2196 1.6 2-22-85 NE 5 45 8 257 4229 2347 5.0 3- 1-85 NE 20 46 8 258 4234 2103 5.4 3- 8-85 NE 20 46 8 258 4234 2196 0.4 3-14-85 SW 8 46 8 257 4237 2286 2.6 3-22-85 3-31-85 4- 5-85 4-11-85 4-19-85 4-27-85 5- 3-85 NW SW NE NW SE NE SW 18 10 13 20 33 32 8 46 46 46 46 46 46 46 7 8 8 7 7 7 8 265 260 264 267 269 267 258 4235 4237 4235 4234 4230 4231 4237 2591 2256 2438 2621 2560 2377 2012 8.7 5.7 4.2 2.1 4.7 1.6 11.4 5-10-85 5-16-85 NW SE 15 34 45 46 8 270 261 4226 4230 2776 2225 20.0 9.9 5-25-85 NE 33 46 8 259 4231 2225 1.7 5-31-85 6-10-85 6-14-85 6-21-85 6-28-85 NE NW NW SE SE 32 24 24 28 19 46 46 46 46 46 7 267 264 266 269 265 4231 4234 4233 4231 4233 2316 2286 2469 2408 2347 8.0 4.6 1.9 3.6 3.3 7 8 7 7 Uncompahgre River Horsefly Horsefly Uncompahgre River Uncompahgre River Uncompahgre River Uncompahgre River Uncompahgre River Uncompahgre River Uncompahgre River Deer Ck Cow Ck Deer Ck Martin Ck Na te Ck Lou Ck Uncompahgre River Flume Ck Uncompahgre River Uncompahgre River Lou Ck Deer Ck Martin Ck Lou Ck Martin Ck Rating Signal Location G F G F G F F F G F F F G F F F F G G F F G G G F G F G F G F G F p F F F F G F G F G F G G �266 Appendix Table L. Aerial telemetry locations of subadult male puma #23. Local descr. U.T.M. Approx. Distance (km) elev. between Major Date 1/4 S T R X Y (m) locations drainage 1- 9-85 1-14-85 NE 32 145 99W 718 4297 2195 Signal Location capture Big Dominquez About 1 km E of Big Dominquez Canyon in bottom or lower 5. slope of Big Dominquez 1-18-85 NW 14 15S 100W 714 4290 2195 7.4 1-25-85 NE 17 15S 100W 709 4291 2286 4.5 2- 1-85 2- 8-85 2-15-85 2-22-85 3- 1-85 NW 5W NE NE NE 12 23 22 30 29 145 135 135 135 12S 100W 100W 100W 100W 101W 714 713 712 707 699 4303 4309 4309 4308 4317 1951 1920 2012 2377 2042 12.6 6.1 1.4 5.0 11.5 3- 8-85 3-14-85 NW 5E 29 20 115 101W IN 3W 697 687 4327 4243 1829 1647 9.7 12.9 3-22-85 4- 5-85 4-19-85 5E SW SE 2 2 32 11S 103W 13S 101W 12S 101W 682 703 699 4332 4313 4314 1555 2073 2256 5.1 28.3 4-27-85 5- 3-85 5-10-85 5-16-85 6- 1-85 6-10-85 6-14-85 6-28-85 NW NE SE NE SE SW NE NE 28 9 32 35 10 14 23 15 13S 13S 13S 13S 13S 13S 13S 13S 709 691 709 704 693 693 694 702 4308 4312 4306 4305 4311 4309 4309 4311 2347 2164 2256 2621 2225 2316 2377 2347 12.5 18.0 19.0 4.0 10.0 1.6 1.5 9.2 100W 102W 100W 101W 102W 102W 102W 101W Ra t i ng 3.8 Ground telemetry Big Dominquez Big Dominquez Unaweep NE Ck Bang's Can. Bang's Can. No Thoroughfare Can. Monument Can. Rattlesnake Can. Mee Can. Ladder Can. No Thoroughfare Can. Bang's Can. Clark Wash NE Ck NE Ck Clark Wash Clark Wash Clark Wash Ladder Can. Date Aerial telemetry locations of subadult male puma #26. Legal descr. U.T.M. Approx. Distance (km) elev. between Major 1/4 5 T R X Y (m) locations drainage 3-20-85 3-22-85 NE 5W 14 35 50N 50N 14W 14 727 729 4275 4269 2073 2256 capture 6.0 3-31-85 SE 27 51 13 734 4281 2196 14.5 4- 5-85 4-11-85 4-19-85 NE NW SE 12 28 25 50 50 50 14 15 15 728 713 718 4277 4271 4271 2256 2438 2530 7.8 15.6 5.4 4-27-85 5E 23 50 14 726 4272 2134 8.2 5- 3-85 5-16-85 5-25-85 NW 2256 13 50 14 727 4275 Vicinity East Fk Escalante and Middle Fk Escalante 5W 36 50 15 718 4269 2377 6- 1-85 SE 33 50 14 723 4269 2438 5.5 6-10-85 NW 16 49 15 713 4265 2682 10.7 6-14-85 NW 10 48 15 716 4256 2621 10.0 6-28-85 Very faint mometary signal over lower Tabaguache Ck, GMU 61. G F G F G F G F G F G F P{odd) F G F F F G F F F P F G G F F F F F F F F G G G F G F G F Appendix Table M. 2.5 Escalante East Fk Escalante Dry Fk Escalante Esea1an te Kelso Middle Fk Escalante East Fk Escalante Escalante Ratinq Signal Location G F G G G F G F G F G F G p 10.6 Middle Fk Esca1ante Middle Fk Escalante Middle Fk Escalante East Branch Shavano Ck F none F F G G G F G F �267 Date Aerial telemetry locations of subadult female puma #27. Approx. Distance (km) Legal descr. U.T.M. elev. between y (m) locations S R x 1/4 T 4-24-85 5W 29 51 14 721 4281 2377 capture 4-27-85 SW 20 15S 98W 728 4289 1981 10.0 5- 3-85 5-10-85 5-16-85 NE 5W SW 22 28 9 51N 51N 50N 14W l4W 15W 724 723 713 4283 4281 4275 2225 2347 2499 7.0 2.9 11.3 5-25-85 NE 8 50N 14W 721 4277 2286 8.6 6- 1-85 6-10-85 NW 5W 35 17 51N 50N 15W 15W 716 711 4279 4274 2560 2438 6.5 7.2 6-14-85 SW 9 50N 15W 713 4275 2530 2.4 6-21-85 6-28-85 5E 5E 9 14 50N 50N 14W 15W 723 717 4275 4274 2134 2408 9.8 6.1 Appendix Table N. Appendix Table O. 7- 6-84 5W 8-24-84 SW 5W 9- 3-84 Aerial telemetry 29 47 8 35 49 13 35 50 14 locations of subadu1t male puma #17a. 258 4241 2195 15.5 735 4260 726 4270 Major drainage North Fk Escalante Gunnison Gulch Palmer Gulch Palmer Gulch North Fk Escalante North Fk Esca 1ante Corral Gulch North Fk Escalante North Fk Escalante Kelso North Fk Escalante Uncompahgre Cri swe 11 East Fk Escalante Rating Signal Location G F G G G F F F F F G G F G G G F G F G G P P Ground telemetry p P aShot by sport hunter 1-11-85 at NW 1/4, 57, T6, R90, 87.2 km from capture site on Alkali Ck., GMU 42. Appendix Table P. Aerial telemetry locations of subadu1t female puma #19. Loca 1 descr. U. T.M. Approx. Distance (kill) elev. between Major x y Date 1/4 5 T R (m) locations drainage 7-18-84 7-20-84 7-27-84 8- 2-84 5W 5E 5W 5E 18 8- 9-84 8-24-84 8-25-84 33 28 48 48 49 49 13 13 13 13 732 734 733 733 4254 4258 4260 4261 2806 2682 2560 2469 4.2 2.5 1.6 5W NE NE 12 26 26 47 49 49 13 11 11 739 756 756 4246 4263 4263 2928 16.5 8-28-84 NE 26 49 11 756 4263 1890 25.8 Dry Ck 9- 3-84 9- 4-84 5E 5E 19 19 155 155 96W 96W 746 746 4290 4290 1494 1494 28.0 Roubideau Roubideau 11-10-84 5E 6 48 14 722 4257 2682 38.5 11-13-84 NW 14 48 11 757 4255 5 o Criswell Criswell Potter Potter Tabaguache Dry Ck Dry Ck North Fk Tabaguache Coal Rating 5igna1 Location {release site)a G G P F Ground telemetry G F G F P P Ground telemetry G P Ground telemetry G G G F Ground telemetry P F P F Ground telemetry G P aTransplanted from NW 1/4, 52, T47, 9W, 1890 m elev., lower Horsefly Ck. after it was trapped at kill site of domestic sheep on 7-18-84. 5hot by ADC, U5FW5 trapper on 1-15-85, 5W 1/4, 514, T14, UTM 728-422S-for 1 verified sheep kill. � https://cpw.cvlcollections.org/files/original/9fd9e5b9703a92c26130939b91bcb640.pdf 5b5218f61c92e5e081901391e2f8aa19 PDF Text Text 1 Colorado Division of Wildlife Wildlife Research Report January 193G JOB PROGRESS REPORT State of Colorado --~~~~-------- Project N-l-R-3 Work Plan (II) ----~~~~------ Job (SE-3-5) Nesting Performance of Peregrine Falcons in Colorado (I) --------~--~~-------- Pe riod Covered: Author: Personnel: 1 January - 31 December 1984 G. R. Craig D. Berger and G. Craig, Colorado Division of Wildlife; J. Enderson, The Colorado College. ABSTRACT While the number of occupied peregrine falcon (Falco peregrinus anatum) breeding territories remained at 13 in 1984, the number of egg producing pairs increased to 12. Eggshell condition continued to be poor with 11.4% thinning. Photographic identification established an annual adult survivorship of 0.81. This Job Progress Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the Author. ��3 NESTING PERFORMANCE OF PEREGRINE FALCONS IN COLORADO Gerald R. Craig P. N. OBJECTIVE The objectives of this study are to annually monitor breeding numbers and reproduction of Colorado peregri ne falcons to document further population declines as well as record the population's responses to recovery efforts. Additionally, health of the population will be monitored indirectly by analyzing pesticide residue levels in the falcons' eggs and principal prey. Information obtained from these investigations wi 11 be made avail able through annual reports to the Rocky Mountai n/Southwest Peregri ne Falcon Recovery Team as well as cooperating agencies to aid in evaluation of recovery efforts. SEGMENT OBJECTIVE 1. Annually monitor the number of breeding pairs of peregrines and reproduction in Colorado. 2. Annually monitor organochlorine pesticide levels in wild breeding peregrines. 3. Monitor recruitment of reintroduced breeding population of Colorado. 4. peregrines into the wild Compile data and submit reports to appropriate state and federal personnel and the Rocky Mountain/Southwest Peregrine Falcon Recovery Team for use in evaluating recovery efforts. (These objectives correspond to Jobs 1113., 1114.,212., 221., 3211., 321 2., and 333. of the approved American Peregrine Falcon Recovery Plan for the Rocky Mountain/Southwest population). METHODS AND MATERIALS Methods and materials used in this study have been described previously (Craig and Enderson 1981). RESULTS AND DISCUSSION Territory Occupancy Territory occupancy and breeding success is summarized in Table 1. A lthough the same number of breedi ng terri tories (13) were occupi ed in 1983 and 1984, site occupancy shifted. A lone male which occupied site 36 and 2 pairs comprised of subadult members (site 5 and 9) �4 disappeared 40,41, and territori es eral times the additional in 1984 and 3 additional pairs 42 in 1984. While addition of 3 increased the known sites to 42, in past years and had been found sites represent reoccupation of were present at sites previously undocumented all were surveyed sevto be vacant. Thus, vacant territories. Adult pairs were present at 11 territories (sites 8, 25, 27, 30, 31, 34, 35, 38, 39, 40 and 42) while subadu1t females were paired with adult males at 2 other localities (sites 3 and 41). The male found dying of injuries suffered from a powerline collision near site 38 in 1983 was replaced by an adult male in 1984 and the pair bred successfully. The released male which courted a subadult female at site 3 the previous year returned with another subadult female in 1984 and the pair produced eggs. Reproduction In 1984, 11 of the pai rs were adult compared to 9 adult pai rs the previous year and while only 7 of the pairs laid eggs the previous year, 12 of the pairs produced eggs in 1984. The pair at site 25 may be approaching senility since they courted very little and did not produce eggs. Their failure was compensated by the two pairs with subadult females that did produce eggs. One of the immature pairs (site 41) produced infertile eggs while the female at site 5 laid 2 viable eggs. Eight of the sites (3, 8, 27, 30, 31, 34, 35, and 39) were manipulated to increase productivity and the remaining pairs were left to reproduce naturally. The 12 egg-producing pairs laid at least 35 eggs (2.92 eggs per pair) and 26 eggs were removed for artificial incubation. The low average clutch size can be accounted for by the exi stence of 2 cl utches of 2 eggs produced by the subadult females. Had the fostered pairs been permitted to incubate their own eggs, it is probable that only 9 young would have been produced. The 5 unmanipulated pairs did not fair well since they managed to produce only 4 young (0.8 young per pair) and when the probable natural production of the 8 manipulated pairs is added, the estimated total wild production would have been 13 young (1.0 young per pair). Foster actions increased wild production to 32 young (2.46 young produced per pair) of which 29 successfully fledged (2.23 young fledged per pair). Eggshell Thickness Thi ckness measurements obtained from 33 wil d eggs in 1984 averaged 0.318 mm (with membrane) which was 11.4 percent thinner than preDDT era eggs and was nearly identical to the 1983 value (0.317 mm). One recently discovered site (42) exhibited a clutch thickness of only 0.298 mm (17% thinner) and the eggs of a relatively young female of no more than 3 years were in even worse condi t i on at 0.295 mm (17.8% thinner). As would be expected, the subadult female at site 3 laid a clutch of normal thickness (0.363 mm), but the other yearling female at site 41 already experienced 9.7 percent thinning (0.324 mm). Judging from the average clutch thicknesses in 1984, sites 27, 30, and 34 exhibit greatest potential for failure in 1985. The unexpectedly thin shells of sites 39 and 42 also make them candidates for future egg manipulation actions. �5 Organochlorine Residues in Egg Contents Pesticide residue analysis was performed by the U~S. Fish and Wildlife Service at the Patuxent Wildlife Research Center on contents of 8 nonviable eggs collected in 1983. Six sites (8, 25, 27, 31, 34 and 35) were represented by the samples and DOE values average (geometrically) 14.26 ppm (range 5.5 to 22.1 ppm). This is somewhat lower than the average obtaineci from 6 eggs collected the previous year, but due to the limited sample size, little can be deduced about trends. Photographic Documentation of Wild Adults The photographic study of adults occupying territories now spans 4 years and represents 12 terri tori es , The data inc 1ude 36 cases where adults have been positively identified and of these, 28 involve birds returning from previous years resulting in an annual survivorship rate of 0.78. An additional 6 cases involve birds that were not positively identified, but probably were individuals which returned from previous years. If these are included, a total of 42 cases result, of which 36 returned yielding an annual survivorship of 0.81. These survivorship estimates are conservative since birds may have returned to other unknown territories and did not really die. The photographic work vity. The male, and 30 over the span of 4 34 wer~ present over a also permits investigation of individual probably the female, has been present years. The pair at site 25 and female 3-year period. longeat site at site LITERATURECITED Craig, Prepared G. R. and J. H. Enderson. 1981. Nesting performance of peregrine falcons in Colorado. Job Prog. Rep., Colo. Div. Wildl., Wildl. Res. Rep., Jan., pp 13-23. by: Gef:ra1d ¥aiq ea'i Wildlife Researcher C �Table 1. Total sites on record New sites located Adult pairs Mixed pairs Lone adults Total occupied sites11 Total breeding pairsWild breeding pairs Wild young produced Wild young fledged Successful wild pairs Pairs fostered Total young fostered Fostered young fledged Successful fostered pairs Total young fledged by pairs Tota 1 hack sites Total young hacked Hacked young fledged Total young fledged ., 11 e- Colorado Peregrine Breeding Success and Recovery Efforts , pairs which produced eggs 1972 1973 1974 1975 1976 1977 - 22 0 8 0 3 11 0 0 0 0 0 22 0 11 0 1 12 2 2 2 2 1 -- 22 0 7 0 2 9 6 5 11 11 5 1 2 2 1 13 22 0 5 1 0 6 5 25 3 6 1 1 8 4 2 2 2 2 2 5 5 2 7 29 4 11 0 1 12 9 3 4 4 3 6 14 7 4 11 --- --- 0 -- --- 2 3 7 4 2 2 2 1 1 5 . 1978 1979 1980 1981 ---_ -- -31 2 7 2 2 11 6 1 3 1 1 5 20 15 4 16 1 4 0 2 13 5 7 11 4 20 32 1 6 2 3 13 4 0 0 0 0 4 15 12 4 12 2 12 9 21 34 2 7 3 3 13 5 1 3 3 1 4 16 13 4 16 4 17 11 27 34 0 7 1 3 11 6 0 0 0 0 6 21 15 6 15 4 19 14 29 1982 1983 -- -1984 36 2 9 0 2 11 6 0 0 0 0 6 26 20 6 20 5 23 13 33 38 2 9 3 1 13 8 1 0 0 0 7 27 21 7 21 6 25 23 44 41 3 11 2 0 13 12 4 4 4 2 8 28 25 8 29 7 32 27 56 �Colorado Division of Wildlife Wildlife Research Report January 1986 7 JOB PROGRESS REPORT State of Colorado ~~------------------ Project No. N-l-R-4 (SE-3-6) Work Plan No. 1 (II) --~--~---------- Job No. (I) Nesting Performance of Peregrine Falcons in Colorado Period Covered: 1 January 1985 - 30 June 1985 Author: G. R. Craig Personnel: D. Berger and G. Craig, Colorado Division of Wildlife; J. Enderson, The Colorado College. ABSTRACT Peregrine falcon (Falco peregrinus anatum) eyrie occupancy continued to improve in 1985 with 14 sites occupied by pairs of which 13 produced eggs. The 8 unmanipulated pairs experienced a fledging success of 1.6 young per pair. Eggshell thickness has not improved since 1984 and remained at 11.9% of pre-DDT era eggs. Females at 6 sites experienced sufficient eggshell thinning to require augmentation efforts next year. Photographic identification of adults over a 6 year span yielded an overall annual adult survivorship of 0.79 and an annual survivorship of 0.87 for males and 0.82 for females. This Job Progress Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the author. ��9 NESTING PERFORMANCE OF PEREGRINE FALCONS IN COLORADO Gerald R. Craig P. N. OBJECTIVE The objectives of this study are to annually monitor breeding numbers and reproduction of Colorado peregrine falcons to document further population declines as well as record the population·s responses to recovery efforts. Additionally, health of the population will be monitored indirectly by analyzing pestiCide residue levels in the falcons· eggs and principal prey. Information obtained from these investigations will be made available through annual reports to the Rocky Mountain/Southwest Peregrine Falcon Recovery Team as well as cooperating agencies to aid in evaluation of recovery efforts. SEGMENT OBJECTIVES 1. Annually monitor the number of breeding pairs of peregrines and reproduction in Colorado. 2. Annually monitor organochlorine pesticide levels in wild breeding peregrines. 3. Monitor recruitment of reintroduced peregrines into the wild breeding population of Colorado. 4. Compile data and submit reports to appropriate state and federal personnel and the Rocky Mount9in/Southwest Peregrine Falcon Recovery Team for use in evaluating recovery efforts. (These objectives correspond to Jobs 1113., 1114., 212., 221., 3211., 3212., and 333. of the approved American Peregrine Falcon Recovery Plan for the Rocky Mountain/Southwest population.) METHODS AND MATERIALS Methods and materials used in this study have been described previously (Craig and Enderson 1981). RESULTS AND DISCUSSION Territory Occupancy In 1985, 1 previously unrecorded breeding territory (site 43) was located and site 40 which was discovered in 1984, was vacant. In addition to discovery of a new pair, a historic territory (site 4) was reoccupied after 11 years of vacancy. In all, 14 sites were occupied by pairs in 1985 (sites 3,4, 8, 25, 27, 30, 31, 34, 35, 38, 39, 41, 42, and 43). After a vacancy of 11 years, site 4 was reoccupied by a pair of released falcons that successfully bred. Adults comprised members of 12 pairs while subadult females were paired with adult males at sites 39 and 43. The subadult feamle which occupied site 39 did not return to breed in 1985 and was replaced by another subadult female. Although actual marker and band numbers could not be read, released falcons comprised members of 6 pairs and both members at sites 4 and 39 were released birds. �Reproduction In all, 13 of the pairs produced eggs, including one pair comprised of a subadult female (site 39) that produced 2 fertile eggs. The pair at site 43, which consisted of an adult male and subadult female, occupied a territory but did not settle down and lay eggs. The pair at site 25 that did not breed in 1984 and was thought to be approaching senility, returned and successfully produced young. Two of the pairs (site 35 and 41) failed to hatch young. The pair at site 35 abandoned their eggs before they were full term, and the eggs at site 41 apparantly were infertile (they were not retrieved for analysis) and did not hatch even though the female incubated them in excess of 40 days. Based upon average clutch thicknesses of individual females in previous years, 6 sites (8, 27,30,34,39, and 42) experienced eggshell thinning in excess of 10 percent thin and were earmarked to be manipulated if the same females returned in 1985. The other pairs were permitted to incubate and hatch their own eggs. In actuality, only 5 sites received foster young (sites 8,27,30,39, and 42), the female at site 34 selected an inaccessible ledge and the eggs could not be manipulated, although we were able to reach the site on a subsequent visit to collect eggshell fragments and band the nestlings. Sites 8, 30, and 35 were recycled to augment anticipated reduced captive production resulting from the Peregrine Fund1s relocation from Fort Collins to Boise, Idaho. Subsequent record production at Boise caused these fears to be unfounded. The 13 breeding pairs produced at least 42 eggs in their first clutches (3.23 eggs per pair) of which 18 were removed for artificial incubation. The 3 pairs that were recycled produced 9 additional eggs of which 5 were removed for artificial incubation. Had the pairs been permitted to incubate and hatch their own eggs, it is probable that only 6 of the 18 first clutch eggs would have hatched. The remaining eggs required special incubation at elevated humidities to counter difficulties induced by eggshell thinning, so artificial incubation actually hatched 11 of the eggs. The 8 pairs that were not fostered produced 17 young (2.1 young per pair) of which 13 successfully fledged (1.6 young per pair). When the probable wild production of the fostered pairs is added, the estimated total wild production was 20 young (1.5 young per pair). Foster activities at 5 sites increased total production to 35 young (2.7 young per pair) with a fledging success of 28 young (2.2 young per pair). Eggshell Thickness A total of 27 eggs were collected from the wild in 1985. Shell thickness measurements were taken of the 14 eggs that hatched and the remaining 13 were shipped intact to Patuxent for chemical analysis. Shell thickness measurements averaged O.316mm which is 11.9% percent thinner than pre DDT era eggs. This value is not significantly different than the 1984 average of 0.319mm (-11.1%) or the 1983 average of 0.317mm (-11.7%). These values negate the gradual improvement in shell thicknesses observed from 1977 to 1982. The 1985 average may change when measurements are available from the 13 eggs which have been submitted for pesticide analysis. Eggshell thicknesses averaged less than 14% for females at 5 sites (sites 30, 31 34,39 and 42). The female at site 34 produced eggs of normal thickness (-2.1%) in her first clutch in 1981 and has experienced gradual shell thinning of -8.5% in 1982, -11.7% in 1983, -12.5% in 1984, and -14.5% in 1985. The eggs of the female at site 30 have vacillated from -12.7 to -16.7% thin over the past 6 years but has not demonstrated a clear trend toward decreased thickness. It is ~ �11 probable that several of the older females may have more or less stable levels of DDT while the younger females are still accumulating body burdens. Due to the history of eggshell thinning at all the above sites, it is important to continue manipulation efforts to sustain reproduction if the same females return in subsequent years. The female at site 8 is well in excess of 7 years of age and judging from behavioral irregularities, may be approaching senility. The females at sites 3,27,35,41,42 and and possibly 25 appear to be producing eggshells averaging greater than -10% thin and may be permitted to breed without interferrence. Organochlorine Residues in Egg Contents Pesticide residue analysis results from eggs collected in 1984 are not yet available. Reprioritization of samples sent to the Patuxent Wildlife Research Center delayed chemical work on the eggs. An additional 13 intact eggs from 1985 have also been submitted for analysis. Photographic Documentation of Wild Adults Photographic identification of adults present at eyries was initiated in 1980. Since then, 65 useable photographs have been obtained of individual falcons at 11 territories. Additional falcons have been identified by subadult plumage (site 3 and 39) or the presence of bands. The cumulative data include 57 instances where individual adults could be positively identified among which 45 were of birds from previous years. This results in an overall annual survivorship rate of 0.79. Males were confirmed to have survived one year in 20 of 23 cases (0.87 annual survivorship) and females in 28 of 34 cases (0.82 annual survivorship. These survivorship figures must be considered conservative since individuals may relocate to other sites and not be identified in subsequent years. Adults were present at sites in all 6 years, 3 were present over a 5 year span, and 3 were present for 3 successive years. Literature Cited Craig, G.R. and J.H. Enderson. 1981. Nesting performance of peregrine falcons in Colorado. Job. Prog. Rep., Colo. Div. Wildl., Wildl. Res. Rep., Jan., pp 13-23. Prepared by: tl.t? a~t Gera ld R. ra ig ��13 Colorado Division of Wildlife Wildlife Research Report January 1986 JOB PROGRESS REPORT State of Colorado -------- Project N-l-R-3 Work Plan Job (SE-3-6) Reintroduction and Augmentation of Peregrine Falcon Production (II) 2 (2) -----------~~-------- Period Covered: Author: Personnel: 1 January - 31 December 1984 G. R. Craig D. Berger, R. Beane, E. Bowden, G. Craig, B. Grebence, J. King, and T. Sisk, Colorado Division of Wildlife; J. Enderson, Colorado College; D. McVean, Bureau of Land Management; and S. Petersburg, National Park Service. ABSTRACT Eight pairs of peregrine falcons (Falco peregrinus anatum) were fostered to augment wi 1d product ion in Co lorado. The effort succeeded in increasing wild fledging success to 3.12 young per pair. Hacking efforts at 7 localities successfully released another 27 young into the wild. Released falcons comprised members of 5 of 13 pairs occupying breeding territories. This Job Progress Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the Author. ��15 REINTRODUCTIONANDAUGMENTATION OF PEREGRINE FALCONPRODUCTION Gerald R. Craig P. N. OBJECTIVE The objective of this program is to sustain the wild breeding peregrine falcon population in Colorado through augmentation of poor natural production and re-establishment of breeding pairs at vacant sites by release of captive-produced falcons. SEGMENTOBJECTIVES 1. Augment Door wil d production wild young ana captive-produced 2. Re 1ease capt i ve-hatched wi 1d young and captive-produced young to the wild through "hacking" at potential and abandoned wild nests. 3. Develop and or abandoned adults. 4. implement release nest sites and by placement of captive-hatched young into occupied wild nests. of at adult falcons at potential wi ld nests occupied by lone Monitor results of the efforts, compile data, and submit reports to appropriate state and federal agencies and the Rocky Mountain/ Southwest Peregrine Falcon Recovery Team. METHODSANDMATERIALS Methods and materials (Craig 1982). for this study have been described previously RESULTS AND DISCUSSION Augmentatiori Efforts Eight (sites 3,8,27,30,31,34,35 and 39) of the 13 pairs of peregri nes encountered in 1984 recei ved foster young. Restri cted resources di d not permit mani pul at i on of addit i ona 1 pai rs present at sites 38 and 40. The pair at site 25 did not produce eggs and thus were not available for fostering and site 42 was discovered too late in the breeding season to be augmented. The pair at site 41 was also discovered late in the season and was still incubating infertile eggs, so an attempt was made to foster young to them. When the site was rev is ited to return young to the pair, they had only moderate interest in the eyrie and failed to accept the young. �16 In 1984, 26 eggs were removed from the 8 manipulated sites (3.25 eggs per pair) and were artificially incubated by the Peregrine Fund. Although the Peregri ne Fund was able to provi de foster young at the time required for placement into the wild, it was difficult to match ages of enough young to provide broods of four, and half of the sites received broods of 3 young. In all, 28 young were returned to the wild pairs (3.5 young per pair) and 25 successfully fledged (3.12 young per pair). Golden eagle predation probably accounted for loss of 2 young and disappearance of the third nestling could not be attributed to a cause. Although the fosteri ng efforts obscure the nesting success that the pairs would have normally experienced, an estimate of natural production can be obtained by multiplying hatch success that the wild eggs probably would have experienced had they not received artificial incubation at the Peregrine Fund by 82% which is the fledging success (survivorship of nestlings from hatch to fledging age) calculated by Ratcliffe (1980). Hence, only 7 of the 9 young hatched would have survived to fledging for an estimated fledging success of 0.87 young per pair. Hacking Efforts Hack sites were operated at 7 locations throughout the State in 1984. The sites recei ved a total of 32 young (4.57 young per site) and 27 successfully achieved independence (3.86 young per site). Golden eagle predation plagued site 37 with 2 and probably 3 of the 5 young being killed by eagles. A young was apparently killed by traffic along a highway ~elow site 14 and another falcon was recaptured with a broken leg after release at site 5. Return of Released Falcons Evidence continued to accumulate indicating that release efforts are succeeding in re-establishing falcons in the wild. Aside from a couple of spec i fi c records, it is diffi cu lt to determi ne whether the marked falcons observed in the wild were released through hacking or fostering. The female hacked at site 36 in 1981 again returned to the eyri e in New Mexi co and bred wi th a wil d male. Two banded falcons bred at site 39 received 3 fostered young and fledged 2 young. Although band numbers could not be distinguished, it is probable that both falcons originated from a hack box at site 36 approximately 18 miles distant. Judging from the color of the bands, neither falcon caul d have been more than 3 years of age. The male that was hacked at site 11 in 1981 and occupied site 3 in 1983 with a subadu1t wild female returned in 1984 with another wild sub adul t female. The pair laid 2 eggs, received 3 fostered ynu.ng, and 'fled_ged all of" them. A second banded subadult female was paired with a wild adult male at site 41, but the eggs were infertile. The male hacked at site 8 in 1980 again returned and paired with the wild female. The pair were fostered and fledged 3 young. Finally, a banded adult male �17 was found paired with a wild female at site 42 several miles downstream from site 27. The pai r were discovered late in the breeding season, had laid at least 3 eggs and successfully fledged 2 young without intervention. In summary, released falcons composed 5 of the 13 pairs occupying breeding territories in Colorado in 1984 which is evidence that release activities are bolstering the poor wild reproduction. LITERATURECITED Craig, Ratcliffe, Prepared G. R. 1982. Reintroduction falcon reproduction, Job Prog. Rep., Jan., pp 53-57. s. D.A. 1980. The peregrine Dakota. 416pp. by: and augmentation of peregrine Rep., Colorado Div. Wi1d1. Res. falcon. Buteo Books, Vermillion, ��19 Colorado Division of Wildlife Wildlife Research Report January 1936 JOB PROGRESS REPORT State of Colorado ~~~~-------------- Project No. N-l-R-4 Work Plan No. II ~~-------------- Job No. (SE-3-7) Reintroduction and Augmentation of Peregrine Falcon Production II Period Covered: 1 January 1985 - 30 June 1985 Author: G. R. Craig Personnel: D. Berger, R. Beane, M. Bertram, G. Craig, B. Grebence, D. Palmer, and T. Sisk, Colorado Division of Wildlife; J. Enderson, The Colorado College; D. McVean, Bureau of Land Management; S. Petersburg, National Park Service; and W. Heinrich, the Peregrine Fund, Inc. ABSTRACT Augmentation efforts were undertaken at 5 wild peregrine (Falco peregrinus anatum ) eyries suffering eggshell thinning. The effort increased production to 3.6 young per pair; estimated natural production was 1.2 young per pair. Hacking efforts released 35 young from 7 sites. Only 24 achieved independence and great horned owl predation caused failure of 1 site. Released falcons were present at 6 of 14 occupied territories and released falcons comprised pairs at 2 of the sites. This Job Progress Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the author.· ��21 REINTRODUCTION AND AUGMENTATJON OF PEREGRINE FALCON PRODUCTION Ger~ld R. Craig P. N. OBJECTIVE The objective of this program is to sustain the wild breeding peregrine falcon population in Colorado through augmentation of poor natural production and re-establishment of breeding pairs a~ vacant sites by release of captive produced falcons. SEGMENT OBJECTIVES 1. Augment poor wild production by placement of captive hatched wild young and captive produced young into wild nests. 2. Release captive hatched wild young and captive produced young to the wild through "hacking" at potential and abandoned wild nests. 3. Compile data and submit reports to appropriate state and federal personnel and the Rocky Mountain/Southwest Peregrine Falcon Recovery Team for use in evaluating recovery efforts. METHODS AND MATERIALS Methods and materials used in this study have been described previously (Craig 1981). RESULTS AND DISCUSSION Augmentation Efforts Based upon average thicknesses of eggshells with thinning of 10% and more in previous years, 6 sites (8,27,30,34,39 and 42) were tentatively identified to receive foster young in 1985. In actuality, only 5 sites were augmented in 1985. The pair at site 34 selected an inaccessible ledge and was not manipulated. Left to their own devices, the pair produced 3 eggs, hatched 2 young and fledged both. Site 8 received 4 young, but only 2 successfully fledged. Apparant cause of death of the young appeared to be starvation within 1 and 2 weeks of fledging. Observation at the time suggested that the adults were not bringing prey at sufficient intervals. Site 27 received and successfully fledged 4 young. Four young were fostered to site 30, but a golden eagle attack at the time of fledging killed 1 young, the remaining 3 fledged successfully. The subadult female at site 39 was replaced by another subadult female in 1985. Due to her inexperience and the possibility that the eggs would be infertile, the pair was manipulated. They received and successfully fledged 2 young. It was found that both of her eggs were fertile, but they required abnormally high incubator humidity and would not have hatched in the wild. The pair at site 40 received and successfully fledged 4 young. As in 1984, their eggs were critically thin and averaged -16.2% of normal. In 1985, 14 first clutch eggs were removed from the 5 sites (2.8 eggs per clutch) and artificially incubated at the Peregrine Fund's Boise, Idaho facilities. Given the low clutch sizes, and poor hatchability caused by eggshell thinning, it is estimated that only 6 young (1.2 young per pair) would have been produced if �22 the pairs had not been manipulated. Augmentation efforts brought 18 young to the pairs (3.6 young per pair) of which 15 successfully fledged (3.0 young per pair). In addition, artificial incubation produced 3 young for release that would not have hatched in the wild. The Peregrine Fund's relocation from Fort Collins to Boise, Idaho did cause logistical problems in transport of eggs to Boise and return of young to Colorado for release. In order to reduce stress on the eggs and young, the only effective method of transport was aircraft. Commercial airline flights were attempted, but regulations forbad carrying young in the passenger section (they cannot be transported in the baggage section without great risk) and an airline strike negated that approach. It was quickly discovered that private aircraft was the only solution. Jim Enderson made several flights to transport eggs and return young both for the fostering work as well as young to be hacked. Project Lighthawk also donated an aircraft to transport eggs and return with young. Due to asynchronous laying dates among sites as well as at Boise, several flights were made to transport clutches to Idaho without return of young. In all, 5 trips were made to transport eggs and young. Both NBC and CBS furnished helicopters for transport of eggs and young to the field, and NBC even furnished a jet to fly eggs and young to and from Boise. Both networks furnished the aircraft as part of a documentary they were producing, and they cannot be relied upon for regular assistance in the future. Hacking Efforts Although 6 hack sites were originally planned to be operated, additional production at Boise permitted the activation of a seventh site. Each site received the maximum number of 5 young, thus 35 young were released in 1985. Unfortunately, the releases were not as successful as past years; only 24 young successfully reached independence. Great horned owl predation completely wiped out all the young at site 2 and young disappeared at 3 other sites. Although golden eagle predation is suspected for loss of 2 young, it cannot be proved. Hence, the 1985 hacking achieved only a 68% in Colorado when previous efforts excedeed 80% success. Return of Released Falcons Presence of banded individuals at 6 (sites 3, 4, 8, 39, 41 and 42) of the 14 occupied territories indicates that release efforts are sustaining and expanding the wild population. Although viewing conditions make it difficult to confirm origin of banded individuals, it was possible to determine that the falcons were released either through hacking or fostering. The male at site 3, which was hacked at site 11 in 1981, returned for a third year to breed with a female of wild origin. The pair laid 4 eggs, hatched 3 and fledged 2 young. The male at site 39 , assumed to have been hacked at Rocky Mountain National Park, returned for the second year and bred with another banded subadult female. Site 4 was reoccupied by a pair of released falcons that may have occupied the site in 1984, but went unnoticed. The male that was hacked at site 8 in 1980 returned for the fourth consecutive year to breed with a banded wild female. Released males at site 41 and 42 returned and bred with wild females for the second consecutive year. An adult male a transmission possible that ascertain the peregrine that was fostered at site 27 in 1983 was found dead below line near Fruita, Colorado in Mesa County in May of 1985. It is he struck the wire but the remains were too badly decomposed to cause of death. �23 Literature Cited Craig, G.R. 1982. Reintroduction and augmentation of peregrine falcon reproduction. Job. Prog. Rep., Colo. Div. Wildl., Wildl. Res. Rep., Jan., pp 53-57. a~ ..... GE{r'a"ld R:trli Prepa red by: _ ..••;;'-'!-:---:Gta~~f'..",..._ g ��Colorado Division Wildlife Research January 1986 25 of Wildlife Report JOB FINAL REPORT S tate of _ _,;;C....;;.o_l o~r...;;a;.;;;d....;;.o _ Pro j e c t _....;..N;_-..;;..1-....;..R~(:l..:.W;_-=12;;;....4;_-....;..R L, Osprey Nesting Studies Work Plan 2 (II) Job I (1) Period Author: Covered: 1 January - 31 December 1984 G.R. Craig ABSTRACT The segment was devoted to data compilation prepared and submitted for publication. and analysis. A manuscript will be ��Colorado Division of Wildlife Wildlife Research Report January 1986 27 JOB FINAL REPORT Colorado State of Project Work Plan Job Title: 01-00-045 (N-l-R) 4: Job Avian Research 1 Greater Prairie-Chicken Transplant Period Covered: 1 January through 30 June 1985 Author: Richard W. Hoffman Personnel: C. Braun, L. Budde, M. Creamer, L. Crooks, K. Dillinger, J. Grimes, R. Hoffman, T. Kroening, B. Linkhart, G. Miller, C. Pabst, F. Pusateri, S. Steinert, R. Van Buren, C. Wagner, Division of Wildlife. ABSTRACT Efforts to restore warm season grasses, reduce densities of sand sagebrush (Artemisia filifolia), and re-establish populations of greater prairie-chickens (Tympanuchus cupido) on the South Tamarack Wildlife Area were initiated in 1978 and continued through 1985. Restoration techniques have included reseeding, tilling, mowing, and burning. In 1984, 36 prairie-chickens (16 males, 20 females) were trapped in Yuma County, Colorado, and released on the Tamarack Prairie. Forty (15 males, 25 females) greater prairie-chickens were captured with cannon nets on 6 leks in Yuma County between 26 March and 16 April 1985 and released on the South Tamarack. A mean of 3.3 birds was trapped per capture attempt in 1985. Peak hen attendance in 1985 occurred between 7 and 11 April which coincided with the most capture attempts (6) and captures (21). Five leks were located adjacent to the South Tamarack, but no leks were found on the property. Number of males per lek ranged from 1 to 7. Males released in 1985 were observed displaying within one week post-release. Five (3 females, 2 males) unbanded birds were observed on leks in 1985, indicating successful reproduction in 1984 and recruitment into the 1985 population. Four males and 9 females were equipped (poncho attachment) with lithium (2) or solar- (11) powered transmitters for post-release monitoring in 1985. An instrumented female released in 1984 was relocated and also monitored in 1985. Six (2 males, 4 females) of the 14 instrumented birds were tracked from early April through mid-July, 3 (1 male, 2 females) died pr ior to la te-Ma y , and radio con tact was los t wi th the remaining 5 birds (1 male, 4 females). Both males moved to the vicinity of a 1ek and one was observed displaying. Five hens were known to have nested and 3 were successful. Mean distance of nests from a lek was 1.5 + 0.8 km. Average clutch size, including a renest, was 10.6 eggs (range 7=12). The maximum documented dispersal distance was 29 km, but this bird eventually returned to within 4 km of the release site. Mean maximum dispersal distance was 6.1 km (N K 7). Although booming grounds were established and successful reproduction- and recruitment were documented, it is still premature to consider the transplant a success. ��29 GREATER PRAIRIE-CHICKEN TRANSPLANT, 1985 Richard W. Hoffman Greater prairie-chickens historically occupied 15 of 63 Colorado counties, but are presently restricted to Yuma, eastern Washington, and southern Phillips counties (Evans 1964, Miller 1981). Recent sightings have occurred in Logan, Morgan, and Kit Carson counties, however, established breeding populations are unconfirmed (Van Sant 1983). The reduction in occupied range has been attributed to changing land use practices that have resulted in habitat alteration. Warm season grasses such as bluestem (Andropogon spp ,), switchgrass (Panicum virgatum), and Indiangrass (Sorghastrum nutans) once provided critIcal nesting, brood rearing, and winter cover because of their resistance to flattening by snows and heavy rains (Duebbert et a1. 1981). These grasses have been replaced by croplands or overgrazed rangelands dominated by sand sagebrush and shorter, less resistant grasses such as blue grama (Bouteloua gracilis), sand dropseed (Sporobolus cryptandrus), and needle-and-thread (~tipa comata) (Miller 1981, Snyder 1984). Loss of nesting and brood rearing abitat appears to be the universal limiting factors for prairie-chickens throughout their range (Kirsch 1974). Efforts to restore warm season grasses and reduce densities of sand sagebrush on the South Tamarack Wildlife Area were initiated in 1978 and continued through 1985 (Snyder 1984). The signif icance of this property is that it represents the only major Division of \·'Lldlifeholding within the historical range of the prairie chicken where a ~eintroduction program is feasible. Restoration techniques have included reseeding, tilling, mowing, and burning. In 1984, 36 prairie chickens (16 males, 20 females) were trapped in Yuma County and released on the Tamarack prairie (Jenniges 1984). Another release was made in 1985. This report summarizes the 1985 trapping operations, lek surveys, and post-release monitoring of transplanted birds. A change of personnel responsible for trapping and monitoring precluded the inclusion of data collected in 1984. P. N. OBJECTIVES Major objectives of this study are to (1) trap and transplant 40 greater prairie-chickens (80% females, 20% males) onto the South Tamarack Wildlife Area, (2) evaluate the success or failure of greater prairie-chickens to establish a breeding population on the South Tamarack and surrounding rangelands, and (3) develop guidelines for the introduction of greater prairie-chickens into new or previously occupied habitats. STUDY AREA Transplant stock was obtained from the Colorado sandhills in northeastern Yuma County, north of Wray.This area, previously described by Evans (1964), supports the highest density of greater prairie-chickens in the state (Evans and Gilbert 1969). Captured birds were released onto the South Tamarack �30 ~. Wildlife Area, Logan County, 2.5 km. south and 0.5 km east of the junction of Interstate 76 and Logan County Road 93 (Fig. 1). The South Tamarack is part of the Tamarack Ranch purchased by the Colorado Game and Fish Department (Division of Wildlife) in 1949. Interstate 76 was constructed through the ranch in the late 1960's. That portion (1,820 ha) south of the interstate is referred to as the South Tamarack or Tamarack Prairie •• Lek surveys and radio-tracking were conducted on the South Tamarack and 330 km2 of adjacent rangelands south of 1-76 extending from Logan County Road 55 east to Colorado 59 (Fig. 1). The area is best characterized as a rolling sandsage-bluestem prairie interspersed with dryland wheat and irrigated corn fields (Miller 1981, Jenniges 1984). Livestock grazing was terminated on the South Tamarack in 1977, but varies from moderate to intensive on adjacent rangelands. METHODS Status of prairie-chickens on the Tamarack and surrounding areas was ascertained by (1) field reconnaissance, (2) personal communications with individuals knowledgeable about the area, and (3) review of prior reports (Van Sant 1983, Jenniges 1984, Miller 1984). Plant communi ties were preViously described by Miller (1981). Locations where males were observed booming in 1984 were revisited in 1985. In addition, 135 sections, including the South Tamarack property (Fig. 1), were systematically surveyed for booming activity by driving or walking major ridgelines between 0400 and 0700 MDT, and listening for booming males. If any booming activity was heard, the lek was visually verified and revisited to determine the number of displaying males, identity of marked birds, and presence or absence of hens. Leks were observed using binoculars (7 x 50) or a spotting scope (20X)from a vehicle parked within 50-100 m away. Lek locations were recorded as UTM coordinates and plotted on 7.5 minute U. S. Geological Survey topographic maps for future reference. Prairie chickens were captured on leks with cannon nets (Dill and Thornsberry 1950, Braun 1976, Giesen et al. 1982). Specific trap sites on the lek were selected by observing the location of dominant males and identifying female attendance patterns. In some cases where the same leks were trapped in 1985 as in 1984, no pre-trapping reconnaissance was conducted and nets were set based on 1984 trapping success. Two nets were set on larger leks or when hen attendance patterns were inconsistent. If necessary, nets were shifted on the lek as dominant males were removed. Both 10 x 20 m (3 cannon set-up) and 12 x 23 m (4 cannon set-up) nets were used. Nets were reset on the same day they were fired and, except for one occasion, were not discharged on consecutive mornings. Captured birds were weighed, classified to age and sex, and marked with a serially-numbered aluminum band on the right leg and a red plastic, numbered bandette on the left leg. Thirteen birds were also equipped with solar (11) (Wildlife Materials model SPCB-1250-3X) or lithium (2) (Telonics model RB-2) powered transmitters mounted on a poncho collar (Amstrup 1980). The birds were held.in burlap sacks, transported by truck, and released within 5 hours of when trapped. Birds trapped on the same day were released simultaneously. All releases were made at the same location on the South Tamarack (Fig. 1). �x; 1 ....-,--+-110---- I '" --''. -r--' l\ ,\ I~ I~. • \( I -IB . \ .... 1 +--1 " I -.~1 ~.. ~tR'<r" I ~I \,\f. I i~ 1), . ... ~ ~ : : h\~ . -, • e" eee ••• :2: --- 34 D~ 0 • t "J I if';:~;' ' ··1\,····, \~ I • ~'J ~ .. I~- ..;; 1 \ '\ -- 1 --+-----, .' ••• •• ~.l-I·_J_-: 2 ._, --_ •• '-. - .".. ~. • ,. I I I • MI., • , ...+....:.___. 4 ••• X X LEGEND STUDY AREA SOUTH TAMARACK RELEASE SITE NEsT SITE LEK I· !_ X I r--- • ! .' 0 ,. \ Figure 1. •••••• • X • "OK.·- ":0 I. , J 2 3 , 4 SCAlE •• Kl.OM£TERS __,__ r , SCAlE •• MtlES ,. 5 \ " N -===t==H:::.-~--1 l" ..-,,1.--••... t_·· ...•.. I It 5 '6. 4 • P H 1 : 9 r. T~'1~-~~r; Greater prairie-chicken release site and nest and lek locations, northeastern Colorado, 1985. LV f-' �32 Radio-marked grouse were located at least twice per week while they were in signal contact. Considerable time was spent searching for grouse from which signal contact was lost. The principal method of radio tracking was from the ground using a hand-held, 3-element yagi antenna and Telonics TR-2 receiver. An aerial search was made on 22 May 1985 to locate missing birds. Locations of instrumented grouse were determined by visual observation and recorded to the nearest 50 m as UTM grid coordinates. RESULTS AND DISCUSSION Trapping and Transplanting Forty greater prairie-chickens were captured using cannon nets on 6 leks in Yuma County between 26 March and 16 April and released onto the South Tamarack Wildlife Area (Table 1). Five additional males were trapped and immediately released as they were not needed to fulfill the transplant requirements. Another male died from stress within 2 hours of capture. No other mortalities occurred during trapping and transplanting operations. Nine leks were monitored as potential trap sites (Table 2). Nets were set on 8 of the 9 leks and birds were actually trapped on 6 leks (Table 1). Repeated attempts to survey Kitzmiller III resulted in the birds shifting their lek attendance pattern. Thus, a reliable trap site could not be determined. Nets set on Conrad 116 were continually disturbed by cows and were eventually removed. A net was set on Brian 01 but there was no opportunity to trap 2 or more birds simultaneou~ly. Table 1. Age and sex composition of greater prairie-chickens South Tamarack Wildlife Area, Colorado, 1985. Trap location a Kitzmiller 05 Tuell Conrad 08 Conrad III Brian 02 Homestead Dates traEped Adult 4,6,8,10,11,16 Apr 26 Mar; 3,11 Apr 9,16 Apr 1 Apr 9 Apr 4 Apr Totals aLeks where birds were trapped. Male Yearling Adult released on the Female Yearling Total 5 4 3 2 0 0 1 0 0 0 0 0 13 6 0 2 0 1 1 0 0 0 2 0 20 10 3 4 2 1 14 1 22 3 40 �33 Table 2. Lek attendance by greater prairie-chickens on 9 leks monitored for trapping in Yuma County, Colorado, 1985. Peak count Males Females N Lek counts Kitzmiller 15 Tuell Homestead Conrad #la Conrad 116b Conrad D8 Kitzmiller III Brian III Brian 112 13 11 10 10 4 10 3 2 5 12 8 6 8 8 16 15 6 6 Mean count Females Males 10.0 6.9 5.7 6.4 4.8 14.4 14.5 5.5 4.7 9 5 5 8 4 9 0 1 3 3.7 3.1 1.4 2.3 1.8 3.8 0.0 0.5 2.0 aAlso referred to as Jill's lek. bAlso referred to as Cow lek. A mean of 3.3 birds was trapped per capture attempt. The peak of hen attendance occurred between 7 and 11 April which coincided with the most capture attempts (6) a~c captures (21) (Table 3). There were 5 instances when the nets did not fire correctly. In all cases, the misfires were attributed to condensation or san.; in the cannons causing the projectile to stick upon firing. Fortunately, at least 1 bird was trapped during each misfire, but an estimated 15 birds escaped. Table 3. Hen attendance County, Colorado, 1985. Observation period Females/lek x 18-22 Mar 23-27 Mar 28 Mar-01 Apr 02-06 Apr 07-11 Apr 12-17 Apr aIncludes captured. patterns and trapping males that were a N captured Capture attempts 1.4 2.4 1.7 2.4 4.1 2.4 5 success on 6 leks in Yuma 0 1 1 4 6 2 released and 0 4 4 13 21 4 1 that died after being �34 Other than a few broken primaries and missing tail feathers, all birds were released in good condition. Thirty-seven of the 40 birds transported to the Tamarack flew upon release, including 13 of 14 radio-marked birds. Average flight distance was about 100 m. Radios were not attached until just prior to releasing the birds. Originally, 14 prairie chickens were equipped with radios, but a lithium radio weighing 29 gm was removed from an 882 gm adult female after she would not fly. The other hen equipped with a lithium radio weighed 982 gm and flew about 200 m after being released. Solar ponchos were 7-12 gm lighter than the lithium ponchos and had no apparent influence on the bird's flight behavior. No radio package exceeded 4% of the bird' s total weight (Table 4). Within 2 days post-release, flights in excess of 1 km were frequently observed when birds were accidentally flushed during monitoring. Table 4. Weights (gm) of greater prairie-chickens Tamarack Wildlife Area, Colorado, 1985.a Descriptive statistic N Sf SD Range released on the South Adult male Adult female Yearling female 14 1,051 63 943-1,195 22 895 43 829-982 3 922 51 885-980 aOnly 1 yearling male weighing 975 gm was captured. Lek Surveys Five leks were located (Fig. 1) and surveyed (Table 5) on areas surrounding the South Tamarack. No leks were found on the property, but leks 1, 2, 3, and 4 were within 2.3 km (range 1.0-2.3 km), while lek #5 was 10.4 km away. It was difficult to identify birds on leks due to the structural characteristics of the vegetation surrounding the leks and the tendency for males to shift their territories if approached too closely. No males on leks 3, 4, and 5 were identified as being banded or unbanded. At least 2 males on lek III were marked with green bandettes (1984 release), 2 were unbanded (offspring from 1984 release) and 3 were of unknown status. Three of 7 hens observed on lek III were survivors from the 1984 release (green bandettes), one was unbanded, and 3 were of unknown status. Five males attending lek #2 were marked with red bandettes (1985 release) including an instrumented male (TX 1730). No females were observed on lek #2, but 4 instrumented hens (TX 0888, 0974, 1101, 1302) were consistently located within 1.5 km of the lek. �35 \. Table 5. Lek /I a Lek surveys, Logan County, northeastern Colorado, 1985. Observation date K birds observed Males Females Unk 1 05 09 24 25 02 17 24 04 18 Apr Apr Apr Apr May May May Jun Jun 7 7 7 4 3 5 lb 5 0 1 3 0 3 0 0 0 0 0 0 0 2 2 0 0 0 0 0 2 17 25 30 02 09 04 13 Apr Apr Apr May May Jun Jun 5 9 6 6 6 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 3 01 May 06 May 07 May 1 1 1 0 0 0 0 0 0 4 07 May 17 May 1 1 0 0 0 0 5 16 May 22 May 2 1 0 0 0 0 aLek locations are depicted in Fig. 1. bOnly one male on the lek, but heard others off the lek. Leks 1 and 2 were established in 1984, although 1ek 2 shifted about 0.5 km north-northwest of its 1984 location. One male was observed on lek 1 in 1984 and 5 on lek #2. It is unknown, but assumed, that they were males released in 1984. No other leks were verified (males observed displaying at the same location on 2 or more occasions) in 1984. There were 2 other sites where booming was heard in 1984, but no males were found. On repeated visits to the same sites in 1984 and 1985, no booming was heard. Males released in 1985 were attracted to lek #2 possibly because it was closest (3.7 km) to the release site. The next closest lek (111) was 5.4 km away. Booming males can be heard from both leks from the knoll overlooking (south) the release site. Displaying males were present on lek #1 prior to �36 releasing birds in 1985, but it is uncertain if males were displaying on lek #2. L. Crooks observed 6 males at the old site of lek 112 on 18 March 1985. L. Crooks, B. Linkhart, and J. Grimes flushed one male from the same site on 21 March 1985. No males were found on 5 April 1985. On 17 April, 5 males were heard and observed 0.5 km north-northwest of the old lek site. There must have been booming males in the vicinity of lek #2 prior to the 1985 release otherwise it is uncertain why the 1985 released males were attracted to the area around lek 112. However, no unbanded birds or birds banded in 1984 were identified on lek #2. Telemetry Investigations Radio transmitters were placed on 13 greater prairie-chickens (4 adult males, 8 adult females, 1 yearling female) to obtain data on dispersal, breeding behavior, nesting, and survival. In addition, an adult female instrumented and released in 1984 was relocated in 1985 with her radio still functioning. A male carrying a backpack-mounted radio was flushed on 25 April. This male was released in 1984 as backpack mounts were not used in 1985. His radio was not transmitting and he was not observed on a lek. Several mornings were spent listening for booming activity in the general vicinity where this male was flushed, but no booming was heard. When located, he was about 3.5 km north-northeast of lek 112. One hundred and sixteen radio locations verified by visual observations were obtained from April through early July; 8 birds (2 males, 6 females) accounted for 92% of the radio locations (Table 6). Radio contact was lost with 3 birds (1 male, 2 females) within 2 days post-release and with 2 other females within 30 days post-release. It is doubtful these birds would have escaped detection unless their radios malfunctioned. A 3.5-hour aerial search on 22 May 1985 was unsuccessful in locating any missing birds. Signals were received from all other radios known to be functioning at the time of the flight. Only 3 mortalities were documented, 2 from avian predators and 1 caused by a mammalian predator. Six birds (2 males, 4 females) were tracked throughout the monitoring period. The maximum distance any bird was located from the release site was 29 km (Table 6). This bird was an adult male (TX 0859) that eventually returned to within 3.8 km of the release site and remained there through the duration of the monitoring period. The other male (TX 1730) for which dispersal data were available was never located more than 6.4 km from the release site. TX 0859 was eventually found in association with TX 1730 near lek 2 on several occasions. The mean maximum dispersal distance for 6 females and 1 male (excluding TX 0859) tracked at least 30 days post-release was 6.1 km (range 5.2-9.0 km) (Table 6). Comparing total distances traveled (j ••• 11.1 km, range 9.1-16.2 km, excluding TX 0859) to the mean maximum dispersal distance (6.1 km) indicates that once the birds dispersed from the release site they became relatively sedentary. Dispersal southeast from the release site was oriented towards in the direction of leks 1 (SW) and 2 (SE). the southwest and Three birds moved �Table 6. TX Freg. Movements Ase of greater-prairie chickens released on the South Tamarack Wildlife Area, Colorado, Sex Date released N radio iocations Distance from release site (km) Max Fina1a Total movements (km) 0859 2+ M 09 Apr 13 29.0 0920 2+ M 08 Apr 1 -- - - 1088 2+ M 16 Apr 2 -- - - 1730 2+ M 11 Apr 10 6.4 4.0 13.7 0887 0888 2+ 2+ F F 11 Apr 04 Apr 0 7 -- 5.4 4.9 11.3 0974 1+ F 09 Apr 23 5.0 4.4 7.9 1001 1018 2+ 2+ F F ot Apr 26 Mar 4 20 4.8 9.0 1264 1101 2+ 2+ F F 11 Apr 08 Apr 0 15 1302 2+ F 08 Apr 1500 2+ F 1125 2+ F aDistance 3.8 - - 67.5 -- 9.0 8.1 16.2 -6.9 5.7 13.3 6 5.2 3.8 7.0 11 Apr 2 -- - -- Apr 84 13 5.0 4.9 6.4 from release site where the bird finally localized. -- 1985. Fate Observed within 50m of 1ek 02 in association with TX 1730. Still alive at termination of study. Observed 0.3 km from release site on 9 Apr. No signal received after 9 Apr. Found dead on 15 May 1.5 km from release area. Did not receive signal between 17 Apr and 15 May. Observed displaying on 1ek 112. Still alive at termination of study. Not located after release. Localized by 30 Apr as if starting to nest. Lost contact after 4 May. Nested successfully on second attempt. Still alive at termination of study. Lost contact after 23 Apr. Nested successfully. Still alive at termination of study. Not located after release. Nested successfully. Still alive at termination of study. Nested - killed 5 m from Still incubating at nest. time of death. Lost contact after 17 Apr. Found radio on 4 June 6.1 km from release area. Apparent mortality. Nested unsuccessfully. Lost contact after 25 June. VJ " �38 southwest (2) or south-southwest (1) and 7 moved east-southeast (2), southeast (3) , or south-southeast (2)• TX 0920 was only found on the day after his release and not thereafter. He was southwest of the release site when located. TX 0887 and 1264 were not relocated after their release. The only evidence of a bird moving in a northerly direction across the interstate was the recovery of a partially chewed transmitter (TX 1500) (no feathers or carcass) 2.1 km west and 200 m north of the junction of Colorado 138 and Logan County Road 93. Six hens were tracked until the nesting period of which 5 definitely nested, including 1 renest (Table 7). TX 0888 localized and behaved like a nesting hen, but contact was lost with her on 4 May before a nest could be located. TX 0974, a yearling female, was flushed from a clutch of 3 eggs on 29 April after which she abandoned the site; by 15 May she was incubating another clutch of 7 eggs. Average clutch size of the other 4 nests was 11.5 eggs (range 10-12). Eggs in 3 of 5 clutches (60%) hatched. Hatching success was 90% (26 of 29 eggs in successful nests hatched). Two of the unhatched eggs contained fully developed chicks and 1 was infertile. No hens released in 1985 were observed on a lek. However, the mean distance of nests from a lek was 1.5 + 0.79 km suggesting that hens were attracted to the leks (Fig. 1). TX 1125, released in 1984, was also not observed on a lek in 1985, yet she nested. Thus, it's possible the newly introduced hens were visiting leks after they were released. Whether they successfully copulated before or after their release is unknown. Copulations were observed on leks during trapping operations in Yuma County. Little use of the South Tamarack was documented despite the vegetative restoration efforts. Birds released there dispersed off the property within 2-3 days and didn't return. This seems to be a common reaction of transplant birds; I.e., to disperse from the release site regardless of the suitability of the habitat. Both the 1984 and 1985 releases were made on the South Tamarack. If in 1984, some birds had been released on private land to the south, they may have dispersed onto the Tamarack, especially if tape-recorded calls had been used to attract birds to potential lek sites. This approach would not have worked in 1985 because lek #1 and possibly lek #2 were already established and acted to lure the 1985 transplanted birds off the property. CONCLUSIONS Attempts to introduce prairie-chickens into formerly occupied ranges have been mostly unsuccessful and poorly documented (Kruse 1973). Consequently, when the reintroduction program in northeastern Colorado was proposed, guidelines regarding timing of releases, stocking levels, number of releases per area, distribution of releases, and age and sex composition of released stock were lacking and needed to be developed. The procedures followed appeared to work as booming grounds were established, and successful reproduction and �Table 7. Nesting activity of greater prairie-chickens released on the South Tamarack Wildlife Area, Colorado, 1985. TX II Age Date of nest1n~a Date of hatch - Nearest lek(km) Distance from release area(km) Clutch size N eggs llatched 1.2(lek 112) 5.0 12 0 16 Jun 0.7(lek #2) 4.4 7 7 14 May 07 Jun 1.0(lek 113) 9.0 12 11 2+ 11 Jun ,07 Ju1 1.9(lek #2) 5.7 10 8 2+ 15 May 2.7(lek #2) 3.8 12 0 1125 2+ 05 Jun 0974 1+ 15 May 1018 2+ 1101 1302 - Fate of nest Nest d~predated. No si.~ of eggs. Flushed from a clutch of 3 eggs on 29 Apr. Abandoned first nest and renested. Last observed on 10 Jul with 6 chicks. Last observed on 10 Jul with 5 chicks. Last observed on 10 Jul. Did not find any chicks, but she acted broody. Killed by a Swainson's Hawk 5 m from nest on 28 May. aDate when first observed on a nest. W 1.0 �40 recruitment were documented. However, post-release evaluations have not verified that recruitment is exceeding losses in the breeding population. It is therefore premature to consider the transplant a success or to recommend guidelines for future releases. RECOMMENDATIONS 1. No further releases should be made on the South Tamarack as enough birds have been released to insure establishment provided the habitat is suitable (Kruse 1973). 2. Emphasis in 1986 (and preferably for 3 years post-release 1986-88 ) should be placed on monitoring existing leks (i.e., recording the number of displaying males/1ek) and searching for new leks (Le., recording total leks/area). Both reconnaissance techniques will provide an index to population size (Cannon and Knopf 1981) and address the question of whether recruitment is exceeding mortality in the breeding population. Surveys should be conducted from late March through early May. Leks should be counted a minimum of 3 times at 7-10 day intervals. All lek locations should be plotted on U.S.G.S. topographic maps for future reference. Records should be kept of the number of females observed/count and the number of marked (red or green bandettes) and unmarked birds observed/count. Since all birds released in 1984 and 1985 were marked, a count of marked and unmarked birds on leks will provide an index of recruitment into the breeding population. 3. Vegetation restoration efforts should continue on the South Tamarack, and if possible, expanded to surrounding rangelands by promoting land use practices that are beneficial to or at least compatible with prairie-chicken requirements. However, habitat requirements of greater prairie-chickens in Colorado are poorly understood and in need of better quantification. A study should be implemented to compare the floristic, structural, and topographic features of grouse use sites and random sites within occupied ranges, and between random sites within occupied and unoccupied ranges. The purpose of this study would be to identify habitat components lacking in unoccupied habitats that could be managed to enhance the suitability of the habitat for prairie-chickens. 4. Releases in other areas should be postponed until the success or failure of the Tamarack introduction is more clearly documented and the characteristics of occupied ranges are quantified. 5. Brood surveys as a means of monitoring production are not recommended because of the difficulty in finding broods and in interpreting the results of brood surveys. 6. Data collected in 1985 should be incorporated with data from 1984 into a manuscript entitled "Experimental introduction of greater prairie-chickens in northeastern Colorado". This manuscript should include guidelines for the introduction of greater prairie-chickens into new or unoccupied habitats. �41 LITERATURE CITED Amstrup, S. C. 214-217. 1980. A radio-collar for game birds. J. Wildl. Manage. 44: Braun, C. E. 1976. Methods for locating, trapping, and banding band-tailed pigeons in Colorado. Colorado Div. Wildl. Spec. Rep. 39. 20pp. Cannon, R. W., and F. L. Knopf. 1981. Lek numbers as an index to prairie grouse populations. J. Wildl. Manage. 45:776-778. Dill, H. H., and W. H. Thornsberry. 1950. A cannon-projected capturing waterfowl. J. Wildl. Manage. 14:132-137. net trap for Duebbert, H. F., E. T. Jacobson, K. F. Higgins, and E. B. Podoll. 1981. Establishment of seeded grasslands for wildlife habitat in the prairie pothole region. U.S. Dep. Inter., Fish and Wildl. Servo Spec. Sci. Rep. - Wildl. 234. 2lpp. Evans, K. E. 1964. Habitat evaluation of the greater prairie chicken in Colorado. M.S. Thesis, Colorado State Univ., Fort Collins. 98pp. , and D. L. Gilbert. 1969. A method for evaluating greater ---chicken habitat in Colorado. J. Wildl. Manage. 33:643-649. prairie Giesen, K. M., T. J. Schoenberg, and C. E. Braun. 1982. Methods for trapping sage grouse in Colorado. Wildl. Soc. Bull. 10:224-231. Jenniges, J. J. 1984. Greater prairie-chicken transplant, 1984. Intern Program. Colorado State Univ. Unpubl. Rep. l3pp. Kirsch, L. M. 1974. Habitat management considerations Wildl. Soc. Bull. 2:124-129. Student for prairie chickens. Kruse, A. D. 1973. Prairie chicken restoration projects. Pages 40-46 in W. D. Svedarsky and T. Wolfe, eds. Proc. Conference on the prairie chicken in Minnesota. Univ. Minnesota, Crookston. Miller, G. C. 1981. Development of a preservation program for three species of prairie grouse. Pages 38-52 in Colorado Div. Wildl. Fed. Aid Rep. SE-3-3. Jan. 1984. Development of a preservation program for three species of prairie grouse. Pages 129-170 in Colorado Div. Wildl. Fed. Aid Rep. Part 1. Jan. Snyder, W. D. 1984. Sandsage-bluestem prairie restoration. Colorado Div. Wildl. Prog. Narr. Fed. Aid Proj. W-37-R. Work Plan 21, Job 3. Van Sant, B. F. 1983. Census of the greater prairie chicken in Colorado. Colorado Div. Wildl. Unpubl. Rep. l8pp. Prepared ��Colorado Division of Wildlife Wildlife Research Report January 1986 43 JOB PROGRESS REPORT State of Colorado Project 02-06-200 (N-5-R) Work Plan Job Title: 23: Job Avian Research 2 Investigations on the Effects of Public Viewing on Selected Species Period covered: 1 January 1984 through 30 June 1985 Author: Janet L. Schreur Personnel: Eugene Decker, Ron A. Ryder, and Janet L. Schreur, Colorado State University; Clait E. Braun, Dave Langlois, and Bert Widhalm, Colorado Division of Wildlife. ABSTRACT The effects of public viewing on selected marsh bird species were investigated at Russell Lakes State Wildlife Area during 1984 and 1985. Fourteen persons in 4 groups participated in guided birdwatching tours from 26 May through 16 June 1985. Sixty-four percent of the participants visited Russell Lakes to birdwatch. The highlight of the visit for 13 of the 14 participants was observing a marsh bird species. Mean number of marsh bird species per ha (4.2) and individuals per ha (42.0) counted on undisturbed wetlands was twice that counted on disturbed wetlands (2.1 species/ha and 19.6 individuals/ha)' On undisturbed wetlands, the number of broods and young (2.2 broods/ha and 12.9 young/ha) was twice the number counted on disturbed wetlands (1.0 and 5.1, respectively). Mean flushing distance of snowy egrets (Egretta thula) from a disturbance was 96 m in 1984. During April-June 1985, snowy egrets and white-faced ibis (Plegadis chihi) flushed at slightly more than 90 m from a person on foot, but a vehicle could approach to 56.8 and 53.7 m, respectively. Quantified responses of redheads (Aythya americana) were dependent on the agent of disturbance. Snowy egret and white-faced ibis reproductive success was monitored at 3 heronries in the San Luis Valley during 1984. Snowy egret and white-faced ibis mean clutch size was 4.16 and 3.56, respectively. The highest snowy egret hatching and fledging success was 66.2 and 61.8% at Monte Vista National Wildlife Refuge. White-faced ibis were most successful at Head Lake with 62.8% of the eggs resulting in fledglings. The lowest reproductive success for both species was at Trites Lake. Disturbance by photographers in 1984 caused abandonment of most nests at this site. This Job Progress Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the Author. ��45 INVESTIGATIONS ON THE EFFECTS OF PUBLIC VIEWING ON SELECTED SPECIES Janet L. Schreur Historically, wildlife management has been directed toward consumptive uses of game species. The justification was that virtually all funding was derived from hunting, fishing, and trapping license fees and excise taxes. However, with accumulating evidence that a large percentage of the U.S. population actively participates in nonconsumptive wildlife recreation, some attention has been redirected (Schreur 1984). By examining demographic trends, investigators predict participation to .continue to increase. In response to this trend, the Colorado Division of Wildlife developed management stratgegies to encourage and provide opportunities for nonconsumptive wildlife recreation (Colo. Div. Wildl. 1983). With the expected increase in nonconsumptive use, there should be corresponding increases in impacts on wildlife resources. Nonconsumptive use may have a larger impact on wildlife populations than hunting because it can occur year around. At present, spring and early summer receive most pressure because viewing and photographic opportunities are. maximized by concentrated bird migrations, dramatic breeding rituals, and the presence of attractive, approachable young. In spite of its potential impact, little work has been done to investigate the impacts of wildlife viewing on wildlife populations. Information about the amount, type, and timing of public viewing is necessary to minimize its effects. Therefore, the Colorado Division of Wildlife in cooperation with Colorado State University initiated an investigation into the effects of public viewing on selected avian species. Experimental disturbances and guided birdwatching tours were conducted at Russell Lakes State Wildlife Area (SWA). A heronry comprised of nesting black-crowned night-herons (Nycticorax nycticorax), snowy egrets, and white-faced ibis was highlighted. These 3 Ciconiiformes were selected as sensitive species because they are large, relative uncommon, and sho~~ birds that nest colonially. These attributes make them desirable for public viewing (Gray 1975). There has also been concern about the status and future of black-crowned night-herons, snowy egrets, and white-faced ibis. Incidences of poor reproductive success in these species have been linked to DDE (Capen and Leiker 1979, Steele 1980, Findholt 1984, Henny et a1. 1984, McEwen et a1. 1984), changes in water levels (Graul 1977, Alford 1978, Ryder et ale 1979), and human disturbance (Kaneko 1972). The objectives of this study were to identify public demand for guided birdwatching tours, establish guidelines for public viewing activities which minimize disturbance to Ciconiiformes and associated marsh birds, and test the effects of different methods of viewing marsh birds. �46 STUDY AREA The study was conducted at 4 sites in the San Luis Valley, Colorado. Work was concentrated at Russell Lakes in Saguache County and at Monte Vista National Wildlife Refuge (MVNWR) in Rio Grande County. San Luis Lake and Head Lake in Alamosa County were also visited. The San Luis Valley, Colorado's largest intermountain park, is in the south-central part of the state. The Sangre de Christo Range on the east and the San Juan Mountains on the west border the relatively flat ancient lake bed. The valley, ranging from 2,285 to 2,440 m in elevation, is 240 km long and has a maximum width of 80 km (Svoboda 1984). The climate is characterized by cool, sunny, short summers and cold dry winters. Annual preciptation averaged 26.8 cm from 1931 through 1960, with July and August being the wettest months (U.S. Dep. Commerce 1978). The growing season is 95 to 120 days in length (Svodoba 1984). Although the climate is arid, portions of the valley have high water tables and many artesian wells. A low divide just north of the Rio Grande River separates the valley into 2 basins (Ryder 1951). Streams draining the northern San Juan and Sangre de Christo mountains disappear into the porous soil of the northern closed basin or are used for irrigation. Water resurfaces and evaporates at San Luis Lake, the low point of the closed basin. The southern half of the valley is drained by the Rio Grande River. The valley floor is dominated by an association of black greasewood (Sarcobatus vermiculatus) and rabbitbrush (Chrysothamnus spp.). Inland saltgrass (Distichlis stricta) is also common on dryland sites. Many shallow alkaline lakes and sloughs are scattered throughout the valley. Wetlands are dominated by hardstem bulrush (Scirpus acutus), baltic rush (Juncus balticus), and common cattail (Typha latifolia). Associated wetland plants include sago pondweed (Potomogeton pectinatus), duckweed (Lemna minor), and water milfoil (Myriophyllum spicatum). Snowy egrets, black-crowned night-herons, and white-faced ibis nest in bulrush wetlands at Russell Lakes SWA, Harrence Lake, Head Lake, San Luis Lake, Trites Lake, and at MVNWR. One of the 2 heronries at MVNWR was in a mixed stand of bulrush and cattail. In this case, 22% of the white-faced ibis nests and 4% of black-crowned night-heron nests were in cattail. Wetlands containing heronries were ei,ther natural (Harrence, Head, San Luis, and Trites lakes) or artificial reservoirs (Russell Lakes SWA and MVNWR). METHODS Tours Birdwatching tours were scheduled for Saturdays and Sundays beginning 20 April and ending 30 June. Reservations for tours were taken in advance. Tours began at 7:00 a.m. Participants were guided on foot along a designated trail at Russell Lakes SWA passing within 85 m of the heronry. Each tour lasted approximately 180 minutes although only 10 minutes were spent near the heronry. Responses of snowy egrets, white-faced ibis, and black-crowned night-herons were recorded. Questionnaires were distributed, completed, and collected at the conclusion of the tour. �47 Comparative Marsh Bird Counts Wetland study units were selected in the Russell Lakes area during 1984 and were divided into treated and untreated sites. Units were selected based on similarity of size and emergent vegetation. Three units on Russell Lakes SWA were treated sites due to control of access, while 4 untreated units were on nearby private property. Treated areas were open to public viewing while untreated were not. Marsh birds were counted on study units from 1/2 hour before to approximately 2 hours after sunrise 3 times per week. Counts were accomplished with a 20x spotting scope from a vehicle on a vantage point. Birds were identified and counted by species. Brood size and age-class were also recorded. Aquatic vegetation was sampled in all 9 units during August with a method modified from the Atlantic Waterfowl Council (1972). Transects were 50 m apart with sampling points every 30 m. A rake was used to sample submerged vegetation. Water depth was measured at each sampling point. Disturbance Trials Disturbance trials were conducted during July through August 1984 and April through June 1985 at Russell Lakes SWA. Marsh birds on the study wetland were counted from a blind before and for one hour after a disturbance. Time between counts was 15 minutes in 1984, but was shortened to 5 minutes in 1985. The agent of disturbance was either 1 or 2 persons on foot or a motor vehicle. Disturbance agents traveled along a dike on one side of the wetland and returned at a slow steady speed. Responses of selected marsh birds to the disturbance were quantified (Table 1). Table 1. Quantification Valley, Colorado. Response values o 1 2 3 4 of marsh bird responses to disturbance, San Luis Response None Watched disturbance Moved away from disturbance Rapidly moved away or dove Flew from unit Flushing Distance Flushing distances of snowy egrets and white-faced ibis resting and feeding at Russell Lakes SWA were measured with a Leitz rangefinder during June through August 1984 and April through June 1985. Flushing distance was the distance from the disturbance agent to the subject when the latter flushed. Disturbance agents were people on foot or in a vehicle that directly approached the subjects at a slow steady speed. �48 Ciconiiformes Reproductive Success Reproductive success of snowy egrets and white-faced ibis nesting in heronries at Trites Lake, MVNWR, and Head Lake was monitored from June through August 1984. Nests were located by wading ~.uiet1'y through the heronry and were marked with a 2 x 2-cm tag. Nests ver.e relocated every 9-11 days and the number of eggs and young were recorded. RESULTS AND DISCUSSION Tours Four tours were conducted between 26 May and 16 June 1985. 100% and no tours were cancelled. Attendance was Responses to questions 1-7 of the Russell Lakes guided tour questionnaire were combined to measure satisfaction (Table 2). In questions 1 and 2, the participants were asked to fill in ~he blanks. There were 2 responses possible for questions 3 through 5 ~ith the first response indicating satisfaction. Questions 5, 6, and 7 referred to monetary values placed on .nonconsumptive uses. Table 2. Responses to questions 1-7 of the birdwatching Russell Lakes SWA, Spring 1985. Question l. 2. 3. 4. 5. 6. 7. Purpose of visit Birdwatch Guided tour See a snowy egret Historic site Close to home Highlight of visit Heronry Variety of species Marsh wren American avocet Cinnamon teal Knowledgable guide Viewing distance Satisfactory Too far Viewing unobstructed Yes No Return visit Yes No Pay to view or photograph wildlife Yes No Nongame Income Tax check-off contributor Yes No N responses 14 8 3 1 1 1 14 8 2 1 1 1 1 14 14 0 14 14 0 12 12 0 13 11 2 14 9 5 tour questionnaire, % 57 21 7 7 7 57 14 7 7 7 7 100 0 100 0 100 0 85 15 64 36 �49 Sixty-four percent of the participants visited Russell Lakes SWA to birdwatch and the highlight of the visit for 13 of the 14 participants was viewing a marsh bird species. All participants were satisfied with viewing opportunities. Twelve of the 14 responded that they would make a return visit. Eighty-five percent would pay $3.00 or less to view or photograph wildlife, and 64% had contributed to Colorado's nongame income tax check-off (two participants were not Colorado residents and did not have an opportunity to contribute). Thirty-six percent of the participants fishing license in Colorado during the percent had hunted and fished and 21% had percent had not purchased either a hunting had purchased either a hunting or last 5 years (Table 3) • Fourteen fished, but not hunted. Sixty-four or fishing license. Table 3. Hunting and fishing Russell Lakes SWA, Spring 1985. of birdwatching activity N respondents (N •• 14) - Activity - Hunted or fished Hunted and fished Fished, did not hunt Hunted, did not fish Did not hunt or fish 5 2 tour participants, % 3 36 14 21 9 64 o o Demographic characteristics of the participants varied (Table 4). All participants were 28 years of age or older and had college training. Participants traveled from 11 to 202 km to visit Russell Lakes SWA with the average being 74 km. Table 4. Age, sex, and education of birdwatching tour participants, Russell Lakes SWA, Spring 1985. N Age S 16 17-19 20-29 30-39 40-49 50-59 ~60 Sex Male Female Education High School College Graduate responses 14 0 0 3 4 2 1 4 14 6 8 13 0 8 5 % 0 0 21 29 14 7 29 43 57 0 62 38 �50 Comparative Marsh Bird Counts The mean number of avian species counted on each treated wetland (7.2) differed slightly from untreated wetlands (6.3) (Table 5). The density of species (4.4/ha) and individuals (42.0/ha) on untreated wetlands was twice that observed on treated wetlands (2.l/ha and 19.6/ha, respectively). Table 5. Indices of marsh bird use of study units at Russell Lakes Colorado, 1984. Category Species Mean SEecies Untreated WE-13 KLS WE-12 KL All 3.5 6.0 6.4 9.1 6.3 4.7 4.8 5.2 2.7 4.4 24.7 50.7 57.1 35.6 42.0 Treatment WE-3 WE-l WE-4 All 4.8 8.4 8.4 7.2 1.3 1.8 3.4 2.1 6.7 20.6 31.5 19.6 SWA, Individuals This same trend also occurred in brood size and density (Table 6). The average number of young per brood differed only slightly between treated and untreated wetlands. On untreated wetlands, the 2.2 broods/ha and 12.9 young/ha recorded was twice those on treated wetlands (1.0 and 5.1, respectively). Table 6. Indices of marsh bird reproductive success at Russell Lakes SWA, Colorado, 1984. Category Brood size Mean Broods/ha Young/ha Untreated WE-13 KL KLS WE-12 All 3.3 3.4 3.8 3.9 3.6 3.4 1.8 0.8 2.7 2.2 13.5 7.6 11.1 19.4 12.9 Treatment WE-4 WE-1 WE-3 All 3.3 4.0 4.0 3.7 2.1 0.5 0.5 1.0 10.1 2.4 2.8 5.1 �51 Selected physical and vegetati ve parameters differed between untreated and treated study units (Table 7). Treated study units were larger and had more open water where birds could be counted than untreated areas. Treated units were also deeper and had denser aquatic vegetation than untreated areas. Table 7. Physical and vegetative SWA, Colorado, 1984. Category Area censused (ha) Untreated WE-13 KLS WE-12 KL Treatment WE-4 WE-3 WE-I parameters Total area (ha) of study units at Russell Lakes .!. water depth (cm) .!. aquatic veg.density 0.74 1.24 1.24 3.43 0.74 1.24 1.24 3.43 20 10 25 15 3.8 0.3 0.6 1.2 2.46 4.12 4.65 9.11 5.78 4.65 53 56 29 3.9 3.7 2.8 Disturbance Trials Responses to disturbances and resulting changes in the number of marsh birds on wetlands at Russell Lakes SWA were recorded in 1984. The number of redheads counted on a wetland usually declined after a disturbance, but many returned within 1 hour (Fig. 1). Redhead response differed (!_ < 0.05) when disturbed by a person on foot vs. when disturbed by a motor vehicle (Table 8). Redheads most frequently responded to a person on foot by swimming to the far side of the wetland. In contrast, the most frequent response to a motor vehicle was watching the disturbance while maintaining their position. �52 10 9 8 7 6 tf.) p 5 ::t:: 4 ~ 3 < I>J P zl ..'" . ..... .'. ...... .... .."., •..~ . ' 2 •••<.~ .•.• 1 0 B D 15 30 45 60 TIME Fig. 1. Number of redheads at treated areas by time intervals. The disturbance occurred between Band D. Each line represents a separate trial on a different date. �53 Table 8. Redhead duck response frequencies when approached motor vehicle (X2 - 21.98, 2 d.f.) on foot or in a Frequenc~ of resEonse Vehicle Foot Response 1 2 4 Obs EXI> Obs Exp 37 99 39 52 82 41 47 32 27 32 49 25 Flushing Distance Mean flushing distance of snowy egrets was 96 m in 1984 (Table 9). Both snowy egret and white-faced ibis flushing distances from persons on foot and vehicles were measured in 1985. Both species flushed from persons on foot at slightly more than 90 m, but could be approached more closely by a vehicle (56.8 and 53.7 m, respectively) (Tables 10 and 11). Table 9. Flushing distance (m) of snowy egrets approached in a vehicle or on foot, San Luis Valley, Colorado, 1984. N j'_ flushing distance 19 Range S.D. 46 27-200 96 Table 10. Flushing distance (m) of snowy egrets approached in a vehicle or on foot, Russell Lakes S.W.A., Colorado, 1985. l- DisturbcDce Foot Vehicle 16 28 flushing distance 93.3 56.8 Range S.D. 57-150 18-110 42.7 26.6 Table 11. Flushing distance (m) of white-faced ibis approached or on foot, Russell Lakes S.W.A., Colorado, 1985. Disturbance li .x flushing distance Foot Vehicle 42 14 94.5 53.7 in a vehicle Range S.D • 38-205 18-105 48.2 26.8 �54 Egrets, ibises, and black-crowned night-herons did not flush from nests as groups of 2 to 16 persons or a vehicle passed at 85 m. Snowy egrets normally stood and watched the disturbance pass. Ibises and herons usually remained motionless in the bulrush during the disturbance. Heronry Reproductive Success Snowy egret and white-faced ibis hatching and fledging success was monitored at Head Lake, Trites Lake, and MVNWR in 1984. Mean clutch size of snowy egrets differed slightly between heronries and was 4.16 overall (Table 12). The mean clutch size of white-faced ibis was 3.56 eggs/nest. Table 12. Reproductive parameters for snowy egrets and white-faced ibis at 3 heronries in the San Luis Valley, Colorado, 1984. Snowy egrets Heronry N nests x clutch size Range S.D. N nests White-faced ibis x clutch size Ranse Head Lake Trites Lake MVNWR 17 23 43 4.12 4.39 4.05 2-6 3-6 2-6 1.17 0.78 0.87 20 4 7 3.45 3.75 3.86 2-5 3-4 3-4 Totals 83 4.16 2-6 0.92 27 3.56 2-5 S.D. 0.69 0.50 0.38 Hatching and fledging success of snowy egrets and white-faced ibis varied markedly between heronries. Young egrets and ibis were considered to have fledged when they ran from their nest and hid in the vegetation at my approach. This occurred at about 2 weeks of age. The highest snowy egret hatching (66.2%) and fledging (61.8%) success were at ~~ (Table 13). Ibis were most successful at Head Lake with 62.8% of eggs laid producing fledglings (Table 14). Hatching and fledging success were lowest for both species at Trites Lake as only 7.6% of the egret and 14.8% of white-faced ibis eggs hatched. Table 13. Hatching and fledging success of snowy egrets in 3 heronries in the San Luis Valley, Colorado, 1984. Heronrx: Head Lake Trites Lake MVNWR N nests 16 28 41 !! eggs laid Eggs hatched 55 170 157 18 13 104 Young fledged 17 13 97 Percent Hatched Fledged 32.7 7.6 66.2 30.9 7.6 61.8 F1edged/ hatched 94.0 100.0 93.3 �55 Table 14. Hatching and fledging success of white-faced ibis in 3 heronries in the San Luis Valley, Colorado, 1984. Heron!I Head Lake Trites Lake MVNWR N nests 13 8 6 N eggs laid 43 27 23 Eggs hatched 28 4 12 Young fledsed 27 4 10 Percent Hatched Fledsed 65.1 14.8 52.2 62.8 14.8 43.5 Fledged/ hatched 96.4 100.0 83.3 The largest loss of reproductive effort was in the egg stage. If an egg hatched, the resulting young had a good chance of surviving to 2 weeks of age. Eighty-three to 100% of the eggs that hatched resulted in fledglings (Tables 13 and 14). The single event causing the greatest loss of reproductive effort was the abandonment of 89% of the snowy egret nests and 75% of the white-faced ibis nests at Trites Lake. The most probable cause of abandonment was the activities of 2 photographers within the heronry. They entered the heronry early on 23 June 1984, photographed nestlings and placed a camera near egret nest 1/40. This camera was triggered remotely from 80 m. Approximately 3 hours were spent near the heronry that day. The same activities were repeated on 24 June. Most nests were found abandoned during the regularly scheduled count on 27 June. The few nests that remained active were on the periphery of the heronry farthest from nest 1/40. LITERATURE CITED Alford, E. H. 1978. Early nesting by white-faced ibis in relation to habitat: an adaptive advantage. M.S. Thesis, Brigham Young Un1v., Provo, UT. 42pp. Atlantic Waterfowl Council. 1972. Techniques handbook of waterfowl habitat development and management. Habitat Devel. and Manage. Comm., Atlantic Waterfowl Counc., unpaginated, 2l7pp. Capen, D. E., and T. J. 1.eiker. 1979. DDE residues in blood and other tissues of white-faced ibis. Environ. Pol1ut. 19:163-171. Colorado Division of Wildlife. 1983. Today's strategy ••• tomorrow's wildlife: comprehensive management plan for Colorado's wildlife. Colorado Div. Wi1dl., Denver. 96pp. Finho1t, S. L. 1984. Organochlorine residues, eggshell thickness, and reproductive success of snowy egrets nesting in Idaho. Condor 86:163-169. Graul, W. 1977. Nesting success of the white-faced ibis in Colorado - 1976. Unpubl. Rep., Colorado Div. Wildl., Denver. 5pp. �56 Gray, G. C. 1975. Nonconsumptive demand for wildlife by municipal conservation commissioners in Massachusetts. M.S. Thesis, Massachusetts, Amherst. 94pp. Univ. Henny, C. J., L. J. Blus, A. J. Krynitsky, and C. M. Bunck. 1984. Current impact of DDE on black-crowned night-herons in the intermountain west. Wildl. Manage. 48:1-13. J. Kaneko, K. D. 1972. Nesting of the white-faced ibis (Plegadis chihi) of Utah Lake. M.S. Thesis, Brigham Young Univ., Provo, UT. 76pp. McEwen, L. C., C. J. Stafford, and G. L. Hensler. 1984. Organochlorine residues in eggs of black-crowned night-heron from Colorado and Wyoming. Environ. Toxicol. and Chem. 3:367-376. Ryder, R. A. 1951. Waterfowl production in the San Luis Valley, Colorado. M.S. Thesis, Colorado State Univ., Fort Collins. 130pp. ___ , W. D. Graul, and G. C. Miller. 1979. Status, distribution and movements of Ciconiiformes in Colorado. Pr oc , Colonial Waterbird 3:49-58. Group Schreur, J. L. 1984. Literature review: the demand for nonconsumptive wildlife uses. Colorado Div. Wildl. Res. Rep. Part 2:387-397. Steele, B. B. 1980. Reproductive success of the white-faced ibis: the effects of pesticides and colony characteristics. M.S. Thesis, Utah State Univ., Logan. 64pp. Svoboda, P. L. 1984. Natural history and geographic relationships of the Brazilian free-tailed bat in the San Luis Valley of Colorado. M.S. Thesis, Fort Hays State Univ., Hays, KS. 69pp. United States Department of Commerce. 1978. Gale Res. Co., Detroit, MI. 606pp. Prepared by Approved by J~L. ~c~ Janet L. Schreur o.a Graduate Research Assistant ~avfE" ~ ClatE:Br~ Wildlife Research Leader Climate of the states. �Colorado Division of Wildlife Wildlife Research Report January 1986 57 JOB FINAL REPORT State of Colorado __ ------~~~--------- Project No. 02-06-200 (N-5-R) Work Plan __ 23 Job Title: ..;::..:._ lb Development of Sage Grouse Viewing Tours in North Park, Colorado Period Covered: Author: Job No. Nonconsumptive Use of Wildlife 1 February 1984 - 30 June 1985 JoAnn Profera Personnel: JoAnn Profera and Eugene Decker, Colorado State University; Clait E. Braun, John F. Corey, Steve H. Porter, and Steve F. Steinert, Colorado Division of Wildlife ABSTRACT Tours to a sage grouse (Centrocercus urophasianus) lek were conducted to measure public demand for viewing tours. Ten guided tours were conducted in 1984 and were attended by 66 people. Poor weather and road conditions limited participant numbers. Participants were satisfied with tours. Sixteen self-guided tours were conducted in 1985, with 85 people attending. Participants were satisfied with the tour and 97% followed the directions given and remained in their vehicles and on designated roads. Trials were conducted on the day before and after each tour as grouse were approached on foot and in a truck to determine the distance they first reacted to approach. In both years, females flushed sooner than males and the distance at which males reacted was inversely related to the number of females present. In 1985 all grouse approached reacted as no males continued to display when approached. Females generally flushed when approached. ��59 DEVELOPMENT OF SAGE GROUSE VIEWING TOURS IN NORTH PARK, COLORADO JoAnn Profera Wildlife management programs historically have been directed toward consumptive users (1.e., hunters and fishermen) because they provided state agencies with the majority of their revenue. Despite agency emphasis on wildlife used for sport, there is a growing interest in non-consumptive uses of wildlife. In 1980, 54% of the people in the United States participated in non-consumptive wildlife activities, outnumbering consumptive users 2:1 (U.S. Dep. Inter. 1982). Although interest in non-consumptive wildlife activities is increasing, opportunities to participate in these activities have been limited as state agencies offer few such opportunities. Shaw and King (1980) found that 66% of non-consumptive users believed that management benefitted mostly hunters and game species. Wildlife agencies and managers should be aware that public interests are shifting toward more non-consumptive uses such as wildlife viewing (Hendee 1969). To prevent a loss in their support base as well as comply with legal mandates to manage all species of wildlife for the benefit of all citizens, agencies must find ways to provide the public with non-consumptive wildlife use opportunities. In 1984, a pilot program was hitiated by the Colorado Division of Wildlife in coordination with Colorado State University to measure demand for and feasibility of wildlife viewing tours. Guided tours to sage grouse leks were selected for rbe :initial effort. Guided viewing tours conducted in 1984 stimulated public interest and participation while providing maximum control over participants and minimizing grouse disturbance. However, the use of guides and guided tours may not be economically feasible. Thus, in 1985 a self-guided tour program was initiated as an alternative to guided tours. Objectives in 1985 were to investigate the feasibility of self-guided tours and identify their impacts on sage grouse. Recommendations were developed for viewing tours that minimize disturbance to sage grouse. STUDY AREA The study was conducted in North Park, Jackson County, Colorado. North Pak is a large intermo;,~·tain park at an elevation of about 2,500 m surrounded by mountains rising sharply to 3,800 m. Small streams in the park flow in a northerly' directicn and are the headwaters of the North Platte River. Over 1,870 km2 of sage grouse habitat lie within North Park, of which 65% is dominated by sagebrush (Beck 1977) primarily Artemisia tridentata vaseyana and A. 1. wyoingensis. Herbaceous vegetation consists primarily of cespitose perennial forbs and bunchgrasses (Beck 1977). The climate is characterized by cool temperatures, low precipitation, and a short growing season. The study was conducted at Coalmont Lek, < 1 km southeast of coalmont. The lek is on the northwest side of a bench east of Pole Mountain Reservoir and is bounded on the west by Little Grizzly Creek and on the east by Grizzly Creek. The arena is approximately 200 m wide and 300 m long. Herbaceous vegetation �60 dominates the relatively flat display area with dense sagebrush stands on the north, east, and south sides. The western edge is bounded by a steep hill sloping to an abandoned coal mine. A north-south jeep trail bisects the lek about 10 m from the center. A 2nd north-south jeep trail occurs about 90 m west of the center. Sage grouse arrive at the lek from the east-northeast. Males are distributed on territories in a general north-south direction. As the season progressed, males also displayed in the dense sagebrush surrounding the lek. Birds departed the lek to the west-southwest. METHODS Lek Selection In March 1984, several leks in North Park were considered for developing opportunities for sage grouse viewing. The leks considered were Coalmont, Delaney Butte, and Perdiz. Criteria for lek selection included vehicle access in early April, expected number of males, and viewing distance to birds. Due to unusually late snowmelt and poor road conditions, Coalmont was the only lek meeting the criteria, therefore it was used for the guided viewing tours. This lek was also used for self-guided viewing tours in 1985. Guided Tours 1984 Guided public viewin~ tours were scheduled on Saturdays and Sundays from 5 April to 27 May 1984. News releases were prepared and released, and an interpretive handout and questionnaire were developed prior to 1 April. Reservations were made by phone on a first-come basis and participants were contacted the week of the tour to confirm reservations. Tours began at 0430 with a bird presentation and distribution of interpretive handouts. Participants drove their own vehicles 27 m from the meeting place to the lek. Vehicles were led single file to the lek along the jeep trail 90 m west of the lek and participants observed gruse displays from their vehicles. The tour leader moved among vehicles on the side away from the grouse answering questions and relaying information. As grouse activity diminished, questionnaires were distributed and completed. Tours concluded with collection of questionnaires. Self-guided Tours 1985 In November 1984, the lek was examined and a tour route was established (Appendix). This tour route incorporated the existing county road and jeep trails, 3 wooden reflective directional signs, and a parking area. Steel posts designated the parking area and restricted traffic on the jeep trail bisecting the lek, 20 m from the center of grouse activity. Signs were wired to four-foot high steel posts. Two small aluminum signs were used to identify and direct participants to the parking area. An observation site was located 1 km southeast of the parking area on Jackson County road 26A. Both the lek and the tour participants were visible from �61 this site. Press releases were sent to all Colorado newspapers and National Audubon Society chapters in Colorado. An information packet was developed. prior to 1 April. This packet included an interpretive brochure, a questionnaire, and a map (Appendix). Tours were scheduled for Saturdays and Sundays from 6 April to 26 May 1985. Due to limited space at the lek and to facilitate observation of tour participants, a reservation system was established limiting the number of vehicles present on any date to 8. Participants made reservations by phone at which time a variety of information was taken. This included tour date, and nu mber of vehicles and participants in the group (Appendix). Information packets were individually prepared and included 1 map per vehicle, 1 brochure per 2 individuals, and 1 questionnaire per participant. Packets were sent by mail. Participants were contacted by phone the week of their tour, confirming their reservation and receipt of the information packet. On tour dates, I arrived at the observation site at 0415 in an unmarked truck and recorded tour group arrival time, activity , and departure time (Appendix). The number if visible male sage grouse along with display activity was also recorded. Sage grouse activity in response to any disturbance was recorded as either walking away or flushing. All observations were made with a 20x spotting scope. On 20 April and 11 May poor weather reduced visibility and the lek was not visible from the observation site. Consequently, participants were observed from the parking area. Disturbance Experiment An experiment designed to measure distances at which grouse reacted to approach was conducted the day preceding and following each tour. In 1984 the experiment was conducted between 27 April and 28 May. In 1985 the experiment was conducted between 07 april and 27 May. Distance was marked at 10-u: intervals in a straight line from the viewing location to the center of grouse activity. Upon arrival approximately 1/2 hour before sunrise, a count was made of all males and females present (Jenni and Hartzler 1978). Two trials were conducted by approaching birds along the marked line at specific intervals in a truck and then on foot. Birds were observed for 20 seconds and distance to the nearest male and female along with their activity were recorded. Bird activities were recorded as either strutting for males or as walking away (males and females). Distances that males and females first fluished were also recorded. Birds were not forced to flush if still displaying when approached to 10 m. RESULTS Guided Tours 1984 Ten tours were conducted between 28 April and 27 May. Tours scheduled prior to 21 April were cancelled because of deep snow on the lek and poor road conditions. Reservations were made by 95 people; 66 (69.5%) actually attended. Tour length ranged from 80 to 155 minutes. Birds were present on the lek at time of departure on all tour dates except 20 Hay when birds fluished at 0526 MDT due to movements of a participant from the vehicle. Distances from vehicles to the center of sage grouse activity ranged from 90 to 120 m. �62 Questionnaires Most participants were satisfied with tour length, type, and amount of information presented (Table 1). A substantial percentage (37) would have preferred to be closer to the grouse and some participants (22%) wanted to visit more than 1 lek. Eighty-seven percent of the participants were interested in self-guided tours. However, when directly asked, 88% of the participants preferred a guided tour. Seventy-eight percent of the participants were willing to pay to view sage grouse (Table 1). Of those willing to pay, 83% would pay up to $5.00 each to view sage grouse. Seventy-three percent of the participants had contributed to the Nongame Income Tax Checkoff Program. Table 1. Sage grouse Spring 1984. tour participant satisfaction, North Park, Colorado, N Category Tour length Satisfied Too long Too short Information presented Satisfied Too technical Too simple Amount of information Satisfied Too much Not enough Viewing distance Satisfied Too far Too close Car caravans Effective Not effective Information handout Useful Not necessary Sage grouse viewing Remain at 1 lek Visit 2 or more leks Take self-guided tour Yes No Pay to view sage grouse Yes No Contribute to Nongame Income Tax Checkoff Program Yes No responses % 51 4 2 89 51 91 5 9 o 7 4 47 1 82 9 16 36 21 63 37 o o 46 11 81 19 37 18 67 33 2 50 78 14 22 55 87 8 13 50 78 14 22 48 18 73 27 �63 Forty-five percent of the participants did not purchase a hunting or fishing license. Twenty-five percent purchased only a fishing license while 27% purchased both hunting and fishing licenses (Table 2). Table 2. Consuomptive wildlife use by sage grouse tour participants, North Park, Colorado, Spring 1984. N respondents (H - 65) Category % 45 27 25 3 29 18 16 2 Did not purchase hunting or fishing license Purchased hunting and fishing license Purchased fishing license Purchased hunting license There was no difference in percent of the participants that were male or female. All participants were at least 20 years of age and 38% were at least 50 years of age. The participants were well educated as 86% had attended college and 50% had gone to graduate school (Table 3). Participants were also well motivated as they traveled an average of 307 km from their residence to the start of the tours in Walden. Most participants were from the denver metro (31%), Larimer (22%), and boulder (20%) county areas. The other participants originated from 5 other counties in Colorado with 1 participant from Wyoming. Almost 22% (21.5) of the total participants were from western Colorado. Table 3. Age, sex, and education Park, Colorado, Spring 1984. Category of sage grouse N responses tour participants, North % Sex Male Female 34 32 51 49 9 22 10 11 14 14 33 15 17 21 1 8 24 33 2 12 36 50 Age 20-29 30-39 40-49 50-59 ~60 Highest education completed Elementary High school College Graduate school �64 Grouse Disturbance 1984 Distances at which grouse reacted to approach varied (Table 4). Approach distance to males was inversely related to the number of females present. A maximum of 59 hens was counted on 7 May and males continued to display when approached on foot to 20 m. On 25 May no hens were observed and males flushed at 90 m when approached on foot. Females usually began walking away and flushed sooner than males. Both males and females began walking away and flushed sooner when approached on foot than when approached in a truck. No males or females flushed upon arrival or during the initial 1ek count. On 18 May, all birds flushed from the lek at 80 m when approached in the truck. Table 4. Distance (m) at which male and female sage grouse reacted approached in a vehicle and on foot, North Park, Colorado, Spring 1984. N Date 27 30 04 07 Apr Apr May May II May 14 May 18 May 21 May 25 May 28 May N uTes fWles 32 25 31 35 37 32 34 40 29 30 52 42 47 59 22 4 2 1 0 2 Continued to d1sElaI 20 30 30 20 50 75 AEEroached bI truck Females Hales Began Began Continued to walk to walk to Flushed Flushed disElaI awaI awaI 30 40 30 20 40 20 25 40 70 70 60 90 40 80 50 55 40 70 20 70 30 50 80 65 40 20 50 70 when AEEroached on foot Females Hales Began Began to walk to walk Flushed Flushed awaI awaI 30 60 60 20 40 65 30 50 90 100 90 80 40 30 35 Not all birds began walking away when approached. Late in the season (14 May), all females immediately flushed from the 1ek when approached by truck. Early in the season, if individual birds were approached either on foot or by truck, they would remain in the lek area. On 21 and 28 May when the individual bird that was approached flushed, the rest of the birds also flushed and left the 1ek area. Self-guided Tours 1985 Participants attended 13 self-guided tours beginning on 14 April and ending on 26 May. Reservations were made for the first 3 tour dates (6-7 and 13 April) but scheduled participants did not attend. Fifty-three reservations were made for 196 people in 61 vehicles. In April, reservations were made by 77 people in 26 vehicles while in May, reservations were made by 119 people in 35 vehicles. Of those individuals making reservations, 85 (43%) actually attended tours. Twenty-four people in 9 vehicles attended in April and 56 people in 22 vehicles attended tours in May. . 20 30 �65 Sage grouse were present and displayed on all tour dates. Participants remained an average of 1 hour and 18 minutes, with 87% of the participants remaining over 1 hour. On 11 May, a golden eagle (Aquila chrysaetos) flew over the lek and all birds flushed and did not return. Participants remained for an additional 15 minutes then departed. Forty-eight percent of the participants arrivedbefore 0500 and 75% arrived before 0530. Sage grouse were present on the lek at departure on half (50%) of the tour dates. Most participants followed the directions given in the brochure. Ninety-four percent of the participants remained in their vehicles at all times. A single individual was observed leaving his vehicle on 3 separate dates. On 28 April, an individual moved to the back of his truck and then returned to the cab. On 11 Mayan individual moved to the back of his truck to photograph sage grouse and remained in back for the duration of the tour. No sage grouse walked away or flushed during these 2 instances. On 12 May, 3 vehicles were observed in the parking aea. At 0700 MDT the sage grouse had left the lek and were dispersed throughout the sagebrush surrounding the lek. Two vehicles left the area while the third left the parking area and stopped on the jeep trail. The individual left his vehicle with a camera and tripod and began moving northeast through the sage. He continued walking in a clockwise direction around the lek flushing 30 sage grouse. This was the only instance when sage grouse were approached and flushed from the area. Ninety-three of the participants parked in the designated parking ara. On three occasions, participants parked on the jeep trail southwest of the lek. On all three occasions participants remained on the trail and in their vehicles. The average number of vehicles was 2.4 per tour. Maximum number of vehicles present was 6 on 5 May. Questionnaires Questionnaires were completed by 51 participants (60%). Participants were satisfied with the brochure, signs, viewing distance, and map (Table 5). Most participants (77%) preferred a reservation system for self-guided tours. Participants heard about the self-guided tours from a variety of sources (Table 6). Most (80%) were willing to pay an average of $2.90 per person to view sage grouse. Forty-six percent of the participants had contributed to the Nongame Income tAx Checkoff Program. Participants expressed a concern over viewing position as 27% would have preferred to view grouse from the eat to avoid looking into the sun. Forty-eight percent of participants did not hunt or fish. Twenty-six percent hunted and fished and the same percentage of participants fished but did not hunt (Table 7). Age, sex, and education of participants on self-guided tours was similar to guided tour participants (TAble 8). Fifty-two percent were female. Age varied with all categories being represented. Seventy-seven percent of the participants attended college or graduate school. Participants traveled an average of 271 km to view sage grouse. Most participants were from the Denver metro (40%), Larimer (19%), Boulder (11%), and El Paso (8%) county areas. Eleven percent were from the western slope. Five participants were not residents of Colorado and traveled frm Texas, South Dakota, Michigan, and Maryland to view sage grouse. �66 Table 5. Sage grouse self-guided Colorado, Spring 1985. tours' participant satisfaction, North Park, N resEonses Catesor;x: Information in brochure sufficient Agree Disagree Signs Useful Distracting Viewing distance Satisfactory Too far Directions on map Easy to follow Needed more detail Sage grouse viewing Reservation First-come basis Pay to view sage grouse Yes No Contribute to Nongame Income Tax Checkoff Program Yes No Table 6. Source of information Spring 1985. Source Colorado Outdoors Wildlife News Division of Wildlife personnel Newspaper articles U.S. Forest Service Audubon Society Friends for self-guided % 48 2 96 4 51 0 100 0 40 11 79 21 49 0 100 0 38 11 77 23 40 10 80 20 22 26 46 54 sage grouse tours, North Park, N responses 3 5 7 15 3 4 7 % 7 11 16 34 7 9 16 �61 Table 7. Consumptive wildlife use by sage participants, North Park, Colorado, Spring 1985. grouse self-guided N responses Category Did not hunt or fish Hunted and fished Fished only Hunter only % 48 26 26 0 22 12 12 0 Table 8. Age, sex, and education of self-guided participants, North Park, Colorado, Spring 1985. tour sage grouse tour N Category responses % Sex Male Female 22 24 48 52 5 1 5 7 8 13 9 10 2 10 15 17 27 19 3 8 24 12 6 17 51 26 Age < 16 17-19 20-29 30-39 40-490 50-59 ;;:.-60 Highest education completed Elementary High school College Graduate school Grouse Disturbance 1985 The distance at which sage grouse reacted to approach in 1985 was consistent with distances obtained in 1984 (Table 9). As in 1984, hens generally reacted sooner than males and both males and females reacted sooner when approached on foot than when approached in the truck. Also, males did not react to approach as quickly when a high number of females was present (15 Apr, 19 Apr, 22 Apr). In contrast to 1984, in 1985 sage grouse reacted to each approach and no males continued to display. On 15 April, with 57 hens on the lek, males began walking away when approached on foot to 50 m and in the truck at 30 m. Throughout both 1984 and 1985, females reacted by imediately flushing when approached. Late in the season on 20 and 27 May when females were present, they walked to the far side of the lek away from approach but did not lush. In 1985, the observed center of grouse activity had moved 15 m east of the observed ceLter of 1984. �0- ex> Table 9. Distance (m) at which male and female sage grouse reacted when approached in a vehicle and on foot. North Park, Colorado, Spring 1985. Date 07 15 19 22 29 06 10 13 20 24 27 Apr Apr Apr Apr Apr May May May May May May N males 31 34 43 33 61 24 22 3 53 58 40 Continued N to females display 28 57 23 51 7 0 0 0 3 0 3 40 60 50 70 , Aeeroached bX truck Males Females Began Began Continued to walk to walk to away away Flushed Flushed diselay 60 30 55 45 65 30 40 50 30 30 30 80 70 60 70 60 65 80 50 50 45 45 85 Aeeroached on foot Males Females Began Began to walk to walk away Flushed away Flushed 60 50 60 50 60 80 80 30 35 40 80 70 70 80 65 70 60 50 75 45 80 30 35 30 80 75 90 75 60 50 80 70 �69 Wooden signs used near the lek were checked after each tour date for signs indicating their use as perches by raptors. Throughout the tour season these signs show no evidence of raptor use. DISCUSSION In both years of the study, the public showed considerable interest in sage grouse viewing tours with 95 reservations in 1984 and 196 reservations in 1985. Although there was a substantial number of reservations, actual attendance at tours was low with 69% participation in 1984 and only 43% participation in 1985. In 1985, 4 tour dates were full with the maximum of 8 vehicles with reservations. At no time was this limit reached at the lek as maximum attendance at any time was 6 vehicles. The distance to area, time of viewing, and unpredictable weather and road conditions are factors that apparently limited actual public attendance. Although the potential for sage self-guided tours, little disturbance remained in their vehicles (97%) and occasion when an individual did leave participants had left the area. grouse disturbance was greater with occurred. Self-guided tour participants did not harass the grouse. On the one his vehicle, he waited until all other Participants on guided tours were interested in getting closer to sage grouse (37%) while self-guided tour participants were satisfied with the viewing distance (79%). Both groups were concerned with viewing direction and suggested viewing grouse from the east side of the lek. Due to the presence of dense sagebrush and the general topography of the area, it would be difficult to view birds from the east. Other leks in North Park should be investigated as possible alternatives for viewing tours. Although distance at which sage grouse first reacted to approach was similar for both years, data for 1985 indicate that sage grouse were more likely to flush from the lek if approached. Sage grouse are tolerant of activity near the lek (Patterson 1952, Connelly et ale 1981), but sufficient distance must be maintained to prevent grouse from flushing. C. E. Braun (pers. commun.) has documented changes in the location of grouse activity on a lek. If tours are conducted at Coalmont Lek in the future, sage grouse activity should be monitored to determine if grouse are reacting to human presence. �70 RECOMMENDATIONS 1. Viewing tours should be made available grouse viewing. 2. Guides need not be present for sage grouse tours and emphasis placed on self-guided tours. 3. Participants sage grouse. 4. Alternative leks should be investigated as possible tour sites so that sage grouse can be viewed from the east (for good light). 5. Sage grouse activity should be mnitored at the tour long-term response of sage grouse to human disturbance. should remain to meet in their vehicles public demand for sage should be at all times when viewing site to document LITERATURE CITED Beck, T. D. I. 1977. Sage grouse flock characteristics and habitat selection in winter. J. Wi1d1. Manage. 41:18-26. Connelly, J. W., W. J. Arthur, and O. D. Markham. 1981. Sage grouse leks on recently disturbed sites. J. Range Manage. 34:153-154. Hendee, J. C. 1969. Appreciative versus consumptive use of wildlife refuges: studies of who gets what and trends in use. Trans. North Am. Wi1d1. and Nat. Resour. Conf. 34:252-263. Jenni, D. A., and J. E. Hartzler. 1978. Attendance at a sage grouse 1ek: implications for spring censuses. J. Wi1d1. Manage. 42:46-52. Patterson, R. L. 341pp. 1952. The sage grouse in Wyoming. Sage Books, Denver, Colo. Shaw, W. W., and D. A. King. 1980. Wildlife management and nonhunting wildlife enthusiasts. Trans. North Am. Wi1d1. and Nat. Resour. 45:219-225. Conf. u.S. Departments of Interior and Commerce. 1982. 1980 national survey of fishing, hunting and wildlife-associated recreation. Colorado supplement. U.S. Dep. Inter., Fish and Wi1d1. Serv., Washington, D.C. 156pp. Prepared by aa... ~ - . - J6AIUl Proiera Wildlife Technician lA Approved by -;:-:;-::~~~~_. -;;-::-::c:r:":":"'~~'-=;"'__ C1a1t E. Braun Wildlife Research Leader _ �71 APPDH>IX �72 Tour route and sign placement for self-guided tours to Coalmont Lek, North Park, Colorado, 1985. N i Colo. 14 JC26 2.6 Km Legend 10 I Sage Grouse •• I 1 I::. No Vehicles Beyond This Point Do Not Drive Beyond This Point 21::.~ Sage Grouse Viewing---. Please Stay in Your Vehicle 31::.1 Parking Area 1 * Questionnaire Box x Steel Posts (wire between posts) • Observation Site �- - - Map to Sage Grouse lek N t Colo. HIghway 14 Hebron ParkIng Area 0 Colo. HIghway 125 x sIgn m ·e ": N sIgn x • JC 26 1.6 miles •• '-J W ��OBSERVER DO's AND DON'Ts Much can be learned about sage grouse and their surroundings by observing their impressive display. We are fortunate that their mating system allows human observation. At. the same time, however, this accessibility can invite human disturbance and interference. This self-guided tour was designed so that with your cooperation in following the listed guidelines and regulations, all may enjoy the display and allow for continued enjoyment in the future. To best enjoy the tours, you should: 1. Arrive at the lek between 4:30and 5:00a.m. (Arriving later may disturb the birds.) SAGE GROUSE OTHER ANIMALS Sage grouse are an integral part of the sagebrush community as are many other birds and mammals. While on your visit to view sage grouse you should watch for these other species: Sage thrasher Golden eagle 2. Dress for cold weather including a hat, gloves, and a blanket. Sage sparrow Rough-legged hawk 3. Be prepared for muddy road conditions. Brewer's sparrow Badger 4. Bring binoculars, spotting scope, camera, and field guides. Horned lark Pronghorn Northern harrier Coyote Prairie falcon White-tailed jackrabbit 5. Observe birds as long as you like, ~ut PLEASE OBSERVE THE FOLLOWING: A Self-Guided Viewing Tour 1. Remain in vehicles at all times. 2. Park in the designated parking area only. 3.' Do not park as to restrict the access of other visitors. 4. No overnight camping on or near the lek. 5. Leave dogs at home. Colorado Division of Wildlife Wildlife Research Center 317 West Prospect Road Fort Collins, Colorado 80526 484-2836 ..._, Vl �THE SAGE GROUSE Sage grouse tCentrocercue urophasianus meaning "spiny-tailed pheasant" in Latin) are the largest grouse in North America. They are birds of the semi-arid plains and are as western as the cowboy and sagebrush. Their range includes parts of 10 western states and 2 Canadian provinces from eastern California and central Colorado north to southern Alberta and Saskatchewan. Sage grouse, or "Cock of the Plains" as d'escribed by Lewis and Clark, are identified by their long, stiff, pointed tail feathers, white breast." and black belly. Sexual dimorphism is quite apparent with males twice as large as females and weighing about 6-7 pounds. Males have a conspicuous black bib and long, thin black filoplumes around the back of the neck. Two yellow-green air sacs are visible during the courtship display. They also have small yellowish-green combs above each eye. Males mature in two years and live only 2-4 years. Their size and bold markings make them highly visible to females and predators including the golden eagle. Females are far less spectacular in appearance. Their gray brown color pattern ~ blends well with sagebrush which helps to conceal them while nesting. Females weigh about 3-4 pounds and live 4-6 years. Sage grouse depend upon sagebrush for food and cuver vear round. THELEK The communal display ground, or lek, provides a vital setting for the sage grouse mating system. The word lek (derived from the Swedish word leka, to play), means to gather or assemble. Leks vary in size and shape but overall have similar characteristics. The lek is an open area relatively free of sagebrush. This prevents deep snow accumulation and allows early spring access for grouse. Grouse activity usually centers around some natural landmark, such as a small bush. Adult males begin to establish territories on the lek in {ate March. One to 3 males will dominate the center territories on the lek and will mate with most of the females. As females arrive at the lek, they associate and mate with the dominant males. Males return to their same position on the lek every day and will display from late March into May. In Colorado, a typical lek may have 25-50males with some leks having as many as 100 males. Once leks are established, they are used for many years if suitable habitat for nesting, brood rearing, and breeding is available. Some leks have been in continuous existence for over 30 years. THE DISPLA YB . Male sage grouse arrive at the lek in evening and early morning, and bt displaying about one hour before sun: When females are ready for mating, the) arrive on leks in the evening and e morning. Males strut with tail feat: fanned and gulp in air to inflate and dis; their yellowish-green esophageal air I As the wings of males move back and fon a brushing motion, the large air eacs moved quickly up and down. As the 8 released, a resonant "poik-poik" or "I plop" sound is made. When females present on the lek, males display I frequently to attract females to territories. A commonly observed display is tb. territory defense. If a male ventures close to a neighbor's territory, the 2 n may stand side by side facing in 0PP directions. Suddenly both may flap wings in an attempt to strike the ot head and back. Injury seldom occurs a intruder usually breaks off the attack returns to his own territory. �77 SAGE CROUSE SELF-GUIDED TOUR RESERVATIONS Date - SPRING 1985 _ Home phone ----------------------------- _ Address Work phone Ci ty & Zip Code Nu:ber of vehicles Name _ _ Number in group _ (No more than 8 vehicles at lek) Tour date: 1st choice -------- 2nd choice _ *Confirmed date Packet sent Questionnaires returned (\~ewill coniirc your reservations 1 week before the date.) _ �78 SAGE GROUSE SELF-GUIDED TOUR DATA SHEET - SPRING 1985 Time Date _ Weather Visibility _ Time of 1st arrival ------- Max. number of vehicles last arrival ---------------- individuals N of questionnaires returned first departure last departure t-1ax.males _ Birds present at last departure? Y Distance from vehicle Number of vehicles ~utside parking area Apparent observer disturbance stopped displaying N females walking away N females flushed N males flushed Notes: N Max. females Number of indo outside vehicle !males _ _ _ Intensity of male ·display Males mostly displaying (> 75%) Males mostly sitting (> 75%) _ �79 SAGE GROUSE SELF-GUIDED TOUR QUESTIONNAIRE Thank you for participating in the self-guided tour to a sage grouse lek • ..This questionnaire is to learn more about your preferences for observing sage grouse. By completing this questionnaire you will provide valuable information that will help us to improve sage grouse viewing in the future. Please place the questionnaire in the box on the si2n as vou leave the lek. Please check the appropriate blank: 1. The brochure provided enough inforcation on sage grouse and their display: ___ 2. The 3. The agree ---disagree? signs at the lek were: ---useful ---distracting? viewing distance to the grouse was: ---satisfactory ---too 4. The directions on the map were: ---easy to fo 11ow ---needed far? core detail? 5. When viewing sage grouse on a self-guided tour, would you prefer: -----a reservation system or ----on a first-come basis? 6. How did you learn about the tour? 7. Would you pay for the opportunity to enter a wildlife area to view sage grouse? Yes No. If yes, how much? 8. Have you fished in Colorado within the past 5 years? Yes No. 9. Have you hunted in Colorado within the past 5 years? Yes No. Income Tax Check-off Program? 10. Have you contributed to the Non~<lr:le .- ----Yes-- No. �80 11.l-1hat town do you live in? 12. Your sex: 13. Your age: ---Male ---Female. ---16 or below __ 17-19 __ 40-49 __ 50-59 __ 20-29 ---60 __ 30-39 or over. 14. What is the highest level of formal education you have completed? ___ E.lementary --....;High ____ College or technical school Do you have any suggestions? school or vocational school ---Graduate Please explain. school! �Colorado Division Wildlife Research January 1986 81 of Wildlife Report JOB PROGRESS State of Project No. Work Plan Job Colorado 86-039-01 1 1 Period Covered: Author: G. C. Miller Personnel: REPORT Ecosystem 1 January M. Bowman, 1985 through G. C. Miller, 31 December Colorado Management 1985 Division of Wildlife ABSTRACT The project was initiated on 01 January 1985, with the objective of integrating ecosystem management principles into Colorado Division of Wildlife (CDOW) habitat management practices. Training, implementation, and monitoring were 3 types of activities designed to lead to that qbjective. Training of CDOW personnel in the process described by Hoover and Will (1984) in Managing Forested Lands for Wildlife (MFLW) began when 241 copies of the book were distributed, primarily to DWMts, AWMts, and biologists. Initial training was provided to 183 CDOW employees, and -hands on- training using Forest Service computers was given to personnel of the Northwest, Southwest, and Southeast Regions (similar training for Northeast and Central Region personnel will take place early in 1986). More intensive training will follow the adaptation of ~ to CDOW's WANG system. Implementation of MFLW on CDOW properties began in June 1985 on lowland riparian properties of the Northeast Region. The Cottonwood and Dodd Bridge SWAts were used to develop standards and guidelines that are lacking for that ecosystem in MFLW. Management units can be described and inventoried for $1. 10 with over 90% accuracy, using 20 habitat types within the ecosystem. Initial findings are that as much as 75% of the areas currently forested on these SWAts will be treeless by the year 2000 unless actions are quickly taken to establish forest stands. Stands must be arranged in forested blocks of at least 25 acres, and mesic haymeadows must comprise at least 400 acres of 15 mile-long riparian segments to attain ecosystem management objectives. In general, however, CDOWappears to have much latitude in manipulating vegetation on its properties without risking detriment to ecological indicator species. Techniques for establishment of cottonwoods are provided. lacre , �82 USDA-Forest Service has scheduled 22 management areas, encompassing approximately 360 mi2 of National Forest lands, for treatment under MFLW processes. AllCDOW Regions are represented by these areas.--ristings of management units and affected DWM's are provided. At present, CDOW personnel should have the expertise to influence management area design through species selection, establishment of habitat objectives, and/or species data on site-specific bases. �83 ECOSYSTEM MANAGEMENT IN THE COLORADO DIVISION OF WILDLIFE G.C. Miller Habitat Investigations The primary objective of the project is to put ecosystem management principles into practice through Colorado Division of Wildlife (CDOW) habitat management activities. Activities designed to attain the objective are of 3 types: Training, Implementation, and Monitoring. In calendar year 1985 progress was made in all 3 areas. SEGMENT OBJECTIVES 1. (Training) Ensure that all permanent CDOW personnel involved in wildlife habitat management activites are trained in the application of Managing Forested Lands for Wildlife (MFLW) (Hoover and Wills 1984) to a level of expertise compatiole witn their responsibilities. Such training may include introductory classes, "hands on" workshops, and/or planning and implementation exercises on CDOW properties. - 2. (Implementation) Assist operation personnel in the implementation of ecosystem management approaches on CDOW properties, including development of appropriate ecological indicator speCies lists. 3. (Monitoring) Monitor USDA-Forest Service implementation of MFLW and insure linkage between implementation and CDOW objectives. ----- RESULTS AND DISCUSSION Training - Distribution of the book, MFLW, to CDOW personnel began on 14 February 1985 with receipt of the Division's copies. As of 31 December 1985, 241 Division personnel, including all DWM's, AWM'St regional terrestrial and habitat biologists had received copies. Recipients of the book who have primary responsibilities for representing CDOW interests on Forest Service lands also received a letter for Director Ruch (Appendix A). In addition to CDOW personnel, copies of MFLW were distributed to the Wildlife Commission, the Nongame Advisory CounCil, and other Colorado agencies (DNR, CSFS, universities and colleges) and wildlife agenctes and associations throughOut the United States. As of 1 July 1985 all but approximately 250 (15%) of the initial printing of 1596 books had been distributed by USDA-Forest Service and CDOW. At least 2 universities wish to use the book as a course text, and 1 state wildlife agency (Arizona) has reQUested copies for their districts. . Most COOW personnel received their initial training in the use of MFLW on 08 and 15 March 1985 at the in-service training sessions in Greeley. Training sessions by Jack capp (USDA-FS) and Gary Miller (CDOW) described the book's contents and demonstrated the manner in which emphaSis and indicator species shOuld be selected (see Graul and Miller 1984). The manner in which CDOW personnel may have greatest influence in the ~ process (selection of �84 emphasis and indicator species, establishing goals and objectives, providjng site-specific information) was described. Tne Forest Service implementatjon schedule, and the 1 April 85 input deadline, was discussed. Processes by whjch management prescriptions may be selected and modified were demonstrated. One hundred eighty-three CDOW personnel have attended at least one introductory training session to date. "Hands on" training sessions, using Forest Service Data General computers, have been held throughout the state for CDOW personnel. Most notable of these were sessions conducted on 19-20 June 1985 at the White River N.F. supervisor's office in Glenwood Springs for 31 Northwest Regional personnel. SUCh sessions also have been held in the Southwest and Southeast Regions, and Northeast and Central regional personnel will receive training NLT March 1986. In 1986 an automated MF'LW program compatible wi th CDml's WANG system wi11 be developed and training-of regional personnel will continue. Implementation - Implementation of MFLW processes on CDm'l lands began in .l,me 1985, with lowland riparian the selected ecosytem. The rationale for using these areas was: 1) high priority of this ecosystem in COOW's habitat management operations and 2) lack of prescription (standards and guidelines) for this ecosytem in MFLW. The results of the exercise is included as Appendix B of this report. Appendix B includes: a system (with estimated costs) to describe CDOW management area; Standards and guidelines for management of forested and non-forested protion of the lowland riparian ecosystem; Potential emphasis and ecological indicator species, with statements about their habitats; Habitat manipulation techniques compatible with ecosystem management Objectives. Monitoring - USDA-Forest Service has scheduled at least 22 management areas for implementation of the process described in MFLW (Table 1). In some cases local DWM's were not involved in the selection process but no CDOW personnel are known to have Objected to any of the selections. Approximately 360 mi2 of Forest Service lands will be affected by implementation -- management unjts range in area from 2,372 to 49,000 acres. (More detailed descriptions of management units are on file at the Habitat Investigations office, Fort Collins, phOne 482-6575). �85 Table 1. USDA-Forest Service Managment areas scheduled for implementation of the Managing Forested Lands for Wildlife process. National Forest Ranger District Area Completion Date DWM Arapaho-Roosevelt Estes-Poudre Clear Creek Red Feather Panorama Ridge Evergreen N. t-tiddleMt. N. Bald 9/85 7/85 6/85 5/85 Hoover Werner Jackson Jackson GM,U,G Grand .l.Jncti on Cebolla Taylor River Middle Point Rambouillet Cr. Squirrel Cr. 6/85 6/87 6/87 S. Platte Salida S. Park Kenosha Pass Herring .):)nesHi11 5/86 2/87 10/87 Rio Grande Saguache Almosa Conejos N. Taylor canyon Archery Range Bighorn 2/86 2/87 2/88 Routt Hahn's Pk. N. Park Middle Park Coulton Cr. Green Ridge Troublesome 6/85 4/86 6/87 Middleton Porter Firth San .).Jan Dolores Mancos Twin Eagle Silver Creek Upper First Fork 6/86 6/87 6/88 McLain EllislReid carron Dillon Eagle Rifle N. Fk. Swan Mt. Ragged Lakes Deep Creek 10/85 10/86 10/87 Chappell Heicher Crane Pike-San lsable Pine White River Bock (?) P. Mason (?) Coghill R. Mason ? Jones (?) �86 •• At present, it appears most "front line" CDOW personnel may not have the expertise or equipment necessary to execute the computer programs of MFLW. However, they should now have enough knowledge of the fundamentals to---. influence species selection and habitat objectives, and to recommend modification of species' matrices when justified. These factors have the greatest influences upon the final design of the management unit. At present, the most critical need for CDOW is to adopt MFLW to CDOW's WANG system and train personnel in the execution of the prograffiS7This should result in further refinements to CDOW's input to USDA-FS management process to other (non-Forest Service) forested lands jn the state. Literature Cited Graul, W.o. and G.C. Mlller. 1985. Strengthening ecosystem management approaches. Wild. Soc. Bull. 12: 282-289. Hoover, R.L. and D.L. Wills eds. 1984. Managing forested lands for wildlife. Colo. Div. Wildl. in cooperation with USDA Forest Service, Rocky Mountain Region, Denver, 459 p. Prepared by_G-=.::~~r-::c.::--~r~nJkc-;..;;;....:~_ GaIJ C. Miller Habitat Investigations �87 APPENDICES �88 APPE~DIX A STATE OF COLORADO .U"" GowerftOr D£N.RTMENT OF NATU"-lL RESOURCES D. La"'''', DIVISION OF WILDLIFE ••••••••• R~. DtNCtor eoeo aroedway Deft •••.• CoIorMo 8021. (HT·11IZ) February 27, 1985 Dear This letter accompanies your copy of Managing Forested Lands for Wildlife, describing a new planning and management process to improve wildlife habitat. Tni s represents our finest opportunity yet to take positive action for the betterment of wildlife on a large scale. First, however, you must understand the philosophy, theories, and concepts which form the book's framework. I urge you to participate in one of the series of workshops t9 be given in the next few weeks. Gary Miller of the Habitat Resources Section, (phone 482-6575 or 667-0296), is your contact person for these workshop~. The next step is, of course, to put the process into practice. Each National Forest will implement the process on at least three management units over the next three years, beginning this spring. At least one of these management units, therefore, may well be in your District or Area--and almost certainly will be in your Region. To make a difference you need to get in on the ground floor--from management unit selection through the planning, implementation, and monitoring phases. Finally, I hope you share my excitement ~hat the Forest Service and Division of Wildlife now have the same "rule book" and can conmunicate in the same terms. And don't forget that, at least on a site-specific basis, the "rules" can be changed. If species-habitat relationships, or density estimates need to be refined, or a species other than one listed needs to be addressed, the process allows that to happen, but only if you get involved. The time is right, the opportunity is here--let's make a difference. Sincerely, ~~ .Jlm Ruch Director �89 •• APPENDIX B PROGRESS REPORT June - August 1985 Gary C. Mjller and Mjchelle D. Bowman Objectives 1. Apply the process described in Managing Forested Lands for Wildlife (Hoover and Wills, 1984) to Colorado Division of Wildlife (CDOW) properties within the lowland riparian ecosystem. a. As part of Step 1 in that process (Lipscomb et al., 1984), characterize understory shrub and grass-forb composition relative to aerial photo interpretation of habitat types on the Dodd Bridge, Cottonwood, Brush, and Berry easement properties (COOw). b. Also as part of Step 1, characterize avian species composition and use of 12 habitat types within the lowland riparian ecosystem on the same properties. c. Provide assistance and suggestions to operations personnel in performance of Steps 2-5 of the ecosystem management process. METHODS AND MATERIALS Habitat types of the Cottonwood and Dodd Bridge properties and surrounding lands (to at least 0.5 miles beyond property boundaries) were identified and mapped from aerial photographs (scale: 1:24,000), most of which were taken in 1979. Aerial photo interpretation was performed by the Colorado State Forest Service. The classification system was similar to that described by Buttery and Gillam (1984) except that canopy closure classes were slightly different, and four, rather than three, size classes of cottonwoods (Populus deltoides var. sargentii) were used. The classification system employed 8 habjtat types with a 9th, cottonwoods, further subdivided into 12 structural stages such that a total of 20 different habitat types theoretically were possible (Table 1). Areas of each habitat type were computed, and costs of photo interpretation were tracked. The accuracy of photo interpretation was calculated from on-ground verification data collected in July and August, with allowances made for the Six-year age of the photographs. Stands of the grassland and cottonwood habitat types were selected for sampling of grass-forb, shrub, and avian characteri~tics only when they were > 200 m from a public road and >100 m from a property boundary to minimize "edge effects" of human influences. Stands which were clearly definable on the ground tended to be selected for sampling over those which "graded" from one type to another. Although sampling of stands was roughly proportionally allocated, an attempt was made to obtain representative stands from all properties. In addition to the Dodd Bridge and Cottonwood areas, stands on the Brush property were sampled because of their "burned" (by COOW personnel in 1984), "unburned", and "plan-to-burn" status. Berry easement stands were sampled because of the high number of C2 stands. �· 90 Sampling within stands was based upon randomly-placed transects with random orientations which traversed the stand from boundary to boundary. Transects did not intersect and were variable in length. Sampling points were established at 50 m intervals along each transect. Each transect was sampled for grass-forb height, foliar cover, litter depth, five most common species, shrub foliar cover, and avian characteristics. TABLE 1 Habitat type and structural stage classification system for the lowland riparian ecosystem. ECOSYSTEM: HABITAT TYPE: LOWLAND RIPARIAN H - Mesic haymeadows and emergents G - Grasslands S - Shrublands R - River and other open water L - Lakes, ponds Ag - Agriculture Nv - Non-vegetated o - Developed C - Cottonwood STRUCTURAL STAGES: (for habitat type C) 1 1 - < 6" dbh Canopy Cover 2 - 6-16" dbh A - 10 - 35% 3 - >16-30" dbh B -) 35 - 55% 4 - ) 30" dbh C - > 55% UE - uneven aged or multi-storied stands 1 Not used in initial photo interpretation. UE habitat types may contain cottonwoods of different heights and diameters and/or be a C3 type with a well-developed understory of coyote willow (Salix exigua), green ash, (Fraxinus enns Ivanica var. subintegemna), russian olive (Elaeagnus angustjfolla , or tamarisk (Tamarix glauca) �91 Grass-forb height and litter depth were measured at each sampling point. Grass-forb cover was ascertained by a line-intercept technique (Canfield, 1941) with 10 m sections of the transect, centered on the sampling points serving as sampling units (at the edge of stands, sampling units extended from the sampling point to 10 m inside the stand). The five most common species were ocularly estimated along the entire transect. Shrub foliar cover (% coverage) was ascertained from strip transects of 4 m width, centered on the randomly-placed transects. The foliar diameter of shrubs within the strip was measured at heights of 0.5, 1.0, and 1.5 m; percent coverage at each height was then computed (under the assumption that shrubs were circular). Occurrence of avian species was recorded as ancillary data during all sampling activities. In addition, between 0600 and 1000 h, birds were censused at each sampling point. A 3-minute "quiet" period was immediately followed by a lO-minute census, during which bird species, sex, number, activity, and distance at which detected were recorded for all birds seen ~ithin the stand. RESULTS Habitat types of the Cottonwood and Dodd Bridge study areas were classified and mapped for areas as small as 1 ha (Fig. 1, 2). Forested portions of both areas were comprised mostly of mature (>16 in dbh) cottonwood stands (Tables 2, 3). Eighty-two percent of the total forested area was comprised of stands dominated by trees >16 in dbh. The Cottonwood area also contained a monderate amount of Old-growth (>30 in dbh) and pole-sized (6-16 in dbh) cottonwoods--habitat types which were rare in the Dodd Bridge area. The Dodd Bridge area contained no shrubland habitat type, and the Cottonwood lacked any dense-canopied ()55% cover) cottonwood habitat types in the mature and Old-growth classes. �\0 N 1. Vegetation map. ottonwood State Wildlife Area, rom 1979 aerial photograph. , • - V COTTONWOOD HAYMEADOW GRASSLAND SHRUBS AGRICULTURE DEVELOPED - NON-VEGETATED - LAKES - RIVERINE SIZE CLASSES I - undft 6· db" 2 - 6 - 16· dbh 3. 16- 30· db" 4 - ov~, 30· dbh CROWN A - 10 B - 35 C - 55 DENSITY _ 35 °/. - 55·1. - lOO·A. ~ �Fig. 2. Vegetation map, Dodd Bridge State Wildlife Area, from 1979 aerial photograph C - COTTONWOOD H - HAYMEAOOW GR S AG o - GRASSLAND - SHRUBS - AGRICULTURE - DEVELOPED L R - LAKE - RIVERINE NV - NON-VEGETATED SIZE CLASSES I 2 3 4 - undtr 6- dbh 6 - 16- dbh 16 - 30· dbh OYf, 30· dbh CROWN DENSITY A - 10 - 35-/. B - 35 - 55-.4 C - 55 - 100·.4 I.() VJ �• 94 TABLE 2 Composition of ecosystem management study area, Cottonwood SWA, Morgan County, Colorado Type No./stands CIA CIB CIC C2A C2B C2C C3A C3B C3C C4A C4B C4C 4 1 0 9 3 1 15 8 0 3 0 0 6 H GR 9 S 11 8 12 6 6 AG 0 NV L R TOTAL Area{ha~ 21.1 1.1 0 55.2 24.6 4.3 145.6 61.3 0 38.3 0 0 355.2 85.7 31.2 707.9 80.9 7.9 6.1 139.9 1766.3 x Area{ha~ 5.3 1.10 6.1 8.2 4.3 9.7 7.7 12.8 59.2 9.5 2.8 88.5 6.7 1.3 1.0 % or total 1.2 0.1 0 3.1 1.4 0.2 8.2 3.5 0 2.2 0 0 20.1 4.9 1.8 40.1 4.6 0.4 0.3 7.9 100.0 i oT roreste:::l 6.0 0.3 0 15.7 7.0 1.2 41.4 17.4 0 11.0 0 0 �95 .. TABLE 3 Composition of ecosystem management study area. Dodd Bridge SWA, Morgan County, Colorado . TYPE No./stands CIA CIB CIC C2A C2B C2C C3A C3B C3C C4A C4B C4C I H GR S Ag 0 NV L R TOTAL 0 0 2 2 1 17 14 4 1 3 1 21 13 0 11 20 11 8 Area(ha5 3.2 0 0 6.5 4.7 3.5 216.0 79.1 11.4 5.9 4.2 2.1 359.1 222.6 0 703.7 92.0 19.8 7.9 143.2 1884.9 x Area(ha) 3.2 3.3 2.4 3.5 12.7 5.6 2.8 5.9 1.4 2.1 17.1 17.1 64.0 4.6 1.8 1.0 % of total 0.2 0 0 0.3 0.2 0.2 11.5 4.2 0.6 0.3 0.2 0.1 19.1 11.8 0 37.3 4.9 1.1 0.4 7.6 100.0 % of forested 1.0 0 0 1.9 1.4 1.4 64.1 23.5 3.4 1.8 1.3 0.6 �• 96 • Overall mean area of cottonwood stands in both areas was 7.7 ha (18.9 ac), ranging up to 12.8 ha (Tables 2, 3), with standard error of the means of 3.6 ha. Mean area of haymeadow, grassland, and shrubland stands was 17.6 ha (43.4 ac), with a standard error of the means of 21.1 ha. e Overall accuracy of the photo interpretation, with no allowance made for the six-year age of the photos, was 84.6~ (n=26 stands inspected), with only cottonwood habitat types misclassified. When the age of the aerial photos was considered, the accuracy for 17 cottonwood stands was 94.1%, with the single error being one of canopy closure, not tree diameter. Cost of photo interpretation, excluding costs of flights, photography, and on-ground verification was $4.05/ha ($1.64/acre). Vegetation of all study areas was measured at 185 sample points and 31 transects, representing 23 cottonwood and haymeadow stands, between 15 June and 23 July, 1985 (Table 4). Mean cover values of the grass-forb strata ranged from 39.3% (C3A types) to 86.1~ (burned haymeadow, Brush SWA). The tallest grass-forb strata were encountered in the UE stands due to a prevalence of ~ardaria sp. ). The shortest grass-forb strata were in the C3A types. The burned stand on the Brush SWA was both denser and taller than the similar unburned stand. Mean litter depths (except for the burned stand, which had no litter) tended to be less than 10 em in cottonwood stands and greater than 10 cm in haymeadow stands (Appendix A'). Herbaceous plant species most frequently among the five most common species in all stands were: wheatgrass (Agropyron sp.), cheatgrass (Bromus tectorum), ~rdaria sp. switchgrass (Panicum Vlr atum), and prairie cordgrass (Spartlna pectinata). Alfalfa (Medicago sative was among the five most common species only on the burned stand at the Brush SWA. Mean shrub foliar cover at the 0.5 m height ranged from 2.l~ (haymeadow) to 30.0% (C2B stands), with no obvious relationship between tree canopy cover and shrub cover (Table 4). Shrub cover at the 1.0 m height occasionally exceeded 10% (in the denser canopied mature cottonwoods), but more commonly occupied less than 5% of the area--virtually no shrub cover reached 1.5 m height. Snowberry (SymphOricarpos sp.) was the dominant shrub. No shrub cover at the 1.0 and 1.5 m height was encountered on the burned stand at the Brush SWA in the first growing season after the burn. Rank correlations (Spearman, 1904 as cited in Snedecor and CoChran, 1967) between canopy closure and shrub cover, grass-forb height, and grass-forb cover revealed no significant trends at the 5~ significance level •. A weak positive correlation (r=0.72, n=7) was noted betwee~ closure and grass-forb height. A strong relationShip existed between shrub cover at the 0.5 and 1.0 m heights (r=0.7335, p<O.Ol). The amount (in percent) of shrub foliar cover at the 1.0 m height, for instance, may be estimated by the equation: Y= -.2271(x) + .4153 where Y=foliar cover (in percent) at 1.0 m height, and x=foliar cover (in percent) at the 0.5 m height. Censuses of avian species were conducted at 198 sample points of 31 transects representing 23 haymeadow and cottonwood stands (Table 5). A total of 31 species were encountered (Appendix B'). Stands with the greatest number of speCies per sample point (C2B) or individuals per sample point (C3B) held no species restricted to these types during the sampling period. Habitat type �TABLE 4 Vegetation characteristics of sampled lowland riparian stands, South Platte River, Colorado, 1985. Range of values shown in parentheses. N (ots.) 12 18 23 31 var. var. var. var. 10 10 10 25 30 16 14 16 \0 -...j �98 • • C3A held the greatest number of species (5) found only in 1 type--3 of those were raptors. A habitat type j~ntifie~ During avian sampling, which was notused during photo interpretatjon, was the uneven-age or multi-storied CUE) stand. Two species found only in the UE nanitat during censuses were great blue heron (Ardea herodias) and yellow-bdlled cuckoo (Coccyzus americanus). American goldfinChes (carduelis tristis) were found only jn haymeadows, hairy woodpeckers (Picoides vi110sus) were encountered only in mature, dense canopied (C3C) stands, and b1ack-headed ~rOSbeakS (Pheucticus melanocephalus) were found only in dense-canopied, pole-sjzed stands (C2C). House wrens (Troglodytes aedon) and eastern kingbirds (Tyrannus tyrannus) were found in all habitats sampled. Except for the haymeadows, detection distances generally approximated 40 m. . When miscellaneous observations were included with avian census data, a total of 36 species were encountered--blue grosbeaks (Guiraca caerulea) were seen only in haymeadows (burned and unburned) types, as were western kingbirds (Tyrannus verticallis) (unburned only). ~ild turkeys (Meleagris gallapavo) (hens wjth broods) were found only in UE types (Appendix C). one hen wlth a brood of 9 chicks was seen repeatedly on the Dodd Bridge area in a UE stand (and never in adjacent or nearby stands) from mid-June through August, and the same stand was used from late July through August by at least one, possibly two, other hens with broods--again, apparently to the exclusion of other stands. The only turkeys seen on the Cottonwood area were a hen with six poults, again in a UE stand. Swainson's hawks (Buteo swainsoni), seen only in type C3A during censuses, also used the UE type. TABLE 5 Avian characteristics of sampled lowland riparian stands, South Platte RJver, Colorado, 1985. Detection Stand x Code C2B 42.5 C2C 37.5 C3A 55.2 C38 39.8 C3C 39.1 43.7 UE H (no burn)88.6 H (burned) 69.0 Dist.(m) SO 34.3 66.2 44.1 41.4 48.8 50.1 69.8 41.6 N(l~s.) 18 30 37 25 30 30 16 ISPP 10 11 23 19 13 21 9 6 ~ I~. N Ilndiv N (0.83) 38 (3.17) (0.61) 23 (1.28) (0.77) 194 (6.47) (0.51) 264 (7.13) (0.52) 105 (4.20) (0.70) 187 (6.23) (0.30) 49 (1.63) (0.38) 43 (2.69) II Unigue 0 1 5 0 1 2 I 0 DISCUSSlDN In order to make knowledgeable, defensible habitat management decisions, a wildlife manager needs an accurate descrjptlon and inventory of the management uni t-Step 1 of the ecosystem management process (LipscOmb et a1 1984). Aerial photo interpretation appears to adequately fulfill much of that need for the lowland riparian ecosystem of the South Platte River. The cost of this exercise ($1.64/acre) was greater than would be incurred with large-scale application-the cost should decrease to $l.lO/acre (T. Owens, CSFS, pers. COI'llTl. ) • �99 The technique allows the inclusion of private land habitat characteristics in the process--CDOW lands may not need to be managed to provide all things to all species at all times, but may be used more efficiently, for example, to provide habitat types which are in shortest supply. One drawback to the • technique is the rapidity with which aerial photos become out-of-date. Photo interpretation does provide baseline information for a known period, however, and changes may then be tracked with a knowledge of cottonwood growth dynamics and periodic on-ground inspections, coupled with mapping and inventory updates. Crouch (1979) found cottonwoods of the South Platte increased approximately 4.5 inches in diameter in 17 years. In the current exercise, 20 different habitat types were possible according to the habitat classification scheme used for photo interpre- tation--one other category, the UE (uneven-aged or multi-storied) stand was not included initially, but could be included in future efforts. Stand delineation could be the same, based upon size and canopy closure of overstory trees, but the stand classification would indicate, by inclusion of a UE suffix, if understory trees comprised 10% of total stand area (i.e., C38/UE) (Table 1). In the current exercise, acreages and proportions of habitat types within CDOW lands were not computed separately. In the future, such computations should be made so ~he relationship of CDOW lands to the entire management unit will be evident It is probably desirable, from a planning and implementation standpoint, to give ~dentifying numbers or codes on the habitat maps for these stands on CDOW properties. Aerial photo interpretation does not adequately address one need of Step I--that of the wildlife species inventory--although it does provide a basis for an -expected species" list using CDOW's Latilong (Wildata) data bases. A moderate effort at on-ground avian survey and census yielded 31 (39%) of the approximately 80 species which would be expected in the study area during the survey period (Graul and Svoboda, Appendix D)--miscellaneous observations increased the species count by 5 (16%). A more detailed inventory of species' abundance and relative abundances (density estimates) mdght be made using either the point technique of Reynolds et ale (1980) or line transect techniques (Burnham et al., 1980), but the long, narrow configuration of many stands in the lowland riparian ecosystem will make the development of quantified species-habitat relationships very expensive (see Verner and Ritter, 1985). Step 2 of the ecosystem management process is the selection of a wildlife habitat goal--Lipscomb et ale (1984) suggested 3 alternatives: 1) species richness; 2) emphaSis species; or 3) .both emphasis species and species richness. Given the Colorado Division of Wildlife's statutory charges, management for both species richness and emphasis species is an appropriate goal (CRS 33-1-101, 33-2-102). Management for emphasis speCies, however, must not override the nUnimum requirements of indicators of speCies richness (ecological indicators) (Graul and Miller 1984). Step 3 of the ecosystem management process is selection of emphasis and indicator species. Examples of emphasis species on Division of Wildlife lands may be deer (Odocoileus sp.), ring-necked pheasants (Phasianus colchicus), �• 100 • northern bobwhite (Colinus virginianus), and wild turkey; a recent workshop (21 September 1985) of citizens interested in nonconsumptive wildlife values documented a public desire for emphasis upon lowland riparian species SUCh as great blue heron, eastern bluebird (Sialia sialis), yellow-billed cuckoo, yellow warbler, and upland sandpiper (Bartramia Iongicauda). Bald eagles (Haliaetus leucoceehalus) are an emphasis speCles candidate because of their endangered status In the conterminous united States. A listing of ecological indicator candidates overlaps somewhat with emphasis species--bald eagles, great blue herons, eastern bluebirds, and yellow-billed cuckoos were possible indicators identified by Graul and Svoboda (n.d., see Appendix 0), and Buttery and Gillam (1984). In addition, yellow-billed cuckoos, great blue herons, and wild turkeys were restricted to one stand type in the current exercise--an indication of possibly restrictive habitat affinities. For Division properties containing large expanses of mesic haymeadows (i.e., Tamarack, Cottonwood), upland sandpipers are appropriate ecological indicators because of their sensitivity to habitat fragmentation (Samson, 1980). Other ecological indicator candidates not generally considered emphasis species are wood duck (Aix sponsa), red bat (Lasiurus borealis), eastern cottontail (Sylvilagus floridanus), and spiny softshell turtle (Trionyx spiniferus) (Graul and Svoboda n.d., Appendix D). Galli et ale (1976) found area-sensitivity to forest fragmentation in the following species also noted during this exercise: blue jay (Cyanocitta cristata), brown thrasher (Toxostoma rufum), yellOW-billed CUCkoo, downy woodpecker (Picoides pubescens), hairy woodpecker, and black-capped chickadee (Parus atricapillus). Their analysis did not deal with stands within forests, and all were fairly consjstent in occurrence at forest sizes of 25 ac--conditions presently met on CDQW properties. It is of interest, however, that yellow-billed cuckoos and hairy woodpeckers were restricted to single stand types during the current exercise. In general, however, there is at present little reason to question Graul and Svoboda's thought (n.d., Appendix 0) that the species of the forested portions of the South Platte lowland riparian ecosystem generally are "flexible" (i.e., eurytopic), with broad regional and vegetative affinities, and should be able to exist in a variety of forested conditions. Step 4 of the ecosystem management process is to develop habitat Objectives which means, for this exercise, first providing the minimum requirements for ecological indicator species. These minimum requirements appear to be maintenance of forested areas (all types combined) in contiguous patches in excess of 25 acres, and mesic haymeadows either approximately 400 acres in area or clusters of smaller meadows, each at least.25 acres and totalling 400 acres in area within a radius of 15 miles (Samson 1980). Habitat types in shortest supply and which demonstrate little likelihood of increasing naturally (Cl, C2, S) should be maintained at least at current levels. A natural increase in type C4 may be expected, from current C3 stands, although attainment of the C4 stage may be dependent upon site capability as well as age of trees. Obviously, C3 types will undergo major declines in extent in the future. Habitat objectives for emphasis species can vary from property to property--there appears to be little problem (from an indicator species standpoint) in promoting habitat interspersion or "edge" within C3 forested stands or small «25 acre) haymeadows. Active management to create �101 heterogeneity in the grass-forb strata, promote understory development, and to create more habitat type S should benefit some, and almost certainly harm none, of the proposed ecological indicators. Approximately 75% of the areas currently forested on the study areas will be treeless (or monotypic Russian olive stands--see Knopf and Olson, 1984) by the year 2000 unless active management to establish trees begins soon. This prediction assumes a liberal estimate of cottonwood senescense--decade 8 or 9 (Fowell 1965), Crouch's (1979) belief that cottonwoods of the South Platte are 50 years old at 12-inch dbh and his finding of approximately 0.25 inch annual increment in mature trees, and assuming 15% natural cottonwood recruitment. This demonstrates the utmost importance of quickly prescribing and performing treatments (Step 5). Stands of cottonwoods or suitable alternate species need to be established on a large scale and continuing basis if an approximation of the current lowland riparian ecosystem is to be sustained. As Miller (1984, Frog. Rep. W-136-R) reported for heron colony areas, and Sedgwick and Knopf (unpubl.) found on the South Platte SWA the critical problem for maintenance of the cottonwood community is not regeneration (germination and seedling establishment)~ather, it is one of recruitment to the sapling and pole size-classes. Obviously, there is a need to discover the factor(s) which allow cottonwoods to recruit naturally •••small areas of the South Platte have recruited cottonwood stands within the past 20 years, and comparative studies of those areas with others which have not recruited cottonwoods could be enlightening. Experiments to enhance seedling survival to the sapling-pole stage should also be an avenue of research. There appears to be little justification, however, for not starting to take corrective actions using current knowledge while such research takes place--research may well reveal that enhancement of natural cottonwood recruitment is not practical. Techniques now exist for establishment of cottonwood stands--cuttings and rooting of l-year wood, pole-cuttings, and cuttings to promote suckering (of younger trees) (Read 1958, Fowells 1965, York 1983, Miller and Pope 1984). The potential of using plantings of tree species other than cottonwoods should be explored. �• 102 • RECOMMENDATIONS . Step 1. Cover maps of all significant CDOW lowland riparian properties (those which are to receive management treatments) should be produced using the classification system of Table 1. These cover maps should include: a) surrounding lands not under CDOW control within at least 1/4 mile of CDOW property boundaries; b) tabulations of acreages, number of stands, and percent of total area for each stand type for the entire area mapped and for CDOW property; c) a numbering system for stands on CDOW property; d) a UE suffix for forested stands which have at least 10% of their area in understory trees or tall (>4 m) shrubs. Step 2. Management of lowland riparian areas should be directed toward maintenance of species richness and management for emphasis species, in that order of priority. Step 3. a) Appropriate potential ecological indicators (for species richness) are: Bald eagle, great blue heron, eastern bluebird, upland sandpiper, yellow-billed cuckoo, hairy woodpecker, wild turkey, wood duck, red bat, spiny softshell turtle, and, perhaps, eastern cottontail and black-headed grosbeak. Some species accounts· not given in Hoover and Wills (1984) are provided in Appendix E. b) Appropriate potential emphasis species are: Bald eagle, great blue heron, eastern bluebird, upland sandpiper, yellow-billed cuckoo, wild turkey, northern bObwhite, ring-necked pheasant, deer, and yellow warbler. Step 4. a) Haoitat objectives should specify maintenance of at least 20% of the lowland riparian ecosystem in forest, structured to provide a sustained yield of 20 . habitat types, and arranged in forested patches of at least 25 acres in area. b) Habitat Objectives should specify maintenance of mesic haymeadows of 400 acres in area, or clusters of smaller haymeadows ()25 acres) which total 400 acres within a Is mile radius. c. Habitat Objectives should not call for major alterations to habitat types Cl, C2, C4, or S. Step 5. Establishment of replacement stands of trees should be given highest priority for habitat management in order to meet the objectives of 4a. a) Such establishment should occur in a manner to allow appropriate monitoring of planting survival and wildlife use--in conjunction with the organizational unit responsible for monitoring and evaluation. �103 b) Cottonwood stand establishment should take place in existing C3A, C3B, or H stands. Establishment with dormant stem cuttings of l-year old wood, pole plantings, cuttings to promote suckering, direct planting, and husbandry of natural regeneration are techniques which may be used. Details of the pole planting teChnlque are given in Appendix F. c) Research to ascertain factor(s) which enhance natural cottonwood recruitment should be undertaken in addition to, not in place of, stand establishment activities. Such research should rely largely on expertise of forest scientists or silviculturists, probably through contract. d) The possibility of developing stands of tree species other than cottonwoods should be explored--again with the assistance of silviculturists. These may include: elm (Ulmus sp.), walnut (Juglans sp.), pecan (~rya sp.), red mulberry (Morus rubra), bur oak (Quercus macrocarpa , and other oaks--all are associates of plains cottonwoods in some areas (Read 1958, Fowells 1965, USFWS 1981). e) Establish understory tree plantings in selected-trial areas with· cottonwood overstory to produce a UE stand configuration. Potential species should be controllable, preferably native, mast-producing, and may include: hackberry (Celtis occidentalis), hawthorn (Crataegus sp.), mountaln-ash (Serbus sp.), or crab apple (Malus sp.) Step 6. Future habitat acquisition proposals within the lowland riparian ecosystem should also evaluate inclusion of upland or xeric shrub habitat types of adjacent areas. Such upland areas may assume importance as refuges during periods of deep snow accumulations or flooding in the riverbottom areas. �• 104 LJterature Cited Burnham, K.P., D.R. Anderson, and J.L. Laake.1980. Estimation of density from line transect sampling of biological population - Wildl. Monogr. 72. 202 D. Buttery, R.F. and B.C. Gillam. 1984. Forested ecosystems. Pages 43-71 in R.L. Hoover and D.L Wills (eds.). Managing forested lands for wildlife.-COlo. Div. Wildl. in cooperation with USDA Forest Service, Rocky Mountain Region, Denver, Colorado. Canfield, R.H. 1941. Application of the line interception method in sampling range vegetation. I. For. 39(4): 388-394. Crouch, G.l. 1979. long-term changes in cottonwoods on a grazed and an ungrazed plains bottomland in northeastern Colorado. USDA Forest Service, Rocky Mountain Forest and Range Exp. Sta. Res. Note RM-379. 4p. Fowell, H.A. 1965. Plains cottonwood (Populus deltoides var. occidentalis Rydb.). Pages 519-522 in Silvics of forest trees of the United States. USDA Forest Service Handb. 271. Galli, A.E., C.F. leck, and R.T.T. Forman. 1976. Avian distribution patterns in forest islands of differenct sizes in central New Jersey. Auk 93(2): 356-364. Graul, w.o. and G.C. Miller. 1984. Strengthening ecosystem management approaches. Wildl. Soc. Bull. 12(3): 282-289. Hoover, R.l. and D.l. Wills eds. 1984. Managing forested lands for wildlife. Colo. Div. Wildl. in cooperation with USDA Forest Service, Rocky Mountain Region, Denver, Colorado. 459 p. Knopf, F.l. and T.E. Olson. 1984. Naturalization of russian-olive: Implications to Rocky Mountain wildlife. Wildl. Soc. Bull. 12(3): 289-298. Lipscomb, J.F., J.C. Capp, S.P. Mealey, and W.W. Sandfort. 1984. Establishing wildlife goals and objectives. Pages 305-321 in R.l. Hoover and D.L. Wills (eds.). Managing forested lands for wildlife.-COlo. Div. Wildl. in cooperation with USDA Forest Service, Rocky Mountain Region, Denver, Colorado. Read, R.A. 1958. Silvical characteristics of plains cottonwood. USDA Forest Service, Rocky Mountain Forest and Range Exp. Sta. Paper No. 33. 18 p. Reynolds, R.T., J.M. Scott, and R.A. Nussbaum. 1980. A variable circular-plot method for estimating bird numbers. Condor 82: 309-313. �105 Samson, F.B. 1980. Island biogeography and the conservation of prairie birds. Pp. 293-305 in C.L. Kucera (ed.), Proc. 7th N. Am. Prairie Conf., SW Mo. State UnTV. Springfield. Snedecor, G.W. and W.G. Cochran. 1967. Statistical Methods, 6th ed. Iowa State Univ. Press, Ames. 593 p. U.S. Fish and Wildlife Service, 1981. The Platte River ecology study: Special research report. U.S. Fish and Wildlife Service, Northern Prairie Wildl. Res. Center, Jamestown, N. Dakota, 187 p. Verner, J. and L.V. Ritter. 1985. A comparison of transects and point counts in oak-pine woodlands of california. Condor 87: 47-68. �•... 0 0- , APPOvrX A Litter Depths and 5 Most Common Herbaceous Species or Lowland Rlpa~ian Stands, South Platte River, June-August, 1985 It: Ol p, • ...• 0 •.•II STAm iE ) OC T WOOO H ilS THANSECT " "'"u Ol ...•• •••••• •••••• •• ••• •• loll loll Ol Ol .," p, ..•" •.." •..•• •• u '0 •••• ..." •..'" •• •• "''0 •• •• •.• 0 '••" •••••• JC u .....• ;J •• III VI •..'" .,•• ....• " ·~t ~. ...• ci. Ol ..•••..u 0 ..." sr '0 .," ;J -" ...• ..• ...•• •... JC o ., 0:: III " '0 "•• c: u ...•." •••• II) ...• ... " Ol )( P, ••• ;J ...•0 ow '" .......•" ...•" 0:: ow '"" < '0 ...•• •• ..•'" •• It: U C2Cl CJA1 C301 C3Ul C2C2 ll:l U::1 H1 1 1 1 2 1 1 2 1 ~0 ~ r." -------------------------------------------No Data COllected----------------------------------------------------------------X X X X 7.8 X 4.2 X X X X X X X X X X 9.0 X X X X X 6.0 X X X X X 2.5 X X X X X :n.0 X X X X 5.7 X CJB2 H2 H2 CJB' C,1\2 C3Cl C'C2 1 1 2 1 1 1 1 8.1 12•• n.5 2.9 1.2 5.9 5.8 ll:' ll:' C2CJ 1 2 1 10.5 8.5 5.8 X H4 H4 H5 H5 C'AJ C'CJ C3CII 1 2 1 2 1 1 19.0 14.5 0 0 0 14.2 X X It: JC ..I :::a u X X X X X X X X X X X 11.8 - '" lID X ..I X X X X , X X X X X 20 15 X X X X X X X X X X X X 14 )( X X X X )( X X X X X )( X X X X X X X 13 8 X lC X )( )( X X X 14 0 ja. X X X X X JC u X X X X X X X X X X X r. )( X X X o..u 7 6 )( X 6 4 , J 2 2 1 1 1 �107 APPENDIX B' Species Occurrence (denoted by X) by Stand Type, South Platte River, Colorado, 1985 Data From Line Transect Surveys Only SPECIES House Wren Northern Bobwhite Mourning Dove American Robin Conrnon Flicker Orchard Oriole Blue Jay Red-winged Blackbird Brown-headed Cowbird Brewer's Blackbird Black-capped Chickadee Hairy Woodpecker Common Yellowthroat Western Meadowlark American Kestrel Northern Oriole American Goldfinch Red-headed Woodpecker Starling Wood Duck Red-tailed Hawk Great-horned Owl Yellow-breasted Chat Yellow Warbler Downy Woodpecker Swainson's Hawk Brown Thrasher Black-headed Grosbeak Great Blue Heron Yellow-billed Cuckoo Eastern Kinebird STAND C3B C2B C2C C3A X X 0 X X 0 X X X X X X X X X X 0 0 0 X X 0 0 0 X X 0 0 0 X X X X X C3C UE Ho X X X X X X X X X X X X X X X X X X X X X X 0 0 0 0 0 0 0 0 0 0 X X X X X X X X 0 X 0 0 0 0 0 0 0 0 X X X X 0 0 0 0 0 0 0 0 0 0 0 0 X X X 0 0 0 X 0 X X X X X X X X X X X X 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 a 0 0 a a a 0 0 0 0 X 1 Suffix b=burned, o=unburned X 0 0 0 0 0 0 0 0 0 0 a X X X X X X X X X X X 0 0 0 0 X X a 0 0 a 0 a 0 0 0 0 0 0 0 0 0 0 0 '0 0 0 X X X X 0 0 X 0 X X X 0 X 0 X X 0 0 0 0 a X a a 0 O. X 0 0 0 0 0 X X X X Ho X X X X 0 0 0 0 0 0 0 0 0 0 0 0 1./ �• 108 APPENDIX C Avian Species Observations (denoted by X) by Stand Type; Census and Miscellaneous Observations Combined SPECIES C2B X House Wren Northern Bobwhite 0 X Mourning Dove American RObin X 0 CorrrnonFlicker Orchard Oriole 0 0 Blue Jay X Red-winged Blackbird X Brown-headed Cowbird Brewer's Blackbird X Black-capped Chickadee 0 Hairy Woodpecke~ 0 X Common Yellow throat X Western Meadowlark X American Kestrel 0 Northern Oriole 0 American Goldfinch Red-headed Woodpecker 0 0 Starling 0 Wood Duck 0 Red-tailed Hawk Great-homed Owl 0 0 Yellow-breasted Chat Yellow Warbler 0 0 Downy Woodpecker 0 Swainson's Hawk 0 Brown Thrasher 0 Black-headed Grosbeak 0 Great Blue Heron 0 Yellow-billed Cuckoo X Eastern Kingbird 0 Western Kingbird 0 Blue Grosbeak 0 Black-billed Magpie 0 Wild Turkey X Mallard C2C E3A X X X X X X X X X X X X X X X X X X X X X 0 0 0 0 X 0 0 0 X X X X 0 0 0 X X X X X X X X X X 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 X X X X X X X X X X 0 0 0 0 X 0 X X 0 0 0 0 X X 0 X 0 0 0 E3~ i1: X X X X X X X X X X X X 0 X X X X X X X X X 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Hb HC'~ X X X X 0 0 0 0 0 0 0 0 0 0 X X 0 0 0 0 0 0 0 0 0 X X X X X X 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 X 0 X X 0 X X 0 0 0 0 X X X X 0 X X X X X X 0 0 0 0 0 0 0 0 0 0 0 0 X X X X 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 X· X X X b 0 0 0 X 0 0 0 0 0 0 11 Suffix b-burned. o-unburned X 0 0 0 �109 APPE~DIX D CHARACTERISTICS OF TERRESTRIAL VERTEBRATES OF THE SOt'TH PLATTE RIVER SYSTEH WALTER D. GRAUL ~~~ PEGGY L. SVOBODA ABSTRACT.-Colorado, The South Platte River system's riparian zone in characterized 209 terrestrial by a cottonwood-willow vertebrate species. bred on the area had widespread breeders were peripheral species, The majority of the species that breeding distributions to the area. contained and 43.7% of the Among the peripheral breeding 53.8% had their main breeding range affinity to the east. Both breeding affinities. history association, and wintering The preceding residents exhibited flexible vegetative features are consistent with the vegetative of the area; large trees have become predominant the last 80 years •. The recent establishment contributed to the amount of hybridization on the area by constituting Mountains recorded in avian populations forest. Suggestions species richness include active management and careful selection Riparian of these trees may have a continuous wooded link between the Rocky and the eastern deciduous vertebrate of management areas, but they are especially for maintaining of cottonwoods indicator species. zones are recognized universally the 100th meridan only within as important wildlife important in the arid lands west of in the United States (Graul and Bissell, 1978: Great Plains Agric. Counc., 1979; Johnson and Jones, 1977; Thomas ~ al., 1979). In these regions lowland river and stream riparian zones �110 frequently are characterized (Ponulus spp.) and willow 1977). by the presence of cottonwood trees (Salix spp.) (Lindauer, 1978; Pase and Layser, In Colorado the cottonwood-willow association only comprises 0.2% of the area ~f the state, but 50% of the species of birds in the state have been recorded there (Bottorff, 1974). (1978) concluded that cottonwood-willow productive and highly diversified - In fact. Beidleman areas are part of the most ecosystem in the West. Because of the relative scarcity and high wildlife values of cottonwood-willow riparian areas it is essential that conservation and/or preservation plans for these areas be based upon a sound biological foundation. In this paper we ~ill describe some important features of the terrestrial cottonwood-willow vertebrate vertebrate species of the South Platte River system in Colorado. assemblage We will then propose that this is the product of relatively recent, major changes in the riparian vegetation. Finally, the implications of these findings will be discussed. STUDY AREA.-- The South Platte River originates in the Rocky Mountains, northeast in central Colorado. as a snowmelt stream Its 724 km course runs to where it joins the North Platte River to become the Platte River near North Platte, Nebraska (Williams, 1978). the South Platte River, and its associated drainages, Colorado to the Nebraska line. Our study area was from Brighton, This includes approximately 61% of the portion of the South Platte River proper that is characterized cottonwoods by and willows. The study area lies within the Great Plains province. Platte River meanders The South through a riparian zone which is about BOOm wide �111 in most places (Crouch. 1979). The riverbottom vegetation throughout ~he system is dominated by an overstory of cottonwood trees (Populus deltoides occidentalis) secondary with willow trees (Salix a~'gdaloides) signifi~ance. spp.), snowberry constitute Scattered patches of other willows (Symphoricarpos the shrub understory occidentalis). being of (Salix and rose (Rosa woodsii) (Crouch, 1978). The herbaceous vegeta- ~ion is dominated by salt grass (Distichlis stricta) and sand dropseed (Sporobolus cryptandrus) METHODS.-- (Lindauer. 1978). To ascertain the terrestrial vertebrate species presumed to be present on the study area, their study area status, -and their habitat affinities, we examined Colorado's Kingery and Graul, 1978; Langlois, most extensive vertebrate as of 1 January 1978. 1978). occurring of 10 latitude and longitude We evaluated data for the three (5,6,7) that encompassed the study area and compiled a reptiles, birds, and mammals that were reported as in the riparian zone. By definition, riparian for the ~hree latilong blocks studied meant areas characterized willow association. the The lati10ng system divides Colorado into 28 lines (Bissell and Graul, 1981). list of amphibians, These data represented records, excluding fish, available for Colorado blocks as defined by the intersections latilong blocks latilong data (Bissell, 1978; by the cottonwood- We excluded species listed as accidental ~han three records for a latilong block). We also eliminated (less species tied strictly to marsh habitat since this habitat was extremely restricted along the South Platte River system and was quite distinct from the deciduous vertebrate tree riparian complex. list was representative We assumed that the resulting of the study area since in the three �112 latilong blocks all drainages were part of the South Platte River drainage system. Thirteen species were added to the list based upon a more recent field investigation (Fitzgerald. 1978~. and scientific A list of the study area species with their status names. is provided in Appendix Overall distributions vertebrate conducted within the study area I. and habitat affinities species that bred or wintered for the study area on the study area were ascer- tained by reviewing the following general references: reptiles -- Collins (1974). Conant (1975). Ernst and Barbour Stebbins (1966); mammals -- Armstrong (1976). Hall and Kelson Ornithologists' (1979). Peterson Union and (1972). (1972). Burt and Grossenheider (1959). Lechleitner (1969); birds -- American (1957), Bailey and Niedrach ~1961), Robbins ~ amphibians (1965). Johnsgard a1. (1966), and Terres (1980). Species listed as definite or likely breeders in the latilong studies were classified as breeders; present in the appropriate although direct breeding RESULTS. reptile, to species breeding habitat during the breeding season, evidence for them was lacking. Species Composition bird, "likely" referred and Status.-- Of the 608 amphibian. and mammal species in the latilong publications, 264(43.4%) had been recorded Colorado. Our study area list (Appendix 1) included 209 (34.4%) of the 608 species. in Colorado in river and stream riparian zones in In comparison the cottonwood-willow study area, was unequaled to other major vegetative association, in vertebrate as exemplified species richness. based upon the latilong data the Colorado vegetative the second highest number of vertebrate associations by our For instance. association with species was pinyon (Pinus edulis)- �113 juniper (Juniperus spp.); it hosted 153 species. One hundred and nineteen (57%) of the 209 species bred on the study area and 73 (37%) were year-round . ., "7'":. \" \.: residents (Table 1). Birds constituted 87% of the specie~ that only used the area on a seasonal basis. ---- .,. .. Breeding Distribution Analvsis.-- species had breeding distributions specific vegetath'e affinity. that encompassed United (Table 2). conservative that went far beyond any regional or In fact; 72.3% of them had breeding distributions States 50% or more of the conterminous If anything, in illustrating the preceding the widespread ranges of the study area species. Snipe, The majority of the breeding throughout of the conterminous For instance, the Marsh Hawk, Common that encompassed much of Canada and Alaska. of Colorado containing species represented an assemblage in all directions, range affinity Vegetative the study area (Table 3). of species with main range affinities but the majority directly These peripheral (53.8%) had their main breeding to the east (Table 3). Affinities.-- None of the 209 vertebrate to the cottonwood-willow association species were throughout its range for any part of its annual cycle, but four appeared obligated association in Colorado during the breeding season. are the lJood Duck, red bat, eastern cottontail, softshell as to the study area since their breeding ranges ended in the quarter restricted less 48 United States, but all three bred Fifty-two (43.7%) of the study area breeders were classified peripheral is scope of the breeding and meadow vole had breeding distributions than one-half statistic 48 turtle. to this The four species and western spiny �• 114 In addition to the preceding four species, based upon an analysis of latilong data it appeared restricted breeding that 17 more of the 209 species were to the cottonwood-willow and/or w~ntering seasons. association For these species, however, the latilong habitat data were incomplete. Sparrow, White-crowned Cuckoo, Black-billed and Niedrach, fisher Cuckoo, American Goldfinch, associated Sparrow, Screech Owl (Bailey 1972) are known situations. 1965), Great Blue Heron pers. commun.), Double-crested (Armstrong, the White-throated deer (Armstrong, in other vegetative (Bailey and Niedrach, and mink Specifically, Sparrow, Eastern Bluebird, Cardinal, Yellow-billed 1965), and white-tailed to occur in Colorado in Colorado for the Cormorant The Belted King(Mark Kandel, (W. D. Graul, pers. obs.), 1972) occur in Colorado in aquatic situations with vegetation other than cottonwood and willow. Bell's Vireo, Winter Wren, and American Redstart Niedrach, 1965) utilize shrub communities cottonwood-willow DISCUSSION. association Vegetative The Fox (Bailey and beyond those found in the in Colorado. Histor\·.-- Williams based upon reviews of historical (1978) and Crouch (1979),' records, described the vegetative changes that have occurred on the South Platte River system since the early 1800's. They reported that in the 1800's trees were rare along the South Platte River, although on the Platte River cottonwood willow and trees were present on the islands and at least at scattered locations description along the main banks. of the potential vegetation system; he concluded association This corresponds with KUchler's that going westward (1964) of the overall Platte River the riparian deciduous ended on about the Colorado-Nebraska line. tree �115 In 1959 the South Platte River riparian system was dominated by a continuous overstory were apparently of mature cottonwood established 1959 and 1978, ho~ever, and willow trees; these trees by the early 1900's (Crouch, 1979). the density of the tree components South Platte River near Crook, Colorado declined reported a 50% decline in the cottonwood Between along the (Crouch, 1978); Crouch component between 1961 and 1978 on a plot grazed by cattle and a 30% decline during the same period on an ungrazed plot. The preceding activities. ~y vegetative Historically, changes likely are related to human tree growth along the Platte River system have been inhibited by fire (Kirsch and Kruse, cutting by Indians and early settlers bison levels (Bison bison) (Crouch, 1979). factors, Human settlement of the South Platte River area and Crouch (1979) noted that the appearance projects. the establishment in water could have altered the first three of the in the early 1900's also corresponded irrigation (Crouch, 1979), and/or grazing by (Bird, 196t), and/or extreme fluctuations in the late 1800's~bviously preceding 1973), and/or with the development of trees of major These projects would have improved conditions of trees by providing for a more stable water flow (Crouch, 1979). Crouch (1979) felt that more intensive water management factor contributing to the recent decline in tree densities study area with intensive grazing being a secondary (1978) has documented was the main factor. on his Williams the dramatic water flow changes along the South Platte River; the main channel width has been reduced 89% in the last 100 years. The relationship between declining cottonwood densities and �• 116 intensive water management may be widespread reported the same phenomenon Vertebrate since Ohmart !! a1. (1977) on the lower Colorado River. Species Characteristics.-- Regardless of the reasons, it is apparent that the South Platte River system has gone from a nearly treeless to a continuously forested condition within the last 80 years. Based upon our analysis of the South Platte River system's terrestrial vertebrates vegetative it appears that this assemblage changes. Specifically, is a result of these the majority of the species that bred on the area had broad breeding ~tributions. Additionally, although some of the breeding and wint~ring residents had a preference for riparian areas. they had rather flexible vegetative fact, it is noteworthy cottonwood-willow obligate that unlike many vegetative association to it on a widespread and flexible vegetative restricted that situations. to the cottonwood-willow in Colorado is consistent with a 'flexible species' theme. The restrictions vegetative the species that was affinities are the very characteristics Having four species apparently In Extensive breeding distributions would enable species to pioneer new vegetative association associations, contained no vertebrate basis. affinities. are a reflection of limited habitat rather than strict preferences of the species. The presence of each of the four species in eastern Colorado represents a western extension of a main range to the east. Outside Colorado the red bat and eastern cottontail occur in a variety of forested, non-riparian (Barbour and Davis, 1969; Schwartz and Schwartz, Colorado the cottonwood-willow streams, and reservoirs 1981). situations In eastern complex associated with lowland rivers, is the only substantial representative of this �117 forest element. variety The western spiny softshell turtle is found in a of aquatic situations throughout Ernst and Barbour, represent 1972). its range (Collins, 1974; In eastern Colorado, the major rivers the only significant aquatic areas with a continuous link to eastern aquatic areas. Thus, this turtle is selecting an aquatic habitat that happens to be adjacent to cottonwoods Finally, throughout situations its range the Wood Duck nests in a variety of with the essential elements being trees near water (Bellrose, 1976; McGilvrey, association 1968). In eastern Colorado, the cottonwood-willow provides the only substantial water complex. the habitats .. and willows. representation The four species, consequently, available of this tree- are merely utilizing to them in Colorado with the only restriction being that they are not occupying or crossing the Rocky Mountains • The development of the overall Platte River system as a wooded lane across the Great Plains may explain why the majority of the peripheral breeding species had their main range affinity to the east. cally, the Platte River system now structurally westward extension represents the forested Rocky Mountains distinct populations and the eastern deciduous forest. Yellow-shafted This between is so prevalent along the South Platte River. the following hybridization along the preceding (1. A· a direct link between may explain, at least in part, why hybridization For instance, a continuous, of the eastern deciduous forest in an aquatic context. The wooded Platte River system also represents connection Specifi- river: Red-shafted Flicker (C. ~. auratus) galbula) and Bullock's Oriole cases have been documented Flicker (£. ~. cafer) and (Short, 1965); Baltimore Oriole (1· A· bullockii) (Sibley and �118 Short, 1964); Indigo Bunting and Lazuli Bunting Rose-breasted Grosbeak and Black-headed races of the Rufous-sided arcticus) Towhee documented hybridization in Boulder, drainages Colorado; Grosbeak (I.~. (Sibley.and West, 1959). (Sibley and Short, 1959); (West, 1962); and two eT\·throphthalmus and Additionally, I. ~. Wheat (1981) has between the Blue Jay and the Steller's Jay Boulder is situated along one of the secondary of the South Platte River. Management.-- If the vertebrate River system is to be maintained, needed. species richness of the South Platte active management probably will be If the tree component declines on a long-term, widespread basis, there likely will be a substantial decline in species richness, since at least avian species diversity has been related to foliage height diversity necessary (~acArthur and MacArthur, 1961). to actively promote regeneration Finally, ecological their maximum species (Graul ~ Special attention densities or widespread indicator species restricted to the association, other criteria must be used to select indicator River system. regional of cottonwoods. since there are no vertebrate cottonwood-willow Thus, it may be al., 1976) for the South Platte should be given to species that attain in this vegetative basis. Regarding association the preceding either on a species the best species would be those with the relati~elv most restricted habitat requirements (Graul ~ al., 1976). We thank Glen L. Crouch, Rocky Mt. Forest & Range Exp. Station; David W. Crumpacker, Miller, Dept. of EPO BioI., Univ. of Colorado, Colo. Div. Wildlife, for critically reviewing and Cary the manuscript, and Robert D. Ohmart, Dept. of Zoology, Arizona State Univ., for c. �119 providing ~eneral comments on it. LITERATURE CITED ~~RI~~ O~~ITHOLOGISTS' UNIO~. 1957. Check-list of North American birds. 5th ed. Port City Press, Inc., Baltimore. 691p. ~~RIC~~ O~~ITH010GISTS' UNIO~. 1973. Thirty-second supplement to the American Ornithologists' Union check-list of North American birds. Auk, 90:411-419. A~RIC~~ O~~ITHOLOGISTS' UNIO~. American Ornithologists' 1976. Thirty-third supplement to the Union check-list of North American birds. Auk, 93:875-879. ~~STRONG, D.M. 1972. Distribution of mammals in Colorado. Univ. Kans. Mus. Nat. Hist. Monogr. 3. 41Sp. ~AILEY. A.M., ~~ R:J. NIEDRACH. 1965. Birds of Colorado. 2 vols. Denver Mus. Nat. Rist., Denver, Colo. 895p. ~ARBOUR, R.~ •• AND W.B. DAVIS. 1969. Bats of North America. Univ. Press Kent., Lexington, 286p. BEIDLEMAN, R.G. vertebrate 1978. The cottonwood-willow riparian ecosystem as a habitat, with particular reference to birds, p. 192-195. In: W.D. Graul and S.J. Bissell (tech. coords.). Lowland river and stream habitat in Colorado: a symposium. Univ. Northern Colorado, Greeley, 4-5 Oct. 1978. Colo. Ch. Wildl. Soc. and Colo. Audubon Coune. BELLROSE, F.C. 1976. Ducks, geese and swans of North America. Stackpole Books, Harrisburg, PA, 540p. �120 BIRD, R.O. 1961. relation Ecology of the aspen parkland of western Canada in to land use. Publ. 1066, Research Branch, Canada Dept. Agr., 155p. BISSELL, S.J. 1978. Colorado mammal distribution latilong study. Colo. Div. Wildl., Denver. 20p. BISSELL, S.J., AND W.D. GRAL~. wildlife BOTTORFF, distribution. R.L. 1974. 1981. The latilong system of mapping Wildl. Soc. Bull., 9(3):185-189. Cottonwood habitat for birds in Colorado. Am. Birds, 28:975-979. BURT, W.H., AND R.P. GROSSEh~EIDER. 1976. A field guide to the mammals. 3rd ed. Houghton Mifflin Co., Boston. 289p. COLLINS, J.T. 1974. Amphibians and reptiles in Kansas. Univ. Kans. Mus. Nat. Hist. Public Ed. Sere 1. 283p. COLLINS, J.T., J.E. HUREEY, J.t. KNIGHT, AND H.M. SMITH. 1978. Standard common and current scientific names for North American amphibians and reptiles. Soc. Study Amph. Rept. Misc. Publ. Herp. Cir., 7:1-36. CONANT, R. 1975. A field guide to reptiles and amphibians of eastern and central North America. Houghton Mifflin Co., Boston. 429p. CROUCH, G.L. 1978. bottomland Effects of protection from livestock grazing on a wildlife habitat in northeastern Colorado, p. 118-125. In: lv.D. Graul and S.J. Bissell (tech. coords.). Lowland river and stream habitat in Colorado: a symposium. Univ. of Northern Colorado, Greeley, 4-5 Oct. 1978. Colo. Ch. Wildl. Soc. and Colo. Audubon Counc. CROUCH, G.L. 1979. Changes in the vegetation complex of a cottonwood �121 ecosystem on the South Platte River, p. 19-22. In: Great Plains Agric. Counc. Riparian and wetland habitats of the Great Plains: proceedings of the 31st meeting. Colorado State Univ., Fort Collins, 18-21 June 1~79. Great Plains Agric. Counc. Publ. 91. £~~ST, C.H., ~,~ R.R. BARBOUR. Press Kentucky, FITZGERALD, J.P. 1978. 1972. Lexington. Turtles of the United States. Univ. 347p. Vertebrate associations in plant communities along the South Platte River in northeastern Colorado, p. 73-88. In: W.D. Graul and S.J. Bissell (tech. coords.). Lowland river and stream habitat in Colorado: a symposium. Univ. Northern Colorado, . Greeley, 4-5 Oct. 1978. Colo. Ch. Wildl. Soc. and Colo. Audubon Counc. GRAUL, W.D., AND S.J. BISSEll (TECH COORDS.). 1978. Lowland river and stream habitat in Colorado: a symposium. Univ. Northern Colorado, Greeley, Counc., 4-5 Oct. 1978. Colo. Ch. Wildl. Soc. and Colo. Audubon 195p. GRAUL, W.D., J. TORRES, Ah~ R. DEh~EY. approach 1976. A species-ecosystem for nongame programs. Wildl. Soc. Bull., 4:79-80. CREAT PLAINS AGRIC. COUNCIL. 1979. the Great Plains: proceedings Riparian and wetland habitats of of the 31st annual meeting. Colorado State Univ., Fort Collins, 18-21 June 1979. Great Plains Agric. Counc. Publ. 91., 88p. HALL, E.R., AND K.R. KELSON. 1959. The mammals of North America. 2 vols. The Ronald Press Co., New York. 1078p. JOHNSGARD, P.A. 1979. Birds of the Great Plains: breeding species and their distributions. Univ. Nebraska Press, Lincoln. 539p. �122 JO~SO~, R.R., ~\~ D.A. JONES. management 1977. Importance, preservation and of riparian habitat: a s~~posium. Tucson, Ariz., 9 July 1977. USDA For. Servo Gen. Tech. Rep. ~~-43. 217p. JONES, J.K., JR., ~.C. CARTER, ~\~ H.H. GENO~AYS. 1979. checklist of North American mammals north of Mexico, Revised 1979 Occa~. Papers Mus., Texas Tech Univ., 62:1-17. KINGERY, H.E., A~D ~.D. GRAVL. 1978. Colorado bird distribution latilong study. Colo. Field Ornith. and Colo. Div. Wildl., Denver. 58p. KIRSCH, L.M., A~ A.D. KRUSE. 1973. Prairie fires and ~ildlife. Proc. Tal: Timbers Fire Ecol. Conf., 12:289-303. KUCHLER, A.W. 1964. Potential natural vegetation United States: Am. Geogr.-Soc. LANGLOIS, D. 1978. Spec. Publ. 36. Colorado reptile and amphibian distribution study. Colo. Div. Wildl., Denver. LECHLEITNER, R.R. of the conterminous 1969. latilong 17p. Wild mammals of Colorado: their appearance, habits, distribution, and abundance. Pruett Publ. Co., Boulder, Colo. 254p. LINDAUER, I.E. 1978. A comparison of the vegetative communities of the South Platte and Arkansas River drainages in eastern Colorado, p. 56-72. In: W.D. Graul and S.J. Bissell (tech. coords.). Lowland river and stream habitat in Colorado: a symposium. Univ. Northern Colorado, Greeley, 4-5 Oct. 1978. Colo. Ch. Wildl. Soc. and Colo. Audubon MACARTHUR, Counc. R.H., AND J.W. MACARTHUR. Ecology, 42:594-598. 1961. On bird species diversity. �123 MCGIL\~EY, F.B. 1968. A guide to wood duck production habitat require- ments. U.S. Bur. Sport Fish. and Wildl., Resour. Publ. 60, 32p. OHMART, R.D., W.O. DEASO~, ~~ the Colorado.River, C. BURKE. 1977. A riparian case history: p. 35-47. In: R.R. Johnson and D.A. Jones (tech. coords.). Importance, preservation and management of riparian habitat: a symposium. Tucson, Ariz., 9 July 1977. USDA For. Servo Gen. Tech. Rep. RM-43. PASE, C.P., ~~i)E.F. LAYSER. 1977. Classification of riparian habitat in the Southwest, p. 5-9. 1£: R.R. Johnson and D.A. Jones (tech. coords.). habitat: Importance, preservation and management of riparian a symposium. Tucson, ~riz., 9 July 1977. USDA For. Servo Gen. Tech. Rep. ~~-43. PETERSO~, R.T. Mifflin ROBBINS, 196). A field ~uide to western birds. 2nd ed. Houghton Co., Boston. 309p. C.S., B. BRu~~, A~D H.S. ZIM. identification: 1966. A guide to field birds of North America. Golden Press, New York. 340p. SCHWARTZ, C.W., AND E.R. SCHWARTZ. Univ. Missouri SHORT~ L.L., JR. SIBLEY, C.G., ~~ The wild mammals of Missouri. Press and Missouri Dept. Conserv., Columbia, 356p. 1965. North America. 1981. Hybridization in the flickers (Colaptes) of Bull. Am. Mus. Nat. Rist., 129:309-428. L.L. SHORT, JR. 1959. Hybridization in the buntings (Passerina) of the Great Plains. Auk, 76:443-463. SIBLEY, C.G., AND L.L. SHORT, JR. 1964. Hybridization in the orioles of the Great Plains. Condor, 66:130-150. SIBLEY, C.G., k~i)D.A. WEST. 1959. Hybridization in the rufous-sided �• 124 towhees of the Gr~at Plains. Auk, 76:326-338. STE~BI~S, R.C. 1966. A field guide to western reptiles and amphibians. Houghton Mifflin Co., Boston. 279p. TERRE 5 , J.K. 1980. The Audubon Society encyclopedia of North American birds. Alfred A. Knopf, New York. 1109p. THO~~S, J.W., C. MASER, AND J.E. RODIEK. 1979. Wildlife habitats in managed rangelands -- the Great Basin of southeastern Oregon. USDA For. Servo Gen. Tech. Rep. Ph~'-80. 18p. l~ST, D.A. 1962. Hybridization in grosbeaks (Pheucticus) of the Great Plains. Auk, 79:399-424. WHEAT, P. 1981. Journal, ~~L1I~~S, G.P. Hybridization of the blue and Steller's jays. c.r.o. 15:9-23. 1978. Historical perspective of the Platte Rivers in Nebraska and' Colorado, p.11-41. In: W.D. Graul and S.J. Bissell (tech. coords.). Lowland river and stream habitat in Colorado: a symposium. Univ. Northern Colorado, Greeley, 4-5 Oct. 1978. Colo. Ch. Wildl. Soc. and Colo. Audubon Counc. Address of authors: Ft. Collins, Colorado Division of Wildlife, 317 W. Prospect, fQ 80526. �125 APPE~~IX I. -- Terrestrial vertebrate species of the South Platte studv area and their studv area status. Status codes are as follows: E c NQ;:breeding, !·-v;ar-round resident, !-breeder, l c likelv breeder, ~ • migrant, W - winter resident. Taxonomv references: herntiles - Collins !.l..!!.. (1978); birds - A.D.U. (1957,1973.1976); mar.cr.als - Jones !.l..!!.. (1979). Common Name Scientific Name Status AMPHIBIANS/REPTILES 1. tiger salamander Ambystoma tigrinum N 2. Woodhouse's Bufo woodhousei N 3. northern Acris crepitans N 4. striped chorus frog Pseudacris N 5. bullfrog Rana catesbeiana N 6. northern Rana pipiens N 7. snapping turtle Chelvdra serpentina R A. yellow mud turtle Kinosternon R 9. Chrvsemvs picta R painted toad cricket frog leopard frog turtle triseriata flavescens 10. western box turtle Terrapene ornata R 11. spiny softshell Trionyx spiniferus R 12. lesser earless lizard Holbrookia 1Ilaculata N 13. eastern fence lizard Sceloporus undulatus R 14. six-lined racerunner Cnemidophorus N 15. great plains skink sexlineatus .•.. Eumeces obsoletus 16. racer Coluber constrictor R 17. western hognose snake Heterodon nasicus N 18. milk snake Lampropeltis R 19. coachwhip Masticophis triangulum fla~ellum N R �·. 126 Common Name Scientific Name Status 20. northern water snake Nerodia sipedon R 2l. Pituophis melanoleucus N 22. plains blackhead snake Tantilla nigriceps N 23. plains garter snake Thamnonhis radix N 24. common garter snake Thamnonhis sirtalis R Phalacrocorax B gopher snake BIRDS 25. Double-crested Cormorant auritus 26. Great Blue Heron Ardea herodias R 27. Cattle Egret Bubulcus ibis B 28. Sno\ry Egret Egretta thula B Aix sponsa b,M Mergus merganser R 3l. Turkey Vulture Cathartes aura b,M 32. Goshawk ACcipiter gentilis W,M ACcipiter striatus R 34. Cooper's Hawk ACcipiter c:ooperii R 35. Red-tailed Hawk Buteo jamaicensis R 36. Broad-win~ed Hawk Buteo platvpterus M Buteo swainsoni B,M 38. Rough-legged Hawk Buteo lagopus W 39. Golden Eagle AQuila chrysaetos R 40. Haliaeetus W,M . 29. Wood Duck 30. Common Merganser 33. Sharp-shinned 37. Hawk Swainson's Hawk Bald Eagle leucocephalus 4l. Marsh Hawk Circus cyaneus R 42. Pandion haliaetus M Osprey �127 Common Name Scientific Name Status 43. Prairie Falcon Falco mexicanus R 44. Peregrine Falcon Falco pere2rinus M 45. Merlin Falco columbarius \~ ,!-1 46. American Kestrel Falco sparverius R 47. Bobwhite Colinus virginianus R 48. Ring-necked Pheasant Phasianus colchicus R 49. Common Snipe Capella gallinago R 50. Rock Dove Columba livia R 5l. Mourning Dove Zenaida macroura B,M COCCYZUS americanus B Coccvzus ervthropthalmus B,M 54. Barn Owl ~~ R 55. Screech Owl Otus asio ---- R Bubo virginianus R ~~ B,M 58. Saw-whet Owl Aegolius acadicus W 59. Poor-Will Phalaenoptilus M 60. Common Nighthawk Chordeiles minor B,M 61. Broad-tailed Hummingbird Selasphorus platvcercus M 62. Belted Kingfisher Megacervle R 63. Common Flicker Colaptes auratus R 64. Red-headed Woodpecker Melanerpes ervthrocephalus B,M 65. Lewis' Woodpecker Melanerpes lewis M 66. Yellow-bellied Sphyrapicus varius 52. Yellow-billed 53. Black-billed Cuckoo 56. Great-horned 57. Long-eared Cuckoo Owl Owl Sapsucker ~ nuttallii alcvon M �128 Status CotTr.'lon Na;!le Scientific Nar.'le 67. Hairy Woodpecker Picoides villosus R 68. Downy Woodpecker Picoides pubescens R 69. Eastern Kingbird Tvrannus tvrannus B 70. Western Kingbird Tvrannus verticalis B 7l. Great Crested Flycatcher Mviarchus crinitus M 72. Ash-throated Mviarchus M 73. Eastern Phoebe Savornis phoebe b,M 74. Least Flycatcher Er.'lpidonax minimus M 75. Western Flycatcher Empidonax difficilis M 76. Western Wood Pewee Contopus sordidulus B 77. Olive-sided Flycatcher ., Nuttallornis borealis M 78. Violet-green swallow 79. Tree Swa 11 0.••.. • • lridoprocne bicolor M 80. Rough-winged Stelgidoptervx B,M 81. Barn Swallow Hirundo rustica B,M 82. Cliff Swallow Petrochelidon B Flycatcher Swallow 83. Blue Jay cinerascens Tachvcineta thalassina ruficollis pvrrhonota M Cvanocitta cristata R Pica pica R 84. Black-billed 85. Common Raven Corvus corax W,}1 86. Common Crow Corvus brachvrhvnchos R 87. Pinon Jay Gvmnorhinus cvanocephalus M 88. Black-capped Parus atricapillus R Sitta carolinensis W,M Sitta canadensis M 89. White-breasted 90. Red-breasted Magpie Chickadee Nuthatch Nuthatch �129 Common Name Scientific Name 91. Brown Creeper Certhia familiaris R 92. House Wren Troglodytes aedon B 93. t\'inter~ren Troglodytes troglodytes W 94. Bewick's Wren Thryomanes bewickii M 95. Mockingbird Mimus polyglottos B 96. Gray Catbird Dumetella carolinensis B,M 97. Brown Thrasher Toxostoma rufUl!l B ,!-1 98. American Turdus migrator ius R 99. Hermit Thrush Catharus guttatus M 100. Swainson's Thrush Catharus ustulatus M 101. Veery 102. Eastern Bluebird Sialia sialis B,M 103. Townsend's Sol~tare Myadestes W 104. Blue-gray Gnatcatcher 105. Golden-cro~ed 106. Ruby-crowned 107. Robin , Catharus fuscescens Kinglet to~sendi Polioptila caerulea Status M M Regulus satrapa Regulus calendula M Water Pipit Anthus spinoletta M 108. Bohemian Waxwing Bombycilla garrulus W 109. Cedar Waxwing Bombvcilla cedrorum b,M 110. Northern Lanius excubitor W Ill. Loggerhead Lanius ludovicianus B 112. Starling Sturnus vulgaris R 113. Bell's vireo Vireo bellii B,M 114. Solitary Vireo Vireo solitarius M Kinglet Shrike Shrike �• 130 Common Nane Scientific Name 115. Red-eyed \'ireo Vireo olivaceus 116. Warbling Vire.o Vireo gilvus 117. Black-and-white 118. Tennessee 119. Orange-cro~~ed 120. Nashville 121. Virginia's 122. Northern 123. Warbler l~arbler Warbler Warbler Warbler Status Mniotil ta varia M Vermivora peregrina M Vermivora celata M Vermivora ruficapilla M Vermivora virginiae M Parula americana M Yellow Warbler Dendroica petechia B 124. Magnolia Dendroica magnolia M 125. Black-throated caerulescens M 126. Yellow-rumped coronata M 127. Black-throated nigrescens M 128. Townsend's Dendroica townsendi M 129. Chestnut-sided Dendroica pensylvanica M 130. Blackpoll Dendroica striata M 131. Ovenbird 132. Northern 133. MacGillivray's 134. Common Yellowthroat Geothlypis 135. Yellow-breasted Icteria virens B ,!-1 136. Wilson's Warbler Wilsonia pusilla 101 137. American Redstart Setophaga ruticilla b,M 138. House Sparrow Passer domesticus R Parula Warbler Blue Warbler Dendroica Warbler Dendroica Gray Warbler Dendroica Warbler Warbler Warbler Waterthrush Warbler Chat Seiurus aurocapillus M Seiurus noveboracensis M Oporornis b ,!-1 tolmiei trichas B,M �131 Common Name Scientific Nane 139. Western Sturnella neglecta R 140. Yellow-headed· Blackbird Xanthoce~halus M 141. Red-winged Agelaius phoeniceus M 142. Orchard Icterus spu-rius B,M 143. Northern Icterus galbula B,M 144. Rusty Blackbird Euphagus carolinus W 145. Brewer's Euphagus cvanoce~halus R 146. Common Grackle Quiscalus ouiscula B 147. Brown-headed Molothrus ater B 148. Western l'anager Piranga ludoviciana M 149. Scarlet l'anager Piranga olivacea M 150. Cardinal Cardinalis cardinalis b 151. Rose-breasted Pheucticus ludovicianus M 152. Black-headed Pheucticus melanoce~halus B,l-1 153. Blue Grosbeak Guiraca caerulea 154. Indigo Bunting Passerina cvanea B 155. Lazuli Passerina Amoena B,M 156. Evening 157. House Finch Carpodacus mexieanus R 158. Pine Siskin Carduelis pinus W,M 159. American Carduelis R 160. Lesser Goldfinch 161. Green-tailed 162. Rufous-sided Meadowlark Blackbird Oriole Oriole Blackbird Cowbird Crosbeak Grosbeak Bunting Grosbeak Goldfinch Hesperiphona Status xanthoce~halus vespertina tristis b,M M Ca-rduelis psaltria M l'owhee Pipilo chlorvrus M l'owhee Pipilo ervthrophthalmus b.M �• 132 Common Name Scientific Name Status 163. Dark-eyed Junco Junco hvemalis w 164. Gray-headed Junco Junco caniceps W 165. Tree Sparrow Spizella arborea W.M 166. Chipping Sparro~ Spizella passerina B.M 167. Clay-colored Sparrow Spizella pallida M 168. Brewer's Sparrow Spizella breweri M 169. Harris' Sparrow Zonotrichia guerula W 170. Zonotrichia leucophrvs W,M Zonotrichia albicollis W.M White-crow~ed 17!. ~nite-throated Sparrow Sparrow 172. Fox Sparrow Passerel1a iliaca W.1'1 173. Lincoln's Sparrow Melospiza lincolnii M 174. Swamp Sparrow Melospiza georgiana W,M 175. Song Spar rev Melospiza melodia R ;, ~rMALS 176. Virginia opposs~~ Didelphis virginiana R 177. eastern mole Scalopus aouaticus R 178. little brown bat Myotis lucifugus N 179. silver-haired bat Lasionvcteris noctivagans M 180. big brown bat Eptesicus fuscus W 18l. red bat Lasiurus borealis b 182. hoary bat Lasiurus cinereus M 183. eastern cottontail Svlvila~us floridanus R 184. rock squirrel Spermophilus variesatus R 185. fox squirrel Sc1urus nie;er R �133 CO!:l::lon Name Scientific Name Status 186. plains pocket gopher Geom,'s bursarius R 187. Ord's kangaroo rat Dipodomvs ordii R 188. beaver Castor canadensis R 189. plains harvest mouse Reithrodontomvs montanus R 190. western harvest mouse Reithrodontomvs megalotis R 191- deer mouse Peromvscus maniculatus R 192. northern grasshopper mouse Onvchomvs leucogaster R 193. eastern woodrat Neotoma floridana R 194. meadow vole Microtus pennsylvanicus R 195. prairie vole Microtus ochrogaster R 196. Norway rat Rattus norvegicus R 197. house mouse Mus musculus R 198. porcupine Erethizon dorsatum R 199. coyote Canis latrans R 200. red fox Vulpes vulpes R 201- raccoon Procvon lotor R 202. long-tailed weasel Mustela frenata R 203. mink Mustela vison R 204. eastern spotted skunk Spilogale putorrbs R 205. striped skunk Mephitis mephitis R 206. mountain lion Felis concolor R 207. bobcat Felis rufus R Odocoileus hemionus R Odocoileus vir~inianus R 208. mule deer 209. white-tailed deer �• 134 TABLE 1.- SUT!ll:lan' of th~ status .£[ ~ vertebrate species. South Platte studv area Herptile Bird t1a~al Total 11 33 29 73 Breeder or likely breeder2 0 45 1 46 Winter resident 0 22 1 23 Migrant only 0 51 2 53 Nonbreeding3 13 0 1 14 Total 24 151 34 209 Status Year-round resident1 1 Present year-round and breed- on the area. 2 Breed on the area, but only present part of the year. 3 Recorded during .the summer months, but does not breed in the cot t onvoc 0willow riparian zone. TABLE 2.-- Summarv of extent of breeding ranges of studv area species listed as breeders or likelv breeders. Percent values are in te~s of --the --area --of --the ~nterr;inous -48 ••••••• United States. ----;,;;;..;;;~ ..;;.,,;;~-=- Species 100% 50-99% < 49% Total 0 6 5 11 10 50 18 78 Mammal 3 17 10 30 Total 13 73 33 119 Herptile Bird �135 TABLE 3.-- Su~ma!'\'of stud'\'~ breeding species 'With breeding ranges endi~~ in the northeast ouarter of Colorado. Species Grou? Direction ~E E Total of Main Breeding Range SE 5 S~ W Herptile o 2 4 1 2 o o o 9 Bird 3 1 15 2 o o 3 2 26 Mamnal 1 2 9 1 1 1 2 o 17 Total 4 5 28 4 3 1 5 2 52 �136 APPENDIX E SPECIES ACCOUNTS YELLOW-BILLED CUCKOO (Coccyzus americanus) The Yellow-billed Cuckoo is an uncommon summer resident in eastern Colorado. The species arrives in early May and departs the area by mid-September. Unlike most other birds in the famjly Cuculidae, the North American species rarely practice brood parisitism. It is a secretive species, mainly inhabiting relatively dense undergrowth and stands of trees; extremely dense woods are avoided. Latilong summary - An unusual bird breeding in 12 latilong blocks, probable breeder in 9 others; restricted to lowland riparian for breeding, but also using riparian transition areas in migration. Ecosystem used - Although YellOw-billed Cuckoos are attracted to the lowlanj riparian by the dense shrub and brush, they are not restricted to this habitat type. Deserted farmland which has been reinvaded by dense shrubs and bushes, as 'well as brushy orchards, are also suitable habitats for the species' needs. Stands used - In the current study, this species was encountered only in the multl-storied stands. Rearing requirements - Nests are located in trees or bushes 2 to 20 feet above the ground. The nest is a fragile, flat structure of twigs, grass, and soft padding such as catkins of oaks and willows. The nest is most typically placed on a horizontal limb. Feeding requirements - Yellow-billed CuCkOOS feed exclusively on insects. Nearly two-thirds of the diet consists of caterpillars, especially hairy and bristly ones which are avoided by other birds. Cover reqUirements - Dense shrub and canopy cover will meet the complete cover requirements of this species. Bibliography Bailey, A.M. and R.J. Niedrach. Birds of Colorado. Denver Mus. Nat. ~st, 1965. Bent, A.C. Life Histories of North American CUckoos, Goatsuckers, Hummingbirds, and Their Allies. New York: Dover Pub. Inc., 1968. Chase, C.A. III, et ale Colorado Bird Distribution Latilong Study. ado Field Drnithol. and Colorado Div. of Wildlife, 1982. The Color- , Johnsgard, P.A. Birds of the Great Plains, Breeding Species and Their Distribution. Lincoln, Neb: Univ. of Nebraska Press, 1979. Martin, A.C., H.S. Aim, and A.L. Nelson. American Wildlife and Plants--A Guide to Wildlife Food Habits. New York: Dover Pub. Inc., 1951. Terres, J.K. York: The Audubon Society EnCYClOpedia of North American Birds. Alfred A. Knopf, 1980. Udvardy, M.D.F. The Audubon Society Field Guide to North American Birds, Western Regjon. New York: Alfred A. Knopf, 1977. New �137 ORCHARD ORIOLE (Icterus spurius) The Orchard Oriole is a fairly common breeding resident throughout eastern Colorado with various reports in the western portion of the state. The species is not territorial, sometimes colonizing with birds of the same or other species. Latilong summary - A rare breeder in 2 latilong blocks in the west; an uncommon to fairly common migrant in the lowland riparian, transitional riparian, urban, and agricultural habitats of the west; a fairly common breeder in 9 blocks, possible in 3 other blocks in the east, using lowland riparian and agricultural habitats. Ecosystems used - Orchard Orioles prefer open, sparsely wooded areas. River bottoms, shelterbelts, residential areas and orchards all may provide breeding habitat for the species. Grasslands and open country may even be used, provided suitable nesting sites are available. Stands used - In the current study, Orchard Orioles were encountered only in cottonwood stands with tree diameters 16" or multi-storied stands. Rearin re uirements - Nests are single or in colonies, often near eastern klngblrds yrannus tyrannus). The nest is a cup suspended from a forked end of a tree branch or in a bush from 6 to 20 feet above the ground. Feeding reguireme~ts - The Orchard Oriole feeds on insects and berries within the foliage of trees. caterpillars, bugs, grasshoppers, ants, beetles, anc spiders comprise the majority of the animal matter consumed, whereas cultivated fruits comprise the majority of the vegetative matter consumed. Cover reguirements - Dense shrubs will provide for the cover requirements of this species. Bibliography Bailey, A.M. and R.J. Niedrach. 1965. Birds of Colorado. Denver Mus. Nat. Hist, Bent, A.C. Life Histories of Blackbirds, Orioles, Tanagers, and Their Allies. New York: Dover Pub. Inc., 1968. Chase, C.A. III, et ale COlorado Bird Distribution Latilong Study. ado Field Ornithol. and Colorado Oiv. of Wildlife, 1982. The Color- Johnsgard, P.A. Birds of the Great Plains, Breeding Species and Their Distribution. Lincoln, Neb: Univ. of Nebraska Press, 1979. Martin, A.C., H.S. Aim, and A.L. Nelson. American Wildlife and Plants--A Guide to Wildlife Food Habits. New York: Dover Pub. Inc., 1951. Terres, J.K. York: The Audubon Society Encyclopedia of North American Birds. Alfred A. Knopf, 1980. Udvardy, M.O.F. The Audubon Society Field Guide to North American Birds, Western Region. New York: Alfred A. Knopf, 1977. New �• 138 .• BLACK-HEADED GROSBEAK (Pheucticus melanocephalus) The Black-headed Grosbeak is a common summer resident throughout Colorado below 8000 feet. It is a late mdgrant, not arriving until May. The species shares the same habitat as the Rose-breasted Grosbeak (unusual migrant in Colorado), and frequent hybridizations are encountered where the ranges overlap (primarily in Nebraska and South Dakota). Latilong summary - A fairly common bird breeding in 20 latilong blocks and posSlbly breedlng in 4 others, utilizing ponderosa pine, scrub oak, pinyonjuniper, aspen, and deciduous forest habitats for breeding while inhabitin~ urban and deciduous habitat types during migration. Ecosystems used - Black-headed Grosbeaks are associated with open stands of deciduouS forest with a well-developed understory. Riparian habitats are preferred, whereas orchards, brushy woodlands, or chaparral and residential areas are used secondarily. Stands used - In the current study, the species was encountered only in dense (55-100% canopy closure) cottonwood pole (6-16" dbh) stands. Rearing re~Uirements - The species nests from 4 to 25 feet above the ground in a treeork, tall shrub, or crown of sapling, usually near an opening. The structure is Shallow and bulky, composed of thin twigs, stems, and rootlets. Feeding requirements - The Black-headed Grosbeak feeds on wild and cultivated fruits and seedS, as well as depending largely upon a variety of insects. . Cover requirements - Open deciduous forests with a well-developed understory will meet the cover needs of this species. Bibliography Bailey, A.M. and R.J. Niedrach. Birds of Colorado. Denver Mus. Nat. Hist, 1965. Bent, A.C. Life Histories of North American cardinals, Grosbeaks, Buntings, Finches, Sparrows, and Their Allies (Vol. 1). New York: TOwhee DOver PG b. Inc., 1968. Chase, C.A. III, et ale Colorado Bird Distribution Latilong Study. ado Field Ornithol. and Colorado Div. of Wildlife, 1982 • .•.. The Color- Johnsgard, P.A. Birds of the Great Plains, Breeding SpeCies and Their Distribution. Lincoln, Neb: Univ. of Nebraska Press, 1979. Martin, A.C., H.S. Aim, and A.L. Nelson. American Wildlife and Plants--A Guide to Wildlife Food Habits. New York: Dover Pub. Inc., 1951. Terres, J.K. York: The Audubon Society Encyclopedia of North American Birds. Alfred A. Knopf, 1980. New �139 BROWN THRASHER (Toxostoma rufum) The Brown Thrasher is a fairly common summer resident and occasional winter resident throughout eastern Colorado. The species has also been reported as a common migrant and possible breeder in portions of western Colorado. It inhabits mainly rivers and streams, usually confined to dense shrubs and thickets due to its secretive nature. Latilong summary - A rare breeder in 2 latilong blocks and a possible breeder in 5 latilong blocks of the west; a fairly common breeder in 10 latilong blocks of the east, and a fairly common migrant using agricultural (orchards, shelterbelts, dwellings, and tree farms) and lowland riparian habitats; a rare winter resident in the lowland riparian and urban habitats. Ecosystems used - Brown Thrashers are associated with dense streamside thickets and agricultural areas. Stands used - In the current study, found only in stands of 35-100% canopy cover, or multi-storied stands. Rearing requirements - Brown Thrashers construct a bulky twig nest lined with rootlets placed on or above the ground in dense brush or thickets. Nest sites average 0.4 to 2.8 m from the ground in Kentucky. Nest site heights typically increase as the season progresses. Feeding requirements - The Brown Thrasher feeds on a variety of insects and berries obtained by scratching on the ground. Beetles are the preferred food of the species, with all other insects and berries eaten secondarily. Cover requirements - Dense shrubs and canopy will meet the cover needs of this species. Bibliography Bailey, A.M. and R.J. Niedrach. 1965. Birds of Colorado. Denver Mus. Nat. Hist, Bent, A.C. life Histories of North American Nuthatches, Wrens, ihrashers and Tneir Allies. New York: Dover Pub. Inc., 1968. Chase, C.A. III, et ale Colorado Bird Distribution Latilong Study. ado Field Ornithol. and Colorado Div. of Wildlife, 1982. The COlor- Martin, A.C., H.S. Aim, and A.L. Nelson. American ~ldlife and Plants--A Guide to Wildlife Food Habits. New York: Dover Pub. Inc., 1951. Terres, J.K. York: The Audubon Society EncyclOpedia of North American Birds. Alfred A. Knopf, 1980. New ~ �, 140 WOOD DUCK (~sponsa) Tne Wood Duck is an uncommon, although apparently beconting more numerous, resident of eastern Colorado. The species has expanded its range from the eastern deciduous forest habitat to the now-suitable habitat provided by the cottonwood riparian zones. Wood Ducks are associated with backwaters of streams, rivers and woodlands in areas open enough to allow flight. Latilon~ summary - A rare breeder in 2 latilong blocks and possible breeder in 4, utillzing river, lake, and riparian lowland habitats; a rare migrant in aquatic and riparian lowland habitats and a rare winter resident on lakes. Ecosystems used - The Wood DuCk frequents slow moving bodies of water with numerous large trees. Flooded deciduous forest with adequate mast production and water less than 18 inches deep is optimum for the Wood Duck. Stands used - In the current study, Wood Ducks were encountered only in open (10-35% canopy cover) stands of mature ( 16" dbh) cottonwoods. Rearing reruirements - Wood Ducks are cavity nesters in deciduous forests. They wi 1 utilize natural cavities or artificial nest structures. Optimum natural cavities occur in trees greater than 16 inches in diameter and have openings 3 1/2 inches wide, with interior cavities 8 inches in diameter.· Preferred nest sites are located above 30 feet from the ground, but have been found from 2 to 65 feet, in open stands w!th slow moving or standing water. Feeding reguirements - The Wood Duck requires an abundant supply of animal matter durlng the flrst 6 weeks of life, later utilizing both vegetative and animal matter. Adult birds eat nuts and fruits of woody plants, as well as a variety of cultivated and natural seeds. Cover reguirements - Open stands of flooded deciduous forest, as well as slow moving streams and rivers with mature deciduous riparian zones provide for the cover requirements of the Wood Duck. However, for brood rearing, the Wood Duck requires an understory of shrubs or low-lying trees within a flooded stand or adjacent to a stream or river for predator avoidance. Bibliography Bailey, A.M. and R.J. Niedrach. 1965. Birds of Colorado. Denver Mus. Nat. Hist, Be1lrose, F.C. Ducks, Geese, and Swans of North America, 3rd edition. Harrisburg, PA: Stackpole Books; WaShington, D.C.: Wildlife Management Institute, 1980. Chase, C.A. III, et ale Colorado Bird Distribution Latilong Study. ado Field Ornithol. and Colorado Div. of Wildlife, 1982. The Color- Johnsgard, P.A. Birds of the Great Plains, Breeding Species and Their Distribution. Lincoln, Neb: Univ. of Nebraska Press, 1979. Martin, A.C., H.S. Aim, and A.L. Nelson. American Wildlife and Plants--A Guide to Wildlife Food Habits. New York: Dover Pub. Inc., 1951. Terres, J.K. York: The Audubon Society Encyclopedia of North American Birds. Alfred A. Knopf, 1980. New �Al'P~DIX 141 !=' Wildlife Resource Notes Information Exchange Bulletin Vol. 2 No. Date: 31 December 1984 Environmental Impact Research Program CONTENTS ~ Effective Technique for Planting Trees in iivarian Habitats -R. Miller and G. E. Pope A Frame Nesting Platform for Ospreys -- R. A. Adair and W. A. Mitchell IiEPas a Wildlife Management Tool at John H. Xe%z k£ervoir -/ J. R. Fulton St. Louis District Holds Lake Managers Meeting -- A. P. Lookofsky and R. S. Wilkins Wetland Ecology Courses Newsletter Update -- C. O. Martin AN EFFECTIVE TECHNIQUE FOR PLANTING TREES IN RIPARIAN HABITATS In 1983 and 1984, personnel from the Army Corps of Engineers and the Colorado Division of Parks and Outdoor Recreation conducted a habitat improvement project by transplanting native trees along a creek near Trinidad Lake, Colorado. The project was undertaken in March and eaTly April. while the trees were still in winter dormancy. Species transp~ted were narrowleaf cottonwood (Populus angustifolia), plains cottonwood ~. sargentii), and coyote willow (Salix exigua). The trees were selected from areas of thick growth along the Purgatoire River. the major source of water for Trinidad Lake. Cottonwoods 4-8 in. in diameter were cut down, trimmed of their branches. and placed into barrels of water containing a rooting hormone; each cutting was from 6-8 ft long. Willow .aplings from 3 to 6 ft in height were alao cut and put in the aolution. True Dorth was notched on each tree ao that it could be oriented correctly when planted. . - The cuttings were taken to Longs Canyon. a licensed Colorado Wildlife Area. and planted along the edge of Lonls Creek. a ripari.n area that has ~ery little releneration of native trees. Bales 3-4 it ~p were drilled as close to the creek as po.sible ao that the tyees could be set directly into the water table. Axe acores were made on that portion ~f the tree that was to be placed below around; this allowed Iround water to easily penetrate the thick bark layer. Soil was tamped firmly around ~ bas&a Gl the transplants. and tar was painted on expoaed cut areas to retard disease and ina~ct infestation. New arowth aprouted from the cuttinls within 2 to 4 weeks, and by July some of " �142 the cottonwoods planted only 4 months earlier had new branches that were 2 ft or longer. By the summer of the year after planting, new trees, already 6-9 it tall, were firmly established. This technique, referred to as "dormant stock planting," has proven effective in protecting streambanks and is an excellent way to quickly establish riparian habitat in arid regions. Details are provided by John C. ""York in U.S. Department of Agriculture Soil Conservation Service Technical Note No. 22, dated June 1983. aobert Miller, Park Aid, Trinidad Lake, U.S. Army Engineer District, Albuquerque (now with the Colorado Division of Parks and Outdoor Recreation) Gregory E. Pope, Reservoir Manager, Trinidad Lake, U.S. Army Engineer District, Albuquerque. A FRAME NESTING PLATFORM FOR OSPREYS A variety of artificial platforms have been designed to facilitate nesting by ospreys (Pandion haliaetus). Most consist of a frame or solid base that can be mounted atop a tree or pole support. The platform described here is a 3- x 3-ft wooden frame mounted on a 25-ft wooden pole; it has been used .uccessfully by the U. S. Bureau of Reclamation to improve osprey nesting habitat in the western United States. Lumber used for the platform should be durable softwood, such as cedar, cypress, or redwood. If platforms are located in humid or marine environments, pressure-treated lumber ahould be used to prevent deterioration. In all cases, poles should be pressure-treated. Design specificat"ions for the frame nesting platform are shown in the accompanying figure (lacing page). The outer frame is built from 2- x 4-in. lumber and includes a 3-ft extension designed to aerve as a perch. The center supports are comprised of 2- x 6-in. boards that are notched and joined to form 4 cross-lap joints; the inside edges of the notches ahould be spaced S in. apart. The bottom of each center .upport ahould be beveled approximately 6 in. from the end to match up with the outer frame. and lalvan~zed metal straps can be nailed over the crou-lap joints to provide additional .upport. After the framework has been constructed. a 3- x 3-ft piece of galvanized welded vire fabric is stapled acrols the top of the platform. Hardwood dowels should be set vertically along the outer edge of the frame to help ospreys .ecure nest materials to the platform. The platform is completed by weaving sticks into the wire mesh; this will belp attract ospreys to the structure and facilitate nest construction. . Because of the size of the platform. it is more feasible to mount it on the pole at the installation site. The top of the pole .hould be trimmed to form 4 sides which fit anugly against the center .upports. Bolt holes for mounting the platform to the pole .hould be predrilled in the center .upports; these boles can then be used as luides for augering pilot holes in the pole. Although the drawing shows 3/8- x IO-in. bolts, lag bolts (l per .ide) are recommended for .aae of aa.embly. The completed assembly can be .et into a hole with a backhoe. The hole can be .xcavated with a power auger and .hould be at l.ast 6 ft deep. The pole must be .et into a dry hole because one .et into a wet hole will eventually lean, thus creating a safety hazard and possibly eliminating a nest �143 Vol. I, No.1 (10 June 1983) Introduction Corps Wildlife Manual -- C. O. Martin (WES) Raptor Management in NPD -- M. F. Passmore (USACE, Walla Walla District) Offset Rental Proves Beneficial to Wildlife Habitat in Tulsa District -L. M. Mason (USACE, Tulsa District) Arkabutla Boasts Bluebird Success -- K. Clossin and T. tiearn (USACE, Arkabutla Lake, Vicksburg District) Guilding Workshop Held -- L. J. O'Neil (WES) Training Brochure Available Vol. I, No.2 (20 September 1983) NPD-SPD Recreation-Resource Management Workshop -- O. D. Beckwith (USACE, North Pacific Division) NPD Wildlife Biologists Meeting -- E. P. Peloquin (USACE, North Pacific Division) Osprey Management at Lost Creek Lake, Oregon -- C. Pierson (USACE, Rogue River Basin Projects. Portland District) Stanislaus River Parks Resource Management Plan -- P. Crowley (USACE, Stanislaus River Parks, Sacramento District) Tensas Refuge Dedicated -- R. A. Haynes (USFWS, Vicksburg) and P. K. Elliott (USACE, Vicksburg District) Testing of Habitat Evaluation Methods -- L. J. O'Neil (WES) Wil~life Disease Workshops -- W. R. Davidson (Univ. Georgia) The Southeast Deer Study Group -- W. A. Mitchell (WES) Vol. I, No.3 (10 December 1983) Ospreys Nest at John H. Kerr Reservoir, Virginia -- J. R. Fulton (USACE. John H. Kerr Reservoir. Wilmington District) Successful Nesting of Bald Eagles at Robert S. Kerr Reservoir. Oklahoma L. D. Isley (USACE, Robert S. Kerr Reservoir, Tulsa District) Wynoochee Wildlife Mitigation -- R. M. Rawson (USACE, Seattle District) Pre-impoundment Environmental Studies of Aquilla Lake, Texas -- L. E. Marcy (Texas A&M Univ.), R. D. Slack (Texas A&M Univ.). and M. Hathorn (USACE. Fort Worth District) Snag Management Symposium -- J. W. Davis (USDA Forest Service, Phoenix). G. A. Goodwin (USDA Forest Service, Flagstaff), and R. A. Ockenfels (Arizona Game and Fish Department) DOD Biologists Hold Training Workshop -- G. G. Stout (Fort Sill. Oklahoma) Barn Owl Study Receives Army Award -- C. O. Martin (WES) Wetlands Functions and Values - A New Training Course Vol. 2, No.1 (30 March 1984) Record Eagle Count on the Arkansas River -- M. S. Johnson (USACE, Russellville Resident Office,' Little Rock District) Chief Joseph Wildlife Mitigation -- J. M. Habermehl (USACE, Chief Joseph Project, Seattle District) Another Successful Transport Season -- lLT J. V. Barker (USACE, Walla Walla District) Great Blue Beron Management at Rend Lake. Illinois -- R. Zoanetti (USACE, Rend Lake, St. Louis District) Visual Data Management in Walla Walla District -- M. F. Passmore (USACE, Walla Walla District) Development of Coastal Species Profiles -- E. J. Pullen (WES) .. �144 Vol. 2, No.2 (30 June 1984) Brush Structures for Wildlife -- C. O. Martin (WES) and J. L. Steele, Jr. (USACE, Ft. Worth District) Brush Piles and Bobwhites - A Case History -- J. L. Steele. Jr. Half-cut Trees and Limbs - A Cost-effective Management Tool -- J. L. Steele. Jr. and C. O. Martin An Elevated Brush Pile Design for Western Quail -- J. Koscuik (VSACE. Walla Walla District) and E. P. Peloquin (USACE. North Pacific Division) Christmas Tree Brush Piles -- L. E. Mettler (VSACE, Walla Walla District) Vol. 2. No.3 (30 September 1984) Canada ~oose Management at Smithville Lake. Missouri -- R. Lenning (U5ACE. Kansas City District) Timber Management. MeH. and Elk Browse Development -- J. R. Kosciuk (U5ACE. Dworshak Project. Walla Walla District) RMIS - A New Raptor Information System -- R. R. Olendorff (U.S. Bureau of Land Management. Sacramento) Habitat Development in the Tombigbee River -- J. C. Mallory (USACE. Mobile District) and A. Miller (WES) Cooperative Aquatic Habitat Studies in Texas -- G. Earls (U5ACE. Southwestern Division) and J. Killgore (WES) Plastic Signs - Inexpensive Resource Protection -- R. Lenning (V5ACE. Kansas City District) Vol. 2, No.4 (31 December 1984) Current issue ~3S3M ooes 'lIsn UWI\U:lcll::IO:l AJ.'WN3d SS3NISna '\f1~1:!::I0 s-!) ON J.IWI:I3d AWt:i\f 3Hl :10 lN3VIIlt:i\fd30 Ol\fd S33:1 Ii 3!)\fJ.SOd 3.1. \f k:I )I:..:_,__:_n__:_s:;:__ _ 0816£ IdcUSSISSIW'!)yneS)l~11\ lt9 XOS Od $Y33NI!)N3 so SdYO~ ·NOI.1.\fJ.SJ.N3WIY3dX3 SA\fMY3J.\fM AW~\f 3Hl :10 l.N3Wl~\fd30 � https://cpw.cvlcollections.org/files/original/47495523cf982d4202f24282d2b26172.pdf b1218b14957cae08cbf01e843f85e6ab PDF Text Text 1 Colorado Division of Wildlife Wildlife Research Report April 1986 JOB PROGRESS REPORT State __ ~C~o~l~o~r~ad~o~ Proj ect· No. _ Migratory Bird Investigations ------------------ Work Plan -Job -Job Tit 1e: __ _;_W.:_:a.:_:t.=e.:_rf.:_o:.:_w.:_l__:_P.:_ro.:_d::.:u::_:c.:_:t_;_i.=..o n;_;___:S.:_:u:.:_r_;_v.=,ey:!..,;s=-- _ Period Covered: 1 May through 30 June 1984 Author: Gerald M. Lorentzson Personnel: M. J. G. S. Bauman, G. Byrne, J. Corey, J. Creasy (Brown's Park NWR), Dennis, E. Dumph, J. Frothingham, J. Kauffeld (M.U. NWR), Lorentzson, D. Masden, W. Russell, G. Saville and Steinert. ABSTRACT Water conditions were good for waterfowl production early in the spring, but severe flooding occurred in late May and early June in the northwestern part of Colorado. The high mountain valleys showed an increase in the number of breeding pairs of ducks over 1983, but the river systems had a marked decline in numbers over 1983. Mallards continue to be the dominant species breeding in Colorado. Green-winged teal showed a noticeable increase in 1984. Canada goose populations continue to increase in all areas of the state. ��3 WATERFOWL PRODUCTIONSURVEYS Gerald Lorentzson P. N. OBJECTIVES 1. To estimate the number of duck breeding pairs, major waterfowl nesting areas in Colorado. 2. To estimate the obtain production 3. Compil e data and submit reports to appropri ate state personnel and the Fish and Wildlife Service for use in monitoring status of the various species and establishing hunting season recommendations. number of goose breeding data on selected goose by species, pairs, nesting on selected and in some cases, areas in Colorado. SEGMENTOBJECTIVES 1. To estimate the number of duck breeding pairs, by species, in the San Luis, Cache la Poudre, South Platte and Yampa Valleys, and in North Park and Brown's Park, using procedures presented in the Proggram Narrative. 2. To estimate the number of goose breeding pairs in the San Luis Valley, on the Yampa, Little Snake and Green rivers in northwest Colorado and in north-central and west-central Colorado using procedures presented in the Program Narrative. 3. Evaluate the present boundaries pairs in the San Luis Valley. 4. Alter the duck breeding pair survey in the Cache la Poudre and South Platte areas to compensate for sample sections which can no longer be counted because of changes in land use patterns. of the area sampled for duck breeding METHODSANDMATERIALS The 1984 breedi ng pai r surveys were conducted duri ng the peri od of May 1 through June 18, 1984. Duck surveys in the San Luis Valley, North Park, Poudre River drainage, and the South Platte River drainage were conducted from the air. Aerial counts in the San Luis Valley and North Park were adjusted for visibility by using a standardized visibility ratio used for 1984. Brown's Park surveys were done from the ground by personnel from the National Wildlife Refuge at Brown's Park. The Yampa Valley duck breeding pair counts were not done this year due to floods on all of the sample areas. �4 The method of sampling the San Luis Valley for duck breeding pairs was revised completely in 1984. Instead of using ground crews for establishing an air to ground visibility ratio, a standardized one was selected by using the most common ratios over the past 15 years. The number of square miles censused was reduced by 777 miles, leaving a total of 647 square mil es censused in 1984. The decrease in square mil es censused resu lted in a reduction of 209 linear miles censused by air. All of the total counts made on lakes in the San Luis Valley were retained in the new method of census i ng, but two, the Dry Lakes area and Smith Lakes, were added to the survey. There were 52 miles of the Rio Grande River added to the survey. New maps were prepared and di stri buted to the regi ons involved in the San Luis Valley survey. Several sets of maps are availab 1e in the Migratory Bi rd Section located in the Research Center in Fort Collins. Canada goose surveys were conducted withi n the May 1 through June 18th period. Estimates of the Colorado, White River, Yampa, Little Snake and Rio Grande river systems were obtained by direct counts from a fixed wing airplane. Production counts in the north-central Colorado area were done from the ground. Brown's Park National Wildlife Refuge personnel continue to do a ground count of Canada geese in their area. All f1yi ng was done with Cessna 185 ai rcraft. Two observers when flying transects, while one observer was used when sampling were used sections. RESULTS AND DISCUSSION Water conditions were good in most areas of Colorado, providing good breeding habitat for waterfowl early in the spring. Cool weather early provided a slow runoff, but a heavy snowpack and warm temperatures brought about flooding in the lowlands later on in the spring, especially along the Colorado and Yampa river systems in Northwestern Colorado. Water conditions in the San Luis Valley were the best since the late 1960's with the northern portion of the Valley having large areas of sheet water. Cool weather in early spring probably held nesting back in the South Park area of the state. The ice did not go off of the large reservoirs in South Park until late in April and early May. Goose banding in South Park in July indicated a very poor production year. The San Lui s Valley and North Park showed an increase of about 20% in breeding pairs from 1983 (Table 1), but in the case of the San Luis Valley, it was still below the long-term average. The South Platte Valley and Poudre Valley showed marked declines in numbers which caused the total number of breedi ng pairs to dec 1i ne 15.8% from 1983 and 9.2% from the long-term average for all of the areas surveyed. �5 Table 1. Summary of Colorado's Duck Breeding in Selected Areas, 1984 Area San Luis Valley North Park South Platte Valley Cache la Poudre Valley Yampa Valley Brown's Park Totals 1984 1983 15, 155 12,621 19,073 15,676 7,539 19,979 8,512 12,218 3,302 (*) 3,302 1,376 1,491 54,957 65,287 Pair Popualtion Long-term Average 25,495 17,699 8,550 4,980 2,649 1,146 60,519 (*) Count was not done in 1984 because of high water. Estimate Percent Change 1983 Long term ave. +20.1 -40.6 +21. 7 + 7.8 -62.3 -11.8 +70.9 -30.3 +20. 1 - 9.2 - 7.7 -15.8 The 1983 count was used in lieu of 1984. Mallards showed a slight increase in the species composition for 1984 and continue to be the dominant species breeding in Colorado (Table 2). Most species showed declines in numbers from 1983 except for the greenwinged teal and American wigeon which showed increases. Gadwal1s and blue-wi nged teal showed increases over the 1954-83 averages. The total breeding pairs in Colorado for 1984 was estimated at 54,957 pairs, which is lower than 1983 but close to the long-term average. Table 2. Species Composition Population S~ecies Mall ard B1ue-wt nged and Cinnamon Teal Gadwall Pintail Shoveler Green-winged Teal Redhead American Wigeon Common Mergansers Scaup Other Divers Wood Ducks Red-breasted Mergansers Totals of Colorado's Number of Breeding Pairs 1984 1983 22,093 19,666 6,595 9,662 8,647 3,807 2,980 5,048 2,686 1,605 312 2,521 1,045 31 14 54,957 1984 Duck Breeding 1954-83 Average 25,850 6,434 11 ,654 5,128 5,846 1,156 3, 187 1,480 297 6,069 3,569 3,657 3,192 3,216 1,178 4,648 122 14 65,287 2,239 55,404 Pair Percent Species Composition 1984 1983 1954~83 Av. 35.8 12.0 33.8 14.8 46.7 11.6 15.7 17.9 6.9 7.9 5.4 8.9 9.2 1.8 4.9 . 4.9 2.9 2.3 .6 .5 4.6 1.9 7. 1 tr. .2 tr 10.9 6.4 6.6 5.8 5.8 2.1 99.9 99.9 100. 1 4.0 �6 Table 3 shows estimated breeding pairs by species and areas. Flooding in 1984 in the Yampa Valley prevented censuses, so 1983 data were utilized. Table 3. Estimated Population of Duck breeding Pairs in Colorado, 1984 San Luis Valle~ Mallard 6,556 2,235 Gadwa 11 Pintail 843 Shoveler 622 Redhead 934 Blue-winged and Cinnamon Teal 3,625 Green-winged Teal 273 American Wigeon 23 Scaup 12 Wood Ducks 30 Other Divers Common Mergansers 2 Red-breasted Mergansers 15,155 Totals North Park Poudre River S.Platte Yampa River Valle~* 4,311 3,556 1,556 639 444 2,908 1,513 1,063 696 393 4,124 771 151 795 767 1.499 358 96 69 28 268 214 98 159 120 19,666 8,647 3,807 2,980 2,686 1,175 4, 111 1,259 1,778 485 182 182 575 30 424 61 405 226 701 206 83 69 204 50 58 51 6,595 5,048 1,605 2,521 31 1,045 312 14 8,512 7,539 244 19,073 36 1 198 65 Brown's Park 149 5 179 14 3,302 1,376 Total 54,957 * 1983 count ut ilized due to flooding. The number of geese showed a decrease in nesting pairs on the Yampa River but similar numbers on the Little Snake River in 1984 (Table 4). Total adults observed increased 253% in 1984 over 1983 on the Little Snake River and 126% from the 1981-83 average. Geese decreased in 1984 by 38% from 1983 and by 36% from the 1981-83 average on the Yampa River area, mostly from the lack of data from Lilly Park. Tab 1e 4. Number of Geese Observed On Aeri a1 Surveys in Moffat and Routt Counties. Nesting PCLirs. 1984 Yampa River Steamboat Spgs-Craig Craig-Juniper Springs Juniper Canyon Juniper Canyon-Lily Park Lily Park Totals Little Snake River Cross Mt.-Powderwash Powderwash-Baggs Totals 15 9 1 9 Non-nesting Pairs I9EJ 81-83 Av 1984 15 28 o 1983 11 1 3 3 o 40 66 171 75 66 276 150 150 1300 17 68 85 34 62 133 14 2 8 64 60 41 93 19 14 22 10 32 16 17 43 43 86 26 25 51 26 20 46 33 4 97 38 81-83 Av 1 19 29 37 65 1983 10 7 23 34 33 Total Adults 54 51 7 90 67 269 9 19 2 22 18 16 62 19 I 8i-83 Av 1984 �7 Total nesting pairs of geese and total adults and estimated goslings decreased somewhat in 1984 in Brown's Park National Wildlife Refuge, but still show numbers similar to 1981-83 averages (Table 5). Table 5. Number of Canada Geese Observed on Ground Counts in Brown's Park National Wildlife Refuge Area Brown's Park 1984 110 Nesting Pairs Total Adults Est.No.Goslings 1983 81-83 Av 1984 1983 81-83 Av 1984 1983 81-83 Av 140 96 325 407 330 296 305 281 The west central population of Canada geese increased, both in singles observed and pairs in 1984 (Tables 6 and 7). The North Fork of the Gunnison and that part off the Gunn ison from Delta to its confl uence were counted in 1984. Table 6. West Central Colorado Canada Goose Breeding Pair Survey, 1984 Area White River Meeker-Rio Blanco Lake Rio Blanco Lake-Rangely Rangely-Utah line Sub-total Colorado River Canyon Creek-Silt SiIt-Rifle Rifle-Parachute Parachute-DeBeque DeBeque-Palisade Palisade-Grand Avenue Bridge Grand Avenue Bridge-Fruita Fruita-Horsethief Canyon Horsethief Canyon-Utah Sub-total Roaring Fork River El Jebel-Carbondale Carbondale-Glenwood Springs Sub-total Gunnison River and North Fork of Gunnison River Hotchkiss-Delta Delta-Mesa County line Mesa County line-Whitewater Whitewater-Grand Junction Sub-total Grand Total Singles Pairs GrouEs 5 14 2 21 6 14 5 25 13 6 5 24 3 25 18 6 2 3 0 4 0 61 3 41 21 16 10 7 1 11 0 110 14 31 3 11 22 14 0 39 0 134 8 1 9 11 3 14 3 4 7 6 3 3 1 13 104 4 9 9 1 23 172 14 0 0 0 14 179 �8 Table 7. Comparison of the Number of Canada Goose Singles and Pairs Observed in Aerial Surveys in West Central Colorado, Spring 1978-84. Area White River Roaring Fork River Colorado River G 1enwood- 5th St. Bridge Bridge-Utah line Gunnison River Hotchkiss-Delta Delta-Grand Junction Totals 1984 21 9 Number Observed Singles 1984 1983 78-83 Av 10 32 25 14 2 3 57 4 36 3 40 5 98 12 51 tr. 3 83 4 19 172 6 7 104 Pairs 1983 78-83 Av. 21 45 3 5 81 14 84 11 119 2 5 152 The results of San Luis Valley goose breeding survey showed about a constant number of nesting pairs over the years 1982-84. It appears that there might have been some distortion in the non-nesting pairs and grouped birds reported in 1983 (Table 8). The numbers do not seem to fit into the overall picture of the breeding pair population in the San Luis Valley. Table 8. Results of the San Luis Valley Canada Goose Breeding Pair Survey Year 1975 Helicopter 1976 Helicopter Fixed Wing* 1977 Helicopter Fixed Wing 1978 No counts 1979 Fixed Wing 1980 Fixed Wing 1981 Fixed Wing 1982 Fixed Wing 1983 Fixed Wing 1984 Fixed Wing Nesting Pairs *Fixed Wing count incomplete. Projected Total Number Non-nesting Pairs Grou~ed Birds 59 59 110 92 77 63 69 64 30 100 100 101 91 130 97 99 96 58 141 226 154 90 159 189 244 376 169 227 811 60 242 489 154 �9 The Northcentral Colorado goose results are shown in Table 9. Table 9. census was done on June 18, 1984. The Results of Northcentral Colorado goose census, June 18, 1984. Production Area Wellington Fort Collins Water Area Terry Lake Water Supply & Storage #4 Deines Pond Launer Pond Douglas Lake Stewart Pond Rocky Ridge Dry Creek Reservoir N. Poudre #3 N. Poudre #2 N. Poudre #5 Bureau of Standards Reservoir #8 Elder Reservoir #8 Annex Van Sant Pond" Cobb Lake Dale Pond Watson Lake Curtis Lake Beghto1 Lake Sub-total Peterson Ponds Dixon Reservoir Miller Ponds College Lake Dean Acres Claymore Sterling Gravel Pits Larimer Co. Ponds Lindenmeier Lak~ Grey Lake Novak's Pond Flatiron Gravel Pits Anderson's Pond Parkwood Kitchel Timnath Romi1y Gravel Pits Fossi1 Creek Horseshoe Wyatt's Pond Andrijeski Marsh Sub-total Total No. Goslings 13 Total No. Adults and Yearlings 76 Total Birds 89 o o 24 24 16 15 31 15 24 17 38 13 28 30 6 o o 23 9 16 18 o 9 24 17 15 4 12 12 o 4 5 1 9 12 16 15 44 17 17 49 31 41 320 23 133 61 61 47 56 364 40 150 61 o o 3 5 229 37 22 877 27 73 5 9 52 12 11 30 358 5 4 22 8 9 43 12 29 93 o 32 o 15 11 137 124 2 63 89 9 45 49 64 46 126 39 65 13 o 7 1 441 1,357 1 8 1,106 64 95 14 410 23 148 154 7 67 111 17 54 92 76 75 219 39 97 13 22 1 1,798 �10 Table 9 (continued) Total No. Goslings Production Area Water Area Loveland 0 Flatiron Reservoir Boedecker 53 Flatiron Gravel Pits 24 Kauffman Gravel Pits 14 Big Thompson River 0 31 McNeil Reservoir Welch Reservoir 60 Reservoir No. 12 12 194 Sub-total Ish Lake 6 0 Crystal Lake Terry Lake 55 34 Faivre Ponds Rest Home Ponds, Sawhill 53 Ponds & Walden Ponds 36 Valmont Reservoir Boulder Valley Farm 6 Angus Ranch Pond 3 193 Sub-total 14 Grant Pond Kettring 35 Centennial 0 Columbine C.C. 29 Chatfield Golf Course (Wohlhurst) 14 11 Bowles Lake King's Pond 29 Tule Lakes 21 Marston Reservoir 29 Pinehurst C.C. 3 24 Clarefield Reservoir Kendrick Lake 0 Sloan's Lake 56 Denver City Park 57 Colo. Blvd. & Quincy 0 0 Blackmer Reservoir Sub-total 322 GRAND TOTAL 1,379 Boulder Denver Total No. Adults and Yearlings Total Birds 0 119 37 11 14 193 119 100 593 16 15 62 38 82 0 172 61 25 14 224 179 112 787 22 15 117 72 135 102 7 2 324 4 38 52 68 138 13 5 517 18 73 52 97 176 158 142 64 64 2 21 8 385 200 20 26 1,428 4,579 190 169 171 85 93 5 45 8 441 257 20 26 1,750 5,958 The number of adu lt Canada geese in the northcentral area increased 2.6% from 1983 and showed a 27.1% increase from the long-term average (Table 10) . �11 Tab 1e 10. Number of Adult Canada Geese Observed in Northeastern Colorado Production Trend Areas, 1984. Area Wellington Fort Collins Loveland Boulder Denver Totals 1984 877 1,357 593 324 1,428 4,579 No. of Adults Av. 1970-83 1983 945 758 1,298 789 624 301 251 500 1,347 1,254 4,465 3,602 Percent ChanBe 1983 from i97 -83 +15.7 -7.2 +4.5 +72 .0 +97.0 -5.0 +29.1 -35.2 +6.0 +13.9 +2.6 +27. 1 The number of gos1ings produced in northcentral Co lorado also showed an increase of 19.3% from 1983 and an increase of 16.4% from the 1970-83 averages (Table 11). The only area which showed a substantial decrease in the number of goslings produced was in the Wellington area. Table 11. Number of.Canada Goose Goslings Produced in Northcentral Colorado Production Trend Areas, 1984 Area Wellington Fort Collins Loveland Boulder Denver Totals No. of Goslings 1983 Av. 1970-83 325 ·262 245 320 198 123 152 200 236 280 1,156 1,185 1984 229 441 194 193 322 1,379 . !- Prepared by ()trOJ.& A0t--litJ:zs;;)·'·· --~G~e~r~a1~d~L~or~e~n~t~z~so~n~~~--------Senior Wildlife Biologist Percent Change 1983 from 1970-83 -29.5 -12.6 +80.0 +37.8 - 2.0 +57.7 +27.0 - 3.5 +36.4 +15.0 +19.3 +16.4 ��Colorado Division of Wildlife Wildlife Research Report April 1986 l3 JOB PROGRESS REPORT State Colorado Project 01-00-045 (W-88-R) Job Avian Research 14 Work Plan 1 Job Title: Ecological Studies of the Flightless Period of Ducks in Colorado Period Covered: 1 January through 31 December 1984 Author: M. Szymczak and J. Ringelman Personnel: J. F. Corey, J. K. Ringelman, S. F. Steinert and M. R. Szymczak, Colorado Division of Wildlife. ABSTRACT Fewer total ducks but a similar number of gadwalls (Arrasstrepera) used Walden Reservoir during the flightless period in 1984 compared to 1983. The location of the major concentration of flightless gadwall on Walden Reservoir was different in 1984 than in 1983. Periodic counts of flightless ducks on selected wetlands indicated numerical and species composition changes between years. Pre-molt mallards (Arras platyrhynchos) (4) in North Park moved to wetlands from 1 to 28 km away from the capture area prior to primary molt. Mallards molted in wetlands having dense patches of Carex nebraskansis or Typha sp , During the premolt stage adult females were significantly heavier and slightly larger than first-year females but there was no difference in condition. Pre-molt female weights and condition did not vary according to wetland or time period. Weight and condition of pre-molt males did not vary by age or wetland. Weights and condition of all mallards declined during the flightless period. Flightless gadwall became progressively heavier during the flightless period in 1984 than in the previous 2 years. However, the same general trends in weight dynamics were noted. ��15 ECOLOGICAL STUDIESOF THEFLIGHTLESSPERIOD OF DUCKSIN COLORADO Michael R. Szymczak James K. Ringe1man P. N. OBJECTIVES 1. Document the species and sex composition and seasonal molting on selected wetlands in North Park. 2. Identify the physical molting ducks. 3. Investigate ducks. 4. Determine ducks. 5. Investigate differences in duration of the flightless species of ducks in relation to sex, body condition, time of molt. 6. Determine condition, 7. Identify spatial the and biological and temporal behavioral time characteristics differences budgets and net duck molting wetlands energy in of ducks of wetlands in use of wetlands the survival rate of molting ducks habitat quality, and time of molt. and quantify abundance balance used by by molting of molting period of selected habitat quality, and relation to sex, body in Colorado. SEGMENT OBJECTIVES 1. Document the species and sex composition and seasonal molting on selected wetlands in North Park. 2. Investigate ducks. 3. Determine ducks. spatial the and temporal behavioral time differences budgets abundance in use of wetlands and net energy balance of ducks by molting of molting METHODS Species Composition and Seasonal Abundance Counts of adult ducks were conducted at weekly intervals at Walden Reservoir beginning on 13 July and ending on 31 August 1984. All counts began shortly after sunrise and were completed by 1037 hrs , Ducks were classified as to molt status and generally only those birds in duck concentration areas were included. When possible the sex of each bird was recorded. Counts were terminated when it became difficult to distinguish young from adults at long distances. �16 Adult ducks on 76 Pond, Elk Pond, Pole Mountain Reservoir, Hebron Pond, and Alkali Pond were counted periodically and molt status of each adult observed was recorded. Pole Hountain was counted on 31 July and 16 August 1984. Alkali Pond was counted 4 times (7/10, 7/26, 8/14, 8/30), while 76, Elk, and Hebron ponds were counted on 27 July and 17 and 30 August. Molt Site Selection Back-pack radio transmitters (Dwyer 1972) were attached to 1 male and 10 female mallards and 2 female gadwalls prior to their wing feather molt. The first radio was attached on 12 July and birds were monitored through 4 October 1984. One mallard was captured on the nest (Coulter 1958), 2 females in decoy traps (Anderson et a!. 1980), and the others in Salt Plains bait traps (Szymczak and Corey 1976). One gadwall was captured with its brood in a drive trap while the 2nd gadwall was captured in a bait trap. Five female mallards received transmitters that weighed from 27 to 30 g; 19-9 radio transmitters were attached to all others. Specific information for each bird is presented in Table 1. Condition/Energy Balance - Mallards Mallards were captured from 12 July through 20 September 1984 at a number of locations (Table 2) using a variety of methods. Most birds were captured in Salt Plains bait traps (Szymczak and Corey 1976). In most instances each bird captured was banded, weighed, and the length of primaries I, V and X recorded. Wing length (Ringe1man and Szymczak 1985) was recorded for birds that had not molted flight feathers (pre-molt) or had completed flight feather growth (post-molt) to the stage at which P IX was completely developed. Mallards in the pre-molt stage were classified to age using a combination of the techniques described by Carney (1964) and Krapu et a1. (1979). Post molt mallards were classified as adults. The presence of a brood patch in pre-molt females was recorded. The development of the alternate plumage in post-molt males was recorded by estimating the percent of belly and head feathers that had been replaced. A condition index (CI) was calculated for those mallards for which a wing measurement was recorded using predictor equations developed by Ringe1man and Szymczak (1985). = For females: fat For males: fat Both sexes: CI = (0.571 x body wt.)-(1.695 x wing length) + 59.0 (0.539 x body wt.)-(1.598 x wing length) + 31.5 fat body wt.-fat �17 Table 1. Data on instrumented mallards in North Park, summer 1984. Radio attached Trap Location Time period Bird Date U505 12 Jul Nest Allard Pond 12-18 Jul 20-23 Jul 23 Jul-IO Aug 20-27 Aug Nesting (Wt = 830g, Wing = 272mm) Movement: 5 km 50. with brood Brood rearing: Burr's Hay meadow Premolt: Hebron: Mush Pond (lost contact) 19 Jul Bait Hebron Pond 19-26 Jul 27 Jul-IO Aug 16-21 Aug Premolt: (Wt • 1010g, Wing· 272mm) Premolt: Mush Pond Premolt: Illinois R ~ 1-2 km S.E. Home Pond (Radio - removed) #507 Ad F Radio Wt • 199 19 Jul Bait Hebron Pond 19-27 Jul Premolt: (Wt • 970g, Wing· 268mm) Big Hebron - Mellon Ponds Premolt: Hopper's Pond Premolt: Mush Pond Area Premolt: Big Hebron Pond Premolt: Mellon Pond (lost contact) 11512 Ad M Radio Wt - 199 20 Jul 1st Yr F Radio Wt • 199 0.854 Unit F Radio Wt ~ 27g 30 Jul-03 Aug 04-14 Aug 15-20 Aug 21-27 Aug Bait Big Hebron 20-21 Jul 24 Jul-02 Aug 03-31 Aug 04-10 Sep #508 1st Yr F Radio Wt • 199 21 Jul 1.435 Ad F Radio Wt • 28g 22 Jul 1.335 1st Yr F Radio Wt • 28g 26 Jul Decoy Hatchery Raceway 30 Jul-02 Aug 07 Aug-ll 5ep Decoy Brewers Pond 24 Jul-17 Aug 21 Aug-04 Oct Bait Elk Pond 06 Aug 16 Aug-12 Sep 04 Oct. 1.463 1st Yr F Radio Wt • 28g 27 Jul Bait No. Allard Pond 30 Jul-07 Aug 15 Aug-13 Sep Status Premolt: (Wt - l290g, Wing = 292mm) Premolt: Big Hebron Flightless: Mellon Pond All locations in/near west end cattails Post molt: Hackley - Germ Pond (lost contact) Premolt: (Wt - 950g, Wing - 274mm) Premolt: Case #3 Pond Flightless: Illinois R ~ 0.4-0.8 km S5W of 50. School Section Pond, Scattered willows-sedges. (Mortality-Predation 11 Sep. PI - 117, V • 136, x - l24mm) Premolt: (Brood patch, Wt - 840g; Wing ~ 268 mm) Premolt: Bluebill - Case #2 Pond Premolt: Case D2 Pond (W. side bullrush) (lost contact) Brood Rearing: (Wt - 920g, Wing - 266mm) Premolt: Lake Creek-Park Ditch 3 km NE of Lake John; NE 1/4 5 28 Flightless: Spring fed seep 50m x 5Om, water ~ 0.5m deep with dense stand Carex nebraskansis. Recap. on 5 Sep., Wt:-;-850g, PI - 102, V - 117, X - 104mm) Post molt: Boettcher Lake Area (lost contact) Premolt: (Wt • 1160g, Wing - 268mm) Premolt: Illinois River, dense willowsedges adjacent to No. and So. Allard Ponds Flightless: Illinois River, dense willowsedges ~ 2 km So. of So. Allard Pond (observed 21 Aug) (lost contact) 0.774 Ad F Radio Wt • 28g 16 Aug Bait Big Hebron Pond 1.375 AdF Radio Wt • 28g 06 Sep Bait Brewers Pond 07 Sep 12 Sep Premolt: (Wt - 1060g, Wing = 27Omm) Premolt: Near confluence of Roaring Fork and No. Platte ~ 2.5 km SE of So. Delaney Butte Lake (lost contact) #510 1st Yr F Radio Wt· 06 Sep Bait Brewers Pond 07 Sep Premolt: (Wt • 970g) (lost contact) 19 g Premolt: (Wt - 1080g, Wing - 278mm) (lost contact) �18 Table 2. Location, capture, and molt status of mallards trapped in North Park July-September 1984. (Premolt = no flight feather shed. Molting = at least one primary still with blood in quill. Post molt = all primaries with hard quill. ) Wetland AM Age and sex bl:sta~e Premolt Moltin8 FYM FYF AM AF AF Hebron Pond Brewer's Pond Allard Pond Mellon Pond Lake John Annex Elk Pond Walden Reservoir Pole Mt. Reservoir 20 1 1 0 0 0 0 0 15 8 8 1 1 1 0 0 4 0 0 0 0 0 0 0 15 6 13 0 0 2 2 0 42 28 24 6 2 1 0 0 5 8 5 5 15 0 2 1 16 49 13 31 0 0 0 0 0 6 2 5 3 0 0 0 117 106 66 48 21 4 4 1 22 34 4 38 103 41 109 16 367 Totals Post molt AM AF Totals aAge derived following Carney (1964) and Krapu et al. (1979). bBirds hatched in the spring/summer of 1983. For analysis of molt stage in relation to any other variable, mallards were placed in 4 categories: 0 = had not mol ted flight feathers; 1 = first soft (blood in quill) primary I, II, or III; 2 = first soft primary IV, V, VI, or VII; 3 = first soft primary VIII, IX, or X; 4 = all primaries hard (no blood in quill). These categories classified birds to early, mid, and late molt categories. Only mallards in stages 2 and 3 were considered flightless. Condition/Energy Balance - Gadwall Adult gadwall trapped in conjunction with a banding operation were weighed and measured to classify stage of molt and relate stage to physical condition. The lengths of primaries I, V, and X were measured but in analysis to date, only P X has been used in establishing molt stage. Girth and uropygial gland to tip of the nail of the bill measurements were taken as described by Szymczak and Ringelman (1983) to use in condition index estimation (Szymczak 1984). If a bird was subsequently recaptured, only weight, girth, and primary measurements were recorded. Birds were captured at Walden Reservoir (7, 13, 22, and 19 Aug; 17 and 27 Sep), MacFarlane Reservoir (8, 16, 23 and 30 Aug; 18 and 26 Sep), Lake John Annex (9 and 28 Aug; 19 Sep), and Pole Mountain Reservoir (14 Aug). �19 RESULTS Walden Reservoir Census Fewer adult ducks were recorded on Walden Reservoir in 1984 in late July than in 1983 (Table 3). In early August 1984 there were more ducks but fewer flightless birds than in 1983. Water releases into Walden, which were hypothesized as the cause of the reduction in the number of birds at Walden in large July 1983, began in early July prior to the first 1984 count. The water increased in depth ~ 1 cm/day from 13 July through 24 August. As in 1983, the molting population was predominantly gadwa11s (Table 3). After July 1984 the number of gadwa11s on Walden Reservoir was similar to the number recorded in 1983 (Fig. 1). The peak of the gadwall flightless period, occurred about 15 August in both years. A higher percent of the birds counted in late August 1984 were flightless, possibly indicating increased numbers of females in the population. The location of concentrations of flightless gadwall was somewhat different in 1984 than in 1983 (Fig. 2). In 1983, most of the flightless gadwall observed were near the small islands in the central portion of the reservoir while in 1984 the major concentration area was west of the former location (Fig. 2). In 1984, few flightless gadwall frequented the north bay but the area south of the large island in the central portion of the reservoir increased in importance (Fig. 2). In contrast to 1983, a few flightless gadwall were located in the southeast bay, but those birds were there only during the final count of the year. Considering that radio-marked flightless birds were not located in the southwest bay, birds recorded there may have been classified incorrectly. The distribution of flightless gadwall on Walden Reservoir in 1984 is presented in Fig. 3. The shift in major concentration areas is difficult to locations were in areas of predominantly open water although overlap into a stand of 50% Potomageton fi1iformis and exa1bescens. In 1983, the birds were adjacent to patches of 2 aquatic plants into which they may have made periodic Analysis of telemetry data may substantiate those movements. interpret. Both the 1984 area did 50% Myriophyllum mixtures of these foraging forays. Gadwall in 1983 used the small island at the south end of the concentration areas as a land roost but a combination of wave action and increased water levels changed the exposed structure of the island in 1984. Water was added to the reservoir in both years beginning in late July. About 55 and 40 cm of water, as measured at the spillway, were added to the reservoir in 1983 and 1984 respectively. Prior to the water additions, spillway depth was 510 cm in 1983 and 550 cm in 1984. Increased water levels during this period affected the availability of aquatic vegetation in both years but more water in 1984 may have affected duck distribution more than in 1983. t101tChronology and Species Composition Periodic counts of adult ducks on selected ponds to document chronology of molt by species in 1984 provided results similar to those obtained in 1983 �N o Table 3. Species composition of adult ducks recorded during weekly counts of waterfowl concentrations on Walden Reservoir, 1984. (Percent flightless in parentheses.) Date of count 27 03 10 334 (0.9) 335 177 156 14 12(16.7) 25 36 7 0 0 0 0 0 370 (7.0) 271 (0.4) 74 50 18 13 (7.7) 23 (8.7) 18(38.9) 5 1 0 0 0 0 381(34.1) 80 (8.8) 51 37 17 18 (5.6) 3 14(42.9) 0 0 0 0 0 0 391(44.8) 156(16.0) 59(16.9) 95 (2.1) 75 (4.0) 4~(26.1) 7 17(23.5) 18 1 0 0 0 0 411(65.5) 174(18.3) 88 (8.0) 36 (5.6) 138 (8.0) 82 24 (4.2) 17(11.8) 9(11.1) 1 6 0 0 0 472(80.9) 108(15.7) 143(12.6) 21(23.8) 128 (9.4) 58 23 5(20.0) 11 1 5(20.0) 1 0 0 1,083 (0.5) 843 (4.4) 601(24.0) 865(27.6) 986(33.0) 976(44.6) 1,362(25.5) 1,697(16.7) 1,871 (2.0) 1,864(15.2) 614(54.6) 702(74.1) 1,132(66.9) 1,361(16.2) 1,405 (6.2) SEecies Gadwall Am. wigeon L. scaup Redhead Ruddy duck Mallard BWT/CT No. pintail GWT Canvasback Ring-necked duck Common merganser C. Goldeneye No. Shoveler Totals Totals (1983) August 17 13 Ju1l:: 20 -- 24 31 527(53.9) 374 (9.9) 85(14.1) 61 (8.2) 172 (4.7) 79 26 (3.8) 19 16 0 3 0 0 0 352(53.4) 748(12.0) 76 (2.6) 85 131 (0.8) 37 58 7(14.3) 157 1 10 0 1 34 �21 ADULT WALDEN GADWALLS RESERVOIR rJ) ..J ..J 41( Q == 41( ~ ~ ..J 352 :::;:) Q .....••. 41( rJ) rJ) W ..J 381.··· ···· ···· · ...· ~ J: e ..J L&. ~ Z W o a: w : 20 454 / /1191 1983--- 370/ ...' 1984············· ' Q. 1 15 AUG 1 15 SEPT Figure 1. :>lumberof adult gad~ll present on Ualden Reservoir and the percent estimated as flightless, mid-July through August 1983 and 1984. �22 WALDEN RESERVOIR DISTRIBUTION OF FLIGHTLESS ADULT GADWALL iii 1983 ~ 1984 Figure 2. Distribution of flightless adult gadwall and 1984, with percentage within specified year indicated. comprising 5% or more of the population are indicated. in 1983 Only those areas �23 W ALDEN RESERVOIR DISTRIBUTION OF FLIGHTLESS ADULT GADWALL <1% • 1-5% 1m 5-10% II 10-20% • >20% Figure 3. Distribution of flightless adult gadwall observed on Walden Reservoir from 13 July through 31 August 1984. �24 (Szymczak 1984). Peak molt periods by species were the same in both years, however sample sizes for some species were reduced because Boettcher Lake was deleted as a study area for logistical reasons. The most surprising results were the numerical and species composition changes between years on individual ponds. Alkali Pond, which supported molting populations of predominantly green-winged teal (Anas crecca), pintails (Anas acuta), lesser scaup (Aythya affinis) and blue-Winged/cinnamon teal (Anas spp.) in 1983 supported few flightless ducks in 1984 (Table 4). Hebron Pond had gadwall and greenwings in 1983, but virtually nothing in 1984. Pole Mountain Reservoir supported molters in both years, but the species composition changed. Table 4. Maximum number of flightless adult ducks observed in 1983 and 1984 on ponds in North Park. SEecies Gadwall Mallard Am. wigeon No. pintail Green-winged teal Blue-winged/ Cinnamon teal L. scaup Alkali Pond 83 84 76 Pond 84 83 Elk Pond 84 83 Hebron Pond 83 84 Pole Mtn. 83 84 8 0 1 38 110 1 0 1 0 6 4 0 12 2 16 3 2 2 0 3 11 7 5 1 13 53 2 5 0 8 32 0 8 0 41 1 0 0 0 2 79 3 122 0 2 32 5 23 15 29 16 22 8 0 8 3 3 0 16 1 4 0 8 2 0 2 3 10 9 4 Molt Site Selection Four mallards (3 females and 1 male) were monitored into the flightless period (Table 1). One female moved about 28 km and another moved 8 km from capture sites to molting areas. The other mallards were captured near molting locations. Five other pre-molt females were monitored until they could no longer be found in North Park (1-40 dys) indicating they left the area before molting flight feathers. The radio was removed from one mallard about one month after capture because the bird was in poor body condition. Another bird was located periodically for 75 days before monitoring was stopped. The four flightless mallards used areas of dense emergent vegetation. The male spent 14 days on Big Hebron Pond which has thinly scattered spikerush (Eleocharis sp.) before moving about 1 km south into or near a small cattail patch on Mellon Pond. Female #508 moved from Case #3 Pond, a wetland with a margin of grazed spikerush, to an area of flooded Carex and willow (Salix sp.) along the Illinois River. Female 111.463 was trapped on North Allard Pond which has a margin of dense flooded Carex, spent a 10-day pre-flightless period in a dense Carex-Salix area on the Illinois River adjacent to the pond, and then moved about 2 km upstream into an area of similar vegetation during the flightless period. Female #1.335 moved from Elk Pond, which has a �25 substantial spikerush margin to a small surface hectares, about 0.5 m in depth nebraskensis. Neither of was located Big Hebron Greasewood marking. spring-fed supporting wetland with about 0.25 a dense stand of Carex the 2 female gadwall instrumented during the brood-rearing period during the flightless period. Contact with the female marked on Pond was lost 18 days after marking while the bird captured on Pond on Case Flats moved out of North Park about 14 days after Weight and Condition of Mallards Weight and condition dynamics during the pre-molt through post-molt period were analyzed by comparing (1) changes in individual birds recaptured periodically, throughout the period and (2) change in the entire sample population grouped according to stage of molt. Examination of recapture data indicated inconsistencies in trends of weight and condition, probably the result of using predominantly baited traps to capture birds and weighing birds immediately after capture. Birds recaptured in bait traps did not consistently gain weight as might be expected. However, subsequent analyses were directed toward the entire sample population using only records from the first date of capture. North Park mallards seem to be similar in weight to other populations studied during the mid-late summer period. Mean weight of males (ages pooled) (1,200 g) in Stage 0 was slightly less than those (1,239 g) recorded by Folk et al. (1966) in Europe or Young and Boag (1982) in Manitoba for males ('V 1,265 g) that had just shed their flight feathers or were in the early stage of primary growth. However, during the period following breeding and prior to remige molt, males gain weight (Folk et al. 1966, Young and Boag 1982). North Park males captured during Stage 0 may have attained average weights similar to those recorded for other populations by the time flight feathers were shed. Comparative weights for females specific to the stage of molt are not available for other populations. However average weight for females during July, August, and September in North Park (976 g) compare favorably with those reported in England (991 g) by Owenand Cook (1977) and by Folk et al. (1966) (979 g). Birds in the premolt stage (0) were classified to age (Table 2). Adult females were heavier (P < 0.05) than first-year birds (Table 5) during the premolt stage but differences in structural size as indicated by wing length were slight (AF = 273 mm, FYF = 270 mm) but significant. The variation in structural size apparently accounted for weight differences as there was no difference in condition (P > 0.28, Table 6) of birds in the 2 age groups. Males in the 2 age groups showed no difference in weight (p > 0.31), wing length (P > 0.06), or condition (p > 0.40), but few first-year birds were included:tn the sample. - �26 Neither weight (p > 0.84) nor condition (p > 0.60) varied between trapping locations for either sex during the pre-molt period. Females in the pre-molt stage captured and measured prior to 15 August had similar weights (p > 0.53) and condition (P > 0.20) to those captured after that date. All males were captured during~ate July through early August and therefore were not measured for seasonal variation. Identifying characteristics used to classify birds to age were lost when birds shed their flight feathers and analysis following stage 0 classified birds only to sex. Weights of mallards declined significantly in both sexes following molt of flight feathers (Table 5, Fig. 4). In both sexes pre-molt weights (ages pooled) were significantly higher than weights of birds captured during growth of flight feathers. Birds seem to lose weight until rreaching stage 3, when they regained the ability to fly. Other investigators have also reported weight loss in mallards during wing molt (Folk et al. 1966, Owen and Cook 1977). Young and Boag (1982) could not demonstrate a significant weight loss in male mallards during the wing molt although they did record a reduction in pectoral and leg musculature combined and carcass lipids. We suspect that a closer examination of sample birds according to different stages of remige molt by the latter authors may have indicated a significant but short-term weight loss. Condition followed a similar trend as weight although methodology allowed estimation of condition only during premolt, late molt and past molt stages (Table 6). Declines in condition from premolt levels were noted in both sexes. The condition of late molt (PX soft) males was lower (R < 0.05) than both pre- and post-molt birds. Female condition decline was significant only after reaching the post molt stage. The comparative low condition index of females in the post-molt stage is probably the result of females having a later molt cycle. Most females captured during the post-molt period had most likely recently completed primary feather growth and had not had sufficient time to restore body reserves. Female weights during the post-molt stage showed a similar lag. There was no variation in condition according to sex (p > 0.38), however, differences in post-molt condition did approach signifrcan~e (R = 0.057) (Table 6). Weight Dynamics of Flightless Gadwall Weight/molt stage data indicated that gadwall captured in 1984 were progressively heavier through the flightless period than during the previous 2 years (Fig. 5) indicating possible changes in wetland quality used by molters. Captured birds generally exhibited the same trends in weight dynamics noted in the past; increasing weight during the 1st one-third of the flightless period, decreases in the 2nd one-third, and increasing weight in the late stages. Weight differences were not significant for males at any stage but female pre-molt (stage 0) weights were significnatly less than those in stage 2 (31-60 mm) weights which were in turn greater than stage 4 (91-120 mm) weights. Stage 3 (61-90) and 4 female weights were significantly less than stage 5 (> 120 mm) weights. Recaptures of individual birds within the same increase/decrease phase, although few in number, indicated the same weight dynamics cycle (Table 7). �27 1300 1200 --J: o f 4 1 3 2 MALES 25> •........•.. ...•.....•.. ......... 1100 Cl t- o 1000 W ~ z « w 900 :iE 800 o NO MOLT 1 I-III SOFT 2 IV-VII PRIMARY SOFT 3 VIII-X SOFT 4 ALL HARD MOLT STAGE Figure 4. Relationship of body weight to stage of primary categorized according to progress of primary growth. i.e., least primary 1-3 and maybe complete in primaries 4, 5 and weights. Weights in stages with common underlines did not molt in North Park mallards 1984. Molt stage for all birds in stage 2, growth complete in at 6. Vertical lines indicate sample range in differ (!> 0.05). �28 . 135 62 ...... ..•.....• ....••• ••••••••30 •••••• 23 ..""/ ..... ...••... I ~ ~I I , I .~ sn 68 / ••• : y 92 3.·I :E Old:! a: o - MALES ~ J: -w (!) ;: ~ 0 750 0 OJ 1982 1983 725 1984 •••••••••••• 700 40 675 ---- 27 ---- 86 FEMALES -~~------54 39 - 27 PRE-MOL T 0-30 LENGTH 61-90 91-120 >120 OF P X (mm) Figure 5. Relationship of body weight to flightless stage of gadwall in North Park 1982, 1983, and 1984. �29 Table 5. Mean weights (~ SD) by stage of molt for mallards captured in North Park, July--September 1984. 0 Wt. Ase/sex 1 N N Wt. Males Adult 1st Year Combined 1,212 (+122) 21 1,135 (+161) 4 1,200 (IU8) 25 1,103 (~92) Females Adult 1st Year Combined 1,036 (+114) 34 989 (+ 75) 36 1,012 (I 09) 70 935 (~90) Sta e 2 Wt. 8 1,042 (~79) 17 4 3 N Wt. N 23 1,080 (~82) 873 (.:!:66) 11 930 (~75) N Wt. 74 1,198 (~102) III 14 16 971 (~82) Table 6. Mean condition index (+ SD) for mallards captured in the premo1t, late-molt (p X soft) and post primary molt (all primaries hard) stages in North Park, July-September 1984.a Premo1t ~05 Index Ase/sex Stases Late molt ~px5 Index N N Post molt Index N Males Adult 1st Year Combined 0.212 (+ 0.054) 0.186 (= 0.067) 0.208 (~ 0.056) 21 4 26 0.166 (~ 0.041) 25 0.203 (~ 0.041) 110 Females Adult 1st Year Combined 0.217 (+ 0.062) 0.203 <+ 0.045) 0.210 (! 0.054) 34 35 69 0.177 (~ 0.053) 7 0.172 (~ 0.050) 12 ap IX must be full length to compute the condition index following Ringe1man and Szymczak (1985). Table 7. Weight dynamics of adult gadwall captured more than once during the 1984 trapping (molting) season. Males P X (mm) pre-molt - 60 61-120 120 Wt. gain N S/daz 0 0 3 4.91 Wt. loss N S/daz 1 6 1 6.25 8.12 1.25 Females Wt. gain Wt. loss N N s/daz s/daz 2 0 0 4.71 2 7 1 1.97 5.58 8.57 �30 LITERATURE CITED Anderson, M. G., R. D. Saylor and A. D. Afton. ducks. J. Wildl. Manage. 44:217-219. 1980. A decoy trap for diving Carney, S. M. 1964. Preliminary keys to waterfowl age and sex identification by means of wing plumage. U.S. Fish Wildl. Servo Spec. Sci. Coulter, M. W. 1958. Dwyer, T. J. 1972. 43:282-284. A new waterfowl nest trap. An adjustable radio-package Bird Banding 29:236-241. for ducks. Bird Banding Folk, C., K. Hudee, and J. Toufar. 1966. The weight of the mallard and its changes in the course of the year. Zool. Listy 15:249-260. Krapu, G. L., D. H. Johnson, and C. W. Dane. mallards. J. Wildl. Manage. 43:384-393. 1979. Age determination in Owen, M., and W. A. Cook. 1977. Variations in body weight, wing length and to condition of mallards (Anas platyrhynchos) and their relationship environmental changes. J. Zool., London 183:377-395. Ringelman, J. K., and M. R. Szymczak. 1985. A physiological for wintering mallards. J. Wi1d1. Manage. 49:564-568. condition index Szymczak, M. R. 1984. Ecological studies of the flightless period of ducks in Colorado. Colorado Div. Wi1d1. Fed. Aid Wi1d1. Res. Rep., Oct. Pp. 13-31. ____ ~_' and J. F. Corey. 1976. Construction and use of the Salt Plains duck trap in Colorado. Colorado Div. Wi1dl. Civ. Rep. 6. 13pp. _______ , and J. K. Ringe1man. 1983. Ecological studies of the flightless period of ducks in Colorado. Colorado Div. Wi1d1- Fed. Aid Wi1d1- Res. Rep., Oct. Pp. 13-31. Young, D. A., and D. A. Boag. 1982. Changes in physical condition of male mallards (Anas p1atyrhynchos) during molt. Can. J. Zool. 60:3220-3226. Prepared by 77l,JJ-f,' ~.-L Michael R. Szymcz~ Wildlife Researcher �31 Colorado Division of Wildlife Wildlife Research Report April 1986 JOB PROGRESS REPORT State of Colorado ---------------------------------- Project Work Plan 01-00-045 (W-88-R) 1: Period covered: Job Avian Research 15 01 July 1984 through 30 June 1985 Author: J. K. Ringelman Personnel: J. F. Corey, J. K. Ringe1man, M. R. Szymczak, Colorado Division of Wildlife. ABSTRACT Mean body weights and fat reserves of mallards (Anas p1atyrhynchos) captured during winter 1984-85 were higher than those recorded in previous years, reflecting the relatively mild winter. Body weights were lowest during mid-winter, but juveniles retained total fat reserves nearly comparable to their adult counterparts. Based on fat reserves during 1984-85, survival time for males under starvation conditions would have been about 10 days; survival time for females was estimated at 12 days. During January, adult females had the highest condition index (CI) values at Bonny Reservoir, whereas juvenile females and males of both ages had greater CI values at Sege1ke Slough. A re-examination of CI data from 1982-84 suggested that as mean values of CI increased, standard deviations of CI decreased (rho = -0.462) and some distributions became negatively skewed. Data from wild birds supports the hypothesis that competition during periods of severe winter weather alters the habitat selection of subordinate individuals, particularly juvenile females. Changes in habitat selection cause differential foraging success and lead to a high variability in CI. It is important to consider this variability when estimating the percentage of a population below a given level of CI. ��33 DEVELOPMENT AND USE OF A PHYSIOLOGICAL CONDITION INDEX FOR MONITORING WINTERING MALLARD NUTRIENT RESERVES James K. Ringe1man Michael R. Szymczak Only recently have waterfowl biologists come to appreciate the role of stored nutrients in the life histories of ducks and geese. Earliest evidence of the importance of nutrient reserves was obtained from studies on arctic-nesting snow geese (Chen caeru1escens; Ankney 1974, Ankney and MacInnes 1978), in which it was demonstrated that the size of reserves prior to breeding was directly correlated with the number of eggs in the clutch. The same relationships were later demonstrated for Canada geese (Branta canadensis; Raveling 1979). Nutrient reserves important for maintaining physiological condition are not only obtained in migratory habitat during the spring (Raveling 1979), but may also be accumulated on wintering grounds (MacInnes et al. 1974). Current studies on arctic-nesting geese have progressed beyond the point of demonstrating the importance of nutrient reserves to investigations addressing when and how these reserves are obtained. The role of nutrient reserves during the breeding period of ducks is less evident than in geese. Bengston (1971) provided field data to support Lack's (1967) hypothesis that the clutch size of ducks is related to food availability on the breeding grounds, yet Milne (1976) demonstrated that the weight of female Common eiders (Somateria mo11issima) in winter was directly related to clutch size the following spring. The apparent discrepancy in these results was resolved by more detailed nutritional studies: clutch size in eiders is limited in part by the size of stored protein reserves (Korschgen 1977) whereas many dabbling ducks wait until arrival on the breeding ground to obtain protein needed for egg production (Krapu 1974, 1981; Drobney 1977; Reinecke 1977). For the mallard, there remains a common link between nutrients obtained in winter and spring and subsequent reproductive success: fat reserves provide the "fuel" that enables mallards and similar species to forage for the higher protein but lower energy animal foods (Krapu 1981). If the mallard population is regulated by limitations imposed during winter, as has been suggested for most temperate bird populations (Fretwell 1972), such regulation may also result from increased winter mortality. Most reports of waterfowl mortality during winter cite severe cold and/or snow as the causative factors (Pearson 1934, Gromme 1936, Trautman et a1. 1939). Fat, the most labile endogenous reserve, serves as the principal energy source used by waterfowl to survive severe winter weather. In all likelihood, winter fat reserves of mallards serve dual functions of increasing winter survival probabilities as well as providing nutrients and energy essential for reproduction. The question of foremost interest is what minimum level of winter fat reserves are necessary to optimize both survival and reproduction? With the successful development of a condition (fat) index during the early phase of this study, emphasis during 1984-85 shifted to documenting the age and sex specific, temporal, and spatial variability in physiological condition among mallards wintering in Colorado. �34 P. N. OBJECTIVES The objectives of this study are to: 1. Develop a physiological condition index reserve levels of wintering mallards. to accurately assess nutrient 2. Determine if differences exist in the physiological condition of mallards wintering in several areas of northeastern Colorado. 3. Document temporal changes in mallard physiological condition on a major Colorado wintering area. 4. Relate spatial and temporal differences in mallard condition, if such differences exist, to weather and the availability of waste cereal grain. SEGMENT OBJECTIVES 1. Capture, weigh, and measure 160 mallards (40 of each age and sex) during winter at four northeast Colorado wintering areas. 2. Apply the physiological condition index to determine regional, sex, and age-specific differences in the level of nutrient reserves. 3. Relate nutrient reserves to weather and the abundance of waste cereal grains in the vicinity of capture sites. METHODS Salt Plains bait traps (Szymczak and Corey 1976) were used to capture mallards at Kodak Ponds (3-4 Dec 1984; 10-11 Jan, 22-24 Feb 1985), Bonny Reservoir (3-4 Dec 1984; 15-17 Jan, 23-28 Feb 1985), Valmont Reservoir (14-16 Jan 1985), and Segelke Slough (17-21 Jan 1985). Forty mallards of each age and sex class were captured at each location duririg each trapping period. Body weight and wing length measurements were obtained on all birds. If necessary, mallards were held 24 hours to allow digestion of ingested corn prior to weighing. Sex-specific condition index equations were used to estimate total and relative fat reserves (Ringelman and Szymczak 1985). Conventional analysis of variance procedures were used to compare mean levels of condition index (CI) among locations during January 1985. Summary statistics for CI, total fat, and body weight were also tabulated for 1984-85. The implications of non-normality and heteroscedasticity in sample distributions of CI were explored using data obtained during this study prior to the 1984-85 field season. Trapping and analytical procedures were identical in all years, thus the conclusions reached as a result of this analysis have relevance to the 1984-85 data set but do not include these data. �35 Summary statistics, including measures of skewness and kurtosis, were generated using the condescriptive procedure of SPSS (Nie et a1. 1975). Because we were interested in the variability within a data set, irrespective of the magnitude of CI values, the coefficient of variation (CV) was used as the preferred measure of variability when performing Spearman Rank correlations. Differences between coefficients of variation were tested directly (Zar 1974:104). Normal deviates were calculated to determine the proportion of a normal distribution lying below a given value of CI (Zar 1974:74). Normal curves were fitted to CI data by computing the frequencies to be expected if the data were normal (Zar 1974:81). RESULTS AND DISCUSSION 1984-85 Mallards Winter weather during 1984-85 was relatively mild compared to the previous two winters. Body weights of mallards reflected the effects of high temperatures and low snowfall (Table 1), exhibiting the typical pattern of highest mean winter body weight early in winter and lowest weights during mid-winter. In all cases, juvenile females were lightest and adult males the heaviest. Most change in body weight was attributable to fat deposition or loss (Table 2). Disparaties in fat reserves among sex-age classes was less than those observed for body weight, with juvenile birds retaining reserves nearly equal to those possessed by adults. Table 1. Mean body weights (+ standard deviation) of mallards captured in northeastern Colorado during winter, 1984-85. Each mean value is derived from a sample of 40 birds. Location Month AauIt female Weight (g) by age-sex class Adult male Juvenile female Bonny Dec Jan Feb Dec Jan Feb Jan Jan 1,014 997 956 989 938 983 972 1,006 1,158(109.2) 1,093 (77.5) 1,065 (80.8) 1,119 (72.4) 1,089 (87.7) 1,095 (76.3) 1,103 (85.3) 1,172 (82.9) Kodak Va1mont Sege1ke (88.0) (78.3) (67.0) (76.7) (71.1) (62.6) (53.1) (85.8) 970 (66.5) 916 (67.6) 942 (78.0) 915(102.1) 888 (68.1) 943 (66.0) 953 (73.5) 993 (63.7) Juvenile male 1,101 1,018 1,062 1,044 1,031 1,058 1,037 1,101 (92.1) (85.3) (73.8) (89.9) (69.3) (63.2) (82.1) (86.7) �36 Table 2. Mean estimated total body fat content (+ standard deviation) for mallards captured in northeastern Colorado during winter, 1984-85. Each mean value is derived from a sample of 40 birds. Location Month Bonny Dec Jan Feb Dec Jan Feb Jan Jan Kodak Va1mont Sege1ke Adult female 164 155 137 156 122 140 136 155 Estimated fat (g) by age-sex class Juvenile male Juvenile female Adult male (47.3) (43.1) (36.8) (42.4) (37.5) (37.6) (31.7) (47.8) 178 149 135 165 141 143 144 182 (54.2) (41.3) (41.5) (38.0) (42.0) (39.0) (42.0) (40.5) 147 120 139 124 105 127 136 158 (36.7) (37.1) (43.0) (51.0) (35.5) (33.2) (39.1) (35.4) 163 128 149 139 130 137 128 164 (45.5) (45.9) (39.3) (43.6) (33.9) (30.0) (43.3) (42.6) Consideration of fat reserves in relation to metabolic demands provides an estimate of survival time under starvation conditions. If starvation of mallards occurs when weight loss approaches 45% of mean maximum body weight (Jordan 1953), then males in 1984-85 could lose about 500 g and females about 450 g weight. Since mallards wintering in Colorado lose 0.472 g fat/g body weight (J. K. Ringe1man, unpub1. data), these losses would equal 236 g (males) and 212 g (females) of fat. At 9.45 Kca1/g fat, this equates to 2,230 (males) and 2,002 Kca1 (females). Jorde (1981) estimated daily energy costs of mallards wintering in Nebraska as 225 Kca1/day for males and about 165 Kca1/ day for females. Thus, fat reserves carried by mallards in Colorado during winter 1984-85 were sufficient to provide about 10 days energy reserve in males and 12 days reserve in females. During January, adult females had higher values of CI at Bonny than at Kodak (~ < 0.05, Table 3). Juvenile females were in relatively higher condition at Sege1ke, intermediate condition at Va1mont and Bonny, and relatively poorer condition at Kodak (P < 0.05). Males of both ages had higher mean CI values at Sege1ke than at the other three locations (f < 0.05). Table 3. Mean condition index (CI) values (+ standard deviation) of mallards captured in northeastern Colorado during winter, 1984-85. Each mean value is derived from a sample of 40 birds. Location Month Adult female CI by age-sex class Adult male Juvenile female Bonny Dec Jan Feb Dec Jan Feb Jan Jan 0.191(0.045) 0.182(0.044) 0.167(0.040) 0.186(0.045) 0.149(0.041) 0.165(0.040) 0.162(0.035) 0.179(0.048) 0.179(0.047) 0.156(0.038) 0.143(0.039) 0.172(0.034) 0.147(0.039) 0.149(0.036) 0.149(0.037) 0.182(0.034) Kodak Va1mont Sege1ke 0.177(0.039) 0.150(0.042) 0.172(0.048) 0.154(0.056) 0.132(0.041) 0.155(0.035) 0.164(0.042) 0.189(0.038) Juvenile male 0.172(0.041) 0.141(0.047) 0.162(0.038) 0.152(0.041) 0.143(0.033) 0.148(0.027) 0.140(0.043) 0.173(0.037) �37 Variability and skewness within a CI distribution The variability within a sample distribution of CI values, as well as the extent to which a distribution deviates from normality, may provide as much biological insight into wintering mallard populations as do mean values of CI. Consider, for example, the data set of CI values from January 1983 (Table 4). When comparing all age and sex groups combined, the sample standard deviations for CI increased with decreasing mean values of CI (rho = -0.462, I-tailed test, P = 0.04; Table 4). Standard deviations averaged 0.041, but ranged from 0:031 (CI = 0.195) to 0.055 (CI = 0.174). The frequency distributions of CI values for juvenile female mallards during early, mid, and late winter 1983-84 exemplify how CI and sample coefficients of variation (CV) change over time (Fig. 1). During a period of moderately severe winter weather, CI averaged 0.141 and CV equaled 37.6%. As weather conditions became more severe during mid-winter, mean CI (0.120) changed insiginificant1y (p > 0.05) but CV increased (49.8%; P < 0.05). When severe weather abated in late winter, mean CI increased (o.lil; P < 0.05) while variability in CI showed a marked decline (CV = 18.6%; ~ < 0.001). Table 4. Means (± standard deviation) of a condition index calculated for mallards trapped at four northeastern Colorado locations during January 1984. Each datum represents a sample of 40 birds. Area 1 2 3 4 Adult female 0.152 0.174 0.187 0.211 (0.053) (0.055) (0.038) (0.033) Condition Index Juvenile female 0.166 0.188 0.189 0.208 (0.047) (0.052) (0.048) (0.041) Standard Deviation Adult male Juvenile male 0.173 0.168 0.195 0.188 (0.044) (0.045) (0.031) (0.032) 0.156 0.180 0.175 0.195 (0.035) (0.037) (0.038) (0.035) Although asymmetries in the distributions of CI usually were not statistically significant, there was a tendency for distributions with high mean values of CI to be leptokurtic and negatively skewed. For juvenile female mallards (Fig. 1), late winter birds were not only in better condition, but several individuals had CI values near and to the right of the mean (kurtosis = 0.224, skewness = -0.629). The distribution of CI values among mid-winter females in poorer condition was p1atykurtic (kurtosis = -0.269), but skewness was not severe (skewness = -0.205). Thus, mean values of CI may not provide a good measure of central tendency in some instances. During early, mid, and late winter male mallards was nearly constant and mean values of CI (Table 5). value and CV (S.D. = 10.44) in CI the same period. 1982-83 and 1983-84, the CV in CI for adult (S .D. = 1.72), despite changes in weather In contrast, large changes in both the mean were apparent among juvenile females during �38 10 6 2 10 6 >- o z W :::::l o w 2 a: u, 18 14 10 6 2 0.06 0.12 0.18 CONDITION 024 INDEX Fig. 1. Actual (histogram) and normalized (solid line) frequency distributions of condition index values for juvenile female mallards during early (top), mid (middle), and late (bottom) winter 1983-84. Ducks were captured at Bonny Reservoir, CO. Sample sizes are 40 (early and late winter) and 39 ducks (midwinter). �39 Table 5. Means and coefficients of variation (CV) of a condition index (CI) calculated for four age/sex classes of mallards captured at Bonny Reservoir, Colorado. Sample size equals about 40 birds per datum. Age/sex class Years Adult female 1983-84 1982-83 Juvenile female 1983-84 1982-83 Adult male 1983-84 1982-83 Juvenile male 1983-84 1982-83 Mean CI CV of CI 0.183 0.163 0.168 0.210 0.152 0.170 0.141 0.120 0.171 0.169 0.166 0.150 0.146 0.141 0.157 0.168 0.173 0.156 0.129 0.139 0.154 0.150 0.156 0.140 22.6 32.3 23.6 25.6 35.1 23.4 37.6 49.8 18.6 34.3 28.0 37.1 27.2 25.7 25.7 30.1 25.6 26.8 32.6 32.2 23.5 33.5 22.6 33.0 (%) Standard deviation of CV values 5.28 10.44 1. 72 5.07 Although a continuum of body condition exists in any population of wintering waterfowl, it is reasonable to expect that suboptimal condition may impact population parameters (Le., survival, reproductive potential) only below a threshold level. If that is the case, investigators should carefully examine variability in CI. The variance as well as the mean of a distribution influences the estimate of the number of individuals below a given value of CI. Consider two sets of CI frequency distributions for juvenile female mallards. The first set (Fig. 2, top) depicts the frequencies expected in a normal distribution where sample means and variances are taken to be the actual population mean and variance. The second set (Fig. 2, bottom) illustrates samples with the same number of individuals (40) and mean values of CI, but with a hypothetical common variance equal to the average of the three real sample variances. If a hypothetical threshold of suboptimal condition is 0.09, then 16.8, 30.7, and 0.5% of the individua1s in the population would have CI values < 0.09 during early, mid, and late winter, respectively (shaded area, Fig. 2, top). However, samples with the same means �40 12 8 >- o z 4 W ::J ~ a: u, 0 8 4 o o 0.24 0.12 CONDITION Fig. 2. Normalized frequency distributions female mallards during three winter periods with actual means and sample sizes (~= 40) Shaded areas denote those portions of curves INDEX of condition indices for juvenile (top) vs. frequency distributions but equal variances (bottom). below a condition index of 0.09. �41 but a commonvariance would have 14.5, 26.8, and 4.7% of the individuals below a CI of 0.09, in the same order (shaded area, Fig. 2, bottom). Thus, two populations might have equal mean values of CI but different percentages of birds below a threshold condition. Conversely, samples with different CI means could have the same percentage of individuals below a given CI. A reduction in the CV of CI among female mallards during late winter could reflect a build-up of endogenous reserves preparatory to breeding. To test that hypothesis, we calculated CI values and associated CV for female mallards measured during early, mid, and late winter, 1982-83. No consistent decrease in CV during late winter was noted among either adult or juvenile female mallards at two locations, but CV did increase with decreasing CI described earlier (Table 4). Correlation between the mean value and the variability of CI suggests a biological causation. Condition reflects the energy balance of the individual, so it is necessary to consider both energy expenditure and acquisition when considering a plausible mechanism. The most energetically costly activities, such as flight and· vigorous feeding, often occur synchronously during field-feeding bouts. During interludes between feeding, mallards in Colorado spend nearly all day and night concentrated in large flocks on lakes or rivers. Thus, thermoregulatory costs may differ among local populations but (except for differences attributable to body size) they should be similar within a flock. These similarities in behavior and thermoregulation suggest that only small differences in energy expenditure would be expected among mallards wintering in Colorado. Several hypotheses can be advanced to explain the variability in body condition related to mean values of CI. Since an upper limit exists in the amount of fat reserves a bird can carry, we hypothesize that a reduction in the CV of CI at high levels of CI may be a simple function of the population approaching this maximumvalue. This relationship would also explain the tendency towards a negatively skewed distribution at high levels of CI. Theoretically, the CV of CI should also decrease at low levels of CI due to death through starvation, although we did not observe evidence of this decrease among wild mallards even under the most severe winter conditions. However, the changes we observed in the CV of CI were evident in wild mallards with fat levels far less than maximum,as evidenced by (1) the high CI values achieved by captive mallards (J. K. Ringelman, unpubl. data) and (2) fat levels reported for wild mallards overwintering in other regions (Whyte and Bolen 1984) that far exceed fat reserves carried by mallards in Colorado. Thus, this hypothesis does not explain the changes in CV we observed within a relatively narrow, moderate range of CI values. Weather-induced changes in the behavior of field-feeding waterfowl were observed in early studies of waterfowl ecology during winter (Bellrose 1944, Hochbaum1955, Bossenmaier and Marshall 1958) and continue to be addressed in recent studies (Baldassarre and Bolen 1984). The tendency of mallards to modify their traditional morning and evening field-feeding behavior during cold, blustery weather (Bellrose 1944) or during periods of snowfall (Baldassarre and Bolen 1984) is particularly noteworthy. Our second hypothesis presumes that weather, the ultimate factor regulating condition of mallards in �42 Colorado, causes differential foraging success among individuals. Differences in net energy acquisition are correlated with this variable foraging success, and are the basis for the high variability in condition during severe winter periods. During severe weather, small groups of mallards forage throughout the day, using a wide variety of cornfields located near roosting areas (Be11rose 1944, J. K. Ringelman, unpub1. data). Because of the large differences in waste corn abundance among fields (Baldassarre and Bolen 1984), the increase in the number of different cornfields used results in variable foraging success. If our second hypothesis is correct, and the sex and age composition of small feeding flocks are representative of the local population, variability in CI within all agelsex groups should increase during severe weather. However, variations in the CV of CI were greatest among juvenile mallards, particularly females (Table 5). Thus, this hypothesis does not provide a satisfactory explanation for these age- and sex-specific differences. If field-feeding situations pressure birds to maximize energy intake over time, and juvenile mallards are less efficient foragers than adults, variability in CV over the range of CI values should be higher among juveniles during times of food scarcity. Contrasts between adult males and juvenile females support the third hypothesis (Table 5), although the difference in the variability in CV between juvenile males and adult females is slight. Our final hypothesis is that competition causes differential foraging success by altering the habitat selection of subordinate individuals. Such intra-specific competition is thought to occur among wintering waterfowl (Nichols and Haramis 1980, Saylor and Afton 1981), particularly during periods of severe weather (Jorde et a1. 1984). During periods of good weather and non-limiting food supply, competition may be minimal and subordinate individuals are able to forage effectively. But when food becomes limiting, as occurs after a snowfall, competition increases and subordinate individuals may be excluded from preferred feeding sites. Paired individuals and adult males are the dominant birds in the social structure of wintering mallard flocks (Jorde 1981). Adult female black ducks (Anas rubripes) pair earlier in the winter than juvenile females (Stotts and Davis 1960). If a similar behavior exists among wintering mallards, then juvenile females probably are subordinate individuals throughout much of the winter. Thus, during periods of intense competition, dominant birds should be in better condition and have lower CV in CI than do juvenile females. Data presented here (Table 5) best fit the latter hypothesis, which implies that adult male mallards were not subject to the same variability in foraging success experienced by juvenile birds. Intra-specific competition, which resulted in juvenile females being displaced to suboptimal feeding areas, is one plausible explanation for these results. �43 LITERATURE CITED Ankney, C. D. 1974. The importance of nutrient reserves to breeding blue geese. Ph.D. Thesis, Univ. Western Ontario, London. 212pp. , and C. ------performance D. MacInnes. 1978. Nutrient reserves and reproductive of female lesser snow geese. Auk 95:459-470. Baldassarre, G. A., and E. G. Bolen. 1984. Field-feeding ecology of waterfowl wintering in the Southern High Plains of Texas. J. Wi1d1. Manage. 48:63-71. Be11rose, F. C. 1944. Bull. 23:327-372. Duck populations and kill. Illinois Nat. Hist. Surv. Bengston, S. A. 1971. Variations in clutch size in ducks in relation to food supply. Ibis 113:523-526. Bossenmaier, E. F., and W. H. Marshall. 1958. Field-feeding by waterfowl in southwestern Manitoba. Wi1d1. Monogr. 1. 32pp. Drobney, R. D. 1977. The feeding ecology, nutrition, and reproductive bioenergetics of wood ducks. Ph.D. Thesis, Univ. Missouri, Columbia. 170pp. Fretwell, S. D. 1972. Populations in a seasonal environment. Univ. Press, Princeton, N.J. Gromme, D. J. 1936. 53:324-325. Princeton Effect of extreme cold on ducks in Milwaukee Bay. Hochbaum, H. A. 1955. Travels and traditions of waterfowl. Minnesota Press, Minneapolis. 301pp. Jordan, J. S. 1953. Effects of starvation on wild mallards. Manage. 17:304-311. Auk Unlv. J. Wi1d1. Jorde, D. G. 1981. Winter and spring staging ecology of mallards in south central Nebraska. M.S. Thesis, Univ. North Dakota, Grand Forks. 116pp. _______ , G. L. Krapu, R. D. Crawford, and M. A. Hay. 1984. Effects of weather on habitat selection and behavior of mallards wintering Nebraska. Condor 86:258-265. Korschgen, C. E. 1977. Manage. 41:360-373. Krapu, G. L. 1974. Auk 91:278-290. Breeding stress of female eiders in Maine. in J. Wi1d1. Feeding ecology of pintail hens during reproduction. 1981. The role of nutrient reserves in mallard reproduction. Auk 98:29-38. �44 Lack, D. 1967. 18:125-128. The significance of clutch size in waterfowl. Wildfowl MacInnes, C. D., R. A. Davis, R. N. Jones, C. B. Lieff, and A. J. Pakulak. 1974. Reproductive efficiency of McConnell River small Canada geese. Wildl. Manage. 38:686-707. J. Milne, H. 1976. Body weights and carcass composition of the common eider. Wildfowl 27:115-122. Nichols, J. D., and G. M. Haramis. 1980. distribution patterns of canvasbacks. Sex-specific differences in winter Condor 82:406-416. Nie, N. H., C. H. Hull, J. G. Jenkins, K. Steinbrenner, and D. H. Bent. 1975. Statistical package for the social sciences. McGraw-Hill Book Co., New York, N.Y. 675pp. Pearson, T. G. 1934. Feeding wild ducks in a crisis. Bird-Lore 36:143. Raveling, D. G. 1979. The annual cycle of body composition of Canada geese with special reference to control of reproduction. Auk 96:234-252. Reinecke, K. J. 1977. The importance of freshwater invertebrates and female energy reserves for black ducks breeding in Maine. Ph.D. Thesis, Uni v. Maine, Orono. l13pp. Ringelman, J. K., and M. R. Szymczak. 1985. A physiological condition index for wintering mallards. J. Wildl. Manage. 49:564-568. Saylor, R. D., and A. D. Afton. 1981. Ecological aspects of common goldeneyes wintering in the upper Mississippi River. Ornis. Scand. 12:99-108. Stotts, V. D., and D. E. Davis. 1960. The black ducks in the Chesapeake Bay of Maryland: breeding behavior and biology. Chesapeake Sci. 1:127-154. Szymczak, M. R., and _J. F. Corey. Plains duck trap in Colorado. 1976. Construction and use of the Salt Colorado Div. Wildl., Div. Rep. 6. l3pp. Trautman, M. B., W. D. Bills, and E. R. Wickliff. 1939. Winter losses from starvation and exposure of waterfowl and upland game birds in Ohio and other northern states. Wilson Bull. 51:86-104. Whyte, R. J., and E. G. Bolen. 1984. composition. Condor 86:477-482. Impact of winter stress on mallard body Zar, J. H. 1974. Biostatistical analysis. Prentice-Hall, Inc., Englewood Cliffs, N.J. 620pp. Prepared b~~~ esK. Rige Wildlife Researcher �Colorado Division Wildlife Research April 1986 45 of Wildlife Report JOB PROGRESS REPORT Colorado State Project 01-00-045 (W-88-R) --------------------~------~----- Work Plan 1 ---- Job Title: 16 Field-feeding Ecology of Mallard Ducks Period Covered: Author: Job Avian Research 01 July 1984 through 30 June 1985 J. K. Ringe1man Personnel: J. F. Corey, J. K. Ringe1man, M. R. Szymczak, Colorado Division of Wildlife ABSTRACT Feeding trials with mallards (Anas p1atyrhynchos) were conducted on dirt substrate during December 1984 and March 1985. Trials in December were unsuccessful because ducks were unwilling to feed outside of the pen environment. March trials were successful and resulted in data sufficient to construct functional response curves for each age/sex class. In each case, feeding rates increased rapidly between corn densities of 0-100 km/ha, but remained nearly constant at densities > 400 kg/ha. With few significant differences in feeding rate apparent among age/sex classes, data were pooled to reveal a clearer response curve. In this case, feeding rate at 100 kg/ha < 300 kg/ha < 600 kg/ha = 800 kg/ha. Variability in feeding rates among individuals at the same corn density was large (CV = 21.6%, range = 6.6-51.5). At high corn densities, variability in feeding rates among individuals decreased (r = -0.76). These relationships suggest inherent differences in feeding efficiency among individual mallards which become more disparate at low corn densities. ��47 FIELD-FEEDING ECOLOGY OF MALLARD DUCKS James K. Ringe1man Man has drastically altered waterfowl habitat during the last 100 years. Whereas widespread drainage of inland wetlands and alteration of coastal areas have had detrimental effects, water development projects and irrigated agriculture have created new wintering and migration habitat. In Colorado, relatively few waterfowl were present in winter until irrigated cereal grain crops were introduced in the mid-1800's (Steine1 1926:174). Within a decade, waterfowl were being "short-stopped" along their traditional migratory routes, attracted by newly-created reservoirs and abundant waste cereal grain (Buller 1975). Wintering waterfowl, particularly the mallard, became so plentiful in the 1940's that a special season was held on mallards in Colorado during 1942-43 to help alleviate damage to grain crops (Wagar 1946). Mallards remain the most common wintering duck species today and accounted for 65% of the total duck harvest in 1981 (Colo. Div. Wi1d1. 1981). Wintering mallards, like all inland wintering waterfowl populations, are highly dependent on waste products of agriculture for food (Girard 1941, Reed 1971, Sugden et a1. 1974). In Colorado and adjacent states to the east and south, waste corn comprises from 50 to 90% of the winter diet (Jorde 1981, Gordon 1981, Baldassarre 1982). Despite the importance of corn to waterfowl, little is known of the factors that regulate its use by ducks. Harvester efficiency, slope of the land, and corn moisture content all determine the amount of corn remaining in the field after harvest (Baldassarre et a1. 1983). This "wastage" , typically 3-5% of the pre-harvest corn crop, contributes the bule of cereal grains necessary to sustain waterfowl throughout the winter. The amount of this wastage used by ducks varies depending on such post-harvest treatments as discing, plowing, burning, and cattle grazing (Jorde 1981, Baldassarre 1982). Economic forecasters predict radical changes in eastern Colorado irrigated agriculture within the next 7-17 years. Four factors will play important roles in this transformation: (1) depletion of ground water aquifers, (2) rising energy costs and related impacts on the cost of operating pumps, (3) conversion of farmland to urban use, especially along the Front Range, and (4) corn prices. Nearly 13% of all Colorado cropland is planted to irrigated corn and it ranks second behind wheat in crop value (Colo. Dep. Agric. 1982) • However, under scenarios of rising energy costs and groundwater depletion, corn is the first irrigated crop to become economically unfeasible (Sharp 1979). The break-even point is when electricity to power pumps reaches $0.12/kwh (Oamek 1981:64), and some scenarios predict the disappearance of irrigated corn from the region overlying the Ogallala Aquifer by 1990 (Young et a1. 1982:129). Front Range area croplands, which benefit from gravity flow irrigation water transported by canals, are under increasing pressure from urban growth. In the last two decades, Colorado has seen over 1.3 million acres converted from agricultural use (Colo. Dep. of Agric. 1980). Much of this development occurs on irrigated land (U.S. Dep. of Agric. 1978). �48 The key to preserving wintering waterfowl and subsequent harvest opportunity is the management of water areas coupled with more intensive land management practices, particularly in relation to cornfields on Division of Wildlife and private lands. Based on historical evidence and knowledge of wintering waterfowl food habits, it is believed that loss of cornfields will result in major shifts of birds out of an area. In the case of the Front Range, this would result in severe loss of hunting opportunity where demand is greatest. This would also be true elsewhere in eastern Colorado where hunting pressure and demand is somewhat less pronounced, but nevertheless important. 50 little is known of how mallards use cornfields and waste grain that it is not possible to determine the area of cornfields needed to sustain a given mallard population. In the past, such information was unneeded because of abundant corn crops. However, with the expected declines in irrigated agriculture in the near future, these data will be necessary to maintain wintering waterfowl. Past field-feeding studies have focused on the timing of flights and the relationships between flight times and weather (Hochbaum1955, 5winebroad 1956, Bossenmaier and Marshall 1958, Winner 1959, Gordon 1981, Jorde 1981), or on ways to alleviate crop damage by ducks (Go110p 1950, Hammond1952, MacLennan 1973). Only recently have studies documented the potential value of post-harvest treatments in making fields more attractive to ducks. Jorde (1981:50) described a commensal relationship between field-feeding mallards and cattle which exposed corn in times of heavy snowfall. Baldassarre et a1. (1983) reported that treatments such as burning or discing of fields with moderate amounts of waste corn made these fields more attractive to ducks than untreated cornfields of standing stubble with waste corn densities many times greater. This suggests an "optimal foraging" strategy of field-feeding ducks in which feeding rate is balanced with other considerations such as search time, handling time, and distance to field. A large body of information on optimal foraging in birds exists in the ecological literature, but thus far these findings have not been applied to the management of field-feeding ducks. Efforts must be made to evaluate methods of making waste corn more available to waterfowl, thereby optimizing feeding rates and winter physiological condition. P.N. OBJECTIVES 1. Determine the relationships corn density, group foraging 2. Relate 3. Document the foraging habitat 4. Develop a model for mallards. post-harvest post-harvest cornfield between feeding rates of mallards, size, sex, and age of duck. treatments requirements to foraging efficiency. of wintering management of and waste mallards. cornfields for wintering �49 SEGMENT OBJECTIVES 1. In late summer, hand-pick and remove ears from 1 ha of corn. 2. In fall, harvest the cornfield using standard techniques leaving canes, leaf litter, and stubble in place. 3. During November-February, conduct feeding trials with mallard ducks. Dose hand-picked study plots at corn densities of 25, 50, 100, 200, 400, 600, and 800 kg/ha. 4. Derive a functional response curve for mallards feeding on waste grain at each density. Compare with foraging rates obtained in "black dirt" experiments. Particion out differences in foraging rates based on search time. 5. Compile and analyze data and prepare progress report. METHODS Eight rows of corn, 8 m wide and about 500 m long, were hand-picked on 27 August 1984. Stalks were allowed to mature, then a harvester was run through the entire field (including the hand-picked segment) in early fall. Since this hand-picked segment was devoid of waste corn, it could then be dosed with a known amount of corn to achieve the desired waste corn density. Feeding trials were conducted in mid-December 1984 and again in mid-March 1985. December trials were conducted on plowed then finely cultivated dirt. Panels measuring about 1.2 by 2.0 m (constructed of poultry netting stretched over a metal conduit frame) were used to build square feeding trial enclosures 8 m on a side. Whole kernel corn was broadcast uniformly wi thin .this 64-m2 enclosure using a hand operated seed spreader. Forty bird~ (10 in each age/sex class) were weighed immediately prior to a feeding trial, released en masse into the seeded enclosure at the start of the trial, then interrupted in their feeding and removed from the enclosure after a short feeding interval. Ducks were then reweighed immediately, and the difference in weight before and after the feeding trial was assumed equal to the corn consumed during the trial. March feeding trials followed the same procedure, except that trials were conducted on a dirt and pea gravel surface inside a partition within the primary holding pen. This enclosure measured 3 by 21.3 m and had a surface area (64 m2) equivalent to the enclosures used in December. Whole kernel corn was broadcast at densities of 100, 200, 300, 400, 600, and 800 kg/ha. Duplicate trials were conducted at the 3030 and 800 kg/ha densities. Data on weight gain was maintained on an individual bird basis according to identifying numbers stamped on leg bands of all birds. Analysis of variance procedures were used to examine age, sex, and corn densityspecific differences in foraging rates, as measured by grams corn ingested per minute. Because the objective was to ascertain maximum foraging rates (i.e., "black dirt" with no leaf or stalk litter), only birds with the five highest foraging rates within each age/sex class were used in data analyses. �50 Because of unsatisfactory feeding performance of birds outside of their main pen facility, it was necessary to hatch and rear new ducks for use in winter 1985 feeding trials. A new imprinting and training routine was used for conditioning these birds. After hatching, ducklings were imprinted to humans by nearly constant associated during the first 48 hours after hatching and several hours of handling daily for several weeks thereafter. Seven males and seven females were retained as experimental birds after achieving adult body size. A conditioning regime was established wherein experimental birds were rounded up inside the main pen, transported a short distance by truck to a feeding "corral" 20-25 m2 in size, released into this corral to feed, then transported back to the main holding pen. Birds were fed only outside of the holding pen, and every 3-4 days were handled and placed into a mesh net weighing sack to simulate the weigh-in procedure. This newest group of experimental birds will be used for feeding trials during winter 1985-86. RESULTS AND DISCUSSION Attempts to conduct feeding trials during December 1984 met with failure. Experimental birds, having been reared and fed daily only inside their main holding pen, refused to feed outside of their pen environment. Feeding trials held in mid-March within a partition of hte main holding pen were successful. Birds fed vigorously and usually consumed nearly all corn within 1-2 minutes of their release. Variation in feeding rates was considerable, even among birds 'with the five highest feeding rates. Coefficients of variation (CV) averaged 21.6% and ranged from 6.6 to 51.5%. This highly variable foraging success among individuals, particularly in a simple environment devoid of leaf and stem litter, suggests that variation in feeding rates and consequent fat reserves of wild birds may reflect inherit differences in feeding ability among individual mallards. As hypothesized, the variability in feeding rates decreased at higher corn densities (r = -0.76, df = 4, P < 0.05). This indicates that inherit differences -in feeding rates b;tween fast and slow-feeding birds are lessened at high waste corn densities, probably because slow-feeding birds experience a relatively higher increae in feeding efficiency due to a superabundance of corn. Response curves of feeding rate as a function of corn density were similar among all age/sex groups (Figs. 1-4). Feeding rates increased rapidly at corn densities between 0 and 100 kg/ha, but were similar at densities above 4-kg/ha. Statistically significant differences in feeding rates were difficult to detect due to small sample sizes and high variability in individual feeding rates. Replicate feeding trials at 300 and 800 kg/ha revealed that mean feeding rates differed by replication for both 300 kg/ha (9.86 g/min vs. 11.88 g/min; F = 11.88, P = 0.001) and 800 kg/ha densities (12.22 g/min vs. 15.20 g/min; F = 10.74, p= 0.002). Thus, several replicate feeding trials at each densitymay be necessary to provide precise estimates of mean feeding rates. Several factors, including weather and hunger level of the experimental birds, may be responsible for differences in feeding rates between days. �51 20 • 0 e0 8 16 oB 0 Bo c E .••. Cl 0 12 • 0 ~ W •... • i • 0 0 OB 8 0 0 0 -e 0 a: C) z c • 8 w W IL 0 4 0 ADULT MALE O~--------r--------'---------r--------~ o 200 400 600 800 CORN DENSITY (kg/ha) Fig. 1. The relationship between corn density and feeding rate for adult male mallards for feeding trials conducted on dirt substrate. Open circles represent feeding rates of individual birds. Open squares equal mean feeding rates. Solid circles are individual feeding rates for a replicate feeding trial. Means with the same letter do not differ q. > 0.05) • �52 20 0 • • 0 16 0 §C 0 0 B,Co - - oB,C i • 0 I 8 0 0 c E Cl ..... • 0 0 12 8 w t< 0 0 o 0 a: CJ z c w w 8 ~ 4 JUVENILE MALE O+---------~--------~--------~--------~ o 200 400 600 800 CORN DENSITY (kg/ha) Fig. 2. The relationship between corn density and feeding rate for juvenile male mallards for feeding trials conducted on dirt substrate. Open circles represent feeding rates of individual birds. Opensquares equal meanfeeding rates. Solid circles are individual feeding rates for a replicate feeding trial. Meanswith the sameletter do not differ (! > 0.05). �53 20 0 • 0 16 0 - 0 c E .•.• ...•CI 12 8 A,Bi 0 • oA,B -e a:: 0 <!J 9 0 UJ UJ 8B 0 UJ t- z •• 8 • 9 SA • 0 0 0 0 0 U. ~ 4 ADUL T FEMALE O~--------~--------~---------r---------' 800 600 400 200 o CORN DENSITY (kg/ha) Fig. 3. The relationship between corn density and feeding rate for adult female mallards for feeding trials conducted on dirt substrate. Open circles represent feeding rates of individual birds. Open squares equal mean feeding rates. Solid circles are individual feeding rates for a replicate feeding trial. Means with the same letter do not differ (!> 0.05). �54 20 0 • •" Cc • 16 cB,C 0 ,...c: ....E0) - 0 0 0 i I- -e a: CA (!) 0 z w w • 12 w c • 8 8 8 0 0 0 0 0 cA 0 0 0 0 0 0 0 ,LL 0 4 JUVENILE FEMALE O+---------~------~--------_r------~ o 200 400 CORN DENSITY 600 800 (kg/ha) Fig. 4. The relationship between corn density and feeding rate for juvenile female mallards for feeding trials conducted on dirt substrate. Open circles represent feeding rates of individual birds. Open squares equal mean feeding rates. Solid circles are individual feeding rates for a replicate feeding trial. Means with the same letter do not differ (!> 0.05). �55 In only one case was a significant difference in feeding rate apparent among agelsex groups (Table 1), indicating that feeding rate data might best be pooled across age/sex groups within corn density. Combining data in this manner gives a clearer picture of the functional response curve and more efficient statistical separation of mean feeding rate by density (Fig. 5). In this combined case, feeding rate is less at 100 kg/ha than at 300 kg/ha, which in turn is less than the 600 and 800 kg/ha feeding rates (f < 0.05). Table 1. Differences in mean feeding rates of mallards in each of four age/sex classes in relation to waste corn density. Abbreviations are: AM (adult male), JM (juvenile male), AF (adult female), JF (juvenile female). Sample size is five birds in each age/sex class. Corn density (kg/ha) 100 200 300 400 600 800 Relationship of mean feeding rate among age/sex class AM = AM = AM = AF = AM = AF~ JM = JM = Jt1 = JF < JM = AM = AF = AF = AF = AM = AF = JM~ JF JF JF JM JF JF F = 5.58 2.54 Significance NS NS NS 0.008 NS 0.09 Future feeding trials will feature birds in a cornfield environment, thereby adding search time to search and handling time. It is anticipated that the resulting functional response curve will be more linear than logistic in shape because of the relatively constant search time involved at even the highest waste corn densities. The difference between the functional response curve derived from cornfield trials and the curves presented here will serve as a quantitative measure of search time. It is this search time caused by ground litter which, in combination with waste corn density, can be manipulated to maximize feeding rates of wild waterfowl. LITERATURE CITED Baldassarre, G. A. 1982. Field-feeding ecology of waterfowl wintering on the southern high plains of Texas. Unpublished manuscript. 20pp. , R. ---quality J. Whyte, E. E. Quinlan, and E. G. Bolen. 1983. Dynamics and of waste corn available to postbreeding waterfowl in Texas. Wildl. Soc. Bull. 11:25-31. Bossenmaier, E. F., and W. H. Marshall. 1958. Field-feeding by waterfowl in southwestern Manitoba. Wildl. Monogr. 1. 32pp. Buller, R. J. 1975. Redistribution of waterfowl: influence of water, protection, and feed. Proc. Int. Waterfowl Symp. 1:143-154. Colorado Department of Agriculture. 1980. Agricultural land conversion in Colorado. Resour. Analysis Sect., Colorado Dep. Agric., Denver. 8pp. �56 20 16 •..•e ....eCJ) - 12 °A.B w ~ < a: CJ z c 8 w W LL. 4 O+---------,_--------~--------_r--------_, 400 600 o 200 800 CORN DENSITY (kg/ha) Fig. 5. Response curve of feeding rate as a function of corn density for all age/sex groups combined. Means with the same letter do not differ (~ > 0.05). �57 1982. Colorado agricultural statistics. Rep. Serv., Denver Bull. 1-82. 94pp. Colorado Division of Wildlife. varmint harvest, 1981. Denver. 228pp. Colo. Crop and Livstock 1981. Colorado small game, furbearer and Wildl. Servo Sect., Colorado Div. Wildl., Girard, G. L. 1941. The mallard: Wildl. Manage. 5:233-259. its management in western Montana. J. Gollop, B. J. 1950. Report on investigation of damage to cereal crops by ducks in the prairie provinces. Unpubl. Rep., Can. Wildl. Serv., Ottawa. llpp. Gordon, D. H. 1981. Condition, feeding ecology, and behavior of mallards wintering in northcentral Oklahoma. M.S. Thesis, Oklahoma State Univ., Stillwater. 68pp. Hammond, M. C. 1952. Waterfowl damage and control measures, Lower Souris Refuge and vicinity. Unpubl. Rep., U.S. Dep , Inter., Fish and Wildl. Serv., Washington, D.C. 5pp. Hochbaum, H. A. 1955. Travels and traditions of waterfowl. Minnesota Press, Minneapolis. 301pp. Unlv. Jorde, D. G. 1981. Winter and spring staging ecology of mallards in southcentra1 Nebraska. M.S. Thesis, Univ. North Dakota, Grand 116pp. Forks. MacLennan, R. 1973. A study of waterfowl crop depredation in Saskatchewan. Can. Wi1dl. Serv., Saskatoon, Saskatchewan. Wi1d1. Rep. 2. 38pp. Oamek, G. E. 1981. Economic adjustments to rising energy costs: the case for pump irrigators. M.S. Thesis, Colorado State Univ., Fort Collins. 90pp. Reed, L. W. 1971. Use of western Lake Erie by migratory and wintering waterfowl. M.S. Thesis, Michigan State Univ., East Lansing. 71pp. Sharp, R. L. 1979. Economic adjustments to increasing energy costs for pump irrigation in northeastern Colorado. M.S. Thesis, Colorado State Univ., Fort Collins. Steinel, A. T. 1926. History of agriculture in Colorado. CoIl., Fort Collins, Colo. 659pp. State Agricultural Sugden, L. G., W. J. Thurlow, R. D. Harris, and K. Vermeer. 1974. Investigations of mallards overwintering at Calgary, Alberta. Field-Nat. 88:303-311. Can. Swinebroad, J. 1956. Some aspects of the role of weather in bird migration. Ph.D. Diss., Ohio State Univ. 315pp. �58 u.s. Department of Agriculture. 1978. Urbanization of rural lands in the northern Colorado Front Range. U.S. Dep. Agric., Washington, D.C. 23pp. Wagar, J. V. K. 1946. Colorado's duck-damage, grain-crop problem. Am. Wildl. Conf. 11:156-162. Proc. No. Winner, R. W. 1959. Field-feeding periodicity of black and mallard ducks. J. Wildl. Manage. 23:197-202. Young, R. A., L. R. Conklin, R. A. Longenbaugh, and R. L. Gardner. 1982. Energy and water scarcity and the irrigated agricultural economy of the Colorado high plains: direct economic and hydrologic impact forecasts (1979-2020). Colorado Water Resour. Res. lnst., Colorado State Univ., Fort Collins. 362pp. Prepared ~j(.~ JeSK. Rge Wildlife Researcher �Colorado Division Wildlife Research April 1986 59 of Wildlife Report JOB PROGRESS REPORT Colorado State Project Work Plan Job Title: 01-00-045 (W-37-R) 1 Job Avian Research 23 Evaluation of No-till Wheat Farming Period Covered: 1 January through 31 December 1984 Author: Warren D. Snyder Personnel: T. Lytle, D. Clippinger, T. Davis, M. Telck, R. Towry, D. We yerman , M. Pinkham, W. Snyder, Colorado Division of Wildlife. ABSTRACT Efforts were coordinated with Division of Wildlife management personnel to. initiate experimental no-till biennially-cropped winter wheat farming on the South Republican Wildlife Area in 1984. Slightly above average precipitation in 1982-83 resulted in good to excellent quality wheat stubble available as nesting cover on no-till summer-fallowed tracts in 1984. Annual precipitation in 1984 was above average but averaged below normal through the growing season. The height-density index of stubble quality averaged 0.977 dm with considerable variation among the eight fields. Analyses of pH, percent organic matter, available nutrients, and other characteristics were obtained along with limited monitoring of soil temperatures and soil moisture. Assistance was provided to management personnel in herbicide treatments and fall seeding of winter wheat. ��61 EVALUATION OF NO-TILL WHEAT FARMING Warren D. Snyder The importance of wheat stubble and green wheat as well as their insecurity as nesting covers for ring-necked pheasants (Phasianus colchicus) in the High Plains of eastern Colorado was recently documented (Snyder 1984). Management biologists within the Colorado Division of Wildlife have expressed an interest in initiating a new approach, no-till summer fallowing on CDOW properties in eastern Colorado. No-till summer fallow, applied to biennially-cropped winter wheat, potentially would increase the quantity, quality, and security of nesting cover while reducing problems with soil erosion, snow drift, and lessees while saving fuel, equipment, and labor. Since increased nesting use of habitats is directly related to their increased height-density quality (Snyder 1984), the purpose of this evaluation study is to document relative quantity and quality of cover in nO-J:ill fields and other available nesting habitats. Major equipment deficiencies, personnel changes, and inexperience in herbicide appplication hampered this first-year effort. However, the knowledge gained will improve the success of future efforts. P. N. OBJECTIVES The objective of this study is to test a no-till wheat-fallow biennial-cropping system for increasing the quality, quantity, and security of nesting cover for ground nesting wildlife on Colorado Division of Wildlife properties in eastern Colorado. SEGMENT OBJECTIVES 1. Review literature and contact agronomists knowledgeable about wheat farming. no-till 2. Coordinate site selection and application of treatments with management habitat biologists and property technicians. 3. Monitor environmental and nesting cover conditions including: a. Precipitation data (obtained from nearby weather stations). b. Soil moisture conditions within treatment and control fields at time of wheat planting in fall. Soil temperatures at a depth of 10 cm will be made in treatment and control fallow fields on 1 June and at time of wheat planting. c. Visual obstruction measurements will be obtained in March or April in wheat stubble treatment and control fields and at prescribed times in green wheat and perennial herbaceous cover tracts. d. Timing of spring tillage of wheat stubble fields (controls) will be recorded. �62 e. Timing of wheat harvest on treatment and control tracts will be monitored along with comparisons of wheat yields and effectiveness of herbicides. f. Activity signs indicating presence of monitored in mid-summer and early fall. rodents will be visually 4. A job progress report will be prepared. 5. Study information and progress will be communicated to Division of Wildlife personnel and to farmers through the Landowner Relations Program. STUDY AREA AND METHODS Background information concerning no-till, equipment, and herbicides were collected from the literature, personal contacts, and field demonstrations and disseminated to management personnel. Selection of study sites and fields was coordinated with management personnel who were responsible for application of herbicides on no-till fields and for farming conventionally-fallowed fields. Monitoring of environmental conditions included obtaining precipitation records from the weather station at Bonny Reservoir Dam. An attempt was made to monitor soil moisture conditions using a soil moisture meter, however, readings were not considered reliable and the effort was terminated. Based on recommendations from D. Smika, USDA Agriculture Research Service, a soil probe was acquired and preliminary measurements of soil moisture depth were obtained in fall. Further use of this device will depend upon evaluations of its reliability. Time constraints prevented test plot selection and extensive monitoring of residual and green vegetation height-density in early spring 1984. Limited measurements, primarily of no-till stubble, were obtained in mid-May using the Robel method (Robel et aL 1970) of visual obstruction measurement. Cover height measurements were also collected on most tracts. RESULTS AND DISCUSSION Coordination and Site Selection Literature pertinent to conservation tillage, especially no-till wheat farming, was acquired. Details on specific items were obtained by phone and by direct contacts with Dr. Daryl Smika, USDA Agronomist, stationed at Akron, Colorado. An October 1984 meeting between interested CDOW management personnel and Dr. Smika was attended. Information exchanges among CDOW personnel including T. Lytle, R. Towry, T. Davis, and myself occurred throughout the year. T. Davis and I attended a field demonstration day to observe use of no-till equipment and to hear discussion of its features and operation. Based on coordination with T. Lytle, Senior Habitat Biologist and R. D. Clippinger, Area Wildlife Manager, eight test tracts within four fields were �63 selected for treatment (Fig. 1) within the South Republican Wildlife Area below Bonny Reservoir. Adjacent tracts to be conventionally fallowed (using tillage) were retained within the fields. A partial vacancy over summer because of changes in technicians on the property resulted in an inconsistent fallow treatment of conventionally-tilled fields. Initial tillage was delayed until mid-summer resulting in substantial soil moisture loss to annual weeds. Therefore, wheat growth among no-till and tilled tracts cannot be directly compared through the 1985 growing season. Environmental Measurements Precipitation.--Precipitation in 1983 averaged slightly above normal (Table 1). Successive snowfalls were received from January through April 1984 yielding above average moisture that substantially increased soil moisture levels. However, May through Septemer rainfall was consistently below average (Table 1) and upper soil levels in conventionally-fallowed fields were dry at wheat planting in September. Successive rains in October stimulated new wheat growth and resulted in annual precipitation being considerably above average in 1984. Table 1. Monthly and annual precipitation recorded at Bonny Reservoir in 1983 and 1984 vs. the long-term mean.a 1983 Month Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Totals Inches 1984 Cm Inches Cm Lons term Inches .x. Cm 0.08 0.56 2.05 2.77 4.30 2.44 2.51 0.59 0.21 1.08 1.44 0.29 0.20 1.42 5.21 7.04 10.92 6.20 6.37 1.50 0.53 2.74 3.66 0.74 0.74 1. 75 2.25 4.52 2.19 1.90 1.76 1.02 0.76 3.10 0.15 0.72 1.88 4.44 5.72 11.48 5.56 4.83 4.47 2.59 1.93 7.87 0.38 1.83 0.31 0.33 0.88 1.45 3.03 2.60 2.49 2.14 1.38 0.88 0.55 0.31 0.79 0.84 2.24 3.68 7.70 6.60 6.32 5.44 3.50 2.23 1.40 0.79 18.32 46.53 20.86 52.98 16.35 41.53 aLong-term monthly and annual means obtained from U.S. Weather Service Climatological Data 1983 Annual Summary. Vegetation Sampling.--The height density index (HDI) among the eight no-till tracts averaged 0.977 with considerable variation among fields. HDI' s of 1.3-1.6 in the west tracts (Field 1) provided good to excellent nesting cover for pheasants and were associated with stubble heights of 47.3-55.2 cm (Table 2). Stubble HDI approximated or exceeded most indices obtained during the 1979-81 study on and around the Sand Draw Property in extreme northeast Colorado (Snyder 1984). �o.j> !' J 0' W!·III \ \ \.~ ,~ \\~'~.~--:~O~ " .'-, \\ , " ,\ . U\1 H)~~ .--"-",~ ~< ~l~ ". ~~ ," ~~~ \~, o ( c :» ':,' "\. ) \~~~ " " ,': ,.:.'. " ~,~ ~ \': . : \! ' •••• \. 'I, i ,,', _...._ ~_...'_~._t .--~ ~~'iY:~\\: .._;< .~-,~~.."U';' 1\ 1\ " -uI I ~- ti ~I No-till \ \.,.'; \.l \ • \":~ ", f'I ,'" ..~/' • ' s.,__ "'x '=1 ,. \ rv _.L~"\.--·":: 1. \ { t \~\ ~_:' ", Fig. 'I " '\1 '~','. ,,,-! !(r=::I~::II: ,I {!. I .... ~ .a '\\ ,- ~ field locations within the South Republican Wildlife Area, 1984. ,. �65 Table 2. Height-density (dm) and height (cm) of residual and/or green vegetation within no-till stubble fields and other cover types, South Republican Wildlife Area, Spring 1984. Cover Treatment Tract Field Date Sample size HDI Wheat stubble No-till North South 1-84 1-84 1-84 23 May 15 May 15 May 100 100 16 1.60 1.33 2.00 26 25 55.20 47.35 No-till Middle 2-84 15 May 140 0.62 28 39.09 Convene No-till South Middle North South Middle North 3-84 3-84 3-84 4-84 4-84 4-84 15 15 23 23 16 16 68 72 100 100 100 100 0.58 0.66 0.47 0.96 1.27 0.96 39 25 25 39 35 48.57 46.13 45.72 49.43 46.40 812 0.98 Convene Barley stubble Wheat No-till No-till May May May May May May 8 tracts combined gr/ forbs Sample size Height 47.37 Annu, N Bishop House 23 May 60 0.36 16 14.90 W of Field 4 23 May 60 0.87 17 19.42 SE of Field 1 23 May 60 0.60 Switchgrass/ sand lovegrass S of Field 2 23 May 60 0.74 Alfalfa E of Field 1 23 May 24 3.73 8 47.94 Green wheat Between fields 3/4 23 May 60 4.64 21 60.72 Height-density samples were obtained from other nearby cover types within the South Republican Wildlife Property on 23 May 1984. At that time rapidly growing wheat and alfalfa yielded much higher indices than wheat stubble whereas unfarmed sites containing mixtures of annuals and perennials yielded lower HDI's (Table 2). Time constraints prevented more extensive spring measurements which would have illustrated the growth pattern of different covers available for use by ground-nesting wildlife. Soil Characteristics.--Retention of standing wheat stubble during summer fallow potentially lowers soil temperatures during spring and summer and increases soil moisture retention in contrast to conventional fallow. Forenoon to early P.M. soil temperatures on 30 May 1984 at approximately 10 cm below the soil surface averaged 65.2 F in no-till stubble and 72.1 in nearby fallow fields (! < 0.05, .! = 7.02, df = 18). Soil temperatures on 13 September under cloudy conditions at time of initial wheat seeding averaged 75 F in both no-till and tilled wheat fields. �66 Soil moisture was observably much closer to the soil surface, especially under a mulch layer, in the firm soil of no-till fields. Top soils were usually loose and dry for several centimeters within conventionally-fallowed fields at time of wheat planting. A 1.2 - 1.8-m long steel rod (approx. 0.5 em diameter) is the tool used by agronomists and farmers to obtain an estimate of soil moisture (D. Smika, pers, commun.). Reportedly, the rod when inserted vertically and vigorously into the soil will penetrate to within about 0.15 m of the lowest level of soil moisture. Preliminary testing in sandy rangeland soils indicated this statement is true. D. Smika noted that silt-loam soil will retain about 5.1 cm moisture/0.3 m (2 in./ft) of soil whereas sandy soils retain less moisture (2.5 - 3.8 cm/0.3 m; 1-1.5 in./ft). A soil probe was acquired and used to obtain 30 measurements within four no-till fields and 24 measurements within four conventionally-fallowed fields within the study area on 12 October 1984. Recent, October rains (Table 1) had improved soil moisture conditions in all fields. Analysis comparing the results of soil probe penetrations within no-till and tilled wheat fields showed the average penetration in no-till fields was 0.85 m (33.5 in.) and that in tilled fields was 0.67 m (26.5 in.) (P < 0.05, t = 2.77, df = 52) providing evidence of greater moisture content in no-till fallowed fields. The greatest penetration was 1.4 m in a no-till tract. Scattered large annuals consumed moisture potentially increasing variability among random sites. Soil samples obtained from three no-till fields within the study area in fall 1984 were submitted to the Colorado State University Soil Testing Laboratory for routine analysis. Results (Table 3) show all soils were nearly neutral in pH, predominately sandy loam in texture, low in organic matter, and low in nitrogen and phosphorus essential for dryland crop production. Soil pH, organic matter, and texture are important variables when selecting and applying herbicides (Smika and Sherman 1982). Table 3. Characteristics of soils from no-till wheat fields, South Republican Wildlife Area, Fall 1984. Characteristic Soil pH Salts Lime Organic matter (O.M.) Nitrogen (NO s-N) Phosphorus (P) Potassium (K) Zinc (Zn) Iron (Fe) Manganese (Mn) Copper (Cu) Texture Field 1 Field 2 Field 3 7.4 0.2 Low 1.3 7 6.4 381 0.9 4.4 2.7 0.9 Silt loam 6.6 0.2 Low 1.4 12 9.4 405 0.6 10.7 4.4 0.7 Silt loam 6.7 0.3 Low 1.0 21 2.8 385 0.3 5.7 1.9 0.8 Silty clay loam �67 Herbicide Treatments Assistance was provided to management personnel in selection and application of herbicides to the no-till tracts. Time and monetary constraints prevented acquisition of persistent herbicides for spring application. Thus, glyphosate (Roundup) was the only herbicide applied on 17-18 May 1984 to approximately 19.39 ha (48 ac ,) at the rate of 0.32 kg/ha (0.29 Lbs, active ingredient/acre). Difficulties in attaining adequate sprayer pressure apparently led to cali'bration problems so that the recommended rate of application, 0.42 kg/ha (0.37 lbs./ac.) was not attained. In addition, spraying efforts were rained out and additional rainfall was received the evening following application. A 30 May inspection showed that the contact herbicide was effective in killing all downy brome (Bromus tectorum) and volunteer wheat but did not completely kill annual mustard (Brassica sp.) or wild lettuce (Lactuca sp.). A second treatment applied on 9 July 1984 included a tank mix of glyphosate and dicamba (Banvel) applied with a surfactant at respective rates of 0.42 kg/ha and 0.56 kg/ha. Again, as in spring, low sprayer pressure was a major problem and an evening rain undoubtedly reduced herbicide effectiveness. Wild lettuce was growing rapidly and had attained heights up to 0.3 m making it difficult to kill at the time of the July treatment. It remained partially alive, but stunted through the remainder of the summer and probably consumed little additional soil moisture. Apparently, because of low sprayer pressure and resultant large droplet size and poor coverage, some young annuals including Russian thistle (Salsoli kali) and witchgrass (Panicum capillare) were not killed by the July treatment and continued to grow in sparse stands. The thistles, while sparse, attained considerable growth by late summer increasing wheat planting difficulties. Thus, the herbicide treatments were only marginally effective, but it is uncertain whether the primary problems were with timing, low sprayer pressure, conflicts with rainfall, or other variables. It is probable that all were factors. It is recommended that a short residual « 90 day) herbicide be included in future spring applications. Both herbicides used in 1984 were of low toxicity and registered for use on summer fallow. In anticipation of no-till replication through 1985, the moderately persistent herbicide atrazine was applied to 8 ha of new wheat stubble on 10 August 1984. The application rate was 1.1 kg/ha (1 lb./ac.). Success of this post harvest application was dependent upon receipt of rainfall within 1-2 weeks to place it into the top soil. Adequate rainfall was not received subsequent to the treatment so it is assumed the treatment was unsuccessful. Both fields contained considerable volunteer wheat when inspected 12 October 1984. Wheat Seeding A small "Tye" brand no-till drill was borrowed from another CDOW property and initial wheat seeding efforts were initiated on 13 September. This drill lacked coulters needed to cut through residual vegetation and therefore did not penetrate the ground sufficiently to achieve satisfactory stands of wheat. Through the efforts of T. Davis, two other brands of demonstration drills were acquired for temporary use and were tested on the no-till tracts. Both performed satisfactorily, however, due to work conflicts among personnel only partial seeding was completed. �68 Early October inspection of the seeded tracts, following rains indicated good stands of wheat were attained on all tracts where planted, including those where the Tye drill was used. Results of this first year demonstrate that if no-till farming is to yield satisfactory results, proper equipment, maintained in good working condition and properly calibrated is essential. Increased knowledge of herbicides, their effectiveness on different weed species, timing of applIcation, and related soil factors are also necessary. Additional knowledge will be gained through field experience. LITERATURE CITED Robel, R. J., J. N. Briggs, A. D. Dayton, and L. C. Hulbert. 1970. Relationships between visual obstruction measurements and grassland vegetation. J. Range Manage. 23:295-297. weight of Smika, D. E., and E. D. Sherman. 1982. Atrazine carryover and its soil factor relationships to no-tillage and minimum tillage fallow-winter wheat cropping in the central Great Plains. Colorado State Univ. Exp , St.n , Tech. Bull. 144. 4pp. Snyder, W. D. 1984. Ring-necked pheasant nesting ecology and wheat farming on the High Plains. J. Wildl. Manage. 48:878-888. �69 Colorado Division of Wildlife Wildlife Research Report April 1986 JOB PROGRESS REPORT State of Project Work Plan Job Title: Colorado 01-03-045 1 Avian Research : Job ~2~3~ Evaluation Period Covered: (W-37-R) of No-till Wheat Farming 01 January through 31 December 1985 Author: Warren Snyder Personnel: C. Braun, T. Davis, M. Telck, and W. Snyder, Colorado Division Wildlife of ABSTRACT Several factors associated with conventional tillage and no-till farming of winter wheat in 1984 in combination with severe weed competition in 1985 made it impossible to compare the 2 procedures on the South Republican Wildlife Area. Spring 1985 was phenologically early prompting early wheat growth that surpassed wheat stubble in height-density quality on about 1 May. However, HDI values of wheat stubble averaged 1.24 dm indicating it too provided good nesting cover which was better than that in upland perennial grasses, alfalfa, and sites dominated by annuals into mid-May 1985. Thus, if all wheat stubble was no-till fallowed and available for nesting through spring and summer it could potentially have significant impact on pheasant production. Early 1985 weed growth prompted a mid-May herbicide treatment within a l2.l-ha tract of wheat stubble. Satisfactory weed control was provided throughout the summer from this single ~reatment, however the tract was not planted to wheat in fall 1985 nor was any land conventionally fallowed or planted. Therefore, efforts to evaluate no-till on the South Republican Wildlife Area had to be terminated. Information concerning biennially cropped no-till farming of winter wheat and wildlife relationships was disseminated to the farming public within the Central High Plains region. ��71 EVALUATION OF NO-TILL WHEAT FARMING Warren D. Snyder P. N. OBJECTIVE The objective of this study is to test a no-till wheat-fallow biennia1cropping system for increasing the quality, quantity, and security of nesting cover for ground nesting wildlife on Colorado Division of Wildlife properties in eastern Colorado. SEGMENT OBJECTIVES 1. Coordinate site selection and application of treatments with management habitat biologists and property technicians. 2. Monitor environmental and nesting cover conditions including: a. Precipitation data (obtained from a nearby weather station). b. Soil moisture conditions within treatment and control fields at time of wheat planting in fall. Soil temperatures at a depth of 10 cm will be made within treatment and control fallow fields on 1 June and at a time of wheat planting. c. Visual obstruction measurements will be obtained in March or April in wheat stubble treatment and control fields and at prescribed times in green wheat and perennial herbaceous cover tracts. d. Timing of spring tillage of wheat stubble fields (controls) will be recorded. e. Timing of wheat harvest on treatment and control tracts will be monitored along with comparisons of wheat yields and effectiveness of herbicides. f. Activity signs indicating presence of rodents will be visually monitored in mid-summer and early fall. 3. A job progress report will be prepared. 4. Study information and progress will be communicated to Division of Wildlife personnel and to farmers through the Landowner Relations Program and through other farm media. RESULTS AND DISCUSSION Coordination, Site Selection, and Treatment Eight test "no-till" tracts within 4 fields were selected for treatment in 1984 within the South Republication Wildlife Area below Bonny Reservoir based �72 on coordination with management personnel of the Southeast Region (Snyder 1985a). Adjacent tracts to be conventionally fallowed (using tillage) were retaIned within the fields. Because of personnel changes of property technicians on the site, initial tillage was delayed until mid-summer 1984 resulting in substantial soil moisture loss to annuals and loose dry topsoil for fall seeding. At the same time initial no-till herbicide treatments were only partially effective in weed control because of lack of equipment, lack of persistent herbicides to use, and lack of knowledge concerning their application. Three brands of no-till grain drills were tried in fall planting and 2 of these provided satisfactory seeding results in the heavy residual stubble. However, time constraints apparently prevented the property technician from completing wheat planting on several of the tracts in fall 1984. Satisfactory wheat stands were attained on planted no-till fields. Moisture conditions in fall and early spring stimulated dense stands of wild lettuce (Lactuca sp.) and downy brome (Bromus tectorum) within both no-till and conventionally fallowed wheat fields. A 2,4-D herbicide treatment in spring would have probably corrected this problem but was not applied. As a consequence, wild lettuce dominated the conventionally fallowed fields to the extent that winter wheat, which was sparse and in low density, did not yield adequately to be harvested in July 1985. The wild lettuce also overstoried the wheat making harvest difficult. Two of the 4 no-till plots that had been planted contained vigorous stands of winter wheat and the other 2 were marginal in quality due to high weed concentrations and nitrogen deficiency. However, all were harvested. Thus, the first year comparisons between no-till and conventionally-fallowed wheat field yielded little information that could be used to recommend either for or against no-till wheat farming based on yields and economics. The moderately persistent herbicide, atrazine, was applied to 8 ha of new wheat stubble on 10 August 1984 in anticipation of no-till replication in 1985 (Snyder 1985~). The application rate was 1.1 kg/ha (1 lb/acre) and its success depended on subsequent precipitation to place it into the top soil. Adequate rainfall was not received for several weeks subsequent to the treatment. As a consequence only partial weed and volunteer wheat control resulted. A second less persistent herbicide treatment was needed in spring 1985 but was not applied. The management technician selected an alternate site for continued no-till treatment. Spring 1985 was phenologically early compared to most years resulting in early growth of both weeds and wheat. In most years, the spring herbicide treatment would preferably have been applied in early June but, because of the rapid foliage growth, the date was moved up to 22 May. Assistance was provided to management technicians in selecting and acquiring herbicides, sprayer preparation, and calibration, field layout and flagging, and in herbicide application. A mixture containing chlorsulfuron (Glean), glyphosate (Roundup), and dicamba (Banvel) was applied with a surfactant using a ground sprayer. Glean and Roundup were applied at rates recommended by agronomists for use on no-till fallow. Banvel was applied at below recommended rates to help kill the larger rapidly growing broad-leafed plants already present. Approximately 12.1 ha (30 ac) were treated which included 3 small tracts that had not contained wheat the previous year. The herbicide mixture was highly effective in killing all existing weeds and volunteer wheat and in preventing �73 new weed establishment in wheat stubble tracts through the summer. The small tracts not containing wheat stubble were essentially bare ground and low growing weed species established on these sites in mid to late summer. These weeds could have been readily eradicated with a single shallow tillage. Management personnel did not have access to a no-till drill so the weedy patches were not tilled and none of the 12.1 ha tract was planted in fall 1985. Environmental Measurements Precipitation and Soil Moisture.--January through September precipitation in 1984 totaled 42.9 cm (~6.9 in.) whereas only 25.4 cm (10.0 in.) of moisture were received for the same interval in 1985 at the Bonny Dam Weather Station (Table 1). The long-term mean for January through September was 37.1 cm (14.6 in.). Thus, precipitation prior to and through the primary 1985 growing season was below average. An attempt to measure spring soil moisture accumulations using the soil probe method (Snyder 1985a) was made on 8 May 1985. However, dry upper soil prevented probe penetration of the soil. Since wheat was not planted in fall 1985 within either no-till or conventially fallowed fields in the South Republican Management Area, fall soil moisture comparisons could not be conducted. Table 1. mea,na• Precipitation at Bonny Dam in 1984 and 1985 vs. the long-term 1984 1985 Inches Cm Inches Cm Lon~-term x Inches Month Cm Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec 1.88 4.44 5.72 11.48 5.56 4.83 4.47 2.59 1.93 7.87 0.38 1.83 0.74 1.75 2.25 4.52 2.19 1.90 1.76 1.02 0.76 3.10 0.15 0.72 0.99 1.14 0.94 2.41 2.24 3.15 9.47 1.47 3.63 2.08 1.52 1.35 0.39 0.45 0.37 0.95 0.88 1.24 3.73 0.58 1.43 0.82 0.60 0.53 0.79 0.84 2.24 3.68 7.70 6.60 6.32 5.44 3.50 2.23 1.40 0.79 0.31 0.33 0.88 1.45 3.03 2.60 2.49 2.14 1.38 0.88 0.55 0.31 Totals 52.98 20.86 30.39 11.97 41.53 16.35 aLong-term means obtained from U.S. Weather Service Climatological Data, 1983 Annual Summary. Vegetation Sampling.--Spring 1985 height-density indices (HDI's) averaged 1.24 dm (Table 2, Fig. 1) for 6 stubble fields or above the 0.98 dm indices recorded for stubble fields in 1984 (Snyder 1985a). These relatively high indices were a product of above average precipit~tion received in 1984 and indicate that cover conditions for ring-necked pheasants (Phasianus colchicus) �74 were generally good. However, observations provide evidence that pheasants were in low density within the South Republican Wildlife Area in both 1984 and 1985. Green wheat attained excellent early growth which exceeded the concealment value of wheat stubble by early May (Table 2, Fig. 1). Winter wheat began heading out in mid to late May, 1-2 weeks in advance of normal phenology. Table 2. Height-density (dm) of residual and/or green vegetation within wheat stubble, green wheat, and other covers, South Republican Wildlife Area, Spring 1985. N Cover type Date Fields Samples X HDI 95% CI Wheat stubble (residual) 23 Apr 6 360 1.241 0.104 Green wheat 23 Apr 8 May 29 May 6 6 6 340 300 260 0.523 1.886 7.969 0.055 0.129 0.259 Annuals and Perennials (Unmanaged) 23 Apr 8 May 29 May 4 4 4 240 208 160 0.313 0.542 1.158 0.067 0.060 0.117 Perennial grasses 23 Apr 29 May 3 3 180 120 1.553 2.294 0.226 0.297 Alfalfa (dry1and) 23 Apr 8 May 29 May 1 1 1 60 60 32 0.388 1.125 3.609 0.098 0.216 0.270 Only 3 tracts of perennial grass were sampled and considerable variation in height-density quality existed among them. Two upland sites containing clumpy stands of switchgrass and sand 10vegrass (Panicum virgatum - Eragrostis trichodes) yielded HDI's of 0.6 and 0.9 dm on 23 April and 1.5 and 1.3 dm on 29 May. Almost no new growth was noted on 8 May so they were not sampled at that time. A riparian (sub-irrigated) site dominated by intermediate wheatgrass (Agropyron intermedium) had a HDI of 3.1 dm on 23 April that increased to 4.1 dm on 29 May. The upland sites ranked below wheat stubble in height-density quality until late May but, when the bottomland tract was included, the overall index exceeded that for wheat stubble (Fig. 1). The 4 unmanaged tracts were dominated by downy brome, but contained a diversity of other annual and perennial vegetation. Downy brome is one of the earliest herbaceous species to begin spring growth and to mature. In the relatively dry weather in 1985, height-density indices ranked below other covers including wheat stubble throughout the sample interval (Table 2, Fig. 1). �75 8 7 • ..--. ::E: C!J STUBBLE E;!J GREEN WHEAT rms PERENNIAL GRASS ~ UNMANAGED (ANNUALS) em ALFALFA 6 ~ .....•. :x: I%l § 5 · 4 · H :>-< E-t H C/) z ~ ~ I E-t ::t: o H ~ ::t: 3 2 1 o ~--;---------. 0° 0 00 0°0 00 0 0 ffil11::: ~ 23 APR "- tEl - .... _--------:::: ---- .. - ·t :::: ~ I ~~~ 29 8 MAY Fig. 1. Height-density indices of vegetation types in comparison to wheat stubble, South Republican Wildlife Area, Spring 1985. �76 One field of alfalfa that was irrigated and had been shallowly disked in early spring yielded rapidly increasing HDI's from 23 April through early and late May. Another tract of alfalfa had been more vigorously cultivated and not included. HDI's there were approximately one-half of values recorded in the first field. Rodent Populations.--Inspections of no-till stubble fields revealed no evidence that rodents were becoming excessively abundant, however, continuance of monitoring over a several year interval would be needed to more accurately assess this potential problem. Dissemination of Study Information Tim Davis, Area 3 Biologist, worked closely with the author on this no-till effort. He was instrumental in acquiring 2 demonstration drills for use on the South Republican Wildlife Area in fall 1984 and in transporting them to the site. He also attempted to initiate no-till wheat farming in Area 3 within the northeast region but lack of equipment and funds have thus far stymied this effort. T. Davis prepared an article encouraging farmer acceptance of no-till noting its benefits to soil conservation and wildlife which was published in the Sterling "Journal Advocate" newspaper. The article was reprinted in the Division's "News for Landowners" which is mailed to farmers throughout Colorado (Davis 1985). An article titled "Pheasants could be a second cash crop on no-tilled ground" was submitted to the High Plains Journal and appeared in a December issue (Snyder 1985b). This article dealt with the conditions necessary for making no-tilled biennially cropped winter wheat produce a high sustained yield of pheasants and other wildlife. Increased wildlife production using no-till would create opportunity for charging hunter access to help defray the costs of converting to no-till farming methods. Factors Influencing Continued No-till Farming on CDOW Lands Efforts to implement and evaluate no-till biennially cropped winter wheat, as a method for increasing wildlife production and harvest, have been discouraging. Lack of capital equipment funds with which the Division could acquire one or more no-till drills is a primary problem. In addition, herbicides are expensive and must be purchased out of the same budget as other management operations. Further, monetary returns for sale of wheat from no-tilled land must go to the general fund for reallocation. They can not be ear-marked for purchase of additional herbicide, fertilizer, or equipment needed to perpetuate no-till. If these obstacles could be overcome, it seemingly would be possible for Division technicians to conduct no-till farming which would involve applying necessary herbicides and seeding. Harvesting and fertilizer applications could be custom contracted. However, no-till farming requires an excellent working knowledge of agronomy and herbicides which would necessitate additional training of property technicians. A second option is to contract sharecrop agreements for no-till farming on Division Properties. However, at present, no-till farming is still in its infancy, most farmers lack experience, are reluctant, or cannot afford to get involved, and most lack the necessary equipment. Therefore, mutually acceptable bids from neighboring farmers probably cannot be obtained. �77 These factors illustrate why evaluation of no-till biennially cropped winter wheat must be discontinued on Division of Wildlife Properties at this time. Pending further assessment, evaluations may be conducted on private lands in eastern Colorado. Further evaluation is needed to determine if the no-till procedure can be an important and effective management procedure for enhancing wildlife production and harvest on both private and Division properties. Under proper management, winter wheat seemingly has high value for use on Division Properties in eastern Colorado. It possesses the high growth vigor, height, lodge-resistent stubble, and feeding values that perennial grasses lack. It provides essential nesting cover that row crops do not provide. LITERATURE CITED Davis, T. 1985. No-till farming benefits both wildlife and farmer. Div. Wildl. News for Landowners 3(4):1. Colorado Snyder, W. D. 1985a. Evaluation of no-till wheat farming. Colorado Div. Wild!., Wildl. Res. Rep. Fed. Aid Proj. W-37-R-38. In Press. 1985b. Pheasants could be a second cash crop on no-tilled ground. High plains Journal l02(5l):11A. Prepared by: lR~ rren ¥d- D.SIlYder Wildlife Researcher C ��Colorado Division Wildlife Research April 1986 of Wildlife Report 79 JOB FINAL REPORT State of Project: Colorado 45-01-506-15050 Work Plan 2: Job Title: (W-88-R) Migratory Bird Investigations Job 10 Distributional Characteristics Inhabiting Colorado Period Covered: of Some Po;eulations of Canada Geese 17 June 1981 - 31 July 1985 Author: Michael Szymczak Personnel: G. Patten and staff, Arapaho National Wildlife Refuge; Tom Kenyon and staff, Colorado State Parks; C. Cesar, Bureau of Land Management; G. Bishop, L. Budde, H. Burdick, G. Byrne, R. Cote, D. Crane, D. Crawford, L. Crooks, T. Davis, R. Desilet, K. Dillinger, J. Jones, K. Kahler, H. Lanning, J. Leslie, S. Porter, C. Reichert, B. Sigler, S. Thorne, J. Wagner, K. Wagner, P. Will, and M. Szymczak, Colorado Division of Wildlife. ABSTRACT Canada geese (Branta canadensis) were trapped and banded on brood-rearing areas in west-central (717), North Park (948), South Park (899), and northeast (985) Colorado. West-central banded geese were recovered predominantly in north-central (52%) and west-central Colorado (27%). Seventy-five percent of the direct recoveries of North Park banded geese were reported from eastern New Mexico, but indirect recoveries were scattered. South Park geese apparently winter predominantly in the San Luis Valley as that area accounted for 71% of all recoveries. Direct recoveries of geese banded in northeast Colorado occurred predominantly in the area of banding (94%) but indirect recoveries indicated northward movement in subsequent years, but probable return to the banding area. First-year recovery rates varied from 1% (South Park) to 9% (west-central) for birds banded as locals and from 1% (South Park) to 3% (northeast) for those banded as adults. January populations of Canada geese in northeast Colorado increased from 1965-69 (x ,. 384) to 1980-84 (x - 6,185). Most large geese banded outside Colorado and recovered or recaptured in northeast Colorado were released in conjunction with breeding restoration projects in northeast Wyoming and western North and South Dakota. The northeast wintering population is within a continuum of wintering Canada goose populations that occupy the Platte River Valley of Wyoming, Colorado, and western Nebraska. January populations of Canada geese in west-central Colorado increased from 56 in 1965 to a high of 4,476 in 1982. Encounters of birds banded post-season in west-central Colorado and those encountered during winter in west-central Colorado but banded outside the state, indicate the primary breeding range for wintering west-central Colorado geese is central Wyoming and northwest Colorado. ��81 DISTRIBUTIONAL CHARACTERISTICS OF SOME POPULATIONS OF CANADA GEESE INHABITING COLORADO Michael R. Szymczak INTRODUCTION During the past 25 years there has been substantial change in the flock size and distribution of Canada geese in Colorado. Historically, Colorado supported 2 populations: (1) the wintering short-grass prairie population in the southeast part of the state whose characteristics were well documented by Rutherford (1968) and Grieb (1970) and (2) a nesting flock of western Canada geese (B. c. moffitti) in extreme northwest Colorado that are members of the Rocky Mountain Population (Krohn and Bizeau 1980). Through a combination of establishing nesting populations through restoration programs in Colorado and other areas, and changes in winter distribution of some flocks, Colorado now contains 7 distinct nesting populations and 5 distinct wintering populations. This project was initiated primarily to document population range characteristics of 5 of the 7 breeding populations and 2 of the 5 wintering populations. All objectives have not been attained and some investigation is continuing. This report was prepared because of a change in funding sources and to meet reporting obligations to Federal Aid in Wildlife Restoration. The report summarizes progress, which includes the achievement of some objectives, through June 1985 and presents recommendations for continued investigation. P. N. OBJECTIVES 1. To document the wintering range and harvest distribution of Canada geese nesting in (1) northwest Colorado, (2) west-central Colorado, (3) North Park, (4) northeast Colorado, and (5) South Park. 2. To ascertain the breeding range of Canada geese wintering in west-central and northeast Colorado. 3. To contribute data to Central and Pacific Flyway management plans for specific populations of Canada geese. SEGMENT OBJECTIVES lao Trap and band at least 150 Canada geese annually on production areas: (1) along the Colorado and/or White rivers in west-central Colorado (4 yrs.); (2) along the Yampa, Little Snake, and Green rivers in northwestern Colorado (4 yrs.); (3) in northeast Colorado (3 yrs.); (4) at least 100 Canada geese annually on production areas in North Park (5 yrs.); and (5) at least 75 Canada geese annually on production areas in South Park. Record the band numbers of all geese recaptured that had been previously banded. lb. Submit banding schedules and recovery reports to the U.S. Fish and Wildlife Service's Bird Banding Laboratory. lc. Plot the recovery locations of all geese reported recovered or �82 recaptured, according to banding location, that result from any of the above banding operations using annual computer printouts provided by the U.S. Fish and Wildlife Service's Bird Banding Laboratory. 2a. Trap and band at 1e~st 250 Canada geese on wintering areas in west-central (4 yrs.) and northeast Colorado (4 yrs.). Each goose captured in northeast Colorado should be weighed and the following measurements recorded: wing chord, middle toe, tarsus, culmen, bill nail length, bill width at base, bill width at posterior edge of nares. Record the band number of all geese recaptured that had been previously banded. 2b. Submit banding schedules and recovery reports to the U.S. Fish and Wildlife Service's Bird Banding Laboratory. 2c. Plot the banding locations of Canada geese reported recovered or recaptured in northeast and north-central Colorado using annual computer printouts provided by the U.S. Fish and Wildlife Service's Bird Banding Laboratory. 2d. Plot the recovery locations of all geese reported recovered or recaptured that had been banded on wintering areas in west-central and northeast Colorado using the computer printouts identified in 2c. 3. Prepare appropriate reports. METHODS Canada geese were trapped on production/brood rearing areas in west-central Colorado (1981-84), northeast Colorado (1981-84), North Park (1981-84), and South Park (1981-84) during their flightless period (late Jun - early Ju1) using standard drive-trapping methods and equipment (Szymczak et a1. 1981). Concentrations of flightless adults and goslings could not be located along the Yampa River in northwest Colorado, therefore birds were not trapped in that area. Recoveries through most of the 1984-85 hunting season of birds banded in the above areas were plotted by degree block of latitude and longitude if recovered outside of Colorado and by 10-minute block if recovered within the state. Recoveries were also recorded by year of recovery according to age. Birds banded in summer 1980 in northeast Colorado and South Park were also included in the analysis. Canada geese were trapped with cannon nets (Dill 1969) in winter in northeast Colorado in 1982 and 1983. Weight, wing chord, tarsus, culmen, bill widths, and bill nail lengths were recorded for geese trapped in January-February 1982. All attempts to trap geese in other years and in west-central Colorado in winter were unsuccessful. Birds trapped in west-central Colorado in January 1981 before initiation of this project were included in the analyses. Recoveries of the 1984-85 hunting season were plotted by degree block and attempts were made to assign the bird to a particular breeding flock. Banded Canada geese recovered or recaptured in northeast and west-central Colorado were recorded by area and, if possible, specific wetland of banding. Probable breeding flocks for some birds wintering in the area of recovery were determined by examining the location of banding, with particular emphasis on birds classified as wild-trapped goslings when banded. �83 RESULTS AND DISCUSSION Summer Populations West-central Colorado Breeding population restoration activities began in west-central Colorado in 1967 with the release of 25 hand-reared fledged goslings on an island in the Colorado River near Loma (Rutherford 1968). In subsequent years an additional 102 goslings were released at the same location. By 1980, nesting geese had pioneered up the Colorado River to its confluence with the Roaring Fork and up the Gunnison River (Fig. 1) system to Hotchkiss with concentrations in the area from Silt to Debeque on the Colorado River (Szymczak 1981). There were 91 indicated nesting pairs (singles, pairs with nests, and paris with goslings) among the 529 geese observed on breeding pair surveys in the entire area in 1984, 55 of those pairs were in the Silt to Debeque area. From 1981 to 1984 over 700 geese were captured in the Rifle-Silt area, 599 of which were goslings (Table 1). Some goslings were moved from the area of capture to Radium and Finger Rock (Table 1) and were not included in this analysis. Distribution of Recoveries. -- All birds recovered were banded at the Silt or Chambers Pond locations. Colorado accounted for 88.7 and 79.3% of the direct and indirect recoveries, respectively. Southeast Arizona and Saskatchewan were the only other areas that had more than 1 recovery (Table 2, Fig. 2). Surprisingly, most Colorado recoveries occured in the Central Flyway along the foothills from Denver north, within the Hi-Line Canada goose population wintering range (Table 3, Figs. 3 and 4). These recoveries included adults as well as birds banded as locals and occurred over a period of years. Of the 31 direct recoveries, 3, 9, and 19 were taken in 1981, 1982, and 1983 respectively, indicating the establishment of a consistent migration pathway. Some birds were in the Front Range area as early as late October but nearly 50% of the recoveries in that area occurred from early to mid-December. Recaptures provided additional information showing the Front Range area's importance to the west-central population. Two birds banded as goslings near Silt were recaptured during summer trapping in the metro-Denver area, 1 at Sloans Lake and 1 at Ketring Park in Littleton. Conversely, 2 birds banded as adults at Sterling Ponds near Fort Collins in 1978 were recaptured near Silt in 1983. Seven birds were recaptured in Wyoming during the flightless period, 4 at Wheatland Reservoir, north of Laramie, a traditional Rocky Mountain population molting area (Krohn and Bizeau 1979) and 3 along the North Platte River near Casper. Seven birds that were banded at Wheatland Reservoir were recaptured near Silt in 1982 and 1983. These encounters indicate some geese of the west-central population use the Wheatland Reservoir molting area. Recovery Rates. -- Both locals and adults banded in west-central Colorado had the highest rate of recovery of any of the 4 banded populations but, although adult recovery rates were only slightly higher. First-year recovery rates of locals approximated 10% (Table 4) but only 2% of the adults were recovered �00 ~ WEST-CENTRAL NORTHEAST White River-l Colorado River-2 Guenzi-l Red Lion-Jumbo Res.-2 Johnsons Pond-3 Haxtun-4 NORTH PARK SOUTH PARK Antero Res.-l Eleven Mile Res.-2 Fig. 1. in Breeding range and specific trapping areas, where applicable, banded summar 1980-84. of Canada goose populations �95 Table l. Canarla geese banded by age and sex, west-central Chambers Pond White River Meeker 1981 Ad Hale Ad Female Loc Male Loc Female Totals 1982 Acl Male Ad Female Loc Male Loc Female Totals 3 2 13 11 29 1983 Ad Male Ad Female Loc Male Loc Female Totals 6 6 20 16 48 Colorado, Silt summer 1981-84. Radium Totals Finger Rock 3 2 11 13 29 3 2 11 13 29 27 27 106 120 280 30 29 119 131 309 11 16 66 44 137 1984 Ad Male Ad Female Loc Male Loc Female TotalR 1 63 44 108 7 5 12 18 22 156 109 305 7 10 17 6 8 25 35 74 14 15 29 57 61 311 288 717 6 8 18 21 39 4 18 1981-84 Ad Male Ad Female Loc Male Loc Female Totals 6 6 20 16 48 3 2 13 11 29 47 53 183 181 464 1 81 65 147 aWhite River, R miles west of Meeker. bChambers Pond, 2 miles east of Rifle. cBirds tranRported ann released. Table 2. Distribution Location of recoveries Ad male Dir Ind Saskatchewan Manitoba Pacific Flyway Idaho Nevada Arizona Colorado Subtotals Ad female Dir Ind A e and sex Loc male Dir Ind Colorado, summer 1981-84. Loc female Dir Ind Totals Dir Ind 10.0 10.0 6.9 3.4 10.0 3.4 3.4 9.1 9.1 Central Flyway Colorado Texas New Mexico Subtotals N Recoveries (%) of Canada geese banded in west-central 0 6.7 23.3 30.0 45.5 50.0 14.3 19.0 33.3 20.0 30.0 36.4 66.7 50.0 67.9 1.9 70.0 9.1 41.7 66.7 50.0 69.8 3.4 44.8 30 12 21 10 53 29 50.0 50.0 60.0 60.0 50.0 100.0 40.0 66.7 3.3 50.0 100.0 40.0 2 2 5 9.4 20.8 30.2 37.9 44.8 41.4 �86 .... ~ Fig. 2. Distribution of recoveries outside of Canada geese banded in westcentral Colorado, 198)-84. Direct recoveries are circled. �37 101 108 109 41 '" 1'1"" 1 r ,~ \' A . 105 "" , I" 10) 104 41 U~ 'I"'" 1 1 hit 2 r.:: '. 40 C0l-0RADO 10 ,... 1'''' .. 1 0 _Ilf-I ",,,sr 2 , /llll w/(. ,'J . 1 1 iI l ~~ I ". •• "'"J f'-U' 2 6 I ~t ~.._ 4o 1 5 1 1 • j2, Io."r- " »o ., 1"", ' ru" ' A ••• I 1 ~ r-: ,•.. 39 v 1< .r J" ~ " i--'iJc: i:;;K 1 1 \ .I;:;;~ II 1 ~ 1"'''"' )8 tI. • . ., T 0, 39 I~ It' r"'~ , II .•. CD I . ., 1..- 1 1"'" ., I )8 r1'" " I 0 I 1"(' , / , V • (JO \ { 37 108 0 19 Fig. 3. Distribution of direct recoveries summer 1981-84. (1 Unknown location). 41 108 109 ., r in Colorado lOb r- I A' 104 105 I 103 of Canada geese banded in west-central COLORADO 107 lr" 31 106 107 105 "" 103 104 , 1\ 1 I_ Jl 't- Colorado, 41 _"'1'$ ~ Ii:;;~ '" ,.. I- 40 r' , .. " .•. "\ 1 1 ,,... 1"'1= ,... 'JI 01. 1/ \ .. r- •• Fig. 4. SWlllDer Distribution 1981-84. , "" , 39 .. . . -r-r- C 107 of indirect recoveries , o .r ., )8 lL I :,... , 106 . r- H· I"" ( lOS .. ~, , , = )7 109 1 1 f'"':' } l/.i 1/ 40 3 'u. i\ "", r"'~ '£, " 1 It' " !=.ur- 1 'r" "" 1 2 1.••" !"'!' ':;:;"'- 1< / 1 1 lI- )8 l-t ~•• ~ tmH " 1 1 rn IJ 1 5 39 .."'. 0 ~ .r 37 105 104 103 in Colorado of Canada geese banded in west-central Colorado, �88 (Table 5). The transmountain migration of these birds into an area of heavy hunting pressure in north-central Colorado is most likely responsible for the comparatively high recovery rates. Summary and Recommendations. -- The artificial movement of birds captured in 1984 unfortunately delayed compiling additional evidence of transmountain migration. A substantial number of birds have been recovered over a period of years along the Front Range identifying that area as the major wintering area for this population. However, all of the birds were banded in the Silt-Rifle area, primarily on one major brood-rearing area. The evidence indicates that segment of the popUlation winters along the Front Range, but the winter range of other segments is still in question. Therefore, I recommend that banding in the Silt-Rifle area be curtailed and any additional trapping should occur along the lower Colorado River or on small pond production areas from Grand Valley to the state line or possibly on the Gunnison River. Trapping attempts on the White River should also be renewed. North Park A concerted effort to establish a Canada goose breeding population in North Park began in 1969 with the release of goslings on Lake John Annex. Canada geese had been released in North Park in 1956 and 1957, but only an occasional pair nested in the park in the years following the original releases. Over a 5-year period, a total of 931 goslings was released at Lake John Annex and Walden Reservoir. Recovery of transplanted goslings occurred predominantly in the Pacific Flyway (Szymczak 1975) along traditional migration routes and in winter areas of the Rocky Mountain Canada goose population (Krohn and Bizeau 1980). At least 20 nesting pairs were observed during an aerial flight in 1975 (Szymczak 1976) and by 1981 geese were nesting throughout the Park, wherever adequate nest sites were available. From 1981 to 1984, 948 Canada geese were captured, banded, and released in North Park (Table 6). Two-thirds of the birds were captured on Walden Reservoir, a production and adult molting area. Other banding locations were Pole Mountain Reservoir, a production area with some molting adults, McFarlane Reservoir, a molting area with some production, and Case Flats, Home Pond, and South McCammon Pond production areas on the Arapaho National Wildlife Refuge. Distribution of Recovery. -- Bandings in North Park resulted in few recoveries. New Mexico accounted for 70% of the direct recoveries but only 17% of the indirect recoveries (Table 7). Direct recoveries in New Mexico occurred in 1981 (10), 1982 (2), and 1984 (5) indicating consistent annual migration. Nine of the 10 direct adult recoveries, however, were of birds banded on Walden Reservoir in 1981 when over 200 adults were banded, many of which were members of the molting population. The origin of the growing molting population on Walden Reservoir which numbered nearly 500 by 1984 (Szymczak, unpubl. data) is unknown. Recoveries in New Mexico were concentrated in the southeast quadrat of the state (Fig. 5) in the area near Bitter Lake National Wildlife Refuge. One bird banded in the winter at Bitter Lake in 1976 was recaptured in North Park. Seven birds (4 locals, 3 adults) banded over 2 years in North Park were recaptured in early December near Springer in northeast New Mexico. �39 Table 3. Distribution summer 1981-84. Location of recoveries (%) in Colorado of Canarla geese banded in west-central A e and sex Loc male Dir Ind Ad female Dir Ind 50.0 60.0 18.5 50.0 60.0 3.7 3.7 25.9 22.2 55.5 22.2 50.0 100.0 40.0 66.7 44.4 61.1 North-central Other Table 4. 1980-84. Year banded 11.1 22.2 7.4 N Recoveries 0 Chronology N banded Loc female Dir Ind Ad male Dir Ind West-central Garfield County Delta County Eagle County Mesa County Subtotals 2 2 of recoveries 5 27 14.3 14.3 22.2 9 Totals Dir Ind 19.1 26.1 13.0 28.6 2.1 2.1 23.4 8.7 47.8 71.4 66.0 52.2 lO.6 16.7 18 7 of Canada geese banded as locals in west-central Year and N of recoveries 1981 1982 1983 1984 Colorado, Colorado, Totals First-year rate recover:!:: 47 23 summer Percent recovered 1981 1982 1983 1984 5 250 265 4 1 0 24 1 9 24 0 4 7 0 2 37 31 0 0.20 0.09 0.07 0.00 40.0 14.8 11.7 0.0 Totals 543 3 24 35 11 73 0.09 13.4 Table 5. 1980-84. Year banded 1981 1982 1983 1984 Totals Chronology N banded of recoveries of Canada geese banded as adults in west-central Year and N of recoveries 1981 1982 1983 1984 Totals Colorado, First-year recover:!:: rate summer Percent recovered 5 59 40 14 1 0 0 1 3 1 0 1 2 0 2 4 3 0 0.20 0.00 0.03 0.00 40.0 6.8 7.5 0.0 118 1 0 5 3 9 0.02 7.6 �90 Table 6. Canada geese banned by age and sex, ~orth Park, summer 1981-84. Walden Reservoir 1981 Ad Male Ad Female Loc Male Lac Female Totals 106 106 59 59 330 1982 Ad Male Ad Female Loc Male Lac Female Unknown Totals 105 1983 Ad Male Ad Female Loc Male Lac Female Totals 12 19 33 24 88 MacFarlane Reservoir Pole Mt. Reservoir Case Flats Home Ponti Totals S. Mcl.ammon 114 115 69 65 363 8 9 10 6 33 2 30 50 7 12 1 100 46 32 11 16 3 5 5 5 10 11 10 31 83 87 31 44 1 246 4 3 13 9 29 10 15 8 8 41 27 38 61 49 175 1 1 7 8 17 1984 Ad Male Ad Female Loc Male Loc Female Unknown Totals 37 41 49 28 3 158 1981-84 Ad Male Ad Female Loc Hale Lac Female Unknown Total 201 198 152 1273 3 681 Table 7. Distribution Location Colorado North Park Larimer County Weld County Logan County Pueblo County North Dakota South Dakota Wyoming New Mexico Texas Arizona California Saskatchewan N Recoveries of recoveries Ad male Ind Dir 37 42 51 31 3 164 1 2 3 6 18 24 18 14 30 50 7 12 1 100 74 6 4 18 17 5 5 10 11 1 1 7 8 45 31 17 (%) of Canada geese banded in North Park, 1981-84. Ad female Dir Ind 9.1 10.0 A e and sex Loc male Dir Ind 20.0 Lac female Dir Ind Totals Dir Ind 25.0 14.8 11.1 11.1 33.3 81.8 261 282 212 1894 4 948 10.0 20.0 10.0 100.0 20.0 fiO.O 30.0 10.0 9.1 33.3 33.3 25.0 22.2 11.1 11.1 7.4 50.0 11.1 11.1 70.4 7.4 100.0 10.0 11 1 2 10 11.1 10 3 4 9 27 4.3 4.3 8.7 4.3 17.4 4.3 4.3 4.3 17.4 4.3 13.0 4.3 8.7 23 �91 Indirect recoveries were scattered with no distributional consistency (Table 7, Fig. 6). The banded population is not Pacific Flyway oriented as were the original transplants. Most recoveries in Colorado (Fig. 7) and New Mexico occurred outside traditional Hi-Line Canada goose population wintering areas and in areas where isolated populations of large Canada geese have been increasing. Recovery Rates. -- Only 3.6% of the locals (Table 8) and 1.9% of the adults (Table 9) were reported recovered the first year following banding from 1981 to 1983. Total recoveries averaged 7% for locals and 4% for adults. Summary and Recommendations. -- Original procedures called for banding 100 Canada geese on North Park production areas annually for 5 years. Over 900 birds, including over 400 locals, were banded from 1981 to 1984. Direct recoveries indicate a consistent movement into New Mexico in fall and winter but indirect recoveries have been scattered. Since recovery rates have been low, few additional indirect recoveries of birds banded through 1984 are expected. Therefore, I recommend that trapping be conducted for another year in North Park with emphasis on capturing goslings and accompanying adults on production areas. Northeast Colorado Canada goose breeding population restoration efforts began in northeast Colorado in 1971 with the release of 102 goslings on Jumbo Reservoir (Fig. 1) near Sedgewick, Colorado. An additional 201 birds were released on Jumbo in 1972 and 1973. From 1972 through 1974, 266 goslings were released at Prewitt Reservoir along the South Platte southwest of Sterling, Colorado in Washington County. During the time of these releases, a small nesting population was present at Johnson's Pond in extreme northeast Colorado. Growth of the population centered around the intensively-managed goose nesting areas on the Red Lion Wildlife Area (Jumbo Annex), Johnson's Pond, and the Guenzi Ranch (Fig. 1), managed by the Northeastern Colorado Rod and Gun Club. By 1975, 16 and 14 nests were noted during aerial surveys on the Red Lion and Johnson's Pond areas, respectively. From 1980 to 1984, nearly 1,000 Canada geese were banded in summer in northeast Colorado, 80% at the above 3 areas (Table 10). The additional birds were banded at a private production area near Haxtun (Fig. 1). Distribution of Recoveries. -- For distribution analysis, banded birds were separated into 2 groups; those banded at Red Lion-Jumbo Reservoir, Johnson's Pond, and Haxtun; and those captured at the Guenzi Ranch. Although few recoveries resulted from the Guenzi bandings, little interchange was noted in Colorado recovery areas between the 2 groups (Tables 11 and 12). The first year following banding, birds from both groups remained in the general area of banding. No recoveries occurred outside Colorado and all in-state recoveries were in the northeast corner (Figs. 8 and 9). Closed hunting areas encompassed the major roosting and some feeding areas. Indirect recoveries indicated some northward movement away from the banding areas, particularly by birds banded as locals. Adults remained as residents while some locals apparently moved north with wintering migrants. Recovery locations in eastern Saskatchewan (Fig. 10) indicate the birds were within the �\0 N Fig. 5. Distribution of direct recoveries of Canada geese banded in North Park, summer 1981-84, and recovered outside Colorado. Fig~ 6. Distribution of indirect recoveries of Canada geese banded in North Park, summer 1981-84, and recovered outside Colorado. �41 107 108 109 0' , \1'" ~ I'P'" 10 0 1• e .•. ~.... 1 1 'r-' 1 OlllJfll 'II r- 40 r-' ," 1'\ ~ I) "'" ~~-',-- r'" , ;;:;:- • :>-. ~ f'\ i'" • /);: w~ ',...'- .. Ne . N \ a ""''* r ,_ 'A"~ 39 lI." ~ / «> I w' , .-' f'" )8 f 10. \ V :'1. < "'" ( . 1:16 107 ~2 ~ ...•. ~I )7 108 ....•. 1\ h / .~ v •.. I.r" , o• , I~ I 1 A ••"1'" , , I . r-; p:, )9 41 ,., '" 1 ) iZ~ U 103 s".~ If 1\ 1 J 10 ,_ 104 w, • )7 104 105 10) Fig. 7. Distribution of recoveries of Canada geese banded in North Park, summer 1981-84, and recovered in Colorado. Direct recoveries are circled. Table 8. Year banded Chronology N banded of recoveries 1981 of Canada geese banded as locals in North Park, Colorado, Year and N recovered 1984 1982 1983 summer 1981-84. Totals First-year recover)! rate Percent recovered 1981 1982 1983 1984 134 75 95 82 3 5 1 1 2 4 1 0 3 6 10 3 7 6 0.02 0.01 0.04 0.07 7.5 4.0 7.4 7.3 Totals 386 3 6 7 10 26 0.04 6.7 Table 9. Year banded Chronology of recoveries N banded 1981 1981 1982 1983 1984 229 170 65 79 7 Totals 543 7 of Canada geese banded as adults in North Park, Colorado, Year and N recovered 1982 1983 1984 1 2 2 3 0 3 5 summer 1981-84. Totals Fir::;t-year recover)! rate Percent recovered 1 3 1 4 11 8 1 4 0.03 0.01 0.00 0.05 4.8 4.7 1.5 5.1 9 24 0.02 4.4 �94 Table 10. 1980-84. Canada geese banded by age and sex, northeast Guenzi Ranch Jumbo Reservoir Red Lion - Johnsons Pond Colorado, summer Haxtun Totals 1980 Ad ~la1e Ad Female Loc Male Loc Female Totals 18 19 20 20 77 22 16 41 41 120 1981 Ad ~la1e Ad Female Loc Male Loc Female Totals 7 6 25 25 63 1 1 13 10 25 21 21 44 58 144 29 28 82 93 232 1982 Ad Male Ad Female Loc Male Loc Female Totals 4 10 19 15 48 4 5 7 9 25 9 6 38 36 89 18 19 37 17 21 82 79 199 1983 Ad Male Ad Female Loc Male Loc Female Totals 3 1 11 9 24 10 11 15 13 49 2 3 20 22 47 24 32 9 8 73 39 47 55 52 193 1980-84 Ad Male Ad Female Loc Male Loc Female Totals 32 36 75 69 212 47 45 ·96 98 286 36 32 112 120 300 42 57 44 44 187 157 170 327 331 985 40 35 61 61 197 �95 Distribution of recoveries (%) of Canada geese banded at Jumbo Reservoir-Red Table 11. Pond, and Haxtun, northeast Colorado, summer 1980-84. Ad male Dir Ind Location Saskatchewan Central Flyway South Dakota Nebraska Kansas Colorado Sedgewick County Haxtun Guenzi Ranch Prewitt Reservoir Morgan C;ounty Unknown 66.7 33.3 Loc female Dir Ind Totals Dir Ind 25.0 21.4 28.6 22.2 7.1 7.1 7.1 14.3 14.3 7.4 7.4 3.7 100.0 21.4 85.7 92.0 4.0 28.6 3 2 of recoveries Ad male Dir Ind Saskatchewan Central Flyway Nebraska Colorado Guenzi Ranch Prewitt Reservoir Haxtun Larimer County 100.0 0 1 10 4 5 11.1 3.7 7.4 14 4.0 7 7 (%) of Canada geese banden at Gllenzi Ranch, northeast Ad female Dir Ind 66.7 33.3 3 0 ,A e ann sex Loc male Dir Ind 50.0 50.0 25.0 25.0 25.0 25.0 2 4 37.0 14.3 14.3 Table 12. Distribution summer 1980-83. N Recoveries A e and sex Loc male Dir Ind 14.3 7.1 14.3 N Recoveries Location Ad female Dir Ind 100.0 75.0 100.0 Lion, Johnsons 27 25 Colorado, Loc female Dir Ind Totals Dir Ind 33.3 12.5 33.3 12.5 100.0 33.3 1 3 66.7 33.3 37.5 12.5 12.5 12.5 6 8 �108 109 41 '" 101 IN'i r ,....,... \ .•, 1"" 10) 104 I~I!I'~ 1\ I,.I 2 1 I) . r.;t.-. 10 "'" 0 rl- 40 I-I ," "'~ ~~:.. J .•. "\ " 39 '" II'"••.•• 1 . u. .,iMij .. , , '" " j" )9 "'" I ,jot \ lIS • I/. r'''I ~h I) ~. 1/ V IL- '" )8 l- ~.••i""' 1/ I•••••• '" Ir-: • I~ [:;:: HO < ''''' . )1 109 I v \ , 1 IJ .1=: '- 0' . II ,,~ If )8 , <D 40 . t"H "s IIWI 1~" • ,. ( •.....•...... :"CJiI 41 'I"'" 108 107 )1 lJ6 lOS 104 10) Fig. 8. Distribution of recovereis in Colorado of Canada geese banded at Red Lion-Jumbo Reservoir, Johnson's Pond, and Haxtun in northeast Colorado, summer 1980-84. Direct recoveries are circled. 41 107 108 109 1""1'" I+'<' I'''''' C ,... I ?oJ- 0 . RA Q09 .•, 10) 104 1\ ~ I,J " R = .• '" , , "'"IJ ~'" _,~ , .... ' , Euf- I !( "\ , 1'-j tII''- I);" F4" 40 !Wi • • fflH or I . .,. )9 I,... rr D~f- 11 " f- Ir' •••• HO < "" l 37 109 108 107 , I•••••• )8 rr "'" /; AU I•...• . ~ 17 1/ . "I '" • r'' : ~h II '1 ~~, ' I? \ 106 41 1 ' ' .. 1 ~rou -. II-J 39 .,I- 'I"'" ./ ~ .. )7 lOS 104 10) Fig. 9. Distribution of recoveries in Colorado of Canada geese banded at the Guenzi Ranch in northeast Colorado, summer 1980-83. Direct recoveries are circled. �97 Fig. 10. Distribution of indirect recoveries outside Colorado banded in northeast Colorado. summer 1980-84. of Canada geese �98 range of the Western Prairie Canada goose population. However, northeast Colorado remained the terminal wintering area as only 1 recovery occurred south of the restoration area. Recovery Rates. -- First-year recovery rates were similar for locals and adults (Tables 13 and 14) unusual in waterfowl populations but probably indicative of this populations' association with a sizeable closed area and first-year non-migrating habits. Over time 8-11% of the banded population was recovered. Recovery rates for locals were generally lower than those for locals banded in west-central Colorado, and similar to North Park locals but higher than for South Park birds. Adult recovery rates were similar to those of west-central adults but higher than for North Park and South Park birds. Summary and Recommendations. -- A comparatively substantial number of recoveries indicate most northeast Colorado geese are harvested in the area of banding. Some birds do move north in subsequent years and are recovered within the breeding range of the Western Prairie population. A portion of those birds may not return to northeast Colorado although the recovery distribution indicates a migratory pathway into the original banding area. Original objectives called for banding 150 geese for 4 years (1981-84), to add to the sample banded in 1980 (197 birds). Banding objectives were exceeded in all years and the resulting recoveries have been adequate to document population range. South Park Canada goose restoration efforts began in South Park in 1972. From 1972 to 1974, 255 goslings were released at Antero Reservoir (Fig. 1). Complete surveys of nesting geese in South Park have not been conducted. However nesting birds have been concentrated at Antero and Eleven Mile reservoirs. Recoveries of the released birds occurred predominantly in the San Luis Valley, Colorado. Other recovery areas of note were the upper Arkansas River Valley near Pueblo, the Texas Panhandle, and the southeast corner of New Mexico. From 1980 to 1984 nearly 900 Canada geese were banded in South Park at Antero and Eleven Mile reservoirs (Table 15). In 1980, nearly 200 members of a molting flock were banded, but in all other years the birds captured were predominantly goslings with accompanying adults. Distribution of Recoveries. -- Few recoveries have resulted from South Park bandings. All but 1 recovery was in Colorado and 74% of the recoveries occurred in the San Luis Valley (Table 16). Recoveries of locals were reported only from Park County, the area of banding, and the San Luis Valley, giving preliminary evidence of a short fall migration, with the flock wintering in the San Luis Valley (Fig. 11). Annually, recoveries in the San Luis Valley began when the goose season opened in early November and continued through the entire hunting season. Members of the molting adult flock that was captured were recovered in the San Luis Valley (2) the first year after banding and in the San Luis Valley (2), Mesa (1), and Larimer (1) counties, Colorado, and central Montana in later years. Five adults captured in South Park, 2 in 1982 and 3 in 1983, had previously been banded as molters at Wheatland Reservoir, Wyoming. �99 Table 13. Year banded Chronology of recoveries N banded 1980 1980 1981 1982 1983 1984 122 175 161 107 93 7 Totals 658 7 Table 14. Year banded 1980 1981 1982 1983 1984 Totals Chronology Year and li of recoveries 1981 1982 1983 1984 1980 Colorado, summer 1980-84. Totals First-year recovery rate Percent recovery 1 2 2 7 5 1 7 3 3 3 2 0 3 2 14 18 8 6 2 0.06 0.01 0.03 0.03 0.02 11.5 10.3 5.0 5.6 2.2 3 14 14 10 48 0.03 7.3 of Canada geese banded as adults, northeast of recoveries N banded of Canada geese banded as locals, northeast Year and li of recoveries 1984 1983 1982 1981 Totals Colorado, First-year recovery rate summer 1980-84. Percent recovery 75 57 38 86 71 2 2 2 1 2 3 1 1 0 2 0 0 0 0 2 6 5 3 2 2 0.03 0.04 0.06 0.02 0.03 8.0 8.8 7.9 2.3 2.8 327 2 4 6 4 2 18 0.03 5.5 �lOU Table 15. Canada geese banded by age and sex, South Park, summer 1980-84. Eleven Hile Reservoir Antero Reservoir 1980 Ad Male Ad Female Loc Male Loc Female Total 84 101 3 2 190 1981 Ad Male Ad Female Loc Male Loc Female Totals 35 45 51 49 180 10 12 10 8 40 45 57 61 57 220 1982 Ad Male Ad Female Loc Male Loc Female Totals 5 3 28 20 56 21 17 S3 57 148 26 20 81 77 204 28 23 33 38 47 49 18 37 1 152 75 72 51 75 1 274 6 5 11 6 5 11 78 78 87 107 1 351 230 250 202 216 1 899 1983 Ad Male Ad Female Loc Male Loc Female Unknown Totals 84 101 3 2 190 122 1984 Loc Male Loc Female Totals 1980-84 Ad Male Ad Female Loc Male Lee Female Unknown Totals Table 16. . Totals Distribution of recoveries 152 172 115 109 548 (%) of Canada geese banded in South Park, Colorado, summer 1980-84. Ad male Dir Ind Ad female Ind Dir Loc male Dir Ind Loc female Dir Ind Totals Dir Ind Colorado San Luis Valley 100.0 71.4 Larimer-Weld County 14.3 Park County Mesa County 14.3 Montana 25.0 25.0 25.0 25.0 100.0 67.7 100.0 66.7 33.3 33.3 100.0 60.0 10.0 20.0 5.0 5.0 Location N Recoveries 2 7 0 4 3 3 3 6 8 20 �101 108 41 109 101 •• r- ~ . .. .. . r •••.•~IM tJ .."..t~ ,. :\ 41 , .. . "1"'" :u.t- or • , I( '\ ii.r Jc: )9 t ••• Hi:-( )9 4 r" _\ • Iw- ... s••• ~ -'" J )8 103 1 1 :;;;"h • , .... 1\ 1 ,. 104 ..• \ ••.. !"" 1'1"" 1 r \ I-- ~ ~ / r-' ~, ••••• os 38 r• 1·- 1oi"A' y /., ..•. J . os • 1 ... ~ £. ' i'" 1/ 1\ ~ l•.•. .,. 2 • II ~~ C2l (j) )7 109 108 107 116 )7 lOS 104 10) ria. 11. Diatr1but1on of reeover1ea in Colorado of canada geese banded in South Park, summer1980-84. Direct recoveries are circled. Recovery Rates. -- South Park Canada geese had the lowest reported recovery rates from the 4 preseason banding areas. Less than 1% of the locals and 1% of the adults were taken the first year after banding (Tables 17 and 18). Recovery rates for this population have been consistent over time. Summary and Recommendations. -- Although few recoveries has resulted from South Park bandings, the distribution of those recoveries has been consistent. The South Park nesting population seems to winter in the San Luis Valley, Colorado. The original procedures called for banding 375 geese over a 5-year period. In 5 years, nearly 900 geese have been banded, but recovery rates have been low. I recommend that an additional 100 goslings and accompanying adults be banded in South Park. Should recoveries through 1985-86 maintain the same geographic consistency, the objective of the South Park banding will have been met. �102 Table 17. 1980-84. Year banded Chronology N banded of recoveries 1980 of Canada geese banded as locals 'in South Park, Colorado, Year and N of recoveries 1982 1984 1981 1983 Totals summer First:-year recovery rate Percent recovered 1980 1981 1982 1983 1984 5 118 158 126 11 0 0 2 0 1 2 0 0 1 2 0 1 2 4 0 0 4 5 6 0 0.00 0.02 0.01 0.02 0.00 0.0 3.4 2.5 4.2 0.0 Totals 418 0 2 3 3 7 15 0.01 3.6 Chronology of recoveries Table 18. 1980-84. Year banded N banded 1980 of Canada geese banded as adults in South Park, Colorado, Year and N of recoveries 1981 1982 1984 1983 summer Totals First-year recovery rate Percent recovered 0.01 0.00 0.00 0.00 4.3 2.9 2.2 0.7 0.004 2.7 1980 1981 1982 1983 185 102 46 147 2 2 0 1 0 0 1 1 0 0 2 2 1 1 8 3 1 1 Totals 480 2 2 1 2 6 13 �103 Winter Populations Northeast Colorado Until the 1970's extreme northeast Colorado was not considered a terminal wintering area for Canada geese. The few geese encountered in this area were normally members of the arctic nesting shortgrass prairie population (Grieb 1970). The increase in winter numbers began about the same time as breeding goose restoration efforts in 1971. Between 1965-69 and 1980-84, average January numbers increased about 20~fold. There was considerable variation in numbers between individual years (Table 19); probably a response to changes in annual weather conditions. Table 19. Average number of Canada geese in northeast Colorado by 5-year intervals according to the January inventory. Period 1965-69 1970-74 1975-79 1980-84 Numbers of Canada ~eese Ranse Averase 384 1,381 2,944 6,815 128-724 80-2,810 1,985-4,003 2,991-14,573 Many of the wintering geese were large Canada's. Plotting the banding location of Canada geese recovered in northeast Colorado indicated that some of the wintering birds were members of the Hi-Line population or restoration flocks in North and South Dakota, and Wyoming (Szymczak 1982). Post-season banding, initiated in 1982, was designated to help identify the breeding derivation of geese wintering in this area. �104 Breeding Areas. -- Post-season trapping in Colorado resulted in only 137 banded geese (Table 20). Six geese captured had been banded previously; 5 in experimental restoration programs (2 in Alberta, 2 in North Dakota, and 1 in Saskatchewan) and a wild-trapped local from north-central Montana. Sixteen of the geese banded post-season in northeast Colorado were subsequently encountered before June 1985. Eight of those encounters were on winter ranges and 5 of the 8 were in the Hi-Line Canada goose population range. Four birds were recovered in early fall within the Hi-Line range (western Saskatchewan) and 3 others were recaptured on Hi-Line breeding/molting areas in Wyoming. Seventy-nine Canada geese banded on breeding areas outside Colorado were reported recovered in northeast Colorado from 1972 to 1984. Including the 6 birds reported recaptured in post-season banding operations, 51% of the birds had been released in conjunction with breeding population restoration efforts. Restoration birds originated primarily in southwest North Dakota, northwest South Dakota, northeast Wyoming and western Nebraska (Fig. 12). Only Alberta contributed a substantial number of wild-trapped birds. The origin of the northeast Colorado population of large Canada geese is difficult to assess from available band encounters. Band recoveries indicated Hi-Line population breeding areas, primarily Alberta, contributed birds to the northeast Colorado harvest. Only 3 birds banded in the heart of the Hi-Line breeding range in Montana were recovered in northeast Colorado. Most Colorado recoveries from Alberta bandings occurred in the 2 degree blocks of latitude and longitude directly west of northeast Colorado. From 1974 through 1978, only 3 of 211 eastern Colorado recoveries of geese banded in Alberta occurred in northeast Colorado. It is logical to expect some Hi-Line population birds to drift east along the South Platte River from the core wintering area. Most other banded birds recovered, were released in restoration areas directly north of the Colorado wintering area. Many birds released in such programs have a tendency to wander, but a movement straight south is not unexpected. Unfortunately, there has been little, if any banding of goose populations established in these restoration areas, and migrational pathways of the 2 groups may be different. Birds released in North Park, Colorado, migrated into southern California and Arizona while their progeny now move south into central New Mexico. Few banded birds recovered originated in eastern Saskatchewan, within the breeding range of the Western Prairie population. A substantial number of birds of that popualtion were banded in the late 1970's. Summary. -- The evidence indicates northeast Colorado is on the fringe of the Hi-Line population wintering range and, although birds of that population are present, they do not compose a significant segment of the population. I hypothesize that most large geese in northeast Colorado originate from newly restored populations in states directly north of the wintering area. Further, the northeast population is not discrete, as it is near the center of a somewhat continuous goose wintering range along the Platte River Valley in southeast Wyoming, northeast Colorado, and western Nebraska. Interchange should be expected. Banding of restored populations north of Colorado would define the limits of the winter range for those birds. �105 Table 20. Age, sex and location of large subspecies of Canada geese banded in January-February, 1982-83 in northeast Colorado. A!5eand sex Prewitt Reservoir Johnson's Pond Jumbo Annex Adult Males Females 26 0 21 22 3 4 5 7 2 10 10 Immature Males Females 1 7 1 Totals 54 25 Unknown age Males Females 24 1 10 0 3 1 37 Totals 55 66 16 137 0 Fig. 12. Origin of banding of Canada geese recovered or recaptured in extreme northeast Colorado 1972-84. Numbers circled were birds released in breeding population restoration efforts; those enclosed in a square were known to be banded on a molting area. �106 West-Central Colorado As in northeast Colorado, the wintering population of Canada geese on the Colorado and Gunnison rivers in west-central Colorado began to grow simultaneous to breeding population restoration efforts in the area. West-central terminal wintering populations, according to the January inventory, underwent steady growth reaching nearly 4,000 birds in 1981 (Fig. 13). January numbers generally have not dramatically fluctuated. Many of the birds counted in winter were thought to be members of the west-central breeding population while others were probably from historical breeding areas on the Yampa and Little Snake rivers in northwest Colorado. Attempts to document the winter ranges of these populations were presented earlier. All birds were considered members of the Rocky Mountain Canada goose population (Krohn and Bizeau 1980) but specific breeding locations for these wintering birds were not known. Attempts to band Canada geese post-season in west-central Colorado were successful only in 1981. Band encounter data of birds trapped in other areas has provided some information. Breeding Areas. -- In January and February 1981, 93 Canada geese were trapped at Highline Lake northwest of Grand Junction, Colorado. Twenty of those birds had been previously banded, all on production and molting areas in central Wyoming. Twenty of the birds banded in west-central Colorado were subsequently encountered through the 1984-85 hunting season. All but 3 were encountered in Wyoming or west-central Colorado. Plotting the banding location of geese recovered or recaptured during winter in west-central Colorado provided additional evidence that breeding areas in central Wyoming provide a substantial portion of the geese wintering in west-central Colorado (Fig. 14). Specific production areas of note are Yellowtail Reservoir on the Big Horn River near Lovell, Ocean Lake northwest of Riverton, and Pathfinder Reservoir and Soda Lakes southwest of Casper. Production areas in western Wyoming on the Bear and Salt rivers were not represented in the west-central harvest. Only a few birds have been banded in the Green River drainage (L. Serdiuk, pers. commun.) and whether those birds migrate into west-central Colorado has yet to be determined. A few birds from the Rocky Mountain population breeding area in Alberta and Utah were recovered in west-central Colorado but those popUlations are not consistent contribuitors to the west-central wintering flocks. The largest number of banded birds encountered were captured as molters at Wheatland Reservoir in southeast Wyoming. Birds captured at Wheatland have included members of many segments of the Rocky Mountain and Hi-Line breeding populations. However, historically most birds have been members of the Pacific Flyway-oriented Rocky Mountain population, including those birds that breed on the river systems in northwest Colorado. (Twenty-three of 26 banded geese recovered on the Yampa River in 1979-84 were banded at Wheatland Reservoir). Attempts to capture birds in northwest Colorado were unsuccessful as reported earlier and definitive information about wintering areas for these populations is still lacking. However, circumstantial evidence indicates that many breeding geese in northwest Colorado winter in the west-central part of the state. �5,000 4,000 3,000 W en W w CJ ZI 2,000 1,000 65 70 75 80 85 YEAR Fig. 13. Growth of the winter Canada goose population Colorado according to the January inventory. 1965-84. in west-central Fig. 14. Origin of banding of Canada geese recovered or recaptured in west-central Colorado 1972-84. NUMbers circled were birds released in breeding population restoration efforts; those enclosed in a squsre were known to be banded on·a molting area. ~ o ••..~ �108 Summary. -- The steady increase in wintering numbers of Canada geese in west-central Colorado indicate the population is composed of birds from growing breeding populations, such as those in central Wyoming, that terminate migration in west-central Colorado. Birds wintering in this area are not transitory migrants whose numbers fluctuate with weather conditions. Geese breeding in Wyoming join the northwest Colorado population that historically wintered in the Imperial Valley of California. These birds apparently now terminate migration in west-central Colorado. LITERATURE CITED Dill, H.H. 1969. A field guide to cannon net trapping. and Wildl. Servo 18 pp. (mimeo). U.S. Dep. Inter., Fish Grieb, J.R. 1970. The short grass prairie Canada goose population. Monogr. 22. 49 pp. Wildl. Krohn, W.B., and E.G. Bizeau. 1979. Molt migration of the Rocky Mountain population of the western Canada goose. Pages 130-140 in R.L. Jarvis and J.C. Bartonek, eds. Management and biology of Pacific Flyway geese, Northwest Section, The Wildl. Soc., Oregon State Univ. Bookstores, Corvallis. __________ , and 1980. The Rocky Mountain population of the western Canada goose: its distribution, habitats, and management. U.S. Dep. Inter., Fish and Wildl. Servo Spec. Sci. Rep. - Wildl. 229. 93 pp. Rutherford, W.H. 1968. Experimental studies on improving the status of Canada Goose populations. Colorado Game, Fish and Parks Dep., Game Res. Rep., Oct. Pp. 65-73. Szymczak, M.R. 1975. Experimental studies in improving the status of Canada goose populations. Colorado Div. Wildl., Wildl. Res. Rep., Oct. Pp. 39-51. 1976. Experimental studies on improving the status of Canada goose populations. Colorado Div. Wildl., Wildl. Res. Rep. Oct. Pp. 31-40. 1981. Waterfowl production surveys. Wildl. Res. Rep., Oct. Pp. 1-14. Colorado Div. Wildl., 1982. Distributional characteristics of some population of Canada geese inhabiting Colorado. Colorado Div. Wildl., Wildl. Res. Rep., Oct. Pp. 19-31. _____________ , R.C. Staffon, and J.F. Corey. 1981. Distribution and harvest of Canada geese nesting along the foothills of Colorado. Colorado Div. Wildl., Spec. Rep. 49. 25 pp. �109 Colorado Division of Wildlife Wildlife Research Report April 1986 JOB PROGRESS REPORT Colorado State Work Plan Job Title: Avian Research 01-00-045 (W-88-R) Project 3 Job 10 Some Population Characteristics Park Period Covered: of Adult Gadwall Molting in North 30 July 1984 through 30 March 1985 Author: Michael Szymczak Personnel: J. Corey, J. Ringe1man, S. Steinert, and M. Szymczak, Colorado Division of Wildlife. ABSTRACT Trapping of gadwall (Anas strepera) in North Park resulted in 684 birds being banded. Projected estimates of survival obtained from real and simulated banding and encounter data indicate the objective of estimating survival (+ 5%) will be achieved by continuing banding through 1985 and analyzing encounter data through the 1985-86 recovery year. ��111 SOME POPULATION CHARACTERISTICS OF ADULT GADWALL MOLTING IN NORTH PARK Michael R. Szymczak P. N. OBJECTIVES 1. To document the distribution of harvest of gadwall banded as flightless young or molting adults in North Park, Colorado. 2. To estimate recovery and survival rates of adult gadwall molting in North Park, Colorado. SEGMENT OBJECTIVES 1. Trap and band 400 adult males, 350 adult females and, if available, up to 100 local gadwall on molting and brood rearing areas in North Park. 2. Submit banding schedules and recapture reports to the U.S. Fish and Wildlife Service's Bird Banding Laboratory. File return information at the Colorado Division of Wildlife Research Center. 3. Prepare progress report. METHODS AND MATERIALS Birds were captured from an air thrust boat at night using a hand-held l2-volt landing light and a long-handled net. All flightless adults were captured, weighed, and measured prior to being banded and released in conjunction with a study of the flightless period in ducks (Szymczak and Ringelman 1984). The band numbers of birds captured that had been previously banded were recorded. Banding schedules and recapture reports were prepared and submitted to the U.S. Fish and Wildlife Service's Bird Banding Laboratory. Informa tion on returning birds recaptured in the same 10-minute grid of banding were filed at the Colorado Division of Wildlife Research Center. Recovery and return and recapture matrices were constructed for adult males and females banded since 1975 and recovered through the 1983-84 hunting season or recaptured through the 1984 banding seasons (Mardekian and McDonald 1981). Data on previous capture success and rates of recovery, return, and recapture were used to construct matrices simulated through the 1985-86 recovery year. Banding and recovery matrices, banding and recapture and return matrices, and the 2 matrices combined were subjected to program ESTIMATE (Brownie et al. 1978) to evaluate the precision of the survival estimate using specific types of band encounters. �112 RESULTS Banding Trapping resulted in 581 adults, 71 locals, and 32 immatures being banded (Table 1). The number of birds trapped at MacFarlane Reservoir exceeded those captured at Walden Reservoir for the first time since banding of gadwall on molting areas began in 1975. The total numbers of adults exceeded the number banded in 1983, but was still short of established sex quotas. Table l. Number of gadwall banded in North Park by location, July through September 1984. Location MacFarlane Reservoir Walden Reservoir Lake John Annex Pole Mountain Reservoir Hebron Ponds Case Flats Totals aCaptured simulations. during Age and sex Local Local males females Adult males Adult females 143 106 40 7 167 93 16 7 1a la 21 0 7 2 34 0 6 1 9 12 0 1 6 1 0 3 380 212 69 21 1 1 296 285 30 41 22 10 684 brood rearing; not used Imm. males in Imm. females survival Totals estimate Survival Estimate Simulations Since 1985 will be the last scheduled year of adult gadwall banding in North Park, probable survival rates were calculated based on real and simulated data to ascertain if goals of estimating survival at +5% would be met on schedule. The confidence intervals of survival estimates (Table 2) indicates objectives will not be met for either sex using only band recoveries; recaptures and returns alone yielded adequate estimates for males but not for females. Recoveries plus recaptures and return resulted in estimates at or exceeding the objective levels for both sexes. It is interesting to note that recapture and return rates combined were equal to recovery rates for males and exceeded those of females. �Table 2. Estimates of survival and recovery rates of adult gadwall banded in North Park. Years 1975-841> Estimate (95% C.l.) 1975-85~ Estimate (95% C.l.) Ag,e/sex Encounter method Parameter 1975-83a Estimate (95% C.l.) Adult male Recovery Survival Recovery 64.89(58.38-71.40) 2.28 (1.84- 2.72) 63.68(57.81-69.55) 2.33 (1.91- 2.75) 62.96(57.61-68.32) 2.38 (1.98- 2.79) Recaptures & returns Survival Recovery 73.03(67.02-79.05) 2.22 (1.80- 2.63) 72.33(67.02-77.63) 2.25 (1.85- 2.64) 71.80(67.05-76.54) 2.29 (1.91- 2.67) Recovery, recapture & return Survival Recovery 69.30(64.90-73.71) 4.48 (3.89- 5.07) 68.45(64.53-72.37) 4.56 (3.99- 5.12) 67.90(64.36-71.44) 4.64 (4.10- 5.18) Recovery Survival Recovery 61.80(51.87-71.73) 1.94 (1.42- 2.47) 60.08(51.33-68.83) 2.02 (1.52- 2.52) 59.06(51.11-67.02) 2.05 (1.57- 2.53) Recaptures & returns Survival Recovery 62.22(48.33-76.11) 3.15 (2.32- 3.99) 64.76(50.93-78.59) 3.48 (2.67- 4.29) 63.87(51.67-76.06) 3.48 (2.73- 4.22) Recovery, recapture & return Survival Recovery 58.83(52.64-65.03) 5.10 (4.21- 6.00) 59.16(53.79-64.53) 5.21 (4.38- 6.04) 58.73(53.89-63.56) 5.32 (4.54- 6.11) Adult female aActua1 banding and recovery data. bActua1 banding and recovery data except simulated 1984 hunting season recoveries. CActua1 banding and recovery data except simulated 1985 bandings, 1985 recaptures and returns, and 1984-85 hunting season recoveries. I-' I-' W �114 LITERATURE CITED Brownie, C., D. Anderson, K. Burnham, and D. Robson. 1978. Statistical U.S. Dep. Inter., Fish inference from band recovery data -- a handbook. and Wildl. Servo Resour. Publ. 131. 2l2pp. Mardekian, S., and L. McDonald. 1981. Simultaneous analysis of band recovery and live-recapture data. J. Wildl. Manage. 45:484-488. Szymczak, M., and J. Ringe1man. 1984. Ecological studies of the flightless period of ducks in Colorado. Colorado Div. Wi1d1., Wi1d1. Res. Rep. Oct. Pp , 13-31. Prepared by 77JJV'g? ~ Michael R. Szymcz~ Wildlife Researcher C � https://cpw.cvlcollections.org/files/original/ec6f8f52233e6ace5b8e8ae26eb920b7.pdf 402e004fcbe054dcb8e310b43702106b PDF Text Text Colorado Division of Wildlife Wildlife Research Report April 1986 115 JOB PROGRESS REPORT State of Project Work Plan Job Title: Colorado 01-03-045 (W-37-R-38) 3 Personnel: l3a _.;;;;.;.=-- Responses of Sage Grouse to Vegetation Period Covered: Author: : Job Avian Research Fertilization 01 January through 31 December 1985 Clait E. Braun David C. Bowden, Marilet Graham, JoAnn Profera, and Tom Remington, Colorado State University; Lee Benson, Wes Boyce, Clait Bra~n, Jack Corey, Warren Cummings, Jack Depperschmidt, Keith Kahler, Steve Porter, Steve Steinert, and Randy VanBuren, Colorado Division of Wildlife ABSTRACT Long-term sage grouse (Centrocercus urophasianus) baseline data collection continued in North Park, Colorado in 1985. The winter of 1984-85 was relatively "normal" but winter 1985-86 started early with heavy snowfall occurring in early November. Numbers of active leks (27) and males counted per lek (25.7) increased from levels in 1984 but were still below "the long-term average. Sage grouse hunter pressure and success (51%) were similar to levels measured in 1984, however, birds per hunter (1.1) decreased. Percent chicks in the fall harvest (54%) was above the long-term average as was percent yearlings (24). However, nesting success (49%) was the lowest since 1981. Timing of hatching was "normal" and about 2 weeks earlier than in 1984. Annual turnover estimates were 53% for adult males and 42% for adult females. The direct (harvest rate) recovery rate for all birds banded .(325) in 1985 was 10%, within the range (7-11%) measured from 1973 through 1984. About 800 sage grouse were harvested on North Park in 1985 by about 700 hunters. The estimated fall population size was slightly over 8,000 sage grouse. A total of 330 ha of sagebrush-dominated rangeland was fertilized at the .rate of 100 lbs of nitrogen (ammonium nitrate) per acre. �117 RESPONSES OF SAGE GROUSE TO VEGETATION FERTILIZATION Clait E. Braun This long-term study to examine the responses of sage grouse to vegetation fertilization in North Park, Colorado was initiated in 1980-81 when it appeared that surface mining of coal would be expanding. The study was scheduled in 2 phases. The objectives of Phase I prior to fertilization/ mitigation by coal companies related to collectiion of baseline data against which the effects of treatment (fertilization) would be evaluated. Thus, the feeding preferences of sage grouse in January-April including the dietary composition of sagebrush (Artemisia spp.) by species and/or subspecies at feeding and random sites was examined. This portiion of the study was completed in 1983 (Remington 1983). With the virtual shutdown of the Kerr and Wyoming Fuels mines in 1983, only the Walden mine has continued to operate. Also, the State Mined Land Reclamation Board has failed to recognize vegetation fertilization as a possible mitigation tool. The Kerr mine increased its production in 1985 when it reached agreement with the Walden mine to jointly fill contracts for coal. Agreement was also reached with the Bureau of Land Management (BLM) and the Colorado Division of Wildlife (CDOW) to jointly fund application of nitrogen fertilizer on about 330 ha of sagebrush-dominated rangelands. Thus, Phase II of the study was initiated in October 1985 with design of the fertilization experiment, application of the fertilizer, and initial selection of a graduate research assistant. P. N. OBJECTIVE Objectives in 1985 initially were throughout North Park through lek data. Once agreement between the added to design the fertilization to monitor the sage grouse population counts, banding, and collection of harvest BLM and CDOW was reached, objectives were experiment and apply the fertilizer. Segment Objectives: 1. Use night-lighting techniques to capture adult and yearling male sage grouse roosting on leks during March-May. Trap and band 300 males and at least 100 females within North Park from March through May. Obtain body weight and other measures of body size from sage grouse at time of capture. 2. Make counts (4/lek) of males and females on all known leks in North Park. 3. Survey areas in North Park to locate new or relocated leks. 4. Collect 10 adult male sage grouse from 4-5 leks during the early (20 Mar 10 Apr) period of the display season. Collect an additional 10 adult males from leks during the latter period of display (15-30 May). Estimate body lipid reserves of collected individuals. Evaluate depletion of lipid reserves during the display season by comparing carcass composition of early and late-collected males (reported in Work Plan 3, Job 15). �118 5. Clear established pellet group transects (30) during September-October and June. 6. Collect population data through use of wing barrels and check stations during September-October. 7. Fertilize up to 990 acres or area equivalent to total area under permit to coal mining companies of sagebrush-dominated rangelands with ammonium nitrate (33.5% N) at the rate of 100 pounds of nitrogen per acre. Fertilizer application will be in fall (between 15 Sep and 10 Oct) with areas to be fertilized randomly selected. Fertilizer will be applied on up to 31 strips, 528 feet wide by 2,640 feet long with each treated strip being paired with an adjacent untreated strip of equal size. 8. Prepare progress reports. scientific meetings. Present pertinent findings at appropriate METHODS Sage grouse were located where they roosted along roads, trails, and on leks and captured in late winter and spring using night-lighting techniques as described by Giesen et al. (1982). All birds captured were classified to age and sex (Beck et al. 1975), weighed to the nearest 1 or 10 grams, and selected birds were measured. A sample of males was collected by asphixiation during 2 time intervals for development of physiological indices (Work Plan 3, Job 15). Counts of birds present on leks in April and May were made following procedures outlined by Braun and Beck (1976). Pellet group transects were cleared and all pellets were classified following Schoenberg (1980). Harvest data were collected through use of hunter-check stations and volunteer wing collection stations (Hoffman and Braun 1975). The fertilization experiment was designed in consultation with David C. Bowden and Len Carpenter. Bids were solicited in September for fertilizer and application. Study areas (3) were selected in consultation with Chuck Cesar and Lee Upham of the BLM, J. Allen White of the U.S. Department of Agriculture, Soil Conservation Service (SCS), and Tom Remington. Treatment and control blocks were randomly located within study areas and oriented on an east-west basis. Plots were located on aerial photographs (overlays of soils maps) and starting points were at USGS section corners. Directions were determined with a transit and all distances were measured with a 300-foot steel tape. All plot corners were marked with steel posts. Fertilizer was applied with a spreader mounted on a truck (Area C and part of Area B) or spreaders pulled by tractors (Area A and part of Area B). Fertilization was accomplished between 5 and 12 November. RESULTS AND DISCUSSION Lek Counts Number of active leks and males counted per lek increased (22 to 27 and 21.2 to 25.7, respectively) from 1984 to 1985 (Tables 1, 2). However, the number of males counted per lek was still the 2nd lowest count recorded in the 1973-85 interval (Table 2). Winter 1984-85 was "normal" and over winter sage grouse survival should have been "normal". This, coupled with good production , �119 Table 1- Counts of sage grouse on leks, North Park, 1985. Lek , Counts Number of Males Females Alkali Lake Arapahoe Aspen Bighorn Boettcher Jct. Buteo Canuck Case Flats Cherokee Cheyenne Coalmont Deer Creek Delaney Butte Denmark Eagle Fish Hatchery Hawk Hound Lost Creek III Migan Owl Creek Perdiz Peregrine Prague Ptar Railroad Ram Raven Ridge Road Riley 6 4 4 3 7 5 4 1 2 5 7+ 5 7 4 2 5 4 6 5 3 2 6 4 5 3 7 1 6 9 5 34 18 8 4 41 11 14 0 0 54 61 45 68 0 0 26 0 11 0 2 0 10 0 0 8 46 0 14 30 20 24 18 0 1 35 9 1 0 0 13 57 5 36 0 0 17 0 31 0 1 0 6 0 0 0 42 0 1 53 0 Spring Creek III Spring Creek 112 Thrasher Turkey Ute Walden Wattenburg 112 7 5 4 5 2 5 5 66 12 37 35 2 7 11 57 0 25 1 0 34 2 Date of high count Females Males 09 May 24 Apr 16 May 21 Apr 15 Apr 16 & 23 Apr 25 Apr 12 May All dates .2 & 16 May 29 Apr 02 May 02 May All dates All dates 12 May All dates 03 May All dates 09 Apr All dates 28 Apr All dates All dates 30 Apr 25 Apr 25 Apr 28 Apr 12 May 29 Apr, 15 May 02 May 17 Apr 30 Apr 28 Apr 16 Apr 11 Apr 29 Apr 12 Apr 24 Apr All dates 21 Apr 15 Apr 23 Apr 25 Apr 12 May All dates 18 Apr 15 Apr 17 Apr 12 Apr All dates All dates 12 Apr All dates 11 Apr All dates 09 Apr All dates 16 Apr All dates All dates All dates 14 Apr 25 Apr 28 Apr 13 Apr All dates 11 Apr All dates 15 Apr 28 Apr All dates 11 Apr 10 Apr �Table 2. Trends in peak counts of male sage grouse, North Park, 1973-85. f-' N 0 Lek Alkali Lake Arapahoe (1977) Aspen (1977) Bighorn (1976) Boettcher Lake Jet. Buteo (1981) Canuck (1974) Case Flats (1978) Cheyenne (1978) Coalmont Cowdrey #5 (1973) Deer Creek Delaney Butte Denmark (1977) Eagle (1978) Fish Hatchery Hound (1974) Lost Creek 11 Lost Creek 12 Migan (1979) Monahan Draw Owl Creek (1976) Perdiz (1979) Peregrine (1978) Prague (1978) Pronghorn (1977) Ptar (1978) Railroad (1975) Raven (1977) Ridge Road Riley (1973) Roth (1974) Spring Creek 11 Spring Creek (12 Spring Creek #4 Thrasher (1978) Turkey (1982) Ute-North (1979) Ute-South (1978) Walden Reservoir (1973) Wattenburg #2 Average/1ek 1973 1974 81 --- ----59 ----- -- 47 30 37 4 --- 72 -49 -29 --- 27 19 11 13 -- 1975 68 ---50 -22 -- --29 9 27 0 -- 67 --62 --69 10 69 0 1 33 18 0 --13 -- 11 -- ----- --- --3 --- ------ --36 --33 12 15 10 33 15 10 -46 39 9 -- -- -- -- 38 24 33.1 --- -- -- 37 22 27.7 --6 ------- 86 -- 27 12 14 49 14 3 1976 1977 ..·1978- 39 36 46 21 52 76 -- - 46 62 -- 30 -- -- 28 5 36 0 -- -- 81 27 33 0 -27 --- 21 5 31 0 58 -- 64 21 47 0 --1 --1 14 ----15 --41 -------87 --32 18 9 49 11 8 16 73 32 9 8 45 23 3 56 61 24 47 86 --21 4 127 21 1 41 4 80 11 97 28 43 0 --0 15 --17 35 10 50 35 94 65 5 1 76 63 3 65 --34 ------37 --26 18 16 12 18 21 30.9 31.9 31.1 39.5 --- -- -- -- -- -- -17 Years 1979 1980 1981 72 43 12 31 107 39 29 10 14 113 21 8 137 25 0 43 12 136 2 82 25 30 0 70 0 9 16 20 43 10 55 62 63 67 41 2 59 63 0 84 0 94 32 0 28 28 144 0 78 27 20 0 53 0 2 8 1 34 0 20 40 43 60 54 1 73 47 0 51 68 32 38 26 106 22 16 0 80 36 0 52 23 109 0 67 23 29 0 53 0 3 23 9 26 0 19 44 49 41 48 0 53 54 0 50 7 13 30 21 0 26 17 0 11 42 16 43.5 -- -- --18 -- 40.1 --0 40.9 1982 60 21 27 27 92 19 33 0 104 52 0 66 60 71 0 50 20 36 0 27 0 0 27 13 7 0 24 33 44 30 58 0 59 56 0 55 22 0 14 42 27 41.2 1983 62 25 22 15 100 28 31 0 101 62 0 47 72 24 0 63 22 26 0 11 0 0 21 12 3 0 1 64 29 53 40 0 59 37 0 59 59 0 5 40 27 39.4 1984 1985 36 18 11 0 22 0 0 0 38 43 0 14 54 8 0 10 19 2 0 0 0 0 8 11 0 0 0 11 5 31 0 0 50 8 0 NC 47 0 2 0 18 34 18 8 4 41 11 14 0 54 61 0 45 68 0 0 26 11 0 0 2 0 0 10 0 0 0 8 46 14 30 20 0 66 12 0 37 35 0 2 7 11 21.2 25.7 �121 (57% chicks, 1.9 chicks/hen in the fall 1984 harvest) in 1984 probably resulted in the increased number of active leks and males counted per lek in 1985. Earlier (Braun 1984), it was hypothesized that number of active leks and males per lek decreased in Spring 1984 because males had poor energy reserves following the harsh winter of 1983-84. This hypothesis suggests that similar numbers of males were alive in Spring 1984 as in Spring 1983. Further, if the hypothesis was true and the winter of 1984-85 was "normal", counts of males on leks and active leks would return to the 1983 levels (Braun 1984). This did not happen as Spring 1985 counts were still 35% lower (25.7 vs. 39.4) than in 1983. Therefore, it is concluded that significant numbers of male (and probably female) sage grouse died over winter 1983-84. Those that did survive displayed late and only sporatica11y on leks in Spring 1984 (Braun 1984). Harvest Data Collection r The sage grouse hunting season in North Park opened one-half hour before sunrise on the 2nd Saturday in September (14th) in 1985 and closed at sunset on 6 October, and was 1 week shorter than in the 1982-84 interval (Table 3). The 1 week shorter season in 1985 was the result of the change in when the 2nd Saturday in September occurred. The daily bag limit was 3 with a possession limit of 6, the same limits that have been in effect since 1976 (Table 3). Table 3. 1973-85. Sage grouse season length and bag limits, North Park, Colorado, Years Season length (days) 1973-74 1975 197& 1977-80 1981 1982-84 1985 3 9 9 16 23 30 23 Bag/Eossession limit 2/4 2/4 3/6 3/6 3/6 3/6 3/6 Three check stations were operated, one at Willow Creek Pass on Colorado 125, one east of Gould on Colorado 14, and one at Muddy Pass near the junction of Colorado 14 with U.S. 40 during both days (Sunday only at Muddy Pass) of the opening weekend versus both days of the opening weekend and the 2nd Sunday in earlier years (1975-83 except neither the Gould nor Stateline check stations were operated on the 2nd Sunday in 1976; the Stateline check station was not operated on the 2nd Sunday in 1979, and the Muddy Pass check station was operated only on the first Sunday in 1980). The Stateline (operated, from 1974 through 1979) check station was not operated. As in previous years of operation (1974-84 for Willow Creek, 1976 and 1979-84 for Gould, 1974 and 1980 for Muddy Pass), each station was open from about 1000 to 1800 hours MDT �122 depending upon traffic volume. These stations were staffed with 1-2 research and 2-3 regional personnel (except Muddy Pass). In addition, either 2-3 students from the Wildlife Management Techniques class at Colorado State University assisted at each check station. Data obtained per party were: county of origin, number of hunters, hours hunted (total of all hunters in each party), birds observed, birds bagged, birds lost, number of banded birds and location where each was harvested, and area hunted within North Park. One wing was obtained from all birds possible. Data collected from grouse hunters who hunted in areas other than North Park were tabulated and analyzed separately. Volunteer wing collection barrels and signs were placed along Colorado 14 northeast of Muddy Pass and near the top of Cameron Pass, north of Three-way on Colorado 127, and at Willow Creek Pass on Colorado 125. Volunteer wing collection barrels were available to hunters during the entire season near Three-way and for all days that check stations were not operated east of Gould, Muddy Pass, and at Willow Creek Pass. Barrels were not placed at Arapahoe Creek (only in 1982-83) nor Seymour Reservoir (1982). Some wings were also obtained through field checks by Arapahoe National Wildlife Refuge and CDOW personnel. Hunter and Harvest Data, Check Stations.--Numbers of hunters checked increased in 1985 due in part to operation of the Muddy Pass check station (9 hunters checked) while number of birds harvested and birds per hunter declined (Table 4). Number of grouse observed remained essentially unchanged while crippling loss doubled (5.5%) from 1984. These data (Table 4) suggest that the fall population of sage grouse was similar to that in fall 1984. Hunter pressure during opening weekend remained significantly lower than in 1974-75 while the sage grouse population was substantially lower than in 1979-83. Table 4. Sage grouse harvest statistics, North Park, Colorado, 1974-85. Year N hunters checked N birds observed N birds harvested Hunter efficiency (%) 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 730 738 595 353 350 521 567 523 515 474 396 465 6,062 5,735 3,393 3,303 4,922 6,910 5,000 6,189 3,886 4,961 2,814 2,894 785 551 459 385 480 982 794 695 570 792 517 492 12.9 9.6 13.5 11. 7 9.8 14.2 15.9 11.2 14.7 16.0 18.4 17.0 Crippling loss (%) 5.1 7.1 5.7 10.6 5.7 4.7 4.3 5.4 4.0 4.8 2.6 5.5 Birds per hunter 1.1 0.7 0.8 1.1 1.4 1.9 1.4 1.3 1.1 1.7 1.3 1.1 �123 Hunter Success.--Hunter success in 1985 decreased slightly (51 vs. 53%) from 1984 (Table 5). However, the proportion of the hunters achieving the bag limit of 3 birds during at least 1 day in 1985 decreased markedly (11 vs. 20%) from 1984. Birds per hunter and hunter success have correlated well in all years. Of interest are the data that indicate that no more than 30% of all hunters achieve the bag limit in 1 day and that only 1-5% of all hunters achieve a 2-day limit on the opening weekend. Table 5. Sage grouse hunter successa, North Park, Colorado, 1974-85. Year Successful (%) 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 59 46 40 54 56 68 60 62 52 65 53 51 b Achieved bag limit (%)c 2 days 1 da! 27 20 9 15 19 30 24 20 16 27 20 11 5 1 <1 1 5 5 2 2 2 3 3 3 Birds per hunter Limit 1.1 0.7 0.8 1.1 1.4 1.9 1.4 1.3 1.1 1.7 1.3 1.1 2/4 2/4 3/6 3/6 3/6 3/6 3/6 3/6 3/6 3/6 3/6 3/6 aDerived from check station data. bHarvested at least 1 sage grouse. cPercent of total hunters checked. Hunter Origin.--Vehic1e license prefixes were recorded at check stations to ascertain origin of hunting parties. About one-third to one-half of all hunters originated in the Denver metropolitan area each year with Larimer County being the single most important point of origin (20-33% of all hunting parties). Boulder and Weld counties were also important contributors of sage grouse hunters to North Park (Table 6). It should be noted that few hunters were checked from Jackson County even though they comprised about 10% of all persons obtaining hunting permits during the 1974-77 special seasons in North Park. Local hunters were not checked as check stations were placed at exits to North Park. Time Distribution of Harvest.--Sage grouse wings were obtained at check stations, wing barrels, and field checks. In 1985, 70% were obtained during the first weekend and 6% from the last weekend (Table 7). With liberalization of the season starting in 1977 (9 to 16 days), the proportion of the harvest that occurred in the opening weekend decreased from 92-99% to 61-73% (Table 7). This decrease reflects reduced hunter pressure during the opening weekend. It is important to note that some sage grouse hunters are active throughout the entire season even though hunter pressure is low after the 2nd weekend. �124 Table 6. 1974-85. Origina of sage grouse hunting parties, North Park, Colorado, Countx: (%) Year Denver Metrob Larimer Boulder 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 50 52 53 47 52 33 42 39 43 50 50 44 24 20 20 23 21 31 26 33 27 21 22 26 10 11 13 16 12 17 13 11 12 8 6 11 All other Weld 5 6 6 5 4 9 7 7 6 6 8 7 11 11 8 9 11 10 12 10 12 15 14 12 aAscertained from vehicle license prefixes. bAdams, Arapahoe, Denver, Douglas, and Jefferson counties. Time distribution of sage grouse wings received, North Park, Table 7. Colorado, 1974-85. Year 1 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 99 94 92 67 62 73 69 72 61 72 70 70 a Weekend (% received) 2 3 4 4 7 13 6 11 12 8 13 6 7 9 8 10 6 9 6 7 3 6 4 2 4 2 4 6 5 Season length (days) 2 2 3 (closed) 3 9 9 16 16 16 16 23 30 30 30 23 aTotals do not equal 100% as wings collected during the week are not included. �125 Harvest and Hunting Pressure.--Hunter pressure was greatest in the Lake John, Peterson Ridge-MacFarlane Reservoir, Ridge Road, and Spring Creek-Owl Ridge areas in 1985. However, harvest did not follow hunter pressure as the Ridge Road, Peterson Ridge-MacFarlane Reservoir, Lake John, and Walden Reservoir areas were the leading harvest sites in 1985 (Table 8). Of interest was the increase in hunter pressure and harvest in the Walden Reservoir area even though the 2 known leks in this area (Hound, Walden) had low counts of males in 1985 (Table 2). However, hunter pressure and harvest has not paralleled changes in sage grouse populations (as measured by lek counts) in North Park during 1974-85. Age and Sex Structure.--Immatures comprised 54% of the fall harvest in 1985, slightly less than the excellent production recorded in 1983 and 1984 but above the long-term average of 51% (Table 9). The percentage of yearlings in the harvest (23%) was also above the long-term average while the percentage of adults (23%) increased from that measured in 1984. These data indicate that nesting success and survival of young in 1985 were good and that survival of young produced in 1984 was also good. The sex ratio at hatching appears to approximate 1:1 although chick females predominate (52:48) in fall harvest samples. Swenson (1985) suggests that this disparity is because of increased energetic demands for growth of chick males. Also, he suggests the disparity is greatest during years unfavorable for juvenile survival and in poorer habitats. The argument that the disparity is greatest in years of poor juvenile survival is not supported by data from North Park as juvenile males comprised 50% of the juvenile proportion of the harvest in only 1 of 12 years despite variable reproductive success. The long-term data from North Park indicate that in the average year about 51% of the fall population of sage grouse is comprised of young of the year, 22% are young of the previous year, and 27% of the birds are at least 2 years of age. With this age structure, it is obvious that poor production in 1 or more consecutive years would result in a markedly reduced population of sage grouse. The data (Table 9) also document that sex ratios of yearlings and adults strongly favor females (yearlings = 63:37, adults = 72:28). Thus, poor production would most noticeably affect males as yearlings comprise about 52% of the spring population of this sex each year. Poor production should be immediately noticed in counts of males on leks the following spring. Without recruitment, numbers of males on leks could decrease by 50% in 1 year. Nestin Success and Production.--Primary feather molts of hens harvested were classified number of primaries from the previous year still retained) and compared to the primary feather molt patterns of males of the same age class harvested in the same time interval. Estimated nest success of both adult and yearling hens decreased markedly in 1985 (Table 10) from levels measured in 1984. The data available indicate that at least one-half of the adult hens are successful in most years (except 1981) while success of yearling hens is highly variable (>50% in only 3 of 12 years). In years when at least 50% of the yearlings are successful (1979, 1983, 1984), chicks per hen increase as does hunter success as measured by birds per hunter data from check stations (except in 1984). It is probable that the percentage of successfully nesting females was overestimated in some years (1974, 1975, 1977, and possibly in 1984) and underestimated in others (1980, 1981, 1985). Because this estimate is based on the molt patterns of primary feathers which are affected by timing of breeding (males) and nesting (females), it is logical to assume that timing �t-' N (J'\ Table 8. Sage grouse harvest and hunting pressure (% of total) within North Park, Colorado, 1974-85a,b. -Year 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 Independence Mountain Hary. Hunt. 4.7 3.1 2.3 1.4 0.9 3.5 2.0 3.3 2.3 4.7 1.0 0.9 6.0 3.8 3.0 0.5 0.0 2.7 2.3 1.7 3.2 5.5 5.2 0.0 Lake John Hunt. Harv. Walden Reservoir Hary. Hunt. RidGe Road Hunt. Harv. 18.1 24.3 28.2 28.9 29.4 23.0 16.6 15.9 19.7 26.6 26.0 28.4 12.4 14.2 13.0 15.8 18.3 8.3 10.2 4.2 7.8 9.6 6.9 11.8 9.6 11.1 13.0 16.9 14.6 10.2 14.0 14.8 18.4 11.9 15.3 18.5 16.4 24.7 21.3 23.9 18.1 19.7 15.5 14.4 20.6 26.9 22.5 17.7 15.4 16.7 12.0 12.2 10.6 7.5 14.9 4.0 8.2 12.0 5.0 13.8 11.3 15.6 21.1 20.8 13.7 13.8 8.1 27.6 22.1 8.8 20.6 24.4 Pole Mountain Harv. Hunt. 4.7 1.8 2.9 3.2 2.0 4.0 1.4 1.7 1.6 0.2 1.0 1.3 3.8 2.0 4.5 6.5 2.5 4.2 2.6 0.7 0.1 0.8 0.0 2.0 aData from check stations only. bTota1s may not approximate 100% a9 in most years some hunters and birds harvested hunted in more than one area. Peterson Ridge-MacFar1ane Res. Hunt. Harv. Spring Creek-Owl RidGe Hary. Hunt. 20.3 17.2 18.5 16.3 21.1 30.0 29.0 27.7 21.9 18.9 20.6 23.0 17.9 13.6 10.9 8.0 6.0 12.5 14.1 14.9 18.6 15.7 15.5 15.5 19.6 14.3 21.3 12.7 36.7 33.3 32.7 26.3 21.2 21.5 24.7 22.6 could not be allocated 13.4 6.2 6.3 8.1 3.1 9.4 13.0 12.9 12.1 13.9 13.4 10.8 to a particular Michigan River SE Hary. Hunt. 2.2 2.6 3.1 2.0 1.7 2.5 2.8 1.1 1.4 3.6 1.8 1.3 1.3 3.8 5.9 2.1 5.6 3.7 1.5 1.7 2.7 3.4 1.7 1.6 EaGle H111 Harv. Hunt. 10.0 10.4 8.0 7.4 6.3 6.0 9.9 11.5 8.4 8.7 9.4 7.7 zone, and some hunters 12.8 9.8 5.0 13.2 9.6 5.7 9.4 10.5 9.3 7.2 6.7 7.1 �-~ Table 9. Age and sex composition of the sage grouse harvest, North Park, Colorado, 1974-85. Immatures Females Males Year N 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 171 101 104 136 184 306 207 234 197 295 236 163 Avg. % 48.9 47.6 49.5 46.9 47.8 49.0 44.9 47.9 50.1 45.6 48.5 46.2 47.6 .t! 179 111 106 154 201 318 254 255 196 352 251 190 % 51.1 52.4 50.5 53.1 52.2 51.0 55.1 52.1 49.9 54.4 51.5 53.8 52.4 Total Yearl1nss Females Males N % 1i 350 212 210 290 385 624 461 489 393 647 487 353 50.1 42.0 42.3 45.9 53.2 57.7 49.1 47.4 50.6 57.4 57.0 53.6 49 52 46 47 62 102 80 78 53 90 68 47 51.4 % 35.5 46.8 39.3 38.2 43.4 39.8 32.0 34.1 39.8 36.7 34.0 29.6 36.8 Total Ii % li % N 89 59 71 76 81 154 170 151 80 155 132 112 64.5 53.2 60.7 61.8 56.6 60.2 68.0 65.9 60.2 63.3 66.0 70.4 138 111 117 123 143 256 250 229 133 245 200 159 19.8 22.0 23.5 19.5 19.8 23.7 26.6 22.3 17.1 21.7 23.4 24.2 45 55 49 48 67 72 70 87 81 53 37 33 63.2 Adults Females Males 22.1 % 21.4 30.2 28.8 21.9 34.2 35.8 30.7 27.7 32.2 22.5 22.0 22.6 27.6 Sample Total N % N 165 127 121 171 129 129 158 227 170 183 131 113 78.6 69.8 71.2 78.1 65.8 64.2 69.3 72.3 67.7 77.5 78.0 77.4 210 182 170 219 196 201 228 314 251 236 168 146 72.4 % 30.1 36.0 34.2 34.6 27.1 18.6 24.3 30.4 32.3 20.9 19.6 22.2 size 698 505 497 632 724 1,081 939 1,032 777 1,128 855 658 26.5 I-' N -...J �128 of breeding events (weather controlled) is the most important factor involved. Thus, underestimates are derived following "early" springs while late springs (for example, 1984) may provide the most reliable estimate. The evidence indicates that, at least in North Park, the percentage of juveniles in the fall harvest closely approximates overall nesting success. Also, hunter success (birds/hunter) closely follows the young per hen ratio in the harvest. Table 10. Sage grouse nesting success and production rates, North Park, Colorado, 1974-85. Year 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 Estimated nestin~ success YearAll Adults lings hens 65 53 53 59 60 65 56 37 59 68 71 54 46 39 27 33 38 56 31 22 38 51 55 44 59 49 43 50 51 60 43 31 52 58 63 49 Young in harvest (%) 50 42 42 46 53 58 49 47 51 57 57 54 Young per hen 1.4 1.1 1.1 1.2 1.8 2.2 1.4 1.3 1.5 1.9 1.9 1.6 Young per Birds successful . per hen hunter 2.3 2.3 2.5 2.0 3.6 3.7 3.3 4.1 3.0 3.2 2.9 3.1 1.1 0.7 0.8 1.1 1.4 1.9 1.4 1.3 1.1 1.7 1.3 1.1 Timing of Hatching.--Timing of hatching was "normal" in 1985 and started 2 weeks earlier than in 1984 (Table 11). Timing of nesting events was later each year from 1982 through 1984 because of heavy and late lying snow cover. In contrast, 1981 was an "early" year and over 80% of the hatch occurred prior to 21 June. Data in Tables 10 and 11 suggest that nesting success and production are better in "late" years (1979, 1982, 1983, 1984) than in "early" years (1976, 1977, 1981) although 1975 appears to be an exception. This relationship appears to be related to moisture conditions which could affect (improve) vegetative cover during the nesting period and/or provide adequate forage (succulent forage, insects) near nesting sites for young chicks. Theoretically, early chick survival should be enhanced if successful hens did not immediately travel to meadows along streams to forage following hatching of their clutches. 1 , �129 Table II. Estimated timing of hatching of sage grouse eggs, North Park, Colorado, 1974-85. 74 Interval 75 76 Year (Percent of all chicks) 77 78 79 80 82 81 11-17 May 18-24 May 83 84 85 2 <1 <1 6 1 <1 <1 22 4 2 6 <1 <1 2 14 1 12 21 25 1 4 31 30 20 9 3 35 28 9 <1 16 21 24 27 12 27 19 19 26 22 1 31 20 14 10 8 34 43 6 19 27 16 25 21 11 8 8 17 21 8 8 19 51 15 15 6 2 7 9 11 2 8 15 26 6 10 7 <1 <1 4 <1 <1 2 <1 <1 3 <1 <1 <1 <1 <1 3 2 5 2 <1 3 2 <1 <1 3 1-7 Jun . ~ 16 8-14 Jun 18 15-21 r r 34 22-28 Jun 29 Jun-5 6-12 Jul Ju3 16 } 6 13-19 Jul 20-26 Jul 27 Jul-2 Aug 6 Annual Turnover.--Estimates of annual turnover rates were calculated for adult males (Table 12) and adult females (Table 13). The estimated annual mortality rate was 53% (1974-85 average) for adult males and 42% for adult females. These estimates are based on the premise that the percentage of yearlings in the poplation should equal the annual mortality of adults if the population is stable. However, if the population was increasing then the percentage of yearlings in the population would overestimate adult mortality rates. Because the sage grouse population in North Park obviously increased in the 1974-85 interval (except 1984), the calculated mortality rates for both adult males and females may be biased upward by several percentage points. �l30 Table 12. Estimated annual turnover of adult male sage grouse, North Park, Colorado, 1974-85a• In harvest Adults Year N % N % Sample size 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 45 55 49 48 67 72 70 87 81 53 37 33 48 51 52 50 52 41 47 53 60 37 35 41 49 52 46 47 62 102 80 78 53 90 68 47 52 49 48 50 48 59 53 47 40 63 65 59 94 107 95 95 129 174 150 165 134 143 105 80 Avg. Year1inss 47 ~ , , 53 Estimated annual mortality of adult males in a stable population ::z: 53% aData from wing collections only. Table 13. Estimated annual turnover of adult female sage grouse, North Park, Colorado, 1974-85a• In harvest Adults Year N % N % Sample size 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 165 127 121 171 129 129 158 227 170 183 131 113 65 68 63 69 61 46 48 60 68 54 50 50 89 59 71 76 81 154 170 151 80 155 132 112 35 32 37 31 39 54 52 40 32 46 50 50 254 186 192 247 210 283 328 378 250 338 263 225 Avg. Yearlinss 58 42 Estimated annual mortality of adult females in a stable population = 42% aData from wing collections only. ~ �131 Banding and Band Recoveries.--During 1985, 325 sage grouse were banded in North Park of which only 74 were hens (Table 14). This was a substantial decrease from the 234 hens banded in 1984, the result of a severe winter and a late spring which concentrated hens and kept them in flocks later in the Spring. The high number of hens banded in 1977 (234) was the result of good access not severe winter weather. Number of males banded in 1985 (251) increased from 1984 (194), the result of more males on grounds and which roosted there at night unlike the situation in 1984. Table 14. Sage grouse banding data, North Park, 1973-85. Number banded D f r Year 1- Females 2+ 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 41 22 62 71 101 32 48 72 31 42 33 123 46 68 27 68 74 133 22 52 94 38 69 32 111 28 All 1- Males 2+ All 109 49 130 145 234 54 100 166 69 102 65 234 74 80 54 138 120 183 106 111 127 110 110 152 102 163 99 88 153 114 123 98 146 173 190 190 157 92 88 179 142 291 234 306 204 257 300 300 300 309 194 251 Recoveries were received from 64 banded sage grouse in 1985 (23 females, 41 males) of which all were shot by hunters. Recoveries represented bandings in 1980-85 for males and 1982-85 for females (Tables 15, 16). Of interest was the recovery of the oldest (7+) male reported to date. Of the male recoveries, 85% (35 of 41) were from 1984-85 bandings while 87% of the female recoveries were from 1984-85. In 1984 following the severe winter of 83-84, 87% (34 of 39) of the male recoveries were from bandings in 1983-84 while 75% (15 of 20) of the female recoveries were from bandings in 1983-84. These data do not suggest that banded birds survived the 1983-84 winter poorly. Of the total recoveries in 1983 prior to the severe winter of 1983-84, 55% (31 of 56) were from bandings in 1983. The respective first-year recoveries in 1984 (following the severe winter of 1983-84) and 1985 were 54 and 53% again suggesting that the severe winter of 1983-84 did not result in large losses of banded birds. This strongly suggests that the low counts of males on leks in 1984 was not the result of low survival of adults. Older males were in the population at the rate experienced in earlier years of the study. Thus, it can be reasonably concluded that the low counts of males on leks in spring 1984 was because males failed to attend leks. This failure to attend leks was probably the result of poor body condition caused by the harsh winter of 1983-84 (J. Hupp, unpub1. data). �132 Table 15. 1973-85. Age Year Male sage grouse banding and recovery data, North Park, Colorado, N banded 1 2 N recovered (lear) 3 4 = Yearlings 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 80 54 138 120 183 106 111 127 110 110 152 102 163 6 6 18 16 19 13 13 13 14 7 16 12 17 4 5 6 5 10 4 4 5 5 1 10 4 6 3 7 6 7 4 0 3 4 3 2 Adults 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 99 88 153 114 123 98 146 173 190 190 157 92 88 7 8 10 16 12 9 13 9 16 19 15 8 9 4 5 4 3 2 9 10 5 5 6 4 5 1 1 2 2 3 3 3 2 2 2 0 1 1 2 1 2 1 1 1 0 1 5 6 0 0 0 1 1 0 0 0 0 1 0 1 1 1· 0 0 0 1 1 0 0 0 0 3 0 3 1 0 1 1 0 1 0 0 0 0 0 1 0 0 1 0 0 0 0 1 �133 Female sage grouse banding and recovery data, North Park, Table 16. Colorado, 1973-85. r D Age Year N banded 1 2 3 Yearlings 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 41 22 62 71 101 32 48 72 31 42 33 123 46 2 2 6 2 6 5 7 8 4 2 0 6 6 2 1 2 4 5 0 4 4 3 0 2 8 4 1 1 2 6 1 0 1 68 27 68 74 133 22 52 94 38 60 32 5 2 6 2 13 1 3 8 5 5 0 6 2 1 0 2 2 2 0 3 6 1 3 1 4 0 0 4 1 4 0 0 2 0 0 1 Adults 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 III 28 0 2 0 N recovered (lear) 4 5 6 1 0 2 1 1 0 0 3 0 0 2 0 0 0 0 0 1 0 0 1 1 1 3 2 0 0 1 0 2 1 0 1 0 1 0 0 1 0 0 0 0 1 1 0 0 0 0 0 0 1 0 0 1 0 7 8 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 �134 Direct recovery rates (i.e., harvest rates) of sage grouse banded in 1985 varied from 7 (adult females) to 13% (yearling females) and averaged 10% for all birds banded. These rates are within the range documented in the 1973-84 interval (Table 17) and indicate that liberalization of season length and bag limits have not markedly affected the harvest rate of sage grouse ~n North Park. Table 17. 1974-85. Direct recovery rates of banded sage grouse, North Park, Colorado, Year Females (%) 2+ III 1- 1- 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 5 9 10 3 6 16 15 11 13 5 0 5 13 7 11 13 13 11 13 12 10 13 6 11 12 10 7 7 9 3 10 4 6 9 13 8 0 5 7 6 8 9 3 8 11 10 10 13 7 0 5 11 Males (%) 2+ III 7 9 7 14 9 10 9 6 8 10 10 9 10 7 10 10 14 10 12 10 8 10 8 10 10 10 All birds (%) 7 9 10 9 9 11 10 8 11 8 8 7 10 Season Limits Lensth 3 3 9 9 16 16 16 16 23 30 30 30 23 2/4 2/4 2/4 3/6 3/6 3/6 3/6 3/6 3/6 3/6 3/6 3/6 3/6 Estimate of Total Harvest.--Tota1 harvest in 1974-77 was estimated from a 100% survey of all sage grouse hunters in North Park. This was possible because all sage grouse hunters were required to obtain a free permit prior to hunting. Because the Willow Creek Pass check station was the only one that was operated each year from 1974 through 1985, the number of hunters checked at it on opening weekend (2 days) was used to estimate total hunters from 1978 through 1985. From 1974 through 1977, an average of 30.7% of all North Park sage grouse hunters were checked at Willow Creek Pass. Thus, 0.307 was divided into the number of hunters checked at Willow Creek Pass each year from 1978 through 1985 to derive an estimate of total sage grouse hunters in North Park (Table 18). This number was then multiplied by the birds per hunter value calculated from check station data each year to derive an estimate of total harvest (Table 18). These data indicate that total number of hunters declined from about 1,000 in 1974-76 to about 600-700. This is in agreement with the total number of hunters contacted at all check stations (Table 4) as total number of hunters contacted at check stations has declined from about 700 to 400-500. The data also suggest that total harvest has declined (Table 18). However, slightly more wings were received from hunters in 1979, 1981, 1982, and 1984 than the estimated harvest. Thus, the harvest was underestimated by the method used. This underestimate could be in the magnitude of 200-300 birds each year. It is reasonable to assume that the total harvest of sage grouse in North Park approximates 1,000-1,500 birds per year. 4 ~ ! , �135 Estimated total number of sage grouse hunters and total harvest, Table 18. North Park, Colorado, 1974-85. ~ r r D r Year Na hunters contacted 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 378 374 317 149 226 173 234 208 203 216 200 219 Total hunters 966b 1,187b 979b 845b 736d 564d 762d 678d 66ld 704d 65ld 713d Birds per c hunter 1.1 0.7 0.8 1.1 1.4 1.9 1.4 1.3 1.1 1.7 1.3 1.1 Total harvest 1,193b 1,075b 847b 1,070b 1,030e 1,072e 1,067e 88le 727e 1,197e 846e 756 Total wings received 698 505 497 632 724 1,081 939 1,032 777 1,128 855 658 aWillow Creek Pass Check Station only during opening weekend. bDerived from questionnaire survey of all sage grouse hunters in North Park. cDerived from check station data (total birds checked f total hunters checked) • dDerived by dividing percent of all hunters checked at Willow Creek Pass in 1974-77 (0.307) into number of hunters checked at Willow Creek Pass in each year. eDerived by multiplying total number of hunters by birds per hunter. Estimated Population Size.--Population size in spring was estimated assuming that peak lek counts represented 60% of all males associated with each lek and that 90% of all leks were located and counted. The estimated total number of males was multiplied by 2 to derive the total number of hens as harvest data (Table 9) indicate that females comprise 68% of the total adults and yearlings. Fall population size was estimated by dividing the percent of adults and yearlings in the fall harvest into the spring population estimate. The data available (Table 19) indicate that spring sage grouse populations in North Park increased from about 3,000 birds in 1974-75 to 7,000-8,000 in 1978-79 and were relatively stable at 6,000-7,000 birds from 1980 through 1983. The spring population markedly decreased in 1984 even though harvest, production, and recovery data suggest that the decline was not marked. Lek counts in 1985, while slightly up over 1984, did not return to 1983 levels. Thus, the decrease in Spring 1984 had to be real. Given "normal" weather in winter 1985-86, the spring 1986 counts of males on leks should increase. Size of the fall sage grouse population in North Park has fluctuated from about 6,000 to 20,000 birds. This reflects the relatively short life span and high annual turnover (53% for males, 42% for females) of sage grouse and the importance of nesting success and production. �136 Table 19. Estimates of spring and fall sage grouse population size, North Park, Colorado, 1974-85. Year N ieks Males per 1ek Total ma1esa 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 19 19 21 26 34 35 30 31 31 31 22 27 28 31 32 31 40 44 40 41 41 39 21 26 987 1,092 1,219 1,508 2,479 2,774 2,211 2,312 2,312 2,210 805 1,256 Estimated Spring Total fema1esb ~o~u1ation c 1,974 2,184 2,438 3,016 4,958 5,548 4,422 4,624 4,624 4,420 1,610 2,512 2,961 3,276 3,657 4,524 7,437 8,322 6,633 6,936 6,936 6,630 2,415 3,768 Young in harvest (%) 50 42 42 46 53 58 49 47 51 57 57 54 Estimated fall ~o~u1ation d 5,922 5,648 6,305 8,378 15,823 19,814 13,006 13,087 14,155 15,419 5,616 8,191 aDerived by dividing ma1es/1ek by 0.6 (percent of males present at high count) and multiplying by number of leks (number counted plus 10%). bDerived by multiplying number of males by 2 as females comprise 68% of the adult and yearling population. CTota1 males plus total females. dDerived by dividing percent adults and yearlings in fall harvest into the spring population estimate. Pellet Transects All pellet transects were not counted nor cleared in spring or fall 1985 because of other assignments, early snowfall, and lack of apparent patterns (Braun 1984). It appears that random pellet transects (30) do not accurately assess sage grouse use or distribution. It is probable that larger samples are needed which are stratified by some feature of the habitat. Fertilization Three areas were selected for use in the fertilization experiment based on past sage grouse use (Beck 1975, Schoenberg 1982, Remington 1983). These areas were: A - classified as moderate use by sage grouse in Section 34, T10N, R79W and Sections 3 and 11 in T9N, R79W (Fig. 1); B - classified as heavy use by sage grouse in Sections 16, 17, 20, and 21 in T9N, R78W (Fig. 2); and C - classified as low use by sage grouse in Section 33, T9N, R78W and Sections 3, 4, and 9 in T8N, R78W (Fig. 3). All areas were on land administered by the BLM in regular grazing allotments. The domestic livestock grazing season varied from 25 May to 12 July depending upon allotment. Areas within the 3 selected areas were grided into 20-ha blocks and numbered. Eleven (plus alternates) 20-ha blocks in each area were randomly selected using a table of random numbers. Areas (10 ha) to be treated in each 20-ha 1 , t 4 , �137 block were randomly selected (even or odd) using a table of random numbers. Thus, sampling was without replacement for blocks and was with replacement to obtain which one-half of the block was to be treated. Blocks that were to be used in the experiment were either 600 x 3,588 ft (Areas A, C) or 1,200 x 1,794 ft (Area B) totaling 20 ha/block of which one-half was to be treated. Block size in Area B differed because the ruggedness of terrain prevented many areas from being used as the fertilizer applicators could not get their machinery over the desired areas. Starting points for plot marking were U.S.G.S. section corners. Some plots were offset to avoid fences. Plots were east-west (length) because prevailing westerly winds could affect fertilizer drift if the fertilizer was applied in northrsouth strips. All plots were marked with steel posts at the 4 corners and the 2 ends of the plot centers. Locations were ascertained using a transit and steel tape. r r r Two different applicators (Agrichem, Area C and part of Area B; Jirdon Agrichem, Area A and part of Area B) were used as the BLM paid $9,422.00 and the CDOW paid $13,959.00 of the total cost. Agrichem used a truck spreader that applied fertilizer in 60-ft swaths while Jirdon Agrichem used 2 tractor-drawn spreaders that applied fertilizer in 50-ft swaths. Area C and the south portion of Area B were treated on 5-7 November. Light snow «2 in. accumulation) fell at times during the application. Jirdon Agrichem started applying fertilizer in Area A on 7 November and continued on 8 November until noon when snow (3-4 in.) and low visibility caused operations to cease. Operations continued on 9 November (~6 in. snow) in Area A and part of Area B. Fertilization of plots in Area B resumed and was completed on 12 November despite heavy snow (~12 in.) and poor visibility because of fog. Fertilizer was applied at the rate of 100 lbs. of nitrogen per acre (271 lbs. of product per acre). The design was that 330 ha (11 10-ha plots in each of 3 areas) would be treated (fertilized). Several problems occurred that resulted in possible uneven application, mislocation of one plot, inability to physically treat small areas, and the apparent undertreatment of the last plot to be treated (49N). Fertilization was started on plot 16 of Area C. Plots 16 and 13 may have been underfertilized because the applicator made only 5 instead of the desired 6 strips. This was corrected on plot 12. Also, as the wind was from the south, there appeared to be fertilizer drift north on plots 16 and 13 resulting in the south edge of the control (north one half of plot 13) being affected by fertilizer. There also appeared to be a small missed area in fertilizer application near the center of the treated area (north one half) of plot 16. Plot 40 in Area B had a large irrigation ditch (abandoned) along the southeast corner that prevented fertilization of a small area (~159 x 138 ft). Plots 21 and 48 in Area B had small fenced areas that could not be fertilized. The largest problem occurred in plots 37 and 38 where the south one half of plot 38 and the north one half of plot 37 were to be treated. Because of heavy fog, plot 38 was treated at an angle from the southeast corner to the west center and the east center to the northwest corner. Plot 37 was omitted because the northwest corner could not be "located". (The west center was thought to be the southwest corner and the northwest corner was thought to be the west center). The reSUlting plot (38) that was treated was 10 ha. Because of the confusion and remaining fertilizer, an additional plot (49N) was added north of 49 (Area B). Thus, the north one half of plot 49 and �138 LOCAT\ON T A "3 c T {ott c 34\ T c fob 35 , ~. N. T C T C T 70 11 7J.. C 3 T .2.. 7~ 1 c C T T c T C C T 10 Fig. 1. Fertilized plots in Area A, North T = treated, C = control. 7g 7t:t ~o ~I 11 Park, Colorado. �139 LOCATiON T'lN R78W e r- ~'"' T T 49N T c c 33 l.f9 T r c 17 [to '-f~ r c c T 37 c T 3S T .5'l T Y6 C 53 2.1. 20 T c (" Fig. 2. c T T 40 c '18 Fertilized plots in Area B, North T = treated, C = control. Park, Colorado. �140 lOCATlOW C R1SW T c c /8 34 :>:> al T c ~a. T 7,9N. "i.3rL c 3 T c 3 "'T 5 T T 3d c T S r- C T 1/ c T c T Q , /~ a 10 T c Fig. 3. /6 Fertilized plots in Area C, North Park, Colorado. T = treated, C = control. , �141 most (possibly short 4 ha) of the south one half of plot 49N were treated. Despite these problems, the overall fertilizer application appeared to be fairly uniform. LITERATURE CITED Beck, T. D. I. 1975. Attributes of a wintering population of sage grouse, North Park, Colorado. M.S. Thesis, Colorado State Univ., Fort Collins. 49 pp. __________ , R. B. Gill, and C. E. Braun. 1975. Sex and age determination of sage grouse from wing characteristics. Colorado Div. Wildl., Game Inf. Leafl. 49 (Rev.). 4 pp. Braun, C. E. 1984. Responses of sage grouse to vegetation fertilization. Prog. Rep., Colorado Div. Wild1. Fed. Aid Proj. W-37-R-37. Pp. 5-30. r I Job ----....,.......-- , and T. D. I. Beck. 1976. Effects of sagebrush control on distribution and abundance of sage grouse. Final Rep., Colorado Div. Wildl. Fed. Aid Proj. W-37-R-29. Pp. 21-84. Giesen, K. M., T. J. Schoenberg, and C. E. Braun. 1982. Methods for trapping sage grouse in Colorado. Wi1d1. Soc. Bull. 10:224-231. Hoffman, R. W., and C. E. Braun. 1975. A volunteer wing collection station. Colorado Div. Wi1d1., Game Inf. Leaf1. 101. 3 pp. r Remington, T. E. 1983. Food selection, nutrition, and energy reserves of sage grouse during winter, North Park, Colorado. M.S. Thesis, Colorado State Univ., Fort Collins. 89 pp. Schoenberg, T. J. 1980. Potential impacts of strip mining on sage grouse movements and habitat use. Job Prog. Rep., Colorado Div. Wi1d1. Fed. Aid Proj. W-37-R-33. Pp. 245-264. Colorado. 1982. Sage grouse movements and habitat selection in North Park, M.S. Thesis, Colorado State Univ., Fort Collins. 86 pp. Swenson, J.E. 1985. Differential survival by sex in juvenile sage grouse and gray partridge. Ornis Scand. 17:14-17. Prepared by _ ___..,.tKc~:.=:..:.-___;;z._. -U~"-=.:.....;.....:.....;....----Clait E. Braun Wildlife Research Leader �143 Colorado Division of Wildlife Wildlife Research Report April 1986 JOB FINAL REPORT Colorado State 01-00-045 (W-37-R) Project Work Plan 3 Job Avian Research 13b Job Title: Responses of Sage Grouse to Vegetation Fertilization Period Covered: 01 September 1980 through 30 June 1983 Author: Thomas E. Remington Personnel: R. A. Ryder, Colorado State University; Kevin Berner, C1ait Braun, Len Carpenter, Kurt Hundgen, Steve Porter, Tom Remington, Joan Ritchie, Tom Schoenberg, Lyn Stevens, John Wagner, Carol Ann Wein1and, Colorado Division of Wildlife. ABSTRACT Sage grouse (Centrocercus urophasianus) food selection, diet quality, and energy reserves were studied in North Park, Colorado during January-April 1981-82. Radiotelemetry was used to locate flocks of feeding birds for observations and locations of feeding sites. Big sagebrush (Artemisia tridentata spp.) comprised 97% of the shrubs at feeding sites and 87% of the shrubs at random sites. Canopy cover ranged from 19% at random sites to 22 and 72% at feeding sites in 1981 and 1982, respectively. Sagebrush height averaged 24 cm at random sites and 17 and 30 em at feeding sites in 1981 and 1982, respectively. Variation in cover and height of sagebrush at feeding sites between years was related to snow depth. Wyoming big sagebrush (!. !. wyomingensis) comprised 86% of the sagebrush at feeding sites and 48% at random sites. Mountain big sagebrush (A. t. vaseyana) comprised 12 and 41% of the sagebrush at feeding and random sites, respectively. '. Ninety percent of the plants identified as fed-upon by sage grouse were Wyoming big sagebrush. Mountain big sagebrush (7%) and alkali sagebrush (!. longi1oba) (3%) were the only other plants fed-upon. Wyoming big sagebrush (ATW) contained more (p < 0.05) protein (14.1 VB. 10.8%) and a lower (p < 0.05) level of monoterpenes-(1.2 vs. 2.8%) than mountain big sagebrush (ATV). There was significant (p < 0.05) variation in protein content within subspecies, fed-upon plants -> non fed-upon plants > random plants. Monoterpene content did not vary (p > 0.05) between fed-upon, non fed-upon, and random plant samples, at least within ATW. Ether extract levels were not related to sage grouse food preferences. �144 Discriminant function analyses were used to identify variables discriminating between fed-upon, non fed-upon, and random plants within subspecies of big sagebrush, and to quantify the magnitude of group differences. Plant vigor and protein content distinguished between fed-upon, non fed-upon, and random ATWplants. Group distinctions were weak as only 31% of the variation in plant vigor and protein content was related to feeding status (fed-upon, non fed-upon, random). Within ATV an oxygenated monoterpene, protein, and plant vigor weakly (38% of the variation explained) distinguished between fed-upon, non fed-upon, and random plant samples. Good separation (81% of the variation explained) of fed-upon and non fed-upon plant samples was evident after random samples were eliminated. All (17 of 17) ATVcases were correctly assigned to fed-upon or non fed-upon categories based on their content of 3 oxygenated monoterpenes and protein. Crop and gizzard contents differed (p < 0.01) in chemical content. Gizzard contents contained less protein, cell contents, ash, and minerals, . and more ether extract, neutral and acid detergent fiber, and lignin than crop contents, apparently because of partial digestion of leaf material in the gizzard. Fat content varied among and within sex and age-classes of sage grouse. Adults had higher (P < 0.05) fat content than juveniles (4.67 vs. 2.87%), and birds collected in 1982 had more (P < 0.05) fat than birds collected in 1981 (4.03 vs. 3.43%). Fat content increased (~ = 0.06) from early to late winter. Estimated energy reserves of sage grouse fasting juveniles, 4-6 days for fasting fasting adult males. would last from 3 to 4.5 days for adult females, and 5-8 days for Petroleum ether extracted less (P < 0.001) fat than diethyl ether (3.60 vs. 4.04%). Diethyl ether extracts- were predictable (P < 0.001, r2 = 98.4%) from petroleum ether extracts. Nutritional quality of the winter diet, and by inference, requirements grouse were higher than those of other grouse. The lack of a grinding may explain how sage grouse can eat a diet of 100% sagebrush in winter they must. of sage gizzard and why �145 ACKNOWLEDGMENTS The financial support provided Division of Wildlife. Federal and the U.S. is gratefully r Department Laboratory Service. in Provo. graph of the Interior. on several I also thank sagebrush r measuring· confirmed vegetation under E. Remington I thank helped Dr. to the successme in the use discussions. of representative samples thankful in picking C. E. Braun of is gratefully acknowledged. grouse leaves. for initiating this and editorial assistance: interest The assistance and For this I E. T. Olson and sagebrush to him for his sincere L. H. Carpenter. sage K. Htm d gen , M. Oehlke r s , T. and T. J. Schoenberg. ual and as a professional. Drs. or collecting less than ideal conditions. for field supervision. especially bers. in capturing P. D. Curtis. E. A. Remington. funding. Forest gas chromato- was critical in many meaningful identifications people assisted K. Berner. Olson. of Agriculture. taxa. Several thank of the Shrub B. L. Welch instructed and took part W-37-R. of Land Management personnel This support ful completion of this project. E. D. McArthur Project for allowing me to use their occasions. of this equipment Bureau of the U. S. Department Utah. by the Colorado Aid to Wildlife Restoration acknowledged. Sciences this project study. for acquiring but I am in me as an individ- of my other committee mem- D. Heiri , C. F. Nockels, and R. A. Ryder Special thanks are extended to �146 Dr. L. H. Carpenter sions about Dr. for enduring sagebrush R. A. Ryder and animals that for his administrative this study, and countless I thank J. Ritchie, pleting eat sagebrush, support, is sincerely and discusand to continual interest in of literature. and C. A. Weinland for their The assistance under constant and friendship competent deadlines, assistance and for their of J. Wagner, in com- friendship. and J. Black S. Porter, appreciated. D. W. Hamar and M. Gerlach of the Department Colorado State a great donations questions Colorado Division of Wildlife employees L. Stevens, labwork Dr. innumerable University, deal of interest donated laboratory to this study. supplies For this, of Pathology, and space and and for accepting me as one of thei::- own, I am grateful. L't'harrk my wife, Liz, for her cheerful in every facet of this study, for the past 3 years. support, and for tolerating for her assistance a part-time husband 1 �147 TABLE OF CONTENTS INTRODUCTION 1 STUDY AREA •. 4 METHODS 10 RESULTS 18 Winter Use Areas • Vegetative Characteristics at Feeding and Random Sites • . . . .• Food Plant Selection ...• Reasons for Food Selection Univariate Analysis . . • Multivariate Analysis Chemical Characteristics of the Winter Diet Body Composi tion DISCUSSION r , 18 18 22 25 25 34 41 44 . • . . . Winter Use Areas . . . Vegetative Characteristics at Feeding Random Sites • . • . . • Food Plan t S el ec tion. • . . • . . . • Reasons for Food Selection • . • • • Chemical Characteristics of the Winter Body Composition • • • . . 52 . • • and 52 53 55 . • • • Diet 58 67 . ·0 • 69 MANAGEMENT IMPLICATIONS 75 LITERATURE 77 CITED �148 LIST OF TABLES Table 1 2 3 4 January-June precipitation and temperature, Walden, Colorado, 1981- 82 ••••••••• 9 November-March snowfall and snow depth, Walden, Colorado, 1978-79 to 1981-82 9 Gas chromatographic parameters used to separate and quantify monoterpenes • 14 Mean canopy cover, plant height, and plant vigor of sagebrush at feeding and random si tes, North Park, Colorado , Jan uary- April, 1981-82 . . 5 6 7 8 9 10 . . 20 Occurrence and cover (%) of shrubs at random and sage grouse feeding sites, North Park, Colorado, January-April, 1981-82 .••• 21 Occurrence and cover (%) of 3 sagebrush taxon at random and sage grouse feeding sites, North Park, Colorado, January-April, 1981-82 • 22 Frequency (%) of 3 sagebrush taxon as sage grouse food plants at all feeding sites and mixedspecies feeding sites, North Park, Colorado, January-April, 1981-82 •••••••• 24 Analysis of variance of crude protein content of Artemisia tridentata between subspecies and years and among fed-upon, non fed -upon, and random plant samples, North Park, Colorado, January-April, 1981-82 • • • • • • • • . ••• 28 .... Analysis of variance of monoterpene content of Artemisia tridentata leaf samples collected in North Park, Colorado, January-April, 1981-82. 31 Mean monoterpene levels (% DM) and univariate F tests between subspecies of Artemisia tridentata collected in North Park, Colorado, January-April, 1981-82 •••••••••••••••• 33 �149 LIST OF TABLES (Continued) Table 11 • •••• 35 Ether extract content (% DM) of fed-upon and non fed-upon samples of Artemisia tridentata, North Park, Colorado, January-April, 1981-82 •• 35 Discriminant function analyses of characteristics of fed-upon, non fed-upon, and random samples of Artemisia tridentata, North Park, Colorado, January-April, 1981-82 . . • . . • • . . • • • • • 36 Chemical analysis of sagebrush leaves from sage grouse crops (N 33) and gizzards (N 23), North Park, COiorado, January-April,-1981-82 •• 43 Average lengths and weights of selected sage grouse body components. . • .• .•.. 45 Relative lengths (mm/ gO. 698 body wt) of the caeca (combined) and small and large intestines of sage grouse collected in North Park, Colorado, JanuaryApril, 1981-82. • • • • . • . . • •.• 46 Total body fat content (diethyl ether extract) of sage grouse collected in North Park, Colorado, January-April, 1981-82 .••••.••••• 47 Two-way analysis of variance showing age and year differences in sage grouse body fat with date of collection as an explanatory covariate, JanuaryApril, 1981-82, North Park, Colorado ••••••••• 48 Petroleum and diethyl ether extracts of paired sage grouse carcass samples. . . . • . • • . • • 51 Winter protein levels (% OM) of leaves and leaves and stems of Artemisia tridentata compiled from the literature . • • • • • • • • • • • • • • 61 21 Chemical 70 22 Estimated energy reserves (Kcal) of sage grouse, January-April, North Park, Colorado, 1981-82. • 12 13 r Ether extract content (% DM) of 3 sagebrush taxon collected at random sites, North Park, Colorado, January-April, 1981-82 • • •• 14 15 16 17 18 19 20 = analyses = of the winter diets of grouse 72 �150 LIST OF FIG URES Figur~ 1 North 2 Winter use areas of radio-marked in North Park. Jackson County. 3 4 5 6 7 8 Park. Jackson County. Colorado 5 sage grouse Colorado. 1981- 82 . . . . . . . . . . . . . . . . . 19 Range. mean. and 95% confidence interval of crude protein content of Artemisia tridentata wyomingensis and A. !_. vaseyana leaf samples collected at random sites in North Park. Colorado. January-April. 1981-82 • • • • •• •• • • • •• 27 Crude protein content of fed-upon. non fed-upon. and random samples of Artemisia tridentata wyomingensis and A. !: vaseyana collected in North Park. Colorado. January-April. 1981-82. 29 Range. mean. and 95% confidence monoterpene content of Artemisia wyomingensis and A. !_. vaseyana collected in North Park. Colorado. April. 1981-82. • • • • • • • • • 32 interval of tridentata leaf samples January• • • • Range. mean. and 95% confidence interval of discriminant function scores separating fed-upon, non fed-upon. and random samples of Artemisia tridentata wyomingensis • • • • 38 Range. mean. and 95% confidence interval of discriminant function scores separating fed-upon, non fed-upon. and random Artemisia tridentata vaseyana samples • • • • • • • • • • • • 40 Range. mean. and 95% confidence interval of discriminant function scores separating fed-upon and non fed-upon samples of Artemisia triden ta ta vaseyana • • • • . • • • • • • • • • • 42 �151 INTRODUCTION Sage grouse of western North America. 2 Canadian provinces. the distribution r are widely distributed 1952, Aldrich brush also provides alteration province have been extirpated (British Sagebrush since herbicides, 1979). reduced 1975). cover (Patterson Columbia) began of the West. In Colorado, between 1900 and directed against (Patterson on sage- 1937, Griner (Rasmussen result 1952, Aldrich 1954, and 1969, Eng 1974, Beck 1977, and distribution of elimination 1963). 1974 (Braun big sagebrush. and and 1 1963). plowed, disked, Extensive chained, and effective and application cut, 30%of all sagebrush et al. 1976). lands Nearly and "con troll! of chemical 2,4- D, followin g World War II (Evans at least of Sage grouse (New Mexico and Nebraska) with the development particularly to Big sagebrush Both numbers (Aldrich has been burned, linked 1952, Klebenow as a direct from 2 states settlement of sagebrush (Girard and 1952, Leach and Hensley and brood 1982). closely dependent 1972, Wallestad and Schladweiler of sagebrush have been are completely 1941, Patterson 1980, Schoenberg sage grouse in 11 states big sagebrush 1958, Wallestad et al. nesting, the rangelands has been much of the year 1938, Keller et al. Petersen beat Sage grouse loafing, and Schladweiler occur particularly et al , 1942, Patterson Leach and Browning Griner distribution of sagebrush, for food throughout 1939, Dargan r They presently Their 1963). throughout et al , were treated all control has been �152 Concern about impacts tion has been frequently 1975, Braun practices et al. 1970, Martin Surface habitat expressed 1976). have been or disturbance on wildlife of sagebrush Detrimental demonstrated of seasonal 1970, Pyrah that States source of sage grouse coal accounted in 1978 compared wildlife managers on less area in the future. This creates when management populations habitats 1969, Klebenow for 20% to less than 1980). It is apparent critical control due to elimination 1967, Higby surface-mined of all coal mined in the United sage grouse (Carr 1968, Vale 1972, Wallestad 1975). Western 2%in 1968 (Slatick of sagebrush for sage grouse habitats altera- 1960, Kufeld effects mining of coal is an increasing disturbance. managers (Anderson habitat will be managing a serious dilemma for goals call for maintaining (Colo. Div. Wildl. 1983). and enhancement of remaining sage grouse or increasing Identification habitats of will be required q I to achieve these and structural Between goals. characteristics and within Food habits Previous season October describe food habits (use in relation period. A few reports, of species Monoterpenes, oil of sagebrush, deter of sage grouse. et al. 1982). that monoterpenes described. in the May April period. and subspecies components 1980~, Farentinos by sage grouse studied of the "volatfle" (Connolly et al , 1981, Radwan et al. or nutrients is unknown, Selection of sagebrush feedin g by herbivores et al , 1980, Schwartz on food selection the physical mostly observational, in the November through to availability) The influence habitats have been extensively has not been investigated. or "essential" of seasonal has described movements have been adequately of sage grouse through research in sagebrush yet this knowledge have is �153 essential if nutritional enhancement Baseline levels of nutrients Objectives winter (Jan-Apr) of this study brush plants fed-upon plants at randomly-located sites, and acid detergen t fiber, lignin, carcass grouse feed selectively brush; (2) sage grouse species of sagebrush protein, (3) sage grouse or alternatively, lower than average terpene contain feed disproportionately levels. protein species levels, and of neutral and crude and (5) measure were: (1) sage and subspecies or alternatively, feed disproportionately greater than average monoterpene on individual or alternatively, of sage- on the species/sub- the most protein plants than average tested feed disproportionately where sagebrush grouse mineral, con tents, Hypotheses food con tent of sage- cell contents, ash, crop and gizzard containing winter by sage grouse, (4) measure from available sage grouse sage grouse and monoterpene and not fed-upon fat levels. are unknown. (1) identify (2) identify (3) measure protein least monoterpenes; r were to: food habits, fat levels of sage grouse is to be attempted. in diets of sage grouse preferences, sage grouse of sagebrush protein levels; plants the at sites levels, and (4) sage containing less than average greater mono- �154 STUDY AREA This study was conducted in North of intensive investigations Colorado. Areas movements of radio-marked marked within the northeast quent winter study area was bounded the south movements quarter North (Fig. of North this In depen dence northwest principal 2,758 m. study on the south Mountain Floodplains are separated Vegetation sagebrush Platte encircled of North ridges River, or showed on the on the west by and by is drained to the area is drained Topog raphy at an elevation by of the of 2,420 to and tributaries and and benches. sites, and by grasses collected These rivers. Park is dominated ranges Park the major drainages hay meadows bordering Park. 1 125, on bounded Ears Range, The study was flat to rolling border ranges, North River. irrigated North The extensive by mountains; by the Rabbit and Illinois by numerous on upland and all subse- Lake and Eagle Hill to the north on the north. Michigan, area and radio- 79W). by the North the Canadian, on on the east by the Canadian Cowdrey Park is completely Range, Park, quarter. County, 1) depended were trapped east by the Medicine Bow and Neversummer the Park Jackson on the west by Colorado Highway by the Michigan River, ION, R78 and Birds were within and by a line connecting. (T9 and birds. Park, by 1 or more species and sedges major drainages. of 6 taxa of sagebrush were Artemisia tridentata on native Beetle (1960) extending vaseyana, into A. ~, of and , �155 N o 5 10 KILOMETERS r COALMONT. DENVER. COLORADO Fig. 1. North Park, area (stipled) Schoenberg Jackson was between 1982). County, Colorado. the Canadian The intensive and Michigan rivers study (after �156 A. ~~. A. ~ Smith (1966) sagebrush vfscidula , A. lon giloba , and A. argillosa. reported type. that A. !.: !: be amended together occurs North to read occupy in areas McArthur in areas !: A. vaseyana about '90% of the with deep, with dry. et al. predominates shallow. 1979. Winward in draw bottoms accumulates. A.!_. ing ridgetops and benches greasewood he collected 1980}. et al. slopes areas occurs 1965. where snow sites includ- in association I observed A. longiloba northwest Smith with black the northeast found deep on alkaline in areas can a and A. cana viscidula occurred drained, draw bottoms. known only from the Coalmont area quadrat. in limited of North the Park. A. longi- with clay pan soils is not possible. and well-drained soil as well as at obvious seasonally-flooded throughout root penetration sites of Coalmont" (1966) did not find which were spread within 1966) where gravelly 1966. A. !_. vaseyana on drier County, A. nova. sagebrush (Robertson wi th coarse, (Smith Park, predominates "Jackson A. ~ loba is a low-growing vaseyana and Young In North should wyomingensis wyomingensis {Beetle or in alkaline statement A.!_. and on east-facing at any of his 40 plots I did not encounter A.!_. area vermiculatus}. (1960: 30) listed as a site where type. loamy soils soils wyomingensis {Sarcobatus Beetle coarse !: and A. well-drained and Young in the study (1966:57) sagebrush 90% of the by Beetle both Smith's et ale 1979, Winward 1980). McArthur A. ~ Thus, about of a 3rd subspecies described was abundant Park. that occupied unaware wyomingensis. wyomingensis and throughout vaseyana Smith was apparently of A. tr-iderrtata , A. (1965). !: A. ridge clay pan sites. areas, A. argillosa Park usually tops A. cana poorly- was previously (Beetle 1960), but I �157 found it in limited areas similar to within the study area those where A. cana occurred. Beetle the morphological characteristics A. cana viscidula and A. lon giloba , although species sites the study area. with A. cana viscidula. common to find plants A. cana viscidula At several sites, A. argillosa In areas that be more accurate described species leaves plant associated occurred, leaves typical it was typical of of A. argillosa. forms predominated. A. argillosa between mixed at the same was frequently both linear 3-lobed mixed-leaf to consider (1960) he did not find the 3 where both contained and deeply these site conditions of A. argi110sa as intermediate I foun d the 3 species at the same site. within under It may as a form of A. cana viscidula. Other shrubs area included antelope with limited distribution black bitterbrush (Symphoricarpos (Purshia rabbitbrush tridentata), vaccinioides), and willows (Salix consisted primarily forbs. greasewood, Schoenberg (1982) whortle-leaf 1982). spp.), Herbaceous montigenum), vegetation and low-growing the plants the study snowberry curran t (Ribes bunchgrasses listed within (Chrysothamnus gooseberry spp . ) (Schoenberg of perennial occurring he identified perennial within the same area as in this study. At least between 1958 and ous because sagebrush 9% of the study 1964 {Schoenberg of the dead sagebrush 1982}. to remove sagebrush These areas plan ts which remain, are conspicualthough some has reinvaded. The climate of North average area was treated frost-free January-June period precipitation Park is cold. dry, of only 46 days and windy, {U.S. with an Dep . Commerce 1979}. was above the long-term average during �158 both years of the study 1981, 1982). (Table Precipitation (1982) of the long-term and June accounted temperatures average in 1982 (Table from November low (Table 150 of the 151 days during this snow depth snow depths 1981-82. period 1). to April was 78 (1981) and than during precipitation There was little tempera- maximum snow depths 1980-81. on Snow depths than in 1980-81, but below 1978-79 or 1979-80. on 48 of the May totals. (~10 cm) snow cover to March. 102% in 1981 and about Low snowfall and above-average in 1981-82 were higher levels in either rainfall average 1980 to March 1981 kept 2). Commerce 1979, 1980, were above average from November occurred Dep. above average for the greater to June extremely from January average; January tures 1) (U.S. 151 days Significant from November (~10 cm) to March �.....". Table 1. January-June precipitation Precipitation and temperature, Walden, Colorado, 1981-82. (em) Mean temperature (C) Year Jan Feb Mar Apr May Jun Totals Jan Feb Mar Apr May Jun 1981 0.17 1. 07 2.18 0.86 5.59 4.34 14.21 -4.8 -4.7 -1.8 4.8 6.3 13.6 1982 a Avg 2.13 0.41 2.01 1. 07 5.54 1. 52 12.68 -7.1 -6.9 -1.8 0.4 5.7 10.5 1. 30 1. 07 1. 27 1. 83 2.59 2.82 10.88 -9.2 -7.7 -4.6 1.8 7.1 11.6 depth, Walden, - aThirty-year Table 2. average, 1941-70. November-March snowfall Total Year Nov 1978-79 9.9 1979-80 Dec snowfall Jan Feb 56.1 30.7 9.9 18.8 11. 4 75.7 1980-81 3.0 5.3 1981-82 20.3 31.0 and snow 1978-79 to 1981-82. Maximum snowdepth days with snoweover (em) Mar' Colorado, Totals Nov 32.0 138.6 5.1/30 14.7 22.9 143.5 3.8 15.5 28.4 29.2 8.6 21. 8 (em) / !>10 em Dec Jan Feb Mar 30.5/1 38.1/0 38.1110 7.6/31 12.7/29 10.2/31 45.7/20 38.1/0 56.0 2.5/30 2.5/30 Tr/31 7.6/28 5.1/31 110.9 7.6/30 20.3/27 30.5/0 5.1/15 7.6/31 20.3/21 t-' VI \0 �160 METHODS Radiotelemetry was used could consistently tions. be located Sage grouse at night using numbered, size plastic captured 1975), and stage mounted on a poncho winter male was fitted sage Radio-marked flocks rather than of individuals fied by visual 7.5-minute observation U.S. Vegetation loca ted sites color-coded as to sex, rectrices to year (Pyrah with serially- grouse Inc.) to the nearest using Carbondale, a modified bolt and tagging and hand-held fabric using individual or occasionally a portable used All locations topographic to locate thus, daily were veri- and plotted on maps. were made at winter-feeding usin g a modification yagi movements, by flushing, radio (Amstrup 3-element were primarily was not stressed. 20 grams. by Bray and Corner were relocated to measure measurements Inc.; with a 17-g solar-powered sage grouse Geological Survey of capture. age (Eng 1955, Beck et al , described made of vinyl-coated (Wildlife Materials, relocation nets were marked (20-22 g; Wildlife Materials, Radio-marked antenna. grouse similar to the tail-clip One adult receiver and for collec- and on leks were captured molt and weighed to the central (1972). 1980). bandettes of primary were attached clamp device of feeding and long-handled Captured was classified Radio transmitters Ill.) flocks of sage grouse 14 {females} or size 16 {males} aluminum leg bands and unnumbered Each bird along roads spotlights 1959, Giesen et ale 1982). that for observations roosting hand-held to ensure and randomly- of Can field IS (1941) line-in tercept �161 method. Three or four randomly measured at each site. intercept distance. that intersected assessment Crown length and plant the line. 5-unit increments Plants which intercepted leaves is light the leaf. of the plant. Thus. from 5 (nearly Sage grouse The exposed and contrasts fed-upon maximum height. for each plant dead) sites cut rather interior sharply plants were visual Values were recorded the line at feeding plants. green and width. vigor were recorded and ranged of feeding. from sagebrush 10-m transects Plant vigor was a subjective of the condition for evidence oriented to 100 (vigorous). were examined than pick leaves of freshly eaten with the dark can be quickly in surface and reliably of identi- fied. Sagebrush intersected leaf samples were collected the line transects samples were individually at feeding tagged. and frozen until used Sagebrush samples were identified teristics (Beetle A. tridentata extract to subspecies and McArthur (Young 1974). in fluorescent Representative pattern USDA Forest and storage Research Service. ethanol Geneticist of reference pattern charac- of a water light (Stevens or methanol leaf extracts 1969) gave similar results. between Provo, bags. by morphological ultra-violet subspecies samples of each taxa observed McAr thur , Principal cations under A method using plastic 1979, Winward 1980), and by the fluorescent leaves observed Leaf or chemical analysis. to species 1965. Winward and Tisdale differences tory, sealed in air-tight et al. that and random sites. for plant identification 1960. McArthur of crushed from each plant Utah. specimens. were not as great. were sent at the Shrub but to E. D. Sciences for confirmation Labora- of identifi- All identifications were �162 confirmed. Reference Wildlife herbarium Transects intense feeding selecting sites were centered activity. Random sites resulting The distance point were prepared weight) of leaves of leaves nitrogen (to retain pestle and feeding monoterpenes) powder by micro-Kjeldahl was estimated by multiplying were extracted with 50 ml of diethyl extract standard transects status was reduced of carvone in a mortar analysis adding ether. Monoterpenes 2 ug of the extract or non pooled samples group. (wet Most groups and ground 1981). (Horwitz by 6.25. with a Nitrogen 1980) and protein (1981). into a Hewlett-Packard levels Monoterpenes apparatus The volume of pressure and an internal The extract flask and the volume brought were identified in liquid Monoterpene as solvent. (2. 5 ug Zml) was added. to a 50-ml volumetric were pooled by from 5 -g samples in a Soxhlet ether of of the equal amounts below 25 ml with reduced ferred corner Samples were immersed nitrogen anhydrous numbers (fed-upon Twenty-gram following Welch and McArthur for 6 hours 2 3-digit a determined. (Welch and McArthur was measured over (azimuth) in a particular of 10-20 plants. to a uniform were determined only). by hand-picking from all plants consisted and direction road were then transects for analysis transposed selecting from vegetation subspecies, feeding-site by randomly and west of the northeast (in paces) leaf samples species, fed-upon, south were located randomly from the nearest Sagebrush transect, and then to meters of on the area of most to 1-km2 grids corresponding area Division Colorado. at feeding map of the study the grid. were filed in the Colorado in Kremmling, a number corresponding specimens and quantified was transto 50 ml by by injecting 5830A Flame Ionization, �163 Reporting Gas Chromatograph. stainless A 1. 22 m by 3.2 ern (4 it x 118 in) steel column packed chromosorb WHP was used programming with 10%carbowax to separate specifications ether caliber rifle at, feeding periods were collected or just enough leaf samples. contents (CC), 1967). was partitioned lignin, and ether concentrations matters extract were determined were obtained by ashing then solubilizing fiber ether on a dry and (NDF), cell as solvent of protein, Lignin Crude fat in a Soxhlet CC, NDF, ADF, in duplicate. matter as sug- (1982). (1963!:!,,\2; 1967). were performed a 0.5-g for hand- (ADF) following Van Soest All analyses by drying and pestle. from crops (1981) and Van Soest anhydrous are expressed crops were freeze- from the NDF residue from ADF (Van Soest 1980). because as described detergent fiber ADF was extracted (Horwitz or evening and body with a mortar leaf samples into neutral with diethyl con tents and gizzards consistency and acid detergent or .22 for analyses. were determined by Mould and Robbins apparatus so of morning were unsatisfactory The sagebrush were partitioned was extracted the conclusion food material protein gizzards gested ether with a shotgun of crop and gizzard to a uniform and crude (l963~,Q; to, leaf samples from crops and ground picked Petroleum monoterpenes, by shooting Morning collections Sagebrush Nitrogen prior for analysis seldom contained dried for extracting Temperature was used. Sage grouse composition. monoterpenes. used are in Table 3. was found to be a poor solvent diethyl 20 ml on 80/100 Nutrient (DM) basis. at 75 C for 24 hours. Dry Minerals sample at 550 C for 12 hours the ash with hydrochloric (HCI) or nitric acid. and The �164 Table 3. Gas chromatographic parameters used to separate and quanti-· fy monoterpenes. 70 C Ini tial oven temperature Ini tial time 1.0 min Ini tial temperature Programmed rate 5 C/min temperature rate changes: At 2.5 min 1.0 C/min At 5.5 min 10.0 C /min At 15.0 min 25.0 C/min Final oven temperature Hold at final oven 200 C temperature Flow rate 30 ml/min of N2 Attenuation 8 Slope sensi tivity Injection 5 min port Flame ionization 20.0 temperature detector 250 C temperature 250 C �165 solution was diluted an d water diluted 1: 50 with a 5% solution (1 g: 5 ml: 15 ml, respec ti vely) • 1: 10 with de-mineralized Model 303 Atomic Absorption calcium, verted magnesium, to parts concentration. curve. (distal the intestinal grinder. of the dried and crushed altered for birds and increase collected precision gizzard, I-X, weight, caeca, oviduct, small and largest outer portions with a meat in 1981 were passed at 80 C for 24 hours and pestle. (6 hours) Kerr et al. of and ground (1982) found up to 120 C did not cause from duplicate Sample preparation in 1982 to reduce of the technique. through Two 100-g subsamples material in a Soxhlet as solvent. liver, to homogenizing samples at temperatures Fat was extracted ether were converted the feet, 6-8 times. were dried loss of fat. anhydrous 20 and the head were removed and collected meat grinder carcass of known Spectronic and testes primaries were plucked, from birds to a fine powder in a mortar that drying of to ppm by compari- heart, of the ovary, emptied prior carcass weight, wing, to radiale/ulnare), contents each homogenized of bill, Carcasses a small hand-driven on a Beckman M. supracoracoideus, and diameter Carcasses to standards were converted on total body and lengths follicle of females. levels for each sample and con- All mineral concentrations Musculus pectoralis, intestine, units Elmer DM. Data were collected of wings was measured absorption HCI, into a Perkin Absorption and sodium were charted from ppm to percent and large and injected Spectrophotometer. Phosphorous son to a standard total length, water oxide, This sample was further per million (ppm) by comparison Spectrophotometer; crop, of lanthanum apparatus 10-g samples with diethyl techniques were sample preparation Carcasses were ground time while �166 still partially entire frozen homogenate with a large was then at 75 C for 24 hours in a Wiley Mill until extracted spread they passed replicates overis ,' through both to determine needed to accurately metal trays a 2-mm screen. The and dried Fat was and diethyl Ten duplicate precision 7 times. Dried samples were ground petroleum were used as solvent. were extracted meat grinder on foil-lined in convection as in 1981, although (separately) electric ether samples from 1 bird of this method and number estimate individual fat content of (Brisbin 1968). Programs Package used in statistical for the Social Sciences characteristics pared analysis using (cover, student's taxa at feeding analysis. height) at feeding and random sites was tested respectively. Because of number of comparisons Probabilities accepted as significant. between fed-upon, levels Reasons and multivariate (ANOVA) was used content lower than to test years performed, the probability were compared or growth and multiple a single forms ~ tests, test of obtaining from tables for food selection techniques. for differences as of signi- X signi- in Rohlf and Sokal in protein of variance and monoterpene of big sagebrush subspecies were were investigated Analysis and random sagebrush between a chi-square 5 times out of 100 comparisons and subspecies non fed-upon, were com- level of a test increases in N random comparisons (1969). Vegetative of sagebrush using for taxon analysis the significance ficance was made by determining with univariate of occurrence was compared by chi-square 1975). and random sites (use v s , availability) of sagebrush fican t ~ values (Nie et al. ~ test •. Frequency Selection a function (SPSS) were from the Statistical and among samples. of big sagebrush Ether extract and years �167 by !. test. group Stepwise (fed-upon, ences in protein, (considered discriminant non fed-upon, ether extract, simultaneously) function within subspecies tested multiple!. values test for differences year of collection. from bird sidered at P < 0.05 unless ANOVA was used was used otherwise were the probability due to grouse measurements. Differ- and gizzards and determining fat content Multiple regression weight and structural significant tests of crops differ- monoterpenes of big sagebrush. in N random comparisons. in carcass samples) and total. and IS individual of chemical analyses X significant!. were used to examine and random sagebrush ences in mean values by performing analyses sex, to predict All tests noted. age, carcass were con- of to and fat �168 RESULTS Winter Use Areas Relocations were used were of radio-marked to delineate radiomarked both winters. 1982 largely captured winter north (Fig. quadrat in locations where and observations use areas in the northeast Differences reflect birds radio transmitters indicated that both areas All sage and remained of use areas and east of Walden remained Field observations 2). of other grouse there between during 1981 and were attached. within these flocks Birds 2 areas. were used during both winters. Vegetation Structural Characteristics characteristics feeding and random sites habitat selection. at Feeding and sagebrush were measured Canopy cover sites canopy cover, than values Shrubs sagebrush species and sagebrush and vigor at feeding at random or feeding at feeding to subspecies. ing and random sites sites in percent at feeding (Table 4). of sites height at Sagebrush sites in 1982 were higher in 1981. and random sites Species vigor at as indicators while sagebrush was lower than at random sites height, composition and compared in 1981 were similar to random site values, feeding and Random Sites were identified composition occurrence varied or percent to species little between cover (Table and feed5). �169 N REGULAR USE 1981 REGULAR USE 1982 INTENSIVE USE 1981, o I 1 2 3 I 4 I I 5 I I KILOMETERS 8« a: o ...J o o J.e. Fig. 2. County, Winter use areas Colorado. 12 of sage 1981-82. grouse 1982 in North Park, Jackson �170 Mean canopy Table 4. brush at feeding April, 1981-82. Site Feeding, and random N 1981 - SE Range 1982 SE Range x SE Range sites, Canopy cover (%) height, North and plant vigor Colorado, Park, Height (cm) of sageJanuary- Plant vigor 21.8 17.4 75.2 1.6 1.5 2.0 10-36 10-33 59-94 26.7 30.4 78.1 2.2 1.7 1.5 14-51 20-46 57-87 19.5 24.1 73.1 1.3 1.5 1.7 8-42 8-42 45-95 20 x Random plant 20 x Feeding, cover, 36 �Table 5. Park, Occurrence Colorado, AT AL AA AC 2.1 Random 86.8 11.3 = feeding sites, North Percen t cover C SV ~ plants encountered 0 0 0.2 0.6 97.2 1.3 0 0 0.3 1.2 1,547 0.5 0.2 1.2 0 91.7 6.1 0.3 0.4 1.5 0 1,062 Artemisia tridentata, thamnus spp , , SV at random and sage grouse SVa 97.1 = of shrubs Percent occurrence ALa AAa ACa Ca Feeding aAT (%) 1981-82. January-April, ATa Site and cover Sarcobatus AL = A. longiloba, AA = A. argi11osa, AC = A. cana, C = Chr-y so- vermiculatus. ~ ......, ~ �172 Feeding sites slightly less A. longiloba contained little Sarcobatus species contained !.: sites and AL than area. random Both types feeding and cover and random sites level, by subspecies, (ATV) (Table more (~< 0.05) proportionally of these at the species vaseyana of sites spp., or was segregated !.: (ATW) and A. contained sites. (AT) and the limited distribution Although composition tridentata Chrysothamnus when A. tridentata wyomingensis Feeding reflecting the study they were distinct random A. argillosa, vermiculatus, within more Artemisia (AL) than A. ~, were similar in shrub A. slightly 6). ATW and less ATV sites. Table 6. Occurrence and sage grouse feeding Percent occurrence and cover sites, (%) of 3 sagebrush North Park, taxon at random Colorado, January-April, 1981-82. ATWc Site coverb ATVc a c AL ATW ATV AL N plants encountered Percent Feeding 86 12 2 87 12 1 1,508 Random 48 41 11 48 46 6 1,042 a 2 AL ~ = b 2 X = 433, 2 df, ~ <0.005. . 10,319, 2 df, ~< 0.001. cATW = = Artemisia tridentata wyomingensis, ATV = A. t. vaseyana, A. Ion giloba . Food Plant Selection Sagebrush subspecies nated grouse plants at feeding and examined as fed-upon if they and non fed-upon sites for evidence were identified of feeding. showed signs to species Plants of 2 or more bites if they had 0 or 1 bite. and were desigby sage In 1982, plants were �173 designated as non fed-upon the snow indicated the plant. that only if they sage grouse This was not possible had 0 or 1 bite and tracks had been close enough for the most part in to feed on in 1981 because of a lack of snow. ATW, ATV, and AL were the only sagebrush fed-upon. They were also the only sagebrush ing sites. A definite was observed (Table plan ts identified 7 and sites 7). at feeding containing was selected lor, 12%of the plants AL was not strongly sagebrush random at a feeding taxa to sage sites. 48, 41, and an even stronger Pooling data Twenty-eight feeding tained sites at feeding against, it was eaten indication sites, comprising plants. readily when of of these taxa sites at were ATW, ATV, and AL comprised 90, 7, respectively. indicates This comparison for ATW and against (5%) contained ATW of availability may be the percentages from all transects There feeding of ATW, ATV, and AL at random plants. both ATW and ATV. for feeding but only 7% of the fed-upon A better grouse selection Even within for or against; of 39 (72%) feeding transects must choose. sites 11%, respectively. and 3% of the fed-upon Selection ATV was selected site. Proportions plants. of ATW was due to the preference plants. selected at feed- ATV and AL comprised 86% of the sagebrush at feeding found as and ten of 676 (90%) fed-upon of the fed-upon comprising species identified for ATW as a food plant were ATW• as a food plant. 90% of the fed-upon encountered sites a high proportion subspecies 0.05) (~< Six hundred 3%, respectively, for this but preference species obscured transects some relationships. contained only ATV. It was at these was weak (P = 0.17) ATV and AL. only ATW. Two The remainder (23%) con- sites grouse selection that sage for ATW and against �174 Table 7. Frequency plants at all feeding Park, Colorado, (%) of 3 sagebrush sites and mixed-species feeding grouse food sites, a North 1981-82. January-April, Percent Plant taxa as sage occurrence N sagebrush plants encoun tered status C All feeding sites Food plants 90 7 3 676 All plants 86 12 2 1,508 Food plants 70 30 164 All plants 62 38 336 sitesd Mixed-species aMixed-species feeding sites were those that contained both ATW and ATV. b ATW = Artemisia triden ta ta wyomin gen sis, AL = A. Ion giloba . c 2 X = 39.32,2 dX 2 = df, P<O.05. 1. 94, 1 df, P = 0.17. ATV = A. t , vaseyana, �175 ATV at mixed-species plants identified at these sites Possible basis (Table 7). The proportion as ATW was 69.5%, whereas selection for individual attributes of fed-upon 62.5% of the sagebrush plants within was investigated plants. (P < 0.05) in 9 of the 43 comparisons (21%). can t differences would be expected out of 100 (Rohlf and Sokal 1969). 1981. were significant Larger differences plants between 2 of 7 height were selected fed-upon comparisons selection was for larger vigorous plants all cases, (~<0.05). alone less than Most of these 1 time com- (5) occurred in 4 of the 7 significant plants. in height Within ATV, (~< 0.05). Sage grouse plants of signifi- Within ATW, 7 of 36 height were significant for taller height This number by chance to feed upon within both selection on the Means were different and non fed-upon plan t s , subspecies by comparing and non fed-upon parisons of fed-upon was ATW. of physical values sites In both appeared cases to select subspecies. more In most, but not was also selection for more vigorous plants. Reasons Univariate for Food Selection Analysis Selection as a possible for protein explanation or against for sage grouse taxa and growth forms of sagebrush. was used for differences between upon, protein to test subspecies non fed-upon, was tested monoterpenes Analysis of variance years, samples. (1) between for certain (ANOVA) and total monoterpenes between and random plant at 3 levels: food preferences in protein of A. tridentata. was hypothesized and among fed- Thus, selection big sagebrush for subspecies, �176 (2) between plants feeding within sites a subspecies (P < O.05) in protein fed-upon, There and feeding content non fed-upon, status (fed, apparent by year (Fig. 4). protein significant locate difference differences content among fed-upon, and non fed-upon samples. In 1982 samples, protein than random from non fed-upon fed-upon samples, samples. t test, between P < 0.05) . plants but neither Pairing A was used fed-upon status All 3 Fed-upon samples; both more protein than of these random more (~<0.05) differed (P > 0.05) and non fed-upon indicated and non fed-upon to non fed-upon, contained mean deviations fed-upon sites. separately. non fed-upon All above snow (1981 samples). samples con tained from each site and comparing tein differences than in 1982. of the year-feeding from each other fed-upon This differ- procedure means from 1981 and 1982 were tested more protein status and non fed- only plants interaction, samples contained and feeding snow depth whereas Because feeding becomes slightly. sagebrush (P <0.05) between of fed-upon and random means differed samples. 8). Main effects were sampled at feeding content (Table from 1981 to 1982 for both multiple comparison in protein samples of species by increased grouse) 3) and among The interaction markedly were sampled at random sites, to sage (Fig. years. site samples increased be explained were differences 2-way interaction and year. The crude (3) among individual There subspecies the main effects while random (and available site. 0.29) between samples declined ence can probably plants = random) graphing upon sagebrush subspecies (P by a significant non-fed, upon between and and random sagebrush was no difference were complicated least and random sites, there samples samples were pro(paired �177 20 19 18 17 - ?f!. z iii I- 0 16 N=27 15 14 a: a.. 13 w 12 ::> 11 N=20 0 a: 0 10 ATV ATW Fig. 3. content Range, mean, of Artemisia (ATV) leaf samples January-April, and 95% confidence tridentata collected 1981-82. interval wyomingensis at random sites of crude (ATW) and A. in North Park, protein !_. vaseyana Colorado, �178 Table 8. between Analysis of variance subspecies and years random plant samples. Source of variation North a Subspecies statusb Year Two-way interactions Subspecies status by feeding Species by year Feeding status Three-way by year interaction Residual Park. Colorado. df aSubspecies bFeeding = status of Artemisia tridentata non fed-upon. January-April, and 1981-82. MS F P 655.62 4 163.90 39.55 0.001 414.51 1 414.51 100.02 0.001 100.16 2 50.08 12.08 0.001 4.68 1 4.68 1.13 0.29 71.69 5 14.34 3.46 0.006 15.54 2 7.77 1.87 0.16 5.21 1 5.21 1. 26 0.26 58.12 2 29.06 7.01 0.001 1. 41 2 0.71 0.17 0.84 513.87· 124 4.14 135 9.20 1,242.60 Total protein and among fed-upon. SS Main effects Feeding of crude A. t. wyomingensis = fed-upon, and A. t. non fed-upon, vaseyana. and random. �179 18 1981 1982 ATW 17 ATV ~ 16 'if!. 15 z w t- O 14 a: a.. w Cl ::J 13 a: o 12 11 10 FEDUPON Fig. 4. random A. !.:_. April. Crude RANDOM NON FED-UPON protein leaf samples vaseyana 1981-82. content of fed-upon. of Artemisia tridentata (ATV) collected in North FEDUPON NON RANDOM FED-UPON non fed-upon. wyomingensis Park. Colorado. and (ATW) and January- �180 Lack of a significant and subspecies fed-ueon. indicated and random that by examining and multiple comparison plant samples non fed-upon of 7 within 0.16) ATV. of the deviation fed-upon between fed-upon samples This ANOVAs significant tial for selection) protein content plant and non fcd-upon were higher within than ATW and accounted and non fed-upon that samples variation for 79% in 1981 (and may only occur 7 poten- at a limited of sites. Analysis samples of variance indicated no differences random times that of total monoterpenoid differences between samples si gnifican t (P years (P> 0.05) (!: > O. 05) . species (!: < O. 05) 10). (Table contained unknown Neither monoterpenes of individual 15 comparisons differences alone less 9). ATW and ATV, but non fed-upon. and of the interactions was 5). Amounts in 8 of the of significant 0.001) between of sagebrush The mean monoterpen e con ten t of ATV was 2. 4 of ATW (Fig. (Table = content or among fed-upon, The amount of individual chance non subspecies. 4) and by separate Fed-upon and 83% in 1982. indicating number for both Within ATW. 35% of the transects in crude status among fed-upon, in 28 of 36 (78%) comparisons between feeding for each sub species , content were consistent. samples (Fig. procedures between differences were similar means in protein interaction protein samples was verified Differences = (P than also varied monoterpenes between in 15 comparisons subspecies. between sub- differed This number would be expected by 1 time out of 100 (Rohlf and Sokal 1969). more a-pinene. at 13.20 (minutes camphene. compound 1.8-cineole. retained ATV B thujon , camphor, within chromatographic �181 Table 9. Analysis of variance triden ta ta leaf samples collected of monoterpene in North Park. content of Artemisia Colorado. January- Aprtl , 1981-82. Source of variation Main eff ec ts Subspecies Feeding a status b Year Two-way Three-way interactions interactions Residual aSubspecies bFeeding = A. status F df MS 70.39 4 17.60 70.12 0.001 64.17 1 64.17 255.70 0.001 0.20 2 0.10 0.39 0.68 0.08 1 0.08 0.32 0.57 1. 02 5 0.20 0.82 0.54 0.09 2 0.05 0.19 0.83 31.12 124 0.25 SS !_. wyomingensis = fed-upon, and A. non fed-upon, !: P vaseyana. and random samples. �182 3.5 ~ c 3.0 N=37 ?f!. en w z w a.. a: w I- 2.5 2.0 0 Z 0 ~ 1.5 N=99 1.0 ATW ATV Fig. 5. content Range. mean. of Artemisia (ATV) leaf samples 1981-82. and 95% confidence tridentata collected wyomingensis in North Park. interval of monoterpene (ATW) and A. Colorado. !: vaseyana January-April, �183 Table 10. between Mean monoterpene subspecies of Artemisia January-April, Colorado, - levels (% DM) and tridentata univariate collected ~ tests in North Park, 1981-82. Mean Monoterpene a ATW Ub_I. 04c 0.003 U-1. 44 AT Va F P 0.000 2.56 0.118 0.001 0.002 2.87 0.098 U-1. 76 0.015 0.022 2.00 0.165 a-Pinene 0.026 0.051 4.23 0.047 Camphene 0.013 0.121 54.38 <0.001 Phellandrene 0.072 0.000 10.49 0.002 1,8-Cineol 0.155 0.495 52.40 <0.001 S-Thujon 0.023 0.482 39.65 <0.001 U-I0.54 0.058 0.063 0.05 0.816 U-l1.03 0.033 0.032 0.01 0.948 Camphor 0.317 0.753 32.67 <0.001 0.064 0.079 0.93 0.342 U-12.49 0.376 0.376 0.00 0.999 Borneol 0.000 0.055 65.24 <0.001 Terpineol 0.000 0.086 48.62 <0.001 Totals 1.160 2.617 44.16 <0.001 Fenchyl alcohol aATW tata = Artemisia tridentata vaseyana. bU = Unknown. cDenotes retention time. wyomingensis, ATV = A. triden- �184 column), and terpineol at 4.72 than Ether than ATW. extract levels of fed-upon, plan t samples were measured levels were related between sagebrush related to sage (Table There fed-upon 11). function examine sage grouse are ranked and. (3) cases the discriminating vigor possibly is a good index to resin fed-upon and non 1981 or 1982 (Table 12). within subspecies of the discriminant can be considered to their Plant height, (total and discriminating samples. of function analysis are simultaneously, (2) power to discriminate based on their vigor, 15 individual) values crude content among fed-upon, Discriminant function to between for protein con- were entered non fed-upon, analyses were done for ATW and ATV. significant vigor and crude (~ <0.10) discriminating was slightly coefficient) levels of big (Nie et al , 1975) was used can be classified Within ATW, plant 'with extract subspecies (P > O. 05) betwen food selection va rrables . and random plant separately extract for ATW does not seem to be extract analysis according and monoterpene as variables Ether samples in either The advantages (1) many variables variables tent, if ether ether Analysis A. tridentata. cases, levels, and random sage- whether food selection. was also no difference A discriminant that: to ascertain The preference ATW and ATV plant Multivariate further grouse non fed-upon, (P < O. 01) but not between years to plan t resin levels. more of the unknown ATV. brush varied ATW contained more important than crude protein. protein power were the only variables (Table to the function Overall quality 13). (higher Plant function of the derived function �185 Table 11. collected Ether extract at random content sites, North (% OM) of 3 sagebrush Park, Colorado, taxon January-April, 1981- 82. ATVa a ATW 1981 x 16.13b 12.02 7.75 1.11 0.81 1. 08 0.86 0.92 11 = Combined 1982 10.23 aATW AL 1981 1982 16.87b SE N ALa = 15 Artemisia 11 9 tridentata wyomingensis, ATV 5 = A. t. vaseyana, A. Ion grloba , bOifferent Table 12. samples Ether (P <0.05) than extract content of Artemisia value tridentata, for 1982. (% OM) of fed-upon North Park, and non fed-upon Colorado, January-April, 1981-82. ATWa fed x SE N ATVa fed ATW non-fed ATV non-fed 1981 1982 1981 1982 1981 1982 1981 1982 12.67 11.11 11. 80 11. 25 13.68 14.70 13.86 13.68 0.68 0.49 0.78 0.42 0.81 1.58 0.50 1. 82 3 4 6 4 17 aATW 19 = Artemisia 17 19 tridentata wyomingensis, ATV = A. t. vaseyana. �Table 13. Discriminant function samples of Artemisia tridentata, Sub species't analysis North Park, Significans variables Groups of characteristics Colorado, of fed-upon, January-April, non fed-upon, and random 1981-82. Function coefficients value 0.46 0.56 59.6 (59 of 99) 0.61 0.62 59.5 (22 of 37) 4.52 0.90 Eigcn Canonical correIa tion Correct classification (%) ATW Fed-upon Plant vigor 0.75 Non fed-upon Crude protein 0.66 Random ATV Fed-upon U-12.49c Non fed-upon Crude protein 0.74 Random Plant vigor 0.49 Fed-upon U-12.49 3.02 Non fed-upon Crude protein -1.88 Fenchyl alcohol -1. 51 -0.81 ATV = Artemisia tridentata bVariables plant vigor, with discriminating and content of crude cUnknown monoterpene (17 of 17) 0.80 Camphor aATW 100 wyomingensis, ATV :;: A. !: vaseyana. power (~< 0.1) from among the following: mean plant height protein, which exited total monoterpencs, the gas chromatograph and 15 individual and monoterpenes. column 12.49 minutes after injection. I-' 00 0"\ �187 was poor. tively The eigen value and canonical low at 0.46 and 0.56, squar-ed can be used the data In this 0.562 by the function. according to their over 59.6% were correctly the 33% expected and ranges of the discriminant the extent 6). Although of 95% confidence correlation (Cooley and Lohnes in 1971). ranges scores for each group of the groups that variables. This supports of variance in crude protein content. ing at sites where ATW plants select the most vigorous plants there is no overlap non fed-upon, to the dis- appear Within these analysis to be feed- and contain containing to space of the univariate are more vigorous at random sites. intervals, with respect Sage grouse to were computed fed-upon, results vigor of cases in multivariate to some extent, indicating criminating grouse (plant This is a modest 95% confidence and random ATW leaf samples were distinct than ATW plants variables by random assignment overlap intervals, were classified classified. centroids), of overlap protein cases for the discriminating The mean (group (Fig. rela- of the amount of variance When the original 3 groups. determine The canonical by the function values protein), improvement were both = 0.31, so about 31%of the var-iation was explained and crude respectively. as an approximation that is explained case, correlation sites, the highest more sage protein content. Within ATV, an unknown retention time), discriminating samples (Table crude protein, (P < 0.1) between 13). and plant fed-upon, The oxygenated influence on the function influence than plant oxygenated vigor. than crude Crude monoterpene vigor entered protein 12.49 min the function non fed-upon, monoterpene protein (at and random had slightly and considerably and plant vigor as more more values were �188 rn z w iIi 0 0 rn 0 a: z I- a.. a: 0 0 Z > ::J u, I- Z N=36 a: 0 i= 1 N=27 o I- Z ~ c{ 0.. c{ z o z ~ rn 0 0 a: -c 0 w rn a: z FED-UPON Fig. 6. function Range, scores of Artemisia mean, and 95% confidence separating tridentata NON FED-UPON fed-upon, wyomingensis. interval non fed-upon, RANDOM of discriminant and random samples �189 positively weighted mono terpene poor ~ith (positive was negatively function coefficients) weighted. Quality an eigen value and canonical respectively. The square approximately 38%of the variance func tion . classified based ment over cases Fifty-nine on their the 33% that of this correlation of the canonical percent while the oxygenated function of 0.61 and 0.62. correlation was 0.38. in the data can be explained of the ori ginal cases discriminant function would be expected was thus by the were correctly scores. a modest improve- by a random assignment of to 3 groups. Plotting and examining 95% confidence was poor random intervals (Fig. 7). 3 groups from non fed-upon overlap function increased markedly (fenchyl crude (Table protein. fenchyl entered alcohol. alcohol the coefficients (~= between o. 88. ~ < 0.05). and contrasting was diminished 12.49 (3.02) 2 monoterpenes. resulted effects was twice as large as that monoterpenes The oxygenated variable Fed-upon protein followed by samples in opposite at 12.49 and signs for The contrast- for the unknown for fenchyl were content. the unknown on the function. since the coefficient and random of the function the function. and higher to and the dis- oxygenated and camphor. and were distinct but non fed-upon Two other and function attempts samples The quality by a lower monoterpene A high correlation ing effect The function were eliminated repeated. 13). why this at 12.49 was still the most important characterized fenchyl samples. scores of non fed-upon Fed-upon Random samples alcohol and camphor) monoterpene limits of scores completely. and random analysis function the mean illustrated when only 2 exist. samples were similar. criminant about Confidence ATV samples separate the mean discriminant alcohol at (1.51). �190 w z W en w a: o o en z o t= o Z ::J U. •••• Z ~ z ::E a: o en o Q. a: w •••• o z o ::E o C) > I- Z -c ...J o Q. ~ W I- w z w C) >X Z Q. C) C) Ci5 -c w z w a: o z N=10· o a: o en -c N=7 a: Z a: o z ·3L---~----------'_--------------FED· UPON Fig. 7. function Artemisia Range. scores mean. and 95% confidence separating tridentata NON FED·UPON fed-upon. vaseyana samples. interval non fed-upon. RANDOM of discriminant and random �191 The eigen value and canonical 4. 52 and 0.90. the data respectively. This indicates samples. about separation This was verified the mean discriminant The mean (± SD) values discriminating between U-12.49. fed-upon (± 0.2); fenchyl fed-upon plants plants alcohol. = measuring = believed that gizzard samples could be gained differed 14). apparently detergent of crop evening (~< because This number by chance Gizzards poorly digestible components lignin, and gizzard non thus of partial digestion of significant alone less than higher It was and 2 independent Twelve and gizzards of the leaf material differences in 13 comparisons 1 time out of 100 (Rohlf levels of indigestible such as NDF. ADF. lignin. and lower levels of digestible ether feeding shot in the evening. crops by material. morning 0.05) between contained was assessed fiber, feeding, from each bird and Sokal 1969). extract. non fed-upon = 0.67 (± 0.2), grouse samples would represent samples would represent would be expected were: = 0.41 plants 0.05 (± 0.03), diet of sage and mineral levels in the gizzard. 8). of the Winter Diet ash. (Table (Fig. 0.81 (± 0.3). acid and neutral of 13 mean values intervals ATV plants fed-upon plants ATV monoterpenes non fed-upon extract, crop 3 oxygenated fed-upon classified. for each group = 0.22 (± 0.1). of the winter protein. scores in and non fed-upon the confidence plants at the sample were correctly and non fed-upon camphor. Chemical Characteristics The quality function fed-upon Although of fed-upon (% DM) for the = 0.09 (± 0.04); plants cases by examining high 81% of the variation for by the function. all of the original complete were reasonably Approximately can be accounted size was small (17). correlation components such or and ether as protein. cell �192 3 (/) (/) w ex: 0 w z w a, ex: w o •••• 0 Z 0 z ~ (/) 0 .•... c o w z •••• -c => LL. •••• z -c z ~ ex: o so c Z Z in •••• 0 " " <{ 0 e 0 Z (/) x 1 ex: c, W >- N=10 2 w ex: o -1 z N=7 z (/) -c w ex: o z FED-UPON Fig. 8. function Artemisia Range. scores mean. and 95% confidence separating tridentata NON FED-UPON fed-upon vaseyana. interval of discriminant and non fed-upon samples of �14. Chemical analysis Table North Park. Source Colorado, leaves from sage grouse 1981-82 (expressed January-April, P rot ei. n a OM of sagebrush crops (N = 33) and gizzards (N = 23). as % OM). EEb CCc d NOF AOF d 12.84e 77.82e 22.18c 13.73e 6.16e Lignin Mg Ca P Na 5.12e 1. 25e 0.12e 0.54 1.SSe 0.35e Ash K Crops 34.6ge x 15. ne 0.96 _O.37 0.90 0.26 0.26 0.53 0.17 0.32 0.03 0.01 0.03 0.04 0.01 23.20 13.62 23.50 72.78 27.22 21. 15 8.66 4.22 0.70 0.15 0.44 1. 21 0.23 0.32 0.51 0.89 0.49 0.49 0.45 0.15 0.32 0.02 0.00 0.07 0.08 0.02 SE Gizzards x SE aN b c = 31, 2 samples eliminated because EE = ether CC = dNDF of contamination by blood in crop. extract. cell contents. = neutral eValue differs detergent fiber. ADF (P < 0.05) from gizzard = acid detergent fiber. value. ,_. '" w �194 contents, ash, and minerals. samples probably the sagebrush The lower dry represents leaves stomach). Gizzard nutritional studies addition pass through contents of sage matter of digestive content enzymes and HCI as the proventriculus are therefore grouse of gizzard (glandular of limited usefulness in foods. Body Composition Collection contents of sage provided of nutritional large intestines their was: female (Table weight relative sex and age variation, to those of other (combined), adult male > yearling 15). This pattern yearling 16). particular nutrients female adult grouse > adult female> that females have a greater adult males have the least. leaves ability Weights of the Musculus generally male > adult weights (gIg female> lengths yearling (length a different yearling to do this than and extract needs. It appears males and that M. supracoracoideus, female (Table 698) did not vary body wtO. male to how well similar to body weight: yearling pattern male > adult can digest to their pectoralis. followed a pattern yearling relative intestine as total body as an index sex and age groups from sagebrush female> would be expected, When relative and grouse. and small and large male> This could be in terpreted sage body compo- to investigate by body weightO. 698) were compared, developed: (Table and weigh of the caeca and small and lengths of caecal of crop and gizzard Lengths follows a similar pattern. divided liver to measure significance. were measured The pattern lengths for chemical analysis an opportunity nents to compare grouse appreciably 15). and adult male> Relative among sex and �15. Average lengths Table and weights of selected sage grouse body components (± SE). Sample size is in paren theses. Age Sex a SI Lengths a Ll (mm) s Caeca Weights (g) Total ~. pect. ~. supra. Liver Fat Carcass Drvf Males l,S44±34 Adult (11) Yearling 17S±4 (11) 2,OOl±39 3,006±S4 (11) (11) 262.1±ll. 3 60.4±2.4 33.6±l. 2 (11) (11) (8) 1,760±32 163±3 1,890±37 2,370±60 244.1±6.3 54.8±l.7 29.3±l.4 (13) (13) (12) (14) (14) (14) (11) IS0.9±IS.9 (10) 63.0±9.3 (13) 33.9±0.6 (10) 32.4±0.2 (13) Females Adult Yearling 1,196±33 148±4 I,S18±34 I,S32±49 IS3.1±3.9 38.2±l.3 19.4±1.S (10) (10) (10) (10) (10) (10) (7) l,41S±27 139±4 1,45S±22 1,409±42 143.6±4.7 34.O±l. 4 20.8±l. 2 (9) (9) (9) (9) (7) (9) aSI = small intestine, bCombined c Carcass lengths dry matter LI = (9) 67.6±8.0 (10) 46.4±7.8 (9) 3S.1±0.7 (10) 33.3±0.S (9) large intestine. of both caecum. expressed as a percentage. f-' \0 VI �196 Table 16. bined) Relative (mml g lengths and small and large Park! Colorado, 0.698 intestines Jan uary- April, body wt) of the caeca of sage grouse (com- collected in North 1981- 82. Sex and age Sra LIa Caeca Adult male Yearling male Adul t female Yearling female 6.9 7.8 7.6 9.0 0.6 0.7 0.9 0.9 7.5 8.3 9.1 9.2 = aSI small intestine, age groups. This indicates the form of glycogen the breast (body LI muscles = large that in the liver intestine. availability of energy and breast muscles and protein would be constant among groups reserves relative in in to needs size). Fat con tent of sage grouse varied (Table with 1981 samples, ether extraction Ten, 10-g samples estimate 17). The number of the technique estimate required used. used individuals This variability the variability sexes, ages, to obtain ± S.D. Ten independent = ± 0.018). 2.85 fat con tent to be less than was precise, an and how many samples were to estimate was determined of the used was evaluated. of fat content. (x encountered precision were extracted from 2.83 to 2.89% fat The technique variability techniques from the same bird of samples sex and age-classes sex and age-classes, (with 95% confidence) between of large and sample preparation to get a precise ranged among and within Because even within of the variance required samples greatly of each bird 1, so 1 was the variability in fat con tent must be real. was explored could be accounted and years (Table using ANOVA to learn for by significant 18). Julian if some of differences date of collection between was �197 Table grouse Total body 17. collected fat content in North Park. Total body (diethyl ether Colorado. January-April. fat (percent of sage 1981-82. live weight) Males Adult extract) Females Juvenile Juvenile Adult 1981 1982 1981 1982 1981 1982 1981 1982 2.62 2.85 0.85 2.68 2.88 3.79 1.78 1.45 2.95 4.97 0.90 2.76 3.05 4.02 2.07 2.91 4.41 5.26 1.00 2.88 3.13 5.71 3.45 3.44 5.81 6.07 1.15 2.89 3.44 4.73 3.75 6.94 8.44 3.64 3.00 5.44 3.14 5.72 4.16 6.09 6.18 4.45 x 4.55 5.52 1.51 3.24 4.25 4.51 3.01 3.55 SE 0.82 0.90 0.54 0.24 0.54 0.61 0.68 0.76 �198 Table 18. ences in sage covar'ia te, Two-way analysis grouse body January-April, of variation Source of variance fat with date 1981-82, SS North df showing age of collection Park, Ms a and year differ- as an explanatory Colorado. F P 47.83 2 _23.91 13.04 0.001 Age 42.22 1 42.22 23.02 0.001 Year 12.95 1 12.95 7.06 0.012 0.01 1 0.01 0.01 0.927 6.69 1 6.69 3.65 0.064 67.87 37 1. 83 122.41 41 2.99 Main effects Interaction Covariate (date) Residual Totals aSex was omitted F (P = 0.72). from the ANOVA because of a non-significant �199 entered as a possible explained Julian (P = significant date Multiple for approximately birds (~<0.05) of collection 0.06). Adults collected Attempts tent scaled by structural among juveniles ~ predictive females. regressions for less variation x. provided X for juveniles were also scaled the best = for 4.80%, 1981 ~ ~ = = 0.12. 0.67). grouse fat con- or from body weight unsuccessful. years The best ranged from 22% females to 37% for adult males to of fat content on weight divided while weight divided the best fit for juveniles. were not significant (~> and bill length. did scaling be- adjusting adults. for sage fit for adults, provided by total length than (1982. for combined Regressions after equations males to 31% for juvenile of primary = in the juvenile in fat content juveniles. for juvenile by length shot in 1981 (4.03 vs. with date of collection; were largely (r2) age accounted 2.87%). and 2.75%. adult and date of collection. measures. vs. 0.29. of determination by wing length = as well of collection. (4.67 birds increased coefficients 69% for adult as year Differences (juvenile 1982. adults. weight that were most pronounced Fat content to derive of collection while sex did not. indicated were even more pronounced as a covariate 0.57; from body years among males. was most pronounced = analysis than juveniles between = 4.30%). ~ of variation. in 1982 had more fat than date of collection juveniles. Age and year for some of the variability classification tween ages and years this accounted particularly 3.16%.1982 amounts fat content Differences age-class. covariate. twice as much variability had higher 3.43%). explanatory by wing length 0.10). but However, Weights these or length accounted of primary = �200 Studies of avian carcass or petroleum ether as solvents of both are apparently there carcass samples. studies P < 0.001) crude was used or whether diethyl extract. if there ether A highly with petroleum have used diethyl extracted ether (Table was a relationship extract (paired 19). Linear between regression .•. The regression ether extract) cien t (slope) then diethyl likely that ether equation + 0.678. ether as an energy these 0.934x ether are structural lipids, to the animal. t test, regression was obtained • (petroleum The regression than other coeffi- to be 1, plus a constant. amount that is not extracted chemical structure reserve = as from petroleum If this coefficien t is assumed was equal to petroleum this constant that extract) ether and the 2 extracts, (!:<O.OOl) The r2 value was 98.4%. was close to 1. has a different possible was y. (diethyl ether and compared less could be predicted significant abilities (to my knowledge) of both were tested ether fat than diethyl to learn The extracting in which this has been docu- studies properties Petroleum have used diethyl to be equal although composition the extracting on paired ether fat. Since 1981 samples were extracted most grouse solvent, typically to extract assumed have been no published mented. composition It is by petroleum lipids. It is which may not be available �201 Table 19. carcass Petroleum samples (N and diethyl = extracts ether Diethyl 3.60a 0.37 x SE (paired of paired sage grouse 21). Petroleum aSmaller ether t test P < 0.001) ether 4.04 0.35 than value for diethyl ether. �202 DISCUSSION Winter Use Areas Si tes within winters the study were delineated 1974-75 by Beck Highway Consistency strongly North Park. area along Jackson grouse area to wintering observed caused that sage habitats heavy grouse where the study area within particularly Gill areas during birds did not occur during (but in sage grouse 1972). may be strongly snow depth. as intensively period traditional by marked quadrat to use less preferred during are and Schladweiler was not used (982) over a 17-year winter-use snow accumulations Field observations 1980. of Walden along Colorado areas (Eng in the southeast sagebrush both 1973-74 and areas Movements of marked areas during during use areas 12, identified (1982), by Schoenberg is likely grouse County 1981-82. north winter-use in Montana conditions, by Schoenberg 1980-81 and during (982) use of wintering of sage by weather of winter grouse documented Distribution influenced sage grouse 1964-65. of locations Traditional has also been areas use of the area winter implies that by sage and by Schoenberg heavy 125 during used as winter-use (975) (1965) documented area Much of the used by sage the winters of from the northeast of North during Park as this study. Schoenberg's perhaps It study more critical) was still available. indicated winter that relatively few birds 1981 when snow cover wintered was absent or in �203 minimal. appears The impetus for sage grouse to be snow accumulation et al. _l963, Beck does not cover (Girard 1975, Autenrieth the sagebrush, movements to wintering 1937, Patterson 1981, Schoenberg sage grouse areas 1952, Dalke 1982). may not leave If snow fall use areas. Vegetative Canopy Characteristics cover loafing grouse reported sites. in North Park 1975, Schoenberg at sage was lower than feeding-loafing sites 1982). Autenrieth winter-use 1972) was similar to values heights Canopy have ranged areas for feeding-loafing Average and Random Sites at 1980-81 and 1981-82 winter and 26.7%, respectively) for winter at Feeding previously cover at winter (1981) found height loafing in North Park. heights sites at winter the range The canopy in Montana be lower than canopy cover average I measured loafing or "winter sites use areas by Schoenberg in Idaho heights feeding use" areas. are in heavier, taller sites, sagebrush 1981), but feeding- in this in other loafing than within 1972). while others cover of 17.4 were also lower than reported Presumably, sites were lower than documented and height only feeding of 27.7% (1982) for winter (Autenrieth and height cover cover cover in this study. at winter Average feeding- (Eng and Schladweiler found in Montana (Eng and Schladweiler Average because reported reported 38.1% canopy and 30.4 cm in 1980-81 and 1981-82, respectively, the average (21.8 from 37.6 to 50.9% (Beck documented of sagebrush sites values in Idaho. sites feeding study studies measured or daytime feeding may sites. feedingroost �204 Typically, sage grouse that I observed would leave roosts cover in draw bottoms or on slopes and move upslope relativ..ely open areas with shorter during was probably the 2 winters areas with relatively Tall, dense brush stands of sagebrush factor and height were the primary above deep snow and available 1979-80 (Schoenberg height snow depths cover 1982). Sage grouse and cover at winter were moderate, to feed in Lack of deep snow an important low sagebrush was exposed sagebrush sagebrush. in heavy in the use of by sage grouse. areas where sage- to sage grouse also used areas feeding sites in with higher in 1981-82 when than in 1980-81 when snow depths were negligible. Shrubs at winter sagebrush. Sage grouse has been extensively et al. feeding and random sites were predominately dependence documented 1963; Eng and Schladweiler 1975; Autenrieth percent 1981; Schoenberg occurrence occurrence Feeding spp. sites was not encountered 1982). Feeding big sagebrush 1952; Dalke some Sarcobatus at random sites, had greater and lower percent probably A. cana, because of species as food plants. vermiculatus. This species probably Sage grouse (ATW) growing in alkaline sites A. argillosa, than random sites, within alkaline areas. winter 1972; Beck 1975, 1977; Wallestad or lack of use of these contained during 1937; Patterson and cover of big sagebrush (A. longiloba) distribution (Girard and cover of Artemisia longiloba, and Chrysothamnus limited on big sagebrush big areas because of its limited occasionally in association fed upon with greasewood. The large feeding difference in composition of ATW, ATV, and AL between and random sites was undoubtedly due to the strong preference �205 of sage grouse for ATW as a food plant. of the sagebrush at feeding feed along upper slopes. 1965. Schoenberg sites because rtdgetops 1982. pers. ATW comprised sage grouse in North , and well-drained observ.), sites where the majority benches Park (Gill ATW predominates. Food Plan t Selection My results from available support species ATW and against identify these the hypothesis and subspecies subspecies et al. 1979, Winward tural at sites method used where by sage ing individual plants. encounter mostly grouse to choose species sites plants (Table when a choice Preference food preferences 7). identify birds of ATW (McArthur decreased plants use struc- to feed upon. feed on ATW than a choice identify- the tops of bottoms. sage may be the best The proportion that from 90% overall that can visual1y grouse and in alkaline food plants. more linear may be a more important Avoiding indicating the way that on rid getcps , along flat benches. ATW. for leaves sage selectively The selection an observer that to selectively By feeding grouse of fed-upon to visually grouse on well-drained prised than ATW predominates slopes, for sage appearance It is possible of the plants but feed ATV has longer. With practice, 2 subspecies. features Feeding 1980). grouse of sagebrush. is less obvious. with more of a whorled the sage ATV as food plar. ts is obvious. leaves identify that way ATW com- to 70% at mixed- misidentification may be common must be made. for ATW and against of mule deer area of Utah and Oregon. deer over ATV as food plan ts differs (Odocoileus hemionus) ATV was strongly ATW and A. 1. tridentata (Sheehy in the Great preferred as forage from Basin by 1975. Scholl et al , 1977. �206 Sheehy and Winward 1981, Welch et al. Brunner (1972) rarely grazed possibly described Mule deer Park appeared rabbits eaten by sage grouse area northeast leaves sagebrushes have been Rasmussen and Griner Browning 1958, Sage grouse feeding edges, irrigation of the sagebrush that sage grouse noted when available, (1972:207) listed in an primarily (1963) but 1975). upon that that black snow alkali sagebrush grouse." A. ~, eaten primarily in the summer. and A. tripartita as well as big by sage grouse 1938, during Leach and Hensley summer (Girard 1954, 1937, Leach and 1970). selected on vegetation has been 1963, Walles tad et al. Dalke et al. seem to be eaten Martin big sage- diet of sage grouse Wyoming. were feeding Brunner of A. arbuscula, sagebrush 1982g.). Big sagebrush Dalke et al. Pygmy for ATV have not identified over big sagebrush by sage (White et al. 100% of the winter in winter. in North of ATV. equal preference from observations limited availability. as "seldom eaten road 1952, One ecotype, americana) exclusion to subspecies. concluded sage was preferred Leaves (Antilocapra researchers of Eden Valley, sagebrush to deer. 207) as a "choice fall diet for sage showed virtually (Patterson (1952) Other palatible (p. grouse as comprising in most areas depth as ATV as ATW was not tested sage black identified idahoensis) Previous Patterson he tentatively to use ATW to the virtual tridentata; identified in Nevada, and pronghorn (Brachylagus or A. t. However, and ATW as highly ATW, was described grouse." brush plants 1981). larger, growing ditches was apparent. more vigorous on what resemble or other plants primarily by mima mounds or along disturbance sites where release Mirna mounds are low, dome-shaped �207 mounds of earth found in the western glacial activity activities (Akley and Brown by pocket and well-formed, (Spermophilis toward 1954). gophers whereas to the latter shape squirrels theory, (~. squirrels with no tendency richardsonii) resemble those are numerattributed squirrels. Gill (1965) following Keller et al. North Park into 3 types inches (12.7 cm); inches (25.4 A-2, sagebrush cover. Barber according 5-10 inches in height. crn ) shorter types et al. observations tion for taller, (1965) type (1941) grouped to growth appear descriptions preferred plants). feeding type sage results that associations, Thus, type type. (selecfrom Gill's consists A-2 almost entirely cf ATV. these interpreted for ATW and against observations for with study it is clear the A-I in the primarily grouse of this However, and A-3 almost entirely as selection up to 5 to feed in the A-I and photographs of ATW or ATW-A. longiloba (A-3) in and A-3, >10 sage grouse penned to contradict more vigorous big sagebrush A-I, cm); the taller (1969) found that types form: (12.7-25.4 He observed and using to all 3 sagebrush These by earth-moving mounds formed by ground Ground either (Thomomys spp . ) are particularly ous in North Park and mounds in this area access created by rodent According spp , ) are of indiscriminate dome formation. to ground States (Newcomb 1952) or more likely, mounds created large United of ATW, can be ATV as food plants by sage grouse. Selection been noted that of food plants for other red grouse heather (Calluna on the basis species (Lagopus vulgaris) of grouse. lagopus of physical attributes has Moss et ale (1972) observed scoticus) pr'efer-r-ed to feed on 20-30 cm tall and 3 or more years old. �208 Gullion (1966) and Svoboda grouse (Bonasa aspen that umbellus) (Populus selected tremuloides) capercaille stunted and Gullion (Tetrao pines (Pinus urogallus) species within potential grouse (Huff herbivores this plant tested by Mattson species intake winter 1980). has been of to 1 or a few Selection among and for other and Moss 1970. Pendergast and Boag 1972. Moss 1972. Moss et al , 1972. and Salo 1973, Evans to explain to grouse. in this 1980) states of nitrogen. food selection These of grouse theories study. The that herbivores and Dietz et al . 1974. Pulliainen selecting nitrogen nitrogen intake results from my study (crude is an important "nitrogen II select parameter theory foods to way to achieve con tent. There are potential food (Miller 1968, Huff and Robinson 1981). clearly of were the basis among and within and Iivanainen and multivariate patterns The most efficient foods with the highest to maximize their protein) diets documented 1970. Miller et al , 1971, Moss 1972, Gurchinoff Doerr or 1978). advanced examples to feed in diseased their 1972. Pulliainen seem to apply is to select numerous species 1976. Herzog maximize their (1962) reported feed upon a wide range restrict and Robinson for the hypotheses (reviewed preferred of grouse 1970, Gardarsson Two theories Seiskari quaking for Food Selection yet food plant and Gullion 1974. Ellison ruffed more decadent 1966. Moss and Hanssen 1971, Gurchinoff Svoboda species in the fall, (Gullion that spp.). Most. if not all. materials older, to feed upon. Reasons plant (1972) found The univariate indicate in sage 1972. that grouse nitrogen food selection. �209 Sage grouse gen, fed primarily (2) at sites more ~itrogen, preferred (1) on subspecies where plants non fed-upon, largely These against There on their protein tion for taller However, fed-upon of 2.2 percentage whereas fed-upon average of only 0.83 percentage plants. Selection in monotypic not vary. and other disturbance and apparently relatively points release or higher factors could also increase because plant Regressions or vigor and crude where contained contained selecan plants, an than non fed-upon or vigor did not occur plant size and vigor did was at mima mounds sagebrush because in height, at these con ten 1. vigor, sites of a nutrient due to the disturbance. protein of protein for both subspecies. was variability Perhaps of water plants occurred and more vigorous availability if high protein than non fed-upon height there 2-4. were more protein of plant content. fed-upon. parameters occurred could and did occur and taller these at transects more protein sites where in protein young, plants of tall sagebrush Where selection protein within subspecies plan t height where no selection on the basis stands plants (~>0.05) ATW plants points between to learn between average Crude of hypotheses hei gh t or vi gor. or more vigorous plants parts and vigor, or vigor were non-significant content. and non- samples in the discriminant content is no simple relationship contained within preferred individual height protein subspecies discriminating support selected of plant crude were iden tifi ed based on height results sage grouse on the basis regressed variable the most nitro- the most nitrogen. and random sagebrush analyses. Because plants which contained was the 2nd most important function of the preferred and (3) on individual subspecies containing These is �210 Crude to other but protein values content reported gen era lly higher (Table 20). be lower than fiber for sagebrush than Protein protein gests reviewed that of plant defense compounds inhibiting bacteria by Bryant digestive (Rhoades tial digestibility 1977. Kelsey 1982), coumarins Kelsey et al. and other unidentified Brown et al , 1975). factor in sage indicating phenolic grouse sesquiterpene However, Monoterpenes in a variety et al. are components (Hanks et al. et al. et al. et al. similar of literature of mono- properties 1976, Pic man et al. of plan t families including 1978), as a possible properties "essential" 1978). 1971, of the depth deterrent of the (Buttkus 1975, Kelsey 1971. Kelsey because et al. poten- 1970, Brown et al , 1975, work has indicated (Rodriquez several 1981, Kelsey were investigated and feeding These digestive terpenoids compounds food selection recent lactones or killing including et al. Monoterpenes anti-microbial terpenes. found (Shafizadeh low levels with proteins. contains (Brown sug- reducers. Sagebrush and Melnikoff theory constituents). 1978. Welch and McArthur flavanoids lactones plant This containing act by complexing 1976). (Shafizadeh 1978), (1980). to grouse or as digestibility compounds et al. should of the higher pertinent or by inhibiting and Cates et al. sesquiterpene compounds reducing because theory (secondary enzymes, in winter, leaf and stem samples feed on plants act as toxins reducing collected for leaf and stem samples and Kuropat selectively compounds was similar of stem material. food selection can either Digestibility reported of leaf samples content herbivores leaf samples leaf samples of combined content 2nd herbivore has been values content and lower protein The of big sagebrush or Pinaceae, for 1982). "volatile" oils Cupressaceae. �211 Table 20. Winter protein stems of Artemisia levels tridentata reported Leaves and stems 14.1c 10-11.S in the literature. 12.0 ATa Leaves and stems Leaves c 10.8 13-l4k 14.0c k and leaves and ATVa AT\Va Leaves (% DM) of leaves. Leaves Leaves and stems n.2d 8.1b 16-17g u.c" 10.2e 11.7e j 11. s" 9.4e lS.2 11. Of IO.Of 13.6j 11. 2f II.9f 10. Of 12.9j 12.9f 11. ::/ i 10.4 IO.Sf 11. 2£ f 11. 3 lI.7f 11. 7£ 12. Of Means 12.1 12.4 aATW ~. AT = A. tridentata = ~. tridentata bMilchunas c 12.1 Present et al. (no subspecies ATV = 9.8 A. tridentata vasey- specified). (1978) (Middle Park. Colo.). study. Sheehy (1943). (1975) (3 areas fWelch and McArthur Colo. [II. 13.9 wyomingensis. dKinney and Sugihara e 10.9 in Dreg.). (1979) (location of 14 accessions; Nevada [1), Utah (111). gPatterson (1952) (Wyo.). hSmith (1950) (Utah). iDietz et al , (1962) (Colo.). jBarber et al , (1969) (North kKelsey et a1. (1982) (Mont.). Park. Colo.). Ariz. [II. �212 Myrtaceae, Labiatae, been identified Terpenoids tures plants adapted compounds penes carbon compounds their is uncertain. function have been allelopathy (Schlatterer Weaver and Klarich (primarily and Tisdale 1977), toxicity to suppress have potent menzeisii) the in-vitro and the rate growth 1970; Nagy and Tengerdy Radwan and Crouch and groups (2) oxy- to the physiology as energy of the plant reserves functions for the external These to plant include 1969, Klarich and Weaver 1973, to bark beetles (Cobb et al. anti-bacterial (Smith properties. from sagebrush. (Juniperus spp.) fatty Volatile oils Douglas-fir have been acid production (Nagy et al. 1965, 1968). of rumen microorganisms, of volatile rumen microorganisms (Loomis and for :monoterpenes. fungi or juniper and sesquiter- of as metabolic byproducts, Several monoter'penes ) extracted (Pseudotsuga steer may serve hypothesized struc- aldehydes). thought 1973). oil- of these into 2 general (1) hydrocarbons 1979), and pathogenic Monoterpenes quantities respectively and relationship Monoterpenes large volatile specialized Monoterpenes residues, esters, are no longer classical in having are subdivided (alcohols, physiology digestion, 1973). skeletons: (Loomis and Croteau Sturgeon primarily or accumulating Monoterpenes Terpenoids plant may be unique plants; have 1972). (Devon and Scott in higher 2 and 3 isoprene on their al though products (Loomis and Croteau 1973). genated plant to secreting contain Croteau based as natural may be ubiquitous containing Over 400 monoterpenes and Compositae. rate of deer shown of and 1964; Oh et al. 1967, 1968, 1967e_.!;!. 1968; Eller 1971; Radwan 1972; 1974; Schwartz et al. 1980~). �213 It is obvious that lems to herbivores. rent effects Mule deer Several seem to exhibit fermentation slight to feeding or no selection based within on monoterpene 1974, 1978; Sheehy and higher (198012) found a strong oils added Tassel-eared squirrels (Sciurus fed on twigs and seedling terpene hydrocarbons Preferences sagebrush Gill (1965) and Barber volatile oil levels however, monoterpenes studies have not accounted content between containing plants. species, selectively average 1982). of big (White et al. of volatile by sage grouse A-3 sagebrush reports mono- 1982~). oils in were lower type. were comparing It ATW volatile oil levels. to make conclusions in determining gophers of 2 subspecies content in the non-preferred micro- et al. 1981, Radwan et al. preferred that both of these microbial levels of volatile lower than et al , (1969) found levels types by by tame mule deer. for 15 accessions and A-2) and ATV (A-3) It is difficult between to monoterpene the A-I and A-2 sagebrush browsed Schwartz and pocket (Faren tinos et al. of pygmy rabbits composition which promoted and preference aberti) Douglas-fir which inhibited stems containing were unrelated is likely, correlation to food pellets deter- by herbivores. trees hydrocarbons). inverse prob- 1975; Scholl et al , 1977). levels of monoterpenes (monoterpene potential have in vesti ga ted possible lower levels of monoterpenes bial fermentation (A-l serious Connolly et al, (1980) found Douglas-fir deer contained than studies of sagebrush (Radwan and Crouch juniper present of volatile oils or monoterpenes or among species However, monoterpenes about food preferences the general of herbivores. for or have mixed variability subspecies, The univariate and populations analysis role of Many in monoterpene of monoterpene- (ANOVA) of monoterpene �214 content (both monoterpene genated subspecies content mixed) and feeding monoterpenes of fed-upon. ences plants and random within grouse selection for low volatile high levels by mule deer oil treatments of volatile It is interesting were oxygenated strongly hydrocarbons were strongly Variable preferwhere mono- fed and non fed-upon 2.62%). by Schwartz Oxygenated bacteria. Similarly. et al , (1980Q_) due to the relatively that oil components. Some monoterpenes with glucoronic monoterpenes to feeding containing to eliminate are monoterpene by small mammals. volatile oils may or detoxify can be detoxified acid discrimi- by sage grouse Conversely. to plants to differen tial abilities reaction = level at low for individual and not fed-upon be due in part condensation was total 1.16%). but were the between inhibitory animal responses study. the 3 monoterpenes ATV fed-upon to ruminant than treatments. that monoterpenes. inhibitory = may have been to note most between in this level noted oils in their discrimina- Monoterpenes preferences discriminating within ATV (mean monoterpene the strong nated sage were many or may not influence ATW (mean monoterpene variables samples This monoterpenes were better in the ANOVA). This was illustrated did not influence analysis. ATW and there non fed-upon. oxy- non fed-upon. subspecies: monoterpenes may not be detectable most important plants within between several fed-upon. Oxygenated of herbivores. terpenes between status (which was used concentrations even though in the discriminan t function on feeding more ATW samples. monoterpenes no relationship to mixing of data from both had no influence tors status. discriminated and random ATV samples was due in part indicated (Dean et al. volatile in mammals by a 1967). The �215 amount of glucoronic cinereus) fication has been acid excreted estimated of more than (Eucalyptus fication daily by the koala to be sufficient leaves (Eberhard of volatile oil components been indicated (Eberhard terpene (Eberhard that White et al , 1982Q). steers that (Eller detoxi- values has through mastication and eructation the skin and sur- et ale 1975) has been hypothesized. have been animals were feeding Whatever 1971) and koalas only 16.6 and Selective germicidal 1981). or through levels in stomach contents levels in sagebrush with eucalyptus et al , 1975). 1971. Welch and Pederson face of the lungs for the detoxi- et ale 1975). with higher Loss of volatile oil components (Eller to account 50%of the volatile oils ingested punctata) (Phascolarctos shown to be about 20%of upon 1982. the mechanism. (Eberhard 15%. respectively. Mono- (Cluff digestion et al. trials with et al , 1975) have shown of the oils ingested are excreted in feces or urine. Sage grouse drive do not have the ability off monoterpenes. is greater than that the non-grinding high intake terpenes. Their for any other gizzard unique of terpenoids. to this species the leaves which could then be absorbed leaves size. Galliform caecal orifices are unable caeca is filtered and Boag 1974). dry of monoterpenes vertebrate. Grinding Intact little intake matter 1975. 1976). to pass to masticate would release By not grinding (Fenna sagebrush that to its the or shun ted into the caeca. are small. and material the caeca likely is an adaptation section of their entering of the caeca Even in galliforms with grinding enters to (as a percentage) It appears into the caeca because by villi in the proximal sagebrush gizzards. large the (Fenna relatively and Boag 1974; Gasaway et al , leaves. sage grouse can �216 prevent terpenes from entering tion by caecal bacteria. through the digestive Barber (1968) found grouse caeca. excluded It is likely that tract that cellulose oils were concentrated studies Barber et al. Barber fecal droppings were more concentrated found insignificant was insignificant that volatile relative volatile to levels oil levels in than in fecal droppings. Both volatile oil levels in caecal contents. Volatile oil components could easily have evaporated pings as droppings in both studies. in sage material was (1969) found that (1968) found pass on in fecal droppings. if leaf structural in sage grouse contents. crop contents digestion This would be expected VFA produc- most of the terpenes and are excreted from the caeca. in gizzard the caeca and inhibiting This would underestimate the extent analyzed from fecal drop- were several of concentration days old. of oils in fecal droppings. Lack of a grinding gizzard may be advantageous for limiting exposure to te rpen es , but lower food digestibility is a probable quence. Sagebrush grouse Thus. leaves the surface is minimal. pass area of leaf material The storage capacity sate for poor digestibility. capacity of the gizzard contents of the 3 heaviest group. Gizzard crop. probably digestive the sage exposed An approximation crops (DM basis) averaged seem to have evolved strategy. makes this possible. gizzard The high weights storage of the for each sex and age 1.44 times that of the a high intake. quality enzymes may compen- of the relative by averaging and gizzards gut whole. to digestive of the sac-like was obtained capacity Sage grouse passage through conse- high rate of of the leafy winter diet �217 Total monoterpene monoterpenes identified ATV (Buttkus McArthur et al. 1977; Kelsey et al. comparable and sesquiterpenes (Buttkus terpene extracts of plant and phenolic resin patterns of herbivores extracts fed-upon 1981). components material levels (Bryant samples within Monoterpene levels are Volatile oils contain di - as well as monoterpenes have been used and correlated and Kuropat subspecies an index phyl'ls , carotenes, terpenes, In conventional mate fat content to be useful. and fatty proximate of plant as indicating extracts and phenolic material. materials compounds (high contain in gizzard contents than in crop contents. The detergent analysis as an alternative is chemically and non Ether ether extract extract energy) extract chloroin ether. is used to esti- levels are inter- forage. more indigestible than fat (Moss 1973. Bryant extractable 1981). are all soluble High ether this contention of ether of of the Winter Diet analysis. a high quality of plant acids support developed 1980, Bryant Waxes, phenolics, My results digestion as indices to food selection did not differ. Chemical Characteristics material for ATW and of ATW and ATV samples and of fed-upon may be too crude ether values 1978. 1982; Welch and to volatile oil levels. and other of individual et al , 1977). Ether preted percentages were similar to published 1981; Welch and Pederson not directly Ether levels and relative as ether plant resins and Kuropat ext rac t levels indicating However. little 1980). were higher or no components. system of partitioning to the traditional partitioned feedstuffs proximate into cell contents. was analysis. Plant NDF, ADF, and �218 lignin. Cell con ten ts consist soluble protein, and starch, cellulose, lignin, partitions lignin protein and cellulose 1982). into cell contents, Procedures to ruminants; of other extent, Protein forages brush et al. winter the to assume these of grouse diets of leaf and stem range (Wallmo et al. cell contents in sagebrush result leaves. from comparing or readily To a large leaf samples of sage- leaf and stem samples or stem samples of other content available Milchunas to evaluate values By difference, are higher differences to combined forages. materials leaf samples from sage grouse from sagebrush et al , 1978). components these brush plant the quality lower than comparable forages 1977, Milchunas digestible ADF but caecal digestors). were generally samples 1982). and fiber-bound it is reasonable NDF and ADF levels of sagebrush crops (Van Soest to partition to evaluate sugars, of hemicellulose, from hemicellulose can also be used (monogastrics protein lipids, NDF, and ADF were developed of foodstuffs procedures digestible while NDF consists and fiber-bound (Van Soest quality of highly of crop samples was high compared on sagebrush 1978), but winter ranges similar to other to other (Wallmo et aI. 1977, winter-collected sage- leaf samples. These results suggest that winter forage. The extent these nutrients is unknown. digestible by ruminants inaccessibility ing that on a plant bearing It may be that fiber leaves. high quality are able to digest plant constituents by sage grouse to caecal bacteria. which lack a grinding green is a relatively to which sage grouse are indigestible of the plant sage grouse, sagebrush of the It is not surpris- gizzard, The high crude because partially exist protein in winter and other �219 readily digestible cell contents of this leafy material poor digestion. to compensate for apparently explanation for the well+documen t ed dependence of sage grouse on sagebrush, since are unavailable in forages This are necessary of similar high quality may be a nutritional winter. It is difficult grouse to that However, of other comparing the quality sage (Table 21). grouse exceeds diets grouse The crude species. of other grouse requirements grouse in view of the high levels tive to winter diets that a high to sage quality (i.e., grouse. of grouse diet of sage of 2. those Mineral of other in protein in sagebrush underscores rela- the concept winter diet is critical grouse. Baseline nutrient levels in the winter have more meaning when related (weight pared to illustrate for plan ts highest content) differ. grouse species by a factor of protein This high protein may indirectly also exceed grouse is interesting of other diet of sage of the winter grouse by sage probably are from other content diet of sage diet of sage requirements of the winter protein Selection of the winter since their those of most other in the winter grouse grouse the quality of winter how different levels to compare the quality gain, survival, to nutrient clutch diet of sage to some aspect size. chick grouse of bird survival, will performance etc , ) and com- levels above or below baseline. Body Composi tion The relative large intestine, in other grouse length 698 (mm/gO. body weight) of the small intestine, and caeca of sage grouse (Leopold 1953. Beer was generally 1955, Pendergast smaller than and Boag �N N o Table Chemical 21. analyses of the win ter diets of grouse (% OM). Cpa Species Centrocercus urophasianus La~ 15.9 leucurus La~ ~gopus scoticus Lignin 22.2 13.7 6.2 6.0-S.2 Oendrasapus canadensis 25-27 ur()sall~..! aCp = crude CA P Na K Mg Reference 1. 25 0.12 0.54 1. 55 0.35 Present study 0.54 0.18 0.01 0.68 0.16 May 1975 0.2-0.5 0.0-0.1 0.0-0.1 0.3-0.6 0.1-0.2 Moss 1968 Gasaway 17-lS 1976a Pendergast 1971 and 8.9 Gurchinoff Robinson and l'JU 6.3 Ellison 6-7 Hoffmann 1961 13.7 Korshgen 1966 11.4 Tetrao 25-26 6-S Oendrasapus obscurus Bona sa umbellus AOF 9.5 10-14 La~~ a -~ - :-:,-=-==-~=~--,='" = a NOF 16-26 50.S 27.1 7.0-7.3 protein. NOF = neutral detergent fiber. 0.1-0.4 O. I 0.45 0.25 0.2-0.3 0.1-0.2 AOF = acid dcrer-gen t fiber. 0.1-0.2 0.03 0.4-0.5 0.1 0.54 0.4-0.5 Huff 0.1 1966 1970 Pulliainen 1978 Boa g �221 1973, Moss 1974, May 1975, Gasaway ably due to the high and needles 1953). comprising Differences and female sage Yearling fiber in relative of other tract is prob- of the buds, grouse that also had longer twigs, (Leopold tract of male for egg laying for low fat levels a digestive ptarmigan difference of the digestive may be an adaptation by having Female white-tailed diets lengths females may compensate to some extent This con ten t and low quality the winter grouse 1976~). by females. (relative to adults) is relatively digestive long. tracts than males (May 1975). Sage grouse Braun weights were similar (1978) for sage grouse March from 1965 to 1976. Beck and Braun of grouse (Pendergast of grouse lose or maintain protein, reserves tively have winter have been winter those of yearlings this assuming complete of M. pectoralis Average of breast energy and males have larger species some species by summing the reserves. 4.4, et al , 1975). and Fat, 4.2 Energy of fat, catabolism and M. supracoracoideus of glycogen muscles reserves study. 1982). of 9.3, catabolism by 1947, West and and protein 1973 in Thomas noted for other can be estimated 1978), and complete oxidation 22). noted (Bump equivalents January- during 1976~, Thomas glycogen, energy 0.38% of the wet weight (Table over over grouse of fat, 19%, respectively, (Grammeltvedt in weights 1975, Gasaway (Dargolts were estimated 0.43 and weight of sage equivalents Kcals/g, respectively of 26 and during with trends by Beck and and Boag 1973, May 1975) although and glycogen reserves Park over winter Meng 1968, Thomas et al. (Kcal) in North (1978) were consistent in body weight energy reported trapped Increases Increases Energy to those which comprises and liver, respec- of adul ts are larger reserves than than females within �222 Table 22. April, Estimated North Park. energy reserves Colorado, a of sage grouse, January- 1981-82. Adult male Fat (Kcal) Yearling male Adult female Yearling female 583.3 625.9 429.6 1,397.2 Proteinb M. pectoralis 88.1 82.1 51. 5 M. supracoracoideus 14.6 13.3 9.2 8.2 M. pectoralis 4.7 4.4 2.7 2.6 M. supracoracoideus 1.1 1.0 0.7 0.6 Liver 0.6 0.5 0.3 0.3 Glycogen c 1,506 Totals aAssumes9.3 bAssumes and 48.3 Kcal/g 4.4 Kcal/g 19% of M. pectoralis (Grammeltvedt cAssumes and liver, fat 685 (Thomas protein 690 490 1982). (Thomas 1982) and catabolism and M. supracoracoideus', of 26 respectively 1978). 4.16 Kcall g glycogen respectively, is glycogen and 0.43 and (Thomas 0.38% of muscle et al . 1975). �223 age classes. Energy the total energy of total energy protein of fat reserves reserves. reserves contributed comprised Breast muscles compared to energy comprised reserves sage grouse as a percentage et al. assessed size of the energy by comparing metabolism. Standard body weight (Zar formula: energy was low, but for other of similar to grouse (Thomas 1982, Brittas = (Kendeigh poorly SMR for galliforms to free-living and West (1968) found (-20 C) willow ptarmigan also applied SMR plus the costs associated to sage grouse cost of free-living living was computed the true 1968). Existence metabolic to using the SMR state, and metabolism of locomotor activity includes for the additional predator escape, metabolism for winter-acclimated I assumed from SMR. warmer that these the existence at -20 C by West (1968), Park are substantially assume that (Zar with foraging, and computed of sage grouse in (SMR) is related to be 1. 22 times SMR and estimated as 1. 72 times SMR. can be expended and in a post-absorbtive conditions. existence grouse can be computed The metabolic cost of free-living cost of locomotor activity of free-living to energy 72.6 (body weightO.698) 1970) includes caged birds. of sage metabolic rate animals at complete rest relates reserves reserves (or basal) 1968). SMR ± 15.3 describes in North Fat content 1982). The relative etc. reported 1982). 1975, Thomas and Popko 1981, Thomas Marcstrom thus of body weight than values muscle in willow and rock ptarmigan. (Thomas and Popko 1981, Thomas higher only 7 to 14% to the 37 and 31%breast respectively or slightly from 85 to 93% of than the cost relationships metabolism and Since the cost of freeand winter this, rate of sage grouse temperatures it is reasonable in North to Park lies �224 somewhere between Given these existence assumptions, metabolism and cost of free-living. energy reserves last from 3 to 4.5 days. Reserves about of fasting 4-6 days and those unlikely content that fasts and body weight food energy this period. during breeding of fasting adult occur Energy and nesting sufficient reserves adult and weight loss for sage grouse would females would last It is winter. 1978) increase Since fat over winter, to meet metabolic needs are probably activities, juveniles males 5-8 days. during (Beck and Braun must be more than during demands of this duration of fasting period more important of high energy (Beck and Braun 1978). �225 MANAGEMENTIMPLICATIONS Resul ts of this lished guidelines Autenrieth feeding North Park, 1982). should sage (Beck nutritional characteristics. should brush terpenes possible (Regelin of snow depth and quality and well-drained in is most topographic diversity ridgetops areas genotypes with sageof big 1979) and low in mono- be selected. use of ATW stands by use of snow fences of sagebrush of sagebrush (Carpenter recognize forage 1976, Laycock wit hin a subspecies should topographic. be delineated of these ~n It may be normally or carbon unavailblack and Wallmo 1975). Fertilization sible grouse 1977. particularly availability If possible. 1981) should et al. composition. should (Welch and McArthur sage specific species Revegetation be encouraged. (Welch and McArthur able because have to create pub- from disturbance. must be disturbed. of ATW. high in protein to encourage 1982). in reclamation the growth (ATW) should sagebrush use areas (Braun use areas. snowfall when habitat use areas be encouraged winter Winter use areas to heavy If winter which favor winter to previously habitats and protected 1975, Schoenberg of moderate limiting. Sage grouse grouse support sage grouse be identified structural years add further for managing et al. sites, research with nitrogen available 1982). to enhance to sage grouse Sage grouse that contain the highest treated plants. Nitrogen amounts appears the quantity may be pos- can select of protein plants and thus to be an important �226 factor in sage grouse food selection must be physiologically that increases resulted nitrogen (Moss et al. increased levels, in dietary in increased Dietary critical. protein clutch size, and, by fertilizing monoterpene and Middleton of captive female ruffed egg quality, sagebrush. grouse reproductive Effects (1982) found grouse and chick survival. to red Sage grouse content, high levels Beckerton has also been related 1974, 1975). by inference, breeding success potential may be of fertilization and palatability, on protein particularly of ATV, should be investigated. Further ences research where species is needed and subspecies from that in North Park. food selection relationships ductive performance dietary investigated nitrogen, by sage grouse sage grouse between energy should or during egg laying. differs monoterpenes in these areas. reserves, and Inter- and repro- be investigated. atten tion should be given to the timing of nutritional winter food prefer- composition of sagebrush The relationship should be further between to document Particular bottlenecks. i. e. , �227 LITERATURE Aldrich. J. W. 1963. J. Wildl. Manage. CITED Geographic orientation 27: 529-545. Amst rup , S. C. 1980. A radio-collar Manage. 44:214-217. of American Tetraonidae. J. Wildl. for game birds. Anderson. A. E. 1960. Effects of sagebrush eradication by chemical means on deer and related wildlife. M.S. Thesis. Colo. State Univ .• Fort Collins. 72pp. Arkley. R. J .• and H. C. Brown. 1954. The origin (hogwallow) microrelief in the far western states. Am. 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Oreg. State 1980. Taxonomy and ecology Univ. Agric. Exp , Stn. Bull. of sagebrush 642. 15pp . in Oregon. • and E. W. Tisdale. 1969. A simplified chemical method ----::-for sagebrush identification. Univ. Idaho. For , , Wildl. and Range Exp . Stn. Note l I . Zpp . Young. A. 1965. A chemical study of the taxonomy of section Tridentatae of the genus Artemisia. Wyo. Range Manage. 198:2-9. Zar , J. G. bi rds. Prepared by 1968. Standard metabolism Condor 70: 278. /~CV) E. ~-dZ;;:; Thomas E. Remington ~ Graduate Research Assistant Approved by -:-,-&",-~",,::;~",,-,,--,---=L~.-L~~=-:"":~ Clait E. Braun Wildlife Research Leader __ comparisons among orders of �241 Colorado Division of Wildlife Wildlife Research Report April 1986 JOB PROGRESS REPORT Colorado State of Project 01-00-045 (W-37-R) Work Plan Job Title: 3: Avian Research Job.....!L Sage Grouse Distribution and Habitat Use in the Gunnison Basin Period Covered: 1 July 1984 through 30 June 1985 Author: . Jerry Hupp Personnel: C. Braun, C. Coghill, R. Griffin, T. Henry, J. Houston, D. Masden, P. Mason, J. 01terman, T. Sherrill, Colorado Division of Wildlife; J. Hupp, R. Ryder, Colorado State University; M. B1ymeyer, J. Capodice, S. Hayes, U.S. Bureau of Land Management; C. Mo1itoris, B. Shuster, B. Wallis, E. Zieroth, U.S. Forest Service; K. Lair, Soil Conservation Service. ABSTRACT Sage grouse (Centrocercus urophasianus)distribution and habitat use were studied near Gunnison, Colorado during July 1984 to June 1985. Radio-marked sage grouse remained in sagebrush uplands during summer 1984. Chicks comprised a large portion (66%) of harvested birds during the fall hunting season indicating that production in 1984 was good. Estimated female nesting success (69%) was also good in 1984. Sage grouse were widely distributed across the Gunnison Basin between January and March 1985. There was no apparent preferred wintering region. Sage grouse flocks were not randomly. distributed among terrain categories during winter. Drainages and topographically low sites with slopes < 50 were preferred. Sage grouse rarely used slopes with north or east aspects or steep (> 150) slopes. Most feeding sites were in stands of Artemisia tridentata vaseyana. There was little feeding activity in stands of A. nova. Snow depths in 1985 were approximately 60% of depths measured in f984~s a result, sagebrush was more available and rarely snow covered in 1985. Sage grouse attendance was surveyed at 17 leks during April and May 1985. Mean peak male attendance (25.5 ma1es/1ek) was slightly higher than in 1984 (23.3 ma1es/1ek). Fifty-two sage grouse were captured and marked in 1985. Body weights of adult (x = 2,085 g) and yearling rx = 1,761 g) males were not different from those (adults = 2,038 g; and yearlings ~ = 1,786 g) measured in 1984. x �243 SAGEGROUSE DISTRIBUTION ANDHABITAT USE IN THEGUNNISON BASIN Jerry Hupp P.N. OBJECTIVES The primary objectives of this study are to: (1) evaluate winter and spring distribution of sage grouse in the Gunnison Basin, Colorado; (2) describe vegetation and structural components of seasonal sage grouse use sites; (3) compare spring lipid reserves of male sage grouse in Jackson and Gunnison counties; and (4) develop a sage grouse habitat management plan for the Gunnison Basin. SEGMENT OBJECTIVES 1. Monitor sage grouse movements between summer and fall 2. Evaluate sex and age composition of hunter-harvested birds. 3. Evaluate 4. Describe shrub structure and sagebrush spring sage grouse use sites. 5. Describe 6. Compare snow depths terrain categories. 7. Locate previously undiscovered grouse 1ek attendance. 8. Capture sage grouse on or near leks in spring. Obtain size and weight. Mark captured individuals with leg transmitters. 9. Obtain information Gunnison Basin. sage grouse winter topographic distribution characteristics and winter and analyze data. fall population species sagebrush leks in Present adult composition at winter and among years and use sites. availability the clutch of from examination in the Gunnison Basin. of winter on sage grouse 10. Evaluate spring lipid reserves Jackson and Gunnison counties. 11. Evaluate meetings. of the use areas. Gunnison Basin. size male pertinent success grouse results sage measures of body bands and radio and nesting sage Survey at in the collected professional in �244 DESCRIPTION OF STUDY AREA The study area in Gunnison and Saguache counties has previously been described by Hunter and Spears (1975) and Hupp (1984) and will be described in the final report. METHODS Summer and Fall Movements Movements between leks or nest sites and areas of summer use were evaluated by relocating sage grouse radiomarked between March and June 1984 (Hupp 1984). Radio-marked sage grouse were not intensively monitored during summer and fall 1984. Individuals were relocated 1-2 times monthly until radio transmitters failed or were recovered. Sage Grouse Harvest Wings of harvested sage grouse were obtained at 17 wing barrels and 2 hunter-check stations in the Gunnison Basin during the 8-23 September hunting season. Check stations were only operated during 8-9 September. Age and sex of harvested birds were classified by primary characteristics (Beck et al. 1975).. Age of harvested chicks was estimated from primary molt and growth. Nesting success of female sage grouse was estimated from primary molt progression. Sage Grouse Distribution and Habitat Use Sage grouse winter distribution was evaluated by searching 143 O.79-km2 circular survey plots between 3 January and 15 March 1985. Random selection of plots was stratified among 8 regions of the Gunnison Basin (Fig. 1). Boundaries of plots were drawn on 7.5 minute topographic maps. Terrain within survey areas was classified by slope and aspect. An Abney level was used to verify slope steepness. I identified 7 terrain categories (Table 1). Boundaries of terrain types were drawn on topographic maps and each category was searched on foot for evidence of grouse use. Sage grouse use sites were determined from flock observations or tracks in snow. Track observations were either feeding or roost sites. Based on track observations, sage grouse usually landed, foraged in a localized area, and then flushed. Therefore feeding areas were discrete and it was possible to identify mUltiple areas of feeding activity in some survey plots. Terrain characteristics were described at all use sites. Chi-square goodness-of-fit tests were used to evaluate distribution of used vs. unused survey plots among 8 regions of the Gunnison Basin. Use of the 7 terrain categories was compared to availability in survey plots following Neu et al. (1974). Availability of terrain categories was determined from dot-grid acreage estimates obtained from topographic maps. I compared shrub structure parameters at winter use sites to other seasonal habitats (spring use and nest sites) and random sites with analysis of variance procedures. �~ ~ SCALE 0123'5 deB IN E3 MILES f STEUBEN N ~ \J1 Fig. 1. Boundaries of 8 regions in the Gunnison Basin in which systematic ground surveys of sage grouse winter distribution were conducted. �246 Table 1. Categories of slope and aspect used to classify terrain in random1yselected s~ge grouse survey plots, Gunnison Basin, Colorado. Terrain category Description 1. 0-50 slope, high topography Areas of < 50 slope; broad (> 100 m) mesa or ridge tops topographically higher than surrounding terrain. 2. 0-50 slope, low topography Areas of < 50 slope; broad (> 100 m) flood plains and stream drainages adjacent to areas of higher topography; terraces that occur on slopes. 3. Drainages Narrow « 100 m) flood plains of permanent or intermittent streams; shallow, eroded gulches on slopes. 4. 6-150 slope, Slopes between 6 and 150 with primarily north or north or east aspect east aspects.a 5. 6-150 slope, Slopes between 6 and 150 with primarily south south or west aspect or west aspects.b 6. > 150 slope, north or east aspect Slopes of aspects.a 7. 150 slope, south or west aspect South of aspects.b > > > 150 with primarily north or east 150 with primarily south or west aAspects with bearings of 316 to 1350• bAspects with bearings of 136 to 3150• Shrub structure, sagebrush species composition, and snow depth were measured at a sample of winter foraging sites. I used line transects to evaluate shrub structure. Transects were approximately centered in the area of foraging activity. One l5-m transect was oriented in the direction sage grouse primarily traveled while feeding. A second 15-m transect bisected the center �247 of the first line at a perpendicular angle. Sagebrush canopy intercepted by transects was measured to provide a percentage estimate of canopy cover. Height of individual sagebrush plants beneath transects was measured at the tallest living stem. Crown length was measured along the longest crown axis and width was measured along an axis perpendicular to length. Each plant was examined for evidence of sage grouse feeding activity. Browsed or non-browsed status of each plant was recorded along with measures of crown size. At 15 sites, adequate foliage samples were obtained for comparison of nutritional value of browsed vs. non-browsed sagebrush plants. Leafy stems were clipped from each plant. Foliage samples from a single plant were sealed in an air-tight bag. All samples were frozen 2-6 hours after collection. Sagebrush species composition was identified at all sites. Sagebrush density was measured wi thin 0 .5 m of transects • Twelve measures of snow depth were obtained at 3-m intervals along transects. Sagebrush structure and species composition were also measured at sites used by sage grouse in spring (1 Apr-30 May) and at nest sites. Vegetation analysis techniques were similar to those used to measure shrub structure at winter use sites. However I did not attempt to evaluate foraging activity at spring use sites. Measurements of shrub structure and species composition were also obtained at 100 randomly selected sites. Random sites were stratified among the 5 terrain categories with < 150 slope. Twenty random points were selected in each category. Winter Sagebrush Availability Mid-winter sagebrush availability in the Gunnison Basin was evaluated during aerial transects on 7 February. Approximately 416 km of transects were flown via fixed-wing aircraft at approximately 150-200 m elevation. Air speed was 130 km/hr. Transects were oriented north-south and spaced at 3.2-km intervals. An observer on the passenger side of the cabin looked through a 25-cm2 viewing square marked at shoulder level on the window. At 15-second intervals the presence or absence of exposed sagebrush in the viewing square was recorded. This provfded a percentage estimate of exposed sagebrush in each region of the Basin. Sagebrush exposure and snow depths were measured along transects during ground searches in randomly selected 0.79-kmf survey plots. Five measures of sagebrush availability and snow depth were obtained at 50-m intervals along transects in each terrain category present in a survey plot. Sagebrush exposure was based on the presence of leafy foliage within a 1.Q-m radius. t Lek Locations and Surveys Locations of known sage grouse leks were plotted on 7.5 minute topographic maps. Leks were visited during early morning (0430-0700 hours MST) between 1 April and 17 May. Numbers of attending birds were recorded. Searches for previously unlocated leks were conducted from a helicopter during early morning on 15-16 and 29-30 April. I also used a parabolic listening device during ground searches for previously unlocated leks. �248 Sage Grouse Trapping and Marking Night-lighting techniques were used to capture sage grouse on or near leks between 18 March and 13 May (Giesen et al. 1982). In addition, an incubating female was successfully nest-trapped and radiomarked following Smith et al. (1980). Measures of body weight and size were obtained from captured individuals. All birds were marked with colored and aluminum leg bands. Radio transmitters were attached to all captured females and some captured males. Nesting Biology Radio-marked females were monitored during the display season to evaluate distance between probable breeding sites and nest locations, and to obtain information on nesting success and clutch size. Clutch size and fate of incidentally discovered nests were also recorded. Sage Grouse Lipid Reserves Male sage grouse were collected in the Gunnison Basin and Jackson County during April and May 1984 for analysis of lipid reserves. Birds were hand-plucked and the breast muscles on 1 side and internal organs were removed from each carcass and weighed. Intestine and caecal lengths were measured and gut contents were removed and discarded. Organs and muscles were returned to each carcass and the whole body frozen and then ground 6 times in a commercial meat grinder. The ground material was dried at 80 C for 24 hours and then finely ground in a Wiley Mill. Diethyl ether will be continuously dripped through 10-g homogenized samples from each carcass for 6 hours to extract lipids. Additional samples of 10 adult males in each area and each time interval were collected in both Jackson County and the Gunnison Basin during April and May 1985 for lipid analysis. RESULTS AND DISCUSSION Summer and Fall Movements Movements of 4 radio-marked male and 8 radio-marked female sage grouse were monitored between June and December 1984. Male sage grouse remained near (i = 1.8 km, SE = 0.68) leks in June and July. Radio transmitters of all marked males were recovered or had failed by 9 August (Table 2). Radio-marked females were relocated between 7 and 9 August. Distances from nest sites varied (x = 3.8 km, SE = 1.27). There was no difference (P > 0.05) in August distances from nest sites between females that successfullY-hatched young (x = 3.4 km) and females that did not successfully nest (x = 0.81 km). All radio-marked sage grouse remained in sagebrush uplands during the summer. I did not observe use of hay meadows or quaking aspen (Populus tremuloides) stands. The number of radio-marked females followed declined between July and December due to mortality and transmitter failure (Table 2). Three radio-marked females began movements away from summer use areas during November and December. One radio-marked female moved 34 km from her summer use area in the Ohio Creek Valley to a site 2 km southwest of Doyleville. 1 �249 Table 2. Radio-marked sage grouse in the Gunnison Basin, March-December 1984. Radio frequency (MHz) Age Males 01 02 03 04 05 06 07 15 23 57 151.052 151.285 151.000 151.910 150.939 150.968 151.013 151.358 150.980 151.018 Yrlg Yr1g Yr1g Ad Yrlg Yrlg Yr1g Yr1g Yr1g Yr1g 21 10 11 11 13 17 24 30 11 18 Mar Apr Apr Apr Apr Apr Apr Apr May May 01 17 19 24 17 26 23 09 29 10 Apr Jun Apr May Jun May May Aug May Ju1 Predation Tail mo1tb Tail molt Tail molt Unknown Tail molt Unknown Tail molt Tail molt Tail molt Females 5701 5702 5703 5704 5705 5706 5708 5709 5710 5711 5712 5704 150.876 151.131. 150.908 150.863 151.041 150.955 151.343 151.254 151.238 151.159 151.283 151.311 Yrlg Ad Yr1g Ad Ad Ad Ad Ad Yr1g Ad Yr1g Ad 29 10 17 22 22 22 24 01 06 06 07 28 Mar Apr Apr Apr Apr Apr Apr May May May May May 17 08 25 11 22 23 08 20 08 20 27 26 Dec Aug Nov May Apr May Aug Dec Sep Dec Oct Oct Predation Harness breakage Predation Predation Unknown Predation Unknown Predation Hunter harvest Unknown Unknown Unknown Band a Date captured Last relocation or date recovered Cause of recovery aAd = adult, Yr1g = yearling. bTransmitter recovered following breakage or molt of central rectrices. Sage Grouse Harvest Hunters deposited 348 sage grouse wings in wing barrels. An additional 84 wings were gathered at check stations for a total of 432 sage grouse wings collected during 1984. The 1984 sample was smaller than in 1979 and 1983 but exceeded samples from 1980, 1981, and 1982. The decline in wing barrel deposits from the 1983 season could indicate lower harvest due to fewer hunters or poor hunter success. Sex and age of harvested sage grouse were identified from wings (Table 3). Chicks comprised 66.2% of the harvest indicating excellent production of young in 1984. With the exception of 1981, chick production has been good (> 60% juveniles in the harvest) in the Gunnison Basin each year since 1977. This could indicate increasing population size or differential vulnerability between juveniles and older birds. �250 Table 3. 1977-84. Hales Age and sex composition of harvested sage grouse, Gunn;f.sonBasin, Juveniles Females Total s Yearlings Females Males Totals Males N % -T----r -N-%- ~-_--%- N--~C-:N--%--J-i-T Year 1977 67 47.2 1978 151 47.3 75 52.8 142 60.4 18 34.0 35 66.0 53 22.6 15 37.5 25 62.5 17.0· 235 517 38 70.4 54 10.4 56 38.9 88 61.1 144 27.9 66 51.2 129 18.6 41 32.0 87 68.0 128 18.5 692 144 53.1 271 64.5 41 48.8 43 51.2 84 20.0 25 38.5 40 61.5 15.5 420 43 57.3 32 42.7 75 35.9 19 42.2 26 57.8 '45 21.5 209 13 50.0 13 50.0 26 9.7 23 33 37.1 1982 9.4 54.3 79 45.7 89 42.6 173 64.6 46 66.7 69 25.7 268 73 67.6 108 16.0 676 49 66.2 432 64.5 53 40.2 79 59.8 132 19.5 1984 111 39.2 174 60.8 286 66.2 29 40.3 43 16.7 25 33.8 19.2 36.1 1983 205 47.0 231 53.0 436 50.7 60.9 43.6 59.7 56.4 72 65 33.3 35 32.4 49.3 40 16 29.6 56 62.9 avo , Total wings % 63 48.8 1981 S-yr Totals _N_ 62.9 168 52.7 319 61.7 1979 216 49.) 219 50.3 435 1980 127 46.7 Adults Females J:!. % 74 17.1 19.9 63.9 Chick production was similar among different regions of the Basin (Table 4) although a high percentage (83%) of juveniles occurred among harvested birds on Sapinero Mesa (N = 47). However, because of the small sample size this may be the result of-sampling error and may not reflect extremely high chick production on Sapinero Mesa. Table 4. Age composition of sage grouse harvested in different regions of the Gunnison Basin, 1984. Juveniles Resion Doy1evi11e. Ohio Creek Six-mile Lane, Gold Basin, and Willow Creek Sapinero Year1in~s % N. Adults % N % 99 88 68.8 60.2 24 30 16.7 20.5 21 28 14.5 19.2 60 39 63.2 83.0 15 3 15~8 6.4 20 5 21.1 10.6 N Nesting success of females was estimated from primary molt progression. Females that had molted primaries 8, 9, or 10 were considered unsuccessful. Nesting success among yearlings (67.3%, N = 49) and adults (69.7%, N = 43) was similar (Table 5). Overall nesting success among females was 68:5% and was comparable to nesting success in most years since 1977. High nesting success is consistent with the high chick proportions observed in the harvest. �251 Table 5. Estimated Basin, 1977-84. sage grouse nesting success Successful All hens Adults Yearlinss and Juveniles per hen production, Juveniles per ~ successful hen Gunnison Percent juveniles in harves t Year .!! 1977 25/35 68.6 23/25 92.0 47/60 78.3 2.4 3.0 60.4 1978 23/38 60.5 67/88 76.1 90/126 71.4 2.5 3.5 61.7 1979 48/66 72.7 59/87 67.8 107/153 69.9 2.8 4.1 62.9 1980 30/43 69.8 27/40 67.5 57/83 68.7 3.3 4.8 64.5 1981 ' 10/32 31.3 12/26 46.2 22/58 37.9 1.5 4.0 42.6 1982 ,' 6/13 46.2 30/46 65.2 36/59 61.0 2.9 4.8 64.6 1983 40/79 50.6 53/73 72.6 93/152 61.2 2.9 4.7 64.5 1984 , 30/43 69.7 33/49 67.3' 63/92 68.5 3.1 4.5 66.2 % .!! % % .!! Hatching dates of 285 harvested juveniles were estimated from primary growth and replacement. The peak of hatching (77.9% of all chicks) was between 29 June and 12 July (Table 6). This is .approximately 2 weeks later than in 1983. Deep and extensive snow cover during spring 1984 probably delayed breeding. In 1984 female flocks were not observed on strutting grounds until the last week of April. The extensive snow cover may have resulted in synchronized breeding among females. In 1984 a higher percentage of chicks hatched over a shorter time period than in 1983. Hatching dates were consistent among different regions of the Basin although 47% of the harvested chicks hatched prior to 29 June on Sapinero Mesa. Table 6. Hatching dates of juvenile sage grouse harvested Basin, 1983-84. 1983 Period ~ 01-07 Jun 08-14 Jun 15-21 Jun 22-28 Jun 29 Jun-05 Jul 06-12 Jul 13-19 Jul 20-26 Jul in the Gunnison 1984 N % N 5 33 156 145 65 22 8 2 1.1 7.6 35.8 33.3 14.9 5.0 1.8 0.5 5 33 151 71 20 5 % 1.8 11.6 53.0 24.9 7.0 1.8 �252 Sage Grouse Winter Distribution Sage grouse use sites were observed in 51 (35.7%) of 143 randomly-selected survey plots in 7 of the 8 regions in the Basin (Fig. 2, Table 7). The distribution of used plots did not differ from an expected distribution that would result if sage grouse flocks randomly occupied the Gunnison Basin (P > 0.05, Table 7). Therefore, strong preference for one or more regions of the Basin was not apparent. Sage grouse were more widely scattered during winter 1985 than during winter 1984. I observed a greater number of use sites in the Doy1evi11e, Tomichi, and Sapinero regions than during winter 1984. Table 7. Observations of sage grouse use in randomly-selected survey plots in the Gunnison Basin, 3 January-15 March 1985. ! plots with use sites Expected number of plots with use sites 11 20 10 10 17 27 27 21 6 4 2 0 6 11 11 11 4 143 51 N Region random plots Tomichi Parlin McIntosh Stueben Sapinero Beaver Chance Doy1evi11e Totals x2 = 0.79-km2 8.82 with 7 d.f., P > 7 4 4 6 10 10 7 51 0.05. aRegion boundaries are delineated in Fig. 1. Winter and Spring Habitat Use A total of 74 sage grouse feeding sites was observed in randomly-selected plots between 3 January and 15 March. Use sites were not randomly distributed among terrain types (Table 8). Drainages and topographically low sites of <50 slope were preferred. Sixty percent of feeding site observations were in these 2 terrain categories although they comprised only 21% of the area available. Topographically high sites of <50 slope and south or west slopes of 6-150 were also used (37.8% of feeding sites) although use was proportional to availability of these categories (Table 8). I observed little use of north or east slopes of 6-150, or slopes of > 150 (2.8% of feeding sites). Frequent use of sagebrush stands in drainages was also observed during winter 1984 (Hupp 1984). �-- ~ A\ o survey plot • SCALE OI23'~ survey plot with use site HUH IN MILES "........., F9 '" • • 0 • 0 • o o N Ln W Fig. 2. Distribution of sage grouse winter flock survey plots in the Gunnison Basin. �254 Table 8. Distribution of sage grouse feeding sites among terrain categories in the Gunnison Basin, 3 January-15 March 1985. Proportions of observed and expected observations are in parenthesis. Confidence intervals (95%) of observed proportions were calculated following Neu et a1. (1974). Failure of the expected proportion to fall within confidence limits indicates that observed use differed < 0.05) from expected use. (K Terrain category 0-50, low topography 0-50, high topography Drainage 6-150, south or west aspect 6-150, north or east aspect > 150, south or west aspect > 150, north or east aspect Observed use 95% CI E!j2ected use 27 (0.365) 12 (0.162) 17 (0.230) 0.214<P<0.516a 0.047<P<0.277 0.098<P<0.362a 12.4 (0.167) 9.1 (0.123) 3.4 (0.047) 16 (0.216) 0.087<P<0.345 21.2 (0.286) 1 (0.014) 0.0 < P<0.051b 17.1 (0.231) 1 (0.014) 0.0 P<0.051b 4.2 (0.058) _b 6.5 (0.088) 0 (0) aobserved use was greater than expected use and indicates preference for terrain category. bObserved use was less than expected use. sage grouse Snow depth, shrub structure, and species composition were measured at 38 winter feeding sites. Grouse primarily foraged in stands of mountain big sagebrush (!. .!. vaseyana). Black sage (!. ~) occurred at only 2 (5.3%) feeding sites where shrub structure and species composition was measured. Black sage was available at 24% of the randomly selected sites. Shrub structure was also measured at 24 sites used by grouse in spring and at 2 nest sites. Comparisons of shrub structure among seasonal use sites and random locations will be presented in the next progress report. Winter Sagebrush Availability Snow depth in February 1985 was approximately 60% of depths observed in (Table 9). As a result, sagebrush exposure in 1985 was much greater during the previous winter (Table 10). Aerial surveys indicated sagebrush was available across a much larger area (84.8% of the Basin) in than in 1984 (6.7%, Table 11). 1984 than that 1985 �255 Table 9. Mean snow depths (em) in 7 terrain categories in the Gunnison Basin. Snow depths in 1984 are from randomly-located transects in Chance and in Graf1in gulches. Snow depths in 1985 are from ground transects randomly-located 0.79-km2 survey plots. 1984 Februa!1 % N Terrain category 0-50, low topography 0-50, high topography Drainages 6-150, south-west aspect 6-150, north-east aspect > 150, south-west aspect > 150, north-east aspect } 1985 Februa!1 JaIlua!1 % N % March 1i % N 108 17.5 18.2 21.0 62 62 62 34.1 28.3 39.0 134 140 244 31.6 25.0 29.4 114 85 149 39.1 56 9.2 91 18.4 230 9.3 150 57.3 132 23.6 92 37.3 229 37.0 165 c 3.9 57 11.0 110 5.7 70 c 26.2 34 34.6 70 35.5 49 53.5 176 65.0 aTerrain categories in Table 1. ~o distinction was made between areas of high topography vs. topography in 1984. cNo measures of snow depth were obtained on slopes > 150 in 1984. low Table 10. Percentage of ground transect survey points where exposed sagebrush foliage was available above snow cover. Sagebrush exposure in 1984 was estimated from randomly-located transects in Chance and Graf1in gulches. Exposure in 1985 was estimated from transects in randomly-lo.cated 0.79-km2 plots. 1984 Februa!1 % N Terrain category 0-50, low topography } 0-50, high topography Drainage 6-150, south-west aspect 6-150, north-east aspect > 150, south-west aspect > 150, north-east aspect 1985 February % N. -JanuarI % N March % N 108 75.0 80.9 88.6 32 42 44 91.8 50.7 90.2 134 140 244 88.6 77 .5 96.7 114 80 150 8.9 56 94.3 70 90.0 230 92.0 150 3.0 132 80.6 62 71.7 230 80.6 165 c 94.6 37 71.8 110 90.0 70 c 77 .8 19 57.1 70 70.0 50 9.7 176 16.7 aTerrain categories are in Table 1. ~o distinction was made between areas of high vs. low topography in 1984. cNo measures slopes in 1984. of sagebrush exposure were obtained on steep (> 150.) �256 Table 11. Mid-winter availability of sagebrush in 8 regions of the Gunnison Basin, 1984 and 1985. Availability was estimated during aerial transects. Exposed sagebrush in mid-winter(%) Region Tomichi Parlin McIntosh Steuben Sapinero Beaver Chance Doyleville Average 1984 1985 6.7 7.6 81.0 84.1 84.2 72.5 86.3 95.4 84.7 81.0 6.7 83.7 11.8 3.5 o 7.9 11.7 11.8 aRegions as identified in Fig. 1. Lek Locations and Attendance Four leks were observed during 1984 aerial searches but were not confirmed prior to termination of the display season. In 1985, strutting activity at 2 of these leks (Kezar and Sugar Creek) (Fig. 3) was confirmed during ground surveillance. However strutting activity was not observed in the vicinity of the other 2 unconfirmed strutting grounds (Beaver and Alkali). During helicopter surveys in 1985, displaying sage grouse were observed at 5 sites where leks were not previously known to occur (Fig. 3). In addition I heard displaying sage grouse near Roundup Basin (Fig. 3) but was unable to visually locate the lek before the courtship season terminated. Sage grouse attendance was surveyed at 17 leks in 1985. Mean peak attendance of males (25.5 males/lek) was slightly higher than in 1984 (Table 12). Female attendance at leks primarily occurred during the 2nd week of April. This is earlier than the period of female attendance in 1984. Sage Grouse Trapping and Marking Fifty-two (49 males, 3 females) sage grouse were captured and marked between 18 March and 13 May 1985 in the Gunnison Basin. In addition 3 males marked in 1984 were recaptured in 1985. Radio transmitters were attached to 3 female and 8 male sage grouse. Body weights of adult and yearling male sage grouse captured in 1985 did not differ (~ > 0.05) from body weights of males captured in 1984 (Table 13). �~ ____,.,.. .~ • confirmed ~ A unconfirmed SCALE OI23.~ IN MILES IflIl{"-----.--F9 !" & ugarCr I, I I, I N Vl '-l Fig. 3. Locations of saee grouse leks discovered in the Gunnison Basin during 1984-85. Unconfirmed leks were observed during aerial surveys and ground surveillence, but are sites where display activity has been observed for < 2 consecutive springs. �258 Table 12. Trends in peak 1ek attendance of male sage grouse in the Gunnison Basin 1980-85. Lek 1980 Gold Basin Chance Gulch Woods Gulch Needle Creek Parlinb Allen Ohio Creek Sapineroc Razor Creek Antelope Blue Mesa Upper Six-Mile McCabe Signal Peak Lost Canyon Mashburn 1 Haahbu'rn 2 Kezar Basin Sugar Creek 1981 NCa 28 10 27 72 16 53 60 48 63 2 ° 15 7 36 43 46 112 68 43 °° 33.6 x Peak male attendance 1982 1983 ° 21 15 18 47 27 38 NC 39 52 NC 7 37.9 26.4 1984 NC 38 8 14 88 51 82 NC NC 62 NC 10 5 11 5 7 4 5 9 66 37 18 17 19 53 NC NC NC 3 NC 22 43 34.0 23.3 1985 ° 53 2 9 94 33 32 41 12 46 NC NC ° 4 12 54 25 12 5 25.5 aNC = No count. blnc1udes North, South, and Upper South Parlin leks. clnc1udes Sapinero 1, 2, and 3 leks. Table 13. Mean body weights (g) of sage grouse Basin, Apri1-r~y 1984 and 1985. Age Adult Yearling Adult Yearling Sex Male Male Female Female N 42 18 9 4 1984 x 2,038 1,786 1,210 1,084 captured SE N 16.5 27.6 23.2 38.4 43 9 2 in the Gunnison 1985 x 2,085 1,761 1,196 SE 20.4 32.3 44.0 ~ ~ �259 Nesting Biology Two sage grouse nests were located during the 1985 nesting season. Clutch size of 1 completed nest was 9 eggs. Both nests were destroyed by mammalian predators. Nest sites were 5.2 and 0.1 km from probable breeding sites. Nesting biology data from 1984 have been previously reported (Hupp 1984). Sage Grouse Lipid Reserves Body weights of adult male sage grouse collected in the Gunnison Basin in 1984 were 27-38% lighter than weights of adult males collected in Jackson County during similar periods (Table 14). This is consistent with earlier observations that Gunnison Basin sage grouse are physically smaller than birds in other Colorado sage grouse populations (Hupp 1984). Among Gunnison Basin sage grouse, there was a slight decrease in body, muscle, and internal organ weights between early (11 Apr-Ol May) and late (18-21 May) periods of the display season. Body, organ, and muscle weight differences among time periods and regions will be evaluated with analysis of variance and multiple range tests. Relative gut lengths of Gunnison Basin sage grouse were greater than relative gut lengths in Jackson County (Table 14). Relative gut lengths were calculated by dividing combined caecal and small intestine lengths by a corrected body weight. Corrected body weight was live weight minus feather, head, tarsal, carpal, and ingestia weights. The difference in relative gut lengths may mean that digestive efficiencies vary between the 2 populations. Necropsies of sage grouse collected in 1985 have not been completed. �260 Table 14. Co.mparative gut mo.rpho.lo.gy,bo.dy, breast muscle, and internal o.rgan weights o.f adult male sage gro.use co.llected in the Gunniso.n Basin and Jackso.n Co.unty, April-May 1984. Gunniso.n Basin Weights (g) Live weight ~. Eecto.ralis ~. sUEraco.reco.ideus Gizzard Heart Testes (co.mbined) % Lipids ~ ~ ~ ~ 11 Apr-l May (N=lO) 18-21 May (N=lO) 26 Apr-3 May (N=lO) 17-21 May (N=lO) 2,104 173 44.1 22.9 24.2 1.69 2,037 170 41.5 19.7 23.2 2.04 2.4 Gut mo.rpho.lo.gy(mm) Small intestine Caeca (co.mbined) Relative small intestine lengtha Relative caecal lengthb Jackso.n Co.unt~ 1,580 1,727 1.00 1.09 0.8 1,520 1,609 0.96 1.01 2,916 244 54.3 39.2 35.0 2.74 2,854 250 57.7 41.5 34.4 2.86 2.3 1,821 2,090 0.8 1,838 2,064 0.80 0.91 0.77 0.87 aSmall intestine length f co.rrected body weight. Co.rrected body weight is live weight minus feather, tarsal, carpal, and ingestia weights. bCo.mbined caecal lengths f co.rrected bo.dy weight. LITERATURE CITED Beck, T. D. I., R. B. Gill, and C. E. Braun. 1975. Sex and age determinatio.n o.f sage gro.use fro.m wing characteristics. Co.lo.rado. Div. Wildl. Game Inf. Leafl. 49 (revised). 4pp. Giesen, K. M., T. J. Scho.enberg, and C. E. Braun. 1982. Metho.ds fo.r trapping sage gro.use in Co.lo.rado..Wildl. So.c. Bull. 10:224-231. Hunter, W. R., and C. F. Spears. 1975. So.il survey o.f the Gunniso.n area, Co.lo.rado..U.S. Go.vt. Print. Off., Washingto.n, D.C. 85pp. Hupp, J. 1984. Sage gro.use distributio.n and habitat use in the Gunniso.n Basin. Jo.b Pro.g. Rep., Co.lo.rado.Div. Wildl. Res. Rep. April 1984. 107-128. Pp. Neu, C. W., C. R. Byers, and J. M. Peek. 1974. A technique for analysis o.f utilizatio.n-availability data. J. Wild1. Manage. 38:541-545. �261 Smith, L. M., J. W. Hupp, and J. T. Ratti. 1980. Reducing abandonment of nest-trapped gray partridge with methoxyflurane. J. Wildl. Manage. 44 :690-691. Prepared by ~ W J.LI~_ JrnytW OH~pp7r!!£ Graduate Research Assistant Approved by t/J r~ Clait E. Braun Wildlife Research Leader �263 JOB FINAL REPORT Colorado State Project 01-03-045 (W-37-R) Work Plan _..;..__ 9 Job Title: Avian Research Job _..;..__ 7 Winter Habitat Preferences and Migration Patterns of Blue Grouse in Middle Park, Colorado Period Covered: 1 January 1981 to 31 December 1983 Author: Brian S. Cade Personnel: C. Braun, M. Crosby, J. J. Jeanson, R. Hoffman, S. Steinert, Colorado Division of Wi1d1ifej A. Cade, B. Cade, K. Medve, B. Piske, R. Ryder, Colorado State University. ABSTRACT Winter habitat preferences and migration patterns of 2 populations of blue grouse (Dendragapus obscurus) were studied in Middle Park, Colorado from April 1981 through November 1983. Radiotelemetry was used to follow adult and yearling grouse between breeding and wintering areas and to locate winter-use sites. . Distances between breeding and wintering areas (N = 30) ranged from 0.1 to 29.5 km with elevation increases of 0 to 760 m. - There were short and long distance migrants in each population. Females moved shorter distances than males (P = 0.002). Directional orientation between breeding and wintering areas was non-random for grouse within breeding populations, with greater variation among those migrating short « 3.0 km) distances than among those migrating long distances. Males left the breeding range 5-7 weeks earlier than females, but both sexes arrived on wintering areas at approximately the same time between early October and mid-November. Blue grouse exhibited fidelity to both breeding and wintering areas. Winter home ranges of 3 juvenile grouse (median = 18.7 ha) were larger (p < 0.001) than home ranges of 10 adult grouse (median = 3.0 ha). During winter, grouse were observed alone, in small flocks of 2-6 birds, and in occasional large flocks of 12-20 birds. Median flock size was 3 birds at Green Mountain and 4 birds at Whiteley Peak. There was a tendency to observe more females, 59% at Green Mountain and 66% at Whiteley Peak, than males at wintering sites on the study areas, but the null hypothesis of a 1:1 sex ratio in flocks was not rejected. �264 Blue grouse occupied a broad structural range of conifer stands during winter including dense (900+ trees/ha) 2nd-growth (50-70 years old) and mature forests (100-200 years old) to open « 100 trees/ha) old-growth forests (200-600 years old). Douglas-fir (Pseudotsuga menziesii) was the preferred species on both study areas, but grouse that wintered off the study areas used lodgepole pine (Pinus contorta) and spruce-fir (Picea engelmannii/Abies lasiocarpa) forests. No preferences for topographic features (elevation, aspect, slope) were detected at winter-use sites. Blue grouse consistently used the largest conifers available within occupied stands. Preference for large conifers was hypothesized to be correlated with food selection. �265 INTRODUCTION Demand for timber resources has generated life and land manag ernen t agencies tions on animal populations. may have first upon species understand 1981) . habitats period abundance (Fretwell species 1972). for blue grouse, Pacific coastal southern occupy ranges Alaska) from mature Great types Bendell 1968; Martinka deciduous 1972; Harju and coniferous States and (including coastal forest forests to and shrub 1960; Blackford 1963; 1968; Zwickel et al. 1974; Weber 1975; Bendell forests true in habrtat s ranging mesic Pacific 1954; Mussehl 1983). require- 1963). Rocky Mountain are restricted 1903, Marshall habitat This is especially 1966, 1967; Boag 1966; Rogers but blue grouse (Anthony (Aldrich regenerating, 1978; Lewis 1979; Stauffer and breeding may be the critical understood. United to (Wiens and Rotenberry from sea level to treeline 1946: Caswell and El'Iott alterations it is necessary wintering winter of the western Basin and southern (Marshall habitats perturba- endemic to the Rocky Moun tarns and Canada or young habitat (Lack 1954), winter are poorly breed the effects different and although a species Blue grouse fall, use their among wild- the impact of forest and distribution, for most avian populations ments of many species xeric To predict how species Many avian about concern and Zwickel A variety of shrub types may be used during spring through to conifer forests during winter 1946, Bendcll 1955, Hoffmann and 1956, Bendell �266 and Zwickel 1978. Stauffer almost exclusive (Beer winter 1943. Stewart Habitat ing areas focused variety grand 1944. Hoffmann have heen to available fir adequately habitat. during (A. grandis), (Pinus Caswell use of large conifers suggesting that determine habitat is an important (Cody (1954), factor for winter although pine than selection rear- have in relation blue grouse white or dense rather in habitat have (Mussehl been reported 1960, Bendell some may migrate use a fir (Abies concolor-) , Douglas-fir, (Caswell 1954, Hoff- 1974, Stauffer (1983) all reported clumps of conifers, species composition grouse. Habitat for other avian tween breeding and wintering move up, habitats (Hoffmann down, using band 1967, Zwickel et al. wintering areas and Elliott (Rogers 1956). areas because to higher may structure species elevations 1967, Zwickel et al. 1968) or reside The spatial 1972). Indirect recoveries spring most recoveries be- why blue and winter efforts to understand (Mussehl 1960, Bendell 1968) failed to document 1968) at the relationship may be the main reason or not at all between (Zwickel and Bendell relationship to migrate downwards round Elliott particularly (A. lasiocarpa), for wintering same area year this few studies and Stauffer in open forests suitability and brood 1968, King 1971, Harju structure of conifers 1981). Blue grouse grouse fir King (1971), forest but including and lodgepole mann 1956, Boag and Kiceniuk 1983) . on breeding winter. subalpine flexilis), and needles It is known that winter to their 1961. King 1968). described. use during coniferous limber pine twigs. of blue grouse habitat of conifers This can be attributed diet of buds. preferences on their 1983}. movements were from birds shot and to during �267 fall. Band recoveries juveniles also were biased and did not reflect between sexes documented or age-classes for other (Seiskari et al. Sex segregation gested by several (Skinner 1954. King 1971) and could result or habitat grouse preferences are dispersal than rather frequently 1962. Hale and Dorney during and Hammer- winter 1927. Marshall migration seasonal patterns Movements of juvenile movements of adults than has been sug- 1946. Caswell from differential for males and females. may be greater ments a behavior 1975. Beck 1977. Herzog and Keppie of blue grouse authors females and movement patterns 1967. Hammerstrom 1973• Hoffman and Braun 1980) . adult differential of blue grouse. tetraonids 1963. Weeden 1964. Irving strom possible towards if first-year migration (Baker move- 1978. Green- wood 1980). Fidelity torial to breeding male blue grouse others). a factor (Bendell Fidelity in migration The principal t ur and Elliott habitat objective Emphasis of this use for blue grouse features availability) study (Jamieson is unknown habitat occupancy. by quantifying attributes and but could be was to identify at winter-use was placed on structural (use vs. areas and winter for terri- 1967. Lewis 1979. among for females to wintering patterns al , and topographic ences has been well documented but is less well established Zwickel 1983~). winter areas patterns floristic. struc- and random sites. associated both among and within with prefer- conifer stands . . ' The theoretical study is that proximate basis for habitat measured factors that habitat sampling procedures characteristics elicit habitat selection used in this are correlated (Rotenberry with 1981). of �268 Identifying patterns of habitat determining the underlying with habitat selection. movement patterns areas, occupancy process Additional of blue grouse determine winter of blue grouse Hypotheses included: blue grouse structural tested winter-use sites characteristics occupied conifer use sites do not differ (ultimate objectives between home range and age composition sizes, do not differ adult and juvenile breeding associated and wintering and ascertain characteristics characteristics from random sites, (4) winter (5) migration (6) winter do not differ tion of males and females on specific in size, wintering sites, do not differ (3) topographic grouse the sex areas. from random sites do not differ, for were to investigate (1) structural by males and females do not differ, and female blue grouse factors) on wintering at winter-use stands, is a prerequisite at (Z) among at winterhabitats patterns used of male home ranges of and (7) the propor- areas do not differ. • �269 STUDY AREAS Research was conducted In Middle Park (Fig. km west of Denver. Middle Park at Green Mountain 1) in northcentral Unlike other is mountainous and Whiteley Peak Colorado intermountain and locally heavily tain and Whiteley Peak are distinct forested approximately parks 161 in Colorado, forested; mountains Green within Moun- the park. Open areas brush (Artemisia consist and quaking occurs cities. Records - 8.1 C in January forests dominated 2). serviceberry antelope to 16.8 C in July areas area (including tremuloides) (Amelanchier (Purshia is continental forests. and wind veloa total temperature mean annual extremes encompassed 332 ha; of 1980). conifer 41% of the area, aspen with scattered conifers) shrub in non-forested (Symphoricarpos rabbitbrush tridentata), elevation with seasonal Commerce areas by sage- 1970, Carpen- (U. S. Dep. with snowberry spp.), et al. Dam indicate cover is the dominant in association bitterbrush study by Douglas-fir Sagebrush and occurs above this precipitation, and mean monthly ern Mountain and non-forested (Fig. in temperature, from Green Mountain of 40 The Green (Populus Climate in Middle Park fluctuations precipitation aspen vegetation from 3,350 to 3,500 m (Gilbert et al , 1979). and daily below 2,700 m are dominated spp , ) communities; of conifer Timberline ter in Middle Park 2%, 57% areas spp.), (Chrysothamnus spp.), and common chokecherry �270 N t • DENVER 12.8 COLORADO Fig. 1. Green Middle Park. Mountain Colorado. and Whiteley Peak study areas, Km �271 o NON-FORESTED 188 ha (57% D ASPEN [IIJ) DOUGLAS-FIR 50 ha (15%) ~ DOUGLAS-FIR/ASPEN 27 ha (8%) 6 ha (2%) 11m DOUGLAS-FIR/JUNIPER 61 ha (18%) r= o Fig. 2. Colorado. Habitat types at Green Mountain, 1 Km Middle Park, �272 (Prunus virginiana). elevation ranges and west. Maximum relief from 2.390 to 2.863 rn , Cliffs, rocky the west face and occur ledges. portion 1880 and 1890. and the east Present aspects slopes extent are east are prominent on all aspects. of Green Mountain was burned activity area is 470 m; Principal and steep to a lesser northern 1930's. on the study The and logged face was selectively logged on between during on the area is limited to moderate the grazing by cattle. Vegetative prised (Fig. cover on the of 27% conifer 3). mixtures; Conifer forests and associations mann spruce blue spruce non-forested forests. include areas rock outcrops. in shrub Terrain and talus marily in the non-forested pine. Engel- limber pine. is the dominant associations occurring of grazing by cattle to heavy and aspen fir. types. and shrub in similar to those the summit is rugged slopes types fir /aspen is 575 m with elevations near to moderate Douglas with subalpine lodgepole Maximum relief This area has a long history use is restricted Douglas-fir; Sagebrush and occurs area is com- and 50% non-forested of Douglas-fir (Picea pungens). from 2.500 to 3.075 rn . sent 23% aspen. (Picea engelmannii). on Green Mountain. cliffs. 503-ha Whiteley Peak study ranging with on all aspects. and sheep. grazing by cattle. Prepri- �D (:}) NON-FORESTED 253 ha (50%) ASPEN 116 ha (23%) [I] DOUGLAS-FIR/LIMBER I~5j MIXED CONIFER PINE 14 ha (3%) 39 ha (8%) • MIXED CONIFER/ASPEN 51 ha (10%) ~ DOUGLAS-FIR/ASPEN 30 ha (6%) N t -- o ~ 1 Km N --..J W Fig. 3. Habitat types at Whiteley Peak. Middle Park. Colorado. �274 METHODS Blue grouse recorded dogs, calls were located (Stirling captured and banded unique combinations fied as territorial known attempt females, (hatched to November 1983. the breeding Ninety-four Mercator percent to the nearest (UTM) grid (lSD-lSI Mhz) placed life Materials, Inc.; 1980); mounted Carbondale, Inc.; Ill.) Mesa, Ariz.). harnesses weights for battery-powered (no (l08 from April 1981 occurred during Capture 50 m as Universal and 11 in 1983 were either with backpack sites grouse (Apr-Aug). 24 in 1982-83. (Telonics, and loca- Transverse coordinates. Radio transmitters units 1967), or non-breeding of the captures periods Sex and unsuccessful and seventy-six on the 2 study and brood-rearing tions were determined 1 egg), any eggs), and Males were classi- and Elliott at least One hundred 68 males) were banded (1971). (Bendell but failed to hatch to nest). {Gullion (l965). followed Braun nesters pointing aluminum bands bands or non-territorial females as successful tape- noose pole (Zwickel and Bendell of color-coded of grouse and summer using 1966) and lor trained with serially-numbered age classification (nested, spring and Bendell with a telescoping 1967), nesters during (Brander were units. on 23 grouse in 1981-82, solar capacitor-assisted (Wild- or lithium battery-powered Transmitters 1968) or poncho were mounted collars 15-27 g for solar-powered (Amstrup and 29-34 g �275 Radio-marked grouse while they remained the study areas were located on the study were located areas. moved and accessibility. radiotracking was from the ground 15 January grouse Aerial searches 1981 and of marked and recorded to the 50 m as UTM coordinates. areas of individual habitats during were defined areas as November through grouse were defined this period. as either Breeding male territories where movements of breeding spring (Lewis 1979, Hannon et al. non-breeding and Elliott and areas females may wander 1967, Hannon et al. they spring occupied between grouse were measured nearest UTM locations minimum elevational tion changes between require from locations areas 1982). in conifer of individual or female nest grouse and wintering grouse locations, i. e. , were localized during Non-territorial males and widely on breeding ranges (Bendell 1982, Jamieson and Zwickel 1983~), during May through breeding June were defined at the different breeding and Mielke 1983). ranked data, areas as the minimum straight-line difference between and wintering between areas: as of individual distance elevation areas. and wintering males and females using (Berry March, range. Distances dure 3-element were made on 3 November Locations upon method of a hand-held, by visual observation Winter was defined their depending The principal using 2 weeks that moved off intervals 1983 to locate missing birds. were determined nearest Grouse at irregular the distance yagi antenna. at least once every changes Distances areas the multiple response were and eleva- were compared a nonpar arnetr-ic permutation Unlike other between nonparametr-ic permutation procetests procedure that �276 (MRPP) can use continuous tics. Non-random breeding directional to wintering 1974). measurement or'ierrtation areas) first-year migration grouse movements (Greenwood (compass was measured Movements of juvenile because data to calculate direction with Rayleigh's were excluded may be dispersal 1980, Herzog test from test (Zar from analyses rather and Keppie statis- than seasonal 1980, Jamieson and Zwickel 19832), Throughout by relocating sign stands against and tracks) the study birds counted. birds 2 or more birds within located, individual radio-marked (droppings considered value for leg bands, using Fisher's (1957). All conifer searched. classified Once to sex, and was compared binomial distribution flock observations for Flocks were in flocks of grouse exact were found searching themselves. were periodically sex ratio for individual using and by concentrated following Koskimies were examined a 1: 1 sex ratio and flocks of grouse or the birds areas The observed Probabilities lative winter, probabilities. were pooled into a cumu- method of combining probabilities (Sokal and Rohlf 1969). Winter home ranges 5 or more times during delineated the same winter. by the minimum-area, and the enclosed parametric were measured for grouse Home range convex-polygon area was measured that boundaries method with a planimeter. MRPP was used to compare home ranges were located were (Mohr 1947), The non- of adults and juveniles. Major habitat study areas types (conifer, were delineated aspen, on enlarged and non-forested) aerial photographs on the and the �277 area of each type was determined conifer ture stands (discrete and species species, sured spatial composition) relative tree on enlarged with a planimeter. units of forest were defined densities, occurred occupied in any part Structural, fer stands according Stand to dominant area was mea- and area corrections C.stands on slopes ~24° (an area correction was considered with the same struc- and location. aerial photos Individual by grouse ~ 10%) • were applied A conifer if 1 or more grouse to stand observations of the stand. floristic, and topographic were sampled with randomly characteristics located of all coni- 0.01- ha circular plots. The location of a random plot was determined by walking number of paces (1- 36 by 10° increments) from an initial puter plots in a random direction grid location. generated sections (5-20) random number on an overlay for stands Grid locations S10 ha and 20 plots Peak and 200 in 16 conifer by stands of forest were observed Grouse plots centered (Stauffer describing forest the following variables measured for tree and grouse-use number of saplings of trees >10 ha , A total of stands at Whiteley Stratifying variation winter-use on the trees to be includsites were in which the 1983). variables sites: to grid inter- at Green Mountain. structure. sampled with 0.01- ha circular Sixteen for stands 180 i n 13 conifer stands from com- Sample sizes were 10 allowed within and among-stand ed in the analyses grouse were determined corresponding of the aerial photos. 380 random plots was sampled, plots pairs a random ~7 em dbh , and canopy height structure species were derived in plots < 7 cm dbh, (height at random number of tallest from tree and dbh above �278 the plot). Point-centered each of 4 quadrats measure geneity (8 cardinal slope one-third species, dbh , height, rounded canopy. 1956) provided served (1-15. position of slope, ridge were marked hereafter with colored of spatial hetero- were aspect bottom, (Stauffer elevalower, 1983). Tree and form (conical, were recorded were observed, (CV) for 46-60°), (valley top) age (1982-83 only), deformed) 31-45, in an additional recorded 16-30, tree of variation as an index variables 30 m, and vertical mid. and upper to the nearest coefficients Topographic directions). which grouse spacing; measures 1976). tion to nearest use trees tree quarter (Roth distances (Cottam and Curtis of horizontal point-centered quarter for the actual referred flagging trees in to as use trees. All and serially-numbered al urnin urn tag s . Basal areas dominance centages calculated (Mueller-Dombois (IP) for each tree by summing relative and then from dbh measurements dividing univariate 1974). Importance species a stand were calculated by 3 (Risser between and multivariate species and Ellenberg dominance, ages were compared defined within relative frequency, and Rice 1971). occupied relative density, Importance and unoccupied nonparametric per- percent- stands MRPP tests with (Berry and Mielke 1983). Structural sites. and random variance (~ = variation (ANOVA). sites among conifer was analyzed A variance stands for use trees, with one-way stabilizing analysis use of transformation I ~+ 1) was applied to tree count data prior to ANOVA (Sokal and Rohlf 1969). Nevertheless, comparisons the assumptions still violated some subset of variables of normality (skewness in all and �279 kurtosis) and/or homogeneous Therefore, all vari.ables Wallis test (Dueser parametric tests be rejected between sites with males, Kruskal-Wallis Structural with varimax summarizes sites (principal as gradients To calculate factor at these habitat sites defined as points among conifer Principal multivariate analysis component analysis data set as fewer vari- structure of variables (Rotenberry and for each random site from the principal for individual of the new combinations of habitat stands component combinations were calculated stands component were plotted against axes. scores for grouse-use were multiplied component and Wiens 1981). preferences gradients 1975). coefficients from the principal Rotenberry ceived scores component and ANOVA and non- to a principal These and mean scores variables 0.05) (~< 1981). on scoring the principal differed (Johnson (case) and both sexes parametric are linear Factor derived that in a large Wiens 1981). analysis, ·were compared that can be interpreted based characteristics components) variables should ANOVA are using (Nie et al. information of the non- null hypotheses females, were subjected rotation Results tests. variables for the random original conifers Kruskal- of the parametric Structural use and random with a nonparametric whether but results ~< 0.05). test, 1978, Noon 1981). to decide for comparison. parametric ables and Shugart were used among grouse-use (Bartlett's were reanalyzed or accepted, presented variances analysis coefficient s on random sites factor scores space, Since factor the nonparametric (cf. map the onto the multidimensional by the random sites. in Euclidean standardized by the scoring The resulting of the grouse sites, scores habitat are per- MRPP (Berry �280 and Mielke 1983) was used factor scores putes a test at grouse-use statistic from a permutation distances in this data scores case) set. metric (Berry an ANOVA . Program on the average a priori, disjoint from an approximation and Mielke 1983). between MRPP com- between distances of no difference point (Euclidean subgroups of the in clustering of with the Pearson A more conventional to compare mean scores at grouse-use The nonparametric position Comparison distances type para- at grouse-use sites. Elevations vertical sites. of all possible within ANOVA also was used and random based Probabilities were determined III distribution and random (delta) distances given to compare Euclidean MRPP test and slope categories of aspect MRPP analysis and random categories (Mielke 1984). sites were compared with was used to compare at grouse-use and random was done with a circular sites. arc-distance �281 RESULTS Movements from Breeding Movements between breeding and wintering for 30 radio-marked adult tracked 13 in 1982-83, in 1981-82, and yearling grouse females) from statistical a greater probability on the study areas Distance. wintering that Green occurred birds (Table transmitters wintering areas. female (hunter 3 from 1981 to 1983. (1 male, 4 because there that remained grouse moved off the study (Table between of short was areas. breeding and to long distances 2). grouse Five banded moved 0.4 to 2.3 km between bands kill) and an adult at male (unknown grouse breeding were recovered 10.4 and 20.0 krn , respectively. distances 11.1 krn , N = between breeding males (MRPP. ~ = 0.306) between between were withand from an adult cause of death) from their areas. Because differ 8 birds 1) and from 0.3 to 29.5 km for 17 radio- In addition, moved at least breeding that on a continuum at Whiteley Peak out radio that grouse; analyses banded were obtained from 0.1 to 11. 8 km for 13 radio-marked Mountain marked those areas transmitters - Movements of blue grouse areas ranged without of observing than blue Areas 6 in 1983, and Movements of 5 banded were excluded to Wintering 4) and Whiteley Peak females (MRPP, P = 0.468) and wintering ar e as dio not at Green Mountain (median = 10.6 krn , N at Green Mountain (median = 7) or {median = = �282 Table Distance, 1. and wintering breeding grouse, elevation Green areas Mountain, gain, and directional of banded Middle Park, and radio-marked Colorado, Breeding Distance (km) Breeding status Band Sex Age 184 Ma Ad Terri torial 244 M Ad Territorial 245 Mb Ad 374 M Ad 375 M Yrlg 162 F Ad 196 F Yrlg 266 F 299 orientation blue 1981-83. to wintering Elevation gain (m) 2.3 between area Direction 90 99 11. 8 600 199 Territorial 1l.5 240 162 Territorial 10.6 600 200 Non-territorial 7.0 480 31 Successful 0.1 0 180 Non-breeder 8.7 480 204 Ad Successful 6.7 330 184 F Ad Unsuccessful 0.1 0 180 306 F Ad Successful 0.5 120 103 313 F Yrlg Successful 0.1 30 153 317 F Ad Successful 1.2 240 277 359 F Ad Successful 0.3 0 338 379 F Ad Successful 5.6 120 184 aBanded bWintered grouse without a radio in same location transmitter. in consecutive years. (0) �283 Table 2. Distance, breeding Whiteley elevation and wintering Peak, areas Middle Park, gain, and directional of banded Colorado, and radio-marked Sex Age Breeding status between blue grouse, 1981-83. Breeding Band orientation Distance (km) to wintering area Elevation gain (m) Direction 570 . 89 202 M Ad Terri torial 241 Mb Ad Territorial 7.6 480 77 242 M Ad Territorial 17.9 600 75 305 Mb Ad Territorial 1.0 330 346 310 M Ad Terri torial 10.6 450 81 312 M Ad Terri torial 29.5 420 89 366 M Ad Territorial 2.8 450 43 201 F Ad Successful 0.4 90 81 224 F Yrlg Non - breeder 1.9 390 265 225 Successful 0.3 120 270 265 F Ad Fa,b Ad Successful 0.4 120 56 267 F Ad Successful 28.0 760 83 304 Fa Ad Successful 2.0 o 234 308 Fb Ad Successful 0.5 60 o 309 Fb Yrlg Non-breeder 1.8 60 263 316 F Yrlg Unsuccessful 5.0 450 104 Yrlg Non-breeder 0.7 210 348 a 14.1 322 F 342 Fa Ad Successful 0.7 240 8 344 F Ad Successful 15.1 360 58 367 Fb Yrlg Non-breeder 1.8 210 291 371 F Successful 0.9 360 87 Ad aBanded bWintered grouse without a radio-transmitter. in same location in consecutive years. (0) �284 0.5 km , N from both sexes = 9) and Whiteley Peak study (Table distances areas 3). than the number shortest were females. of females movement long distances combined Male blue grouse to test = 9) that moved less (>5.0 km) comparable 10), for differences difference by a male (1. 0 km). = 1. 8 km,.E moved greater Most of this (N· = (median data between !: = (MRPP, 0.002) can be attributed distance Several to than the females moved to distances traveled by most males. Elevation Change. - Elevation = 0.88) with distance (Spearman's!:. areas and differed (Table 3). ranging Radio-marked from the study wintering between area. directional occurring orientation areas 52° (northeast) orientation areas - There between Mountain (Rayleigh's ing short males and females at elevations at elevations at wintering The corresponding Orientation. at Green wintering and wintering elevations of 2,745 - areas away at Green Mountain and 2,500 - 3,380 m for areas. Directional Peak elevations breeding correlated at Whiteley Peak bred 2,380 - 2,780 m for breeding grouse between 0.002) grouse was positively from 2,470 to 2,830 m and wintered 3,320 rn , the highest were !: = (MRPP, change test, breeding (Rayleigh's !: < 0.01) . was 175° (south) for grouse distances between ing long distances was a significant and wintering test, !:< Mean direction for grouse at Whiteley Peak and wintering (>5.0 km) had the least from Whiteley Peak moved east areas from breeding (Fig. Mountain 4). variation areas, Grouse Ears to and migrat- in directional while those All long-distance into the Rabbit for and at Whiteley at Green « 3.0 km) had the greatest breeding 0.05) non-random Range. migratmigrants Grouse �285 Table 3. areas of radio-marked Color ado, Distance male and gain between female blue breeding grouse, Male (N = 11) Middle Park, k Female 19) (MRPP) 0.002 (N = (km) Median 10.6 1.2 x 11. 3 4.2 Max 29.5 28.0 Min 1.0 0.1 Elevation and wintering 1981- 83. Descriptive statistic Distance and elevation gain (m) Median 450 120 x 438 220 Max 600 760 Min 200 o 0.002 �N 00 0'\ N N WI I I I I If (i (~ ))j I I J ~ W ,E I I I I ( (( ('(~ ) )) ) ) ) I I E MALES FEMALES S WHITELEY Fig. 4. Distance radio-marked areas, dashed blue PEAK and directional grouse, arrows S GREEN MOUNTAIN orientation Middle Park, are mean directions. between Colorado, breeding 1981-83. and wintering Dots represent areas wintering for �287 migrating long distances Gore Range northeast with the exception between = Territorial late June Non-territorial adult areas. October males were on winter Distances between summering earliest males. (May), (winter from the breeding hens with broods ally towards remaining between localized areas at the study late September successful) remained range off the study than (1 non-breeding, areas areas breeding during areas. Median departure through areas to winter Two adult areas and then females in early there areas (both August, returned Two yearling and remained summer. and females to females moved to summer locations areas 4). moved gradu- moved rapidly until October, more for females left the August-October, for winter 1 unsuccessful) varied areas from the study in late October. off the study Summering for males (Table and mid-November. areas All for 5 males ranged (Oct). moved 2 and 4 km from the study on the study wintering areas areas. near wintering ?) areas the latest near long distances wintering 4). at summering 4). and unsuccessful for females were 5-7 weeks later Females migrating (Table 1. 8 krn) , Non-breeding Most females remained (Table range and wintering = from 0.9 to 4.5 km (median females than range localized by mid-November habitat areas It took males 1-3 weeks They remained areas Dates of departure breeding when they moved to wintering were in coniferous winter traveled and left the breeding males also left at this time. until early dates male that males abandoned and late July to move to summering areas of a yearling into the into the Williams Fork Mountains. Timing. areas from Green Mountain moved south near through their �N CJ:) CJ:) Table 4. Dcp ar t ur e dates rnar k cd blue grouse. from breeding Middle Park. Departure Sex Year Males 1981 Females Colorado. and arrival dates on wintering areas for radio- 1981-83. from breeding Arrival range Earliest Median Latest 6 26 Jun 07 Ju] 23 Jul 1982 6 24 Jun 18 Jul 1983 2 27 Jun All 14 1981 area Median Latest 4 10 Oct 23 Oct 03 Nov 27 Ju] 4 10 Oct 16 Oct 21 Nov 27 Jun 27 Jun 2 14 Oct 14 Oct 15 Oct 24 Jun 07 Jul 27 Ju] 10 10 Oct 14 Oct 21 Nov 4 25 May 28 Aug 10 Oct 5 11 Oct 21 Oct 26 Oct 1982 6 29 Jun 30 Aug 30 Oct 8 24 Sep 23 Oct 05 Nov 1983 3 20 Jul 14 Aug 15 Aug 4 29 Sep 10 Nov 16 Nov 25 May 15 Aug 30 Oct 17 24 Sep 23 Oct 16 Nov - 13 N - on wintering Earliest All N range �289 October. Females arrived and mid-November Fidelity grouse occupied survived yearling areas range during between the same time as males. - All radio-marked season to the next in consecutive 3 successful late September Areas. from one breeding females returned areas approximately and 1 non-breeding used areas and Wintering adult males, female, yearling specific 4), the same breeding (5 territorial specific (Table to Breeding that at wintering adult females, yearling the preceding years. Ten grouse 1 successful female) returned year. to the Two non-breeding, to the same breeding (1.2 and 1.4 km difference) (N= 12) range, occupied but not the during the pre- vious spring. Of 8 radio-marked adult males, ing areas 2 adult grouse females, both years. between 2 yearling One yearling ing area in 1982-83 than but distance tracked used females) during during 7 (3 the same wintera different winter was only 0.5 km. at the same areas winters, used female occupied as a juvenile the 2 areas females were observed in consecutive winter- 1981-82, Two banded consecutive adult winters. Winter Home Ranges Home ranges grouse (N = located were calculated at least 5 times between 4) and 19&2-83 (N = 9) (Table mates may be poor and variable grouse only were located moved short during for 1 male and 12 female radio-marked distances, winter, reasonable and, 5). November and March 1981-82 Accuracy among grouse because 5-13 times each winter. most <200 m, between therefore, approximations home range of the actual of home range individual However, consecutive estimates magnitude esti- blue grouse locations are probably of home range sizes. �290 5. Table grouse. Winter home ranges Middle Park. (Nov-Mar) of 13 radio-marked blue 1981-83. Colorado. Conifer standsa Band Sex Age Home range size (ha) 305 M Ad 1.9 1 8 162 F Ad 3.1 2 12 201 F Ad 1.8 2 10 266 F Ad 3.0 1 5 299 Fb Yrlg 3.5 2 9 306 F Ad 7.1 2 11 308 F Ad 2.6 1 7 309 F Yrlg 1.6 3 6 317 F Ad 4.4 2 11 359 F Ad 3.3 2 12 299 Fb Juv 18.7 2 7 358 F Juv 42.2 4 13 360 F Juv 9.2 3 10 aNumber of different conifer stands contained range. b Same female but during different winters. N locations within a home �291 Juveniles home ranges (N than stand. restrictive located their observed years movements 3 male territories stands winter birds (7 males. flocks. 25 females. 1 had only males. birds consisted and 15 unknowns) of unidentified of females only. study area. 5). The most of a female and encompassed portions and included parts of sex. 44 flocks Winter of lone of 9 had males and females. of unidentified 3 at Nine flocks sexes. and 2 flocks and (10 males. of flocks. both sex. of 234 grouse of lone birds contained sex. during 47 observations Observations of unidentified Mountain and 44 observations and 39 observations 4 flocks had males and sex. Flock size did not differ either only females. of 25 observations 5 unknowns) of unidentified conifer January on Green 22 had females and birds had females and birds birds of 216 grouse contained Whiteley Peak consisted 10 females. During 1981-82 and 1982-83 included Nine flocks contained between Mountain Except a single were those home at Whiteley Peak. observations (Nov-Mar) 3.0 ha). ha). (Table of blue grouse Pop ulat ion Characteristics Ninety-one (l8.7 within winter at Green = had a smaller winter times in the same tree territories (MRPP. P <0.001) 10. median as a juvenile during Winter home ranges of 7 male breeding = (N moved among several movements 1983. at least than which restricted 6 consecutive March consecutive (3.5 ha) most grouse 18.7 ha) had larger and yearlings during as a yearling for 3 birds = 3. median adults A female tracked range = between winters (MRPP. Median flock size was 3 birds and 4 at Whiteley Peak (Table 6). The proportion !:> 0.100) at Green at Mountain of males and females �N -0 N Table 6. Size and sex composition of winter Flock Green flocks of blue grouse, Middle Park, Sex size Colorado, 1981-83. composi tion a N - Median x - 1981-82 14 3 3.2 2 6 7 11 16 1982- 83 27 3 4.3 2 20 8 11 71 1981-82 19 4 4.4 2 12 21 29 15 1982- 83 20 4 6.0 2 14 21 52 16 Min Max Males Females Uniden tified Mountain Whiteley Peak aMarked grouse that were observed more than once during a winter were only counted 1 time. �293 in flocks did not differ from a 1: 1 sex ratio at Green Mountain (combined exact binomial probabilities, !:> O.900) (combined exact binomial probabilities, P > 0.500) . flocks and number of birds ( ~4), the probability of observing 1: 1 binomial distribution number identified 66% at Whiteley Peak All age-classes tion in each cohort were observed a significant of grouse wintered was not determined and few birds 1 juvenile male, 1 juvenile identified females, male, 9 adult total on the study because winter females, conifer stands ranging were sampled at Green Mountain by blue grouse observations to 27.5 ha, during of grouse I-VIII. Conifer winter. areas. Propor- few marked during birds winter. included Grouse 1 adult females. and 4 juvenile male, Three adult females were stands (Table 8). tions at Whiteley Peak (~ each at Green Mountain of winter Preferences in size from 0.7 to 107.3 ha (Table 7) of which Ninety-four on Green Mountain and 9 of 13 stands winter evidence was a greater at Whiteley Peak. Sixteen during from the and 2 juvenile Winter Habitat stands departure 59%at Green Mountain and were captured at Green Mountain during 5 adult most 6). identified males, There in flocks, (Table Because to sex in flocks were small was minimal. of females identified or Whiteley Peak percent (N = of all winter 100) occurred at Whiteley Peak ranged sampled were occupied Ninety-six = 12 were occupied percent 104) occurred (XIII-XVI) in size from 1. 3 by blue grouse of the winter in stands I-III. and Whiteley Peak use by blue grouse in observaFour stands (X-XIII) and were classified had no as unoccupied. �N \0 ~ Table Stand 7. winter. Green size. basal Mountain. area. and Middle Park. species importance Colorado. percentages (IP) of conifer stands occupied and unoccupied Stand size {hal ~ basal area Species IP 21. 0 (m1/ha) Douglas-fir 88 Subalpine Limber number aThis VII VIII IX X XI XII XIII XIV XV XVI 40.8 5.3 6.5 27.5a 6.8 1.5 2.3 2.6 0.7 5.7 9.5 107.3 10.0 15.8 12.4 l3.1 l3.6 16.5 12.0 10.4 27.7 20.1 19.1 22.1 11.8 0.9 4.8 95 52 82 71 BB 93 n 96 B9 IV 2.9 3.7 33.6 100 B7 100 91 79 9 21 3 pine pine Engelmann Rocky Unoccupied VI III 100 during V II fir Lodgepole Quaking 22.7 grouse 1981-83. Occupied Stand by blue 5 spruce aspen Mtn. 3 12 3 juniper stand extended 48 beyond the study area 15 boundaries; 29 7 25 9 total size 13 4 11 was @ 50 ha. �Table 8. Stand size. during winter. basal area. and species importance percentages Whiteley Peak. Middle Park. Colorado. (IP) of conifer stands occupied and unoccupied by blue grouse 1981-83. Stand number Unoccupied Occupied I II III IV V VI VII VIII IX X XI XII XIII 3.5 19.3 14.4 1.5 3.8 10.7 3.0 6.0 20.8 27.5a 21.6 1.3 13.9 46.7 22.5 24.5 28.7 25.4 10.8 4.1 35.3 24.4 35.5 37.3 35.4 5.2 Douglas-fir 34 55 38 36 89 26 53 21 10 24 68 Subalpine fir 41 39 25 38 45 19 7 Stand size (ha) ~basal area (m2/ha) Species IP Limher pine 9 2 19 Lodgepole pine Engelmann spruce 10 Quaking aspen aThis stand extended 6 43 11 19 100 47 15 19 19 11 15 2 5 10 2 2 15 5 28 beyond the boundaries 54 of the study area; total size was @ 30 2 27 4 42 17 39 60 ha , N \.0 \.J1 �296 Most blue grouse Grouse were observed 9% of the sites roosts observed were in conifers. and in snow roosts at Green Mountain and Whiteley Peak. during fed in the same trees to adjacent conifers roost to feeding trees winter on the snow or ground were encountered often during winter 1982-83. in which they during evening areas. feeding. as noted Many old snow Although roosted. they grouse also moved Movements by Caswell at >100 m from (1954), were not observed. Species stands Composition. - Douglas-fir at Green Mountain. Limber pine. pole pine were present but occurred with quaking in association Rocky Mountain juniper the mountain. Species and unoccupied Species diverse did not differ as 7 stands occurred composition contained in 10 of the stands conifer of at least 13 conifer (MRPP. ~ = stands 0.560) conifer face and with on the west face of were identified. 0.720) between occupied use was equivalent in the random at Whiteley Peak was more predominated pine in 1 stand species in 8 stands. (Table (Douglas-fir). 3 species. Quaking and predominated between stands At Green Mountain blue grouse Douglas-fir = Douglas-fir and lodgepole a single and unoccupied however, on the east (MRPP, ~ and lodge- Douglas-fir of Douglas-fir of conifer had associations was no difference spruce, stands. fir in 4 stands, Only 1 stand aspen 7). scopulorum) stands than at Green Mountain. subalpine occupied (Juniperus in all conifer Engelmann uncommon (Table Only 3 pure composition predominated species 8). where- aspen in 1. composition There in at Whiteley Peak. were observed to availability sample of conifers only in Douglas-fir; as percent frequency was 99% (Table 9). of Use �297 Table 9. those used Conifer as blue species in the random (%) grouse winter-use trees, sample compared Middle Park, to Colorado, 1981-83. Green Species Douglas- fir Mountain Use Random (~ = 96)a (N = 884) Whiteley Peak Use (N = 97)a Random (N = 993) 100 99 86 31 0 <1 7 41 Limber pine 0 0 7 7 Lodgepole 0 <1 0 18 0 <1 0 3 Subalpine fir pine Engelmann a time. Trees spruce used more than once by grouse were only counted 1 �298 \. trees at Whiteley Peak were subalpine fir. Grouse used 86% Douglas-fir, Douglas-fir 7%limber pine, in greater proportion 31%) than its occurrence in the random sample of conifers suggesting for this a preference Forest differed Peak Structure. (Table not differ variation at Green 10, Appendices among stands characteristics A and B). study quarter area (1976) for clumped coefficients Table ~ l. 0) - principal at Green Mountain Whiteley Peak (Table Principal as a gradient Mountain density A matrix components by sites of correlation (with P < 0.05 in - were derived from random gradients from random sites at 11). (PC) I at both study areas of conifers. of large conifers (2::39 ern dbh). was interpreted PC II for Green and PC III for Whiteley Peak represented a gradient of non-conifers. of low PC III for Green and PC II for Whiteley Peak were gradients The 4 structural reported Four major structural from low to high density of low to high density did were done on random and 3 were derived component to high density Mountain to those for 15 variables 10) was used in each analysis. (eigenvalues sites analyses variables) that spacing. and for Whiteley Peak. (standardized variable coeffi- Roth component at Whiteley means for this distances; to random sites of from 42 to 69% and were comparable for Green Mountain 9), was the coefficient ranged principal at random The single cient Separate (86 vs. (Table Mountain- and among stands at either for point-centered 7% species. - Structural among stands and of increasing PC IV at Green Mountain was a gradient of medium-sized gradients at Green conifers (24- 38 em dbh). Mountain and the 3 at Whiteley �Table 10. random Parametric sites, Middle and nonparametric Park, tests for differences among stands on structural Trees (mnemonic) ~7 cm dbh Iha Conifers (DETR) '?,7 em dbh/ha Non-conifers Non-conifer Conifer (DECO) ?,7 em dbh/ha saplings saplings (DEOT) <7 em dbh/ha (OT06) <7 em dbh Iha (C006) ANOVA KruskaIWallis ANOVA KruskalWallis <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 0.003 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 7-15 cm dbh/ha Conifers 16-23 em dbh Iha (C023) <0.001 <0.001 <0.001 <0.001 Conifers 24-38 ern dbh/ha (C03B) <0.001 <0.001 <0.001 <0.001 Conifers '?,39 cm dbh/ha (CO>39) <0.001 <0.001 <0.001 0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 0.001 O.Oll 0.003 <0.001 0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 (DIST) <0.001 <0.001 <0.001 <0.001 (CVDIST) 0.192 0.162 0.71B 0.B39 CV (%) (cm) of conifers for dbh Mean dbh (cm) CV (%) for dbh Canopy height Mean (CODBH) of conifers (CVCODBH) of non-conifers (OTDBH) of non-conifers em) above em) of point-centered CV (%) (C015) Whiteley Peak P of no difference among 13 stands Conifers Mean dbh for point-centered plot (CVOTDBH) (CAHT) quarter quarter at 19B1-83. Colorado, Green Mountain P of no difference ;'mong 16 stands Variable variables distances distances N 1.0 1.0 �w o o 11. Table Correlation from 200 random sites between structural at Green Mountain variables and and varimax-rotated 180 random sites principal at Whiteley Peak, components derived Middle Park, Colorado, 1981-83. Principal Green Whiteley Peak Mountain IV -0.00 0.34 0.28 0.70 0.61 -0.02 O.92 0.06 -0.08 0.27 0.96 -0.05 0.08 0.04 -0.12 0.82 0.07 -0.03 0.87 -0.11 0.23 -0.12 0.43 -0.00 -0.22 0.43 -0.29 0.74 0.07 -0.06 -0.36 0.57 -0.04 -0.07 II I II III III I Variable Components ?,7 cm dbh/ha Trees a ~7 cm dbh/ha Conifers ?7 cm dbh/ha Non-conifers Non-conifer Conifer 0.85 saplings saplings < 7 cm dbh/ha <7 em dbh/ha Conifers 7-15 cm dbh/ha 0.90 -0.11 0.01 -0.11 0.84 -0.04 -0.18 Conifers 16-23 cm dbh/ha 0.62 -0.00 -0.09 0.35 0.77 0.08 -0.10 Conifers 24-38 cm dbh/ha 0.15 0.19 -0.17 0.81 0.52 -0.10 0.35 Conifers ~39 ern dbh/ha -0.08 0.82 -0.04 -0.13 -0.18 -0.11 0.78 (cm) of conifers -0.12 0.75 0.04 0.43 -0.14 -0.03 0.83 0.54 0.57 -0.01 -0.11 0.57 -0.18 0.44 -0.12 0.11 0.85 -0.09 0.11 0.79 0.01 -0.06 0.18 0.80 -0.15 -0.08 0.82 0.00 0.23 0.81 0.03 0.40 0.51 0.09 0.64 -0.59 -0.34 -0.27 -0.32 -0.52 -0.40 -0.30 30.4 16.7 16.4 7.6 29.8 19.7 13.3 30.4 47.1 63.5 71.1 29.8 49.4 62.7 Mean dbh CV (%) for dbh Mean dbh of conifers (cm) of non-conifers CV (%) for dbh Canopy height of non-conifers (m) Mean (m) of point-centered Percent quarter of total variance Cumulative percent aUnderlined of variance coefficients indicate distances key variables defining each component. �301 Peak accounted respectively for 71 and 63%of the variation (Table Factor gradients were calculated for each conifer defined by blue grouse Occupied stands (stand III, with a few large these extremes many large stand component included a broad (;;:39 em dbh) Figs. range (stand had moderate or medium-sized included (stand in others stands I, Figs. conifers densities 5). low density and 8). of large Stands (Figs. Conifer stands (stand conifers 2nd- Figs. (occupied) stands stands including between few to Quaking aspen in some occupied stands habitats at Whiteley Peak conifers aspen and few aspen density 7 and 8), and numerous size of large and stands (stand VI, Figs. (occupied-intensive of blue grouse with a use) 7 and were similar in 5-8). unoccupied and XIV at Green Mountain stands Occupied with a moderate III. similar to and different occupied types. old-growth 5 and 6). of large with >5 observations stands at Whiteley Peak Occupied stands those with < 5 ob ser-v ations structure of structural of conifers (Figs. with a high density 7 and 8), stands from high density IV, Fig. 6). and or medium (24-38 em dbh) conifers (Fig. and few aspen Conifer 5 and 6) to low density conifers (case) on the structural axes. and lor Rocky Mountain juniper were abundant but absent site were ordinated at Green Mountain varied with few large conifers for each random by the principal occupied growth structure, 11). scores mean scores in forest (Figs. stands. and winter These by grouse during from occupied (Figs. winter included habitats. Stands 5 and 6) and stands XII and XIII 7 and 8) were similar in structure stands were immediately use by blue grouse adjacent was expected XIII to several to occupied but not observed. �302 a ._ CI) z cc WwW • OCCUPIED-INTENSIVE 0 OCCUPIED • UNOCCUPIED 2 I XO .11 Z(!)!:!: OCCZ .1 0..<0 ..J() ~LL- IV. o OOI O>-~ _Jt-:::E XIV. VII O XII XIII • ..OVI VIII••• V OXI .111 OIX c:~() - USE • XVI W 0> OOC") Z a: 1\ n, ~ g-2~------'_---------~-------L------~ -2 o 2 LOW HIGH DENSITY OF CONIFERS PRINCIPAL COMPONENT Fig. 5. Conifer stands principal components Colorado. 1981-83. ordinated I and II. by mean factor Green Mountain. scores I on Middle Park. �303 I o • OCCUPIED-INTENSIVE o OCCUPIED • UNOCCUPIED 2 I > J- CJ) Z a: W w_ Z !!:I .(]) • XIII II.OXII .1 0 ~c a. ~ 0 o xo o~ VIII•• 0 lJ..O °eo III. • a. zw .IV XIV OXI OVI xv 0 a. VII V.OIX XVI" >-('1) ...J t-' _"I:t <{ CJ)C\J o z a: USE ~ 0 ...J -2 -2 LOW 0 2 HIGH . DENSITY OF NON-CONIFERS PRINCIPAL COMPONENT Fig. 6. principal Colorado, Conifer stands components 1981-83. ordinated by mean factor III and IV, Green Mountain, scores III on Middle Park, �304 :c 2, o :c C/) I- 0: Z OCCUPIED-INTENSIVE • OCCUPIED • UNOCCUPIED 0 W W LL Z Z 0.. I OVI Z ~ 0 0 o Z « 0.. o Z a: 0 0.. .x .XI .11 0 0 o _. USE OIV 0 LL XIII. >- ~ C/) XII VIII V 0 Delli • o 0, 'x o VII Z w 0 ~ 0 _J -2 -2 LOW 0 2 HIGH DENS'ITY OF CONIFERS PRINCIPAL COMPONENT Fig. 7. principal Colorado. Conifer stands components 1981- 83. ordinated I and II. by mean factor Whiteley Peak. I scores on Middle Park. �305 I o • OCCUPIED-INTENSIVE o OCCUPIED UNOCCUPIED 2 • I ••••• z Cf) a: ww Z ~u. a:0 «z o, ~ 0 ., VIII W 0 X". o V ..J U LL_ 0.", II. o OJ: >co ~o -' « Cf)~ n, Zu - w00> o Z a: - USE .XI 0 X OIX 0 • IV XIII.OVI 0 ('I) 1\ VII o, ~ 0 ..J -2 0 -2 LOW 2 HIGH· DENSITY OF CONIFERS PRINCIPAL COMPONENT Fig. 8. principal Colorado, Conifer stands components 1981-83. ordinated I and III, by mean factor Whiteley Peak, scores I on Middle Park, �306 Unoccupied stands that differed from occupied XV and XVI at Green Mountain (Fig. low densities and stands Peak (Fig. 7), There trees of small conifers, used. by grouse use sites 5), both both with high densities were no structural differed 16 variables) and among 3 stands 16 variables) (Table 12). sagebrush of conifers with and aspen. Forest at Whiteley Peak Samples in the other sites or among structure at winter- (P < o. 05 for 14 of at Green Mountain (!: < O.05 4 occupied Green Mountain and 6 at Whiteley Peak were inadequate stands at for inclusion Structural similar among stands at Green Mountain were not the same as those were similar among stands at Whiteley Peak. 24-38 em dbh and the coefficient quarter tain, distarices whereas conifer stands at Green Mountain differed Appendix gradients and age of use trees (Table 12, Appendix among occupied stands stands were of conifers (aspen) (Table for did not differ 12, Appendices differed E). at Green Moun- of variation C and among occupied stands Height and age of use at Whiteley Peak by blue grouse derived were ordinated from the principal (Table 12, scores at grouse-use within conifer stands to examine habitat on a within-stand basis. along the structural component Factor ability Density that F). Sites used sites. coefficients at Whiteley Peak sites for point-centered among occupied dbh , and mean dbh of non-conifers Dbh , height, trees of variation mean dbh of conifers, among occupied D). did not differ at grouse-use for 13 of in this comparison. that features stands types among winter stands. among 8 stands included X and XI at Whiteley similarities in different stands analyses and random sites preferences Comparisons on random were compared relative to avail- were made for 6 �Table 12. grouse Parametric winter-use and nonparametric sites and use trees. tests of differences Middle Park. among stands Colorado. for structural variables at blue 1981-83. Green Mountain P of no difference among 8 stands Whiteley Peak P of no difference -among 3 stands Variable ANOVA Krusk~~ Wallis ANOVA KruskalWallis Trees <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 0.002 <0.001 ~7 cm dbh /ha Conifers ~7 cm dbh/ha Non-conifers Non-conifer Conifer ~7 ern dbh/ha saplings saplings <7 ern dbh/ha <7 em dbh/ha Conifers 7-15 cm dbh/ha <0.001 <0.001 0.022 0.016 Conifers 16-23 em dbh/ha 0.004 0.003 <0.001 <0.001 Conifers 24-38 cm dbh/ha n .107 0.146 <0.001 <0.001 Conifers ~39 cm dbh/ha <0.001 <0.001 0.020 0.012 <0.001 <0.001 0.341 0.260 <0.001 <0.001 0.445 0.222 0.080 0.039 0.406 0.058 CV (%) for dbh of non-conifers <0.001 <0.001 <0.001 <0.001 Canopy height <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 Mean dbh (cm) of conifers CV (%) for dbh of conifers Mean dbh (cm) of non-conifers (m) Mean (m) of point-centered CV (%) of point-centered Dbh (cm) of use trees Height (m) of use trees Age (years) of use trees quarter quarter distances distances 0.101 0.079 0.004 0.004 <0.001 <0.001 0.770 0.943 0.003 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 w o ......• �308 occupied stands sufficient at Green Mountain and 3 at Whiteley Peak that samples Blue grouse of large total density ences for only grouse-use Use trees used had larger 11 and 12). a lower 9 and 10). between gradient Differ- conifer grouse-use Grouse-use 24-38 em dbh) and and random within within between for density were (PC 1) were significant of non-conifers) relationship the largest 1 stand 1 stand grouse-use of non-conifers Forty-eight and (Table 13). and random or density with broad, sample) of this form of conifer feeding rounded or there of at Whiteley Peak. between use trees was not detected. crowns 15). evidence stands at Green overmature Availability was not determined. (random Grouse were of feeding (clipped at Green Mountain and 27 of 97 use Dbh and height that were large, (Table stands. within occupied and 58%of the use trees was direct in 27 of 96 use trees within occupied conifers respectively, conifers observed conifers dbh than random Mountain and Whiteley Peak, feeding density 24-38 em dbh , Blue grouse 0.200) extent, on the large 13 and 14). of conifers srtes along the gradients conifers density (Tables was no consistent trees (Figs. differences on PC III (density on PC IV (density needles) with a greater to a lesser and random sites on the conifer 3 of 9 stands differed (Figs. and, (7 of 9 stands) for 5 of 9 stands; random sites There sites PC II at Green Mountain and PC III at Whiteley Peak, significant sites preferred (9 of 9 stands) of conifers between gradient, (~7) of use sites. consistently conifers had did not differ (!_ test, were fed upon and use trees Large accumulations of grouse P > in which droppings �309 • RANDOM SITES GROUSE-USE SITES * ::r: 2 o *\ * • ::r: I- 00 a: Z ww W IV <!)LL z a:z 0 n, ~ 0 o _. « a.. o Z a: ....J{) ~I • 0 >-CO ....0 * .~ • *-. 1\ <0 *-----. V III oo~ z{) we)) O(Y) 1\ - a.. ~ 0 ....J -2 -2 LOW 2 0 HIGH - DENSITY OF CONIFERS PRINCIPAL COMPONENT Fig. 9. Mean factor for grouse-use stands. scores and random Green Mountain. on principal components sites within occupied Middle Park. Colorado. I I and II conifer 1981-83. �310 • J: 2 o * RANDOM SITES GROUSE-USE SITES J: ••••• Z W Z C/) • " " a: WW (!)LL 11 • • Z 0 a: «0 a, ~II ...JU ~ LL 0 oJ: co o 0 >-0 _J ••••• « n, zU o We> OM C/)~ Z a: 1\ n, ~ 0 ...J ·2 ·2 0 2 LOW HIGH DENISITY OF CONIFERS PRINCIPAL COMPONENT Fig. 10. Mean factor scores on principal components for grouse-use and random sites within occupied Whiteley Peak. Middle Park. Colorado. 1981-83. I I and III conifer stands. �- ""- Table 13. Factor scores on the first 4 principal components for grouse-use and random sites within 6 conifer stands. Green Mountain. Middle Park. Colorado. 1981-83. Stand I I! III IV V VIII Grouse Random sites Principal component ~ SD I II III IV I I! III IV I II II! IV I I! III IV I II II! IV I I! III IV -0.24 1.09 -0.47 0.24 0.40 1.46 -0.33 0.58 1.79 -0.59 P sites ANOVA N MRPP 0.49 20 0.230 0.717 0.88 20 0.119 0.083 0.93 20 0.027 0.025 1.06 20 0.628 0.663 0.27 0.52 10 0~009 0.021 12 0.95 0.98 10 0.161 0.132 0.16 12 -0.54 0.11 10 0.453 0.444 -0.20 1.57 12 0.63 1.35 10 0.216 0.204 1.11 0.68 14 1.65 0.99 10 0.240 0.125 N ~ 0.79 24 -0.17 1.34 24 0.47 24 0.01 24 0.57 12 1.43 SD -0.27 0.23 14 -0.44 0.26 10 0.095 0.095 -0.44 0.10 14 -0.44 0.12 10 1.0{)0 0.988 -0.43 0.66 14 -0.64 0.85 10 0.340 0.495 0.084 0.007 -1.30 0.38 9 -1. 08 0.26 20 0.174 1.69 1.19 9 0.02 1.49 20 0.006 0.67 1.23 9 0.70 1.47 20 0.574 0.962 0.34 0.39 9 -0.25 0.59 20 0.009 0.011 0.11 0.39 7 0.43 0.45 10 0.385 0.158 -0.11 0.35 7 -0.14 0.32 10 0.520 0.832 0.66 0.58 7 0.20 0.78 10 0.138 0.197 0.171 0.113 0.53 0.53 7 -0.06 0.80 10 0.60 1.28 11 -0.15 0.61 10 0.167 0.111 0.016 0.021 0.77 1.08 11 -0.13 0.36 10 -0.47 0.26 11 -0.24 0.69 10 0.592 0.316 -0.49 1.72 11 0.17 0.78 10 0.164 0.297 Vol t'-' t'-' �W I-' N Table 14. Factor scores on the first 3 principal components conifer stands, Whiteley Peak, Middle Park, Stand I Principal component SD N MRPP ANOVA 0.71 34 0.85 0.74 10 0.038 0.038 -0.39 0.41 34 -0.33 0.47 10 0.742 0.705 1.43 0.69 34 0.78 0.93 10 0.019 0.020 -0.53 0.40 23 -0.80 0.36 20 0.032 0.026 II 0.19 0.87 23 0.80 1.07 20 0.070 0.045 III 0.77 0.46 23 0.20 1.30 20 0.001 0.057 I -0.52 0.61 31 -0.22 0.56 20 0.071 0.084 II -0.61 0.19 31 -0.64 0.17 20 0.342 0.519 0.87 0.80 31 0.21 0.93 20 0.018 0.009 I III 0.30 P sites N III III Random - I SD sites within 3 1981-83. sites x - and random x - II II Grouse Colorado, for grouse-use �~ ~- - N=88 Ii I I I. ~=70 I I a: w I tfl- ~IIII :::> I N=13 I H=10 E <0.001 N=97 CD RANDOM TREES USE TREES , «0.001) 1:<0.001 «0.001) I Z o -c 1:<0.001 «0.001) .-~~---------------------------------- II Z N=27 I I IIV N=13 I M=14 I E=0.062 (0.036) -1 ~ t!l=53 c:p V I tJ=7 '--,J I VIII ~=35 1:=0.004 «0.001) I !:':I=ll E<0.001 «0.001) 60 70 >.y.:/ o 10 20 30 40 50 80 90 DBH (eM) Fig. 11. Means (± 2 standard I errors) and ranges random conifers within occupied stands. for dbh of blue grouse winter-use Green Mountain. Middle Park. Colorado. and 1981-83 W t-' W Probabilities are for ANOVA and Kruskal-Wallis. �UJ .,.. I-' ( .J RANDOM TREES USE TREES N=93 ----------.--1 --...----------------1 - I a: w P<0.001 «0.001) N=33 I I CD N=37 ~ ::::> II z I- I I I N=22 P<0.001 «0.001) c z -c N=87 1- • III ~ I I o 10 30 20 40 N=34 50 e<0.001 60 70 80 DBH (eM) Fig. 12. Means (±2 standard errors) and ranges for dbh of blue grouse winter-use and .\ random conifers Probabilities within occupied stands. Whiteley are for ANOVA and Kruskal-Wallis. - .•. - - Peak. Middle Park. Colorado. 1981- 83 «0.001) 90 �315 occurred under nearly were fed upon than Table 15. all use trees. my observations Forms of blue grouse Colorado, More of these trees probably indicated. winter use trees, Middle Park, 1981-83. Tree form (%) N Conical Green Mountain 96 35 48 17 Whiteley Peak 97 24 58 18 Study area Forest Because Structure at Sites reason winter-use to believe sites were observed. both sexes (Table saplings sities differed 0.145) tests. difference males, Density was their only males females, any important had highly and differences non-significant of non-conifer at Whiteley Peak with greater only males occurred. to this where there No biological since blue grouse (aspen) den- importance do not use aspen winter. Topography. ranged Most variables among use sites where can be attributed among sites where - together winter, between were few sites did not indicate for nonparametric at sites during G). areas during differences There Comparisons 16, Appendix probabilities structural would exist. were observed were commonly observed on the study that Deformed top Used by Male and Female Grouse. male and female blue grouse and used the same. habitats little Rounded - Elevation at grouse-use from 2,402 to 2,858 m (~= from the elevational sites on Green Mountain 2,644 m) and did not differ distribution at random sites (Table (~= 17). �W I-' 0'\ Table 16. Parametric males. females. and nonparametric and both sexes. tests Middle Park. of differences Colorado. among blue grouse winter-use sites for 1981-83. Green Mountain P of no difference among use sites Krusk~Wallis Whiteley Peak P of no difference 'among use sites ANOVA KruskalWallis Variable ANOVA Trees 0.544 0.401 0.092 0.058 0.667 0.650 0.286 0.260 0.691 0.509 0.313 0.608 0.569 0.509 0.097 0.014 0.942 0.951 0.095 0.588 ?;7 em dbh Iha Conifers ?7 cm dbh/ha Norr-coriifer-s R7 em dbh/ha Non-conifer Conifer saplings saplings <7 em dbh Iha <7 cm dbh Iha Conifers 7-15 ern d bh Iha 0.929 0.628 0.251 0.399 Conifers 16-23 cm dbh/ha 0.414 0.380 0.138 0.263 Conifers 24-38 ern dbh/ha 0.889 0.884 0.100 0.136 Conifers ?39 em dbh/ha 0.860 0.820 0.399 0.363 0.890 0.582 0.259 0.304 0.165 0.182 0.302 0.487 0.587 1.000 0.552 0.519 0.334 0.773 0.369 0.198 0.063 0.172 0.176 0.080 0.017 0.174 0.125 0.123 0.423 0.302 0.260 0.782 Mean dbh CV (%) for dbh of conifers Mean dbh CV (%) (cm) of conifers (cm) of non-conifers for dbh of non-conifers Canopy height (m) Mean (m) of point-centered CV (%) of point-centered quarter quarter distances distances �Table 17. Elevation ,(m) at blue grouse Grouse use winter-use and random sites, Middle Random sites N x - SE Max Min Park, Colorado, 1981-83. P sites N x - SE Max Min ANOVA Study area Green Mountain 100 2,644 10 2,854 2,402 200 2,631 8 2,858 2,402 0.145 Peak 104 2,867 8 3,040 2,614 180 2,821 8 3,070 2,584 < O. 001 Whiteley Vol I-' -...J �318 The mean elevation 2,867 m (range most conifer sample 2,614-3,040 habitat (Table Slope, used by grouse m). This was higher and vertical and random percent sites of grouse-use sites on northwest, north, sites west, 41%of random sites Large stands elevations percent of conifers east, located by tracking radio-marked areas. Wintering marked grouse 1 in lodgepole pine, These were mature forests 19). Spruce-fir Areas. analyses and 80% of and vertical sites were on south, of grouse-use one-third that areas forests sites sites and of Whiteley Peak. occurred at low slopes of Whiteley Peak. wintering areas were migrated from the study were visited infrequently; were not attempted. forests, Five radio- 6 in spruce-fir and 1 in a Douglas-fir that of and random aspect - Additional grouse in spruce-fir pine, (Table one-third but only 38%of random percent off the study habitat wintered sites grouse-use and southeast Off Study detailed 74%of grouse-use of grouse-use on the upper Winter Habitat therefore, were that were not used by grouse on northeast, areas sites However, aspects, Eighty- were on and 74%of random between Eighty-seven occurred 18). 18) as 8.9% of grouse-use (Table and northwest aspects. between (Table were on the mid- and upper Eighty were on these did not differ aspects; were on 16- 45° slopes. differed. southwest, sites Slope did not differ at Whiteley Peak position than from the random and 77%of random sites and northeast and 75% of random sites the mountain. position at Green Mountain 81%of grouse-use random (~<0.001) 17). 16-45° slopes; sites at Whiteley Peak was on Whiteley Peak as determined aspect, grouse-use three at sites /spruce-fir /lodgepole type. had low to high densities of conifers used by wintering were grouse �Table Topographic 18. Colorado, characteristics (%) winter-use and random sites, Use (~ = 100) Variable Whiteley Peak Mountain Random (~ = 200) (0) pa MRPP Use (!'i = 104) Random (N = 180) 17 23 10 16 16-30 59 61 53 41 31-45 24 16 34 39 46-60 0 0 3 4 Aspect 1.000 <0.001 N 21 27 2 9 NE 42 29 7 30 E 1 11 2 14 SE 1 1 9 8 S 1 4 24 12 SW 7 7 13 8 W NW 7 3 16 9 20 18 27 9 position <0.001 0.200 14 6 10 12 35 41 87 41 mid 1/3 39 34 2 32 lower 1/3 12 19 1 15 ridge upper 1/3 aprobability of no difference between grouse-use and random pa MRPP 0.227 0.104 1-15 Vertical Middle Park, 1981- 83. Green Slope at blue grouse sites. w I-' 1.0 . �320 Table 19. habitats Characteristics off study at blue areas grouse in Middle Park, Conifer density (per ha) Forest type Lodgepole N pine Lodgepole/ winter-use Colorado, sites in 1981-83. Conifer dbh (cm) Canopy height (m) Min Max x - Min Max x - Min Max 300 1,700 20 14 30 16 14 17 7 400 100 800 27 8 43 16 9 23 4 400 200 500 25 8 49 17 14 19 2 600 200 1,000 31 10 61 27 25 30 4 x 1,100 aspen/subalpine fir Lodgepole/ spruce-fir Spruce-fir �321 uneven-aged conifers. stands Where lodgepole poles were large, pole forest firs, but grouse trees in these fir, the lodgepole old-growth habitat types and Engelmann A mature than used was similar to stand dense, included spruce. the lodge- (> 50 em dbh) pine rather Douglas-fir. (~39 ern dbh) with spruce-fir, had a few large forest and large with 16-30 ern dbh. B) as moderately beneath subalpine used {spruce-fir (Appendix growing mature pine occurred used by grouse grouse Douglas-fir Peak with small (7-15 cm dbh) mature Douglas- the firs. The I at Whiteley spruce- fir was Conifers lodgepole lodge- actually pine, used by Douglas-fir, �322 DISCUSSION Movements from Breeding Movements of blue grouse can be characterized Adult and yearlings spring used way) during previous during and Elliott the previous varied (Marshall non-migratory population their traversed ing areas. suggested as defined of a spruce grouse migrants by 1960, Bendell km) and pop ulation , were year- did not occupy and, therefore, by Baker (1978). (Dendragapus was identified the A canadensis) by Herzog (1980). Blue grouse nearest the year (1- from the were short-«3 birds to the traveled were' females that These in returning Distances There throughout which also contained and Keppie areas within the same breeding as non-migrants segment in fall, areas 1978). km) among grouse 1968). areas. or habitats were not classified to breeding year. migrants of the study same home range (Baker 1946, Hoffmann 1956, Mussehl (> 5 km) migrants residents and wintering but were within the range Most (13 of IS) short-distance round Areas migration areas (0.1-29.5 1967, Zwickel et al. long-distance return to wintering population, studies breeding moved from wintering migration same breeding between as a seasonal and from breeding same areas to Wintering migrating breeding apparently short areas. suitable Long-distance distances wintered Grouse migrating winter migrants habitat in conifer forests long distances en route to their within each population winter- moved in �323 the same general winter near djrection. each other. marked and others described (N this 1955). migration Migration on my study altitudinal moved laterally Thus. downward Previous dates 1947, Bendell unsuccessful studies and Elliott areas. followed during October-November. Most habitats below usually involved distances and wintering areas. Rogers (1968) occurred on the Uncompahgre below breeding movements Most authors areas later arrived on the winter areas. failed to document (Marshall 1946, Wing 1968) assumed during in fall. on wintering Males and some unsuccessful females moved to locations (1972). short arrived my study of seasonal migrating on wintering fall. while females with broods associated Some grouse 1967. Zwickel et al. females arrived change areas 1946, Wing in Middle Park in open shrub of blue grouse conifers been of proximity to wintering of blue grouse on wintering of blue grouse were breeding fall movements of 1-2 km 1903, Marshall by Zwickel and Bendell breeding in Colorado where within have previously a function movement. by Bendell Limited observations the elevational movements between from Whiteley were reported (Anthony However, areas an upward grouse of migrants some wintered patterns a view also expressed for them to areas. migration was strictly zone. arrival that of blue grouse contention. the conifer Plateau range Similar findings 5) indicated as an altitudinal with their noted = movements 1947, Bendell grouse winter up to 20 km from natal Seasonal habitats, was no tendency (1967) and Zwickel et ala (1968). juveniles support e. g., >300 km", Peak encompassed and Elliott but there range males and summer or early Radio-marked areas during or non-breeding during summer, �324 remained localized moved 0.9-4.5 areas at these through km to wintering areas. to be spatially distinct appeared Blue grouse in conifer areas may select forests because they are terrestrial tation, berries, and leaves During winter, blue grouse areas ing areas based The migration the typical comparable 1980). to" males, during and others ing areas. banded There birds breeding winter territories, in spring Studies nested i.e., that habitat used by other wintering and occupied of other on breeding territories tetraonids Beck 1977, Herzog and Keppie explain why this pattern et al. 1980). summering and winter- (Baker long distances comparable to they inhabited to their winter- or observations on or adjacent included grouse of to their suitable during territories winter. vacated these elsewhere. have documented for females (Weeden 1964, Irving that evidence territories needles for females immediately adjacent even when their 1982). did not conform to distances in forests males wintered vege- overstory. movements was no radio-tracking Adult males observed areas greater females nested indicating habitat, understory of the forest few males moved short Several winter may select While some females migrated many females. requirements. and feed on conifer of the forest i. e., locations (King and Bendell of male and female blue grouse avian pattern. 1978. Greenwood habitat range. and feed upon herbaceous Blue grouse on characteristics and summering on the winter seasonal are arboreal on characteristics and in October summer and winter of shrubs 1961, King 1968). based areas of different summer. (Hoffmann Wintering different During September, greater movements 1967, Hoffman and Braun Empirical evidence was not observed 1975, is lacking for blue grouse. Fidelity to �325 to breeding areas is considered systems based toriality in grouse. ptarmigan on resource winter of competing defense near their strong breeding space. fidelity their areas stable strategies maximize individual population selection tageous migration (Swingland fitness. chances (Bendell and Elliott wintering 1983). Stability strategies to migrate for blue grouse or age and appeared year. There are stabilized (i. e , , relative migrants may be maintained short whether in a 1978, dependent it is advandepends each strategy. were not conditional fixed for an individual, were more long- than (Baker or long distances are pursuing to fitness) by frequency 1976), i.e., (Maynard Smith and Parker for an individual mixed evolu- 1983) used by blue grouse for long- and short-distance tion strategies second male are territorial may represent Migration upon how many in the population sex, their does not include when the net costs and benefits are equivalent Swingland that territories. Short- and long-distance tionary to increase Male blue grouse to breeding mating 1980), e. g . , terri- (1975) suggested areas 1967, Lewis 1979), but this apparently near for males having (Greenwood Hoffman and Braun for breeding and exhibit advantageous Migra- on breeding at least short-distance status, by their migrants within a population. Several frequency factors may affect dependent manner resulting breeding areas during breeding areas and if availability with increasing of fitness, relative density winter. of blue grouse in fewer grouse If winter of grouse, for some grouse fitness resources of resources remaining to other near are limited near per individual it may be advantageous, to migrate in a habitats decreases in terms for winter �326 (the ideal free distribution dence suggesting ber of grouse that phagous winter are abundant in certain than areas. quantity Quality areas, but may affect their conifers was little evi- However, for grouse distribution and, although conifers food may not be limit- by Zwickel and Bendell during thereby, a mono- may feed preferentially of winter as suggested food may be having Thus, blue grouse and quantity of blue grouse them to certain There (Miller and Watson 1978). on wintering trees. 1972). of cover or food was limiting the num- on my study rather diet ing abundance (1972), quantity wintering limited by quality of Fretwell winter by restricting limiting densities within occupied habitats. Another areas possible factor limiting year-round may be summer food resources. of herbaceous vegetation, leaves (King and Bendell 1982). on breeding by early areas Dessication Both a decline in quantity dessication. herbaceous develops breeding areas. ing to higher close proximity parental care to suitable for chicks, their at higher forests suitable winter habitat. elevations. fitness from plant elevations coincident and lush with dessi- are limited on advantage by migrat- summer food is abundant Because they may have greater relative occurs in Middle Park. at higher may gain a survival elev ations where females to increase to habitats is delayed summer vegetation If summer food resources some grouse during of food may result in subalpine cation at lower elevations. of shrubs in most years of breeding feed on a variety of herbaceous and quality Plant phenology growth Blue grouse and berries August occupancy and in males provide opportunity by migrating Females have greater no than long distances reproductive �327 investment in their to restrict their offspring, movements, and it may be advantageous thereby for them minimizing chick mortality. Winter Home Ranges Movements of blue grouse an area adults <10 ha for adults during winter and <45 ha for juveniles. were smaller than those of juveniles, eral avian pattern (Greenwood areas of greater 1980). than adults because juveniles ing sites. were not observed Population Wintering small groups patterns grouse blue grouse (2-6 birds), than associated in selecting with adults suitable aggression or winter- between grouse winter. Characteristics During Winter in Middle Park were observed and an occasional and disassociated large with other and size fluctuated have been <6 distinct adults more on wintering interactions and overt of with the gen- throughout flocks on each study as lone birds, flock (12-20. birds), also found by Caswell (1954) and King (1971). Flock membership during may wander experience displays during consistent of intraspecific lack prior Territorial grouse within Home ranges movements for juveniles Juvenile because were restricted grouse Radio-marked during winter. winter. There may area at any given time winter. Although at the study did not differ there areas was a slight during from 1: 1. in the same tree winter, tendency the observed Both sexes and occupied to observe more females sex ratios in flocks were commonly observed the same conifer stands together on and off the �328 study areas. These who observed observations are contrary only males on subalpine to those wintering areas of King (1971),. in British Colum-. bia. Winter Habitat Douglas-fir occurred on study areas preferred over However, migrant uncommon. subalpine One grouse observed blue grouse 1983- 84. forests dense occupied (100-200 years Previous old), winter (Caswell densities studies observed, Douglas-fir areas was spruce. used spruce- was nonexistent pine even when large T. E. Remington or Douglas- (pers. commun.) pine on Whiteley Peak during species preferred a broad structural 2nd growth by grouse «100 trees /ha) species have associated of conifers mature forests range (50-70 years may vary blue grouse 1983). of Douglas-fir occupied as a result of selective Seven additional in stand conifer forests old) or mature old-growth for-est s associations. with open mature Open to moderately were commonly used by blue but they were not restricted variation of conifer or mixed species 1954, King 1971, Stauffer stands Some annual off study lodgepole with a single in Middle Park, Two mature by blue grouse and Engelmann Douglas-fir old) to open (190-530 trees Iha) grouse where conifer (900+ trees Iha) (200-600 years used areas. Blue grouse from dense wintered in the stand. Thus, among contiguous pine, used lodgepole using sites At Whiteley Peak, lodgepole that pine forests were present forests fir, grouse fir trees winter at all wintering in Middle Park. fir or lodgepole Preferences by blue grouse logging occupancy stands to these during types. had lower the by blue grouse were occupied 1930's. was the 2nd �329 winter of the study, 1982-83. and 3 stands Variation of radio-marked in observer grouse most of the variation conifer stands easily searched for grouse ation in stand Several conifer similar in structure unoccupied able. stands may not be unsuitable i.e., habitats winter populations all suitable habitat. An alternative were unsuitable stands in some unmeasured territorial beyond of blue grouse stands wintering areas males and nesting level of use was apparent. between extent of use during used intensively structural of habitats types during as those vari- that were when numbers (Fretwell were insufficient in 1972), to saturate is that these unoccupied differed from occupied stands were used by the breeding season. was no evident and forest during and These were not determined, winter by but simply less suit- some threshold included used infrequently. used by blue grouse winter were used. habitat, There winter areas most annual during that females during relative a to juveniles. These stands and occupancy to wintering explanation attribute. Grouse d en s itie s in forest stands in stand may become occupied increase 2 small «1. 5 ha) 1981, Wiens 1981) and not used by grouse habitats account for si /e may affect than juveniles, and location to stands Less suitable more suitable fidelity may be attributed stands in number were used one winter variation exhibited home ranges occupancy However, (Rice et al. for some of the observed smaller winter probably in population among habitats Since adults and an increase the 2nd winter Annual variations blue grouse. occupied during occupancy. distribution may account efficiency in stand not the other. species used in 1981-82 were not used in winter, structure. but a relationship Conifer the same wide range Thus, within grouse the range densities may of �330 be unrelated habitat to habitat structure, by Zwickel and Bendell and overwinter survival rates as also suggested (1972). A better for grouse for breeding estimate within each habitat needs to be determined before each type can be rated tance for perpetuating blue grouse Conifer stands habitats (50-70 years growth provides lar topographic suitable winter selection Maturing habitat through winter trees. included and those The former but the latter 2nd-growth stage were conifers of stand develop- for blue grouse. winter-use Topography as to its impor- of small conifers the earliest did not select feature. during of mature at any time. type (Van Horne 1983). and brood rearing, old) may represent Blue grouse habitat trees/ha) for nesting used by grouse ment that «70 (>1,200 trees Iha) were used populations not used by blue grouse with low densities with dense rarely stands of density sites based may indirectly its effect on forest on any particu- relate development to winter and stand structure. Conifers actually among different preferred used by blue grouse stands, the largest by Caswell (1954), but within «45 by blue grouse Spruce grouse caillie (Tetrao conifer forests greatly blue grouse conifers. Use of large conifers King (1971), and Stauffer (1983). ?;7 cm dbh has limbs stout 7-15 ern dbh a stand varied years enough of age), during winter. (Gurchinoff urogallus) during to support although Saplings and Robinson (Seiskari winter, •...: consistently also was noted Any conifer a grouse, abundant, in size but conifers were rarely used <6 em dbh were not used. 1972, Ellison 1962), both species also do not use saplings 1976) and caperwhich occupy and small �331 conifers during winter. conifers and lor food for grouse concealment, Preference selection. Large for large Blue grouse food during winter conifers during dependent 1943, Stewart 1968), and they use the same conifers (King 1973, this study). ferences food selection of secondary Kuropat suggests defense 1980). reducing that compounds Conifers factors can contribute of secondary growing site conditions, of trees (Kozlowski 1971). large, mature conifers take of secondary Protein herbivore fire, food selection. Studies conflicting results. 1972), and processes including and maturation may preferentially to determine needles feed in to minimize their levels as reported current whether high in protein Boag and Kiceniuk in- have been found for spruce needles during grouse grouse of blue grouse have produced for Similar conflicting which also feed on conifer Ellison (1976) found no preferential by spruce theory (1968) found no selection by Hoffmann (1961). results rich needles (Radwan in trees overcrowding, habitats and digestibility- resins (Mattson. 1980) is another feed upon conifer on protein (Bryant constituents. selectively winter. of herbivore Many environmental Blue grouse pre- with high levels potential compounds within occupied plant theory digestion phenolic 1970). defense for and roosting to slowing of the physiological disease, maximization high protein tannins, food animals have definite A current numerous (Maarse and Kepner and reduction for feeding that inhibit contain to winter upon conifers animals avoid plants compounds including and terpenes growth food items. shelter, 1944, Hoffmann 1961, King Most herbivorous among potential better winter. may be related are entirely (Beer may provide as reported feeding by Gurchinoff �332 and Robinson (1972). Further research is needed to clarify the role of plant chemistry in winter food selection and habitat use for blue grouse and other tetraonids. Large conifers may provide better shelter from weather and concealment from predators, but these factors may be of secondary importance in selection of suitable trees. Blue grouse will leave the .shelter of conifers to roost under the snow. Snow roosting may be an important method of conserving body heat during cold winter weather, as observed for ruffed grouse (Bonasa umbellus) by Gullion and Marshall (1968). Only' further investigation will indicate the rela- tive merits of large conifers as food and cover, and whether observed preferences are related to a specific need affecting winter survival. �333 MANAGEMENTIMPLICATIONS Blue grouse and structural largest range conifers specific or floristic an on-site whether of conifer accumulations and browsed habitats. Because area supported a broad winter there conifers are a good indication the were no with occupied to determine blue grouse beneath floristic prefering associated would be necessary of droppings branches during characteristics evaluation or not a specific Colorado occupy forests within occupied structural habitats. Large in Middle Park, during during winter. winter of concentrated use by blue grouse. Some blue grouse tions adjacent wintering resource in conifer to or on breeding areas areas. age landowners to follow activity If ownership areas good If land use practices. on these areas and probably habitat. ized, impacts on breeding detrimental at relatively Most low elevation can do little to manage these impact on winter (Marshall forests in Middle Park are in private agencies the usual winter However. excessive 1946. Mussehl 1963, Rogers low elevabreedingl and public other than Cattle grazing has little, grazing and brood-rearing encouris if any. can have localhabitat 1968. Zwickel 1973. Stauffer 1983). Given the broad during between winter range of conifer and the extensive breeding and wintering forests that blue grouse movemerit s they are capable areas, winter habitat occupy of making is probably not �334 limiting for blue grouse and Bendell grouse populations, (1972) with different currently are abundant, of the fall population reasoning. is harvested towards to blue grouse maintaining, Blue grouse rugged ever, terrain winter rather that winter ing suitable conifers winter should uneven-aged habitat. will have a detrimental small patch Blue grouse during cuts rather areas provided in forests how- selective conifers (~24 em dbh) cutting clear-cut between promotes will be present used by grouse habitat. open areas openings Others, large Selective (as in lodgepole than one large these on steep, could be cut. 40/ha) that mature impact on winter will move across winter forests most compatible with maintain- in all cuts. clear-cuts must be done in wintering habitat. used by grouse, A few (at least ·growth which ensures Large recommenda- for timber harvesting. from forests be left standing on a given area. additional in inaccessible prescription «10%) should be oriented that have been or potentially is the silvicultural blue (Mussehl 1960, Bendell management habitat than creating When timber must be removed cutting Therefore, is unsuitable in areas by Zwickel Furthermore, by hunters winter frequently reached cibd only a small proportion and Elliott 1967, Hoffman 1984). tions relating a conclusion in winter When clear-cutting pine types), several are recommended. standing are <250 m, timber �335 LITERA TURE CITED Aldrich, J. 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Bull. 64. chicken 52pp. in �338 Hannon, S. J., L. G. Sopuck, movements of female blue delayed Harju, breeding H. J. dusky Herzog, 1974. in yearlings. Ph.D. R. S. 1956. at Sage Hen Creek, 1961. Wildl. Manage. Hoffman, blue ---- of the ecology of the Migration 142pp. in a local popula- 82:366-372. on a sooty California. Condor of the winter grouse population 58: 321-337. food of blue grouse. J. 25:209-210. 1984. grouse populations. Tech Press, 1980. Observations R. W. Robertson, induced Univ. Wyoming, Laramie. Condor The quality Spring for socially of some aspects Thesis, grouse. evidence 1982. Auk 99: 687-694. and D":"M. Keppie. tion of spruce Hoffmann, grouse: An analysis grouse. P. W., and F. C. Zwickel. Effects of changes Pages ed . Symposium Lubbock. regulations on 000-000 in S. L. Beasom and S. on game harvest management, Texas (In Press). , and C. E. Braun. tion of white-tailed in hunting 1975. ptarmigan Migration of a wintering in Colorado. J. popula- Wildl. Manage. 39: 485-490. Irving, L., G. C. West, L. J. tion of willow ptarmigan Jamieson, 1. G., fidelity ---- and F. in blue grouse D. H. habitat Can. 1983b. and their ing population. Johnson, grouse. J. Migra- Auk 20: 77-85. 1983a. Dispersal and site Zoo1. 61:570-573. Spatial relationship 1967. patterns of yearling to recruitment male into the breed- Auk 100:653-657. 1981. studies. and S. Paneak. in Alaska. C. Zwicke1. , and blue Peyton, The use and misuse Pages of statistics 11-19 in D. E. Capen, ed. in wildlife The use of �339 multivaria te statistics Agric., King, For. D. G. grouse ton. 1971. Gen. Tech. Feeding 1968. tion dynamics Columbia, M. S. Thesis. habits dynamics Univ. in relation of blue grouse. M.S. Vancouver. (Dendragapus Kos kimies , J. of blue Alberta. Edmon- in the subalpine. (L.). T. T. 1954. Clarendon R. A. 1982. Univ. Foods selected fuliginosus). Flocking British behavior Can. by blue grouse J. Zool. 60: 3268- 3281. in caper caillie , Tetrao tetrix (L.). Finn. 18: 1-32. 1971. Academic Press, D. L. Thesis, and black game Lyrurus Game Res. Kozlowski, to the ecology and popula- 62pp. obscurus 1957. urogallus Pap. of blue grouse Food habits , and J. F. Bendell. Lewis. RM-87. 6:121-125. R. D. Lack, Rep. U . S. Dep , 139pp. Syesis ---- of wildlife habitat. The ecology and population in the sub-alpine. 1973. King, Servo in studies Growth and development New Yor k , N. Y . The natural Press, Oxford, 1979. Suitability by male blue grouse. M.S. Vol. 1. 462pp. regulation U. K. of trees. of animal numbers. 343pp. and selection of territorial sites Alberta. Edmonton. Thesis, Univ. 1970. Changes used 85pp. Maar se , H., volatile Agric. and R. E. Kepner. terpenes in Douglas-fir Food Chern. needles 18: 1095-1101. in composition during maturation. of J. �340 Marshall, W. H. 1946. food habits Idaho. Martinka, Cover preferences, of Richardson's Wilson Bull. R. R. 1972. territories grouse seasonal and ruffed movements, grouse and in southern 58:42-52. Structural in southwestern characteristics Montana. J. of blue grouse Wildl. Manage. 36: 498- 510. Mattson, W. J., content. Maynard Jr. 1980. Annu. Rev. Smith, metric Mielke, J., Ecol. in relation and Syst. and G. A. Parker. contests. P. W. Herbivory 1984. 1976. Miller, 24:159-175. Multi-response permutation 5. 1978. Heather relevance to the regulation of red grouse o. C. O. 1947. small mammals. Mueller-Dombois, vegetation Mussehl, D., T. W. lations Berlin, Table of equivalent Am. MidI. Nat. populations 1960. John Wiley Blue grouse in the Bridger Mountains, eds. of populations and its , . Pages Ecological West Germany. Vol. 27. of North American 37: 223-249. and H. Ellenberg. ecology. Encyclopedia productivity W. Heal and D. F . Perkins, Springer-Verlag, Pages (In press). and A. Watson. studies. Mohr, Vol. procedures. eds. G. R., 227- 285 in The logic of asym- Anim. Behav. sciences. nitrogen 11: 119-161. 000-000 in S. Kotz and N. C. Johnson, statistical to plant & 1974. Sons. production, Montana. Aims and methods New York, N.Y. movements, J. of 547pp. and popu- Wildl. Manage. 24:60-68. 1963. cations. J. Blue grouse brood Wildl. Manage. cover 27: 547-556. selection and land use impli- �341 Nie, N. H., C. H. Hull, J. Bent. 1975. G. Jenkins, Statistical package McGraw-Hill Book Co., Noon, B. R. perate 1981. competition. Radwan, For. Rice, habitats: discriminant Dep . Agric., P. G., analysis. J. forest 1968. Fish and Parks, Tech. T. 1981. ed. of wildlife habitat. ____ Pages of genotypes deer. 1981. Can. in J. Servo Gen. 1971. species. scale in inter- 59-71 in D. E. Capen, Tech. ed. of wildlife habitat. Rep. RM-87. Phytosociological analysis of Ecology 52: 940-945. in Colorado. Publ , 21. 64pp. Why measure bird Dep , Agric., Colo. Dep , Game, habitat? The use of multivariate U.S. Bird community of temporal in studies The blue grouse in D. E. Capen, Rep. by black-tailed statistics and E. L. Rice. G. E. Rotenberry, Douglas-fir the importance For. Oklahoma upland Rogers, between and B. W. Anderson. The use of multivariate Risser, and expression 2:250-255. use of riparian U.S. of an avian guild along a tem- preference R. D. Ohmart, preting 675pp. 51:105-124. Differences to browsing Res. J., 1971. N. Y. the importance Ecol. Monogr. M. A. relation New York, gradient: and D. H. for the social sciences. The distribution elevational K. Steinbrenner, For. Pages statistics Servo Gen. 29-32 in studies Tech. RM-87. , and J. component A. Wiens. analysis D. E. Capen, wildlife habitat. RM-87. ed. 1981. A synthetic of bird Ihabitat relationships. The use of multivariate U.S. Dep . Agric. approach For. Pages statistics Servo to principal 197- 208 in in studies Gen. Tech. Rep. of �342 Roth, R. R. 1976. Ecology Seiskari, 1962. urogallus, 1927. Wilson Bull. species diversity. grouse, of capercaillie, Lyrurus tetrix. Tetrao Finn. Pap. grouse in Yellowstone Park. 39: 208-214. 1983. grouse in southeastern Moscow. ecology Richardson's D. F. blue and bird 22: 1-119. M. P. Stewart. On the winter and the black Game Res. Stauffer, heterogeneity 57: 773-782. P. Skinner, Spatial Seasonal habitat relationships Idaho. of ruffed and Ph.D. Thesis, Univ. Idaho. of the blue grouse. Condor 108pp. R. E. 1944. Food habits 46: 112-120. S'ttr-lirig v T. , and J. F. Bendell. recorded Sokal. R. R.. Census of blue calls of a female. J. Wildl. Manage. and F. J. Rohlf. 1969. San Francisco. Swingland, 1966. 1. R. Calif. 1983. grouse 30:184-187. Biometry. W. H. Freeman. 776pp. Intraspecific differences in movement. 102-115 in 1. R. Swing land and P. J. Greenwood. ecology U.S. of animal movements. Department summary. Atmospheric Van Horne. B. quality. Weber. U . S. Dep. Admin. 1983. response 1975. 85(14). Density Commerce, The Oxford. Climatological as a misleading data: Natl. U. K. annual Oceanic and indicator of habitat 47:893-901. ecology. manipulation Unit Press. eds. Pages 14pp. Blue grouse to habitat Wildl. Res. Clarendon 1980. J. Wildl. Manage. D. A. Coop. of Commerce. Colorado. with Spec. habitat in north Rep. 33. requirements, central 66pp. Utah. Utah and �343 Weeden, R. B. ptarmigan Wiens, 1964. Spatial during J. A. separation winter. 1981. Scale problems birds. , and J. T. ---- tion of avian and J. Studies Wing, L. Zar , J. H. ---- ---- Prepared numbers and the evalua- 522-532 in C. J. numbers of terrestrial Condor Ralph birds. , 1. O. Buss, and their WildI. Manage. 6~ of female blue grouse 1967. A snare Blue grouse, during the for capturing blue habitat, and populations. 15:150-169. H. Brigham. relevance 1968. Autumn movements to populations 32: 456-468. Br;ans:cad~ Inc. 32: 456-468. Congr. and J. Prentice-Hall, 75: 114-119. Bendell. Ornithol. Trans. 620pp. 1972. Int. of the blue grouse. analysis. Dispersion , and by 513- 12: 504-511. J. WildI. Manage. of blue grouse J. N.J. 1973. , and J. F. Proc. movements Biostatistical Cliffs, season. grouse. ---- Seasonal 1974. F. C. brood Estimating Censusing Pages Estimating Am. Wildl. Conf. Engelwood Zwickel, Pages in Avian BioI. 6. 1947. North censusing. eds. 1981. occupancy. eds , and willow in Avian BioI. 6. Rotenberry. M. Scott, in avian M. Scott, Studies habitat in rock Auk 81: 534-541. 521 in C. J. Ralph and J. of terrestrial of sexes Cu4-frtt=: and management. �345 APPENDICES �w .j;:- 0'1 Appendix A. Structural characteristics at random sites in 16 conifer stands. Green Mountain. Middle Park. Colorado. 1981-83. Conifer stand Occupied Variablea I (N = 20) SE ! - II (N = 10) SE ! III (N = 10) SE ! IV (N = 20) SE ! - V (N = 10) SE ! VI (N = 10) SE ! VII (N = 20) SE ! VIII (N = 10) SE ! DETR 520 63 700 109 970 163 170 31 620 66 850 98 660 85 380 85 DECO 440 49 700 109 970 163 70 15 530 47 620 104 610 88 350 90 DEOT 80 40 0 0 100 30 90 41 230 40 50 20 30 30 0 0 OT06 20 16 0 0 130 62 30 13 290 60 560 216 150 40 20 20 C006 240 65 300 121 1250 154 10 7 500 122 530 135 390 135 460 78 C01S 110 29 220 29 650 132 10 5 320 55 430 96 330 69 110 38 COB 130 30 160 60 260 65 20 9 110 28 150 27 150 37 130 33 C038 170 28 250 37 60 31 10 8 100 26 30 15 120 26 110 43 CO>39 30 11 70 26 0 0 30 10 0 0 10 10 10 7 0 0 CODBH 24 0.7 38 1.3 15 1.5 19 1.4 20 CVCODBH 1.8 OTDBH CVOTDBH CAHT DIST CVDIST 1.3 23 0.9 13 36 4.4 49 6.8 36 12 1.5 5 2.2 14 0.8 5.3 57 0.4 5.6 15 5.7 69 1.0 9 0.7 0.8 3.4 0.3 10.2 42 7.9 2.1 4.5 15 5 3.8 39 4.4 46 5.4 37 3.3 25 8.1 18 2.1 8 0.5 9 0.4 9 0.7 15 0 3 13 2.0 0 0 0.6 11 11 5.3 4 2.6 19 6.1 11 1.5 11 1.1 11 0.9 9.6 54 0.8 5.5 3.5 57 0.5 8.9 3.2 48 0.4 5.3 4.0 54 --------~------------------------~-----------------------------------,------ 0.3 4.4 6.4 62 0.7 0.9 8.8 �Appendix A. Continued. Occupied IX (N = Unoccupied 10) (N = (N 10) x SE x SE -x DETR 710 257 490 67 DECO 410 82 490 67 - Variablea - XII (N = 10) XI X = 10) SE -x 1050 201 370 910 216 370 XIII (N = 10) (N = (N 20) = 10) x - SE 102 760 126 310 35 70 31 220 95 102 740 121 260 34 60 27 140 40 80 SE -x XVI XV XIV (N = 10) SE SE x ~ SE DEOT 300 259 0 0 140 88 0 0 20 20 50 34 10 5 80 OT06 130 54 0 0 630 370 0 0 10 10 150 40 10 15 110 99 C006 390 86 360 130 860 257 120 44 320 128 220 80 90 35 90 46 C015 230 42 110 62 660 208 70 50 320 70 150 40 50 22 40 16 C023 150 50 150 34 150 40 150 45 190 57 40 22 10 8 70 21 C038 30 IS 170 63 70 26 130 33 230 56 40 22 0 0 30 15 22 30 15 20 13 0 0 30 16 0 0 0 0 CO>39 0 0 60 CODBH 15 1.0 26 1.7 18 1.9 26 1.7 CVCODBH 27 6.2 43 7.9 47 6.7 20 6.7 OTDBH 9 0.5 7 CVOTDBH 3 2.3 CAHT DIST 10 0.7 CVDIST 48 4.3 ~nemonics 0.6 5.7 18 4.8 55 1.1 0.4 8.3 19 41 1.2 21 4.4 14 1.9 18 2.1 6.2 29 7.6 3 2.1 23 7.9 0.3 10 0 9 2.5 12 0 10 0 3 1.9 0 0 2 1.4 0 0 1 0 14 1.3 3.7 42 0.7 8.2 13 6.6 44 14 0.9 0.6 4.9 3.4 \ 59 0.9 0.5 4.2 10 6.6 65 1.5 5 0.5 7 1.2 0.3 17.4 1.9 13.2 2.7 5.5 46 4.9 51 6.7 of structural variables in Table 10. w .po. .....• �w .j:'00 Appendix B. Structural characteristicsat random sites in 13 conifer stands. Whiteley Peak. Middle Park. Colorado. 1981-83. Conifer stand I (N Variablea ~ DETR DECO 960 DEOT 40 0 990 OT06 C006 C015 C023 C038 CO >39 CODBH CVCODBH OTDBH CVOTDBH CAHT DIST CVDIST 920 300 230 290 100 23 47 15 1 21 3.6 43 = 10) SE II (N = 20) III (N = 20) SE ~ Occupied IV (N = 10) SE ~ ~ SE 95 92 580 190 120 37 450 440 55 55 1350 1080 175 105 22 390 125 10 10 0 453 60 80 52 36 1.9 4.9 3.1 1.1 1.0 910 40 30 60 30 70 31 19 10 16 14 306 13 10 15 12 22 4.0 30 170 170 llO 35 44 270 400 153 130 430 500 410 170 0 17 41 128 68 64 34 0 0.7 1.3 0.9 8.2 5.5 55 5.8 0.8 3.4 1.3 2.7 5.7 90 70 24 43 7 0 12 5.4 60 38 27 20 22 2.3 5.3 0 0 0.9 2.0 6.5 9 5 15 2.7 58 VI V (N ~ 390 = 10) SE (N - ~ 520 390 85 85 0 0 0 0 450 2140 160 90 130 100 28 60 51 42 36 30 3.2 7.1 0.9 2.3 0.4 13 1.4 1.0 5.0 6.1 58 1.0 5.7 70 27 ------------------------------------------------------------------ 70 130 10 10 30 20 31 5 8 8 8 5.7 49 = 10) SE VII (N = 10) -~ SE 165 30 50 50 22 173 0 10 0 10 13 386 52 10 10 21 13 6.5 5.0 0.7 3.3 1.3 0.6 6.5 20 0 20 10 20 22 0 13 10 13 33 3 5.7 2.6 10 1.7 22.0 66 4.0 6.9 �.••. Appendix B. Continued. Occupied VIII (N :::10) Unoccupied IX (N ::: 20) X (N ::: 20) x SE x SE -x DETR 280 53 910 96 1730 DECO 280 53 820 DEOT 0 0 OT06 e006 0 0 '0 0 90 430 530 93 60 e015 40 16 e023 30 e038 eO>39 100 110 eODBH 39 4.6 18 9 1.0 eVCODBH 37 11.3 46 3.3 36 OTDBH 7 0.3 eVOTDBH 1 Variable'' eAHT 15 DIST eVDIST 7.7 59 SE XI (N = 20) -x SE XII (N = 10) SE x XIII (N = 10) SE x - 1240 95 700 101 80 29 1100 167 148 680 680 102 30 15 630 174 560 90 147 20 13 50 31 178 108 290 290 120 47 97 440 30 204 460 78 580 117 190 170 80 310 51 870 108 177 47 230 92 10 10 15 190 37 340 53 250 37 150 45 0 0 40 35 150 20 34 170 10 34 230 30 42 210 57 0 15 90 28 20 0 13 0.6 1.6 14 8~8 50 3.3 32 4.0 38 5.9 0 12 2.8 1.1 15.9 0 II 0.6 0.5 8 0.6 0.6 12 2.6 12 3.3 9 0 0 1 1.3 0.7 19 0.8 0.6 0.1 17 0.8 0.2 19 2.8 0.3 1.6 1.4 5.3 55 5.3 3.9 55 10 15.3 4.9 55 6.3 0.3 4.1 18 5 1.1 2.6 51 23 1.3 25 2.4 40 21 aMnemonics of structuralvariables in Table 10. w .p.. \0 �w VI o Appendix Structural characteristics at blue grouse winter-use sitesin 8 occupied conifer stands, Green Mountain, Middle Park. Colorado. C. 1981-83. Conifer stand I (N Variablea = 27)b SE ~ II 12) = III = 14) ~ SE ~ SE lli ~ (N (N V IV = 14)c SE (N ~ = VI lli 7) SE ~ = 3) SE lli ~ VII = 6)d SE VIII 11) SE lli = ~ DETR 500 70 410 50 770 60 260 30 630 80 540 90 350 120 670 140 DECO 500 70 410 50 770 60 170 20 490 80 370 90 350 120 660 140 DEOT 0 0 0 0 0 90 30 140 40 170 170 0 0 10 10 0 OT06 10 10 10 10 40 20 20 20 370 80 570 380 0 0 0 0 C006 160 60 320 160 940 170 0 0 140 50 70 70 170 130 650 260 C015 160 40 70 30 510 60 0 0 190 60 70 30 70 40 360 130 COB 120 30 70 20 200 40 0 0 190 90 0 0 50 30 150 60 C038 30 140 40 60 20 70 20 llO 40 270 90 170 70 100 40 CO>39 160 60' 10 120 30 0 0 100 20 0 0 30 30 70 20 50 20 CODBH 29 1.1 29 CVCODBH OTDBH 1.8 30 2.1 14 0.6 38 5.9 43 7.2 37 3.0 12 0.0 CVOTDBH CAHT 0 0.0 15 0.7 DIST 7.8 CVDIST 57 aMnemonics bN = 16 0.8 4.2 7.5 52 0.6 0.7 6.6 10 3.4 53 of structural variables In Table 10. 24 for DIST. CVDIST. 50 5.3 19 7 2.5 42 7.3 18 2.7 9 0.7 3.2 42 7.2 23 31 4.3 32 7.2 53 9.5 13 0.0 10 0.0 0 0.0 12 0.7 16 5.0 6 2.6 0 0.0 0.4 16 1.7 12 0.8 12 2.4 16 2.2 0.3 11.2 1.5 0.1 12.5 2.9 9.8 63 51 2.9 4.4 11.0 c~ dN 38 = = 0.5 7.4 4.1 84 9 for DIST. CVDIST. 5 for DIST. CVDIST. 17.9 5.7 39 2.2 0.7 3.4 �351 Appendix sites D. Structural in 3 occupied Colorado. characteristics conifer stands. of blue Whiteley Peak. grouse winter-use Middle Park. 1981-83. Conifer I (N a Variable = stand III II 36)b = 25)c (N (N = 35)d x SE x SE x SE DETR 750 60 480 50 350 40 DECO 710 60 280 30 350 40 DEOT 40 20 200 50 o o OT06 o o 570 no 150 110 C006 390 110 50 20 200 60 C015 190 30 70 20 130 40 C023 120 20 30 10 20 10 C038 260 30 100 30 80 10 CO>39 140 10 80 10 no 10 CODBH 31 1.7 33 1.9 35 2.1 CVCODBH 49 3.2 43 7.7 40 5.4 OTDBH 13 1.1 11 1.2 7 0.0 7 1.9 19 2.3 31 2.3 23 0.7 16 0.6 14 0.7 CVOTDBH CAHT 4.5 DIST CVDIST variables bN 3.4 39 aMnemonics in Table = 0.2 of structural 10. 34 for DIST. CVDIST. 5.8 0.4 39 3.3 c = 23 for dN = 31 for N 7.0 0.4 54 4.0 DIST. CVDIST. DIST. CVDIST. �w Vl N Appendix conifer E. Dbh , height. stands. Green Dbh and age of trees Mountain. used Middle Park. by blue during winter in 8 occupied 1981-83. Colorado. Height (em) grouse (m) Age (years) Min x - SE N Max Min Stand x - SE N - Max Min x - I 37 2.4 27 61 10 15 0.7 27 23 5 182 19.7 19 374 47 II 39 1.7 10 51 29 16 0.6 10 19 13 212 34.0 8 340 96 III 21 1.3 1'3 32 14 10 0.4 13 14 8 63 2.8 9 75 51 IV 50 5.3 14 94 27 15 1.9 14 32 5 268 32.0 2 300 236 V 25 2.6 7 33 14 11 0.6 7 14 10 68 1.4 7 72 61 VI 32 3.0 3 35 29 11 3.2 3 17 6 VII 39 6.3 5 63 28 15 2.7 5 26 11 VIII 36 5.1 11 62 12 11 0.8 11 15 7 139 25.0 10 327 42 SE -N Max �F. Appendix conifer Dbh , height. stands. Whiteley Dbh and age of trees Peak. Middle Park. used by blue Height during winter in 3 occupied 1981-83. Colorado. (cm) grouse (m) Age (years) Max Min x SE N - Max Min x - 33 91 25 21 0.8 33 33 13 180 13.1 14 256 94 3.0 22 79 25 16 0.6 22 21 8 153 73.8 11 284 66 2.6 34 80 16 13 0.8 34 21 3 315 40.0 22 652 57 Stand x - SE I 48 3.1 II 47 III 45 N SE N - Max Min W l./1 W �Appendix G. w Habitat characteristicsat blue grouse winter-use sites for males, females, and both VI ~ sexes, Middle Park, Colorado, 1981-83. Green Moun tain Females (N=38) Both (N=13) SE x SE x Variable'' Males (N=6) x SE DETR DECO 350 350 131 131 500 480 55 57 DEOT OT06 0 0 0 0 20 30 10 13 100 C006 450 C015 150 391 131 300 180 99 47 270 140 C023 20 17 21 80 C038 CO>39 100 26 90 130 38 46 23 110 40 80 34 40 17 80 32 32 17 1 13 90 31 51 13 82 24 80 80 3.1 34.7 0 29 54 10 12.7 0.3 0 1.3 1.1 7 15 3.3 1.1 4 18 6.6 29 0.8 4.2 4.9 32 CODBH CVCODBH OTDBH 5.6 11.5 CVOTDBH CAHT 13 DIST CVDIST 11.9 61 1.9 2.1 10.9 15 7.0 54 2.4 4.6 2.9 0.8 0.6 0.6 3.9 460 420 40 17 7.4 63 aMnemonics of habitat variables in Table 10. Males (N=9) Whiteley Peak Females (N=20) x - SE -x - SE 33 33 600 470 .105 100 670 570 81 86 38 100 130 100 48 490 69 153 130 III 110 48 250 112 430 130 60 24 110 28 210 22 4.2 120 32 41 5.5 10 Both (N=66) SE x 490 430 44 41 18 65 60 240 82 190 140 27 34 110 23 36 42 19, 50 150 13 2.1 120 18 10 1.6 5.8 35 38 0.9 2.2 12 2 1.2 1.0 1.1 19 0.7 0.4 2.1 6.2 35 0.3 1.6 3.3 �355 'JOB PROGRESS REPORT State of Colorado Project 01-03-045 (W-37-R) Work Plan Job Title: 9: Job Personnel: 8 Food Selection and Nutritional Ecology of Blue Grouse During Winter Period Covered: Author: Avian Research 01 July 1984 through 30 June 1985 T. E. Remington C. E. Braun, R. W. Hoffman, T. E. Remington, T. J. Schoenberg, Colorado Division of Wildlife ABSTRACT Winter food habits and preferences of blue grouse (Dendragapus obscurus) were investigated in Middle Park, Colorado. Radio-marked birds fed exclusively on needles and buds; tree species fed-upon depended on stand composition but consisted primarily of Douglas-fir (Pseudotsuga menziesii) or lodgepole pine (Pinus contorta). Limber pine (Pinus flexilis) was fed-upon occasionally. Feeding was not observed within Engelmann spruce (Picea engelmannii) or subalpine fir (Abies lasiocarpa). Most feeding within Douglas-fir and lodgepole pine (90 and 98% of all observations, respectively) was in the upper or middle canopy and on 1-3 year-old needles (90 arid 63%, respectively). Consumption of needles of both species declined progressively with needle age (R <0.05). Preferences of captive birds for 5 conifer species were ranked as Douglas-fir> lodgepole pine > limber pine> Engelmann spruce = subalpine fir. Captive birds ate more (p < 0.05) needles from branches from "old" (> 75 years) Douglas-fir trees than-from "young" « 75 years) trees and more (P < 0.05) I and 2 year-old needles than 3 and "4 or 5 and 6 year-old needi~s. Preference for needles from old trees was reversed (p < 0.05)· after the monoterpene content of needles from old and young trees was reduced (by heating), and restored when monoterpenes were added back. Palatable needles (those from old Douglas-fir, lodgepole pine, and limber pine) were lower in monoterpenes or benzoic acid than unpalatable needles (those from young Douglas-fir, subalpine fir, and Engelmann spruce). Nutritional quality (measured as crude protein and cell contents) was not consistently related to palatability. Preferences for younger needles may be related to nutritional quality as 1 and 2 year-old needles contained more (p < 0.05) crude protein than other needles. Blue grouse consumed more (p < 0.05) of a pelle ted ration treated with baseline (level in fed-upon Douglas~fir) levels of 3 monoterpenes (camphene, limonene, bornyl acetate) than food treated with 2 or 4 times baseline levels. This was not true of food treated with benzoic acid. �357 FOOD SELECTION AND NUTRITIONAL ECOLOGY OF BLUE GROUSE DURING WINTER Thomas E. Remington D The quality and quantity of winter food resources is generally recognized as a critical factor in the distribution and/or productivity of grouse populations (Gullion 1966; Miller et ale 1966, 1971; Moss 1969; Moss et ale 1975; Watson et al. 1977; Watson and O'Hare 1979; Beckerton and Middleton 1982). Few reports exist of food habits of blue grouse during the winter (Beer 1943, Stewart 1944, Marshall 1946, Hoffmann 1961, Boag and Kiceniuk 1968). Food preferences of blue grouse and the nutritional quality of their diets are unknown during winter. Data on winter food habits, food selection, and food quality of blue grouse are lacking from Colorado. Two current theories of herbivore food selection may pertain to blue grouse. Blue grouse may selectively feed to maximize nitrogen intake (Mattson 1980), or to minimize their intake of plant defense compounds which may inhibit their ability to digest foods or otherwise reduce fitness (Freeland and Janzen 1974, Bryant and Kuropat 1980). Blue grouse must contend with a winter diet that is both low in protein and high in fiber, and which also contains digestibility-reducing compounds that may make significant amounts of nutrients unavailable. The overall objective of this study is to test the hypothesis that during winter, blue grouse selectively feed from among available food items to maximize digestible nitrogen and/or energy. P.N. OBJECTIVES The objectives of this study are to investigate (1) winter food habits, and (2) winter food preferences of blue grouse, and (3) measure the nutritional quality (protein, cell contents) and anti-quality components (terpenoids, phenolic resins, neutral detergent fiber [NDF], acid detergent fiber [ADF]) of blue grouse winter foods and their relationship to diet preferences. Specific objectives are to: 1. Identify winter foods of blue grouse. 2. Investigate blue grouse winter use of conifers for food by species, tree ages, and growth forms in relation to their availability. 3. Quantify tree species composition and· physical characteristics grouse winter feeding sites. 4. Measure protein and NDF, ADF content of conifer needles from trees fed upon by blue grouse and from randomly-located trees. of blue �358 5. Measure tannin, phenolic resin, and mono-, di-, and sesquiterpene levels in conifer needles from trees fed upon by blue grouse and from random1ylocated trees. 6. Rank blue grouse preference for Douglas-fir, subalpine fir, lodgepole pine, limber pine, and Engelmann spruce as winter foods. 7. Measure protein, NDF, ADF, and total digestibility by blue grouse of randomly-selected needles of Douglas-fir, lodgepole pine, limber pine, subalpine fir, and Engelmann spruce. 8. Investigate the deterrent effects of specific, phenolic resins, and mono-, di-, and sesquiterpenes to browsing by blue grouse and to blue grouse digestibility of conifer needles. SEGMENT OBJECTIVES 1. Review literature pertaining to nutrition, physiology, winter ecology, and captive maintenance of grouse; chemical and nutritional characteristics of conifers; and laboratory techniques applicable to measuring nutrient or secondary plant constituent levels. 2. Capture and radiomark 8 blue grouse (preferably 3 adults and 1 juvenile of each sex). 3. Relocate radio-marked birds location of feeding sites. 4. Measure height, dbh, and age and determine species of each fed-upon tree. 5. Collect needle samples from fed-upon trees for analysis of nitrogen, NDF, ADF, terpenoid, and phenolic resin levels. 6. Collect needle samples from randomly located trees for nitrogen, NDF, ADF, terpenoid, and phenolic resin levels. 7. Collect 2 grouse/week from January through March for nutritional (protein, fiber, minerals) and compositional analysis of crop contents, and body composition. 8. Measure nitrogen, NDF, ADF, terpenoid, and phenolic resin levels of needle samples from random and fed-upon conifers, and of needle ages 1-6 of Douglas-fir. 9. Capture 4 grouse (preferably digestibility trials. for 3 observation females, 1 of feeding male) for behavior analysis preference and of and a �359 10. Repeat trials to rank preferences of blue grouse for 5 conifer species. 11. Repeat trials to rank preferences of blue grouse for ages of Douglas-fir needles. 12. Conduct trials to rank digestibility (total, nitrogen) of 2 preferred (Douglas-fir, lodgepole pine) and 2 non-preferred (Engelmann spruce, subalpine fir) conifers. 13. Compile data, analyze results, and prepare a progress report covering the 2nd year's activities. DESCRIPTION OF STUDY AREAS Whiteley Peak and adjacent Burnt Mountain, approximately 30 km west of Kremmling in Grand County, Colorado, were the primary study areas. In addition, 2 blue grouse originally radiomarked on Green Mountain were relocated periodically on Blue Ridge and above Guthrie Gulch for observations of feeding activity. Vegetative and physical characteristics of Whiteley Peak have been described (Cade 1985). Vegetative cover on Burnt Mountain consists primarily of homogenous associations of lodgepole pine/subalpine fir or quaking aspen (Populus tremu1oides)/sagebrush (Artemisia spp.). Blue Ridge and Guthrie Gulch are both dominated by lodgepole pine/subalpine fir associations. Isolated pockets of mature Douglas-fir and limber pine are present along the top of Blue Ridge. METHODS Most observations of winter feeding activity were made by locating 14 radio-marked blue grouse and observing them and/or other birds associated with them. Solar (Wildlife Materials, Inc., Carbondale, Ill.) or lithium battery-powered (Telonics, Inc., Mesa, Ariz.) transmitters were attached to vinyl ponchos which slipped over the head of the birds (Amstrup 1980). Weight of the transmitter and harness was about 20 and 30 gms for solar and battery-powered units, respectively. Grouse were captured with a telescoping noose pole (Zwicke1 and Bendell 1967). Data collected at feeding sites included tree species fed upon, pOSition within tree where feeding occurred, and flock composition. Fed-upon trees were marked with orange flagging and numbered aluminum tags. Tree dbh and heights were measured with a dbh tape and clinometer, respectively. Tree ages were estimated by counting annual growth rings from cores obtained by increment borer. Needle samples were collected from browsed areas within fed-upon trees. The age of needles eaten by blue grouse was determined by counting bud scales back from the terminal bud on each branch or branch1et that was fed upon. �360 Preference trials were conducted with 3 wild-captured adult males and 1 juvenile female. The 1st preference trial (tree species) was preceded by an acclimation period of 8, 6, 7, and 5 days for birds 1-4, respectively. Birds were offered branches of each of the 5 conifer species tested during this acclimation period. Branches were changed daily. During the trial, pair wise combinations of Douglas-fir, lodgepole pine, limber pine, Engelmann spruce, and subalpine fir were presented to the birds. The order of treatments (pairs of conifer species) was randomized; birds were then randomly assigned to treatment schedules. The randomization was restricted so that each treatment was replicated once in the morning (0700 to 1200 MST) and once in the afternoon (1315 to 1815 MST). Each treatment was thus replicated 8 times, twice with each bird. Branches of the 2 species tested were alternated and wired to 2 pegboards which were hung on the sides of the cage. Treatment response was the amount of each species consumed (gms), determined by weighing each branch before and after a replicate and subtracting spillage from the difference. Spillage was collected on aluminum trays that slid under the cages. Initially, 1,000 + 50 gms of each species were offered to insure ad libitum quantities. By the-3rd day of the trial it was determined that 600 + 50 gms were adequate for the morning replicate and 750 + 50 gms were adequate for the afternoon replicate. Controls were weighed during the first 2 days, but there were no detectable weight losses due to dessication. Branches used in the trials were gathered from the middle canopy of 5 randomly-located "mature" trees of each species. Needle age preference trials were conducted by presenting 3 containers containing ad libitum (100 gms = male, 75 gms = female) quantities of 1 and 2, 3 and 4, and 5 and 6 year-old Douglas-fir needles to the birds twice daily. Containers were attached to 1 side of the cage in a random order. Needles were removed from branches collected from mid-canopy of randomly located mature trees. Timing of the 2 daily replicates was altered to 0700 to 1445 and 1545 to 1830 MST to reduce the disparity in intake between morning and afternoon replicates. Tree age preference trials were conducted in a similar fashion to species preference trials. Branches collected from 3 "old" and 3 "young" Douglas-fir trees were paired. Pairs were collected from the same area to eliminate site variation. Young and old tree pairs were 27 and 137, 72 and 593, and 27 and 75 years old. The ability of blue grouse to identify and select needles from older trees without structural or visual cues associated with the branches was tested by removing 1-4 year old needles from branches collected from "old" and "young" Douglas-fir trees and presenting both to the birds. Needles were removed from the branches by gently agitating the twigs under liquid nitrogen. Methodology was similar to that used in the needle age preference trial. Beakers were randomly positioned for the A.M. replicate and reversed for the P.M. replicate. Needles were placed in identical 400 ml Pyrex beakers. Monoterpene content in needles was reduced by heating both sets of needles on metal trays at 40C for 8 hours. Because this treatment altered the needles in other ways, the treatment was reversed by adding monoterpenes back to the needles. This was done by steam distilling (14 hours) green needles in a volatile oil distillation apparatus and adding the distilled oil by micro-pipette back to an equivalent amount of heat-treated needles. �361 The ability of blue grouse to detect and avoid individual monoterpenes was evaluated by adding 3 levels of camphene, limonene, bornyl acetate, and benzoic acid to Purina game bird maintenance chow and measuring blue grouse consumption. The 3 levels were: baseline, equivalent to the level of that compound in fed-upon Douglas-fir, 2 times baseline, and 4 times baseline. The benzoic acid was dissolved in alcohol and 2 mls of each of the 3 concentrations applied to 100 g of game bird chow by micro-pipette. The alcohol was then allowed to evaporate. The 3 monoterpenes were added directly by micro-pipette to the game bird chow which was then kept frozen until used. Experimental design used was identical to those used for needle trials. Trials were conducted to measure total metabolizable energy (TME) and total metabolizable nitrogen (TMN) of needles of 2 palatable and 2 unpalatable species. The birds were given ad libitum quantities of needles of a given spec Les for 4 days. Droppings were cleared and collected each morning and evening for analysis. Birds were weighed (+ 1 gram) each morning prior to food presentation. Order of treatment (species) presentation was randomized and birds randomly assigned to treatment schedules. Nitrogen, cell-contents, NDF, ADF, and monoterpene content were measured as indicators of nutritional quality of needle samples. Needles were removed from the branches by gentle agitation after immersion in liquid nitrogen or storage in a super cold (-70e) freezer. Samples were then ground under liquid nitrogen. Samples for nutritional analysis were dried at 100e overnight and ground again with a mortar and pestle. Only 1 and 2 year-old needles were used for analysis. Monoterpene levels were determined using a modification of Welch and McArthur (1981). Ten-gram needle samples (wet weight) were extracted for 8 hours in a Soxhlet apparatus with 50 ml of diethyl anhydrous ether as solvent. The volume of extract was reduced below 8 ml with reduced pressure and an internal standard of carvone (2.5 mg/ml) was added. The extract was transferred to a 10-ml volumetric flask and the volume brought to 10 ml by adding ether. Monoterpenes were quantified by injecting 2 g of the extract into a Perkin Elmer Sigma 2 coupled to an IBl-19000 Data Station. A 30 m by 0.25 mm DB-l capillary column was used. Temperature programming specifications used are in Table 1. Peaks were identified using a VGMM l6F mass spectrometer and comparing mass spectra obtained to published spectra. Table 1. Gas monoterpenes. chromatographic parameters Initial oven temperature Initial time Programmed temperature rate changes: At 2.0 min. At 90 e Final oven temperature Hold at final temp Flow rate Injection port temperature Flame ionization detector temperature used to separate and quantify 70 e 2.0 min 2.0 c/min 10.0 c/min 250 e 10 min 30 ml/min of N2 275 e 275 e �362 Nitrogen was measured by micro-kje1dah1 analysis (Horwitz 1980) and protein estimated by multiplying nitrogen by 6.25. Needle samples were partitioned into cell contents (CC), NDF, and ADF following Van Soest (1963a, b; 1967). ADF was extracted from the NDF residue as suggested by Mould and Robbins (1981) and Van Soest (1982). Lignin was partitioned from ADF (Van Soest 1963a,b; 1967). All nutritional analyses were performed in duplicate and are expressed on a dry matter (DM) basis. Dry matters were obtained by drying ground samples at 100C for 12 hours. Statistical Analyses Much of the data in this report is preliminary and incomplete; many tests of significance have not been completed. Means and standard deviations are given so that group differences can be evaluated. Selection for needles of different ages was evaluated by comparing the percentage of each age class (1-6, from fed-upon trees) browsed by blue grouse using Hote11ings T2 test. This test tests for differences across all groups. If differences were found across groups, pairwise t tests were used to find which groups differed. ANOVA was used to test for differences in intake of different needle ages by blue grouse in captivity. Duncan's multiple range test was used to determine which needle age groups differed in consumption. ANOVAwas also used to evaluate the ability of blue grouse to discriminate among 3 levels of monoterpenes added to a pe11eted ration. Paired t tests were used to test whether consumption of needles of 5 conifer speCies presented in pairwise combinations differed. Paired t tests were also used to test whether consumption of needles from old vs. young Douglas-fir differed before and after heat treatment, and after steam-distilled oils were added. RESULTS ANDDISCUSSION One hundred and thirty-three feeding observations of 14 radio-marked blue grouse were recorded. Species fed-upon depended on availability (stand composition), but most feeding occurred in Douglas-fir, lodgepole pine or limber pine (Table 2). Significant feeding (> 30 sec) within Engelmann spruce or subalpine fir was not observed. Based on a subjective comparison of use to availability, blue grouse preferences were ranked as Douglas-fir> lodgepole pine > limber pine > Engelmann spruce and subalpine fir. For instance, 2 radio-marked birds overwintered (with at least 8 other birds) in a stand that contained only limber pine yet limber pine was rarely used in stands where Douglas-fir or lodgepole pine were present. Blue grouse use of Douglas-fir has been documented by Beer (1943), Stewart (1944), Marshall (1946), and Cade (1985). Feeding on lodgepole pine has been documented by Boag and Kiceniuk (1968). Ninety and 98% of the birds observed feeding within Douglas-fir and lodgepole pine, respectively, were in the upper or middle canopy (Table 3). Boag and Kiceniuk (1968) noted that spruce grouse (Dendragapus canadensis) and/or blue grouse concentrate browsing within mid-canopy of lodgepole pine. Hoffmann (1961) found that blue grouse fed exclusively within the upper portion of white fir (Abies conco10r). Capercai11ie (Tetrao uroga11us) preferentially fed within the upper (male) or middle (female) canopy of Scots pine (Pinus sy1vestris) (Linden 1984). �363 Table 2. Tree species fed upon (number of observations) by radio-marked grouse, Middle Park, Colorado, November-April, 1983-84, 1984-85. Species Bird Ii SA 2+ 2+ 1+ 2+ 2+ 1+ 1+ 112+ 2+ 12+ 2+ 130 171 008 156 168 165 099 150 140 148 013 175 144 008 Totals c fed uEon Pif1 blue a Psme! Pico b Psme Pico Totals 11 6 18 0 0 8 1 0 10 10 0 6 0 6 0 0 0 16 4 0 5 13 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 5 0 8 0 0 0 0 0 0 3 0 0 1 0 0 0 0 0 11 6 20 16 4 11 6 13 11 10 5 6 8 6 76 38 15 4 133 Stand comEosition Psme/Ab1a/Pif1/Pien/Pico Psme/Pif1 Psme/Pif1 Pico/Ab1a Pico/Ab1a Psme/Ab1a/Pico/Pif1/Pien Pico/Ab1a/Pif1/Psme Pico/Ab1a Psme/Ab1a/Pico/Pif1/Pien Psme/Pif1 Pif1 (100%) Psme/Ab1a/Pif1/Pien/Pico Pif1 (100%) Psme/Pif1/Ab1a apsme = Douglas-fir, Pico = lodgepole pine, Pif1 = limber pine, Ab1a = suba1~ine fir, Pien = Engelmann spruce. Fed upon both species during 1 feeding period. cSex and age of bird. Table 3. Tree positions (number of observations) Middle Park, Colorado, November 1983-Apri1 1984. Species Psme Pico a Lower 1/3 of feeding blue grouse in Middle 1/3 11 1 47 22 UEEer 1/3 51 23 apsme = Douglas-fir, Pico = lodgepole pine. No differences in feeding behavior were apparent among sex and age-classes of blue grouse. Birds of mixed sex and age classes were frequently observed within the same tree. �364 Structural characteristics of fed-upon trees varied within and among stands, but birds usually fed in the largest trees available (Table 4). Fed-upon Douglas-fir trees averaged 237 years old, although the range of ages was large (55-600 yrs). Only 8 of 53 (15%) fed-upon Douglas firs for which ages were obtained were less than 100 years old (Fig. 1). Most of these were in a young, mixed conifer stand on the north side of Whiteley Peak where older . trees were absent. Fed-upon lodgepole pine and limber pine were much younger, averaging 78 and 103 years, respectively (Table 4). Ellison (1976) noted that spruce grouse avoided feeding in saplings of white spruce (Picea glauca). Boag and Kiceniuk (1968) observed that spruce and/or blue grouse fed almost exclusively within "older" (> 15 years) trees. Cade (1985) found that blue grouse "use trees (trees used for feeding and/or roosting) on Whiteley Peak were larger (dbh; !< 0.05) than random trees within occupied stands. tl Table 4. Height (m), dbh (cm), and age of trees fed Middle Park, Colorado, November-April, 1983-84, 1984-85. Descriptive statistic x SD Range N Ht Douslas-fir Dbh Age 15.9 5.2 9-31 36 49.3 17.3 12-90 59 237 138 55-600 52 Lodse:eole :eine Ht Dbh Age 15.2 3.2 5-20 28 28.0 8.2 9-49 28 78 15 27-111 26 upon by blue Ht grouse, Limber pine Dbh Age 40.1 17.4 24-82 10 103 67 55-253 10 Blue grouse use and selection of needles by age-class were investigated by counting browsed and unbrowsed Douglas-fir and lodgepole pine needles from 1 to 6 year old. Needle age-classes were defined as to 1 to 6 years for simplici ty. Current-year growth needles described as "1" year-old needles were from 5 to 10 months old over the period of study; "2" year-old needles were 17 to 22 months, etc. Six years was chosen as the cutoff point because needles of both species are generally persistent to 6 years (Harlow and Harrar 1969), and no browsing was detected on older needles. One thousand needles were counted from each fed-upon tree. The percentage of needles of each age-class browsed by blue grouse was used to test for selection. Blue grouse selectively fed among age-classes of both Douglas-fir and lodgepole pine (Hotellings .!?, ! < 0.001). Consumption of needles of both species declined progressively with needle age. This was most apparent within Douglas-fir (Fig. 2). Each age-class within Douglas-fir was preferred (higher percentage browsed) to those older except needle ages 5 and 6 were equally non-preferred (pairwise t tests, P < 0.05). Within lodgepole pine, 1 and 2 year-old needles were preferred -( P < 0.05) to ages 4-6; 3 year-old needles were preferred to ages 5-6; and 5 year-old needles were preferred to 6 year-old needles (Fig. 2). �LODGEPOLE PINE 65 @ 60 \I (/) DOUGLA::I-fIK -- >-' u z r-- ttl Z - c::: .•.. • o !z r-- t-IS8 258 358 25 ~ 20 - S8 35 30 LL 2 • •• 50 o r- w 55 (/) 45 ~ 40 w 6- , 3: ffi o r-- 458 I I naI sse LODGEPOLE PINE ~ 15 10 ffi 5 a. ~ (/) 65 60 3:: 55 0 >- I O. V/A ~ u, Z c ttl w' 0::: II. 0 • a: w • II 38 ,. a. 71 fa III TREE AGE CLASS Fig. 1. Distribution of ages of Douglas-fir and lodgepole pine browsed by blue grouse, Middle Park, Colorado, 1983, 1984. v«.! vaa 56 40 35 30 - 25 ~ 20 cC ••. 15 Z ~ 10 z V//A - L&J :::t c' ""La DOUGLAS-FIR r--- ~ 50 (/) 45 u' Z V//A 23.4 r--- 5 0 1 2 3 4 nr=:l 5 6 NEEDLE AGE (YEARS) Fig. 2. Percentage of needle age classes 1-6 of Douglas-fir and lodgepole pine browsed by blue grouse, Middle Park, Colorado, 1983, 19~4. W 0\ VI �366 Ellison (1976) found that during winter spruce grouse browsed age-classes of white spruce needles in proportion to their availability; no selection was evident. One year-old needles of lodgepole pine were the age-class "most heavily used" by spruce grouse and/or blue grouse in Alberta (Boag and Kiceniuk 1968:28). The net effect of this selection is that 1-3 year-old needles comprise the bulk of needles consumed from both Douglas-fir (90%) and lodgepole pine (63%) (Table 5). These estimates are minimal since usually only branches containing needles of all (1-6) age-classes were selected for counting. Thus, use of 1-3 year-old needles on branches less than 6 years old wouldn't have been counted. Table 5. Percentage of 1 to 6 year-old needles browsed by blue grouse from Douglas-fir (Na = 19) and lodgepole pine (Na = 22) in relation to total needles browsed, November 1983-April 1984. - x SD 1 2 44 17 32 10 Douglas-fir Needle ase 3 4 14 7 6 6 5 6 1 3 4 1 2 20 8 Lodge~ole ~ine Needle age 2 3 4 24 9 19 6 aN = number of trees for which needles were counted. 1,000 needles were counted/tree. 18 7 5 6 12 6 7 5 Approximately Apparent selection for certain species and age-classes of conifers and for needles of certain ages (Tables 1, 3-4) was tested in trials using captive birds. By presenting ad libitum quantities of each treatment to captive birds, problems of measuring and interpreting use and availability in the field were eliminated. Selection for certain species, ages or growth forms of trees on the basis of perching ability, thermal cover or other factors related to feeding efficiency rather than chemical properties of needles was eliminated in captivity. Blue grouse responded well to captivity and the experimental designs used. Differences of biological significance were detected in 9 of 10 comparisons of conifer species (Table 6). Because of the small number of birds tested (with correspondingly low degrees of freedom) and multiplicity of t tests (requiring a probability of error of 0.01 to test significance at '0.1), statistical significance (~< 0.1) was achieved in only 3 of 10 comparisons. Species preference trials will be repeated next year with 4 more birds. The additional degrees of freedom and reduced variability due to larger sample sizes should (if preferences remain similar) make 9 of 10 comparisons statistically significant. �367 Table 6. Mean total consumption and mean difference in grouse of needles from conifer species offered in pairs. Pairb Psme/Alba Psme/Pien Psme/Pif1 Psme/Pico Pico/Ab1a Pico/Pien Pico/Pifl Pien/Ab1a Pien/Pifl Ab1a/Pifl Total consumed (sm) x 100 106 102 105 102 104 104 88 104 101 Difference s= 118d 100d 67 41 90 93d 74 3 -50 -46 consumption by blue between pair members (§m) SD 10 15 14 18 17 15 23 10 23 11 aCorrected for morning, afternoon differential (morning intake f [morning intake/afternoon intake]). bPsme = Douglas-fir, Ab1a = subalpine fir, Pi en = Engelmann spruce, Pif1 = limber pine, Pico = lodgepole pine. cSD = SD IN, N = 8. dDifference between pairs significant at P < 0.1 (tested at P < 0.01 to adjust for mUltiple [10] ! tests). Douglas-fir was preferred (R_ < 0.1) over both subalpine fir and Engelmann spruce, and lodgepole pine was preferred over Engelmann spruce. A defendable ranking of species preferences is Douglas-fir> lodgepole pine> limber pine> Engelmann spruce = subalpine fir. There seemed to be no real difference in preference between the 2 least preferred species, subalpine fir and Engelmann spruce. Except for this pair, all species in the above ranking were apparently preferred to those ranked below it. Since all 5 species were offered to all 4 birds, once in a morning trial and once in an afternoon trial, the total amount of each species consumed can be used as a check of this ranking. The amount consumed was 2,492, 2,350, 1,493, 927, and 909 gms of Douglas-fir, lodgepole pine, limber pine, Engelmann spruce, and subalpine fir, respectively. This ranking compares well to what radio-marked birds fed upon in the wild. All radio-marked birds subsisted primarily on the 3 most preferred species, and avoided the 2 least preferred species entirely. Selection for larger, older trees (or avoidance of smaller, younger trees) was tested by offering birds equal, ad libitum amounts of needles from "old" or "young" trees by alternating branches from each within cages. Preference for older trees was confirmed (p < 0.05) (Fig. 3). Needles from "old" trees were consumed in greater quantities in 21 of 24 replicates. Selection may have been even greater than indicated. In all 3 replicates where consumption of needles from young trees exceeded consumption of needles from older trees, browsing was concentrated on terminal or lateral leaders of branches (from young trees) exhibiting extremely rapid growth. It may be that grouse are selecting needles at a finer level of resolution than just tree age. �w 0.6 0.4 0 0.3 60 l- r--1 0.2 551- I I 0.1 50 l- I I OC w 45 ~ 0.5 ::::E 0.4 40 >- ~ 0.3 35 "#. 0.2 w ~ :;:) 0 Gl E Q. Gl 011 e: .9! o >- .Q t:. EI e: Gl e: ii 1!I 011 ;, 0 0 V) 2 >- ~ z w ~ I- a: .. 25 c( CJ n 20 151- I I I I 10 l- I I I I 5t- I I I I 0 "OLD" DOUGLAS-FIR "YOUNG" DOUGLAS- FIR z 0 ~ I 8 ~ E3 ~ 0.3 0.5 0.4 ~ 0.1 0 Fig. 3. Amount (x, gm) of needles consumed by 4 captive blue-grouse from branches of "old" and "young" Douglas-fir trees, March, 1984. z S m 0.5 tfi 0 o 0 z z -I :::0 - (I 0.5 -< .»:!l: o =l m ,2.0~ LIMBER r· ~ 5 0.3 0.2 :!l: 1.5 _ "#. 1.00 Gl - I - II o r» 2.0 ~ e: e3 S m QI a: ~ 0.2 ~ 0.1 0 ::::E 0 I ~ jO.5 1.0 ~ -- z -11.0 1.5 LODGEPOLE PINE 0.5 w 0.4 2.0 -11.5 2.00 - I- 30 00 011 DOUGLASFIR - 0.1 J2.5 e: V) z '" I~ Ii ~I ~J I I I e: e: 011 QI .t: e: c. ._ 0.5 65 r- 011 SUBALPINE FIR 1.0 I- I - § ~ - II -iO.5 "'0 Fig. 4. Monoterpene content of 1 and 2 year-old needles from randomly located trees of subalpine fir, Douglas-fir, lodgepole pine, and limber pine, Middle Park, Colorado, December-March, 1983-84. �369 While it is apparent that blue grouse selectively feed on younger needles, especially within Douglas-fir (Fig. 2, Table 5), it is not clear whether this reflects preference. For instance, it's possible that blue grouse move to the tips of branches to eat buds, and then feed on needles in the vicinity of the twig tip. Buds of Douglas-fir contain substantially lower levels of monoterpenes than needles (Von Rudloff 1971). It is also unclear how blue grouse identify palatable needles to feed upon. Preference for needle ages was tested by randomly ordering 3 containers containing ad libitum quantities of 1 and 2, 3 and 4, and 5 and 6 year-old needles within each cage. Blue grouse strongly preferred (F = 450, 2 and 23 df, P < 0.001) 1 and 2 year-old needles (Table 7). Table 7. Mean intake (gm) of 1 and 2, 3 and 4, and 5 and 6 year-old Douglas-fir needles during 6 trials with 4 captive blue grouse (~ = 24), 11arch 1984. Statistic 1 and 2 X SD 72 7 Needle age 3 and 4 3a 3 5 and 6 6 5 aMeans sharing the same letter do not differ (p > 0.05, Duncan Multiple Range Test). There was no difference (1: > 0.05) in intake between 3 and 4, and 5 and 6 year-old needles. It is significa~t not only that blue grouse preferred 1 and 2 year-old needles but also that they were able to identify them when they were removed from the branches. There were no discernible differences in needle size, color, or texture. This is a strong indication that blue grouse use chemical cues to identify palatable needles. Reasons for Food Selection Blue grouse preferences for different species and ages of conifers, and for younger needles were hypothesized to be related to selection for nutritional quality (high nitrogen and cell contents, and low NDF, ADF) or avoidance of monoterpenes. Species preferences were not clearly related to nutritional quality (Table 8). For instance, Douglas-fir was most preferred yet contained significantly less crude protein than 3 of the other 4 species. Lodgepole pine was second only to Douglas-fir in preference yet contained the least cell contents and most indigestible ADF and lignin. �370 Table 8. Nutritional characteristics of randomly located conifer needles (1 and 2 years old) collected during winter, 1983-84. Mean ± S.D. expressed as % DM. Speciesa Psme Pico PHI Pien Abla Crude protein 6.0 7.3 6.9 5.6 6.7 ± ± ± ± 0.4 0.8 0.7 0.7 ± 0.7 Cell contents 51.0 38.8 44.9 42.5 47.7 ± ± ± ± 3.0 4.4 3.3 2.0 ± 3.0 NDF 48.9 61.2 55.1 57.5 52.3 ± ± ± ± ± ADF 3.0 4.4 3.3 2.0 3.0 36.8 47.0 39.7 40.6 36.4 ± 3.7 ± 3.1 ± 3.2 ± 1.5 ± 2.4 Lignin 19.6 27.2 19.9 19.7 20.7 ± ± ± ± 3.0 4.2 3.6 1.0 ± 1.9 apsme = Douglas-fir, Pico = lodgepole pine, Pifl = limber pine, Pien = Engelmann spruce, Abla = subalpine fir. There were substantial differences in monoterpene content among species that may relate to preferences (Fig. 4). Subalpine fir contains twice the monoterpene content of Douglas-fir. Lodgepole pine and limber pine contain few monoterpenes relative to Douglas-fir (ranked above them), so if monoterpenes are involved in preferences some level must be acceptable. Engelmann spruce is non-preferred yet it contains only 1 volatile constituent in measurable quantities, benzoic acid, a terpene-like organic acid. Qualitative differences in toxicity of monoterpenes may be important in selection. Preference for needles from older Douglas-fir was not correlated to selection for nutrients as older Douglas-fir contained less (p < 0.05) crude protein and cell contents than needles from younger Douglas-fir (Table 9). Needles from mature, randomly-located Douglas-fir were similar (p > 0.05) in protein and cell contents to same age needles from fed-upon Douglas-fir, indicating no selection on this basis for individual trees. Selection may be related to avoidance of monoterpenes. Needles from younger Douglas-fir contained about twice the monoterpene content of needles from older Douglas-fir; a level similar to the level in (unpalatable) subalpine fir needles (Fig. 5). Needles from random (mature) Douglas-fir trees were similar in monoterpene composition to needles from fed-upon trees, indicating no selection on this basis for individual trees within Douglas-fir (Fig. 5). �DOUGLAS-FIR NEEDLE SAMPLES ,2.5 (& = MATURE FED-UPON 190, RANGE = 74-347) 2.0 Gl Gl r:: Gl s: r:: Gl _ O.2~ .~ a: w 0.1 ~ 0 2 t: § ::::2: >- a: 0.5 Gl Q CD r:: (ij CD r:: E r:: a:: '" o •• I CD CD r:: CD s;r:: .... ~ r:: Gl r:: Gl 0 >::; ~ 0 E :.:::i ~ a;0 ]i 0 I- DOUGLAS-FIR NEEDLE SAMPLES 1.5 c( II I b 1.0 f:: 0.5 3:: 0 MATURE RANDOM ~ = 163, RANGE = 65-348) 00.4 0 z Sm 2.0 1.5 ~ ~ I ~ I:E:I a I m Z 1.0 m o 0.50 ~ 0 w z ~ 0.8 YOUNG (8,= 37, RANGE = 22-72) z -I m z -I ~ a: 0 :::0 -< ~ 0.7 2.5 ~ 0.6 0 ::::2: 0.5 0.3 0.2 0.1 0 3:: :l> 2.0 ~ m 1.5~ 0.4 ~I ~ 1.0 [3 I 0.5' -I 0.5 0.4 r:: CD r:: Gl -.0.3 r:: Gl W 0.1 ::::2: r:: a::• E r:: '" o Gl r:: a:: .... ~ >a: o 0.5 '#. -:- 0.4 ~ ~ 0.3 ~ z 0.2 0 o 0.1 w z 0 w a. a: 0.5 3 ~ I CD r:: Gl r:: CD e>. ::; sa + e:3 + 0 E :.:::i ~ E CD 0 ]i 0 I- ~ 1.5 11 I c( 1.0 0 a I I 0.5 0 6 YEARS 2.0 1.5 0.1 0 ~ I ~ c::::I ~ z 0 -I m I o 1.0 0 0.4 0.3 ::::2: 0.2 f:: 2.0 ~ m 1.5 ~ W b b -I 0.5 ~ 4 YEARS I~ 5 .,2.0 2 YEARS Ii. !i Gl Gl a: ~ + 1 Gl z ~ z -I '#. 0 :::0 -< 3:: :l> 1.0 ~ m 0.5 ~ ~ -'0 0 Fig. 5. Monoterpene content of 1 and 2 year-old needles from mature fed-upon, mature random, and young Douglas-fir trees, Middle Park, Colorado, December-March, 1983-84. Fig. 6. Monoterpene content of 1 and 2, 3 and 4, and 5 and 6 year-old needles of·Douglas-fir, Middle Park, Colorado, December-March, 1983-84. w .•...• ~ �372 Table 9. Nutritional characteristics of needles from young, mature random, and mature fed-upon Douglas-fir. (:! ± S.D., % DM.) Needle Age GrouE Young Random Fed-upon 1&2 1&2 1&2 3&4 5&6 Crude Erotein 6.8 6.0 5.9 6.1 5.3 ± ± ± ± ± 0.6 0.4 0.6 0.7 0.3 Cell contents 52.7 51.0 49.8 54.0 55.1 ± ± ± ± ± 1.6 3.0 1.6 1.9 2.0 NDF 47.2 48.9 50.2 46.0 44.9 ± ± ± ± ± ADF 1.6 3.0 1.6 1.9 2.0 35.7 36.8 38.2 34.8 34.0 ± ± ± ± ± DM Li~in 1.3 3.7 1.0 1.4 1.7 20.3 19.6 21.1 18.9 18.3 ± ± ± ± ± 1.2 3.0 1.2 1.4 1.0 0.452 0.445 0.476 0.5013 0.508 Selection for younger needles within Douglas-fir may be related to selection for crude protein. Protein content of 1 and 2 year old needles was higher (p < 0.05) than the protein content of 5 and 6 year old needles, although sImilar to 3 and 4 year old needles. Interestingly, cell-contents were lower and NDF, ADF, and lignin were higher in younger needles (Table 9). This is in conflict with the hypothesis that younger needles are selected for because of higher nutritional quality (digestible energy). It is possible that the higher cell-content value of older needles is an artifact of a higher phenolic or tannin content, both of which are soluble in neutral detergent (Mould and Robbins 1981). Lowry (1970) and Ellison (1976) reported that nitrogen (and crude protein) content of black (Picea mariana) and white spruce declined with needle age. Higher nitrogen content in young needles can result from translocation of nitrogen from older to younger needles, especially in conifers growing on nutrient deficient sites (Gosz 1981). Monoterpene composition was constant across needle ages (Fig. 6), and was probably not related to selection. The ability of blue grouse to detect and avoid monoterpenes was investigated experimentally by reducing monoterpene levels in needles and evaluating preferences, and by adding specific monoterpenes to a pelleted ration and measuring consumption. It was hypothesized that if blue grouse can select for nutritional quality and detect and avoid monoterpenes, then relative preferences for needles from young and old Douglas-fir should be reversed if monoterpene levels in needles from young trees are reduced below a tolerance threshold. It is significant that even after needles were removed from the branches, the birds still consumed more (p < 0.05) needles from old trees than from young trees. After heat treatment,-preferences were reversed (p < 0.05) for 3 of 4 birds (Fig. 7). Heating caused other changes in the needles besides reducing monoterpene levels (reduced moisture content, altered color). Because of this, monoterpenes (obtained by steam distillation of fresh needles) were added back to the heat-treated needles and the trial repeated. The 3 surviving grouse again preferred (p < 0.05) old needles (Fig. 7). It appears that blue grouse can assess the monoterpene content of needles and avoid high levels. Selection of needles from young trees after heat treatment may indicate that blue grouse can also assess the nutritional value (protein, cell contents vs. NDF, ADF) of needles and make choices on this basis. Sage grouse (Centrocercus uroEhasianus) selectively fed among and within sagebrush (Artemisia spp.) taxa to maximize protein and minimize monoterpene intake (Remington and Braun 1985). �50 Ii 40 ffi ~ w 30 fa 20 Hl z 10 15 o w ::i::J \-18 ~ 1-15 o ~ i-12 h»"L" o J: •.... ······f 0"'9 YOUNG o : [6 50 50 (BIRD 4) HEAT TREATED (BIRDS 1·3) 1['40 z: w 40 ~ ~ m o !;; CJ) ~ 30 30 ~ 20 m 20 ~ m w o , o 10 9 ['-0-'-'-'-, ""h'" .- t·········.. I o 0.25 0.50 0.75 1.00 1.25 1.50 LIMON ENE 10 '»»>' , 3 <!) 0 0.25 0.500.751.001.251.50 24 z o' BORNYL ACETATE ~~21 CJ) oI ttlz I _r24 CAMPHENE 0 ~t21 I I I ::!: 18 ::J YOUNG BENZOIC ACID CJ) ~ 1-15 0 VOLATILE OILS ADDED Ii 40 ~ 1-12 0 z J: w ~ 30 fa 20 w ..J 01..9 o ~t 6 0 W ~ 10 0 ~ o OLD YOUNG Fig. 7. Amount Q[, gm) of needles from old and young Douglas-fir trees,eaten by blue grouse before and after heat treatment and after addition of volatile oils, February 1985. o 0.25 0.50 0.75 1.00 1.25 1.50 CONCENTRATION (gil OOgDM) 3 o 0 0.25 0.50 0.75 1.00 1.25 1.50 CONCENTRATION (gil OOgDM) Fig. 8. Consumption ex, gm) by 4 captive blue grouse of a pe11eted ration treated with 3 levels of monoterpenes. w -...j w �374 It appears that monoterpenes within young Douglas-fir (and presumably subalpine fir) inhibit feeding by blue grouse. It was hypothesized that 1 or a few individual monoterpenes were responsible for most of the feeding inhibition. Three monoterpenes (camphene, 1imonene, borny1 acetate) and benzoic acid were selected as likely feeding deterrents because they were in relatively low quantity or absent in palatable needles (those from old Douglas-fir, and lodgepole and limber pine) and in relatively high quantities in unpalatable needles (those from young Douglas-fir, subalpine fir, and Engelmann spruce). Blue grouse ate less (p < 0.05) of the 2 times and 4 times baseline-teated game bird chow than of-the baseline treatment of all 3 monoterpenes (Fig. 8). Consumption was inversely related (p < 0.05) to monoterpene concentration. Camphene was the only compound to -have a large effect on consumption. Whether this was due to qualitative differences among monoterpenes or to the larger baseline quantity of camphene is not known. Consumption of game bird chow treated with benzoic acid showed the same trend as monoterpene-treated game bird chow but differences among treatment levels were not significant (p = 0.20). If deterrent effects of these monoterpenes are additive, then monoterpene levels may explain the low palatability of young Douglas-fir and subalpine fir. It does not appear likely that benzoic acid levels within Engelmann spruce are the cause of its low palatability. Trials were conducted to measure total metabolizable energy (TME) of palatable (Douglas-fir and lodgepole pine) and unpalatable (Engelmann spruce and subalpine fir) needles. These trials were not entirely successful because the extent of weight loss resulting from 4 days exposure to unpalatable foods was not anticipated. Three of 4 birds died prior to completion of the trials. Information gained from laboratory analyses, when completed, should allow an evaluation of the methodology used and suggest alternative designs for next winter. Weight losses were greatest when birds were fed unpalatable species (-40 g/day on Engelmann spruce, -21 g/day on subalpine fir) and lowest for the palatable species (-4 g/day on Douglas-fir, -19 g/day on lodgepole pine). Greater weight losses on unpalatable species were at least partially due to decreased intake, as birds ate an average of 0.20 gm per gm body weight of the 2 palatable species daily and only 0.15 and 0.18 gm per gm body weight of Engelmann spruce and subalpine fir, respectively. LITERATURE CITED Amstrup, S. C. 214-217. 1980. A radio-collar for game birds. Beckerton, P. R., and A. L. A. Middleton. levels on ruffed grouse reproduction. Beer, J. R. 1943. 1982. J. Wild1. Manage. 44: Effects of dietary protein J. Wi1d1. Manage. 46:569-579. Food habits of the blue grouse. J. Wildl. Manage. 7:32-44. Boag, D. A., and J. W. Kiceniuk. 1968. Protein and caloric content of lodgepole pine needles. For. Chron. 44:28-31. Bryant, J. P., and P. J. Kuropat. 1980. Selection of winter forage by subarctic browsing vertebrates: the role of plant chemistry. Annu. Rev. Ecol. and Syst. 11:261-286. �375 Cade, B. S. 1985. Winter habitat preferences and migration patterns of blue grouse in Middle Park, Colorado. M.S. Thesis, Colorado State Univ., Fort Collins. 101pp. Ellison, L. 1976. Winter food selection by Alaskan spruce grouse. Manage. 40:205-213. J. Wildl. Freeland, W. J., and D. H. Janzen. 1974. Strategies in herbivory by mammals: the role of plant secondary compounds. Am. Nat. 108:269-289. Gosz, J. R. 1981. 405-426. Nitrogen cycling in coniferous ecosystems. Ecol. Bull. 33: Gullion, G. W. 1966. A viewpoint concerning the significance of studies of game bird food habits. Condor 68:372-376. Harlow, W. M., and E. S. Harrar. 1969. Textbook of dendrology. McGraw-Hill Book Co., New York, N.Y. 5l2pp. 5th ed. Hoffmann, R. S. 1961. The quality of the winter food of blue grouse. Wildl. Manage. 25:209-210. Horwitz, W., Editor. 1980. of official analytical D.C. 1018pp. J. Official methods of analysis of the association chemists. Assoc. Off. Anal. Chem., Washington, Hrivnak, J., M. Mahdalik, E. Vaadiova, and L. Sojak. 1973. Analysis of monoterpenes from the needles of Pinus sylvestris and Picea excelsa using capillary gas chromatography. Holzforschung and Holzverwertung 25:24-26. Linden, H. 1984. The role of energy and resin contents in the selective feeding of pine needles by the capercaillie. Annu. Zool. Fenn. 21:435-439. Lowry, G. L. 1970. Variations in nutrients of black spruce needles. Pages 235-259 in C. T. Youngbert and C. B. Davey, eds. Tree growth and forest soils. Oregon State Univ. Press, Corvalis. Marshall, W. H. 1946. Cover preverences, seasonal movements and food habits of Richardson's grouse and ruffed grouse in southern Idaho. Wilson Bull. 58:42-52. Mattson, W. J., Jr. 1980. Herbivory in relation to plant nitrogen content. Annu. Rev. Ecol. and Syst. 11:119-161. Miller, G. R., D. Jenkins, and A. Watson. 1966. Heather performance and red J. grouse populations. I. Visual estimates of heather performance. Appl. Ecol. 3:313-326. , A. Watson, and D. Jenkins. ---to experimental improvement 10:323-335. 1971. Responses of red grouse populations of their food. Symp. Br. Ecol. Soc. �376 Moss, R. 1969. A comparison of red grouse (Lagopus 1agopus scoticus) stocks with the production and nutritive value of heather (Ca11una vulgaris). J. Anim. Eco1. 38:103-112. _______ , A. Watson, and R. Parr. 1975. Maternal nutrition and breeding J. Anim. success in red grouse (Lagopus 1agopus scoticus). 44:233-244. Eco1. Mould, E. D., and C. T. Robbins. 1981. Evaluation of detergent analysis in estimating nutritional value of browse. J. Wi1d1. Manage. 45:937-947. Remington, T. E., and C. E. Braun. 1985. Sage grouse food selection in winter, North Park, Colorado. J. Wi1d1. Manage. 49:1055-1061. Stewart, R. E. 1944. Food habits of the blue grouse. Condor 46:112-120. Van Soest, P. J. 1963a. Use of detergents in the analysis of fibrous feeds. I. Preparation of-fiber residues of low nitrogen content. J. Assoc. Off. Agric. Chem. 46:825-829. 1963b. Use of detergents in the analysis of fibrous feeds. II. A rapid method for the determination of fiber and lignin. J. Assoc. Off. Agric. Chem. 46:829-835. 1967. Use of detergents in the analysis of fibrous feeds. IV. Determination of plant cell wall constituents. J. Assoc. Off. Agric. Chem. 50:50-55. 1982. Nutritional ecology of the ruminant. Corvallis, Oreg. 374pp. 0 and B Books, Inc., Von Rudloff, E. 1971. Chemosystematic studies in the genus Pseudotsuga. I. Leaf oil analysis of the coastal and Rocky Mountain varieties of the Douglas-fir. Can. J. Bot. 50:1025-1040. Watson, A., and P. J. O'Hare. 1979. treated and untreated Irish bog. Red grouse populations on experimentally J. App1. Eco1. 16:433-452. , R. Moss, -----on red grouse J. Phillips, and R. Parr. 1977. The effect of fertilizers stocks on Scottish moors grazed by sheep, cattle and deer. Pages 193-212 in P. Pesson, compiler. Eco10gie du petit gibier et amenagement deschasses. Gauthier-Ni11ars, Paris, France. Welch, B. L., and E. D. McArthur. 1981. Variation of monoterpenoid content among subspecies and accessions of Artemisia tridentata grown in a uniform garden. J. Range Manage. 34:380-384. Zwicke1, F. C., and J. F. Bende11. J. Wi1d1. Manage. 31:202-204. 1967. A snare for capturing blue grouse. �377 Prepared bYTh~m~q-Graduate Research Assistant �� https://cpw.cvlcollections.org/files/original/715c8f9e90748305c9b2c84dee2c8bcf.pdf d256e57459656226c2cee9ea2a013b7b PDF Text Text Colorado Divis~on of Wildlife Wildlife Research Report April 1986 379 JOB PROGRESS REPORT State of Colorado Project: 01-00-045 (W-37-R) Work Plan Job Title: 12: Personnel: 15 Chronology of Breeding and Nesting Activities of Wild Turkeys in Relation to Timing of Spring Hunting Seasons Period Covered: Author: Job Avian Research 1 July 1984 through 30 June 1985 Richard W. Hoffman C. E. Braun, J. Grimes, R. W. Hoffman, R. L. Holder, T. J. Spezze, and R. D. Velarde, Colorado Division of Wildlife; JoAnn Profera, Colorado State University. ABSTRACT A detailed study plan was prepared in 1983-84. Efforts to initiate this study were delayed in 1984-85 due to: (1) lack of support for implementation of a permit system and wing collection program, and (2) failure to incorporate the proposed program into the regulations. Rejection of the proposal was based on the inconvenience it would create for the hunter despite the contention that, because of the growing interest in pursuing wild turkeys, hunters would be willing to cooperate regardless of the inconvenience. A survey was designed and mailed to all 1984 spring turkey hunters to identify their spring hunting activities and to poll their opinions about the permit system and wing collection program. The regulations were revised to include special and limited permits. All holders of 1985 limited permits were mailed a wing envelope and subsequently surveyed immediately following the season to measure their cooperation in returning the wing envelope. ��381 CHRONOLOGY OF BREEDING AND NESTING ACTIVITIES OF WILD TURKEYS IN RELATION TO TIMING OF SPRING HUNTING SEASONS Past research efforts on Merriam's Wild Turkey (Me1eagris &a110pavo merriami) in Colorado and elsewhere throughout its range have been inadequate and poorly designed. These earlier studies were mostly descriptive in nature and none was designed to test specific hypotheses. The traditional approach to managing turkeys in Colorado has been through trapping and transplanting when, ironically, minimal knowledge exists concerning distribution, habitat preferences, population levels, production rates, survival rates, and harvest levels. Baseline data about these and other population attributes are essential for identifying problems, formulating testable hypotheses, and developing management strategies. P.N. OBJECTIVES 1. Document the timing of winter flock dispersal, onset of gobbling, peaks of gobbling, nest initiation, onset of incubation, and peak of hatch in relation to geographic area, year, and timing of the spring season. 2. Describe the gonadal cycle of females and compare the reproductive condition of females in relation to timing of the spring season. 3. Measure the abandonment rate of incubating human disturbance around the nest. 4. Monitor hunter activity and harvest of wild turkeys on a statewide basis. females to varying levels of SEGMENT OBJECTIVES 1. Review literature pertinent to the objectives of this study. 2. Conduct a survey of 1984 spring hunters to determine their willingness cooperate in an experimental permit system and mail wing survey. 3. Select 3 study areas, 1 each in the NE, SE and SW Regions, where adequate numbers (100+) of turkeys exist for trapping, marking, and monitoring purposes. If funding constraints restrict the study to one area, then the SE Region has top priority. 4. Using drop nets and cannon nets, trap and individually mark 100+ birds on each study area. In addition, 40 birds, including 6 males and 34 females, will be equipped with tail-mounted radio transmitters on the study area in the SE Region. 5. Document timing of winter flock break-up. 6. Document onset of gobbling and peaks of gobbling activity. to �382 7. Document onset of egg laying and nest initiation and peak of hatch. 8. Measure the effects of human disturbance on the rate of nest abandonment. 9. Collect 2 hens/week starting the 1st week of April and continuing through the 4th week of May to document the reproductive condition of females. 10. Implement a hunter questionnaire-wing collection program using a permit system-mail wing survey. 11. Compile data, analyze results and prepare a progress report. RESULTS AND DISCUSSION Only segment objectives 1, 2, 10, and 11 were accomplished in 1984-85 as no field work was conducted. Efforts were focused on gathering data on hunter activities, opinions, and participation that would demonstrate the need and support for this study. 1984 Spring Turkey Hunter Questionnaire At the January 1984 Regulations Review meeting a decision was made not to approve a proposal by Avian Research to implement a permit system and wing collection program for wild turkeys. Rejection of the proposal was based on the apparent inconvenience it would create for the hunter; i.e., the hunter would have to obtain a permit in addition to purchasing a license. Our contention was that because of the growing interest in pursuing wild turkey, hunters would be willing to cooperate. A survey was approved to investigate turkey hunter attitudes. The survey (Appendix A) was designed and mailed to all spring turkey hunters to identify their spring hunting activities and to learn their preference regarding the permit system and wing collection program. Table 1. Spring 1984 turkey hunter survey, Colorado. Mailing Surveys mailed Surveys returned Percent returned Non-deliverable 1st 2nd Totals 3,515 1,824 52 126 1,691 623 37 23 3,515a 2,447 70 149 aTota1 license holders sampled. holders (3,518). This represents 99.9% of all license �383 Table 2. Colorado spring 1984 turkey hunter questionnaire data. Projected for Mailing 2nd 1st !! in sample % of total license holders N hunters l' hunters !! not hunting % not hunting N hunters observing turkeys % hunters observing turkeysb N successful hunters % successful huntersb N hunter days Days/hunter N turkeys harvested !! turkeys lost Total kill % crippling loss 1,824 52 1,542 85 282 15 990 64 313 20 5,243 3.4 313 64 377 17 623 18 489 78 134 - 22 289 59 66 13 1,467 3.0 66 19 85 22 2,447 70 2,031 83 416 17 1,279 63 379 19 6,710 3.3 379 83 462 18 1,Ona 30 835 78 236 22 493 59 109 13 2,505 3.0 109 32 141 22 Projected for 3,518 3,518 100 2,867 81 652 19 1,772 62 488 17 9,215 3.2 488 115 603 19 aNon-respondents. bBased only on those license holders who actually hunted. Questionnaires were sent to 3,515 turkey license holders on 23 July 1984 of which 52% were returned by 17 August 1984. On 18 August 1984 a follow-up questionnaire was sent to all non-respondents and 623 additional responses were received. In total, 2,447 (70%) license holders responded (Table 1). Mean values calculated for the follow-up questionnaire were used to project for the 30% (1,071) non-respondents (Table 2). The projected estimates indicate that 2,867 hunters (81% of all license holders hunted) harvested 488 turkeys during the 1984 spring season for a success rate of 17% (Table 2). Reported crippling loss (19%) was extremely high suggesting hunters shoot before birds are within range « 40 yards). Spring hunters averaged 3.2 days of hunting over the course of the 23-day season (21 Apr-13 May). Nineteen percent of the hunters who purchased licenses did not hunt; 75% indicated they also intended to hunt during the fall season. Based on the distribution of harvest by units (Tables 3 and 4), east slope areas accounted for 94% of the harvest with 88% from the southeast portion of the state. �384 Table 3. Wild turkey harvest by county, Colorado, Spring 1984. Harvest County N Las Animas Huerfano Fremont Custer Pueblo Baca El Paso Archuleta Bent Mesa Otero Yuma Larimer Boulder Douglas Teller Clear Creek Jefferson La Plata Costilla Alamosa Saguache Park Kit Carson Crowley Totals 93 63 41 34 31 27 10 9 9 8 8 7 6 6 6 5 4 3 2 2 1 1 1 1 1 379 25 17 11 9 8 7 3 2 2 2 2 2 2 2 2 1 1 <1 <1 <1 <1 <1 <1 <1 <1 100 �385 Table 4. 1984. Wild turkey harvest by small game management unit, Colorado, Spring Harvest Unit 84 79 80 68 70 78 76 81 88 82 32 42 50 56 30 86 83 90 40 58 52 72 10 N % 177 31 30 24 20 13 11 10 9 7 7 6 6 6 47 8 8 6 5 3 3 3 2 2 2 2 2 2 1 1 1 <1 <1 <1 <1 <1 <1 100 5 Totals 4 4 2 2 2 1 1 1 379 Although the greatest pressure and harvest occurred opening weekend (Table 5), turkey hunters did not exhibit the "opening weekend syndrome" common with other upland game bird hunters. The spring harvest appears to be more a function of the timing of breeding activities of turkeys than the number of hunters in the field. After opening weekend, the harvest did not decrease substantially over the remainder of the season. In fact, the success rate increased, suggesting the birds were still responsive to calls when the season ended. Since the peak of hatch probably does not occur until early June, 1-2 weeks of additional hunting may be biologically justifiable. Only 20% of the turkeys harvested were not in flocks (Table 6). Most were taken from flocks comprised of 1-5 birds. However, 37% of the harvested turkeys came from flocks with 6 or more birds. These flocks probably included hens. There was a greater tendency for hunters to encounter larger flocks earlier in the season. This supports the belief that incubation was not underway when the season opened. Spring seasons should be timed to coincide with laying and incubation to insure minimal disturbance of hens and maximum harvest of males. �386 Table 5. Distribution spring turkey season. of hunting pressure and harvest by time period, 1984 Hunter dai:s Time Eeriod N 1st 1st 2nd 2nd 3rd 3rd 4th 931 492 811 283 611 363 488 weekend week weekend week weekend week weekend Totals Harvest % N 24 12 21 7 16 9 102 59 57 27 32 24 41 11 3,939 ~ 30 17 17 8 9 7 12 342 Table 6. Size of flocks from which turkeys were harvested during the 1984 spring season in Colorado. N % Flock size o 1-5 6-10 10 68 20 147 43 70 20 60 17 Hunters indicated support for a permit system and wing collection program (Table 7). Few hunters complained about the permits only being available by mail or at Division of lVildlife offices. Table 7. Willingness of spring turkey hunters to cooperate in a permit system and wing collection program. Response Yes (%) Permit system Wing collection program 1,961 (83) 2,164 (90) N (%) Totals 399 (17) 233 (10) 2,360 2,397 �387 Proposed Changes in Turkey Regulations In January 1985, the survey results were presented to the Colorado Division of Wildlife Management Team. The data were ignored and another dilemma surfaced regarding the regulations. It was observed that implementation of the permit system would require a change in the regulations because there was already a section (#1205) in the regulations dealing with permits (i.e., limited permits for Lake Dorothey, Bonny Reservoir, etc.). The question arose as to how the CDOW could impose a special research permit in addition to an existing limited permit. Since the 1985 turkey regulations had already been approved by the Commission, the Management Team decided it was too late to modify the 1985 regulations. Research was instructed to prepare a draft regulation to accommodate both limited and special permits. The following regulations comply with this assignment. The changes only apply to section #1205. CHAPTER 12 - WILD TURKEY ARTICLE I - GENERAL PROVISIONS Statutory Reference 33-1-106 #1205 - PERMITS, APPLICATIONS, AND DRAWING PROCEDURES. In addition to a valid license, persons hunting turkeys in Colorado must obtain one of two types of permits: 1. Colorado Special Wild Turkey Hunting Season Permit Unlimited in number and free of charge. Special permits will be available on a walk-in basis at all Colorado Division of Wildlife Regional Offices, and area offices .in Pueblo and Durango. In addition, special permits can be obtained by mail application to the Denver Headquarters of the Colorado Division of Wildlife, 6060 Broadway, Denver, Colorado 80216. Application forms will be available at all Division of Wildlife offices and at all license agents who sell turkey licenses. To obtain a special permit, persons must first obtain a valid turkey license. The special permit and license must be in the possession of the hunter at all times while hunting turkeys. Special permits shall be valid only for the season indicated on the permit. a. Special Spring Permit - Available from March 1, 1985 through the end of the spring season (May 19, 1985) and valid only for the spring season (April 20 through May 19, 1985). b. Special Fall Permit - Available August 1, 1985 through the end of the fall season (October 6, 1985) and valid only for the fall season (September 21 through October 6, 1985). �388 2. Colorado Limited Wild Turkey Hunting Season Permit - Limited in number, free of charge, and available only through application. Holders of a valid limited permit need not obtain a special permit. Limited permits shall be valid only in the area and for the time period indicated on the permit (refer to Section #1209). a. b. Applications 1. No person shall submit application per season. more than one 2. Group applications will be limited to four (4) people per group. 3. Incomplete applications will not be accepted. 4. Applications will be accepted only on appropriate application form provided by Division. 5. Applications will be available at all Division of Wildlife Offices. 6. Persons need not possess a valid turkey license to apply for a limited permit. (1) the the Drawings 1. 2. Spring Season (a) Applications for limited permits must be received at the Denver Headquarters of the Colorado Division of Wildlife, 6060 Broadway, Denver, Colorado, 80216 by 8: 00 a.m. March 26, 1985. (b) All limited permits will be issued at a public drawing on March 29, 1985 at the Denver Headquarters of the Colorado Division of Wildlife. Limited permits remaining after the public drawing will be issued on a first-come, first-served basis. Fall Season (a) Applications for limited permits must be received at the Denver Headquarters of the Colorado Division of Wildlife, 6060 Broadway, Denver, Colorado, 80216 by 8:00 a.m. August 27, 1985. (b) All limited permits will be issued at a public drawing on August 30, 1985, at the Denver Headquarters of the Division of Wildlife. Limited permits remaining after the public drawings will be issued on a first-come. first-served basis. �389 1985 Survey of Special Turkey Permit Holders As a pilot study designed to measure hunter participation in a mail wing survey, wing envelopes were distributed to the 140 applicants successful in drawing a 1985 limited turkey permit (Table 8). The wing envelope and permit were mailed to all successful applicants on 29 March 1985. Instructions on the envelope requested that if the hunter harvested a turkey he was to (1) remove the least damaged wing, (2) remove 3 or 4 breast feathers, (3) place the wing and breast feathers in the envelope, (4) include only the wing and breast feathers from the bird they harvested, (5) fill in the date, location (county, SGMU, nearest town), and time when the bird was harvested and his name and address, and (6) mail the envelope as soon as possible. Table 8. Limited turkey permits, Spring 1985, Colorado. Area Lake Dorothey Area 6 Area 42 Area 84 Totals N permits N applications 1st Choice 2nd 3rd 75 25 30 10 140 281 192 122 97 629 75 25 30 10 140 -0-0-0-0-0- -0-0-0-0-0- Questionnaires (Appendix B) were sent of which 106 (76%) were returned by conducted, thus, no attempt was made 106 respondents, 100 (94%) purchased hunted, indicating that hunters did going hunting. The success rate was hunters returned their wing envelope. from successful hunters who failed harvest by permit area was: Unit Unit Unit Unit Success (%) 27 13 25 10 22 to the 140 permit holders on 28 May 1985 14 July 1985. No followup survey was to project for non-respondents. Of the a turkey license and 98 (92%) actually not apply without strong intentions of 33% (32/98); 24 of 32 (75%) successful In addition, 3 envelopes were received to respond to the questionnaire. The 6 (South Platte) 3 84 (Pueblo Reservoir) 4 42 (Republican) 10 80 (Lake Dorothey) 15 Based on examination of the breast feathers, one of the harvested female. It's possible this bird was bearded and therefore envelopes did not contain breast feathers in addition to a contained flank feathers rather than breast feathers. Only 12 birds was a legal. Two wing and 2 of 27 wings �390 examined could be positively assigned an age (9 yearlings, 3 adults) because the key feathers (primaries 9 and 10) for distinguishing age classes were worn off as a result of dragging the wing tips during courtship activities. This problem was more pronounced for adults than yearlings. It is probable that the 15 unclassified wings were from adults. Thus, the harvest was probably composed of 18 adults (67%) and 9 yearlings (33%). �391 APPENDIX A STATE OF COLORAOO Rich.rot D. Umm. Goy.mor DEPARTMENT OF NATURAL RESOURCES DIVISION OF WILDLIFE Wildlife Research Center 317 West Prospect Road Fort Collins, CO 80526 J.m •• B. Ruch. Director &oeo Bro.ctw.y Den •••••• Color8do 8021 IS1297·1 1921 Dear Wild Turkey Hunter: Colorado sportsmen have shown a growing interest in pursuing wild turkeys as hunter participation and harvest have more than doubled over the past decade. By obtaining a license to hunt wild turkey i~ Colorado, you have indicated an interest in the future of this sport. Please assist US in enhancing our management efforts on wild turkevs by completing the questionnaire below and returning it to US in the enclosed self-addressed envelope. Information you supply will be instrumental in establishing future seasons and directing management strategies. Please read the questionnaire carefully and f111 in all requested information completely and accurately. If you did not hunt turkeys during the spring season we ask that you complete Questions 1, 7, 8 and 9 and return the questionnaire. If you did hunt, ans~~r all the questions. Please report only on your bunting activities ..~~A fo, <he 19" sp<iog •• asen, ~>.rh\ ,-~D~~:O: ______________- SPR I NG TURKEV 1. Did you hant turkeys during the 1984 spring season1 If no, proc:ee:J to e•.• estion 7 and complet3 the remainder 2.. Did l. Old you """'est If yes, the Sr."~11 9-- tM.;.-::aer 1-5 -ti NoD que!itlonnaire. vesD NoD unit number vesD NoD • coul'lty • ~nd the turkey ~s harvested 'inCi'""Checkin the appropriate box ci ctl'!er turkeys present ., the time yoy Iwrv~s[e.1 ywr turkey. (!!!!~~.!.!.:.!) o the a lurkev during the 1984 spring se.son? :JIlt.5. give date below t:-:e of SURVEY I:urkeys during the 198ft spring seasonl ct:se.rye yaY HARVEST _ I'I.or. 6-I~ than 10 DDDD S.. Circle the r " d~tes on ..nich you hunted turkeys during the T \I T n 25 26 27 28 2) 2' IlAY 3D 06 07 01 02 0) O~ OS 0' 10 " 05 12 spring season. spring seAson7 21 APRil 2 1984 S 13 ~id you hit Ho.t tU"Y ""keys 7. Do"",.plan on ""ntin9 turkeys durln9 the 1984 f.1I s• .,on? I. Would you be w.llin9 to cooper.te. In an expe.rlmentA' survey next year by obt~inl"9 a Mndatory perr.ait in ~dition to the required IIcense7 Permits would be .vailable by mail and.t Division of Vi Jd. ife off ices •• nd unlimi ted In number and free of charge .. ,. \Jould you b. _iJlin9 ",ith • ;:.-ostac;e-paid but not to cooperate wing en".lop.7 in. retrieve wing during collection the 1981t 6. pr~ra", HoD NoD D NoD ve.D next year If provided vesD v•• DEPARTMENT OF NATURAL RESOURCES, DaVid H. Gelclles. E.eculive Doreclor.WILDLIFE COMMISSION. James C. Kennedy. Ch.inn.n TomOilly \Y Schultz. Vice Chaorman.Mochael K. Higbee. Secrelary.Richard L. Divelbiss. Member.Donald A. Fem.ndez. Member Wilbur L. Redden. Memoer.James T. Smilh. Member.Jean K. Too~ Member �392 �393 STATE OF COLORADO Rlcheret D. umm. G_mor DEPARTMENT OF NATURAL RESOURCES DIVISION OF WILDLIFE Wildlife Research Center 317 West Prospect Road Fort Collins, CO 80526 Jem •• B. Ruch. DI_ 60150 Broedw.., Den_. Colo •• do 802111 (297·11921 Dear Wild Turkey Hunter: We recently sent you a questionnaire pertaining to your hunting activities during the 1984 spring wild turkey season. So far we have not received your reply. Please assist us in enhancing our management efforts on wild turkey by completing the questionnaire below and returning it to us in the enclosed self-addressed envelope. Information you supply, even though you mav not have participated, will be instrumental in establishing future seasons and irecting management strategies. Thank you for your cooperatio SPRING TURKEY HARVEST SURVEY 1. Did you hunt turkeys during the 198' sprln9 'Season? If no, proceed to question 7 .nd complet~ the r~inder of 2. Old you observe turkeys l. Did you ""rvest a turkey during the 198~ spring season? II. If yes, pleAse give the sr,yll 9~e d,ate telow the other the "t,,;~oer of during the turkeY turkeys 19S•• spring unit questionn.ire. -ri NoD yesD HoD yesD HoD season? number .•• 5 Nrvested present at the • county •• nd in"d'Cneck in the ilPj)rooriate box the t ir.\e 't~ h~rye5ted your turkey. (~~~~) D 1-5 6-10 IIore than 10 DODD S. Circle the eaees ~APRIL In hunted turkeys T F S 2' 25 2' 27 21 07 01 02 OJ C, 08 09 10 " I] 6. How Qny 7. Do dur ing the 1518,. spring season. :U ]0 MY 06 you V " 2) on w,lch T turkeys did you hit but OS 12 not I retrieve·durittg you pl.n on hunting turkel'S during the 198~ fall the 198" spring season? seasonl 'esD NoD to cooper.,. In ,an expa:rimenull survey next ye.r by obt.lnlng •••• ndatory s:er::lit in ,addition to the reQuired Ilcense1 Permits would be ,ay.ilable by Nil ,aftd ,at Oivi'Sion of Wildlife offices. ,and unlimited In number ,and free of ch.rge. yesD HoD \:ould you be willing ",ith ,a post.ge-p,aid Ye.D NoD I. Would you be willing ,. to cooper.ta win9 envelop.1 in a wing co)Jectloft prognlla next year If provided DEPARTMENT OF NATURAL RESOURCES. David H. Getches. Executive Director.WILDLIFE COMMISSION. James C. Kennedy. Chairman Timothy W. Schullz. Vice Chaorman.Michael K. Higbee. Secrelary.Richard L. Divelbiss. Member.Oonald A. Femandez, Member Wilbur L. Redden. Member.James T. Sm.th. Member.Jean K. Tool. Member �W \0 .f:'- �395 APPENDIX STATE OF COLORADO Richard D. Lamm, Governor DEPARTMENT OF NATURAL RESOURCES DIVISION OF WILDLIFE James B. Ruch, Director 6060 Broadway B Wildlife Research 317 West Prospect Fort Collins, CO Center Road 80526 Denver, Colorado 80216 Telephone: (303) 297-1192 Dear Wild Turkey Hunter: Colorado sportsmen have shown a growing interest in pursuing wild turkeys as hunter participation and harvest have more than doubled over the past decade. By obtaining a special permit to hunt wild turkey in Colorado, you have indicated an interest in the future of this sport. Please assist us in enhancing our management efforts on wild turkeys by completing the questionnaire below and returning it to us in the enclosed self-addressed envelope. Information you supply will be instrumental in establishing future seasons and directing management strategies. Please read the questionnaire carefully and fill in all requested information completely and accurately. If you did not hunt turkeys during the spring season we ask that you complete Questions 1 and 2 and return the questionnaire. If you did hunt, answer all the questions. Please report only on your hunting activities for the 1985 spring season. S:CZ:lt1f #, ~Chard Wildlife 1. Did you purchase a license 1985 spring season? I . w. HOffm~ Researcher to hunt turkeys during the 2. Did you hunt turkeys during the 1985 spring season? 3. Did you harvest 4. If yes, please give the small game unit number , county , and date the turkey was harvested. (Map on reverse side.) 5. Did you return a wing in the special envelope a turkey during the 1985 spring season? provided Yes No Yes No Yes No Yes No . DEPARTMENT OF NATURAL RESOURCES, David H. Getches. Executive Director. WILDLIFE COMMISSION, Timothy W. Schultz, Oharrman ' James T. Smith, Vice Chairman • Richard Divelbiss. Secretary • Donald A Fernandez, Member. Rebecc·a L Frank. Member Robert L Friedenberger, Member • John Lay, Member. George VanDenBera. Member �W \0 0\ �397 Colorado Division of Wildlife Wildlife Research Report April 1986 JOB PROGRESS REPORT Colorado State of Project Work Plan Job Title: 01-00-045 (W-37-R) 12: Job Avian Research 16 Nesting Parameters, Nest Site Selection. and Reproductive Performance of Rio Grande Wild Turkeys in Lowland Riparian Habitats Period Covered: 01 July 1984 through 30 June 1985 Author: R. W. Hoffman Personnel: C. E. Braun, R. W. Hoffman, Colorado Division of Wildlife; S. S. Rosenstock, Colorado State University ABSTRACT Efforts to initiate this study in 1984-85 were delayed due to lack of financial support and reassignment of the principal investigator to other higher priority work assignments. The decision was made not to proceed with this study until 1985-86. ��399 NESTING PARAMETERS, NEST SITE SELECTION, AND REPRODUCTIVE PERFORMANCE OF RIO GRANDE WILD TURKEYS IN LOWLAND RIPARIAN HABITATS Richard W. Hoffman The wild turkey (Me1eagris ga11opavo) is becoming an increasingly popular game bird in Colorado. Numbers of hunters participating in spring and fall seasons have tripled in the last 10 years (Colo. Div. Wi1dl. 1982). In response to this demand, the Division of Wildlife has made the extension of turkey range in Colorado a primary management goal (Colo. Div. Wi1d1. 1983). Towards this goal, efforts in northeastern Colorado have been directed at establishing Rio Grande turkeys (~. ~. intermedia) in lowland riparian habitats along the South Platte River. Since 1980, 3 releases totaling 60 birds have been made at Hillrose (24), Tamarack Ranch (20), and Masters (16). While none of the releases was a total failure, the transplants have only had limited success. The Hillrose release appears to be the most successful as several flocks now occur between Snyder and Merino. The Tamarack Ranch population has persisted, but with virtually no increase in the population or major dispersal from the release area. Continual sightings of turkeys near Masters indicates at least some birds from that release survived and possibly reproduced. However, it is too early to evaluate the success or failure of this release. Although nests and broods have been found near all 3 release sites, production and subsequent recruitment of young into the population have been lower than expected. Other states, notably Kansas, have experienced high rates of population growth for Rio Grandes introduced into riparian habitats (Peabody 1963; Capel 1967, 1968). A shortage of undisturbed nesting cover may be a factor depressing reproductive success of Rio Grande turkeys along the South Platte River in Colorado. For the past 3 years, riparian habitats adjacent to the river have been inundated by high water during spring (May - early Ju1) making most of the riparian zone unavailable for nesting. Minimal nesting cover exists outside the riparian zone due to intensive farming. Five instances were documented in 1983 where turkeys nested outside of the riverbottom, apparently because of flooding. Turkeys attempted to nest in an alfalfa field (3 nests) or along fencerows (2 nests) • These nests were subsequently destroyed by mowing (3) or by predators (2). Flooding has also been linked to turkey population declines in Kansas (Peabody 1963). Studies are needed to examine nesting parameters, nest site selection, and nesting success of wild turkeys in riparian habitats. This information is vital to (1) understanding the relationship between nesting cover and reproductive performance, (2) guiding future land acquisitions by the CDOW, and (3) identifying habitat management practices that will enhance reproductive success of wild turkeys in riparian habitats. �400 P.N. OBJECTIVES The objectives of this initial planning study are to: 1. Examine and evaluate existing data on Rio Grande wild turkeys in lowland riparian habitats to obtain information pertinent to the development of a detailed study plan. 2. Prepare a detailed study plan to examine nesting parameters, nest site selection, and reproductive performance of Rio Grande wild turkeys in lowland riparian habitats. 3. Continue ongoing literature review. 4. Obtain funding. SEGMENT OBJECTIVES 1. Conduct ongoing review of literature on the Rio Grande wild turkey including habitat use, movements, dispersal, food habiats, reproduction, limiting and mortality factors, effects of harvest, effects of land-management practices, trapping and transplanting, radio-telemetric observation, and other research/study techniques. 2. Interview selected personnel of the Colorado Division of Wildlife concerning Rio Grande wild turkey research needs and study techniques. 3. Interview personnel in other states involved in research and management of Rio Grande wild turkeys to identify potential problems and assist in the development of a study plan. 4. Implement a report system sightings of wild turkeys. 5. Evaluate lowland riparian habitats Colorado as potential study areas. 6. Prepare a detailed study plan on the approved research topic and select a suitable study area to meet research requirements. 7. Obtain funding to support the research project. whereby cooperating landowners occupied by Rio Grande can report turkeys in RESULTS AND DISCUSSION Only segment objectives 1, 2, and 3 were accomplished in 1984-85. No field work was conducted and no effort was made to finalize the study plan due to a lack of funding and need to complete other higher priority work assignments. The decision was made not to proceed until the 1985-86 budget, which included funding for WP12, J16, was approved. Planned accomplishments for 1985-86 include preparing a study plan, selecting a graduate student, selecting a study area, and implementing data collection. �401 LITERATURE CITED Capel, S. W. 1967. Wild turkey population trends and introductions. For., Fish and Game Comm., Job Compl. Rep., P-R Proj. W-23-R-5. 1968. Wild turkey population trends and introductions. Fish and Game Comm., Job Comple. Rep., P-R Proj. W-23-R-6. Kansas Kansas For. Colorado Division of Wildlife. 1982. Colorado small game, furbearer, varmint harvest. Colorado Div. Wildl., Denver. 2l0pp. 1983. Today's strategy ••• tomorrow's wildlife. A comprehensive management plan for Colorado's Wildlife. Colorado Div. Wildl., Denver. 96pp. Peabody, W. 1963. Distribution, habitat and population trends of turkeys in Kansas. Knas. For., Fish and Game Comm., Job Compl. Rep., P-R Proj. W-23-R-1. ��Colorado Division of Wildlife Wildlife Research Report April 1986 403 JOB PROGRESS REPORT State of Colorado Project 01-00-045 (W-37-R) Work Plan Job Title: 13: Job Avian Research 8 Population Characteristics and Habitat Requirements of Columbian Sharp-tailed Grouse in Northwestern Colorado Period Covered: 1 January through 31 December 1984 Author: Kenneth M. Giesen Personnel: Mike Bauman, Clait Braun, Kristi Coughlon, Ken Giesen, Jim Haskins, Jim Hicks, Rick Hoffman, Mike Middleton, Dan Schaad, Colorado Division of Wildlife. ABSTRACT Counts of 12 active sharp-tailed grouse leks (Tympanuchus phasianellus columbianus) in Routt and Moffat counties in 1984 resulted in 61 males, 10 females, and 107 total sharptails being counted. The number of individuals counted was the lowest since 1980 because of the late spring and resultant poor access. Nineteen sharptails (9 males, 10 females) were captured and individually marked; all but 4 were adults. Sixteen radio-transmitters attached to ponchos were placed on selected sharptai1s (7 males, 9 females) and these birds were located up to 27 times during the summer and fall. Females generally dispersed farther from leks and had larger home ranges than males although there was much variation. A sample of 60 wings was received from the hunter harvest of which 31 (51.7%) were from juveniles. Total harvest was the lowest recorded since 1978. ��405 POPULATION CHARACTERISTICS ANDHABITAT REQUIREMENTS OF COLUMBIAN SHARP-TAILED GROUSE IN NORTHWESTERN COLORADO Kenneth M. Giesen The Columbian or mountain sharp-tailed grouse has declined in distribution and abundance throughout its historical range, including Colorado (Aldrich 1963, Miller and Graul 1980). Reasons for this decline are not well documented although changes in land use resulting from agriculture, energy development, and human population growth have coincided with population declines (Hart et a1. 1950, Kessler and Bosch 1982). Although the Columbian sharp-tailed grouse is a game species in Colorado, little information is available upon which to base management decisions. Baseline data on Columbian sharptai1 populations, habitats, and harvest are lacking not only in Colorado but throughout its range. Previous studies on distribution of Columbian sharptai1s in Colorado indicated that the largest populations and apparent best habitats occur in Routt and eastern Moffat counties (Rogers 1969, Giesen and Hoffman 1981) with most of the harvest occurring near California Park and Twentymi1e Park in central Routt County (Colo. Div. Wi1d1. 1984). Research on the sharptai1 resource is needed to obtain basic information on breeding densities, nesting success, production and survival of young, fall population numbers, harvest, population turnover, and recruitment to the breeding population. Information on seasonal movements and habitat use is needed to quantify food and cover requirements. Land use changes resulting from human population growth, agriculture, and energy development are expected to further reduce sharptai1 habitat in the near future while hunting and other recreational uses of sharptai1s and their habitat are expected to increase. This report covers the 4th year of the planned 5-year study. P.N. OBJECTIVES The major objectives of this study are to measure sharptai1 breeding density, production, harvest, survival and turnover rates, and to obtain qualitative and quantitative measures of Columbian sharptai1 habitat in western Colorado. Segment Objectives 1. Review pertinent literature applicable to the objectives of this study. 2a. Locate dancing grounds in March, April, and May by systematic search of suitable habitats using binoculars, spotting scopes, and a parabolic microphone listening device during morning and evening display periods. 2b. Count male and female sharptai1s on known sharptai1 leks in the 2 study areas twice weekly from mid-March through May during morning display periods. Count male and female sharptai1s on leks adjacent to study areas at bi-week1y periods during April and May. �406 2c. Identify individual male and female sharptails on leks using binoculars and spotting scopes to observe color band combinations. 3a. Locate sharptail broods by systematic search of suitable habitats and by using a tape recorded chick distress call. 3b. Ascertain brood size from counts of broods located in July and August. 4a. Trap adult sharptails using funnel traps on leks and brood areas and baited funnel traps in fall and winter, and individually mark with solar or bands. Place aluminum bands and colored plastic battery-powered radio transmitters on 2 males and on all females captured on the study leks. 4b. Capture sharptai1 chicks using walk-in and/or drive traps in areas where sharptails are known to occur. 5. Locate all radio-marked sharptai1s weekly range size and seasonal habitat use. 6. Habitat at sharptail use sites and random sites will be quantified using cover boards and point-center quarter measurements. 7. Estimate nesting success from wing molt of hens captured in summer and from wings obtained from hunter harvested birds. 8. Ascertain distribution of hunters and hunter harvest from field hunter checks, check stations, and wing barrels. 9. Obtain number and location of marked birds shot through use of field hunter checks, check stations, and voluntary mail reporting. 10. Food habits will be ascertained from crops of sharptai1s obtained from hunters and systematic collections. 11. Compile data, analyze results, and prepare progress reports. for documentation of home METHODS Field surveys were conducted on both study areas (Cedar Hill Gulch, Hayden-North) from late-March through early June using binoculars and a parabolic microphone listening device to locate leks within the study areas. Binoculars and spotting scopes were used to obtain counts of male and female sharptai1s on leks and flush counts were used to obtain complete counts of sharptai1s on leks. Walk-in drive traps were used to capture sharptai1s on leks. Intensive searches on foot and a trained pointing dog were used to locate sharptai1 broods in July and August. Sixteen s91ar- or battery-powered radio transmitters were attached to ponchos (Amstrup 1980) and placed on sharptai1s to facilitate periodic relocation. Vegetative structure and species composition were measured on leks, sharptai1 use sites, and random sites using cover board (Jones 1968) and point-centered quarter (Cottam and Curtis 1956) methods. Distribution of hunters and hunter harvest was measured using 2 check stations, field checks, and 12 wing barrels. �407 DESCRIPTION OF STUDY AREA Two areas (Cedar Hill Gulch, Hayden-North) were selected for intensive study. Cedar Hill Gulch is in Moffat County (T9N, R89W, parts of Sections 31, 32 and 33; and T8N, R89W, parts of Sections 4-10 and 15-17) and is primarily dominated by native mountain shrub communities and irrigated hay meadows. Hayden-North is in Routt County (T8N, R88W, parts of Sections 3-5; and T9N, R88W, parts of Sections 19-21 and 28-33). Hayden-North is dominated by native mountain shrub communities, hay pastures, and wheat fields. A complete vegetative description will be included in the final report. RESULTS AND DISCUSSION Lek Counts From 28 March through 12 June, 151 counts of 12 active sharp-tailed grouse leks were obtained in Routt and Moffat counties. Eleven counts were made on an additional 4 historic leks (Bear Creek, California Park Road #2, Cottonwood Creek, Salt Creek 112) on which no sharptails were seen. A minimum of 107 sharptails (8.9/active lek) was counted. The number of leks counted in 1984 decreased 50% from the 1981-83 period (Table 1). Inclement weather conditions and poor access during the breeding season accounted for most of the decrease in counting effort. Because of other research activities and poor access, little effort was directed to locating additional leks this year. Table 1. Sharp-tailed grouse counties, 1964-65 and 1977-84. Year 1964 1965 1977 1978 1979 1980 1981 1982 1983 1984 N active grounds counted 7 15 2 6 15 5 24 26 24 12 dancing ground N sharEtails counted 38 91 16 54 123 36 335 317 311 107 counts in Routt and Hoffat N sharptails/active 1ek 5.4 6.7 8.0 9.0 8.2 7.2 14.0 12.2 13.0 8.9 Because of the ongoing research project there was an uneven counting effort among leks. Five leks accounted for most of the counting effort (120 counts, 79.5%) because they coincided with trapping activities. Peak counts of males generally occurred in mid-April prior to hen attendance; peak counts of females occurred in early May. �408 Numbers of sharptai1s counted on both study leks declined from previous years (Table 2). This was particularly true for Smith's Lek on the Hayden-North study area. Snow conditions on the 1ek may have caused failure of "new" males to recruit as production in 1982 and 1983 (as measured from the fall harvest) was the highest recorded in Colorado. Alternatively, the severe winter of 1983-84 may have substantially increased overwinter mortality although I have no data to support this hypothesis. I suspect that alternative 1ek sites may have been used because snowcover on Smith's Lek remained into May. The failure to detect alternative leks may have resulted in lower counts. Table 2. High counts of sharp-tailed grouse on study leks in Routt and Moffat counties, 1981-84. Year Males 1981 1982 1983 1984 18 10 9 11 Cedar Hill Gulch Females 0 2 1 3 Totals 24 14 19 17 Males Smith's Females Totals ---Not counted in 1981 20 1 3 17 6 2 24 18 7 TraEEin~ and Banding Nineteen sharp-tailed grouse (9 males, 10 females) were captured and individually marked in 1984. All were captured using walk-in drive traps on leks. In addition 2 males originally banded in 1982 and 1 male originally banded in 1983 were recaptured. Only 4 of 22 birds trapped (18.2%) were yearlings suggesting poor recruitment to the breeding population in spite of high production in 1983. Although many yearling males may not recruit to leks until 2 years of age or older (Rippin and Boag 1974), it is surprising that only 1 of 10 hens captured was a yearling. Ovarian analysis of hunter harvested birds (unpub1. data) suggests that nearly all hens breed. Telemetry Investigations Sixteen radio transmitters were placed on sharp-tailed grouse (7 males, 9 females) in May to obtain data on nesting, movements, and habitat use. Mortality and transmitter malfunction resulted in a loss of information beyond 5 weeks for 4 birds; 8 birds had functional transmitters lasting greater than 10 weeks. Four nests were located using telemetry in 1984. Three complete clutches averaged 9.7 eggs and 1 partial clutch had 6 eggs. The average distance from the 1ek to the nest site was 0.82 km (range 0.62-1.00 km). Another hen apparently nested 1.2 km east of Smith's Lek but the clutch was depredated before it was examined. Two hens were depredated prior to incubation and radio transmitters failed on 2 other marked hens. �409 With few exceptions all radio-marked sharptai1s remained within 2.0 km of the 1ek where they were trapped until October (Table 3). Generally males had less dispersion from leks than females and there was little difference between areas. One male dispersed from the Cedar Hill Gulch Lek immediately after the breeding season (4.3 km movement). Females generally left the 1ek after mating and remained dispersed for the remainder of the year. Table 3. Average monthly dispersion grouse from 1ek of capture, 1984. Band Age 227 245 281 282 283 284 285 286 287 288 291 292 293 294 295 296 Sex 4+ 2+ 2+ 2+ M M M M M F F F F M F F M F F F 1- 2+ 2+ 2+ 2+ 1- 2+ 2+ 2+ 2+ 1- 2+ May Jun 180 550 30 700 30 480 430 720 960 160 680 920 220 1,540 1,220 580 130 Ju1 (m) of male and Month Aug Sep female Oct 280 3,000 230 790 900 520 4,300 270 610 450 200 450 200 600 1,430 460 900 1,000 200 2,200 640 650 1,440 1,900 950 2,000 1,600 2,000 3,200 2,300 40 100 420 1,080 500 1,600 1,500 400 sharp-tailed Nov Dec 4,100 Locations of radio-marked sharp-tailed grouse were plotted on USGS topographic maps (scale 1:24,000) and home ranges were calculated. There was no difference in home range size between birds from Smith's Lek (Table 4) and Cedar Hill Gulch Lek (Table 5). There was much variation in home range size among individual birds although females generally had larger home ranges than males. Table 4. Home ranges of sharp-tailed grouse trapped at Smith's Lek, 1984. N Band 227 245 286 293 294 295 Age 4+ 2+ 2+ 2+ 2+ 1- Sex H M F M F F relocations 12 6 5 23 7 17 Period 8 8 11 10 11 11 May-12 May-18 May-30 May-21 May-12 May-21 Home range (ha) Jun May May Aug Jun Aug 18.8 6.9 6.9 28.8 61.8 125.7 �410 Table 5. 1984. Band Home ranges of sharp-tailed grouse trapped at Cedar Hill Gulch Lek, Age 281 282 283 284 285 287 288 291 292 296 2+ 2+ 12+ 2+ 2+ 12+ 2+ 2+ Sex M M H F F F M F F F N relocations 25 8 17 25 13 27 18 5 11 16 Period 5 4 10 7 7 7 10 10 10 14 May-21 May-11 May-24 May-II May-23 May-II May-24 May-IO May-24 May-24 Home range (ha) Sep Ju1 Jul Oct Jun Oct Oct Jun Jun Oct 60.4 N/Aa 29.2 91.0 11.1 85.4 166.0 6.9 20.1 136.1 aThree relocations on lek, remainder of relocations up to 4,300 m from lek. Harvest The hunting season for Columbian (mountain) sharp-tailed grouse in 1984 started one-half hour before sunrise on 8 September and closed at sunset on 7 October on all lands west of the Continental Divide. This was the longest season (30 days) on sharp-tailed grouse in Colorado in modern times. It was also the first year in which all Small Game Management Units on the Western Slope had identical season lengths. Bag and possession limits were 3 and 6, respectively and separate from the bag limits of sage grouse (Centrocercus urophasianus). The sample of wings received (60) was the smallest received since 1978 and was only 27% of the sample of wings obtained in 1983. The reduced harvest may be attributed to fewer grouse hunters and/or reduced sharptail populations in the major hunting areas. Wings were received from 2 check stations (California Park = 15, Cedar Mountain = 2), 6 wing barrels (California Park Road = 1, Hayden Cog Road = 9, Hayden Gulch Road = 7, Ralph White Reservoir = 5, Steamboat Springs = 3, Twentymile Park Road = 5), and field checks of hunters (12 wings). One sharptail was found dead during the hunting season. Of this sample, 37 were from Unit 14, 2 from Unit 16, and 21 from Unit 26. Over one-third of the wings (36.7%) were collected during the initial weekend of the season. Operation of grouse check stations on opening weekend accounted for most (77.3%) of these wings. Harvest on weekdays (21 wings, 35%) was less than on weekends (39 wings, 65%). Twenty percent of the wings were received during the final week of the season indicating hunters took advantage of the more liberal season lengths. Temporal distribution of the 1984 harvest along with comparative data from 1981-83 (Table 6) indicate that more liberal seasons tend to distribute hunting pressure over longer periods. �411 Table 6. Time distribution northwestern Colorado, 1981-84. of 1981 Period First weekend First week Second weekend Second week Third weekend Third week Fourth weekend Fourth week Fifth weekend Totals grouse shar~tailed wings received, 1983 1982 % 1984 N % 104 26 7 3 2 73.2 18.3 4.9 2.1 1.4 90 22 19 18 29 SEASON SEASON SEASON SEASON 50.6 12.4 10.7 10.1 16.3 CLOSED CLOSED CLOSED CLOSED 103 29 37 5 50 46.0 12.9 16.5 2.2 22.3 22 7 10 6 2 0 1 8 4 36.7 11.7 16.7 10.0 3.3 0 1.7 13.3 6.7 142 99.9 178 100.0 224 99.9 60 100.1 N N % % N Age was ascertained from 60 wings examined and gonadal inspection of 19 sharp tails provided sex ratios in the harvest (Table 7). No yearlings were identified, probably because of small sample size and the fact that 78.6% of the adult-yearling class had molted primary 10. Sex ratios in both age classes were skewed towards females but this may be an artifact of small sample sizes. Data on age and sex composition of the hunter harvest from all years for which data are available are presented in Table 8. Table 7. Age composition Colorado, 1984. of harvested sharp-tailed grouse, Males a A~e class N Adults Yearlings Chicks 29 0 31 % 48.3 0.0 51.7 N 3 0 3 aonly those for which gonads were examined. % 27.3 0.0 37.5 N 8 0 5 northwestern Females a % 72.7 0.0 62.5 �412 Table 8. Age and sex composition northwestern Colorado, 1976-83. Year Adults a N % 1976 1977 1978 1979 1980 1981 1982 1983 1984 10 47 13 32 25 83 60 74 29 Totals 374 9-yr. avg. N % 4 25 15 47 38 59 117 150 31 28.6 34.7 53.6 59.5 60.3 41.6 66.1 67.0 51.7 14 72 28 79 63 142 177 224 60 5 5 1 3 14 9 5 3 858 40 484 43.6 sharp-tailed a,b Adults Females Males Total sample Youn~ 71.4 65.3 46.4 40.5 39.7 58.4 33.9 33.0 48.3 harvested of 4 grouse, b Youn~ Females Males 6 4 18 7 7 8 7 3 13 24 18 3 10 3 15 19 23 5 49 74 79 56.4 aIncludes yearlings. bKnown sex only. While we cannot assume a random sample was obtained population we can draw some conclusions from our data. from the sharptail 1. Production of young was only fair in 1983 (51.7% chicks) and below the 9-year average. If we assume an even sex ratio among adults, then the number of chicks per hen in 1984 (2.1) was below the' 9-year average (2.6) and approximately half of the 1982-83 values (4.0 chicks/hen average). 2. Since the percentage of juveniles in a stable population is a measure of total year-to-year turnover, we can be confident that even in years of poor production (Le., 1981, 1984) more chicks are produced than can be recruited into the following years' breeding population. Our long-term harvest data suggests that annual population turnover is 55-60%. LITERATURE CITED Aldrich, J. W. 1963. Geographic orientation of American Tetraonidae. Wildl. Manage. 24:529-545. Amstrup, S. C. 214-217. 1980. A radio-collar for game birds. J. J. Wildl. Manage. 44: Colorado Division of Wildlife. 1984. Colorado small game, furbearer and varmint harvest, 1983. Colorado Div. Wildl., Denver. l86pp. �413 Cottam, G., and J. T. Curtis. 1956. The use of distance measures in phytosociological sampling. Ecology 37:451-460. Giesen, K. M., and D. M. Hoffman. 1981. Distribution and status of mountain sharp-tailed grouse. Colorado Div. Wildl., Final Rep., Fed. Aid Proj. W-37-R-34. Apr. 1981. Pp. 183-189. Hart, C. M., O. S. Lee, and J. B. Low. 1950. The sharp-tailed grouse in Utah. Its life history, status, and management. Utah Dep. Fish and Game, Fed. Aid Div. Pub1. 3. 79pp. Jones, R. E. 1968. A board to measure cover used by prairie grouse. Wi1d1. Manage. 32:28-31. J. Kessler, W. B., and R. P. Bosch. 1982. Sharp-tailed grouse and range management practices. Pages 133-146 in Peek, J. M. and P. D. Dalke, editors. Proc. Wi1d1./Livestock Symp.,lJniv. Idaho, Moscow. Miller, G. C., and W. D. Graul. 1980. Status of sharp-tailed grouse in North America. Pages 18-28 in P. A. Vohs, Jr., and F. L. Knopf, eds. Proc. Prairie Grouse Symp., Oklahoma State Univ., Stillwater. Rippin, A. B., and D. A. Boag. 1974. Recruitment to populations of male sharp-tailed grouse. J. Wild1. Manage. 38:616-621. Rogers, G. E. 1969. The sharp-tailed grouse in Colorado. Fish and Parks Tech. Pub1. 23. 94pp. Prepared by _+~~...:.;....:.=-_;....;_~~~;;:og=-,.~ KeiltlethM.Giesen Wildlife Researcher _ Colorado Div. Game, ��Colorado Division of Wildlife Wildlife Research Report April 1986 415 JOB PROGRESS REPORT Colorado State of Project 01-03-045 (W-37-R) Work Plan 13 Job Title: 8 -~- Population Characteristics and Habitat Requirements of Columbian Sharp-tailed Grouse in Northwestern Colorado Period Covered: Author: : Job Avian Research 01 January through 31 December 1985 Kenneth M. Giesen Personnel: Mike Bauman, C1ait Braun, Kevin Brennan, Ken Giesen, Jim Haskins, Jim Hicks, Rick Hoffman, Mike Middleton, Colorado Division of Wildlife. ABSTRACT Counts of 7 active sharp-tailed grouse (Tympanuchus phasiane11us) leks in Routt and Moffat counties resulted in 73 sharptails being counted (ave. 10.4). Five additional leks had only sporatic attendance by 1 or 2 males and an additional 8 historic leks were not active in 1985. One male and 2 female sharp-tailed grouse were captured on leks and both females were radiomarked. Two of 3 sharp-tail grouse nests located were successful. Harvest data indicated that production of young was excellent (72.5% chicks) and most harvest occurred in the first week of the hunting season. ��417 POPULATION CHARACTERISTICS AND HABITAT REQUIREMENTS OF COLUMBIAN SHARP-TAILED GROUSE IN NORTHWESTERN COLORADO Kenneth M. Giesen The Columbian or mountain sharp-tailed grouse has declined in distribution and abundance throughout its historical range, including Colorado (Aldrich 1963, Miller and Graul 1980). Reasons for this decline are not well documented although changes in land use resulting from agriculture, energy development, and human population growth have coincided with population declines (Hart et al. 1950, Kessler and Bosch 1982). Although the Columbian sharp-tailed grouse is a game species in Colorado, little information has been available upon which to base management decisions. Baseline data on Columbian sharptail populations, habitats, and harvest are lacking not only in Colorado but throughout its range. Previous studies on distribution of Columbian sharptails in Colorado indicated the largest populations and apparent best habitats occur in Routt and eastern Moffat counties (Rogers 1969, Giesen and Hoffman 1981) with most of the harvest occurring near California Park and Twentymile Park in central Routt County (Colo. Div. Wildl. 1984). Research on the sharptail resource is needed to obtain basic information on breeding densities, nesting success, production and survival of young, fall population numbers, harvest, population turnover, and recruitment to the breeding population. Information on seasonal movements and habitat use is needed to quantify food and cover requirements. Land use changes resulting from human population growth, agriculture, and energy development are expected to further reduce sharptail habitat in the near future while hunting and other recreational uses of sharptails and their habitat are expected to increase. This report covers the 4th year of the planned 5-year study. P.N. OBJECTIVES The major objectives of this study are to measure sharptail breeding density, production, harvest, survival and turnover rates, and to obtain qualitative and quantitative measures of Columbian sharptail habitat in western Colorado. Segment Objectives 1. Review pertinent literature applicable to the objectives of this study. 2a. Locate dancing grounds in March, April, and May by systematic search of suitable habitats using binoculars, spotting scopes, and a parabolic microphone listening device during morning and evening display periods. 2b. Count male and female sharptails on known sharptail leks in the 2 study areas twice weekly from mid-March through May during morning display periods. Count male and female sharptails on leks adjacent to study areas at bi-weekly periods during April and May. �418 2c. Identify individual male and female sharptails on leks using binoculars and spotting scopes to observe color band combinations. 3a. Locate sharptail broods by systematic search of suitable habitats and by using a tape recorded chick distress call. 3b. Ascertain brood size from counts of broods located in July and August. 4a. Trap adult sharptails using funnel traps on leks and brood areas and baited funnel traps in fall and winter, and individually mark with aluminum bands and colored plastic bands. Place solar or battery-powered radio transmitters on 2 males and on all females captured on the study leks. 4b. Capture sharptail chicks using walk-in and/or drive traps in areas where sharptails are known to occur. 5. Locate all radio-marked sharptails weekly range size and seasonal habitat use. 6. Habitat at sharptail use sites and random sites will be quantified using cover boards and point-center quarter measurements. 7. Estimate nesting success from wing molt of hens captured in summer and from wings obtained from hunter harvested birds. 8. Ascertain distribution of hunters and hunter harvest from field hunter checks, check stations, and wing barrels. 9. Obtain number and location of marked birds shot through use of field hunter checks, check stations, and voluntary mail reporting. 10. Food habits will be ascertained from crops of sharptails obtained from hunters and systematic collections. 11. Compile data, analyze results, and prepare progress reports. for documentation of home METHODS Field surveys were conducted on both study areas (Cedar Hill Gulch, Hayden-North) from April through June using binoculars and a parabolic microphone listening device to locate leks within the study areas. Binoculars and spotting scopes were used to obtain counts of male and female sharptails on leks and flush counts were used to obtain complete counts of sharptails on leks. Walk-in drive traps were used to capture sharptails on leks. Intensive searches on foot and a trained pointing dog were used to locate sharptail broods in July and August. Sixteen solar- or battery-powered radio transmitters were attached to ponchos (Amstrup 1980) and placed on sharptails to facilitate periodic relocation. Vegetative structure and species composition were measured on leks, sharptail use sites, and random sites using cover board (Jones 1968) and point-centered quarter (Cottam and Curtis 1956) methods. Distribution of hunters and hunter harvest was measured using 2 check stations, field checks, and 18 wing barrels. �419 DESCRIPTION OF STUDY AREA Two areas (Cedar Hill Gulch, Hayden-North) were selected for intensive study. Cedar Hill Gulch is in Moffat County (T9N, R89W, parts of Sections 31, 32 and 33; and T8N, R89W, parts of Sections 4-10 and 15-17) and is primarily dominated by native mountain shrub communities and irrigated hay meadows. Hayden-North is in Routt County (T8N, R88W, parts of Sections 3-5; and T9N, R88W, parts of Sections 19-21 and 28-33). Hayden-North is dominated by native mountain shrub communities, hay pastures, and wheat fields. A complete vegetative description will be included in the final report. RESULTS AND DISCUSSION Lek Counts From 30 March through 24 June, 64 counts of 7 active sharp-tailed grouse leks were obtained in Routt and eastern Moffat counties. A total of 48 additional counts was made on 5 leks which had only sporadic attendance by 1 or 2 males (Bear Creek, California Park Road #1, California Park Road #2, Smiths, Villards). Another 29 counts were obtained on 8 historic leks which were not active in 1985 (Annans 20-mile, Cottonwood Creek, Elk River Cemetary, Fly Gulch, George Gulch, Maneotis, Morapos, Pelly's). The number of active leks and number of sharp-tailed grouse counted have declined since 1981-83 (Table 1). Research personnel accounted for 135 of the 141 counts (96%) of active and historic leks. Only 1 DWM recorded lek counts of sharp-tailed grouse in 1985 (6 counts of 4 leks) (Table 2). Because of research activities and other commitments little effort was made to inventory all known sharptail leks. Sharp-tailed grouse dancing ground Table l. counties, 1964-65 and 1977-85. counts in Routt and Moffat Year N active srounds counted Total birds counted Average lek size 1964 1965 1977 1978 1979 1980 1981 1982 1983 1984 1985 7 15 2 6 15 5 24 26 24 12 7 38 91 16 54 123 36 335 317 311 107 73 5.4 6.1 8.0 9.0 8.2 7.2 14.0 12.2 13.0 8.9 10.4 �420 Table 2. Sharp-tailed grouse dancing ground summary, 1985. Lek N Counts N total birds 23 9 2 12 10 5 9 21 15 Cedar Hill Gulch Elkhead 113 Masiarelli McKinney Ranch Morgan Creek Noland Ranch Wisemans 6 2 10 Date of high count 03 May 18, 22 Apr 17 Apr 18 Apr 20 May 03 May 22 May 6 7 After several years of counting sharp-tailed grouse on dancing grounds in northwestern Colorado it is readily apparent that we know little of their lek dynamics. In some areas where sharptail populations are high (20-mile park) we have not been able to consistently locate birds in spring. Some populations have experienced little or no hunting pressure (many on private property) yet Lek counts have indicated population declines. Some lek sites have remained in the same location for more than 20 years. Generally, counts of sharptails on leks show little daily variation although peak counts usually occur early in the season. Activity of sharptails on leks shows high daily and seasonal variation. Without a better understanding of lek dynamics (percent of males attending, attendance pattern of females, lek site fidelity), it is difficult to recommend lek counts as a management tool for monitoring populations. Numbers of sharptails counted on both study leks declined from previous years (Table 3). This was particularly true for Smith's Lek on the Hayden-North study area. Snow conditions on the lek may have caused failure of "new" males to recruit as production in 1982 and 1983 (as measured from the fall harvest) was the highest recorded in Colorado. Alternatively, the severe winter of 1983-84 may have substantially increased over winter mortality although I have no data to support this hypothesis. I suspect that alternative lek sites may have been used because snow cover on Smith's Lek remained into May. The failure to detect alternative leks may have resulted in lower counts. High counts of sharp-tailed grouse on study leks in Routt and Table 3. Moffat counties, 1981-85. Year Males 1981 1982 1983 1984 1985 18 10 9 11 10 Cedar Hill Gulch Females Totals 0 2 1 3 0 24 14 19 17 10 Males 20 17 6 2 Smith's Females Totals Not counted in 1981 1 24 3 18 2 7 0 2 �421 Trapping and Banding Three sharp-tailed grouse (1 male, 2 females) were captured and individually marked in 1985. All were captured using walk-in funnel traps at Cedar Hill Gulch Lek. The low trapping success may have been the result of trapper inexperience (the primary investigator was assigned elsewhere during the trapping season) and sharptail behavior at the lek. Because the number of males attending the 1ek was low, there appeared to be less interaction among males resulting in fewer chases into the wire leads going into the traps. Because of only sporatic lek attendance at Smith's Lek north of Hayden, trapping efforts there were terminated after 2 weeks. Telemetry Investigations Radio transmitters were placed on both hens captured at Cedar Hill Gulch Lek. Another hen, originally captured in 1984, had a functioning transmitter and these 3 hens provided all telemetry data for 1985. These hens nested 700, 1,000, and 1,150 m respectively, from Cedar Hill Gulch Lek. All nests were in the native mountain shrub community of serviceberry (Amelanchier sp ,) and snowberry (Symphoricarpus sp.). Two of 3 hens were successful in nesting although brood sizes 3 weeks post-hatch were only 2 and 7 chicks. One successful hen was found depredated on 14 August and neither of the remaining radio-marked hens was located after 18 July. All telemetry locations of the hens were in serviceberry-snowberry-sagebrush habitats within 1,600 m of Cedar Hill Gulch Lek. Harvest The hunting season for Columbian (mountain) sharp-tailed grouse in 1985 started one-half hour before sunrise on 14 September and closed at sunset on 6 October in Small Game Management Units 14, 16, 24, 26, 58, 62, and 64. The season length (23 days) was shorter than in 1984 (30 days) but longer than all previous seasons with the exception of a 25 day season in Units 14, 16, 18, and 20 in 1980. The bag and possession limits were 3 and 6, respectively, and separate from sage grouse. The sample of wings received in 1985 (69) was higher than in 1984 but only 40% of the sample received in 1981-83. The low harvest can be attributed to fewer hunters than in previous years and possibly lower sharptai1 populations in California Park, one of the traditionally high harvest areas. Distribution of the Harvest Wings were received from 2 check stations (California Park = 10, Cedar Mountain = 6), 7 wing barrels (Twentymile Park = 22, California Park = 10, Oak Creek = 7, Hahn's Peak = 2, Hayden Gulch = 2, Elk Mountain = 1, 5-Pines South = 1), and field checks of hunters (8 wings). Of this sample, 37 were from Unit 26, 26 from Unit 14, and 6 from Unit 16. Comparative data from 1981-84 are presented in Table 4. Over half the wings (62.3%) were collected during the initial weekend season. Operation of grouse check stations accounted for 37.2% of Harvest on weekdays (13 wings, 18.8%) was less than on weekends (56 81.2%) • Over 10% of the wings were received from the last weekend of the these. wings, of the �422 Table 4. Origin of sharp-tailed grouse wings, northwestern 1981 Location Unit 14 Calif. Park Check Station Gould Check Station Black Mountain Wing Barrel Ralph White Res. Wing Barrel Moffat Co. Rd. 29 Wing Barrel Hayden Cog Rd. Wing Barrel Calif. Park Wing Barrel Elk Mtn. Wing Barrel Hahn's Peak Rd. Wing Barrel Field Checks 1982 % N Colorado, 1981-85. 1983 % !1 N 1984 % N 1985 % N % 35 0 0 5 N/A N/A 2 N/A 0 24 24.6 0.0 0.0 3.5 N/A N/A 1.4 N/A 0.0 16.9 34 0 0 0 N/A 9 3 N/A 1 17 19.1 0.0 0.0 0.0 N/A 5.1 1.7 N/A 0.6 9.6 45 2 1 0 11 3 16 16 17 10 20.1 0.9 0.4 0.0 4.9 1.3 7.1 7.1 7.6 4.5 15 0 0 5 0 9 1 0 0 7 25.0 0.0 0.0 8.3 0.0 15.0 1.7 0.0 0.0 11.7 10 0 N/A 0 0 0 10 1 2 3 14.5 0.0 N/A 0.0 0.0 0.0 14.5 1.4 2.9 4.3 66 46.5 64 36.0 121 54.0 37 61.7 26 37.7 1 0 2 0.7 0.0 1.4 0 0 0 0.0 0.0 0.0 4 5 0 1.8 2.2 0.0 2 0 0 3.3 0.0 0.0 6 0 0 8.7 0.0 0.0 3 2.1 0 0.0 9 4.0 2 3.3 6 8.7 38 6 5 0 9 12 3 0 N/A 0 0 26.8 4.2 3.5 0.0 6.3 8.5 2.1 0.0 N/A 0.0 0.0 24 7 36 2 28 13 0 0 N/A 0 3 13.5 3.9 20.0 1.1 15.7 7.3 0.0 0.0 N/A 0.0 1.7 3 15 23 3 24 9 3 3 N/A 0 11 1.3 6.7 10.3 1.3 10.7 4.0 1.3 1.3 N/A 0.0 4.9 N/A 5 7 N/A 3 0 0 0 0 0 6 N/A 8.3 11.7 N/A 5.0 0.0 0.0 0.0 0.0 0.0 10.0 N/A 22 2 N/A 0 7 0 0 0 1 5 N/A 31.9 2.9 N/A 0.0 10.1 0.0 0.0 0.0 1.4 7.2 73 51.4 113 63.5 94 42.0 21 35.0 37 53.6 0 0.0 1 0.6 0 0.0 0 0.0 0 0.0 Subtotal 0 0.0 1 0.6 0 0.0 0 0.0 0 0.0 Total 142 100.0 178 100.1 224 100.0 60 100.0 69 100.0 Subtotal Unit 16 Cedar Mtn. Check Station Cedar Mtn. Wing Barrel Moffat Co. Rd. 3 Wing Barrel Subtotal Unit 26 Twentymile Park Check Station Twentymile Park Wing Barrel Hayden Gulch Wing Barrel Milner Wing Barrel Steamboat Springs Wing Barrel Oak Creek Wing Barrel Wolcott Wing Barrel Rock Creek Wing Barrel Sage Creek Wing Barrel Five Pines South Field Checks Subtotal Unit 54 Muddy Creek Wing Barrel �423 season indicating hunters took advantage of longer season lengths. Temporal distribution of the 1985 harvest is given in Table 5 along with comparative data for 1981-84. Age was ascertained from 69 wings examined and gonadal inspection of 14 sharptails provided sex ratios of the harvest (Table 6). Only 1 yearling was identified, probably because of small sample size and the fact that 65.0% of the adult-yearling class had molted primary X. Sex ratio was skewed towards females in the juvenile age class but this may be an artifact of small sample sizes. Data on age and sex composition of the hunter harvest from all years for which data are available are presented in Table 7. While we cannot assume a random sample was obtained population we can draw some conclusions from our data. from the sharptail 1. Production of young was excellent in 1985 (72.5% chicks), the highest recorded since we began collecting harvest data in 1976. This level of production is nearly 15% above the 10-year average. If we assume an even sex ratio among adults, then the number of chicks per hen in 1985 was 5.3, the highest ever recorded. 2. Since the percentage of juveniles in a stable population is a measure of total year-to-year turnover we can estimate an annual turnover of 55-60%. This suggests that we cannot stockpile sharptails and that an acceptable harvest level may be 25-30% of the fall population. 3. Most hunting pressure and harvest occurs during the initial weekend of the season. Long seasons have a minor impact on total harvest but provide additional recreational resource. A minimum season length of 30 days will provide adequate recreational opportunity without impacting subsequent breeding populations. �424 Table 5. Time distribution of sharp-tailed 1981 grouse wings received, northwestern 1982 1983 1984 Period N % !i. % N. % First weekend First week Second weekend Second week Third weekend Third week Fourth weekend Fourth week Fifth weekend 104 26 7 3 2 N/A N/A N/A N/A 73.2 18.3 4.9 2.1 1.4 N/A N/A N/A N/A 90 22 19 18 29 N/A N/A N/A N/A 50.6 12.4 10.7 10.1 16.3 N/A N/A N/A N/A 103 29 37 5 50 N/A N/A N/A N/A 46.0 12.9 16.5 2.2 22.3 N/A N/A N/A N/A 22 7 10 6 2 0 1 4 4 Totals 142 99.9 178 100.0 224 99.9 60 Table 6. Age Colorado, 1985. composition Colorado, 1981-85. of harvested sharp-tailed grouse, N Adults Yearlings Chicks 18 1 50 % 26.1 1.4 72.5 li % 36.7 11. 7 16.7 10.0 3.3 0.0 1.7 13.3 6.7 43 11 5 2 1 0 7 N/A N/A 62.3 15.9 7.2 2.9 1.4 0.0 10.1 N/A N/A 100.1 69 99.8 northwestern Females Males Age class 1985 % Ii H. % N. 2 0 3 40.0 0.0. 60.0 2 0 7 % 22.2 0.0 77.8 aOnly those for which gonads were examined. Table 7. Age and sex composition Adults a Year N 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 10 47 13 32 25 83 60 74 29 19 Totals 10-year average of harvested Youn~ % 71.4 65.3 46.4 40.5 39.7 58.5 33.9 33.0 48.3 27.5 393 N 4 25 15 47 38 59 117 150 31 50 % 28.6 34.7 53.6 59.5 60.3 41.5 66.1 67.0 51.7 72.5 534 42.4 alnc1udes yearlings. bKnown sex only. 57.6 sharp-tailed Total saml!le grouse, northwestern Males Adu1tsa,b Females 14 72 28 79 63 142 177 224 60 69 5 5 1 3 14 9 5 3 2 927 42 4 Colorado. Males Younsb Females 6 4 18 7 7 8 2 7 3 13 24 18 3 3 10 3 15 19 23 5 7 51 77 86 �425 LITERATURE CITED Aldrich, J. W. 1963. Geographic orientation of American Tetraonidae. Wi1d1. Manage. 24:529-545. Amstrup, S. C. 214-217. 1980. A radio-collar for game birds. J. J. Wi1d1. Manage. 44: Colorado Division of Wildlife. 1984. Colorado small game, furbearer and varmint harvest, 1983. Colorado Div. Wi1d1., Denver. 186 pp. Cottam, G., and J. T. Curtis. 1956. The use of distance measures in phytosocio1ogica1 sampling. Ecology 37:451-460. Giesen, K. M., and D. M. Hoffman. 1981. Distribution and status of mountain sharp-tailed grouse. Colorado Div • Wi1dl., Final Rep., Fed. Aid Pro j . W-37-R-34. Apr. 1981. Pp. 183-189. Hart, C. M., O. S. Lee, and J. B. Low. 1950. The sharp-tailed grouse in Utah. Its life history, status, and management. Utah Dep , Fish and Game, Fed. Aid Div. Pub1. 3. 79 pp. Jones, R. E. 1968. A board to measure Wi1d1. Manage. 32:28-31. cover used by prairie grouse. Kessler, W. B., and R. P. Bosch. 1982. Sharp-tailed grouse and range management practices. Pages 133-146 in J. M. Peek and P. D. editors. Proc. Wi1d1./Livestock Symp., Univ. Idaho, Moscow. J. Dalke, Miller, G. C., and W. D. Graul. 1980. Status of sharp-tailed grouse in North America. Pages 18-28 in P. A. Vohs, Jr., and F. L. Knopf, eds. Proc. Prairie Grouse Symp., Oklahoma State Univ., Stillwater. Rogers, G. E. 1969. The sharp-tailed grouse in Colorado. Fish and Parks Tech. Pub1. 23. 94 pp. Prepared by ==-..:..~;:::;...~~'---=-:-....;.'-'~~..=.'=.:::-Kenneth M. Giesen Wildlife Researcher _ Colorado Div. Game, ��427 Colorado Division of Wildlife Wildlife Research Report April 1986 JOB PROGRESS REPORT State of Project Work Plan Job Title: Colorado 01-03-045 (W-37-R-38) 17 7 Population Dynamics of White-tailed Ptarmigan Period Covered: Personnel: : Job Avian Research 01 January 1984 through 31 December 1985 C1ait E. Braun and Kenneth M. Giesen, Colorado Division of Wildlife ABSTRACT Long-term studies of populations of white-tailed ptarmigan (Lagopus 1eucurus) were continued at hunted (Mt. Evans) and unhunted (Rocky Mountain National Park) areas in Colorado through 1985. Densities of breeding ptarmigan were stable during this period at Mt. Evans, although increasing from 1982-83 levels, but continued to decrease at Rocky Mountain National Park. Densities at this site were the lowest documented in the 1966-85 interval, the result of decreased adult survival and juvenile recruitment. Nesting success was variable at the 2 sites, being good in 1984 and less than average in 1985 at Mt. Evans, and average to less than average at Rocky Mountain National Park. Harvest was minimal at Mt. Evans in 1984 (6.4% of the fall population was harvested) but increased in 1985 (11.6% harvested). ��429 POPULATION DYNAMICS OF WHITE-TAILED PTARMIGAN Clait E. Braun and Kenneth M. Giesen Long-term studies of trends in population size and investigation of reasons for fluctuations in size of tetraonid populations are lacking. Studies on the population dynamics of unhunted and hunted populations of white-tailed ptarmigan were initiated in Colorado in 1966 and have continued essentially uninterrupted at 2 sites. Studies of the unhunted population (Rocky Mountain National Park) have identified possible short-term cycles of 7-8 years with an amplitude of 25-30% between high and low breeding densities. Conversely, studies of the manipulated population (hunted) at Mt. Evans through 1980 have not indicated any cyclic pattern and it would appear that controlled hunting may mask any long-term trend that may occur. This study is designed to examine the question whether white-tailed ptarmigan are truly cyclic and whether hunting affects the apparent oscillations. P. N. OBJECTIVES The goals of this investigation are to be able to predict the length and amplitude of cycles in white-tailed ptarmigan in Colorado, to examine the impact of hunting on cycles, and to clarify underlying causes of the apparent cycles. SEGMENT OBJECTIVES 1. Conduct breeding (May-Jun) and brood (Aug-Sep) censuses of white-tailed ptarmigan using tape-recorded calls of males (breeding) and chicks (broods). 2. Censuses will be conducted on previously established, defined study areas at Mt. Evans (hunted) and at Rocky Mountain National Park (unhunted). 3. Capture (noose poles) and band (aluminum and plastic color-coded bands) all unmarked white-tailed ptarmigan encountered on study areas at Mt. Evans and at Rocky Mountain National Park. 4. Individually identify all ptarmigan observed on study areas at Mt. Evans and Rocky Mountain National Park through use of binoculars. 5. Make hunting season and bag limit recommendations for Mt. Evans and collect hunting data through use of volunteer wing barrels and hunter field checks. 6. Compile data, analyze results, and prepare progress reports. STUDY AREA AND METHODS Areas investigated were Mt. Goliath-Mt. Evans in Clear Creek County and at Tombstone Ridge-Sundance Mountain to Fall River Pass in Rocky Mountain �430 National Park in Larimer County. The physiography, geology, location, and vegetation of these study areas has been previously described (Braun 1969, 1971; Braun and Rogers 1971; Giesen 1977). Ptarmigan were located through use of tape-recorded calls (Braun et al. 1973), captured through use of telescoping noose poles (Zwickel and Bendell 1967) as described by Braun and Rogers (1971), and classified to age and sex and banded following Braun and Rogers (1971). Age of chicks was estimated following Giesen and Braun (1979). Numbered plastic bandettes were not used as in earlier years (Braun and Rogers 1971) as a color-code system using up to 4 different colored plastic bandettes was instituted in 1977-78. A check station was operated on the Mt. Evans highway during the opening weekend of the ptarmigan season in that area. A volunteer wing collection station was available to hunters in the area when the check station was not in operation until the season closed. RESULTS AND DISCUSSION Breeding Densities Mt. Evans Timing of breeding events in the Mt. Evans area was earlier in 1984 than in 1982 or 1983. During the late May-early June interval, 15 pairs and 2 single males were identified. The breeding density increased over levels documented in 1982 and 1983 and was the fourth highest on record (higher only in 1979-81) (Table 1). Of interest was the finding that 6 of 15 paired territorial males were yearlings. Yearling females totaled 8 of 15 hens located in pairs. In 1985, timing of breeding events in the Mt. Evans area was again early, about 3-5 days earlier than in 1984. During the breeding period, 14 pairs and 4 single males were located on the study area and the breeding density (8.0 birds/km2) remained the same as in 1984 (Table 1). Only 2 of 14 paired territorial males were yearlings in 1985 while only 3 of 14 hens located in pairs were yearlings. Rocky Mountain National Park Surveys of ptarmigan present on the RMNP study units during May and June 1984 and 1985 revealed 13 pairs and 6 single males (5.8 birds/km2) and 14 pairs and 5 single males (6.0 birds/km2), respectively (Table 1). These densities were the lowest recorded since initiation of the long-term study in 1966 and continued the decline first documented in 1977. The population decline has been manifested by below average adult survival (males, <50-55%; females, <25-44%) and poor recruitment of juveniles (0-5% of banded chicks recruited). �431 Table 1. 1966-85. White-tailed ptarmigan breeding densities (birds/km2), Colorado. Studl area Year Rocky Mountain National Park (5.5 km2) Mt. Evans (4.0 km2) 1966 1967 1968 1969 1970 1971 1972 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 11.3 9.8 11.5 12.0 9.6 9.1 8.7 7.8 8.0 11.1 13.5 12.9 10.7 8.7 8.4 8.2 7.8 6.7 5.8 6.0 3.0 2.7 2.7 2.2 2.0 4.2 7.5 6.2 6.2 6.2 6.7 >6.0 7.5 10.3 9.5 9.0 6.5 6.5 8.0 8.0 Nesting Success and Brood Size Mt. Evans Seventeen hens were located during mid July-early September 1984 on or immediately adjacent to the study area. Ten hens (58.8%) were with broods while 7 were apparently unsuccessful nesters (without chicks). Average brood size to 1 September was 3.4 chicks/successful female. Data were available for 31 chicks captured and banded or collected (1), and taken during the hunting season (1). Twenty-five chicks (80.6%) hatched between 3 and 15 July, 2 hatched on 22 July, while 4 hatched on 29-31 July. This latter group probably were progeny of a renest. Nesting success and production decreased in 1985 as only 7 of 17 (41.1%) hens observed from July to early September were with broods. Average brood size to 1 September was 3.5 chicks/successful female. Data were available for 25 chicks captured and banded or taken during the hunting season. Of this total, 21 (84%) were progeny of eggs hatching between 30 June and 11 July. Four were probably the result of renests as they hatched between 15 and 19 July. �432 Rocky Mountain National Park Three successful hens (3/5, 60%) were observed between late July and early September 1984 with average brood size of only 2.2 chicks (range 1-5). The peak of hatch occurred during the second week of July. Production improved in 1985 but only 5 of 9 hens (55.6%) identified during the brood season were with chicks. Average brood size was 4.6 (range 1-7) indicating good chick survival. The peak of hatch occurred during the first week of July (median hatch date = 4 Jul, range 30 Jun-14 Jul). Harvest Mt. Evans The hunting season at Mt. Evans in 1984 opened on 22 September and closed on 7 October (16 days) with a bag and possession limit of 3 and 6. Thus, the season was delayed 2 weeks from the statewide opening as it has been each year since 1978 (except 1981 which was delayed 1 week). A check station was operated from dawn to dusk on the opening weekend (22-23 Sep) similar to operations in previous years. In addition, a volunteer wing collection station was available through 7 October. During the 2-day check, 26 hunters with 9 ptarmigan were checked. Five of the 9 ptarmigan were banded. No other wings or bands were received during or after the 1984 hunting season. Thus, a minimum of 6.4% of the resident fall population of ptarmigan was harvested. This was the lowest rate of harvest since 1977. Harvest rates have decreased each year since 1981 after having increased from 1977 through 1981. In 1985, the hunting season at Mr. Evans opened on 21 September and closed on 6 October (16 days) with a bag and possession limit of 3 and 6. Thus, the season was delayed 2 weeks from the statewide opening as it has been each year since 1978 (except 1981 which was delayed 1 week). A check station was operated from dawn to dusk on the opening weekend (21-22 Sep) similar to operations in previous years. In addition, a volunteer wing collection station was available through 6 October. During the 2-day check, 33 hunters with 14 ptarmigan were checked. Ten of the 14 ptarmigan were banded. No other wings or bands were received during or after the 1985 hunting season. Thus, a minimum of 11.6% of the resident fall population of ptarmigan was harvested. This was almost twice the harvest rate experienced in 1984 and was similar to the harvest rate (12.8%) documented in 1983. LITERATURE CITED Braun, C. E. 1969. Popula~ion dynamics, habitat, and movements of whitetailed ptarmigan in Colorado. Ph.D. Thesis, Colorado State Univ., Fort Collins. 189 pp. 1971. Habitat requirements of Colorado white-tailed ptarmigan. Proc. West. Assoc. State Game and Fish Comm. 51:284-292. _________ , and G. E. Rogers. 1971. The white-tailed ptarmigan in Colorado. Colorado Div. Game, Fish and Parks Tech. Publ. 27. 80 pp. �433 _________ , R. K. Schmidt, Jr., and G. E. Rogers. 1973. Census of Colorado white-tailed ptarmigan with tape recorded calls. J. Wildl. Manage. 37:90-93. Giesen, K. M. 1977. Mortality and dispersal of juvenile white-tailed ptarmigan. M.S. Thesis, Colorado State Univ., Fort Collins. 55 pp. __________ , and C. E. Braun. 1979. A technique for age determination juvenile white-tailed ptarmigan. J. Wildl. Manage. 43:508-511. Zwickel, F. C., and J. F. Bendell. J. Wildl. Manage. 31:202-204. 1967. Prepared by: Clait E. Braun Wildlife Research Leader Kenneth M. Giesen Wildlife Researcher B of A snare for capturing blue grouse. ��435 Colorado Division of Wildlife Wildlife Research Report April 1986 JOB PROGRESS REPORT Colorado State of 01-00-045 (W-37-R) (N-4-R) Project Work Plan Job Title: 21 Job: Avian Research 2 Dynamics of Cottonwood Regeneration· Period Covered: 1 January through 31 December 1984 Author: Warren D. Snyder Personnel: C. E. Braun, D. C. Bowden, S. F. Steinert, W. D. Snyder; Colorado Division of Wildlife. ABSTRACT Inventory of the lower Arkansas River and a segment of the South Republican River was completed under contract with the Colorado State Forest Service. Partial analysis of the data revealed that cottonwood (Populus spp.) stands along the Arkansas River were more severely impacted by lack of natural regeneration (p < 0.05) than cottonwoods along any of the other major rivers inventoried. Loss over an approximate 30-year interval was about 1% per year. Most losses were in the older age classes and within the higher density classifications. Shrubs, primarily tamarisk (Tamarix spp.), were the most abundant cover type present and increased by about 21% during the inventory interval. Cropland, developed land, and standing water also increased. River and unvegetated (sandbars - mudflats), combined to represent river channel, decreased (p < 0.05) during the interval with a much greater loss (P < 0.05) occurring below John Martin Reservoir than upstream. The relative short (15 years) interval for the 4.8 km (3 miles) of the South Republican River inventoried below Bonny Reservoir hampered efforts to assess long-term changes. Total hectares of cottonwoods increased during the interval but there were no young cottonwoods present during the latter (1975) photo period. River channel decreased by about one-half during the interval. Statistical treatment was applied to cover types within the South Platte, Colorado, and Rio Grande rivers and sampled strata were also compared. Intensive transects revealed approximately 98% mortality of cottonwood seedlings germinating after high water in 1983, to fall 1984 along the lower South Platte River. Seedling establishment in 1984 was much reduced and that occurring was in sites of reduced opportunity for survival. Twenty-four random transects were established in fall 1984 to monitor attrition of 1983 and 1984 cottonwood seedlings. Thirty random extensive transects were established to monitor seedling occurrence within the South Platte riverbottom. These latter transects revealed that cottonwood seedlings were present within only 0.6% of 1,667 m2 sampling frames. Over-summer survival was rated fair to good for 4 willow (Salix spp,) species, fair for plains cottonwood (Populus sargentii), and none for other species used in late-winter 1984 preliminary stem cutting propagation trials. Site selection and monitoring of groundwater level changes were made in preparation for proposed 1985 stem cutting trials on several sites in Colorado. ��437 DYNAMICS OF COTTONWOOD REGENERATION Warren D. Snyder This study expands and continues efforts to assess the status and trends of cottonwoods and other vegetation along major drainages in Colorado. Original studies titled "Lowland Riparian Cottonwood Community Studies" were initiated in 1982 under nongame research project N-4-R. Major work completed was by contract with the Colorado State Forest Service wherein a partial inventory of vegetation (primarily cottonwoods) was completed along the South Platte, Colorado, Rio Grande, and Arkansas rivers. Preliminary findings of these inventories were summarized (Snyder 1984); however, data analysis and summation are continuing and some information is presented here. In addition, this study expands efforts to document natural regeneration of cottonwoods along the South Platte River and to test methods for establishing woody cover within streamside riparian zones where natural revegetation cannot be expected to occur. P. N. OBJECTIVES Quantify changes in stand density and changes in area of riparian cover types over a recent time span approximating 30 years within the South Platte, Arkansas, and South Republican (below Bonny Reservoir) rivers in eastern Colorado, the Rio Grande River in the San Luis Valley, and the lower Colorado River in western Colorado. Analyses and evaluations will be based on aerial interpretation by the Colorado State Forest Service. Document conditions conducive to natural regeneration and survival of plains cottonwoods and willows, including site characteristics and frequency of occurrence, in streamside riparian habitats of the lower South Platte River in northeastern Colorado following high water conditions in 1983 and 1984. Test methods for establishing woody vegetation within streamside riparian zones where natural propagation cannot be expected. These include: (a) using stem cuttings of dormant specimens and planting them so their bases extend into the groundwater as a method for propagation of selected tree, shrub, and vine species for use where natural propagation no longer occurs, (b) create exposed bare ground sites using tillage or scarification for natural establishment of plains cottonwoods, peachleaf willow (Salix amygdaloides), or other woody vegetation present near the site. METHODS 1. Pertinent literature was reviewed concerning facets of riparian ecology. Emphasis was placed on acquiring information concerning stem cutting planting techniques. 2. Statistical analysis of riverbottom inventory data provided under contract by the Colorado State Forest Service was continued after review of procedures with D. C. Bowden, statistician. A paired t test was used to compare change in hectares of cover types between early and recent photo intervals and analysis of variance was used to test for variations among strata. �438 3. Two approaches were used to document conditions conducive to natural regeneration and survival of plains cottonwoods and willows along the South Platte River. The first documented the density, attrition, and site factors in selected permanent transects containing seedlings. The second provided a frequency index of seedling establishment within the lower South Platte riverbottom. a. Several grazed and ungrazed sites were selected along the lower South Platte River for sampling. Sites were mapped and low to high densities of cottonwood seedlings were plotted. These were used in random selection of intensive transects. Steel posts were used to mark the ends of transects that varied in length up to 25 m. A tape, marked at l-m intervals, was extended between the posts. Within high density stands of seedlings, a 0.93-m2 sampling frame was used to tally seedlings at l-m intervals on one side of the tape. In low density stands seedlings were tallied continuously along the tape for 1 m on both sides of the tape. Most transects were established in September 1984 at which time observations on causes of seedling attrition were recorded. A 2.5-3.8-cm (Ld.) perforated plastic pipe was inserted about 1.5 m into the ground within 5 m of several of the transect lines to monitor fluctuations in groundwater levels relative to seedling survival. These are to be measured in May, July, and September and at other times as necessary to monitor groundwater level changes. Site characteristic information recorded at each transect included (1) location (edge of gravel bar, side channel, edge of main channel), and distance and direction from marker; (2) overstory (percent of transect under canopy of trees and shrubs); competition (other seedlings, perennial and annual grasses and forbs, shrubs, trees, and percent bare ground); and (3) livestock grazing (season and intensity). A hand auger was used to compare soil samples to a depth of 0.5 m to discern if pronounced differences in soil texture existed among sites. b. Frequency of cottonwood seedling occurrence was sampled using 30 transects stratified among Division of Wildlife properties along the lower South Platte River in northeastern Colorado. Transects were established along the north side of the river on the Sedgwick Bar, Ceres, and Jones properties and along the south side of the river on the Julesburg easement, Tamarack, and Dune Ridge properties. Odometer readings were used to randomly select starting points on all properties except the Tamarack where hunting unit parking signs were used. Locations where the distance from the river channel to the edge of the flood plain was too narrow « 100 m) were excluded. Since seedling establishment tends to be nonrandom in narrow linear strips paralleling the river, transects were begun at the edge of the flood plain inundated by the 1983 high water and proceeded at approximate right angles toward a marked location at the edge of the river channel. Starting points and compass angles were recorded for future reference. Sampling was diverted or offset around water or other obstructions. �439 A 1-m2 (1 m/side) plot was sampled every 5 steps along the transect. The dominant cover present was recorded in all samples where cottonwood seedlings were absent. Categories included sand bar, bare ground, litter, shrub (species), vine (species), annual grass, perennial grass, annual forb, and perennial forb to provide a general index of the understory vegetation composition. Cottonwood seedlings, when tallied, were recorded as to year of origin (1983 or 1984). Other seedlings (willow, ash, e t c,) were also tallied. Since high water conditions prevented sampling in spring 1984, transects were established and transects were inventoried in September and October 1984. 4. Woody Cover Propagation Procedures a. Stem cutting propagation Preliminary trial efforts within the East Tamarack Check Station Meadow were reported in Snyder (1984). Survival status of 1983 stem cutting plantings was recorded in late spring and early fall 1984. Monitoring of groundwater levels was continued along with assessment of other environmental conditions. Site selection for proposed 1985 stem cutting propagation trials continued through late spring and summer. A 3.8-cm diameter perforated plastic pipe was inserted about L 5 m into the ground at each proposed site to monitor groundwater levels and changes. These were monitored several times through summer and fall 1984 at the South Platte and South Republican sites but time constraints prevented successive readings within sites on the Arkansas and Rio Grande rivers. Selected sites for proposed stem cutting propagation trials in spring 1985 include (with possible modification): (1) South Platte Wildlife Area (East Tamarack: north side west from the Red Lion Road, Area 15 (formerly area 11), Check Station Meadow, and within the west meadow at Duck Creek, (2) the north edge of the Union Tract within the R. Elliott Property (South Platte River); (3) Hale Ponds area and north of the Hale Store within the South Republican Wildlife Area; (4) near the sprayed cattail marsh within the John Martin Wildlife Area and within sedge marshes within the Dean Property along the Arkansas and Purgatorie rivers; and (5) near the Rio Grande River within the Rio Grande Wildlife Property in the San Luis Valley. b. Soil scarification to induce natural reproduction By random selection,· 24 plots within 8 sites along the west and east Tamarack were designated for soil scarification treatment. Each plot approximated 5-10 m by 20 m in size. Site preparation by plowing was initiated in fall 1984 and perforated plastic tubes were inserted at several sites to monitor groundwater level changes. �440 RESULTS AND DISCUSSION Analyses of Cottonwood and Riverbottom Inventory Data Arkansas River.--The Colorado State Forest Service was contracted to use aerial photo interpretation to quantify the condition of and changes in cottonwood stands, shrubs, other vegetation, and land use along the lower Arkansas River from east of Pueblo to the Kansas border. A short span of the South Republican River between Bonny Reservoir and the Kansas border also was sampled. This progress report summarizes the major findings of this inventory. More detailed analysis of the cottonwood stands is in progress by D. C. Bowden and will be included within a future report. The inventory sampled 3,433 ha (8,479.5 ac) within 22 linear miles of a ISO-mile segment of the lower Arkansas River beginning approximately 22 miles east of Pueblo and continuing to the Kansas border. Fifteen miles containing John Martin Reservoir were excluded. The original map of the river valley provided by the State Forest service for use in stratified random sample selection showed 120 miles of river (excluding the reservoir) within the total sample. Twenty-two miles were randomly selected representing an 18.3% sample. However, subsequent detailed mapping by the Colorado State Forest Service revealed approximately 150 miles of river within this same distance. Thus, the sample size decreased to 14.7% and, because of shifts in sample miles among strata, only 11.7% of the lower 60 mile-long stratum (below the reservoir) was sampled. The upper stratum (30 miles) was sampled at 20% and the middle stratum (60 miles) was sampled at 15%. In addition to the 22 miles randomly sampled, one non-random mile at the upper end of John Martin Reservoir just above Fort Lyon, was selected for inventory to benefit management personnel working on the John Martin Wildlife Area. Original year aerial photos ranged from 1940 to 1955 and all recent photos were from 1980 yielding average time intervals of 25.7, 29.1, and 38.6 years respectively for the upper, middle, and lower sampled strata. The overall mean interval was 31.2 years. Changes in area and proportions of sampled cover types varied (Table 1). Cottonwoods declined about 30% during the interval for an approximate loss of 1% per year (p < 0.05, t = 2.90). Shrubs, the most abundant cover type along the river, increased 21% during the interval (P < 0.10, t = 1.78) • Hay meadow and grassland both declined at modest rates Whereas a dramatic conversion to farmland was documented (P < 0.05, t = 3.00) • Developed land, while comprising only 1-2% of the-total, nearly doubled in quantity (P < 0.05, t = 3.00). River and unvegetated (sandbars and mudflats) were -combined for analysis since differing water levels between photo periods could bias results. A marked decline in river area was noted (P < 0.05, t = 5.45). There was a greater reduction of river channel below John Marti; Reservoir than above (P < 0.05, F < = 12.80) (Table 2). This is probably the result of the completion of the reservoir in the early 1950's and the subsequent reduction in flooding, water flows, and water level fluctuations. These observations raise the question as to whether the river channel and water level are deeper below John Martin Reservoir than above? The nearly sediment-free water released from the reservoir has a greater scouring effect than water that is already sediment laden. Over time this should deepen the channel and lower the ground water table in adjacent areas. This would impact much of the riparian vegetation including cottonwoods. Plans are to measure the depth of the water level below adjacent lands at random sites below and above John Martin Reservoir to gain further insight into this variable. �441 Table 1. Proportion and changes (ha) in vegetation types over a 31.2-year interval along the Arkansas River, southeastern Colorado 1940(50)-80. Vegetation type Early interval ha % 647.23 859.62 751.78 549.48 33.96 36.57 211.97 340.73 2.20 Cottonwood Shrub Hay meadow Grassland Agriculture Developed River Unvegetated Standing water Totals *p 18.85 25.04 21.90 16.00 0.99 1.07 6.17 9.92 0.06 3,433.54 < Recent interval ha % 449.05 1,041.05 646.86 495.46 426.26 72.65 225.80 51.86 23.06 13.08 30.33 18.85 14.44 12.42 2.12 6.58 1.51 0.67 Chan~e ha % -198.18 -30.62 181.43 21.11 104.92 13.96 -54.02 -9.83 392.30 1,155.18 36.08 98.66 13.83 6.52 -288.87 -84.78 20.86 948.18 ..t.. df 2.90* 1.78 0.87 0.66 3.00* 3.00* 21 21 16 20 14 10 1.10 8 3,432.05 0.05. Table 2. Average change (ha) of vegetation types/linear mile of river from early to recent intervals along the Arkansas River, southeastern Colorado. Vegetation type Upper Stratum Middle Lower Cottonwood Shrub Hay meadow Grassland Agriculture Developed River & unvegetated -9.00 4.39 -2.17 4.85 7.37 0.93 -9.77 -7.06 9.94 -9.89 -1.28 11.77 3.39 -6.86 -11.24 9.37 0.41 -10.23 34.60 0 -22.10 *p < F value 0.17 0.12 0.29 1.28 1.73 0.87 6.41* 0.05. Comparisons among strata for other vegetation types did not reveal significant differences (Table 2). However, increased farming along the flood plain below the reservoir was apparent. Most pronounced increases in shrub abundance were noted in the middle and lower strata. Cottonwoods showed the greatest decline below John Martin, probably because the time span (38.6 years) was greater there. The loss per year probably would not be greater downstream than upstream if the decline was prorated on a yearly basis. �442 The proportion and changes of cottonwoods by age class and canopy cover along the Arkansas River varied (Table 3). Young trees less than 6 inches (15 cm) dbh comprised only a small percent of the total trees present and decreased only slightly in area occupied over the time interval (! < 0.05). However, nearly all remaining in 1980 were in low density stands as few moderate to high density stands remained (Table 3). Cottonwoods in the 6-15 inch (15-38 cm) dbh category showed evidence of moderate declines (P < 0.05) with major declines in the higher density stands. Trees in the 16-30 inch (41-76 cm) dbh class were dominant among the age classes and showed the greatest decline over time in abundance (P < 0.10). Declines in all density classifications were noted (Table 3). old trees greater than 30 inches (76 cm) dbh, while not abundant, declined (P < 0.05) about 70%. Inspections in 1984 documented considerable cutting of older trees for firewood since the last photo period indicating that an accelerated loss of this age class is occurring. Table 3. Proportion and changes by age class and canopy cover (ha) of cottonwood stands over a 31.2-year intervala along the Arkansas River, southeastern Colorado, 1940-80. Age class (in. dbh) Canopy cover(%) < 35 35-55 > 55 6 Subtotals 6-15 < 35 35-55 > 55 Subtotals 16-30 < 35 35-55 > 55 Subtotals 30 < 35 35-55 > 55 Subtotals All ages Totals < 35 35-55 > 55 Ear1:t:interval % ha 16.63 11.16 13.33 41.12 95.02 110.98 26.06 232.06 189.25 103.65 38.76 331.66 15.32 25.20 1.87 42.39 316.22 250.99 80.02 647.23 Recent interval ha % 6.35 37.96 0 1.86 39.82 35.85 115.06 48.64 16.33 180.03 51.24 147.62 51.61 17.17 216.40 6.55 5.11 3.90 3.79 12.80 305.75 104.15 39.15 449.05 Chan~e % ha ..t. df 8.87 21.33 -11.16 -11.47 -1.30 0.66 0.06 14 40.09 20.04 -62.34 - 9.73 -52.03 26.25 1.05 21 48.19 -41.63 -52.04 -21.59 -115.26 58.16 1.77 20 2.85 -10.21 -21.30 1.92 -29.59 14.93 4.66 13 -10.47 -146.84 -40.87 -198.18 aThe time interval ranged from 25 to 40 years with a mean of 31.18 years. �443 Colorado State Forest Service personnel tried unsuccessfully to distinguish tamarisk (salt cedar) from other shrubs along the Arkansas River using photo interpretation. Subsequently, they randomly selected 43 1/100-acre sample plots within shrub communities and tallied species composition. Tamarisk dominated most sites and represented over 75% of the shrubs and small trees present (Table 4). Sandbar willow (Salix interior) was the second most important shrub with low densities of Russian olive (E1aeagnus angustifo1ia), indigo bush (Amorpha fruiticosa), and a few other species. Table 4. Percent composition of 43 1/100-acre shrub plots obtained by the Colorado State Forest Service, Arkansas River, Colorado, 1984. Stratum Sample Eoints 1 2 3 12 19 12 x shrubsI Eoint 10.9 13.8 21.0 Willow 16.7 5.4 16.7 Percent Tamarisk 75.0 78.9 75.0 Mixed shrubs 8.3 15.7 8.3 Inspections in summer 1984 revealed that much of the tamarisk in the Arkansas Valley had died back, apparently because of extremely cold temperatures during the preceding winter. New basal regrowth was occurring indicating most plants were set back and not killed. Similar die-back of tamarisk was noted along the South Platte. Tamarisk has generally been considered of negative value to wildlife because of its rapid invasion, competitiveness that tends to exclude other vegetation, and difficulties in eradication. The resulting continuous stands are apparently of less value to most wildlife than the more diversified structure provided by cottonwoods, willows, and other species in mixed stands. However, with the apparent declining water table below John Martin Reservoir, were it not for tamarisk, much of the remaining flood plain would contain almost no woody cover. There, it provides protection for deer (Odocoi1eus spp.), a few quail (Co1inus, Ca1iEeE1a), and other wildlife and abundant nesting cover for mourning doves (Zenaida macroura) and several passerines. If edge openings and food production could be increased within and adjacent to it, populations of quail, pheasants (Phasianus), cottontails (Sylvi1agus spp.), and other wildlife would be enhanced. Pueblo Reservoir, completed in the late 1970's, had not impacted composition of upper river strata vegetation at the time of the 1980 aerial photos. However, stabilization of river flows below the reservoir will undoubtedly be similar to that below John Martin with similar impacts. The major difference will be in the greater volume of water carried below Pueblo Reservoir compared to that below John Martin. However, numerous side streams feed into the Arkansas River below either of the two reservoirs. Although several side streams have been dammed, occasional flooding may occur to stimulate some regeneration of woody vegetation. In contrast, if a mainstream reservoir was constructed on the South Platte as proposed, only one major tributary, the Bijou, could occasionally enhance natural revegetation. �444 South Republican River .--Three adjoining miles of river west of the Kansas border to Bonny Reservoir were sampled by the Colorado State Forest Service. Regrettably, the time span was only from 1961 to 1975 and the reservoir had been completed several years prior to the early photo date. Therefore, the combination of small samples and the relative brief time interval yielded data of limited value. Proportions and changes in hectares of vegetation types varied (Table 5) with grassland, cottonwood stands, and farmland being dominant. Grassland and river channel were the only types decreasing during the interval. Table 5. Proportions and changes (ha) of vegetation types over a recent (1961-75) l5-year interval on the South Republican River near Hale Ponds below Bonny Reservoir, Colorado. Ve~etation tJ:Ee Cottonwood Shrub Hay meadow Grassland Agriculture River Unvegetated Totals Early interval % ha 128.44 21.75 0.87 288.73 60.42 12.30 14.00 526.51 24.39 4.13 0.16 54.84 11.48 2.34 2.66 Recent interval % ha 188.51 37.17 6.84 168.02 113 .21 7.07 5.69 35.80 7.06 1.30 31.91 21.50 1.34 1.08 Change ha 60.07 15.42 5.97 -120.71 52.79 -5.23 -8.31 % 46.76 70.90 686.21 -41.81 87.37 -42.52 -59.36 526.51 The tracts contained only a small proportion of young cottonwoods in 1961 and no young trees were inventoried from the 1975 photos indicating a lack of cottonwood reproduction (Table 6). Moderate increases of older trees were noted. Stands were more evenly distributed among the three density classes than were those along the Arkansas River. Deepening and narrowing of the stream channel below Bonny Reservoir appears to be a factor that is potentially lowering the water table in adjacent riparian zones. Random measurements to confirm or deny this are in the planning stage. A series of small metal check dams could be installed, at least on CDOW land to raise water levels assuming livestock grazing was discontinued. Attempts to locate an earlier series of aerial photos dated prior to construction of Bonny Reservoir will be made. However, it is assumed that Lfvestock grazing has been continuous along the South Republican for many years. Therefore, it is doubtful that dense stands of cottonwoods were ever abundant even though occasional flooding, essential to their establishment, occurred. �445 Table 6. Proportion and changes by age class and canopy cover (ha) of cottonwood stands over a 16-year interval (1961-75) in the Hale Ponds vicinity, South Republican River, Colorado. Age class (in. dbh) Canopy cover(%) < 35 35-55 >55 6 Subtotals 6-15 < 35 35-55 >55 Subtotals 16-30 < 35 35-55 >55 Subtotals 30 <35 35-55 >55 Subtotals All ages Subtotals <35 35-55 >55 Ear1~ interval ha % 5.63 1.85 0 7.48 22.22 17.41 3.60 43.23 25.20 26.04 23.95 75.19 1.27 1.27 0 2.54 54.32 46.57 27.55 128.44 5.82 Recent interval ha % 0 0 0 0 0 Chan~e ha -5.63 -1.85 0 -7.48 33.66 42.21 21.48 2.08 65.77 58.54 24.56 41.02 46.44 112.02 1.98 0 8.09 2.63 10.72 -1.27 6.82 2.63 8.18 66.77 70.59 51.15 188.51 12.45 24.02 23.60 60.07 34.80 19.99 4.07 -1.52 22.54 59.42 -0.64 14.98 22.49 36.83 South Platte, Colorado, and Rio Grande Rivers.--A summary of the preliminary inventory findings for these rivers was provided in Snyder (1984). More detailed analysis has continued. Currently, a multivariate analysis of cottonwood age class and density classes is in progress by D. C. Bowden. Findings will be presented in a future report. Changes from early to recent photo intervals for all vegetation types were compared statistically (Tables 7-9). Within the South Platte River, hectares of cottonwoods decreased (P < 0.10) whereas the decrease in shrubs was not significant. All other cover types changed markedly (Table 7). Early to recent changes in hectares within cover types along the Colorado River were less dramatic than along the South Platte (Table 8). Cottonwoods decreased (P < 0.10) as did the hay meadow type. Major increases occurred within developed and standing water cover types (P < 0.05). Within the Rio Grande riverbottom, narrow1eaf cottonwoods (Populus angustifolia) increased but not at a significant rate (~ < 0.10). There were no changes in cover of developed and standing water. Major decreases occurred among shrub, hay meadow, and river channel types (p < 0.05) whereas grassland and agriculture showed marked increases (~ < 0.05). - �446 Table 7. Changes (ha) in vegetation types over a 36-year interval along the South Platte River, northeastern Colorado, 1941-79. Vegetation Cottonwood Shrub Hay meadow Grassland Agriculture Developed River + Unvegetated Standing water *p up < < Early interval Recent interval DF t 1,806.5 490.8 2,141.0 128.5 492.6 31.6 343.6 5.4 1,638.8 388.9 1,177.1 392.9 1,068.5 104.0 595.0 80.8 28 28 28 20 20 18 28 25 1.76* 1.67 5.33** 3.23** 3.87** 1.88* 5.45** 2.47** 0.1 • 0.05. Table 8. Changes (ha) in vegetation types over a 25-year interval along the Colorado River, western Colorado, 1955-80. Early interval Vegetation Cottonwood Shrub Hay meadow Grassland Agriculture Developed River + Unvegetated Standing water *p **p < < 377.5 320.9 495.2 104.8 184.3 22.3 313.6 4.5 Recent interval DF t 311.6 341.9 378.0 137.2 170.8 108.8 197.2 77 .5 20 20 17 19 6 16 20 11 1.74* 0.93 1.94* 0.71 0.55 3.38** 0.87 3.00** 0.10. 0.05. Table 9. Changes (ha) in vegetation types over a 3l-year interval along the Rio Grande River, San Luis Valley, Colorado 1941-83. Vegetation Early interval Recent interval DF t Cottonwood Shrub Hay meadow Grassland Agriculture Developed River + Unvegetated Standing water 559.8 210.7 2,208.5 27.9 3.5 22.3 198.2 32.5 611.2 158.6 1,753.5 99.9 435.0 32.6 125.3 37.3 15 19 19 10 9 15 19 17 1.46 2.31* 2.79* 2.42* 3.05* 1.43 6.86* 0.56 *p < O. 5. �447 Analyses were conducted to detect differences for individual cover types among strata along the inventoried segments of the South Platte, Colorado, and Rio Grande rivers. For most, no differences among strata were detected (Table 10), however, cottonwood and river channel types showed differences along the South Platte and river channel and standing water showed differences along the Colorado River (Table 11). Cottonwoods increased in stratum 2 (from near Weldona downstream to near Merino) whereas hectares of cottonwoods decreased both upstream and downstream. The river channel (river plus unvegetated sandbars) also showed less of an increase in stratum 2 than either upstream or downstream. Among strata along the Colorado River, river channel increased significantly in stratum 2 (Rifle-Rulison area) compared to decreased channel width in strata 3 and 4 (Table 11). The amount of standing water increased significantly in the lower stratum (Clifton-Grand Junction area) compared to the 2nd stratum. Table 10. Early to recent changes among sampling Platte, Colorado, and Rio Grande rivers. South Platte DF F value Vegetation Cottonwood Shrub Hay meadow Grassland Agriculture Developed River + Unvegetated Standing water *p < 3,25 3,25 3,25 3,17 3,17 3,15 3,25 3,22 DF 3.92* 0.84 0.11 3.36 1.09 0.57 5.01* 0.85 strata along Colorado F value 13,17 3,17 3,14 3,16 0.52 0.48 2.72 1.90 3,13 3,17 1, 9 0.93 4.73* 5.75* the South Rio Grande DF F value = 2,13 2,17 2,17 2, 8 2, 7 2,13 2,17 2,15 0.85 0.53 3.18 2.89 3.31 2.92 4.48 1.22 0.05. Table 11. Differences among strata for changes from early to recent intervals within certain cover types along the South Platte and Colorado rivers. River Cover type South Platte Cottonwood x Stratum 2 9.74 4 x -13.14 3 -13.15 a Difference River channel 1 0.16 2 0.31 3 1 4 8.51 12.89 14.16 Difference Colorado River channel x Difference Standing water x 2 1 4 3 3.00 1.16 -2.79 -3.60 2 1.11 4 9.85 1 3 Insufficient data aUnderscoring of two or more means denotes no significant difference. �448 Natural Establishment and Survival of Cottonwood Seedlings South Platte River .--Several preliminary 25-m transects were established in early fall 1983 where cottonwood seedlings had become established following flooding in spring-summer 1983. These transects, while not randomly selected, provided a general measure of seedling survival from summer 1983 to fall 1984 (Table 12). Major attrition occurred and fall 1984 survival was only 2.2% of that of the previous fall. Observations indicated that while some seedlings were lost over winter, major mortality occurred during a prolonged period of high water in spring 1984. This was followed by a rapid early July decrease of the river, with accompanying groundwater declines in July and August. Seedlings on higher sites not inundated by spring flooding were desiccated by dry, late-summer conditions. Additional transects established in fall 1984 (Table 13) indicated that most 1983 seedlings were in relatively sparse stands. Comparison of extensive transects (Tables 14, 15) documents the loss as cottonwood seedlings germinating in 1983 occurred in 14.8% of 1,055 m2 samples in fall 1983 but in only 10 of 1,667 samples (0.60%) in 1984. Table 12. Comparisons of cottonwood seedling density/transect and per 0.09 m2 sample along the South Platte River, northeast Colorado, 1983-84. Site Transect Sedgwick Bar Haxtun R&G Tamarack Jones Fall 1983 Total! transect i/0.09m 1 2 3 1 8-9E 2W 4W 4-5W l7W Inter. Exter. Totals Mean 1983 to 1984 survival = Fall 1984 Total! transect x/0.09m 56 29 67 234 68 25 123 81 98 76 49 2.24 0.58 2.67 9.36 2.72 1.00 4.92 3.24 3.92 3.04 1.96 2 0 6 1 0 0 8 0 0 0 1 0.08 906 3.29 20 0.07 2.21% aEach transect contained 25 0.093 m2-samp1es spaced 1 m apart. 0.24 0.04 0.32 9.94 �449 Table 13. Cottonwood seedling density along South Platte River transects in fall 1984. Site Transect Samples/ transect Total seedlings (m2) Seedlings established in 1983 Sedgwick Bar Haxtun R&G Tamarack Bravo 4 2 4 2W 4W1 4W2 4W3 6Wl 6W2 21W 22W 24W 1 2 x density 25 9 25 25 25 20 7 17 25 25 16 25 25 25 69 82 28 3 54 27 35 172 199 13 92 51 268 67 2.76 9.11 1.12 0.12 2.16 1.35 5.00 10.12 7.96 0.52 5.75 2.04 10.72 2.68 (0.09m2) Tamarack 9W 25 48 (m2) Seedlings established in 1984 Tamarack Bravo 13Wl l3W2 3 1.92 25 25 18 360 84 158 14.40 3.36 8.78 (0.09m2) Sedgwick Bar Haxtun R&G Tamarack Bravo 5 3 2E 6W3 6W4 3 25 25 17 20 25 25 94 195 48 216 334 106 3.76 7.80 2.82 8.64 13.36 4.24 aSamples were either 0 •09 or 1 m2 based on seedling density along the transect. �450 Table 14. Frequency of selected transects along October 1983. Site occurrence the South Location (m2) of cottonwood seedlings within Platte River, northeastern Colorado, Samples Seedling occurrence Sedgwick Bar West, north side Middle, north side East, north side 175 140 140 19 9 40 Tamarack 8E, south side 5E 3-4E 5W 150 150 150 150 19 7 23 39 1,055 156 Totals Frequency of occurrence (%) 14.79 �Table 15. Frequency of occurrence (m2) of cottonwood seedlings, other seedlings, and other cover typesa obtained within random transects along the South Platte River, northeastern Colorado, September 1984. Site Transect Julesburg Sedgwick Bar Haxtun R&G Tamarack Bravo Dune Ridge Jones Totals E-1 E-2 H-1 W-2 W-3 1 2 3 4 1 2 3 4 E-6 E-5 E-3 E-1 W-1 1~-2 W-11 W-17 W-18 \01-19 1 2 3 1 2 1 2 Cottonwoods Alive Dead 0 0 0 2 3 2 0 0 0 2 5 1 0 0 2 0 1 0 1 0 4 0 1 1 2 0 1 2 1 1 32 Others Forbs Grass Annu. Perro . Annu. Perro Shrubs Low Tall Vines Trees 1 0 0 1 11 0 0 0 5 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1 0 0 0 1 0 0 0 0 1 0 0 0 0 0 2 0 0 0 1 0 0 0 11 4 4 3 16 7 8 13 10 11 26 32 7 12 29 31 15 4 5 10 16 6 21 11 25 6 10 8 18 20 8 5 9 8 16 1 4 0 2 8 7 15 7 5 12 6 3 4 1 1 6 12 10 2 11 7 3 6 7 6 3 3 1 5 5 2 4 3 4 9 13 15 3 12 13 10 8 2 7 1 2 0 1 8 3 4 2 1 1 1 18 10 13 25 18 4 14 4 12 6 10 10 8 25 21 21 35 8 2 9 19 5 20 0 7 1 3 20 27 27 9 5 2 3 11 0 0 0 0 2 4 1 0 23 15 3 1 8 2 6 4 22 13 0 3 3 0 0 5 10 1 4 3 12 9 2 10 0 1 4 2 0 8 16 15 3 14 2 0 2 8 3 3 5 13 8 0 13 7 7 0 3 1 5 2 0 0 1 0 3 1 0 0 8 1 2 0 1 3 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 20 8 399 192 146 402 155 175 32 4 Bare Grnd. Sand Bar 0 0 0 3 0 0 4 0 1 1 1 5 0 0 0 0 1 0 1 0 0 1 0 1 1 0 0 4 0 0 2 0 0 0 5 10 4 0 2 17 6 4 0 0 8 1 0 0 1 0 0 1 1 0 5 0 2 0 0 0 0 0 0 0 1 1 0 0 0 1 2 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 69 8 24 Litter Totals 53 35 33 67 98 29 49 22 37 65 79 85 33 101 117 77 81 29 23 29 60 50 72 28 70 30 22 55 66 72 1,667 aCover types were classified by the dominant cover present where more than one type was present within a m2 sample. bOn1y 10 of 32 cottonwood seedlings were from 1983. .j:-. lJ1 i-' �452 Spring flooding in 1984 was much less conducive to cottonwood seedling establishment than in 1983. A cold late spring slowed seed maturity and dispersal. Water levels began to decrease in June prior to major seed dissemination and then decreased rapidly in early July. As a consequence, seedlings germinating in 1984 were along main stream and side channels at lower levels than those established in 1983. Therefore, they were much more vulnerable to inundation and destruction by subsequent high water. Some were probably lost during sporatic high water in fall 1984. Seedlings of peachleaf willow and sandbar willow were generally less abundant than those of cottonwoods but were common following 1983-84 high water conditions. Seedlings of frost grape (Vitis vulpina), red ash (Fraxinus pennsylvanica), and Russian olive were less common and primarily were observed following the high water in 1983. The primary key to cottonwood-willow seed germination and seedling establishment is persistent wet soil conditions. Seedlings were observed establishing in sites of dense herbaceous vegetation, such as within the Check Station meadow on the Tamarack. Seedlings have repeatedly been observed growing on a tar and gravel building roof where moistures condensation from an air conditioner provided persistent wet conditions. Seedling growth and survival are dependent on more complex combinations of variables. Soils within the South Platte flood plain in northeastern Colorado are generally classed as Fluvaquents (Amen et al. 1977) and composed of bands and lens of sand, gravel, loam, and clay giving way to sand and gravel subsoil at less than 51 cm below the soil surface. Soil auger inspections revealed that sites containing a thicker strata of loam, clay-loam or similar soils above sand or gravel generally favored seedling survival over sites dominated by sand and gravel. Apparent reasons included better early growth and vigor allowing the larger seedlings to tap deeper into the soil so that they could survive when water tables decreased as in July and August 1984. Heavier soils also have greater capillary action and can hold subsoil moisture for use by seedlings better than sands and gravels. The tradeoff is that the better soils often contain dense herbaceous vegetation making it more difficult for seedlings to establish and to survive competition for available moisture. Perennial grasses were the most common dominant vegetation occurring within sampled extensive transects in September (Table 15). Annual forbs ranked second. The high composition of forbs, many of which provide food for seed-eating wildlife, was in part a product of the soil disturbance created by flooding in 1983 and 1984. Natural Seedling Establishment in Other Locations.--High water and flooding along the Colorado River and tributaries induced limited natural propagation of plains cottonwoods, willows, and tamarisk at lower elevations. However, high water persisted past the primary season of seed dissemination in 1984 and undoubtedly washed out or killed many of the seedlings established in 1983. Livestock browsing of seedlings is a major factor reducing natural regeneration in many western Colorado sites including CDOW properties. Little or no seedling establishment was noted along the Rio Grande or lower Arkansas rivers during late July 1984. �453 Stem Cutting Propagation Trials Trial stem-cutting propagation conducted in late winter 1984 resulted in fair to good survival to fall for willows, fair survival for plains cottonwoods, and poor to no success for other species (Table 16). Cotoneaster (Cotoneaster spp.) showed preliminary leafing but little subsequent evidence of success, whereas elderberries (Sambucus spp.) leafed out and persisted until late spring before dying. Some cuttings of both elderberry and frost grape may have been severely injured or partially dead prior to planting due to the severe temperatures in winter 1983-84 since major die-back of uncut stems was noted in spring. Other possible factors in reduced survival may have included timing of the effort (late Feb), high soil pH and salt content, and high water levels in spring (Table 17). The groundwater level was close to the soil surface within the Check Station Meadow for several weeks through spring and at several sites, primarily those containing willows, emerged above ground so that the stem cuttings were standing in water. Dense vegetation growth, primarily sedge, and perennial grasses, crowded and covered the shorter stem cuttings, primarily willow by late summer so that it was difficult to locate them to assess survival and growth. Some may have been stressed for water in July and August because of competition, however, mortality had occurred prior to that for most. Pumping operations during placement of an underground pipeline along the west edge of the study site in October temporarily decreased the water level but its negative impact on the stem cuttings is doubtful. Table 16. Survival of stem cutting plantings of selected tree, shrub, and vine species on the Check Station meadow, South Platte Wildlife Area, 1984. SEecies Populus sargentii Salix amygdaloides S. interior S. exigua Salix sp- (Golden willow) Amorpha fruiticosa Sambucus sp. Cotoneaster sp. Vitis vulpina Parthenocissus guinguefolia Site 1 2 1 1 1 2 1 2 3 1 1 2 1 2 1 1 Number planted 6 25 6 12 25 10 12 12 15 15 12 12 15 12 12 4 Survival 6 Jun 9 Sep 4 14 1 10 of 12 of 6 of 10 of of 0 0 0 0 0 0 0 5 12 a 9 12 8 of 12 a 4 of 6 a 9b 0 0 0 0 1 9 7 6 2 la 11a 10 14 6 aDense vegetation concealed plantings, especially those that had died making them difficult to find. bMost were in poor condition when inventoried. �454 Table 17. Groundwater levels (cm) below the soil surface at four stem cutting planting sites within the Check Station Meadow, South Platte Wildlife Area, 1984. Site POEu1us Sambucus Cotoneaster Salix exigua 1 1 1 1 1 Mar 28 Mar 27 AEr 20 May 6 Jun 5 Ju1 29 Aug 1 Oct 10 Nov 61 63 43 13 71 43 18 48 46 25 0 28 25 13 0 61 36 42 13 69 76 58 23 69 66 51 25 109 86 64 66 51 18 aThe river began rising a few days prior to the 29 August measurement after being nearly dry since mid-July. bGroundwater was being pumped out temporarily during laying of a nearby pipeline across the river. A July 1984 inspection of the Colorado, Rio Grande, and Arkansas rivers and CDOW properties along them revealed limited opportunity for stem cutting propagation. Test sites were selected for limited stem cutting planting of narrow1eaf cottonwoods within the Rio Grande Property along the Rio Grande River in the San Luis Valley. However, there is no need for additional cottonwoods within the property. Opportunities for stem cutting plantings along the Arkansas are limited. The water table appears to be so low in many sites below John Martin Reservoir, that deep drilling and large samplings would be needed for planting of cottonwoods there. This, however, would be feasible (E. Swensen, pers. commun.) if the proper equipment was available. Nearly all work would have to be conducted on private lands and therefore would be dependent on landowner acceptance, livestock grazing, and other factors. Better opportunities exist on CDOW properties upstream where seepage from irrigation perpetuates wet soil conditions conducive to planting small clumps or thickets of shrubs, trees, and vines. LITERATURE CITED Amen, A. E., D. L. Anderson, T. J. Hughes, and T. J. Weber. 1977. Soil survey of Logan County, Colorado. U.S. Dep. Agric., Soil Conserv. Serv., Washington, D.C. 252pp. Snyder, W. D. 1984. Lowland riparian cottonwood community studies. Job Prog. Rep., Colorado Div. Wi1d1., Wi1d1. Res. Rep., Fed. Aid Proj. N-4-R-2. Jan.:292-370. Prepared by :-:----6*-''''''''""-::~~?_;W-=:-......;.J:;....' --"""""~--+'-=~ Warren D. Snyder Wildlife Researcher �455 Colorado Division of Wildlife Wildlife Research Report April 1986 JOB PROGRESS REPORT State of Colorado --~~~~----------------------01-03-045 (W-37-R) (N-4-R) Project Work Plan Job Title: 21 2 ---- Dynamics of Cottonwood Regeneration Period Covered: Author: : Job Avian Research 01 January through 31 December 1985 Warren D. Snyder Personnel: C. E. Braun, D. C. Bowden, J. Palic, M. W. Stanley, and W. D. Snyder, Colorado Division of Wildlife ABSTRACT Photo-interpretation inventory of a small segment of the South Republican River revealed few plains cottonwoods (Populus sargentii) were present along the stream in the late 1930's and dramatic increases have occurred since then. In contrast, shrubs are less common now than during the initial inventory. Cross-section stream profile sampling indicated the stream channel below Bonny Reservoir was deeper than in upstream samples (p < 0.05). Annual survival to late summer 1985 among cottonwood seedlings along;the South Platte River was 61% for those established in 1983 and 20% for 1984 seedlings yielding an overall average of 31% within 23 transects. Mortality was higher in grazed than in ungrazed sites but desiccation, rather than browsing was considered the primary reason for attrition. Extensive transects conducted along the South Platte supported findings within the intensive transects and showed perennial grasses recovered from flooding in previous years while annuals declined in abundance. Groundwater levels averaged lower along the South Platte and South Republican rivers in 1985 than in 1984 but higher along the Arkansas River within seepage areas. Stem cutting survival was poor in all locations. Most cuttings drowned along the Arkansas River and early summer water table declines along the other drainages apparently desiccated stem cuttings. Few seedlings established within soil scarification plots because groundwater was not high enough to support germination in most plots. Preliminary efforts to evaluate the impact of controlled burns within a sparsely timbered area along the South Platte River were initiated. ��457 DYNAMICS OF COTTONWOOD REGENERATION Warren D. Snyder P. N. OBJECTIVES Quantify changes in stand density and changes in area of riparian cover types over a recent time span approximately 30 years within the South Platte, Arkansas, Colorado, Rio Grande, and South Republican (below Bonny Reservoir) rivers in Colorado. Analyses and evaluations will be based on aerial interpretation by the Colorado State Forest Service. Document conditions conducive to natural regeneration and survival of plains cottonwoods and willows, including site characteristics and frequency of occurrence, in streamside riparian habitats of the lower South Platte River in northeastern Colorado following high water conditions in 1983 and 1984. Test methods for establishing woody vegetation within streamside riparian zones where natural propagation cannot be expected. These include: (a) using stem cuttings of dormant specimens and planting them so their bases extend into the groundwater as a method for propagation of selected tree, shrub, and vine species for use where natural propagation no longer occurs, (b) create exposed bare ground sites using tillage or scarification for natural establishment of plains cottonwoods, peachleaf willow (Salix amygda10ides), or other woody vegetation. METHODS Primary methods used in this study were reported earlier (Snyder 1985). To obtain stream profile data for the South Republican, a map of the stream was gridded at 400-m intervals. These intervals were numbered for random selection of 12 points respectively below and above Bonny Reservoir in Yuma County. A 30-m cord, marked at 1-m intervals was held level across the stream and vertical distances to the stream were measured. Evaluation of controlled burning within the South Platte riverbottom included late May and early September post-burn inventory of survival and regrowth status of trees and tall shrubs within the Brush Wildlife Area burned in April 1985. In addition pre-treatment inventory and status data were obtained for trees and tall shrubs within proposed 1986 burn sites. Pre-burn status of cottonwood seedlings within the proposed spring 1986 burn site on the Brush Wildlife Area was sampled using the intensive transect method (Snyder 1985). RESULTS AND DISCUSSION Analyses of Cottonwood Inventory Data Major Rivers.--Pre1iminary analyses of the photo-interpretation inventory data concerning trends and status of cottonwoods, shrubs, and other vegetation or land use along the lower portions of Colorado's 4 major rivers were reported earlier (Snyder 1984, 1985). However, because cottonwood samples were �458 stratified and inventoried by 3 canopy covers and 4 age-class levels, additional analyses were warranted. David C. Bowden, statistician, completed portions of these analyses and review of the findings revealed that additional analyses were needed. Therefore, results will not be reported until the entire analyses have been completed. South Republican.--An inventory of 4.8 kg (3 mi.) of the South Republican River below Bonny Reservoir in southeastern Yuma County was conducted by the Colorado State Forest Service (Snyder 1985). However, the interval of comparison was 15 years (1961-75) and Bonny Reservoir was about 10 years old at the time of the earliest interval. In an effort to find earlier aerial photos, the U.S.D.A. Soil Conservation Service office in Wray was contacted and photos dated 12 January 1938 were obtained. These provided a time span of 37 years (38 growing seasons) and provided insight into stream, cottonwood, and cover type conditions prior to construction of Bonny Reservoir (Tables 1, 2). The photos and findings showed that few cottonwoods were present within the sampled area in 1938 and those present were primarily in open stands «35% canopy cover). All age-classes were well represented within the 1938 sample (Table 1) in contrast to lack of young « 6" dbh) trees in the 1975 sample. However, cottonwoods increased dramatically from 19.8 ha in 1983 to 188.5 ha in 1975. There were 128.4 ha of cottonwoods in 1961 revealing that substantial increases occurred both prior to and after 1961. Whywere few cottonwoods present within the sample miles in 1938? The reasons are not clear but sandy, highly erodable soils easily trampled by livestock may have been a factor. Browsing of seedlings by livestock undoubtedly was also important. Above average precipitation in the 1940' s may have reduced the intensity of streamside grazing and browsing, and may have increased streamflows and flooding. However, this is uncertain. A major flood occurred along the stream in 1935 and its influence on cottonwoods is uncertain. The impact of the Reservoir is not fully understood because seedling establishment apparently occurred both prior to and following its construction. The South Republican River, based on the 1938 aerial photos, was a broad shallow fluctuating stream with extensive unvegetated sandbars. Apparently, little water was present at the time of the January 1938 photos. The Arikaree River, where it intersects U.S. 385, is assumed to typify the appearance of the South Republican at that time. Sandbars have decreased dramatically in recent years as the upstream reservoir has stabilized stream flow (Table 2). Stream cross-section profile samples obtained below and above Bonny Reservoir in Yuma County reveal the stream channel tends to be deeper below the reservoir. Mean depth of the channel averaged 6.6 dm below Bonny Reservoir and 4.5 dm upstream (! 22 = 1.41, .!: > 0.05). The 7 deepest measurements per sample were averaged and the downstream and upstream means were compared. Downstream means averaged 13.6 dm contrasted to 7.7 dm upstream (t22 = 2.70, .!: < 0.05) indicating the channel has deepened in recent years dOwrlstream from the reservoir. Livestock grazing, which tends to erode stream banks and flatten channels, was present in both upstream and downstream locations. There was a marked decrease in hectares of shrubs from 1938 to 1975 (Table 2). Both sets of data are probably inflated as shrubs in recent years are primarily confined to stream banks and scattered open stands in a few other locations. Livestock grazing has suppressed shrub growth as is readily noted �459 Table 1. Proportion and changes by age-classes and canopy cover (ha) of cottonwood stands over a 38-year interval (1983-75) in the Hale Ponds vicinity, South Republican River, Colorado. Age class (in. dbh) <6 Canopy cover (%) <35 35-55 >55 <35 35-55 >55 Subtotals 16-30 <35 35-55 >55 Subtotals >30 <35 35-55 >55 Totals o o o 42.21 21.48 2.08 o o 34.3 1.31 65.77 2.28 18.1 112.02 2.65 o 1.26 8.09 2.63 o 19.7 10.72 -5.53 35.42 21.48 2.08 34.9 24.56 41.02 46.44 o 3.91 <35 35-55 >55 27.9 Change ha -4.03 -1.50 o 6.79 3.59 Subtotals All ages o 6.79 Recent interval ha % o o 4.03 1.50 5.53 Subtotals 6-15 Early interval ha % 58.98 23.25 41.02 44.16 59.4 108.43 -2.65 8.09 1.37 5.7 6.81 14.78 1.50 3.54 66.77 70.59 51.15 51.99 69.09 47.61 19.82 188.51 168.69 �460 Table 2. Proportions and changes (ha) of vegetation types over a 38-year interval (1938-75) in the Hale Ponds vicinity, South Republican River, Colorado. Earl~ interval ha % Vesetation Cottonwood Shrub Hay Meadow. Grassland Agriculture Developed River Unvegetated 19.82 137.40 25.45 206.71 61.04 0.97 1.49 73.63 3.76 26.10 4.84 39.26 11.59 0.18 0.28 13.98 Recent interval % ha 188.51 37.17 6.84 168.02 113.21 0 7.07 5.69 35.80 7.06 1.30 31.91 21.50 0 1.34 1.08 Change ha 168.69 -100.23 -18.61 -38.69 52.17 -0.97 5.58 -67.94 % 851.11 -72.95 -73.12 -18.72 85.47 -100.00 374.50 92.27 by direct observation. Russian olive (Elaeagnus angustifolia) in open stands may have increased in recent years and may be interpreted as shrubs in aerial photos. A few tamarisk (Tamarix sp.) exist northeast of the Hale Ponds near the Kansas border, but are not healthy or increasing in abundance at present. Natural Establishment and Survival of Cottonwood Seedlings Intensive Transects.--Cottonwood seedling survival for 23 transects averaged 35% from September 1984 to September 1985. Transect 1 on the Bravo Property (Table 3) was excluded from this and subsequent analyses because an upstream sand dam diverted water from a river channel resulting in total seedling mortality at the site. Annual survival from September 1984 to 1985 was better (61%) for 1983 seedlings than for those established in 1984 (20%). Several factors, including larger seedlings with deeper root systems and thinner stands, were believed responsible for better survival of 1983 seedlings. Most 1984 seedlings were on lower sites than those where 1983 surviving seedlings occurred because the river level was lower during seedling germination in 1984. Livestock grazing, primarily in winter, occurred on 11 of 23 intensive transects. Among 1983 seedlings, attrition was 41% in grazed transects and 36.7% in ungrazed. Annual attrition was 91.1% among 1984 seedlings within grazed sites contrasted to 57.2% in ungrazed sites. Based on 0 bservations, most loss of seedlings to livestock, if it occurred, was due to trampling rather than browsing. The seedlings were not leafed-out when livestock were present and therefore, were not attractive as forage. Although the above data indicate increased mortality in grazed sites, this remains uncertain. Observations indicate most seedling attrition occurred due to desiccation during mid- to late summer when the South Platte River dropped to minimal stream flows and adjacent ground water levels declined. Extensive Transects.--Cottonwood seedlings were found within only 13 of 30 extensive transects where 1,779 m samples were obtained in September 1985 �Table 3. Cottonwood seedling density (m2) along South Platte River transects in fall 1984 and 1985. Transect /I Site Grazing status Samp1es/ transect Grazed Ungrazed Ungrazed Ungrazed Grazed Grazed Grazed Grazed Grazed Ungrazed Ungrazed Ungrazed Ungrazed Ungrazed Ungrazed 25 17 25 25 16 25 25 25 25 54 27 35 172 199 13 92 51 268 67 48 Grazed Grazed Ungrazed Grazed Ungrazed Ungrazed Grazed Grazed Ungrazed 25 25 18 25 25 17 20 25 25 360 84 158 94 195 48 216 334 106 Total seedlings 1984 1985 Mean density 1984 1985 a Seedlings established in 1983 Sedgwick Bar Haxtun R&G 4 2 4 Tamarack 2W 4W1 4W2 4W3 6W1 6W2 21W 22W 24W Bravo 1 Tamarack 2 9W 9 25 25 25 20 7 69 82 28 3 29 24 4 1 26 3 24 129 119 8 81 44 Ob 54 27 2.76 9.11 1.12 0.12 2.16 1.35 5.00 10.12 7.96 0.52 5.75 2.04 10.72 2.68 21.33 2.16 12.00 14.40 3.36 8.78 41. 78 86.67 31.33 96.00 148.44 47.11 0.08 0.04 2.83 0.44 42.22 5.89 31.11 16.44 27.56 1.16 2.67 0.16 0.04 1.04 0.15 3.43 7.59 4.76 0.32 5.06 1. 76 Seedlings established in 1984 Tamarack Bravo Sedgwick Bar Haxtun R&G Tamarack Bravo 13W1 13W2 3 5 3 2E 6W3 6W4 3 2 1 51 1 95 9 56 37 62 .s:-- aSamp1es were either 0.09 or 1 m2 based on seedling density along the transect. bNearby channel had been dammed upstream diverting water from near the transect. 0'1 I-' �462 along lines running perpendicular to the lower South Platte River (Table 4). In comparison, seedlings were found in 17 of 30 transects in September 1984. Seedlings established in 1983 or 1984 occurred 32 times in 1984 sampling (Snyder 1985) but only 17 times in 1985. However, several encounters of cottonwood seedlings established in 1985 were recorded bringing the total seedling occurrences to 26. Most seedlings established in 1985 were in low sites where they would be inundated by subsequent high stream flows so they are not expected to contribute significantly to long-term cottonwood stand regneration. Young green ash (Fraxinus pennsy1vanica) were recorded in 5 sample frames in 1985 showing their increasing establishment and importance along the South Platte in northeastern Colorado. Where tree seedlings were not encountered, the dominant vegetation within other sample frames was recorded (Table 4). The primary decrease was in annual forbs which represented 23.9% of the occurrences in 1984 but only 14.6% in 1985. Perennial grass occurrences increased from 24.1% in 1984 to 31.4% in 1985. This transition from annuals to perennials would be expected as the latter recovered from the 1983 and 1984 floods. The 1983 flood inundated nearly all flood plain sites and persisted from early spring until mid summer so the impact was considerable. The 1984 flood was not as extensive nor as long lasting and the impact was much less. River Volume and Groundwater Measurements The lower South Platte River in northeastern Colorado contained considerable water throughout 1984 except for a brief July-August interval when it was depleted by irrigation demands. However, it declined in early winter 1985 and remained low until 28 April. It filled to about one-half full or better in late April and remained fairly stable in volume until mid-June when irrigation demand again depleted it to low flow. Upstream rains in mid to late July provided a temporary flow increase but it remained low through most of August with fluctuating volume increases in September and into the fall with the termination of irrigation. Groundwater levels tended to fluctuate in unison with river levels. Measurements were not made frequently enough to depict actual river level changes but data from the Red Lion, Tamarack 11-E, and Tamarack check station meadow sites show early spring and mid-summer declines in 1985 (Fig. 1). They also show that river and water table levels were higher in 1984 than in 1985. The Duck Creek site received seepage water from the canal leading into Julesburg Reservoir so fluctuations there reflect a combination of canal flows and irrigation demands. Water levels in June 1984 were higher than in June 1985 within stem cutting sites on the South Republican River in southeast Yuma County (Fig. 2). The difference was believed at least partially due to above average January-April precipitation in 1984 when 23.5 cm (0.26 in.) was received contrasted to 5.5 cm (2.16 Ln, ) in 1985. Groundwater levels fell to similar low late-summer levels during both years. Groundwater levels were recorded at several other locations along the South Platte River (Table 5). Tubes were also installed in 1984 at 2 stem cutting sites within the Arkansas Valley. However, they could not be measured frequently enough to yield meaningful data. �Table 4. Frequency of occurrence <m2) of cottonwood seedlings, other along the South Platte River, northeastern Colorado, September 1985. Site Julesburg Sedgwick Bar Transect E-1 E-2 W-1 W-2 W-3 1 Seedlings Cottonwood Other 5c 1b 1 2 3 4 Haxtun R&G 1 2 3 4 Tamarack Bravo 1 2 Jones 3 2 2 E-6 Percent 4 7 2 5 4 21 3 4 3 10 7 7 9 4 3 4 6 1 11 1 3 1 11 11 27 18 5 6 29 45 29 27 5 38 29 3 3 5 5 6 2 1 10 5 10 11 3 6 13 5 5 2 3 1 5 1 9 5 9 11 1 8 3 259 139 14.9 8.0 10 19 4 24 3 29 16 4 5 7 4 14 6 5 15 6 2 7 7 8 18 8.4 6 4 Down timbo 1 1 3 1 4 1 1 2 1 1 4 2 1 2 2 8 1 5 11 7 6 2 6 5 14 9 5 2 1 1 9 2 1 3 1 transects Litter 1 1 1 4 1 1 2 3 7 2 1 20 4 4 10 558 222 199 12.7 11.4 1 40 2.3 15 0.9 38 61 27 43 1 88 59 106 136 75 67 30 23 31 60 53 1 64 1 31 66 33 2 1 1 22 54 1 1 66 74 1 18 63 43 48 73 101 77 3 6 1 Total 67 2 3 1 36 31 32.0 Sandbar 1 4 14 3 4 146 random Bare Grnd. 3 3 9 14 17 5 27 8 5 10 3 within 2 4 1 1 obtained 1 2 10 12 Common reed typesa 5 6 7 cover 2 3 5 9 4 2 20 6 26 6 8 10 18 2 2 4 7 3 1 1 13 10 2 10 E-1 W-1 W-2 W-11 W-17 W-18 W-19 1 2 10 8 Vines 5 6 5 3 Shrubs Low Tall 11 9 2 9 8 1 and other 10 19 11 16 18 1 seedlings, 11 4 1 29 12 12 23 25 9 16 14 19 E-5 E-3 1 2 Totals 3 3 2 8 1 Grass Annual Peren. 3 3 16 7 2 3 Dune Ridge Forbs Annual Peren. tree 52 3.0 aCover types were classified by the dominant cover present where more than one type was present within a m2 sample. blnc1udes 12 seedlings established in 1983, 5 established in 1984, and 9 established in 1985. CAll were green ash. Peach-leaf willow were not encountered. 80 4.6 14 0.8 20 1,779 1.1 .p.. 0\ W �464 o ---- I \ I I I I I I 4 CHECK STATION MEADOW TAMARACK 11-E-: RED LION DUCK CREEK ._._._._ •••..••...•.•... ", 2 - - I ---' ,, , ,, , \ \ \ , \ ..•.•. \..... ,..... 6 ""'\ ,, ' ,'\ \ \ <, \ ,\ , '\ \ \ \ \ \ \ I \'v ~ - Depth \ \ I I ,,, \ .' \ ".i-: ."' '\\ '" .- X Planting \ / \ . \ \ I , , \ ~. _..... ,=- ~-,,:-\.' ,. t - --_I I •• "_ \ \ ••••• ~.I \ \ " ,. • 12 \ i i ., .,) 14 16 ~ > o z U 1984 0 1985 Fig. 1. Groundwater fluctuations within stern cutting planting sites in 1984 and 1985 in relation to 1985 mean planting depth, South Platte River, Logan County. �465 o . ......•...•..... _ 2 Hale Hale Hale Hale Store - North Store - South Pond East Pond West 4 6 8 .. .' ' .,....._... ". 16 ., -, 18 s:: :J to) -, ...... " ,;'- ..... ..... .' ...... . ........./.." ..' .". ./ / :/. , .", ,~./ .". ./ e 0. Q.I CI1 1984 :J to) 1985 Fig. 2, Groundwater fluctuations within stern cutting planting sites in 1984 and 1985 in relation to 1985 mean planting depth, South Republican River, Yuma County. �.j::'- Table 5. Groundwater Colorado, 1985. Location site levels (dm) below the soil surface along the lower South Platte River, northeastern Mar 5 27 Apr 22-26 7 Ma:t 14-20 Jun 10 Ju1 25 16 10.9 13.2 8.4 13.2 15.2 10.7 6.4 10.7 14.0 10.4 8.1 12.2 14.0 ~ 5-23 Oct SeE 3 23-26 22 12.7 14.5 10.9 7.4 10.7 11.4 6.4 9.7 11.2 5.1 4.8 7.6 12.2 8.6 Tamarack Red Liona 11 Easta Chk. St. M.a Coyote W. 4 West 7 West 9 West 13 West 17 West 24 West Duck Creeka 8.4 8.4 6.1 2.3 8.0 13.3 7.8 6.4 10.2 11.4 9.7 8.6 7.9 9.7 5.3 2.0 1.8 7.1 9.9 9.8 5.1 7.4 5.1 1.8 6.9 7.9 8.4 4.3 4.3 9.4 13.7 9.4 13.7 9.4 Haxtun R&G 5.8 Bravo 4.8 7.1 7.4 7.4 5.3 12.2 9.4 Elliott E. 8.1 3.8 6.9 9.4 9.4 Elliott W. 7.4 3.1 5.1 7.4 8.1 Cottonwood 1 5.8 5.8 5.8 Dodd Bridge 1 7.6 7.4 7.1 -aStern cutting planting sites in 1985. 8.9 '" 0'. �467 Stem Cutting Propagation Trials Survival of stem cuttings in 1985 was disappointing. Survival of 1985 plantings to mid-spring and to mid-June was generally satisfactory at sites along the South Platte and South Republican rivers (Table 6). However, a combination of factors (severe heat stress, declining groundwater levels, competition from adjacent vegetation, and soils) caused rapid desiccation of stem cuttings in early July. Almost no rainfall was received through June and early July. Along the South Republican, some mature cottonwoods "fired" and died or partially died because of a combination of heat stress and water deficiency. Consequently, by September a majority of the stem cuttings were no longer alive (Table 7). Comparison of mean planting depth in relation to summer low water table levels (Figs. 1, 2) show that the plantings had not been planted deep enough. Cuttings should be placed at least as deep as the lowest water level which normally occurs in late summer or early fall (Swenson and Mullins 1985). Their data also show that cuttings placed into a fluctuating water table show much lower survival than those placed into a constant water table. This is understandable because the stems use their energy reserves forming new roots and if the water table drops and the roots are desiccated in dry soil there is no other energy source to sustain the cutting. Cuttings placed at progressively greater distances above a constant water table also sustain dramatically lower survival (Swenson and Mullins 1985). All cuttings were measured and the distance they were placed into the ground and into the water table was also determined at time of planting to discern the relation between planting depth, groundwater level, and survival. However, survival was too low within sites and too much variation and bias occurred among species and among sites to ascertain that a relationship between planting depth and survia1 existed. Stem cuttings in both 1984 and 1985 were placed in vegetation sites with no effort made to reduce competition. Some vegetatiion at the immediate base of the cuttings was removed so that they could be located and not be shaded. The root systems of nearby herbaceous vegetation undoubtedly competed with the cuttings for moisture and probably contributed to the mid-summer high mortality. Rabbit girdling of willows was a problem within the Red Lion and check station meadow sites soon after planting and several mortalities were attributed to that factor. Soil type was also considered important in stem cutting survival and establishment. Heavy clay soils at the Duck Creek and Hale store-south sites possibly did not allow rapid moisture transfer to the cuttings and were difficult to tamp into place around the cuttings at the time of planting. High sodium levels at Duck Creek and at sites along the Arkansas River may have prevented rooting. Soil tests for these sites are planned. Declining water tables were not responsible for poor stem cutting survival within the Arkansas Valley planting sites. Water levels rose subsequent to the late March planting and in several instances water emerged above ground to drown the cuttings. Most were already dead or nearly dead when inspected in late May. Survival to September 1985 was noted for cottonwood, all wi110l>1 species, and the 2 vines. The other species, Russian olive, New Mexico Elderberry (Sambucus neomexicana), cotoneaster (Cotoneaster sp.), and lilac �_.,. Table 6. Survival of stem cutting plantings from early to late spring 1985 within the South Platte, South Republican, and Arkansas river planting sites. "I ~I~ 00 III Drainage site South Platte Chk. St. Mead. 3 East (Tam) 11 East (Tam) Red Lion Duck Creek Sedg. Var Subtotal South Republican Hale St. - So. Hale St. - No. Hale Pond - E Hale Pone - W Subtotal Arkansas Rocky Ford Lusk Timpas Purgatoire ~1 ~I~ 4a 4/10 10/10 0/10 3 21/40 5 5 4 4 18/20 5 2 5 5 '" III 0 '" 0 'M ~I'~ XI'" ...• .., ." Q) s:: en'" 'M '" 'M U .c ~I~ 0.'" '"o ."::l ~.!:: 5 3 5 2 5 5 5/10 5 4 5 5 4 3/3 5/10 2 22/30 17/18 15/15 5 5 5 15/15 4 5 5 4 5 5 19/20 J~ 2 0 5 s:: '" III '" U ::l ." U ::l x Q) .c f: '" Q) 0 en'" s:: 0 2 2 0/4 5 5 4 3 17/20 3 2 1 0 2 1 0 5 4 5 14/15 9/20 7/15 4/19 0 0 2 2 5 0 Q) ." 0 u ••.• 0 •..• (/) '" s:: Q) .., 0 o U . 0. III .~ '" 0.. en III 4 4/20 8/10 4/10 3 0 1 0 'M ::l :> :> '" 'M '" ::l Po< 3 2 9/10 3 1 1 15/25 5 2/5 s:: Q) Q) ::l .c .., s:: 3 2 2 3 5/5 8/15 1/5 3 2 5 0 10/15 1 1/10 2 0 0 0 0 0/5 0/5 0/5 2/10 17/20 9/10 6/20 3/15 Combined surv./planted 56/80 39/40 47/70 37/53 14/15 17/55 15/30 8/34 17/30 5/10 20/40 4/20 Survival rate 0.70 0.97 0.67 0.70 0.93 0.31 0.50 0.24 0.57 0.50 0.50 0.20 Subtotal 4/15 III ." Ill •••• ..,'" aFive stem cuttings per trial were used unless otherwise listed. 0/5 c- Combined Surviv./Planted 26 4 35 29 18 7 45 10 50 55 23 19 119 202 33 28 32 24 50 40 45 30 117 165 15 13 8 5 25 30 25 30 41 110 0.37 277 477 0.58 0.59 0.71 �Table 7. Survival of stem cutting plantings from early spring to fall 1985 within the South Platte, South Republican, and Arkansas River planting sites. II) Drainage Site South Platte Chk. St. Mead. 3-East ll-East Red Lion Duck Creek Sedgwick Bar Subtotal ~~ la 1/10 3/10 0/10 0 ~,i ~I~ '" <I) I-< 0 ....• .c .~ ~2 o ~I~ !;Jot .....•.. ••..• XII-< OJ I-< ••..• ;:l '" ••..• I:: til 1 0 1 1 0 0 1/10 0 0 0 5 2 1/3 5/10 1 5 1 1 9 0 0 0 3 0 1 0 0 1 0 0 0 5 0 0 Subtotal 3 1 1 5 Arkansas Rocky Ford Lusk Timpas Purgatoire 0 0 1 0 0 0 1 0 0 0 0 0 0 Subtotal 1 0 1 0 Combined Surv./P1anted 9/80 2/40 3/70 14/53 13/15 Survival 0.11 0.05 0.04 0.26 0.87 South Republican Hale St. - So. Hale St. - No. Hale Pond - E Hale Pond - W rate 0 0 0 I) x ;:l OJ .a e '" til e 0 OJ I:: 0 0 0 I-< CII ... Jl 'CIII::" ...o Po U <I) 0 I-< 0 0 1 1 2 4 0 0 3 5 5 13 til ~ • Po IJ) 0 0 8 0 0 0 0 0 0 0 0 ~ ~ I-< ••..• '" ;:l Po. 0 Combined Survi v , !Pla_nted n 0 4 1 14 3 1 0 45 10 50 55 23 19 0 0 23 202 0 1 4 0 1 12 12 12 50 40 45 30 5 1 37 165 0 0 1 0 1 0 25 30 25 30 2 110 0.02 62 477 0.13 0 0 0 0 ~I~ ... " 0 0 0/4 0 '1 ••..• 0 1)'<-< 0 •...• I:: CII ;:l <I) 0/10 0 0 7 1 0 0 0 0 0 0 0 0 15/55 0.30 0/34 0/30 0/10 5/40 1/20 0.27 0.00 0.00 0.00 0.00 0.12 0.05 0 aFive stem cuttings per trial were used unless otherwise ~J '" '" ;:l ••..• I:: II) I) II) ••..• <1)..-1 0.11 0.22 listed. ~ 0'> 1.0 �470 (Syringa sp.), showed limited survival to mid-spring (Table 6), however, none survived into fall. Some of the cuttings of indigo bush (Amorpha fruiticosa) and coyote willow (Salix exigua) had apparently winter killed or were in poor vigor when planted and never leafed out. Most cuttings of indigo bush were slow to leaf out but, among the native cuttings, this species survived at the highest rate to fall (Table 7). The introduced golden willow (Salix "golden"), that has been planted on the South Republican Wildlife Area, showed excellent survival on the 3 sites where planted. Three of the 5 grape vine (Vitis vulpina) cuttings that survived to fall contained a few roots (because they were rooted into the topsoil when laying prostrate) but there was no clear evidence that the roots increased survival. Others with roots at the same site did not survive. The reason that grape vines on the 2 South Republican sites survived is uncertain but heavy mid-summer rains were believed to be a factor in sustaining them. Survival of coyote willow was poor (Table 7). This tall, bushy willow has excellent growth form for wildlife plantings and would be a major species used if successful stem cutting techniques can be developed. Both the common and scientific names for this plant are uncertain and need to be verified at the CSU Botany Department. Considerable confusion exists among many references in riparian literature concerning sandbar willow (S. interior) and coyote willow (~. exigua), and clarification is needed. The species identified as coyote willow does not seem to fit published species descriptions. At the Red Lion site along the South Platte, low river level and lack of equipment prevented placing the 10 cottonwood cuttings deep enough into the ground water table. Although the groundwater table came up later in spring, none of the cuttings showed signs of survival or leafing through spring monitoring. Soil Scarification Natural Reproduction Eighteen randomly selected sites within 6 locations were moldboard plowed in October 1984 along the east Tamarack bottomlands. These sites were rototilled in early spring 1985 to break up large clods and provide a smooth surface. However, hard rains needed to compact the soil were not received in spring so the soil remained more loose than preferred for good moisture retention. Three additional random plots were selected along the west side of Colorado 55 where soil had been disturbed during a late summer 1984 pipline installation. The unusually warm spring prompted early cottonwood seed dissemination beginning in mid- to late May and continuing until mid-June. The South Platte River remained at relatively stable levels, about one-half full during this interval which was not conducive to seedling germination either within the tilled sites or other higher locations along the river. Irrigation demands reduced stream flow to low levels begdrmf.ng about 18-20 June and the river remained nearly dry for a month. All treatment sites were inspected on 27 June 1985. Cottonwood seedlings were found on all 3 of the pipeline plots where the soil was firm and elevations were low in comparison to river water level allowing moisture to saturate the soil surface. Seedlings were present only in the lower southeast corner of �471 the south plot and the lower northwest corner of the north plot. Seedlings were common throughout the more level and wet middle plot. Relatively low stream flows prevented any seedling establishment on other plots to the east. All remained dry on the surface except in dead furrows along the edge of several plots. In addition, almost no rain was received during the interval of cotton dissemination that would have helped keep soil surfaces moist. Seedlings had been established on or near most of these plots in 1983 and/or 1984 without soil tillage treatment. Most or all seedlings within the pipeline plots died of desiccation in mid to late summer as river levels declined. Most of the plots were subsequently treated with shallow tillage in late summer to remove established vegetation. Observations along the South Platte and South Republican rivers indicate that soil disturbance enhances opportunities for seedling establishment. For example, a dike upgraded along the meadow of the east Tamarack in the early 1970's is almost continuously lined with cottonwoods that are now pole-sized or larger. Similar invasions of seedlings have been noted in disturbed sites below Bonny Reservoir. Thus, soil scarification is apparently an aid to seedling establishment assuming groundwater table levels are high enough and/or rainfall is adequate. F. Knopf (pers. commun.) stated that sprinkler irrigation could be used to keep a moist soil condition conducive to seedling establishment. It would also be essential to sustain irrigation for at least an additional year or 2 to allow seedling root systems to develop adequately. Monitoring of Controlled Riverbottom Burns Management personnel began limited burning of small tracts within open to partially timbered riverbottoms on the Brush Wildlife Area along the South Platte River in Morgan County during early spring 1985. Post-burn inventories of tree and shrub survival and condition were conducted on 20 May and 4 September 1985 (Table 8). A back fire was used along the north side of the tract and a head fire was used on the remainder. The latter burned rather intensely as considerable residual herbaceous vegetation was present. A majority of the mature to over-mature trees sustained little damage but several received moderate to severe damage (Table 8) within the head-fired portion of the burn. The backfire did not appear to severely injure either cottonwoods or peach-leaf willows, however all young trees were moderately to severely impacted by the headfires. One of the primary problems with burning among cottonwoods was that the low hanging branches and fallen branches and trees, which are important for protecting and screening wildlife, especially wintering game species, were lost or killed. Cottonwoods (Table 8) showed no sign of basal sprouting. Young seedlings that had germinated in 1983 were burned off but many, especially the smaller ones resprouted from the base. Although most of the peach-leaf willows were moderately to severely impacted by the headfire, nearly all showed vigorous basal sprouting and regrowth up to 2 m in length. Sandbar willow seemed to be more severely impacted by the fire. Regrowth to late summer averaged about 0.67 m but was not considered vigorous. All top growth of indiobush was killed but regrowth was excellent (1.5 m) indicating this species can sustain the impact for fire exceptionally well. �472 Table 8. Status of cottonwood and peach-leaf willow trees following a controlled burn on the Brush Wildlife Area, Morgan County, Colorado, 1985. Fire impact Young Tree age Mature Little (limited basal scorching; branches above 2-m not killed) Moderate (considerable basal scorching, branches killed to above 2-m) Severe, (killed or most upper branches killed) Totals 2 7 (1) (4) 2 (5) Overmature 2 (2) 18 2 11 (1) (2) 5 (2) 28 (3) Total (3) 5 (6) 4 (3) 34 (11) aCottonwoods bpeach-1eaf willow Three of 4 Russian olives were young «10 em dbh) and were entirely scorched by the fire. One older tree was about 50% killed. Resprouting occurred on all trees from center and basal areas but regrowth was not vigorous. One tamarisk, which undoubtedly was partially dead prior to the fire, showed marginal basal sprouting. Following the 1985 burn on the Brush Wildlife Area, herbaceous vegetation regrowth was more vigorous within the west portion of the tract than in the east part. Possibly the groundwater table was closer to the surface but this is uncertain. Tall, dense stands of sweet clover (Me1ilotus spp.), prairie cordgrass (Spartina pectinata), switchgrass (Panicum virgatum), licorice (Glycyrrhiza 1epidota), and open stands of sunflowers (He1ianthus sp.) and snow-on-the-mountain (Euphorbia marginata) dominated within the west portion of the burn. Snowberry (Symphoricarpos sp.) also made excellent regrowth following the fire. Within the drier locations to the east, open to moderately dense stands of perennial grasses dominated with little if any increase in annual or perennial forbs. The fireguards, which had been fall plowed and disced in spring, provided highly variable stands of annual forbs, primarily sunflowers and cockleburs (Xanthium strumarium). Many of the perennial grasses including wheatgrasses (Agropyron spp.) and sa1tgrass (Distich1is stricta) had not been killed by tillage. If tillage is not continued, the fireguards will quickly become dominated by these and other perennial grasses and by cheatgrass brome (Bromus tectorum). Fireguards were laid out and plowed for 2 proposed burns to be conducted in spring 1986 within the Brush Wildlife Area. The sites were primarily open and dominated by dense herbaceous vegetation. A pretreatment fall 1985 inventory showed 10 cottonwoods, 7 peach-leaf willow, 1 Russian olive, 2 indigobush, �473 1 clump of sandbar willow, 1 Chinese elm (Ulmus parvifo1ia), and 2 Virginia creepers (Parthenocissus quinquefo1ia) on the east site. The west site contained 1 each of peach-leaf willow, Russian olive, American elm (Ulmus americana), and boxe1der (Acer nequndo). General age, height, height of lower live branches, and other data were recorded for each tree or shrub. An intensive 25-m long transect was established in September 1985 within which 226 cottonwood seedlings were encountered. These ranged in height between 0.67 and 1.21 m and had germinated in 1983. Based on monitoring of the 1985 burn, assistance was provided to Northeast Region management personnel in the form of general guidelines that should be following in burning within the South Platte riverbottom. LITERATURE REVIEW Snyder, W. D. 1984. Lowland riparian cottonwood community studies. Job Progress Rep., Colorado Div. Wildl., Wildl. Res. Rep., Fed. Aid Proj. N-4-R-2. Jan:292-370. 1985. Dynamics of Cottonwood Regeneration. Job Progress Rep., Colorado Div. Wi1d1., Wi1d1. Res. Rep., Fed. Aid Proj. 01-00-045 (W-37-R). Apr. Swenson, E. A., and C. L. Mullins. 1985. Revegetating Southwestern floodplains. Pp. 135-138 in R. R. Coord.) Riparian Ecosystems and their management: uses. First North American Riparian Converence. Servo Gen. Tech. Rep. RM-120. 523 pp. Prepared by: ?j{)tt»u:J ~ Warren D. Snyder Wildlife Researcher C riparian trees in Johnson et al. (Tech. reconciling conflicting U.S. Dep. Agric. For. ��475 Colorado Division of Wildlife Wildlife Research Report April 1986 JOB PROGRESS REPORT State of Colorado ---------------------------------- Project Work Plan Job Title: 01-00-045 (W-37-R) 21: Job Avian Research 3 Sandsage-Bluestem Prairie Renovation Period Covered: 1 January through 31 December 1984 Author: Warren D. Snyder Personnel: D. C. Bowden, C. E. Braun, W. Brown, L. Budde, T. Davis, M. Etl, M. Gardner, D. Lukens, W. Miles, M. Stanley, S. Steinert, other Division of Wildlife personnel, and W. D. Snyder, Colorado Division of Wildlife. ABSTRACT Vegetation renovation, using controlled burning, was initiated by Northeast Regional personnel on approximately 112 ha (3 sites) of the South Tamarack Prairie to enhance sandsage-bluestem (Artemisia-Andropogon) prairie for prairie grouse. Pre-treatment spring 1984 residual grass-forb vegetation yielded height-density indices (HDls) approximating 0.25 dm and sand sagebrush (A. filifolia) HDls ranging between 0.75 and 1.00 dm. Blue grama (Bouteloua gracilis), needle-and-thread (Stipa comata), prairie sandreed (Calamovilfa longifolia), sand dropseed (Sporobolus cryptandrus), and sand sagebrush were the dominant species present within the burn sites. Among these species crown cover of needle-and-thread and sandsage was reduced by fire but the fire did not kill sandsage which resprouted quickly and made rapid regrowth. Pre-treatment crown cover measurements were biased for some species because of difficulties in accurately aging residual vegetation in early spring. Perennial forbs increased, apparently due to fire, on Burn 1 but under different conditions in Burn 3 showed little increase. Annual forbs had higher densities within controls than in the burns. Lambsquarter (Chenopodium album) was the most abundant forb within controls. Sites and controls for proposed spring 1985 burns were selected, mapped, and pre-treatment vegetation sampling was conducted. Test plots evaluating interseeding, tillage plus seeding, and tillage-herbicide plus seeding all attained satisfactory grass stands. Preliminary comparisons revealed that use of atrazine herbicide enhanced first-year stand establishment and growth of certain warm-season grasses over other treatments. ��477 SANDSAGE-BLUESTEM PRAIRIE RENOVATION Warren D. Snyder The South Tamarack Prairie within the South Platte Wildlife Area contains approximately 1,820 ha of grassland that has been designated for establishment of greater prairie-chickens (Tympanuchus cupido). Livestock grazing. was discontinued on the site following 1977 and several hundred hectares were interseeded in 1981-82 with varying success. Based on recommendations of a consultant, L. Kirsch, controlled burning was implemented on three tracts totaling approximately 112 ha in spring 1984. Personnel of the nongame research group and the Northeast Region jointly began trapping and transplanting greater prairie-chickens in spring 1984 and released 36 birds onto the South Tamarack Prairie. Monitoring provided evidence of at least partial establishment success and additional releases are planned for 1985. This study was initiated to evaluate controlled burning, and other renovation and revegetation approaches for increasing vegetation quality of the South Tamarack sandsage-bluestem prairie. Results should show which approa~hes are most economical and effective for increasing height-density quality and enhancing composition of native warm season grasses. This report summarizes efforts and findings of the initial year. P.N. OBJECTIVES Test renovation and revegetation techniques for increasing standing residual height-density of grasses, increasing the proportion of tall, warm-season grasses within the composition, and for reducing the quantity of sand sagebrush to < 30% canopy cover in an ungrazed sandsage-bluestem prairie on the South Tamarack, South Platte Wildlife Area in northeastern Colorado. SEGMENT OBJECTIVES concerning prairie renovation and revegetation 1. Review literature techniques. 2. Select treatment sites and controls in coordination with management personnel of the Northeast Region. Controlled burning renovation and revegetation treatments will be enacted as outlined in the Program Narrative. 3. Environmental monitoring will include: a. Precipitation will be monitored throughout the year. b. Soil, plant phenology, and weather conditions will be monitored at time of controlled burning in May. �478 4. c. Visual obstruction measurements will be obtained on treatments and controls where applicable in late March and early ApriL Optional limited sampling may be conducted prior to snowfall in fall. d. Crown cover, species composition, and frequency of occurrence measurements of vegetation will be obtained in late August and early September. e. Photos of treatments will be taken in October. Data compilation and writing the annual conducted from November through April. job progress report will be METHODS Following coordination with management personnel and design coordination with D. C. Bowden, two burn (treatment) sites, Burn 1 and 3, were selected for vegetation sampling and evaluation. The sites were mapped, gridded, and random vegetation sampling sites were located using a measuring wheel and compass. Fire guards were placed within the burn sites to exclude controls. Burn 1 contained nine treatment and nine control vegetation sampling locations and Burn 3 contained four treatment and four control sampling locations. The visual obstruction pole procedure (Robel et ale 1970) as modified by Kirsch (1977) was used to measure the height-density quality of residual vegetation. Obstructions by sand sagebrush were delineated and listed separately to obtain three indices: (1) grass-forb, (2) sand sagebrush, and (3) the combined totaL Readings were obtained in March and April prior to green-up with 25 stations (100 measurements) spaced at five pace or greater intervals around the outside of the vegetation composition transects. These efforts will be duplicated in subsequent spring intervals. Crown cover, species composition, and frequency of vegetation were measured using the metric belt transect system described by Schmutz et ale (1982) in pre-treatment (spring) and post-treatment (late summer) intervals in 1984. A 50-m line placed between two steel posts was divided at 0, 12.5, 25.0, 37.5, and 50.0-m intervals and five 7.9-m transect lines were extended 900 to the left, right, left, etc. at the five points along the line. A 0.1-m2 frame (31.6 cm/side) divided in half and one-half further divided into five 0.01-m2 segments was used for sampling. The frame was positioned 25 times along the 7.9-m transect line yielding a 2.5-m2 sample per line or 12.5 m2 per sample point. A point frame vegetation composition sampling procedure (Floyd and Anderson 1983) was used to sample pre-treatment (fall 1984) vegetation within proposed spring 1985 burn sites and their controls. Thirty-six points were sampled within O.5-m2 and replicated four times at 2-m intervals along three randomly placed lines at each random point. Twelve random points were established in each of the three treatment and three control sites yielding 5,184 tallies per site. Photo hubs were established prior to spring 1984 burning at four controls and eight treatment sites within Burn 1 and at two controls and four treatment sites within Burn 3. Photos were taken again in October 1984. �479 Monitoring of environmental conditions at the time of controlled burning included (1) soil temperature, (2) soil moisture at 5, 15, and 30-cm depths, (3) plant phenology of species present within the range site, (4) wind speed and direction, (5) relative humidity, and (6) air temperature and cloud cover. Precipitation gauges were placed within or near Burns 1 and 3 but could not be read immediately following each rainfall and were thus of limited value. Some rainfall information was obtained from M. Gardner at the Tamarack Ranch Headquarters and information was obtained from H. Hamilton (Crook) and the U.S. Weather Service recording station (south of Sedgwick). Two replications of five linear 0.5-ha revegetation plots were established in early spring 1984 and randomly selected treatments were applied as follows: (1) rototillage and seeding a tall warm-season grass mixture, (2) repeat as in 1 except to add tall wheatgrass (Agropyron elongatum) and alfalfa to the mixture, (3) repeat as in 1 except to apply 0.6 kg/ha (a.i.) of atrazine herbicide between tillage and planting, (4) use the interseeder previously used on the Tamarack to plant a tall warm-season grass mixture, and (5) repeat as in 4 with addition of tall wheatgrass and alfalfa. Although the seed was old and potentially of reduced germination (com~ared to sack tag information), pure Lfve seed application rates/O.09m were: switchgrass (Panicum virgatum) (11.0), yellow Indiangrass (Sorghastrum nutaus) (4.4), big bluestem (Andropogon gerardi) (3.8), sand bluestem (A. hallii) (2.6), little bluestem scoparius) (2.2), tall wheatgrass (5.4), and Ranger alfalfa (10.0). Initial tillage was completed in early April, atrazine was applied on 18 April, and seeding and interseeding was completed on 24-25 April. Since a drill that would handle the light fluffy seeds of the bluestems and Indiangrass was not available, these were hand-broadcast. The plots were then shallow tilled with a spike-toothed harrow and the other grasses and aLfaLfa were drilled. Tillage was not conducted prior to using the interseeder which handled all grass seeds. The seeding rate for interseeding could not be accurately calibrated but was nearly equal to that for the other plots. (A. Because of limited seedling establishment and growth in 1984, and considerable consumption of vegetation by grasshoppers, vegetation sampling was restricted to comparisons between warm-season grass plots untreated and treated with atrazine herbicide in fall 1984. Ten random 7-m transects were established in each of the two plots/treatment and vegetation was tallied within 0.5-m2 frames at 1, 4, and 7-m segments of the transect line yielding a 1.5-m sample/transect or 15 m/plot. Grass seedlings, annual forbs, and other vegetation were recorded as present or absent within 50 0.01-m2 grids within each 0.5-m frame yielding 150 potential hits/transect. The point frame sampling procedure will be used in subsequent years after the vegetation has attained additional growth. STUDY AREA The South Tamarack Prairie (Fig. 1) is that rangeland portion of the South Platte Wildlife Area (Tamarack Ranch) south of Interstate Highway 1-76. It contains approximately l,820 ha bounded on the east by Logan County Road 93, on the west by Colorado Highway 25, and by private and state school lands in the sandsage-bluestem prairie on the south. The elevation approximates 1,160 m. Soils are of the Daily and Valent loamy sands (Amen et ale 1977). Precipitation averages approximately 40 cm/year. �~ CXl o Fig. 1. Location of ~984 burns, proposed 1985 burns and their controls, and 1984 revegetation test plots within the South Tamarack. �481 RESULTS AND DISCUSSION Based on information from the literature and contacts, evaluation of controlled burning has not been conducted in eastern Colorado either on sandsage-mixed prairie (found in sandy soils) or in shortgrass prairie. Findings of Bragg (1978) conducted in the choppy sandhills on the Crescent Lake National Wildlife Refuge in west-central Nebraska (approximately 130 km north of the South Tamarack) probably most closely apply to conditions on the South Tamarack; however, major differences occur in soils and vegetation, and to a lesser extent in climate. Other than general statements that sand sagebrush rootsprouts readily after fire, little information could be found concerning the impacts of controlled burning on sandsage. Thus, findings of this study should fill voids concerning habitat modification efforts in the semi-arid sandsage-bluestem (mixed) prairies of eastern Colorado. Implementation of Controlled Burns - Spring 1984 Management personnel of the Northeast Region, primarily Area 3, with assistance from Wildlife Research personnel, laid out, mowed, and tilled fire breaks around three burn sites (Fig. 1) approximating 50 ha (Burn 1), 42 ha (Burn 2), and 20 ha (Burn 3). Because of time and manpower limitations and because extensive grass seeding was planned in Burn 2, vegetation evaluations were restricted to Burns 1 and 3. Controlled burning was initiated on 4 May 1984 with M. Gardner serving as fire boss. Weather at the time of the burns was clear with a northwest wind of 8-12 mph which gradually diminished to 4-6 mph by noon. Burn 1 was treated from 0800 to 1000 hours with a temperature range of 40-60 F and a relative humidity range from 60 to 32%. Burn 3 was conducted from 1130 to 1230 hours, with a temperature range of 62-65 F and relative humidity at 26-28%. Burn 3 was considerably hotter than Burn 1 because of the higher temperature and lower relative humidity. Soil moisture was considered adequate and, based on 5 replications at 5.1, 10.2, and 20.3 cm respectively, averaged 47,51 and 53 %. These readings should be interpreted with caution because the moisture meter was not considered a highly reliable instrument. Average monthly temperatures were near normal in March 1984 but ranged between 5.5-6.0 F below normal for April at nearby weather stations. May temperatures averaged near normal. The colder than usual April temperatures retarded growth of vegetation so that it was phenologically behind schedule at the time the burns were conducted in early May. Phenological conditions are summarized in Table 1. �482 Table 1. Phenological characteristics or status of vegetation within South Tamarack Prairie at time of controlled burning, 4 May 1984. Species Warm-season grasses Needle-and-thread Sandhill muhly Quackgrass Small Carex sp. Allium (wild onion) Cymopterus (wafer parsnip) Lathyrus (Peavine) Beardtongue and spiderwort Sandsage the Status No green-up of any species noted New growth up to 15.2 cm (6 in.) Beginning new growth Several inches of new growth Some flower heads starting to bloom .Most with flower heads not yet bloomed Many have had flower heads for 2-3 weeks 5-10 cm tall and a few beginning to bud 5-7.5 em tall Partial leafing but mostly basal. Highly variable among plants. New growth of grasses and forbs was observed within 5 days following the burns and major green-up had occurred within 10 days. By 23 May « 3 weeks after the burns) sandsage had sprouted profusely and was up to 10 cm tall and prairie sandreed was up to 25 cm. Showy peavine (Lathyrus spp.) had made extensive regrowth and it, beardtongue (Penstomen spp.), and spiderwort (Tradescantia occidentalis) were near bloom stage. Initial growth following the burns was rapid and then slowed during June and July. Although the preceding fall (1983) had been dry, a major snow in late November 1983 and subsequent above-average precipitation through April 1984 enhanced soil moisture conditions (Table 2). However, precipitation was below average following the burns from May through September (Table 2). Precipitation gauges were placed at Burns I and 3 but they could not be read after every rainfall so on-site data were missing. Limited readings and observations showed that Burn 3 rather consistently received less rainfall than was received on Burn 1. This was readily observed in vegetation growth throughout the summer. �483 Table 2. Monthly and annual precipitation (inches) at Crook, 5 miles south of Sedgwick, and at Sterling, Colorado in 1984. Crooka Month 0.47 1.12 0.66 3.42 2.01 2.80 0.49 0.66 0.44 1.39 0.01 0.80 Jan Feb Mar Apr May Jun Ju1 Aug Sep Oct Nov Dec Annual 14.27 Sedgwick Sterling 0.39 1.58 0.67 3.63 1.22 1.31 1.50 0.47 0.65 2.12 O.Olc 0.65c 0.21 0.08 3.52 1.66 1.99 1.87 1.00 1.39 0.44 2.27 O.Olc 0.50c 14.20 14.94 Sedgwick long term xb 0.30 0.35 0.77 2.16 2.57 2.73 2.51 2.26 1.46 0.99 0.42 0.50 17.02 aData from H. Hamilton. bLong-term weather data from U.S. Weather Service Climatological Data. cApproximate: weather data not yet available. Vegetation Sampling Height-Density Measurements.--Pre-treatment height density sampling, conducted in early spring 1984, is summarized in Tables 3 and 4 for Burns 1 and 3 and their respective controls. Comparison of the two tables reveals that sandsage was the primary vegetation obstruction in about 45% of the Burn 3 readings compared to less than 20% in Burn 1. Grass-forb vegetation HDIs were slightly higher in Burn 1 than in Burn 3 but both were low. Kirsch et a1. (1978) showed increasing duck nesting use of residual vegetation as HDIs increased from 0.5 to 2.5 dm or greater. In personal communications he noted the same trend was true for prairie grouse. Snyder (1984) and others have documented similar findings for pheasants. HDIs < 0.5 provide poor concealment and marginal nesting cover for prairie grouse, ducks, and pheasants. Sandsage consistently provided higher cover readings than those of grass-forb vegetation, however, the higher sandsage density in Burn 3 may have been responsible for decreased grass-forb HDIs there because of competition. A heavy snow in late November 1983 that persisted through much of the winter was an important factor causing extensive lodging and matting of residual cover. In contrast to the low HDI readings obtained in spring 1984, a small sample (N = 46) was obtained on 10 November 1983 yielding a grass-forb HDI of 1.43 dm.- The location was near Burn 2 (Fig. 1) and would have been comparable with vegetation on Burn 1. The height-density indices will be more meaningful when compared with post-burn readings obtained in early spring 1985. This procedure should yield the best index showing the value of controlled burning for renovation of sandsage-mixed grass prairie in eastern Colorado. �484 Table 3. Height-density (dm) of residual grass-forb and sandsage vegetation within treatment and control locations in Burn 1, South Tamarack Prairie, 1984. Grass-forb Transect N Sandsage x- .x li TREATMENT 13-6 14-6 14.6-2.8 15-2 16-1 16-5 17-4 19-1 21-2 48 53 71 55 68 62 63 56 54 530 0.232 95% CI < 0.253 1.134 0.750 1.050 0.944 0.437 0.982 1.111 0.588 0.704 28 20 5 9 4 14 9 17 22 128 0.375 0.321 0.306 0.259 0.202 0.185 0.242 0.192 0.222 < 0.274 0.341 Combined 0.769 < 0.373 < < 0.871 < 0.973 < 0.855 0.405 CONTROL 15-4 14-5 21-4 12-5 15-5 17-7 17-1 16-2 13-4 95% CI Combined 83 87 59 74 66 79 80 97 98 723 0.234 < 0.252 0.426 0.423 0.528 0.875 0.787 1.190 0.600 1.500 1.000 17 13 9 26 34 21 20 3 2 145 0.214 0.250 0.225 0.318 0.299 0.339 0.141 0.201 0.291 < 0.269 0.312 < 0.337 0.669 < 0.372 < 0.762 �485 Table 4. Height-density (dm) of residual grass-forb and sandsage vegetation within treatment and control locations in Burn 3, South Tamarack Prairie, 1984. Grass-forb Transect N Sandsage x N TREATMENT 5-1 5-3 5-4 3.5-6 52 53 52 64 221 48 47 48 36 179 0.226 0.250 0.293 0.137 0.183 < 0.222 < 0.261 95% CI 0.781 0.729 0.995 0.792 0.723 <0.827 <0.931 0.433 < 0.492 < 0.552 Combined CONTROL 4-2 4-4 6.5-1 8-2 95% CI Combined 57 54 57 47 215 1.163 0.989 0.878 0.750 43 46 43 53 185 0.180 0.148 0.241 0.154 0.154 < 0.183 < 0.211 0.840 < 0.935 < 1.030 0.471 < 0.531 < 0.590 Vegetation Crown Cover, Composition, and Occurrence.--Common and scientific names of species tallied or observed on the South Tamarack Prairie are presented in Table 5. This is not a complete list of species p'resentas a few legumes still remain to be identified. Crown cover(O.Ol m2), species composition (%), and frequency of occurrence data for pre- and post-treatment intervals for Burns 1 and 3 and their controls are presented in Tables 6 through 9. The pre-treatment samples, conducted prior to major green-up in early spring 1984, contain several weaknesses due to time of sampling. Overwinter lodging and snow compaction of residual vegetation increased sampling difficulty. Species identification was more difficult and the residual (dead vegetation) from 1983 growth could not be readily distinguished from that of previous years for some species. Subsequent sampling on other sites in fall 1984, using a point-frame procedure, provided evidence that the amount of year-old dead vegetation was markedly underestimated in spring 1984. Annual and perennial forbs were not represented by sampling in relation to their importance and several species were not detected because they had become fragmented or had been defoliated by insects. Identification was difficult for some species. Sampling of major grasses and sandsage was considered most reliable. �486 Table 5. 1984. Grasses, forbs and other vegetation on the South Tamarack Prairie in Scientific name Common name Status Grasses and sedges Agropyron cristatum A. elongatum A. repens A. smithii Andropogon gerardi A. halli! A. scoparius Aristida oligantha Bouteloua gracilis Bromus spp , Calamovilfa longifolia Cenchrus sp. Eragrostis trichodes Echinochloa crusgalli Koleria cristata Muhlenbergia pungens Panicum virgatum Paspalum stramineum Sorghastrum nutans Sporobolus cryptandrus Stipa comata Crested wheatgrass Tall wheatgrass Quackgrass Western wheatgrass Big bluestem Sand bluestem Little bluestem Prairie threeawn Blue grama Cheatgrass brome Prairie sandreed Sandbur Sand lovegrass Barnyardgrass Prairie Junegrass Sandhill muhly Switchgrass Sand paspalum Yellow 1ndiangrass Sand dropseed Needle-and-thread Seeded along 1-76 Seeded in tests Occasional Common, native 1nterseeded Common, native 1nterseeded Occasional Abundant Occasional Abundant Uncommon 1nterseeded Uncommon Common Common Occasional-seeded Common 1nterseeded Abundant Abundant Sedges Sandhill sedge Abundant Abundant Sand verbena Wild onion Western ragweed Prickly poppy Greed sagewort Sand sagebrush (sandsage) Cudweed sage Milkweed Wooly loco Pigweed Lambsquarter Chicory Rocky Mountain beeplant Thistle Canada horseweed Texas croton Cryptantha Wafer parsnip Plains larkspur Hedgehog cactus Annual buckwheat Milk purslane Evolvulus Occasional Abundant Abundant Uncommon Uncommon Abundant Common Uncommon Occasional Occasional Abundant Occasional Uncommon Occasional Common Abundant Common Common Common Common Common Abundant Abundant Carex spp. Cyperus schwainitii Forbs Abronia fragrans Allium textile Ambrosia psilostachya Argemone intermedia Artemisia caudata A. filifolia A. ludoviciana Asclepias sp. Astragalus mollissimus Amaranthus spp. Chenopodium album Cichorium intybus Cleome serrulata Cirsium spp. Conyza canadensis Croton texensis Cryptantha spp. Cymopteris montanus Delphinium virescens Echinocereus viridiflorus Eriogonum annum Euphorbia spp. Evolvulus nuttalianus �487 Table 5. (cont.) Scientific name Froelichia gracilis Gaura coccinea Gutierrezia sarothrae Grindelia squarrosa Haplopappus spinulosus Helianthus sp. Hoffmannseggia jamesii Ipomoea leptophylla Kochia scoparia Lactuca sp. Lathyrus polymorphus Lesquerella ludoviciana Leucocrinum mont anum Liatris punctata Lygodesmia juncea Mammillaria vivipara Mentzelia nuda Oenothera ~ Opuntia humifusa Penstemon angustifolius Pepidium densiflorum Phlox andicola Physalis subglabrata Plantago purshii Psoralea lanceolata P. tenuiflora Psoralea spp. Ratibida columnifera Rumex venosus Salsola kali Sphaeralcea coccinea Thelesperma megapotimicum Tradescantia occidentalis Tragopogan spp. Yucca glauca Common name Status Slender froelichia Scarlet gaura Snakeweed Curlycut gumweed Ironplant goldenweed Sunflower Rushpea Bush morningglory Kochia Wild lettuce Showy peavine Bladderpod Sandlily Dotted gayflower Skeleton weed Ball cactus Evening starflower Primrose Pricklypear cactus Blue beard tongue Peppergrass Prairie phlox Ground cherry Wolly Indian wheat Lemon scurf pea Wild alfalfa Scurf pea Prairie coneflower Winged dock Russian thistle Scarlet glebemallow Greenthread Spiderwort Salsify Yucca Uncommon Occasional Uncommon Uncommon Common Common Uncommon Common Uncommon Uncommon Abundant Occasional Uncommon Uncommon Common Common Common Common Abundant Common Occasional Common Occasional Occasional Uncommon Common Occasional Uncommon Uncommon Common Occasional Occasional Abundant Uncommon Uncommon �488 Table 6. Crown cover (0.01 m2), species composition (%), and frequency of occurrence of vegetation during pre- and post-burn intervals on Burn 1, South Tamarack Prairie, 1984. Pre-burn Vegetation Crown cover Compo Post-burn Freq./ occ. Crown cover Compo Freq./ occ. 2,792.0 778.5 679.0 1,794.0 429.0 6.0 2.0 7.0 0.5 5.0 36.24 10.11 8.81 23.29 5.57 0.08 0.03 0.09 0.01 0.06 0.978 0.978 0.978 0.978 0.578 0.067 0.067 0.022 0.022 0.044 63.0 0.82 0.333 3,479.5 67.5 Bare ground Dead vegetation 1,055.5 1,123.0 Bouteloua gracilis Stipa comata Sporobolus spp. Calamovilfa longifolia Andropogon hallii Agropyron smithii Aristida sp. Panicum virgatum Muhlenbergia sp. Koleria cristata 3,262.0 2,324.0 404.0 1,671.0 258.0 36.96 25.62 4.45 18.42 2.84 0.978 1.000 0.955 0.867 0.489 4.0 0.04 0.022 11.5 0.13 0.089 Cyperus & Carex spp. 926.5 10.21 0.889 682.0 8.85 0.867 Opuntia sp. Mammillaria & Echinocereus 33.0 2.5 0.36 0.03 0.355 0.044 21.5 5.0 0.28 0.06 0.311 0.111 Ambrosia psilostachya Tradescantia occidentalis Phlox andicola Evolvulus nuttalianus Lathyrus polymorphus Penstemon angustifolius Psoralea tenuiflora Allium textile Delphinium sp. Thelesperma sp. Haplopappus spinulosus Mentzelia nuda Froelichia-gr.;cilis Lygodesmia juncea Oenothera sp. Ratibida columnifera Physalis subglabrata 69.0 21.5 11.0 19.5 0.76 0.24 0.12 0.21 0.311 0.333 0.067 0.222 5.5 3.0 0.5 0.06 0.03 0.01 0.111 0.044 0.022 11.0 0.12 0.044 75.5 163.5 17.0 24.5 8.5 8.0 8.0 0.5 1.0 44.0 2.0 0.98 2.12 0.22 0.32 0.11 0.10 0.10 0.01 0.01 0.57 0.02 0.333 0.822 0.178 0.200 0.089 0.089 0.089 0.022 0.022 0.289 0.067 6.0 0.07 0.089 3.0 0.5 5.0 5.0 1.0 0.04 0.01 0.06 0.06 0.01 0.022 0.022 0.022 0.022 0.022 1.5 34.0 33.0 1.5 1.0 0.02 0.44 0.43 0.02 0.01 0.067 0.467 0.600 0.022 0.044 0.5 0.01 0.022 Artemisia filifolia Chenopodium album Euphorbia sp. Croton texensis Plantago purshii Helianthus sp. Eriognum annum Cruciferae spp. Unidentified 0.5 0.01 0.022 2.0 7.0 0.5 18.0 0.02 0.08 0.01 0.20 0.044 0.111 0.022 0.178 �489 Table 7. Crown cover (0.01 m2), species composition (%), and frequency of occurrence of vegetation during pre- and post-burn intervals on Burn 1 controls, Tamarack Prairie, 1984. Post-burn Pre-burn Vegetation Bare ground Dead vegetation Crown cover Compo Freq.j occ. Crown cover Compo Freq.j occ. 822.0 2,105.0 940.0 826.0 4,592.5 1,683.0 309.5 1,753.0 196.5 3.0 61.5 2.0 48.42 17.74 3.26 18.48 2.07 0.03 0.65 0.02 1.000 1.000 1.000 0.844 0.422 0.022 0.222 0.022 2,926.0 1,648.5 332.5 1,723.5 263.0 9.0 26.5 7.0 35.15 19.81 3.99 20.71 3.16 0.11 0.32 0.08 1.000 1.000 0.955 1.000 0.555 0.044 0.155 0.044 5.0 0.05 0.044 32.5 0.39 0.311 572.0 6.03 0.800 605.0 7.27 0.822 Opuntia sp. Mammillaria & Echinocereus 39.5 3.5 0.42 0.04 0.333 0.067 32.0 3.5 0.38 0.04 0.267 0.111 Ambrosia psilostachya Artemisia ludoviciana Tradescantia occidentalis Phlox andicola Evolvulus nuttalianus Lathyrus polymorphus Penstemon angustifo1ius Allium textile Psora1ea tenuif10ra Lygodesmia juncea The1esperma sp. Froelichia gracilis Mentze1ia nuda Cymopteris montanus Cichorium intybus 102.5 1.08 0.222 53.5 15.5 11.0 7.5 6.0 17.5 0.56 0.16 0.16 0.08 0.06 0.18 0.667 0.222 0.111 0.067 0.044 0.378 6.0 0.06 0.044 15.0 7.5 0.16 0.08 0.044 0.222 64.0 14.0 84.5 21.0 12.5 2.5 8.0 0.5 11.5 1.0 23.5 1.0 4.0 0.77 0.17 1.01 0.25 0.15 0.03 0.10 0.01 0.14 0.01 0.28 0.01 0.05 0.200 0.044 0.644 0.200 0.133 0.067 0.111 0.022 0.067 0.022 0.222 0.022 0.044 8.0 0.10 0.044 366.5 35.0 16.5 0.5 3.0 1.0 4.5 20.5 3.0 4.0 1.0 2.5 4.40 0.42 0.20 0.01 0.04 0.01 0.05 0.25 0.04 0.05 0.01 0.03 0.978 0.333 0.333 0.022 0.044 0.044 0.178 0.178 0.044 0.044 0.022 0.022 Bouteloua gracilis Stipa comata Sporobolus cryptandrus Calamovilfa longifolia Andropogon hallii Paspalum stramineum Agropyron smithii Aristida sp. Cyperus & Carex spp. Artemisia filifolia Chenopodium album Euphorbia sp. Croton texensis Salsola kali Amaranthii'SSp. Plantago purshii Cryptantha sp. Conyza canadensis Eriogonum annum Cruciferae spp. Lactuca sp. Unidentified 5.0 0.05 0.022 2.0 0.02 0.022 1.0 6.0 0.01 0.06 0.022 0.111 7.0 0.07 0.155 �490 Table 8. Crown cover (0.01 m2), species composition (%), and frequency of occurrence of vegetation during pre- and post-burn intervals on Burn 3, South Tamarack Prairie, 1984. Pre-burn Vegetation Crown cover Compo Post-burn Freq./ occ. Crown cover Compo Freq./ occ. 2,825.0 125.5 Bare ground Dead vegetation 811.5 486.5 Bouteloua gracilis Stipa comata Sporobolus cryptandrus Calamovilfa longifolia Andropogon hallii Paspa1um stramineum Panicum virgatum Koleria christata Muh1enbergia sp. 665.5 531.0 440.0 390.5 71.0 52.5 41.0 4.0 17.0 17.98 14.34 11.88 10.55 1.92 1.42 loll 0.11 0.46 1.00 1.00 1.00 0.95 0.45 0.40 0.20 0.05 0.20 350.0 95.0 277.5 273.0 36.5 47.5 0.5 17.08 4.63 13.54 13.32 1.78 2.32 0.02 0.95 0.95 1.00 1.00 0.40 0.50 0.05 1.0 0.05 0.05 11.0 0.30 0.55 71.0 3.46 1.00 969.5 26.19 1.00 466.5 22.76 1.00 83.0 2.5 2.24 0.07 0.70 0.10 31. 5 0.5 1.54 0.02 0.80 0.05 322.5 8.71 1.00 9.5 11.5 46.5 0.26 0.31 1.27 0.35 0.10 0.45 85.0 8.0 57.0 5.0 44.0 66.5 2.0 4.15 0.39 2.78 0.24 2.15 3.24 0.10 1.00 0.10 0.75 0.15 0.45 0.30 0.15 1.5 7.5 0.04 0.20 0.15 0.20 0.5 0.01 0.05 13.0 0.35 0.20 2.5 9.0 4.0 3.0 3.5 52.5 0.12 0.44 0.19 0.15 0.17 2.56 0.05 0.25 0.10 0.15 0.10 0.30 0.5 7.0 0.01 0.19 0.05 0.20 1.0 12.0 34.5 9.0 0.5 0.05 0.58 1.68 0.44 0.02 0.05 0.60 0.85 0.20 0.05 Cyperus & Carex spp. Artemisia filifolia Opuntia sp. Mammillaria & Echinosereus spp. Ambrosia psilostachya Artemisia 1udoviciana Tradescantia occidentalis Phlox andicola Evo1vulus nuttalianus Lathyrus polymorphus Penstemon angustifolius Allium textile Psoralea tenuiflora Thelesperma megapotimicum Haplopappus spinulosus Physalis subglabrata Oenothera sp. Mentzelia nuda Ipomoea leptophylla Cymopteris montanus Unidentified perennials Chenopodium album Euphorbia supina Croton texensis Amaranthus sp. Plantago purshii Cryptantha sp. Conyza canadensis Eriogonum annum 0.5 0.01 0.05 2.0 2.0 2.0 0.05 0.05 0.05 0.05 0.05 0.10 �491 Table 9. Crown cover (0.01 m2), species composition (%), and frequency of occurrence of vegetation during pre- and postburn intervals on Burn 3 controls, South Tamarack Prairie, 1984. Post-burn Pre-burn Vegetation Crown cover Bare ground Dead vegetation 805.0 567.5 Boute10ua gracilis Stipa comata Sporob01us cryptandrus Calamovilfa longifolia Andropogon hal1ii Paspalum stramineum Panicum virgatum Muhlenbergia sp. Agropyron smithii Bromus sp. 525.0 679.5 521.5 483.0 58.5 54.0 23.5 38.0 0.5 Compo Freq./ occ. Crown cover Compo Freq. / .occ. 757.0 1,348.0 14:47 18.73 14.38 13.31 1.61 1.49 0.65 1.05 0.01 0.75 1.00 1.00 1.00 0.40 0.30 0.20 0.15 0.05 374.5 348.5 284.5 288.0 37.5 35.0 2.0 19.0 12.94 12.04 9.83 9.95 1.30 1.21 0.07 0.66 0.75 0.95 1.00 1.00 0.20 0.45 0.05 0.20 1.5 0.05 0.10 Cyperus & Carex sp. 46.0 1.27 0.60 80.5 2.78 0.95 Artemisia fi1ifolia 898.0 24.76 1.00 1,173.5 40.54 1.00 48.0 1.32 0.50 37.5 1.29 0.45 195.5 5.39 0.90 11.0 4.5 22.5 2.0 0.30 0.12 0.62 0.06 0.45 0.15 0.25 0.50 25.0 26.0 9.0 44.5 5.5 5.0 3.0 0.5 0.86 0.90 0.31 1.54 0.19 0.17 0.10 0.02 0.25 0.40 0.10 0.70 0.15 0.15 0.05 0.05 4.0 0.11 0.20 7.0 1.5 5.5 12.0 25.0 4.0 0.24 0.05 0.19 0.41 0.86 0.14 0.15 0.05 0.10 0.30 0.10 0.05 26.0 2.0 8.0 1.5 1.0 1.0 0.90 0.07 0.28 0.05 0.03 0.03 0.75 0.15 0.40 0.10 0.10 0.10 Opuntia sp. Lathyrus polymorphus Ambrosia psi10stachya Artemisia ludoviciana Tradescantia occidentalis Phlox andico1a Ev01vulus nuttalianus Penstemon angustifolius Allium textile Psoralea tenuiflora Lygodesmia juncea The1esperma megapotimicum Haplopappus spinu10sus Mentzelia nuda Ipomoea le~hy11a Unidentified perennials Chenopodium album Euphorbia supina Croton texensis Plantago purshii Cryptantha sp. Eriogonum annum Sa1s01a kali 6.0 0.17 0.20 3.0 0.08 0.05 0.5 0.01 0.05 1.0 2.0 0.03 0.06 0.10 0.05 �492 Within Burn 1 grasses, domina ted by blue grama, comprised over 80% of the composition (Table 10); a much greater percentage than in Burn 3. Sandsage and perennial forbs were less abundant in Burn 1, whereas sandsage was the dominant species present in Burn 3. Cacti were much more common in Burn 3. Table 10. Composition (%) of dominant grasses, sandsage, and other covers during pre- and post-burn intervals within combined treatment and control samples, on Burns 1 and 3, South Tamarack Prairie, 1984. Burn 3 Burn 1 Vegetation Bouteloua gracilis Stipa comata Sporobolus cryptandrus Calamovilfa longifolia Andropogon hallii Other grasses & sedges Subtotal (grasses) Artemisia filifolia Cacti Perennial forbs Annual forbs Pre Post Pre Post 42.3a 21.6 3.8 18.4 2.4 0.5 89.1 35.7 15.1 6.3 21.9 4.3 1.0 84.4 16.2 16.5 13.1 11.9 1.8 3.9 63.0 14.6 9.0 11.4 11.3 1.5 5.2 53.0 8.1 0.4 2.1 0.3 8.0 0.4 3.9 3.3 25.5 1.8 9.1 0.1 32.2 1.4 10.4 1.9 alnability to accurately age residual (dead vegetation) may have skewed some species higher than actual on some grasses during pre-burn intervals. Analysis species Burns 1 analyses received of covariance was used to determine if crown cover among species, groups, and other covers differed because of the controlled burns. and 3, although burned on the same date, were not combined for because of differences in soils, vegetation composition, and rainfall following treatments. Blue grama sample means provided evidence of considerable bias resulting, primarily from attempting to obtain the pre-treatment samples in early spring. A major unexpected reduction of blue grama crown cover from spring to late summer was evident in the control whereas a lesser reduction occurred within the burned samples (Table 11). Inability to distinguish 1983 growth in spring 1984, from older residual was considered the primary bias within the blue grama data. As a result, dead vegetation increased dramatically from spring to summer in the control (Table 11). I conclude that the high F value (Table 11) for blue grama should be discounted. - �493 Ta ble 11. Mean crown cover (0.01 m2 )I samplea and analysis of covariance relationships between selected species, species groups, and covers within burn and control samples during pre- and post-burn intervals, Burn 1, South Tamarack Prairie, 1984. Vegetation Bouteloua gracilis StiEa comata SEorobolus Calamovilfa AndroEoson hallii Artemisia filifolia Cacti Perennial forbs Ambrosia & A. ludoviciana Tradescantia Annual forbs Bare ground Dead vegetation Treatment Pre-burn Post-burn x SD x SD Control Post-burn Pre-bum x SD x SD F1.l15 value 362.4 257.7 44.9 185.7 28.7 102.9 3.9 16.3 132.7 83.7 43.8 120.3 28.0 59,7 3.5 13.1 310.2 86.5 75.4 197.3 47.7 75.8 2.9 40.8 108.7 42.8 72.0 109.8 41.9 55.8 2.0 27.6 510.3 187.0 34.4 194.8 21.8 63.5 0.4 27.6 144.2 92.8 19.7 63.2 24.6 42.6 0.8 12.8 325.1 183.2 36.9 191.5 29.2 67.2 3.9 28.8 133.2 89.9 16.4 53.6 29.6 46.0 5.0 15.1 2l.6Sb 38.7s 3.6ns 0.3ns 1.5ns l7.3s 7.7 2.4 3.1 117.3 124.8 9.4 3.1 5.3 63.0 55.6 8.4 18.2 4.1 386.6 7.5 9.9 10.6 3.2 73.9 6.4 11.4 5.9 1.6 104.4 91.7 16.1 4.1 2.3 49.6 25.1 8.9 9.4 50.6 91.3 233.9 10.2 7.0 45.4 50.7 42.7 0.6ns 7.9s 9.8s 132.7s 211.Os r.s= 8.ls aN = 9 for all samples. bS = P < 0.05, ns = not significant The crown cover means and F value for needle-and-thread grass were less suspect than for blue grama (Table 11). This is because a dramatic decrease in needle-and-thread occurred following the fire, whereas samples on the control remained stable. These findings were also supported by visual observations. Needle-and-thread grass made considerable growth prior to the burn and visually appeared to have been set back. These data contradict the information from Kirsch and Kruse (1973). They listed blue grama and needle-and-thread as increasers under controlled burning management. Clearly, more information and more refined sampling on future burns is needed. Evidence of fire impact on other grass species was not detected statistically. Small samples of sand bluestem were possibly a factor because the amount of crown cover almost doubled from spring to summer within the burn, but showed only modest increase within the control (Table 11). Sand bluestem attained much greater height and seed head production within the burns than within their controls based on photo hubs and observations. Sandhill muhly, which was common within Burn 3, from appearance was most severely impacted by the fire although dry soil conditions through the growing season may also have been an important factor. Data in Tables 8 and 9 provide further evidence of the negative impact of fire on sandhill muhly. Sandsage showed marked crown cover decreases as a result of fire (! < 0.05); however, fire removed the old canopy but had little impact in reducing sandsage survival. Sandsage plants were individually marked within both the �494 treatment and control portions of Burns 1 and 3. Only 1 (possibly 2) plants died among 65 marked within the burned areas compared to no mortalities among 60 marked plants wi thin the controls. The rapid and extensive regrowth of sandsage, especially on Burn 3, was considered an important factor in the reduced growth of perennial grasses and forbs' there, especially since summer rainfall was below average. Prickly pear cactus responded to fire in much the same manner as sandsage. Much of the old growth was killed but new growth began soon after the fire. Perennial forbs apparently responded to fire differently within Burn 1 than within Burn 3. Crown cover more than doubled following fire in Burn 1 whereas equal amounts were recorded in spring and summer in the control (Table 11). Greater moisture availability due to more rainfall and less competitive sandsage in Burn 1 may have enhanced perennial forb growth there in contrast to Burn 3 (Table 12). General appearances indicated grasshoppers impacted forbs much more on Burn 3. Perennial forbs decreased on both the treatment and control samples in Burn 3. Table 12. Mean crown cover (0.01 m2)/samplea and analysis of covariance relations between selected species, species groups, and covers within burn and control samples during pre- and postburn intervals, Burn 3, South Tamarack Prairie, 1984. Ve~etation Bouteloua gracilis Stipa comata Sporobo1us cryptandrus Calamovilfa longifolia AndroEo~on hallii Artemisia filifolia Cacti Perennial forbs Ambrosia & A. ludoviciana Tradescantia occidentalis Annual forbs Bare ground Dead vegetation Treatment mean Pre-burn Post-burn 166.4 132.7 110.0 97.6 17.7 242.4 21.4 102.1 80.6 1.0 2.6 202.9 121.6 87.5 23.7 69.4 68.2 9.1 116.6 8.0 85.5 23.2 6.3 14.2 706.2 31.4 Control mean Post-Durn Pre-burn 131.2 169.9 130.4 120.7 15.6 224.4 12.0 61.4 48.9 1.2 1.6 201.2 141.9 93.6 87.1 71.0 72.0 9.4 293.4 9.4 43.4 8.7 4.9 9.9 189.2 337.0 F vaTue 2.3 6.8 0.4 0.3 0.1 23.9 1.7 2.6 0.6 0.3 0.7 280.2 90.9 aN = 4 for all samples. b'S = P < 0.05, ns = not significant. Western ragweed was the most abundant perennial forb within both burn sites and their controls. It was not distinguished from cudweed sage during pre-burn sampling so the two were combined for analysis. Analysis showed little evidence that fire impacted their presence. Major reduction from spring to fall was evident on Burn 3, possibly because of a combination of deficient soil moisture and abundant grasshoppers. Sampling was not conducted but grasshoppers appeared to be in greater concentration or at least more �495 destructive to vegetation on Burn 3 than within Burn 1. Fire possibly enhanced spiderwort, the second most common forb on Burn 1 (P < 0.05) (Table 11). A similar increase was also recorded on Burn 3 (Table 12): Annual forbs were not tallied often during pre-treatment sampling within either Burns 1 and 3. Possibly this was a consequence of over-winter decomposition. Annuals remained sparse following treatment within Burn 1 with Texas croton (Croton texensis) the primary species present in summer. It was most often observed growing near the base of burned sandsage. Annual forbs showed much higher densities in late summer control samples within Burn 1 (P < 0.05) where lambsquarter was by far the most abundant. Milk purslane (Euphorbia spp.), horseweed (Conyza canadensis), and Texas croton were tallied frequently but were of lesser importance (Tables 6 and 7). Annual forbs did not show the dramatic spring to summer increase on Burn 3 controls as evident in Burn 1 (Table 12). Species composition was similar on the two burn sites. As would be expected, removal of dead vegetation and sandsage with fire dramatically increased the amount of bare ground within both burns. Bare ground was greater both prior to and following treatment in Burn 3 than in Burn 1. Dead vegetation was apparently undersampled during pretreatment spring sampling on both sites. As a consequence, high F values were attained. However, significant reductions of residual would have undoubtedly occurred due to fire even if the bias did not exist. Pre-Treatment Crown Cover Sampling of Proposed 1985 Burn Sites Point-frame sampling of pre-treatment vegetation crown cover was completed within three proposed burn sites and 3 adjacent controls (Fig. 1) during fall 1984. Sampling was completed prior to random selection to determine which would receive treatment. Although crown cover and composition varied among sites (Table 13), results show that blue grama (23.7%), needle-and-thread (22.5%), prairie sandreed (24.2%), sand dropseed (4.2%), and sand sagebrush (14.4%) dominated the composition (89%). Other species and their relative importance are summarized in Table 13. All sites contain some sample points within moderate to dense stands of sandsage; however, grasses comprise the dominant overall composition in all. Growth of the preceding growing season was more easily distinguished from old residual than in spring 1984 sampling; however, for some species (primarily blue grama) it was still difficult. Findings (Table 13) indicate almost 60% of the crown cover was dead vegetation (including litter) and it and bare ground combined comprised two-thirds of the crown cover. Revegetation Treatments Fair to good stands of grasses and alfalfa were achieved in all seeded and interseeded plots. Early summer 1984 inspections could not discern differences among plots receiving and not receiving the atrazine herbicide treatment. However, relatively dry weather throughout most of the summer, combine,d with considerable growth of lambsquarter and other forbs caused a pronounced difference between herbicide-treated and untreated plots by late summer. Grass seedlings on untreated plots became moisture deficient and made little growth after mid-summer. Seedlings in atrazine plots continued to grow into early fall, attaining heights of 0.3 m or more and a few produced seed heads. Random samples obtained in October 1984 revealed greater abundance of �496 Table 13. Vegetation crown cover, bare ground, dead vegetation, and overall composition (%) based on point-frame sampling of proposed spring 1985 burn and control sites, South Tamarack Prairie, Fall 1984. Vesetation Bare ground Dead vegetation Subtotal Boutelous gracilis StiEa~ SEorobo1us cryptandrus Calamovi1fa 10ngifo1ia AndroEogon ha11ii PasEa1um stramineum AgroEyron smithii Aristida sp. Koleria cristata Panicum virga tum Muhlenbersia sp. Subtotals 1 west 1 east 2 west 2 east 3 west 3 east 305 2,607 438 2,538 370 3,444 330 3,397 624 3,148 503 3,207 685 489 104 536 20 2 434 564 103 506 42 222 471 62 405 11 318 357 49 414 11 331 152 61 352 14 427 266 46 253 11 48 2 36 5 9 3 1 1 66 2 55 ~sp. Panicum caEil1are 1 3 4 284 323 0Euntia sp. Mammillaria 13 9 Ambrosia Esi10stachya Artemisia ludoviciana Tradescantia occidenta1is Phlox andicola E:vOIVulus nutta1ianus Lathyrus EolymorEhus Psorlea spp. ThelesEerma sp. HaEloEaEEus sEinu10us IEomoea 1eEtoEhy11a Mentze1ia nuda Physalis subglabrata Cichorium intybus Lysodesmia juncea Oenothera sp. SEhaera1cea coccinea Cirsium sp. Unidentified perennials Subtotal 45 51 ChenoEodium album EUEhorbia sp. CrYEtantha sp. Conyza canadensis Salsola kali Helianthus sp. Croton texensis Eriosonum ~ Polysonum aviculare Lesquerella 1udoviciana Subtotal 38 6 3 9 2 CYEerus Artemisia &~ 15 13 sp. filifo1ia 9 3 14 14 2 1 5 4 ComEosition 8.2626 58.9667 67.2293 7.7707 7.3913 1.3664 7.9282 0.3504 0.0064 0.5112 0.0386 0.0032 0.0514 0.0418 25.4596 23.7123 22.5547 4.1695 24.1931 1.0694 0.0196 1.5599 0.1177 0.0098 0.1570 0.1235 77.6865 0.1768 0.0032 0.5396 0.0098 2 2 3 0.0450 0.1373 55 167 408 230 4.7164 14.3922 5 1 11 29 57 0.3987 0.0032 1.2165 0.0098 20 2 4 3 1 1 6 4 3 8 7 1 6 10 2 1 1 18 31 4 0.3761 0.2090 0.0932 0.0161 0.1575 0.0514 0.0418 0.0289 0.0096 0.0386 0.0032 0.0129 0.0032 0.0032 0.0032 0.0129 0.0032 0.0964 1.1603 1.1478 0.6377 0.2845 0.0490 0.4807 0.1570 0.1275 0.0883 0.0294 0.1177 0.0098 0.0392 0.0098 0.0098 0.0098 0.0392 0.0098 0.2943 3.5413 0.5530 0.0072 0.0161 0.0482 0.0096 0.0354 0.0354 0.0193 0.0064 0.0064 0.8070 1.6874 0.2354 0.0490 0.1472 0.0294 0.1079 0.1079 0.0589 0.0196 0.0196 1 1 2 6 4 3 1 6 3 5 2 6 2 1 1 1 4 1 3 27 42 16 Crown cover 45 2 33 5 1 9 3 5 6 8 1 1 1 2 6 2 1 1 1 1 1 ~ �497 seeded grass (P < 0.05) on atrazine-treated warm-season grass plots whereas untreated warm=aeason grass plots contained much greater abundance of annual forbs (p < 0.05) (Table 14). Considerable native perennial grass and forb cover dOminated all sites since the single tillage did not kill all vegetation, especially that which was deep rooted. Table 14. Mean ta11ies/1.5-m2 samples among revegetation plots and treated with atrazine herbicide, South Tamarack Prairie, 1984. Treated Plot 1 Plot 2 Vegetation Seeded warm-season grasses Perennial native grasses Annual grasses Perennial forbs Annual forbs Sand sagebrush Cacti 13.5 33.4 0.3 2.6 0.1 0.9 1.1 10.1 31.1 0.3 1.8 0.6 0.2 1.1 Untreated Plot 1 Plot 2 3.7 17.9 0.5 10.8 7.1 0 0.8 4.0 32.8 0.5 2.9 11.1 0.4 0.8 untreated -F 1,2 value 21.70 0.84 1.37 18.84 Dry summer conditions and grasshoppers defoliated the alfalfa limiting its growth. Random sampling of the warm-season tall wheatgrass-a1fa1fa plots and the interseeded plots will be conducted in 1985. Little vegetation was available there to sample in fall 1984. Wildlife Use of the South Tamarack One pair of upland plovers (Bartramia longicauda) was observed on Burn 1 two or three times in late May and early June but apparently did not nest on the South Tamarack. A female pronghorn (Anti1ocapra americana) was observed on Burn 1 in late June apparently in the act of defending a fawn against a nearby coyote (Canis 1atrans). She was not observed on the property or site again. Up to 25-27 Swainson' s hawks (Buteo swainsoni) were attracted to the burns during treatment and were observed to take advantage of the fire's exposure of small rodents, grasshoppers, and herpti1es. Mourning doves (Zenaida macroura), ring-necked pheasants (Phasianus co1chicus), and greater prairie-chickens (Tympanuchus cupido) were observed or feathers and other sign of their presence were observed within the fire breaks surrounding the controlled burns. Mourning doves were common nesters there and several nests were lost to fire. Three nests of mallards (Anas p1atyrhynchos) were located, one of which was destroyed by the fire. None of the nests was successful. Striped skunks (Mephitis mephitis) were occasionally observed on the South Tamarack Prairie but badgers (Taxidea taxus) were notable by their absence. No evidence of their recent presence was observed. Pocket gophers (Geomys sp.) were abundant as were thirteen-lined ground squirrels (Spermophi1us tridecem1ineatus) as food sources for badgers. There was little evidence of jack rabbits (Lepus spp.) or desert cottontails (Sy1vi1agus audubonii) although both were assumed to be present in low numbers. �498 LITERATURE CITED Amen, A. E., D. L. Anderson, T. J. Hughes, and T. J. Weber. 1977. Soil survey of Logan County, Colorado. U.S. Dep. Agric., Soil Conserve Serv., Washington, D.C. 252pp. + Append. Bragg, T. B. 1978. Effects of burning, cattle grazing, and topography on vegetation of a choppy sands range site in the Nebraska sandhills prairie. Proc. Int. Rangeland Cong., Soc. Range Manage. 1:248-253. Floyd, D. A., and J. E. Anderson. 1983. A new point intercept frame for estimating cover of vegetation. Pages 107-113 in Idaho Natl. Eng. Lab. Radioecology and Ecology Programs. U.S. Dep. Energy DOE/ID 12098. Kirsch, L. M. 1977. Instructions for use of the height-density pole for measuring residual vegetation on grassland wildlife habitats. U.S. Dep. Inter., Fish and Wildl. Serv., Jamestown. Unpubl. Rep. 2pp. ____ ~~, and A. D. Kruse. 1973. Prairie fires and wildlife. Timbers Fire Ecol. Conf. 12:289-303. Proc. Tall _______ , H. F. Duebbert, and A. D. Kruse. 1978. Grazing and haying effects Trans. North Am. Wildl. and Nat. of habitats of upland nesting birds. Resour. Conf. 43:486-497. Robel, R. J., J. N. Briggs, A. D. Dayton, and L. C. Hulbert. 1970. Relationships between visual obstruction measurements and weight of grassland vegetation. J. Range Manage. 23:295-297. Schmutz, E. M., M. E. Reese, B. N. Freeman, and L. C. Weaver. 1982. Metric belt transect system for measuring cover, composition, and production of plants. Rangelands 4:162-164. Snyder, W. D. 1984. Ring-necked pheasant nesting ecology and wheat farming on the High Plains. J. Wildl. Manage. 48:878-888. Prepared by 1i1ct&wJ ~ Warren D. Snyder Wildlife Researcher C �Colorado Division of Wildlife Wildlife Research Report April 1986 499 JOB PROGRESS REPORT State of --~-Colorado -- Avian Research Project _0,;;_;1:;_-__;0_;.3_-...;_04_5;.._..;('-W_-_37.;_-_R....:.) _ Work Plan 21 ------ Job Title: __ 3 ..;;,__- Sandsage-Bluestem Prairie Renovation Period Covered: Author: : Job 01 January - 31 December 1985 Warren D. Snyder Personnel: C. E. Braun, W. Brown, L. Budde, M. Creamer, L. Crooks, T. Davis, K. Dillinger, M. Etl, M. Gardner, T. Kroening, W. Miles, J. Palic, F. Pusateri, and W. Snyder, Colorado Division of Wildlife. ABSTRACT Vegetation renovation and revegetation were continued within the Tamarack Prairie by persouneI of the Northeast Region to enhance sandsage-bluestem (Artemisia-Andropogon) prairie for prairie grouse. Controlled burning of 59 ha was conducted in mid-May 1985. Revegetation was conducted on an additional 20.2 ha of rangeland. Evaluation of these and previous (1984) manipulations was continued and is summarized here. Phenology was markedly early during spring 1985 and precipitation, although deficient in June, approximated the expected annual norm. The height-density quality of residual vegetation with 1984 burn sites was reduced by burning (p < 0.05). Residual grass-forb cover within burn 1-84 changed from 0.25 dm (pre-treatment, spring 1984) to 0.14 dm (early spring, 1985) whereas visual obstruction readings increased slightly within control sites. Sandsage (Artemisia filifolia) pre-treatment obstruction was 0.87 dm compared to 0.31 dm in spring 1985 .within the same burn site (p < 0.05). Pre-treatment height-density indices for grass-forb vegetation within the 3 1985 burn sites averaged 0.38 dm (± 0.02) compared to 0.32 dm (± 0.02) for controls. All grass-forb indices revealed that residual cover was marginal in quality for nesting prairie grouse. Pre- and post-treatment analyses of vegetation crown cover within both 1984 and 1985 burn sites showed no marked changes. Sandsage regrew rapidly within 1984 burns and the 4 dominant grasses showed little change in importance. Prairie sandreed (Calamovilfa longifolia) was the only 1 of the 4 to markedly increase (R < 0.05) within the 1985 burn sites. Switchgrass (Panicum virgatum) and bluestems were not abundant enough within the "native" prairie to ascertain the impact of fire on them. However, they had increased by fall 1985 within an interseeded portion of burn 1-84 when contrasted to controls (p < 0.05). Visual inspections within the 1985 burns also indicated growth and seed head production was markedly enhanced. Perennial forbs were not markedly affected by fire and some increase in annual forbs within the 1984 burn sites was �500 noted. Evaluations concerning altering the time of spring burns to impact sandsage showed sandsage mortality was not increased by delayed spring burning but regrowth and vigor potentially were reduced. An attempt to partially reduce sandsage canopy cover by aerial strip spraying of 2,4-D herbicide in June 1985 caused almost total mortality of sandsage and annual forbs within a 80.8-ha site. Discing of 19 revegetation strips followed by application of a low rate of atrazine herbicide and seeding yielded fair stands of switchgrass which attained excellent growth in 1985. Discing plus atrazine herbicide treatment in spring also released deep-rooted tall warm-season grasses in previously interseeded tracts. The interseeded grasses attained excellent growth (p< 0.05) when contrasted to controls indicating this approach can be used to speed dominance by these species. �501 SANDSAGE-BLUESTEM PRAIRIE RENOVATION Warren D. Snyder The Tamarack Prairie within the South Platte Wildlife Area contains approximately 1,680 ha of grassland that has been designated for establishment of greater prairie-chickens (Tympanuchus cupido). Livestock grazing was discontinued on the site following 1977 and about 120 ha were interseeded in 1981-82 with varying success. Based on recommendations of a consultant, L. Kirsch, controlled burning was implemented on 3 tracts totaling approximately 112 ha in spring 1984. Weather factors delayed 1985 burning and resulted in burning only 59 ha within 3 small tracts. This study was initiated to evaluate controlled burning, and other renovation and revegetation approaches for increasing vegetation quality on the South Tamarack sandsage-bluestem prairie. Results should show which approaches are most economical and effective for increasing height-density quality and enhancing the composition of native warm season grasses. Findings of the second year of study are summarized here. P. N. OBJECTIVES Test renovation and revegetation techniques for increasing standing residual height-densi ty of grasses, increasing the proportion of tall, warm-season grasses within the composition, and for reducing the quantity of sand sagebrush to < 30% canopy cover in an ungrazed sandsage-bluestem prairie on the South Tamarack, South Platte Wildlife Area in northeastern Colorado. SEGMENT OBJECTIVES Monitoring of environmental and vegetation conditions and changes continued as follows: 1. Precipitation was monitored throughout the year supplementing electronic rain gauge data with information from nearby weather stations through the winter months. 2. Soil moisture accumulations, plant phenology, and weather were monitored primarily in spring and especially at time of controlled burns. 3. Visual obstruction (height-density) measurements were treatments and controls where applicable in late winter spring prior to green-up. 4. Crown cover, species composition, and frequency of occurrence measurements were obtained from mid-summer to early fall. 5. Photos of treatment and controls were taken in October. Data compilation and writing the annual during fall and winter 1985-86. job progress obtained on and/or early report was conducted �502 METHODS The major approaches used within this study were previously summarized (Snyder 1985). Supplements or modifications are summarized here. Use of a soil moisture meter to determine percent moisture at 5, 15, and 30-cm depths at time of controlled burning was discontinued in part because measurements were neither accurate or meaningful. A soil probe approximately 1.9 m long and 1.4 cm in diameter was used to measure the accumulated soil moisture. This device, used by agronomists for determining subsoil moisture in croplands (D. Smika, pers. commun.), provided reliable indices of spring subsoil moisture accumulations in the fine sandy loam or loamy sand soils of the South Tamarack. Tests showed that when the probe was inserted by hand into the ground it consistently penetrated the moist soil but stopped in the transition zone to dry soil. The probe could be used in spring before top soils dried hindering penetration and could also be used after summer rains to sample moisture accumulations. Two rain gauges that recorded each 0.025 cm (0.01 in) of rainfall were installed near burn 1-84 and east of burn 3-85 in spring 1985 to monitor rainfall through the growing season. The metric belt transect system used to obtain crown cover, species composition, and frequency of occurrence within burn 1-84 and 2-84 was modified slightly in 1985 to reduce intensity of sampling. In 1985 every other 31.6-cm segment of the 7.9-m transect was skipped yielding 13 placements rather than 25 along each belt. Sampling of the 1984 burns and their controls was conducted in late July 1985. Point frame sampling of 1985 burns and proposed 1986 burns and their controls was initiated on 12 August 1985 and continued through the end of the month. Approximately 21 ha (50 acres) of rangeland dominated by needle-and-thread (Stipa comata) and blue grama (BouteLoua gracilis) within the northeast to middle east portions of the Tamarack Prairie (Fig. 1) were selected for revegetation treatment in late winter 1985. Nineteen strips of varying length each 26 m wide were positioned at approximate right angles to prevailing northwest winds. The strips were double disced by management personnel after the ground thawed in late winter. Supplemental treatments including harrowing with a spike-toothed harrow and application of atrazine herbicide at the rate of 0.75 kg/ha (0.67 1b/ac). Seeding of a switchgrass-dominated tall, warm-season grass mixture followed within 1 or 2 days of the herbicide treatment in late April 1985. Two narrow (disc-width) strips within fair to good stands of interseeded grasses were sha.Ll.ow-df.sced by M. Gardner in March 1985 within each of 2 interseeded tracts. One disced strip was sprayed with atrazine herbicide in late April 1985 at the rate of 0.75 kg/ha (0.67 lb/ac) and the other remained untreated within each site. Six random points were subsequently selected for height sampling of the interseeded grasses in fall 1985. Six samples per random point were subsequently obtained within the disced and disced plus atrazine treatments, and their respective adjacent controls. A herbicide treatment to reduce sandsage canopy cover was implemented by management personnel in spring 1985 within a 83.3-ha site in the east central portion of the Tamarack Prairie (Fig. 1). An aerial applicator was contracted �TAMARACKPRAIRIE (WEST PART) Scale N o [ill. Interseeded 2.1 1 mi. 3/4 1/2 1/4 ..., tract ; ...~......0".....\. Section number . "..-.•..:~..-;.... ?"'B~N ••.....•Burn·tr~ct boundary Goo ---- \ 'l'k'all 50 .\ I '" \. _.,BURN 1-184 '. " ~ •• " 1 1 ••••••••••••••• .~ ~ "\ \ ~ ® t."lIo.. 6P ~ o· I ,".,~ I:I"Q ~ , ;,.-.. 'Y', ~~ " ~, Fig. 1. -~M'" i.a~s- -IPlN'lI!.. ".".(~ .. "'1------ ____ .- --- ·'\ 2-84 \".. Windmill .,., 1 '., / az;::.:J' "0( ~ ~ •••••••••• ~ ~ II ~ I!... ~ ~· Burn sites and 1981-82 interseeded tracts within the west part of the Tamarack Prairie. VI o w �504 to apply 0.84 kg/ha of 2,4-D low vol with 18.7 l/ha of water (0.75 lb/ac. of herbicide in 2 gal. of water) plus a wetting agent in l8.3-m wide strips at 36.6-m wide intervals (50% cover) during the bloom stage of sandsage , The site, containing varying moderate to high sandsage densities, was gridded and 12 random points under the center of the spray plane path were selected for vegetation sampling. At each of the 12 points a transect was laid out to obtain 8 36-point frame samples at l-m intervals. RESULTS AND DISCUSSION Environmental Conditions and Controlled Burning - 1985 In contrast to below normal temperatures that suppressed vegetation growth in April 1984, March through May 1985 temperatures consistently averaged above normal stimulating early spring green-up on the South Tamarack (Table 1). Vegetation phenology was about 3 weeks earlier than in 1984. Warm March temperatures stimulated early green-up and rapid growth of cool-season plants. Continued warm weather in April and May stimulated growth of warm season grasses starting in mid-April. In contrast, warm season grasses had not begun new growth at the time of the 4 May 1984 controlled burns. At the time of the 16 May 1985 burns, needle-and-thread was already producing seed heads, prairie sandreed was about 30 cm tall, spiderwort (Tradescantia occidentalis) was in bud to early bloom stages, and sweet pea (Lathyrus polymorphus) was past major bloom (Table 2). Sandsage was in early leafing stages when burned in 1984 but had leafed and attained considerable new growth at the time of the 1985 burns. The 1985 burns would have had to be conducted in about the third week of April to compare phenologically with the 1984 burns. The 1984 burns were conducted before emergence of prairie sandreed and sand bluestem and were probably timed at least 1-2 weeks too early for optimum enhancement of tall, warm season grasses. Initially, 1985 controlled burns were scheduled for 6 May but wind from the south that would carry the smoke over 1-76 forced repeated postponements until mid-May. Two small test burns completed on 6 May showed that although much new growth was present, enough residual remained to carry fire. When favorable weather conditions for controlled burning occurred on 16 May, much additional vegetation growth had occurred. The fires set on portions of the 3 burns were notably cool and suppressed and did not carry well in locations where residual was sparse (e.g., interseeded tracts). Fire guards were not needed on the windward side of the burns which were quickly suppressed with spray from pumper units. Originally, fire guards had been established to burn 63.8 ha in site 1, 45.7 ha in site 2, and 41.6 ha in site 3. However, because of the uncertain conditions the sizes of the burns were reduced to 17.8 ha in site 1, 26.7 ha in site 2, and 14.5 ha in site 3 yielding 59 hectares (146 acres) combined, or 39% of the original 151.1 ha (Figs. 1, 2). However 17 of the 36 random transects were included within the 1985 burns and an equal number of control transects were used for comparison. Precipitation received in October and December 1984, and April and May 1985 provided soil moisture considered adequate for conducting burns in 1985. Comparisons were made between a home-made and a custom soil probe acquired �505 Table 1. Precipitation (inches) and monthly average temperature (F) departures recorded in 1984 and 1985 in relation to long-term averages, Tamarack Prairie and nearby weather stations. Month Jan Feb Mar Apr May Jun Ju1 Aug Sep Oct Nov Dec Annual PreciEitation b Lons-term ~ 1984 19851: 0.32 0.35 0.78 2.02 2.55 2.52 2.09 2.04 1.37 0.99 0.44 0.49 0.47 1.12 0.66 3.42 2.01 2.80 0.49 0.66 0.44 1.39 0.01 0.80 0.47 0.13 0.38 2.24 2.49 0.82 3.78 0.83 1.42 0.37 1.06 1.20 15.96 14.27 15.19 Departure from x temEerature 1984 1985 -4.8 0.7 0.4 -5.9 0.6 -1.8 0.3 3.5 -0.7 -3.0 2.9 1.2 0.1 -7.1 7.1 4.4 3.6 -1.7 0.5 0.5 -1.8 -2.1 -7.1 -7.8 aAverage of long-term data from Sterling and Sedgwick weather stations. bData from H. Hamilton, Private weather recorder, Crook, Colorado. cTamarack Prairie data from 2 precipitation guages supplemented with weather station data during winter months. �Vl o 0\ Table 2. Phenological conditions of selected vegetation during spring 1985, Tamarack Prairie. April Vegetation 1 Ma 6 16 Full bloom 2.5-5 em Allium textile Ambrosia psilostaehya Artemisia filifolia A. ludovieiana Cympoteris montanus Evolvu1us nutta1ianus Ipomoea leptophylla Lathyrus po1ymorphus Lesquerel1a ludovieiana Leueoerinum montanum Mentze1ia nuda Penstomen angustifo1ius Phlox andieola Basal 20 em Greening Tradeseantia oeeidentalis 10-12 em Andropogon hallii Agropyron repens A. smithii Boute10ua gracilis Calmovilfa 10ngifo1ia Muhlenbergia pungens Panieum virgatum Stipa eomata 22 1 Bud Nearly leafed Emerging - 2.5 em Blooming No growth 5 em Fully leafed Budding 12 em Early bloom 2.5 em Blooming Mid-bloom Early bloom Full bloom 20-30 em Full bloom Early to mid bloom Bud to early bloom 5 em 10-15 ell 8-10 em 5-10 em 20-25 em 15 dm 5-7 em No growth 30 em 30 em 1-2 em 5 em 6 em 20-25 em 25-30 em 35-40 em heading �TAMARAcK N PRAIRIE (EAST PART) c:~-- ...•• _ Scale o 114 0] Interseeded ZI -, 112 3/4 1 mi. tract ill B Section number ........ Burn tract boundary G- Windmill ..---" .. o .-. I.; \. - --- Trail mtf~:~:~:! , .«~tIP. .• I I II , I •...-- ..••. ,.-\&l.......•.... . \ \BURN 2-85 \ \\ \ \I \ \\ \ ._1 \'\ \ ~ •••••• \.-~" \ ,I b... eJl 0 ~ ~ !?__•.. " ,, ,~- •••'\! ~ 20 I ~ 'U. I I ,, f3 , .~ \ \ \ \ \ (j \ za 21 , '~ ,'' ~. \ \ I , , •• ' ••. '. \ 't····· I ••••••• ~ts 3'::C!4. '" -, \.... \.' ,'}pI &1 ,,' " .... ~&-_- ••---.\ '._,,\ . _-- --_ ---' \ ~ 1 \ \ \ \ -, ,,~~- ~ I _ ..__.1 Tamarack Prairie. Fig 2. Burn sites, 1981-82 interseeded tracts, and the sandsage spray site within the east part of the l.n o " �508 from Monsanto Chemical Company within burn 1-84 and its controls on 22 April 1985. Results revealed that either probe could be used with equal effectiveness and accuracy. Sandy soils retain about 2.5 - 3.8 cm of moisture/0.3 m (1 to 1.5 in./ft.) (D. Smika, pers. commun.). Therefore, South Tamarack soils contained at least 5 cm (2 in.) of moisture on 22 April when average soil probe penetration was 0.67 m (N = 88), and at least 7.5 cm (3 in.) on 17 May when average soil probe penetration was 1.1 m (N = 26). Soil probe penetration averaged 1.2 m on 5 and 13 June, however dry soil surface made penetration difficult at some locations at the latter date. Soil probe measurements obtained on 23 October 1985 after completion of the growing season and recent snows and rains averaged 0.4 m. Precipitation received in April and May 1985 was near long-term monthly averages, however, rainfall was markedly deficient in June, a month important for growth of warm-season grasses (Table 1). Rains began in mid-July alleviating the prolonged dry interval and exceeded the expected average for July. August rainfall was deficient but September precipitation approximated the expected and stimulated late growth of most grass species including cool-season species. October 1985 was relatively dry, but about 20 and 35 cm of snow were respectively received on 9 November and the week of 9 December. These snows, and other light snows and cold temperatures retained snow cover on the South Tamarack from 9 November into 1986. April through September precipitation in 1985 was about 2.5 cm below the long-term average for the interval but late fall precipitation brought 1985 total precipitation to near the long-term average (Table 1). Vegetation Sampling and Evaluation On Burned Sites Height-Density Sampling.--Controlled burns conducted on sites 1 and 3 in 1984 markedly reduced the height-density quantity of residual vegetation available to wildlife in spring 1985 (P < 0.05, Table 3). This included impacts on grass-forb cover, sandsage, aDd both in combination. The fires, by removing residual, reduced the overall vegetation density close to the ground. However, increased height and seeding by blue grama, prairie sandreed, and sand bluestem (Andropogon halHi) were noted following the 1984 fires and growing season. No marked difference in seed-head production was observed between 1984 burned sites and their controls during or after the 1985 growing season. Pre-treatment height-density sampling was completed on 1985 burns, their controls, and proposed 1986 burns and controls (Table 4). Data from the 17 transects within the 3 1985 burns were consolidated into one index as were the 19 transects within proposed 1986 burns and controls (Table 4). Height-density indicies ranged from 0.28 to 0.38 dm for grass-forb vegetation and from 0.57 to 0.74 dm for sandsage. Sandsage obstructed 20% of the readings in burn 1-84 but only 13% in that site in 1985 after the fire whereas it obstructed 17% in 1984 and 16% in 1985 within burn 1-84 controls. Within burn 3-84 sandsage respectively obstructed 45 and 46% of the treatment and control height-density pre-treatment readings whereas 26 and 52% of the respective post-treatment readings were obstructed by sandsage. Sandsage density was much reduced in 1985 burn sites where 11% of the pre-treatment height-density readings were obstructed by sandsage. �509 Height-density (dm) means of residual grass-forb and sandsage Table 3. vegetation within 1-84 and 3-84 treatments and controls from early spring 1984 through early spring 1985, Tamarack Prairie. Vegetation Treatment Pre-treat (84) Post-treat (85) F-value Burn Site 1-84 Grass-forb Burned Control 0.253 0.252 0.136 0.296 47.5a Sandsage Burned Control 0.871 0.762 0.310 0.671 40.la Combined Burned Control 0.373 0.337 0.162 0.355 66.la Burn Site 3-84 Grass-forb Burned Control 0.222 0.183 0.021 0.191 22.5a Sandsage Burned Control 0.827 0.935 0.121 0.797 l14.la Combined Burned Control 0.492 0.531 0.047 0.503 l87.0a ap < 0.05. Table 4. Height-density values (dm) for grass-forb, sandsage, and combined covers within 1985 burns and their controls and proposed 1986 burns and their controls, pre-treatment 1985, Tamarack Prairie. 1986 1986 Vegetation variable Burned transects Control transects Burned transects Control traansects 828 0.38 ± 0.02 781 0.32 ± 0.02 902 0.29 ± 0.02 951 0.27 ± 0.015 132 0.74 ± 0.09 179 0.63 ± 0.07 166 0.57 ± 0.08 117 0.68 ± 0.09 960 0.43 ± 0.025 960 0.38 ± 0.02 1,068 0.33 ± 0.02 1,068 0.32 ± 0.02 Grass-forb N x Sandsage N "'X Combined N x �510 Height-density indices < 0.5 cm were considered marginal for nesting by prairie grouse (Snyder 1985). Both pre- and post-treatment height-density indices for grass-forb vegetation were markedly below 0.5 dm (Table 3). Studies by Westemeier (1972) and others show little nesting use of burned grassland by grouse during the first growing season following fire. Nesting use increased as cover quality became optimum during the second through fourth years following fire in areas receiving higher rainfall and attaining excellent grass production. Based on findings of this study in northeastern Colorado where annual precipitation is usually < 40 cm, residual cover conditions remain marginal for nesting into the second growing season following burning. The dominant grasses (blue grama, need1e-and-thread, prairie sandreed, and sand dropseed (Sporobo1us cryptandrus) must be considered in assessing the capacity of this rangeland to respond to fire. Additiona1 years of monitoring are needed before drawing conclusions. At present, it does not appear that fire yields the dramatic response within this semiarid mixed-grass prairie that is obtained in tall-grass prairie in locations receiving higher annual rainfall. Similar findings were reported by Bragg (1978) and Engle and Bultsma (1984). Crown Cover, Composition, and Frequency of Occurrence.--Crown cover (0.1-m2) and other vegetation data were obtained in late July 1985 for burns 1-84 and 3-84, and their controls (Tables 5, 6). June and early July dry weather retarded growth of most vegetation in 1985, which, when combined with the earlier sampling, yielded lower crown cover tallies than were recorded in 1984 (Table 7, 8). Late July and September rains stimulated additional growth of most grass species following sampling. Crown cover (point frame), species composition (%), and frequency of occurrence data from 1985 burns and their controls (obtained in 1985) varied (Table 9). The three 1985 burns included 17 of the original 36 transects (5 transects in burn 1-85, 8 transects in burn 2-85, 4 transects in burn 3-85) and these transects were combined in the preliminary analyses. Vegetative data for the remaining 19 unburned transects and controls proposed for burning in 1986 are summarized in Table 10. Mean crown cover and analyses of covariance relationships for selected species, species groups, and covers within burns 1-84 and 3-84 and burns 1-85, 2-85, and 3-85 (combined) are summarized in Tables 7, 8, and 11. Comparisons of composition during pre- and post-treatment intervals· for more important species within burns 1-84 and 3-84 are respectively summarized in Tables 12 and 13. Blue Grama.--Snyder (1985) reported, pre-treatment (1984) sampling of blue grama was considered biased because growth of the previous growing season could not be distinguished from older residual (pre-treatment sampling was conducted in early spring 1984). Therefore, pre-treatment (1984) and 1985 comparisons should probably be discounted. August 1984 to July 1985 post-treatment comparisons showed no marked changed (p> 0.05, Table 11). The 1985 data are considered more reliable than that collected in 1984. Blue grama had attained considerable growth prior to the 1985 burns contrasted to no new growth at the time of the 1984 fires. Need1e-and-thread.--Crown cover samples showed evidence that fire reduced the amount of need1e-and-thread in burn 1-84 (p < 0.05) and within the 1985 combined burns (P < 0.05) (Tables 7, 11). Evidence of fire impact could not be detected in burn-3-84 (Table 8). Some recovery of need1e-and-thread was noted �511 Table 5. Crown cover (0.01-m2), species composition (%), and frequency of occurrence of vegetation within treatment and control transects on Tamarack Prairie burn 1-84 during July 1985. Treatment Vegetation Bare ground Dead vegetation Boute10ua gracilis Stipa comata Sporobo1us cryptandrus Ca1amovi1fa 10ngifo1ia Andropogon ha11ii Agropyron smithii Aristida sp. Paspa1um stramineum Muh1enbergia sp. Ko1eria cristata Unid. annual grass Cyperus & Carex spp. Artemisia fi1ifo1ia Opuntia sp , Mammi1aria & Echinocereus Ambrosia psi10stachya Artemisia 1udoviciana Tradescantia occidenta1is Phlox andico1a Evo1vulus nutta1ianus Lathyrus po1ymorphus Penstomen angustifo1uis The1esperma megapotimicum Map10pappus spinu10sus Mentze1ia nuda Asclepias Erigeron sp. Chenopodium album Plantago purshii Eriogonum annum Pepidium densif10rum Cryptantha sp. Lesquere11a 1udoviciana sP:- Crown cover Compo Control Freq.! occur. 2,146.0 1,413.0 545.5 483.5 274.0 343.0 77.0 9.0 1.0 23.81 21.10 11.96 14.97 3.36 0.39 0.04 0.978 1.000 0.978 0.978 0.511 0.200 0.022 0.5 1.0 9.0 7.0 393.0 11.0 1.5 5.0 0.02 0.04 0.39 0.31 17.15 0.45 0.07 0.22 0.022 0.044 0.222 0.200 0.867 0.178 0.067 0.111 26.5 6.0 6.0 1.0 3.5 12.0 0.5 1.16 0.26 0.26 0.04 0.15 0.52 0.02 0.489 0.156 0.133 0.044 0.089 0.222 0.022 0.5 0.5 45.5 4.0 0.5 16.0 7.5 0.02 0.02 1.99 0.17 0.02 0.70 0.33 0.022 0.022 0.733 0.178 0.022 0.356 0.133 Crown cover Compo Freq.! occur. 1,082.5 2,814.5 433.5 588.5 86.0 447.5 45.5 8.5 3.0 2.0 22.20 30.13 4.40 22.91 2.33 0.44 0.15 0.10 1.000 1.000 0.822 1.000 0.400 0.067 0.044 0.022 0.5 0.5 6.5 231.5 13.5 0.5 5.5 6.5 29.5 9.5 7.0 1.0 3.5 10.0 0.03 0.03 0.33 11.85 0.69 0.03 0.28 0.33 1.51 0.49 0.36 0.05 0.18 0.51 0.022 0.022 0.133 0.778 0.289 0.022 0.111 0.044 0.467 0.178 0.089 0.044 0.067 0.178 1.0 0.05 0.022 1.0 1.5 1.5 1.5 2.0 2.0 2.5 0.05 0.08 0.08 0.08 0.10 0.10 0.13 0.022 0.067 0.044 0.044 0.067 0.067 0.111 �512 Table 6. Crown cover (0.01-m2), species composition (%), and frequency of occurrence of vegetation within treatment and control transects on Tamarack Prairie burn 3-84, July 1985. Treatment Vegetation Bare ground Dead vegetation Boute1ous gracilis Stipa comata Sporobo1us cryptandrus Ca1amovi1fa longifo1ia Andropogon ha11ii Paspa1um stramineum Muh1enbergia sp. Ko1eria cristata Bromus inernus Unid. ann. grass Carex & Cyperus spp. Artemisia fi1ifolia Opuntia sp . Ambrosia psi10stachya Artemisia 1udoviciana Tradescantia occidenta1is Phlox andico1a Evo1vu1us nutta1ianus Lathyrus po1ymorphus Penstomen angustifo1ius Cichorium intybus Physalis subg1abrata The1esperma megapotimicum Hap10pappus spinu10sus Mentze1ia nuda Oenothera Ipomoea 1eptophy11a Unident. perennial forbs Chenopodium album Croton texensis Plantago purshii Eriogonum annum Pepidium densif10rum Cryptantha sp. Lesquere11a 1udoviciana sp:-- Crown cover Compo Control Freq.! occur. 1,082.0 318.5 73.0 123.5 110.5 179.0 9.5 9.0 1.0 0.5 1.0 8.0 27.0 408.5 15.5 0.5 5.5 33.0 4.0 13.5 29.0 1.5 6.09 10.30 9.21 14.92 0.79 0.75 0.08 0.04 0.08 0.67 2.25 34.06 1.29 0.04 0.46 2.76 0.33 1.13 2.42 0.13 0.90 0.95 0.95 1.00 0.25 0.30 0.10 0.05 0.10 0.45 0.85 1.00 0.65 0.05 0.10 0.85 0.20 0.35 0.25 0.10 1.0 2.0 6.0 0.08 0.17 0.50 0.05 0.20 0.35 2.0 6.5 12.5 86.5 1.0 7.0 0.17 0.54 1.04 7.21 0.08 0.58 0.15 0.10 0.45 1.00 0.10 0.50 5.0 17.0 0.42 1.42 0.30 0.60 Crown cover Compo Freq.! occur. 569.5 958.5 53.5 107.5 87.0 128.5 11.5 5.5 2.0 4.99 10.03 8.12 11.99 1.07 0.51 0.19 0.75 1.00 1.00 1.00 0.30 0.30 0.10 26.5 520.5 9.5 1.5 6.0 20.0 3.0 3.5 28.0 1.0 1.0 2.47 48.55 0.89 0.14 0.56 1.87 0.28 0.33 2.61 0.09 0.09 0.70 1.00 0.30 0.10 0.10 0.60 0.10 0.15 0.25 0.05 0.10 2.0 0.19 0.05 9.5 0.89 0.20 7.0 6.0 15.0 1.0 5.0 0.5 6.0 3.5 0.5 0.65 0.56 1.40 0.09 0.47 0.05 0.56 0.33 0.05 0.05 0.35 0.95 0.10 0.30 0.05 0.30 0.10 0.05 �Table 7. Mean crown cover (0.01-m2) and analysis of covariance relationships for selected species, species groups, and covers within burn and control samples during pre-treatment (1984) and post-treatment (1984-85) intervals, burn 1-84, Tamarack Prairie. Vegetation Boute10ua gracilis Stipa comata Sporobo1us sp. Ca1amovi1fa sp. Andropogon ha11ii Comb. warm seasona Artemisia fi1ifo1ia Perennial forbs Annual forbs Bare ground Dead vegetation Control transects Aug Pre-tr Ju1 1984 1984 1985 Burned transects Aug Ju1 Pre-tr 1984 1984 1985 362.4 . 258.2 44.9 185.7 28.7 213.7 102.9 16.3 3.1 117.3 124.8 alnc1udes Ca1amovi1fa bDenotes P < 0.05. cDenotes P > 0.05. 310.2 86.5 75.4 199.3 47.7 245.8 75.8 40.8 4.1 386.6 7.5 116.5 103.3 58.5 73.3 16.5 89.9 84.0 12.5 15.9 458.5 301.9 10ngifolia, Andropogon 510.3 187.0 34.4 194.8 21.8 216.9 63.6 27.6 1.6 104.4 91.8 325.1 183.2 36.9 191.5 29.2 221. 7 67.2 28.8 50.6 91.3 233.9 92.6 125.7 18.4 95.6 9.7 105.8 49.5 15.9 2.4 231.4 601.4 Pre-tr to 85 .I-value Aug-84 to 85 14.89b 19.26b 9.82b 5.75c 1.54c 3.16c O.Oc O.llc 17.22b 166.6lb 87.05b 4.24c 3.37c 15.56b 7.21b 0.02c 5.55c 14.10b 5.99c 7.92b 0.82c 1.54c hal1ii, Panicum virgatum, and Paspa1um stramineum. VI I-' W �V1 i-' ~ Table 8. Mean crown cover (0.01-m2) and analysis of covariance relationships for selected species, species groups, and covers within burn and control samples during pre-treatment (1984) and post-treatment (1984-85) intervals, burn 3-84, Tamarack Prairie. Vegetation Boute1oua gracilis Stipa comata Sporobo1us sp. Ca1amovilfa sp. Andropogon ha11ii Comb. warm seasona Artemisia fi1ifo1ia Perenia1 forbs Annual forbs Bare ground Dead vegetation Control transects Pre-tr Aug Ju1 1984 1984 1985 Burned transects Aug Pre-tr Ju1 1984 1984 1985 166.4 132.8 110.0 97.6 17.8 128.0 242.4 103.3 2.6 202.9 121.6 alnc1udes Ca1amovi1fa bDenotes P < O. OS. cDenotes P > 0.05. 87.5 23.8 69.4 68.3 9.1 89.4 116.6 85.5 14.3 706.3 3L4 35.1 59.4 53.1 86.1 4.6 95.0 195.9 50.2 63.0 520.2 153.1 longifo1ia, Andropogon 131.3 169.9 130.4 120.8 15.6 155.7 224.4 6L4 L6 201.2 14L9 93.6 87.1 7L1 72.0 9.4 90.6 293.4 57.S 9.9 189.3 337.0 25.7 5L7 4L8 6L8 5.5 70.0 200.0 39.6 17.8 273.8 460.8 Pre-tr to 85 F-va1ue Aug-84 to 85 0.43c 0.95c o.zi= 4.1Sc 0.40c 3.18c 0.07c 0.24c 18.00b 94.49b 3L81b ha11ii, Panicum virgatum, and Paspa1um stramineum. 3.23c L53c 0.30c 3.S5c 0.07c 1.95c 12.10b 0.02c 16.78b L18c O.OOc �515 Table 9. Crown cover (point frame), species composition (%), and frequency of occurrence of vegetation within treatment and control transects on the Tamarack Prairie, 1985 burns, August 1985. Control Treatment Vegetation Bare ground Dead vegetation Boute1oua gracilis Stipa comata Sporobo1us cryptandrus Ca1amovi1fa longifo1ia Andropogon ha11ii Paspa1um stramineum Agropyron smithii Aristida sp. Panicum virgatum Muh1enbergia pungens Panicum capi11are Bronums tectorum Erogrostis ci1ianensis Unident. annual grass Cyperus & Carex spp. Artemisia fi1ifo1ia Opuntia sp. Ambrosia psi10stachya Artemisia 1udoviciana Tradescantia occidenta1is Phlox andico1a Evo1vu1us nutta1ianus Lathyrus po1ymorphus Psora1ea tenuif10ra Physalis subg1abrata The1esperma megapotimicum Mentze1ia nuda Lygodesmia~cea Sphaera1cea coccinea Croton texensis Chenopodium album Euphorbia sp. Pepidium densif10rum Tragopogan sp. Sa1so1i kali Amaranthus sp. Cirsium sp. Conyza canadensis Unidentified Crown cover Compo Freq./ occur. Crown cover Compo Freq./ occur. 13.14 33.05 13.60 17.09 2.47 0.04 2.11 0.11 0.04 0.14 0.18 0.04 0.04 1.00 1.00 1.00 0.94 0.47 0.06 0.71 0.18 0.06 0.12 0.06 0.06 0.06 0.46 13.53 0.63 0.07 0.29 0.82 0.35 0.06 1 11 3 0.46 0.04 0.39 0.11 0.29 0.06 0.24 0.12 3 0.11 0.12 3 0.11 0.42 0.56 0.42 0.04 0.04 0.04 0.32 0.11 0.04 0.11 0.12 0.18 0.24 0.06 0.06 0.06 0.06 0.06 0.12 0.06 0.18 1,875 2,891 360 471 358 926 87 13.96 18.27 13.89 35.92 3.37 0.82 1.00 1.00 1.00 0.59 801 3,705 373 938 386 485 70 30 1.16 0.59 60 37 1.44 0.12 9 0.35 0.47 0.12 0.06 11 1 0.23 0.62 5.39 0.89 0.43 0.04 0.06 0.29 0.65 0.35 0.18 0.06 1 11 0.04 0.43 0.06 0.29 17 8 1 5 2 0.66 0.31 0.04 0.19 0.08 0.35 0.06 0.06 0.12 0.06 2 0.08 0.12 16 0.62 0.29 1 12 6 16 139 23 3 1 4 5 1 1 13 384 18 2 13 28 1 1.09 0.04 0.06 0.06 12 16 12 1 1 1 9 3 1 3 �516 Table 10. Crown cover (point frame), species composition (%), and frequency of occurrence of vegetation within treatment and control transects on the Tamarack Prairie proposed 1986 burn sites, pre-treatment, August 1985. Control Treatment Vegetation Bare ground Dead vegetation Bouteloua gracilis Stipa comata Sporobolus cryptandrus Ca1amovilfa longifolia Andropogon hallii Agropyron smithii Aristida sp. Muhlenbergia pungens Panicum capi11are Bromus tectorum Unident. ann. grass Cyperus & Carex spp. Artemisia fi1ifolia Opuntia sp , Ambrosia psilostachya Artemisia ludoviciana Tradescantia occidentalis Philox andicola Evolvulus nuttalianus Lathyrus polymorphus Psora1ea tenuiflora Physalis subglabrata Thelesperma megapotimicum Ipomoea 1eptophy1la Erigeron sp. Sphaera1cea coccinea Hap10pappus spinu10sus Croton texensis Chenopodium album Eriogonum annum Crypthantha,sp. Plantago purshia He1ianthus sp. Lesquere11a ludoviciana Cirsium sp. Unidentified Crown cover 781 4,587 686 759 260 480 45 81 11 12 20 4 18 308 37 21 31 4 12 2 2 Compo Freq.! occur. 24.15 26.73 9.15 16.90 1.58 2.85 0.39 0.42 1.00 1.00 0.84 0.95 0.26 0.53 0.16 0.05 0.70 0.14 0.63 10.85 1.30 0.74 1.09 0.14 0.05 0.05 0.21 0.74 0.37 0.26 0.16 0.16 0.42 0.07 0.07 0.21 0.05 0.05 6 0.21 0.21 9 8 5 0.32 0.07 0.28 0.18 0.16 0.05 0.16 0.26 1 1 1 1 2 11 0.04 0.04 0.04 0.04 0.07 0.39 0.05 0.05 0.05 0.05 0.05 0.16 2 Crown cover Compo Freq.! occur. 20.96 31.51 8.95 20.48 0.64 1.08 0.06 0.16. 0.03 0.95 1.00 1.00 1.00 0.32 0.42 0.05 0.05 0.05 9 6 3 0.76 11.69 1.15 0.22 0.48 0.25 0.29 0.19 0.10 0.21 0.74 0.63 0.11 0.16 0.26 0.11 0.16 0.11 2 2 2 1 5 0.06 0.06 0.06 0.03 0.16 0.11 0.05 0.05 0.05 0.11 2 13 0.06 0.41 0.06 0.06 0.11 0.37 0.05 0.05 619 4,306 658 989 281 643 20 34 2 5 1 24 367 36 7 15 8 2 2 �517 Table 11. Mean crown cover (point frame ta11iesltransect) and analysis of covariance relationships for selected species, species groups, and covers within burned and controls amp1es during pre-tream~nt (1984) and post-treatment (1985) intervals on combined (burn 1, 2, & 3-1985) sites, Tamarack Prairie. Vegetation Boute10ua gracilis Stipa comata Sporobo1us cryptandrus Ca1amovi1fa 10ngifo1ia Andropogon ha11ii Combined warm-seasona Artemisia fi1ifo1ia Perennial forbs Annual forbs Bare ground Dead vegetation Treatment Pre-tr Post-tr 1984 1985 Control Pre-tr Post-tr 1984 1985 28.5 31.4 4.8 36.8 1.7 39.4 22.5 4.4 4.1 30.9 260.1 30.8 31.3 8.1 33.1 2.1 35.9 24.7 2.9 3.6 42.2 249.3 21.2 27.7 21.1 54.5 5.1 61.8 8.2 3.3 2.8 110.3 170.1 F- Degrees freedom 21.9 55.2 22.7 28.5 4.1 32.8 22.6 2.1 3.5 47.1 217.9 value O.llc 15.99b 0.46c 17.11b 0.68c 19.02b 30.01b 0.18c 0.10c 92.20b 32.35b 1,31 1,31 1,31 1,31 1,15 1,31 1,31 1,26 1,31 1,31 1,31 alnc1udes Ca1amovi1fa 10ngifolia, Andr02ogon ha11ii, Panicum virgatum, and Pas2a1um stramineum. bDenotes P < 0.05. cDenotes Ii >0.05. Table 12. comprising Composition during pre- and post-treatment intervals >1% of the total on burn 1-84, Tamarack Prairie. Vegetation Boute10ua gracilis Sti2a comata Sporobo1us cryptandrus Ca10movi1fa 10ngifo1ia Andropogon ha11ii Artemisia fi1ifo1ia Ambrosia psi10stachya Tradescantia occidenta11is Chenopodium album Pre-tr. 1984 37.0 25.6 4.5 18.4 2.8 10.2 0.8 0.2 Treatment Aug 1984 36.2 10.1 8.8 23.3 5.6 8.9 1.0 2.1 tr. Ju1 1985 Pre-tr. 1984 23.8 21.1 12.0 15.0 3.4 17.2 0.2 1.2 2.0 48.4 17.7 3.3 18.5 2.1 6.0 1.1 0.6 tr. for Control Aug 1984 35.2 19.8 4.0 20.7 3.2 7.3 0.8 1.0 4.4 species July 1985 22.2 30.1 4.4 22.9 2.3 11.9 0.3 1.5 0.1 �518 Table 13. comprising Composition during pre- and post-treatment intervals for species of the total on burn 3-84, Tamarack Prairie. > 1% Vegetation Boute1oua gracilis Stipa comata Sporobolus cryptandrus Calamovilfa longifolia Andropogon hallii Paspa1um stramineum Panicum virgatum Muhlenbergia pungens Cyperus & Carex sp. Artemisia filifo1ia Opuntia sp. Ambrosia psilostachla Tradescantia occidentalis Lathlrus pollmorphus Evo1vulus nuttalianus Ipomoea 1eptophllla Chenopodium album Croton texensis Crlptantha sp. Pre-tr. 1984 18.0 14.3 11.9 10.6 1.9 1.4 1.1 0.5 0.3 29.2 2.2 8.7 0.3 1.3 tr. tr. Treatment Aug 1984 17.1 4.6 13.5 13.3 1.8 2.3 tr. tr. 3.5 22.8 1.5 4.2 2.8 3.2 2.2 2.6 tr. 1.7 Ju1 1985 Pre-tr. 1984 6.1 10.3 9.2 14.9 0.8 0.8 14.5 18.7 14.4 13.3 1.6 1.5 0.7 1.1 1.3 24.8 1.3 5.4 0.3 2.3 34.1 1.3 tr. 2.8 2.4 1.1 0.5 7.2 0.6 1.4 0.6 tr. Control Aug 1984 12.9 12.0 9.8 10.0 1.3 1.2 0.1 0.7 2.8 40.5 1.3 0.9 1.5 0.9 0.2 0.9 0.9 0.3 tr. July 1985 5.0 10.0 8.2 12.0 1.1 0.5 0.2 2.5 48.6 0.9 0.1 1.9 2.6 0.3 0.7 1.4 0.1 from summer 1984 to summer 1985 in both 1984 burns but in neither case was the increase pronounced (p > 0.05). Observations indicated seed head production was dramatically enhanced in 1985 within 1984 burns. Although some green-up of this species had occurred prior to 1984 burns, major growth and seed head production had occurred at the time of the phenologically late 1985 burns. Thus, it would be expected that the 1985 fires would have a dramatic first year impact. Sand Dropseed.--Sand dropseed appeared to be enhanced by fire in burn 1-84 from pre-treatment and August 1984 to 1985 (p <0.05) (Table 7). However, this was less evident in burns 3-84 (Table 8) and there was no evidence of enhancement from the 1985 burns (Table 11). The 1984 burns occurred prior to green-up of this species whereas some growth had started prior to the 1985 burns. Prairie Sandreed.--Comparisons from the 1984 burns are uncertain with little evidence of a trend. Growth in 1985 appeared reduced from that in 1984 in burn 1-84 (Table 7) (p <0.05) but this was not evident in burn 3-84 where the species is much less-abundant. Numerical data (p < 0.05, Table 11) confirm visual observations that prairie sandreed was - markedly favored by the phenologically late 1985 burns and made dramatic growth during the summer. Field notes on 21 and 30 May both record observations that prairie sandreed grew rapidly following the 1985 burns although it had attained about 30 cm of growth at the time of the 16 May fires (Table 2). �519 Sand Bluestem.--Samples were not adequate to allow detection of impacts of burning if they occurred either year. Data for sand bluestem, prairie sandreed, and small samples of switchgrass and sand paspalum were combined (Tables 7, 8) but no clear impact of the 1984 burns could be detected. Some enhancement of these combined species over sandreed alone was noted in the 1985 burns (Table 11). Visual inspections of stands of sand bluestem within burn 3-85 revealed dramatic seed head production following the 1985 fire but vegetative growth form remained relatively sparse. Seed head production with burn 1-84, where pronounced in 1984, was not evident in 1985. Sandsage. --Fires in 1984 were conducted earlier under much drier conditions, before major leafing had occurred, and were much hotter consuming nearly all of the upper portions of the sage than fires in 1985. Rapid regrowth was noted in 1984 which continued into 1985 so that by the end of the second growing season, sandsage had attained much of its original growth form. Pre-treatment to July 1985 comparisons for both 1984 burns did not reveal impact of the fire (p > 0.05, Tables 7, 8), whereas a marked change from August 1984 to July 1985-was noted on both burns (p < 0.05) confirming rapid recovery. The 1985 burns were suppressed by green vegetation, and many tops of sage plants were not completely consumed by the fires as in 1984. However, the cambium layer on nearly all above ground parts was killed forcing basal resprouting. Sandsage had attained more growth in 1985 and, therefore, was more severely impacted by the fire as evidenced by slower resprouting and regrowth. A marked reduction in crown cover following the 1985 fires was noted (p < 0.05). Prickly Pear Cactus (Opuntia sp.).--Quantitative data are inadequate for analyses however, prickly pear, like sandsage was impacted only temporarily by the fire and made rapid recovery. The hotter 1984 burns killed many above ground parts stimulating rapid regrowth. Many of the above ground parts of cactii in 1985 were not killed by fire and resprouting was less pronounced. Perennial Forbs .--In combination, perennial forbs tended to be less abundant on all treatments and controls in 1985 than in late summer 1984. Possibly the dry early summer weather conditions in 1985 were influential, but this is uncertain. There is little evidence that fires in either 1984 or 1985 enhanced or severely hindered this group as a whole (f > 0.05) • Grasshoppers, while plentiful in 1985, appeared much less influential on forbs in 1985 than in 1984, especially in burn 3-84 where their impact was substantial. Most of the perennial forbs appeared to be markedly fire tolerant. Spiderwort (Tradescantia occidentalis) was in to early bloom stages at the time of the 1985 fires but recovered quickly and made rapid regrowth after the fires. Sweet pea (Lathyrus polymorphus) was past major bloom at the time of burning but recovered and bloomed again. Perennial ragweed (Ambrosia psilostachya), one of the most abundant perennials present, did not appear to be stimulated by burning either year although it had not made major growth prior to the fires. Annual Forbs.--This group increased in 1985 from pre-treatment and post-treatment 1984 averages in both 1984 burn sites (p <0.05, Tables 7, 8). Lambsquarter (Chenopodium album) was the dominant species. Pigweed (Amaranthus sp.) and milk purslane (Euphorbia sp.) were potentially enhanced by the 1985 burns whereas lambsquarter was suppressed by the late burn. More species diversity was noted within the controls than within the burned transects �520 (Table 9), although considerable variance was noted in species diversity among proposed 1986 burn sites (Table 10). Species numbers were about the same in 1985 for 1984 burns and their controls (Table 5, 6). Bare Ground and Dead Vegetation.--As would be expected, the fires dramatically reduced dead vegetation (both standing and litter) and increased bare ground from pre-treatment to subsequent intervals. There were no major changes from late summer 1984 to 1985 within sites burned in 1984. The cooler fires in 1985 left more unburned litter on the ground than was present after the 1984 burns. The 1984 fires were phenologically too early whereas those in 1985 were late when contrasted to optimum conditions. The optimum time frame for burning is when cool seasosn species have made considerable growth but before major growth of warm season species, and when there is substantial residual to carry a fire of at least moderate intensity. Although the 1985 fires were suppressed and cool due to the abundance of green vegetation, they appeared to enhance tall warm season species more than the 1984 burns. Precipitation amounts and patterns may have also influenced the results. June rains were much better in 1984 than in 1985 (Table 1), potentially benefiting cool season grasses more in 1984. In contrast, July through September precipitation was better in 1985 potentially enhancing warm season species to a greater extent. If fire is to markedly enhance the height-density quality of the Tamarack Prairie for prairie grouse, it must be through increased plant growth and vigor or through altered species composition toward tall, warm season grasses. The late burning in 1985 seemed to enhance prairie sandreed and suppress needle-and-thread but apparently had little impact on the other 2 dominant grasses, blue grama and sand dropseed. Suppression of these latter 2 species which comprise about one-third of the composition would be desirable in efforts to obtain taller more vigorous cover of increased height-density quality. Impacts of Fire on Interseeded Tall, Warm Season Grasses.--Fifteen metric belt transects were sampled in April 1984 in an interseeded portion of burn 1-84 which contained excellent stands of switchgrass and bluestem. An additional 15 transects were randomly positioned just outside the fire guard within the same interseeded tract. The site had been interseeded in spring 1982 during an interval of above average moisture and seeded grasses were well established. Post-treatment samples were obtained in early October 1985. Analysis of covariance revealed that switchgrass-bluestem crown cover was greater where impacted by the fire than in the controls from pre- to post treatment intervals (!1,27 = 7.27, P < 0.05). Interseeded grasses also comprised a higher proportion of the total grasses after the burn in the treated site than in the adjacent control (!1,27 = 6.39, R< 0.05). These data provide evidence that if more of these tall, warm-season grasses were present within the Tamarack Prairie, controlled burning might be more beneficial for increasing height-density quality and dominance by these species. Com arison Techni ues.--The metric belt system (Schmutz et al. 1982 was initiated at the start of the study on 1984 burns and its use was continued on these transects in 1985. Point frame sampling (Floyd and Anderson 1983) was initiated on sites to be burned in 1985 and 1986 and on spray-treated and revegetation transects. Transect 500-1600 within burn 1-84 �521 was sampled using both methods in August 1985 for comparison of results. The metric belt method tallied 18 species of vegetation plus bare ground and dead vegetation whereas the point frame method recorded hits on only 10 species plus bare ground and dead vegetation. Percentages of dominant grasses using the metric belt method were consistently lower than those obtained using point frame but proportional crown cover relationships among species tended to be similar. Blue grama was most common using techniques followed by about equal amounts of sandsage and needle-and-thread, and decreasing percentages of prairie sandreed, sand dropseed, and sand bluestem. Dead vegetation (including litter) was under estimated using the metric belt technique when compared to point frame sampling and bare ground was over estimated. Point frame sampling tends to be much less tedious than metric belt sampling because of the continuous ocular estimation and mental calculations needed for the latter. However, rapid sampling using point frame requires 2 persons to execute. Influence of Time of Burning on Sandsage.--The 4 May 1984 burns had little effect on sandsage survival (Snyder 1985) and regrowth was rapid. The question arose as to whether burning rangeland later in spring would increase mortality of sandsage. Therefore, sandsage plants were marked within burns conducted on 6 May and 16 May 1985. Lightning set one fire on the Tamarack Prairie, north of 1-76, on 24 June 1985 as well as several fires in adjacent rangelands to the south. Phenologically, the 6 May 1985 burn was 2-3 weeks later than fires conducted in 1984 and the 16 May burns were probably about a month later. Sandsage survival was reduced little if any by these phenologically later burns. Only 3 sage plants of 100 sampled showed retarded recovery from the 6 May 1985 burns were dead contrasted to 100% survival among unburned controls. Late August sampling of sandsage within the 24 June burn revealed that 11 of 50 plants had potentially been killed but some late budding was still occurring so the marking pins were left in place for additional evaluation in 1986. The apparent mortalities occurred where the fire had burned at a moderately hot rate under high afternoon temperatures. The fire started up again in evening when fanned by a moderate wind after a light rain had dampened the vegetation. The fire was cooler as the tops of the sage plants were not completely burned off. Preliminary tallies of 81 plants indicated only 2 dead plants. Regrowth of sandsage was notably retarded but the impact of the fire on other vegetation was also dramatic with little recovery in 1985. Available evidence indicates spring fires timed to enhance warm-season grasses generally will not reduce sandsage density but, if timed phenologically late, they will suppress rapid regrowth of sandsage. In sites dominated by dense sandsage, as in burn 3-84, its competition for available soil moisture seems to be a major hindrance to fire enhancement of grasses. Strip Spraying of Sandsage Aerial application of 2,4-D herbicide to sandsage was completed during the early morning of 6 June 1985. Winds, which preferrably should have been calm, were blowing from the south at approximately 8-13 km/hr (5-8 mph). The undulating or hilly terrain prevented the spray plane from remaining close to the ground in most locations, which in combination with the wind, caused �522 considerable herbicide drift. As a consequence, the herbicide was applied uniformly over the 80.8-ha area rather than in east to west strips as originally planned. No significant precipitation was receiyed for several weeks following the application. Nearly 100% of the sandsage and broad-leafed plants within the site were killed rather than the partial strip reduction of sandsage that was planned. Mortality of sandsage extended 100 m to the north of the tract and stunting of broad-leafed plants was noted 400 m or more to the north. Live sandsage or forbs were not found within the transects in late August. CayfLower (Liatris sp.), which apparently began growth after the spray treatment, was about the only forb surviving in late summer. Aerial application of herbicides should probably not be used on the Tamarack Prairie. Development of ground application equipment for wick application directly to sandsage while not killing understory forbs is recommended. Revegetation Treatments Ti11age-Reseeding.--Severa1 test plots, seeded or interseeded in spring 1984, were established earlier (Snyder 1985). Rototi11age reduced densities of perennial native grasses, but these remained dominant within atrazine-treated plots whereas they remained abundant and annual forbs dominated on plots not treated with herbicide. The atrazine treatment markedly reduced competitive annuals and enhanced establishment and growth of seeded warm season grasses in 1984 (Snyder 1985). Grasshoppers and dry weather in late summer 1984 severely impacted seeded stands on all plots. Dense stands of annuals, primarily 1ambsquarter, occurred on all untreated and interseeded plots in spring 1985 and moderately dense stands of annuals were present on the previous herbicide-treated plots. As a consequence, seeded perennials grew little in 1985 illustrating that atrazine, applied at a low rate, should be used prior to the second growing season. Because of the dominance of all plots by annuals and native grasses in 1985, vegetation sampling was discontinued. Approximately 20 ha (19 strips) were double disced, harrowed, and treated with 0.75 kg/ha (0.61 1bs/ac.) of atrazine in spring 1985 by management personnel. The strips were seeded to a tall, warm season grass mixture, dominated by switchgrass in early May. Precipitation, and cool damp conditions, conducive to grass germination and establishment, were marginal in 1985 but fair stands, dominated by switchgrass, resulted. Atrazine effectively reduced most competitive vegetation and mid- to late summer rains prompted excellent growth and limited first year seed head production. The herbicide had not been mixed thoroughly when applied to 2 strips where application rates were excessive and grass establishment did not occur. Discing and harrowing were much more effective than rototi11age (used the previous year) for destroying competitive native perennial grasses. Soil probe measurements indicated that reduction of competitive vegetation in the revegetation strips allowed subsoil moisture to accumulate enhancing establishment and growth of grass seedlings. The average probe penetration on 2 May 1985 was 7.8 dm (N = 7) in revegetation strips and 7.1 dm in adjacent grass (N = 5) (p > 0.05)-:- Depth of soil penetration increased to 10.7 dm in revegetation strips (N = 6) and 10.9 dm in burned sites (N = 26) on 17 May 1985 showing about equal moisture accumulations (p > 0.05). However, on 5 June average penetration was 15.0 dm in revegetation -strips (N = 20) compared to 11.9 dm (N = 20) in adjacent grass (f < 0.05) • - �523 Atrazine herbicide did not effectively curtail all competitive annuals. Late summer poLnt+f rame sampling at 12 random transects revealed that witchgrass (Panicum capillare) and Texas croton (Croton texensis) were 2 of the most common plants. However, lambsquarter and other weedy highly competitive annuals were uncommon to absent. Seeded switchgrass was the dominant perennial grass. Prairie sandreed and sand bluestem, which are deep-rooted, came back in vigorous stands where previously present (Table 14). Sand dropseed and blue grama were common. Needle-and-thread was almost completely eliminated by the tillage. Several species of native perennial forbs persisted in sparse stands. Table 14. Crown cover (point frame), composition (%), occurrence on 12 random transects within revegetation Prairie, 1985. Vegetation Bare ground Dead vegetation Panicum virgatum (seeded) Calamovilfa longifolia Andropogon hallii Sporobolus cryptandrus Bouteloua gracilis Agropyron smithii Cyperus & Carex spp. Panicum capillare Artemisia filifolia Opuntia sp, Psoralea tenuiflora Sphaeralcea coccinea Physalis subglabrata Ipomoea leptophylla Oenothera sp. Cichorium intybus Evolvulus nuttalianus Croton texensis Crown cover 804 212 303 66 8 126 25 4 11 64 5 2 1 2 8 3 1 6 1 76 and frequency of strips, Tamarack Composition Frequ. of occur. 42.6 9.3 1.1 17.7 3.5 0.6 1.5 9.0 0.7 0.3 0.1 0.3 1.1 0.4 0.1 0.8 0.1 10.7 1.000 0.583 0.167 0.833 0.750 0.250 0.083 0.500 0.083 0.167 0.083 0.083 0.083 0.083 0.083 0.083 0.083 0.250 Height-density comparison sampling was not conducted, however, observations in November 1985 revealed that the new switchgrass stands provided better height-density cover than adjacent native grass stands. These revegetation strips should continue to improve for the next few years. However, a second low-rate herbicide application is considered essential for rapid continued growth of the seeded grasses. Tillage Renovation of Interseeded Tracts.--Approximately 120 ha of the Tamarack Prairie were interseeded to tall, warm-season grasses in spring 1981 and 1982. Success in stand establishment was highly variable although precipitation amounts and patterns were excellent for stand establishment both years. Between-row native vegetation has remained highly competitive suppressing growth of the seeded grasses. �524 Observations in 1984 revealed that discing of fireguards across inter seeded tracts did not eradicate deep-rooted grasses such as switchgrass, b1uestem, and prairie sandreed. Instead, much of the competitive shallow-rooted vegetation was removed permitting vigorous growth of the deep-rooted tall grasses that attained heights of 1 m or more. In further evaluation, M. Gardner shallowly disced 2 narrow strips in March 1985 within 2 interseeded tracts within the Tamarack Prairie. One strip in each was treated with 0.75 kg/ha of atrazine in late April. Preliminary evaluation on 3 October 1985 showed that discing plus atrazine resulted in a mean height of 7.67 dm compared to 6.77 dm for discing alone and 3.43 dm for adjacent untreated interseeded grass. Discing plus atrazine yielded greater height (p < 0.05) than discing alone, and the latter yielded greater height (P < 0.(5) than untreated interseeded grass. Where interseeded grasses exist in satisfactory stands, as they do in about 50% of the interseeded tracts, discing plus atrazine is the most rapid cost-effective method available for dramatically increasing the height-density quality of existing cover. In addition, the treatment will smooth the furrows created by interseeding. Wildlife Use of the Tamarack Prairie Although prairie-chickens were released on the Tamarack Prairie during 1984 and 1985, none was observed during numerous days of activity in 1985. One feather, believed from a prairie-chicken, was found in burn 2-85 in August. M. Gardner reported flushing 3 prairie-chickens in Novembernorth of 1-76 near the east end of the property. A cornfield, just east of the Tamarack, possibly was a factor in their presence. Lack of food within the Tamarack Prairie should not be discounted as a reason released birds have not remained there or established leks there. Ring-necked pheasants (Phasianus co1chicus) were not observed on the Tamarack Prairie in 1985. One mallard (Anas p1atyrhynchos) hen and nest were found within a clump of sand b1uestem in burn 1-84. The nest was apparently destroyed by a predator at a later date. Mourning doves (Zenaida macroura) were not especially abundant in 1985 although several small migrating flocks were observed feeding in the revegetation strips in September. One or 2 pairs of upland plovers (Bartramia longicauda) were observed in the 1985 burns in June indicating possible nesting activity there. Three to 6 pronghorn (Anti1ocapra americana) were observed consistently within 1985 burns and revegetation strips from late June through August. Their pursuit by an archery hunter in August potentially prompted their departure. Mule deer and/or white-tailed deer (Odocoi1eus hemionus and Q. virginianus) were intermittant to almost continuous occupants of the Tamarack Prairie. One or 2 black-tailed jack rabbits (lePus) ca1ifornicus) were observed but no desert cottontails (Sy1vi1agus audubonii were noted. �525 LITERATURE CITED Bragg, T. B. 1978. Effects of burning, cattle grazing, and topography on vegetation of a choppy sands range site in the Nebraska sandhi11s prairie. Proc. Int. Rangeland Cong., Soc. Range Manage. 1:248-253. Engle, D. M., and P. M. Bu1tsma. 1984. Burning of northern mixed prairie during drought. J. Range Manage. 37:398-401. Floyd, D. A., and J. E. Anderson. 1983. A new point intercept frame for estimating cover of vegetation. Pages 107-113 in Idaho Nat!. Eng. Lab. Radioecology and Ecology Programs. U.S. Dep. Energy DOE/ID 12098. Schmutz, E. M., M. E. Reese, B. N. Freeman, and L. C. Weaver. 1982. Metric belt transect system for measuring cover, composition, and production of plants. Rangelands 4:162-164. Snyder, W. D. Wildlife. 1985. Sandsage-b1uestem prairie renovation. Fed. Aid Proj. Rep. 01-00-045. Apr. Colorado Div. Westemeier, R. L. 1972. Prescribed burning in grassland management for Proc. Tall Timbers Fire Eco1. prairie chickens in Illinois. 12:317-338. Prepare by 1IJtJJr!lu) ~~ .IPJ Warren D. Snyder Wildlife Researcher C Conf. ��Colorado Division of Wildlife Wildlife Research Report April 1986 527 JOB PROGRESS REPORT State of Colorado Project 01-00-045 (W-37-R) Work Plan Job Title: 21: Job Avian Research 4 Response of Selected Wildlife Species to Aspen Silvicultural Practices Period Covered: 01 July 1984 through 30 June 1985 Author: R. W. Hoffman and T. E. Remington Personnel: C. E. Braun, R. W. Hoffman, and T. E. Remington, Colorado Division of Wildlife ABSTRACT Literature review and communications with agencies interested in the quaking aspen (Populus tremu10ides) type were continued in 1984-85 as were efforts to identify research needs concerning aspen-wildlife relationships that could be addressed by the Colorado Division of Wildlife. A pilot study was conducted to measure consumption by blue grouse (Dendragapus obscurus) of game bird chow treated with extracts of aspen vegetative buds, and extracts of bud scales and catkins from male flower buds of aspen. All extracts inhibited feeding; however, when the choice was food treated with bud scale vs. catkin extract, blue grouse ate significantly more (P = 0.03) catkin extract treated food. This may explain why blue grouse do not feed on aspen buds until the catkins have emerged. When the choice was flower vs. vegetative bud extract treated food, blue grouse ate more (P = 0.003) food treated with extract from flower buds. It is suggested that blue grouse and other wildlife species may feed selectively among available aspen clones due to interclona1 chemical variations. ��529 RESPONSE OF SELECTED WILDLIFE SPECIES TO ASPEN SILVICULTURAL PRACTICES Richard W. Hoffman and Thomas E. Remington Quaking aspen dominated plant communities comprise nearly 2.3 million acres of commercial forest lands in Colorado (Green and Setzer 1974). Additional acreage is occupied by non-commercial stands and mixtures of aspen and conifer species. Together these types offer important water, forage, wildlife, and recreational values. Aspen typically occurs in clones produced asexually by adventitious sprouting from a single parent root system (Schier 1981). Seedling establishment is rare due to short-lived seed and demanding seedbed requirements (Maini 1968, McDonough 1979). Sprouting can be stimulated by destroying the extisting stand (Crouch 1981, Schier 1981). For centuries, fire was the natural regenerative force responsible for perpetuation of aspen (Gruell and Loope 1974). Twentieth century forest management policies have emphasized fire suppression. The result is that most aspen stands in the Central Rockies have reached the mature to overmature categories and are in need of treatment if they are to be maintained. In the absence of fire, clear-cutting is the most effective regenerative procedure (Crouch 1981, Jones 1975, Schier and Smith 1979). Until recently, there has been little commercial demand 1n Colorado for aspen for manufacturing wood products; consequently, mass treatments (clear-cuts) within the aspen type have been virtually non-existent. This situation is destined to change with the announcement by the Louisiana Pacific Corporation (LP) of plans to process aspen chips into waferboard, a compound type panel that LP feels may eventually replace conventional plywood. LP's plans call for the construction of 2 mills, 1 near Montrose and another in Kremmling, and the harvest of 2,500 acres of aspen/year/mill to maintain an annual production of 25 million board-feet of waferboard. The realization that large scale aspen manipulation programs may soon become common practice in Colorado has generated a pressing need for information on the impacts these treatments will have upon other associated resources. Aspen-wildlife relationships are of particular interest because little quantitative information exists. Aspen has long been recognized for its cover and forage benefits to wildlife, but this understanding has not progressed beyond the descriptive stage. Aspen will be managed for its occur with or without input managers are interested in, and in developing aspen management requirements. timber producing values and manipulation will from wildlife managers. Ironically, forest have requested input from wildlife specialists strategies that are consistent with wildlife �530 P. N. OBJECTIVES The objectives of this initial planning study are to: 1. Examine and evaluate existing literature on aspen-wildlife relationships to identify information needs pertinent to the development of a detailed study plan. 2. Interview selected personnel of the CDOW, USFS, BLM, and Louisiana Pacific Corporation concerning research needs within the aspen type. 3. Prioritize research needs and identify funding sources. 4. Prepare a detailed study plan on an approved research topic dealing with aspen-wildlife relationships. SEGMENT OBJECTIVES 1. Review literature pertaining to aspen ecology, aspen silvicultural practices, aspen-wildlife relationships, effects of aspen manipulation on wildlife populations, and chemical and nutritional characteristics of aspen. 2. Interview selected personnel of the CDOW, USFS, BLM, Colorado State Forest Service, and Louisiana Pacific Corporation concerning aspen-wildlife research needs. 3. Interview selected personnel involved in aspen research and management in other states to identify potential problems and assist in the selection of an appropriate research topic. 4. Visit proposed aspen timber sales suitable for conducting research. 5. Prepare a detailed study plan on an approved research topic and select study areas to meet research needs. 6. Obtain funding to support the research project. in Colorado that are potentially RESULTS AND DISCUSSION Cooperation from the U.S. Forest Service and Louisiana Pacific (LP) , and control over the treatments to be imposed were prerequisites to the further development of a study plan. No commitments were made due to the political and environmental issues surrounding the proposed treatment of aspen in Colorado. The future of LP in Colorado is uncertain. As a result, the decision was made not to proceed with an aspen study in 1984-85, but to continue open communications with other agencies interested in the aspen type and to pursue other avenues of aspen-wildlife research that could be addressed by the CDOW. �531 A community study in the aspen type is presently not feasible within the CDOW Terrestrial research group and it is questionable whether such a study would provide meaningful information for making management decisions. A study designed to measure changes in abundance and distribution of wildlife species in response to aspen manipulation is premature and should not be implemented until political and environmental issues are settled. Such a study cannot" proceed without complete control over the treatments to be imposed. Studies of how and why selected wildlife species use the aspen type have important management implications, fall within the research capabilities of the CDOW, and can be conducted independent of other agencies. By understanding not only what species use the aspen type, but why and how they use it, wildlife managers will be better equipped to predict and mitigate the impacts of aspen manipulation on wildlife populations. This information is also prerequisite to designing studies to measure wildlife responses to aspen manipulation. Aspen-Blue Grouse Relationships A pilot study was conducted to evaluate the possibility that there are compounds present within buds and/or bud scales of aspen which inhibit feeding by blue grouse. This chemical inhibition theory has been advanced to explain ruffed grouse (Bonasa umbellus) feeding behavior within aspen (G. W. Gullion, pers. commun.). In areas where aspen occurs, most blue grouse territories are associated with aspen clones (Hoffman 1981). Blue grouse have been observed feeding on buds and catkins of aspen in spring, even where conifers are present (pers. observ., R. W. Hoffman and T. E. Remington). The relative inhibitory effects of vegetative buds, male flower buds, and bud scales were evaluated by treating a pelleted ration with diethyl ether extracts of these plant materials and measuring blue grouse consumption relative to controls. This was a first step in identifying how blue grouse use the aspen resource. We hypothesized that blue grouse feed only on trees bearing male flower buds and only after catkin emergence when the catkins can be gleaned away from the bud scales. Male flower buds (with emergent catkins) and vegetative buds of aspen were collected on Whiteley Peak in Middle Park, Grand County, Colorado in April. The bud scales and catkins were separated and dried (as were vegetative buds) at 100 C (18 hrs). The dried material was ground in a Wiley mill and 38 g were extracted in a Soxhlet apparatus for 12 hours with diethyl ether as solvent. The extract was concentrated by roto-evaporation and the final volume brought up to 100 mls. The extract was poured over 120 g of Purina game bird chow and the ether was allowed to evaporate. Controls were prepared by pouring 100 mls of ether over 120 g of Purina game bird chow and allowing the ether to evaporate. Four captive blue grouse were used as experimental subjects. These birds had been captured 3 months earlier and maintained on conifer needles and game bird chow. The birds had been on game bird chow for 29 days before the trials. Each of the extracts was tested against a control and then against each other. Twenty-five grams of each treatment pair were placed in beakers. The beakers were then randomly positioned within each cage and left there for 3 hours. Each bud was tested twice daily. Treatment response was the amount of game bird chow consumed. �532 There appeared to be inhibitory compounds present in bud scales, catkins, and vegetative buds since consumption of game bird chow treated with ether extracts from these plant materials was less (P < 0.05) than consumption of game bird chow treated with ether only (contrOls) (Table 1). The level of inhibitory compounds appeared to be highest in vegetative buds and higher in bud scales than catkins based on relative consumption in vegetative bud vs , flower bud and catkin vs , bud scale paired comparisons (Table 1). This finding is also supported by the relative levels of dry matter extracted by ether from bud scales, catkins, and vegetative buds (Table 2). Ether extracts have been used as an index to phenolic resin levels in plant material (Bryant and Kuropat 1980). Phenolic resins are a potent class of feeding and digestibility inhibitors (Bryant and Kuropat 1980). Table 1. Consumption (x gms eaten) by 4 blue grouse of game bird chow treated with diethyl ether (control) and diethyl ether extracts of bud scales and catkins from male flower buds of aspen. Extract pair P N Bud scale 1.31 Control 9.58 12 0.001 Catkin 2.43 Control 7.50 11 0.006 Bud scale 2.53 Catkin 8.07 10 0.032 Flower bud 4.17 Vegetative bud 0.96 12 0.034 Table 2. Diethyl ether extract of plant parts from male aspen trees. Plant part Catkin bud scales Catkins Vegetative buds % DM 9.01 5.53 12.00 These results are consistent with the theory that there are feeding inhibitors in aspen bud scales that prevent blue grouse from feeding on buds until catkins emerge in spring. Future research should concentrate on attempting to isolate and identify compounds involved; investigate their physiological effects on blue grouse; and measure the extent and type of use of aspen for food (feeding on catkins, flower buds, or vegetative buds and extent of selectivity) and cover. �533 LITERATURE CITED Bryant, J. P., and D. J. Kuropat. 1980. Selection of winter forage by subarctic browsing vertebrates: the role of plant chemistry. Annu. Rev. Ecol. and Syst. 11:261-286. Crouch, G. L. 1981. Regeneration of aspen clearcuts in northwestern Colorado. u.s. Dep. Agric., For. Sere Res. Note RM-407. 5pp. Green, A. W., and T. S. Setzer. 1974. The Rocky Mountain timber situation, 1970. u.s. Dep. Agric., For. Servo Resour. Bull. Int-IO. 78pp. Gruell, G. E., and L. L. Loope. 1974. Relationships among aspen, fire, and ungulate browsing in Jackson Hole, Wyoming. U.S. Dep. Agric., For. Serv., Int-unpubl. rep. 33pp. Hoffman, R. W. 1981. Population dynamics and habitat relationships of blue grouse. Colorado Div. Wildl., Res. Rep., Fed. Aid Proj. W-37-R. Work Plan 9, Job 5. April 1981. pp. 103-172. Jones, J. R. 1975. Regeneration on an aspen clearcut in Arizona. Agric., For. Sere Res. Note RM-285. 8pp. U.S. Dep. Maini, J. S. 1968. Silvics and ecology of Populus in Canada. Pages 20-69 in J. S. Maini and J. H. Cayford, eds. Growth and utilization of poplars in Canada. Canada Dep. For. and Rural Dev. Publ. 1205. McDonough, W. T. 1979. Quaking aspen-seed germination and early seedling growth. U.S. Dep. Agric., For. Servo Res. Note Int-234. l3pp. Schier, G. A. 1981. Aspen regeneration. Pages 15-21 in N. V. DeByle, ed. Situation management of two intermountain species: aspen and coyotes. Part 1 - Aspen. Utah State Univ., Logan. , and A. D. Smith. 1979. Sucker regeneration in a Utah ----~~ following clearcutting, partial cutting, scarification, U.S. Dep. Agric., For. Servo Res. Note Int-253. 6pp. aspen clone and girdling. ��535 JOB PROGRESS REPORT Colorado State of 01-03-045 01-00-045 (W-37-R, W-88-R) Project Work Plan Job Title: 22: Job 1 Avian Research (6-1) Avian Research Publications Period Covered: 01 June 1984 through 31 December 1985 Author: Clait E. Braun Personnel: C. E. Braun, P. O. Dunn, R. W. Hoffman, J. Profera, T. E. Remington, J. K. Ringelman, W. D. Snyder, and M. R. Szymczak, Colorado Division of Wildlife. ABSTRACT Publications accomplished under this job in 1984-85 are: Braun, C. E. 1984. Attributes of a hunted sage grouse population in Colorado, U.S.A. Pages 148-162 in P. J. Hudson and T. W. I. Lovel, eds. 3rd Int. Grouse Symp., World Pheasant Assoc., York Univ., U.K. 1984. Biological investigations of white-tailed ptarmigan in Colorado, U.S.A. - A review. Pages 131-147 in P. J. Hudson and T. W. I. Lovel, eds. 3rd Int. Grouse Symp., World Pheasant Assoc., York Univ., U.K. , and --------regulations T. D. I. Beck. 1985. Effects of changes in hunting on sage grouse harvest and populations. Pages 335-343 in S. L. Beasom and S. F. Roberson, eds. Game harvest management. Proc.~rd Int. Symp., Caesar Kleberg Res. Inst., Kingsville, TX. Dunn, P.O., grouse. and C. E. Braun. Auk 102:621-627. 1985. Natal dispersal and lek fidelity of sage Hoffman, R. W. 1985. Blue grouse wing analyses: methodology and population inferences. Colorado Div. Wi1dl. Spec. Rep. 60. 21 pp. 1985. Effects of changes in hunting regulations on blue grouse populations. Pages 327-334 in S. L. Beasom and S. F. Roberson, eds. Game harvest management. Proc. 3rd Int. Symp., Caesar Kleberg Res. Inst., Kingsville, TX. Profera, J., and C. E. Braun. 1985. Sage grouse public viewing tour development in North Park, Colorado. J. Colo.-Wyo. Acad. Sci. 17(1):36. Remington, T. E., and C. E. Braun. 1985. Sage grouse food selection in winter, North Park, Colorado. J. Wildl. Manage. 49:1055-1061. �536 Ringelman, J. K., and M. R. Szymczak. 1985. A physiological for wintering mallards. J. Wildl. Manage. 49:564-568. condition index Snyder, W. D. 1984. Odd areas for wildlife. Pages 59-64B in Wild1. Resour. Comm., eds. Increasing wildlife on farms and ranches. Kansas State Univ. Coop. Ext. Servo and Great Plains Agric. Counc., Manhattan. 1984. Ring-necked pheasant nesting ecology and wheat farming on the High Plains. J. Wildl. Manage. 48:878-888. 1985. Management procedures for ring-necked pheasants in Colorado. Colorado. Div. Wildl. Spec. Rep. 59. 53 pp. 1985. Colorado. Prepared by Survival of radio-marked hen ring-necked pheasants in J. Wildl. Manage. 49:1044-1050. t/~ ~ Crait E. Braun Wildlife Research Leader � https://cpw.cvlcollections.org/files/original/91476127e9ad03fe36ec78a371326c3a.pdf 34a880ea4a91af50b45e1799c24850bd PDF Text Text 1 Colorado Division of Wildlife Wildlife Research Report January 1987· JOB PROGRESS REPORT Colorado State of Project (W-151- R-l) Per10d Covered: Author: Bald Eagle Nest Site Protection and Enhancement Program 1 July - 30 June 1987 G.R. Craig Personnel: G.R. Craig, Colorado Division of Wildlife and Richard Knight, Dept of Fishery and Wildlife Biology, Colorado state University. ABSTRACT Nine bald eagle breeding territories were occupied by 8 adult pairs in 1987. A total of 10 young were successfully fledged by 6 pairs. One pair failed when the nest was blown down and the other pair abandoned their egg prior to hatch. The egg that was abandoned was of normal shell thickness. Several sites are in need of stabilization. This Job Progress Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the author. ��3 BALD EAGLE NEST SITE PROTECTION AND ENHANCEMENT PROGRAM Gerald R. Craig SEGMENT OBJECTIVES 1. Annually visit all known bald eagle breeding territories throughout Colorado and observe them from a distance to establish the presence of breeding adults. Breeding pairs will be kept under surveillance to determine onset of incubation. Occupied sites will be monitored throughout the breeding season to document reproductive success. When a pair's behavior indicates that hatching has occurred, the nest will be visited to document the number of young produced and collect eggshell fragments or any nonviable eggs still present. A second visit will be made to band and color mark nestlings. The nest will be observed at time of fledging to determine actual fledging success. 2. When a pair is encountered that failed during incubation, attempts will be made to vis it the nest and determ ine the cause for fai1 ure. If any are present, eggshell fragments and nonviable eggs will be collected for shell thickness measurement and chemical analysis. 3. Confirmed and potential breeding sites will be surveyed annually to determine the presence of banded or color marked eagles. Band confirmation will be accomplished through observation from a distance with telescopes. 4. Documented nest sites will be mapped on topographic maps, land ownership of the nest site and important foraging areas will be determined, and vegitative communities will be mapped. 5. On occasion, nest sites may be jeopardized because they are placed in dead trees subject to wind throw. In those instances, the nest structures will be mechanically stabilized and artificial nest platforms may be placed in more substantial neighboring trees. 6. Initiate agreements with federal land management agencies and private landowners to protect and enhance nesting and foraging habitat. METHODS AND MATERIALS This work will be a cooperative endeavor between the Division and Dr. Richard Knight of Colorado State University. 1. Annually survey all documented breed ing sites to determ ine presence of bald eagles. Pairs at territories will be determined by DWMs and other field personnel. Previously undocumented pairs will probably be revealed in the course of aerial eagle and waterfowl flights. DWMs will confirm actual incubation from ground visits. 2. Occupied territories will be visited by DWMs periodically throughout the breeding season to determine hatch of young, nesting failures, etc. �4 3. In May and June, a Utility Worker will observe breeding eagles from a distance and endeavor to follow their movements to locate important foraging areas. Responses of eagles to various human activities and land uses also will be recorded. 4. In June, when the young are determ ined to be old enough to band, sites will be visited to place a federal band on one leg and a colored, alpha numeric marker on the other. The color markers will permit identification if the young return to Colorado to breed in subsequent years. During the same nest visit the following will be recorded: Physical parameters such as tree height, DBH, condition, and dominance. Nest condition, size, and location. Vegitative community and land use practices. In addition, collect prey remains, nonviable eggs and eggshell fragments. 5. When necessary, remedial actions will be taken to stabilize nests that may be subject to wind throw. Should the tree be decadent and in danger of falling, an artificial nest base may be placed in a suitable, adjoining tree. Action will be taken only after it has been deemed desireable to encourage the eagles to nest at the same location. RESULTS AND DISCUSSION Territory Occupancy Cumulative bald eagle nesting activities for Colorado are summarized in Table 1. The first nesting attempt since 1950 (Bailey and Neidrach 1965) was documented in La Plata County in 1974. The pair utilized an osprey nest and incubated one egg, but abandoned the effort. The number of breeding attempts gradually increased until 10 pairs were present in the state in 1986. In all, 13 breeding territories have been recorded since 1974. In 1987, 9 territories (Adams, Archuleta, La Plata #1 and #2, Moffat #1 and #2, Montezuma #2, and Rio Blanco #1 and #3) were occupied. The pair that had occupied the Weld County site did not return in 1987 and although it was suspected that the pair were in the vicinity, they could not be relocated despite several helicopter searches. All of the documented nests are associated with riparian systems. Six sites (Moffat #1, #2 and #3, Rio Blanco #1 and #2, and Weld) are located in riparian habitat adjacent to rivers. Four sites (La Plata #1, Adams, Grand and Montezuma #2) are on lake shores, and 2 (Archuleta and Montezuma #1) are in upland areas away from water. Both of the upland sites were within a mile or so of lakes. The Montezuma #1 site was situated in a large cottonwood tree immediatly adjacent to a golf course. Land Status Of the 13 sites on record, 7 sites (Archuleta, Moffat #1 and #2, Montezuma #2, Rio Blanco #1 ans #2, and Weld) are on private property, 3 sites (Grand, La �5 Plata #1, and Montezuma #2) are on Forest Service administered lands, 1 site (Moffat #3) is on BLM land, 1 nest is on Indian land, and 1 nest is situated in a State park. Reproduction Table 2 summarizes breeding effort observed in 1987. Adult pairs were present at 8 sites and a single adult was observed at the La Plata #2 nest. All 8 adult pairs laid eggs, and 7 pairs hatched young. The pair at the Adams County site failed when the male disappeared about 4 weeks into incubation and the female abandoned the egg approximately 5 days prior to hatch. The egg contained a full term embryo. The 7 pairs that hatched their eggs produced 12 young (1.71 young/successful pair, 1.5 young/total pair and 1.33young/occupied site). The young at Moffat County #2 died when the nest was blown down. The pair had experienced the same probl em in 1986 when they used an abandoned heron nest as the base for their nest. In 1987, they rebui lt their nest on another heron nest and experienced the same difficulty. In all, 10 young were successfully fledged by 6 pairs (1.67 young/successful pair, 1.25 young/adult pair and 1.11 young/occupied site). Culmen length, and foot pad length measeurements were take of 6 young at 3 sites (Archuleta, Moffat #1 and Rio Blanco #1) in an effort to determine sex according to criteria developed by Bortolotti (1984) and Garcelon et al (1985). Surprizingly, all fell into the male category. Since the criteria were developed from northgern birds, it is possible that the Colorado birds are smaller and even females would fall into the male category. The six young that were measured were banded with Fish and Wildlife Service bands and red bands with etched alpha numerics were placed upon the other leg. Eggshell Condition The only eggshell information in 1987 was obtained from the abandoned egg at the Adams County site. The shell thickness (with membrane) averaged 0.541mm at the equator which is 8.0 % thin when compared with pre DDT era eggs collected from Florida, Louisiana and Texas (Anderson and Hickey 1972). For all intents, the eggshell measurements fall within the random deviation experienced for normal eggshell thickness. Organochlorine Residues in Eggs Contents of the abandoned egg from the Adams County site were submitted chemical contaminant analysis and the results are not yet available. Nest Stabilization for Efforts In 1987, no actions were taken to stabilize nests although it was noted that at least 3 nests (Adams, Montezuma #2 and Moffat #1) were in jeopardy. The Adams County nest was placed on an old heron nest in a decadent, inundated cottonwood tree. While the nest is securely situated, the tree is dead and subject to wind throw. Failure of Moffat #2 has already been related. To avoid future problems, since the pair has demonstrated the predilection for using abandoned �6 heron nests, an artificial platform should be placed in a more substantial fork. The nest at Moffat #1 is situated on the top of the cottonwood that broke and wedged horizontally in an upper fork. The whole arrangement forms a teeter-totter that should be guyed to the main trunk. In addition to immediate jeopardy, the nest at Montezuma #2 is in a large, apparantly heart rotted ponderosa pine that is partially inundated. It is difficult to say how resistant to wind throw the tree may be. It may be appropriate to place an artificial platform in the vicinity to assure the pair has an alternate should the tree fai1. LITERATURE CITED Anderson, D.W. and J.J. Hickey. 1972. Eggshell changes in certain North American birds. Proc. XVth Intl. Ornithol. Congress 1972. pp514-540. Bailey, W.A. and R.J. Neidrach. 1965. Birds of Colorado (2 vo l.), Den. Mus. of Nat. Hist. 865p. Bortolottu, G.R. 1984. Criteria for determining age and sex of nestl ing Bald Eagles. J. Field Ornithol. 55:467-481. Garcelon, O.K., M.S. Martell, P.T. Redig, and L.C. Buoden. 1985. Morphometric, karyotypic, and laparoscopic techniques for determining sex in Bald Eagles. J. Wildl. Manage. 49(3):595-599. Prepared by.. /: /J () ". o. u . LJt~ Gerald -R. Craig (f.----Wildlife Researcher C �-...J �8 Table 2. Colorado Bald Eagle Nesting Efforts - 1987 Site Age of Birds Male Female Young Produced Young Fledged Moffat Co. #1 A A 2 2 i~offat Co. #2 A A 2 0 Rio Blanco Co. #1 A A 2 2 Rio Blanco Co. #3 A A 1 1 Montezuma Co. #2 A A 1 1 La Plata Co. #1 A A 2 2 A 0 0 La Plata Co. #2 Adams Co. A A 0 0 Archuleta Co. A A 2 2 12 10 TOTAL A = Adult Comments Killed, nest blown out Single adult present at nest 1 egg abandoned prior to hatch �9 BALD EAGLE NEST FORM Nest Letter: Date: --Territory No.: Site Name: . ~--~-----------------------------------------------------Location (legal) Field Contact (DOW and Land Mgmt Agency personnel): Name: Address: PROPERTY OWNER ------------------------- Phone: _ ------------------------------------------------------------- NEST SURROUNDINGS Distance (m) to Water: Distance to Road (main~t-al~·n-e~d~): Distance to Human Habitation: ------Major Vegitative Community: Species: Tree Cond ition: Tree Diameter Breast Height: Nes t Hei ght (above ground) : Nest Position in Tree: NEST AND NEST TREE ---------- Tree Height(m): . Tree Diameter Nest Height: _ _ ----------------------------------- Nest Dominance Status: Diameters and Condition of Supporting Limbs: ----------------------------- Outside Nest Height:-=--.,..--,;-~--Net D iamet er (t0 p}, 0utside :-:--_.,---,,...,--_,:---:--____ Ins ide : Largest Stick in Nest (length and diameter): Nest Material Description: Nest Exposure: Percent Canopy Closure: Distance to Other Nests-:-----Prey in Nest: Prey Below Nest: --PREY REMAINS _ ------------------------------------------------------ _ _ �10 BALD EAGLE NESTlING FORM Territory No. : Nest Location: Nest No. : Date: ----. YOUNG 1 YOUNG 2 YOUNG 3 Band No. (F&~JS) Leg IO Band No. & Color Leg Est. Age Weight (g) Length 8th Primary Bill Depth Culmen Length w. Cere Culmen Length wlo Cere Foot Pad Length Calculated Age Calculated Sex Comments on Condition of Young: ---- Presence and Activity of Adults: _ �11 Colorado Division of Wildlife Wildlife Research Report January 1987 JOB PROGRESS REPORT State of Colorado --~~~~------- Project __ Peregrine Falcon Restoration Program ._;(,_S_E-_l_l..1..) _ Period Covered: Author: 1 January - 31 December 1987 G.R. Craig Personnel: D. Berger and G.R. Craig, Colorado Enderson, The Colorado College. Division of Wildlife; J. ABSTRACT The number of occupied peregrine falcon breeding territories in Colorado continued to increase in 1987. Twenty-nine sites were occupied by 23 adult pairs, 23 pairs bred and 22 pairs successfully produced young. Five wild pairs were fostered to augment anticipated poor productivity. Eggshell condition continued to decline to -15.3% overall thinning. Two female exhibited extreme thinnning of -27.9%. Five hack sites produced 19 young. When wild production, fostering and hacking efforts are considered, 85 young achieved independence in 1987. Reoccupancy of East Slope sites continues to lag and must receive emphasis. This Job Progress Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the author. ��13 PEREGRINE FALCON RESTORATION PROGRAM Gerald R. Craig P. N. OBJECTIVE The objectives of this study are to annually monitor site occupancy, reproduction, pesiticide residues and eggshell thinning, and population turnover of breeding peregrines. Pairs experiencing poor reproduction will be augmented with captive hatched young, captive produced peregrines will be released into vacant territories, and recruitment into the wild breeding population will be monitored. SEGMENT OBJECTIVE 1. Annually monitor the number of breeding reproductive success in Colorado. pairs of peregrines 2. Annually monitor organochlorine grines in Colorado. levels in wild breeding 3. Monitor breeding population turnover through band recoveries, presence of color markers, and telephotographic identification of individual breeding adults. 4. Augment poor wild production by placement of captive hatched wild young and captive produced young into occupied wild nests. 5. Release captive hatched and captive produced young at potential and vacant wild territories. 6. Monitor recruitment of reintroduced population of Colorado. pesticide peregrines into the wild and their pere- breeding (These objectives correspond to Jobs 111., 12., 211., 212., 213., 214., 22., 3221., 3222., 323., 33., 41., and 42. of the approved American Peregrine Falcon Recovery Plan for the Rocky Mountain/Southwest population). METHODS AND MATERIALS Methods and materials and Enderson 1981). used in this study have been described previously (Craig RESULTS AND DISCUSSION Territory Occupancy In the 1987 breeding season, territory occupancy increased 38% from 21 territories the prior year to 29 territories (Table 1). The increase resulted from reoccupancy of 3 historic (pre 1973) territories (sites 1, 7 and 18) and discovery of pairs at 4 sites that had been previously surveyed but were �14 unoccupied at the time. Adult pairs were present at 24 sites, 3 pairs were comprised of immature females paired with adult males, and 2 territories were frequented by lone adults. The addition of 4 new sites increased the sites on record to 51. Of the 14 historic sites that were occupied in 1972-73, 57% (8 sites) have been reoccupied (Table 2). Unfortunately, the female that disappeared at site 8 in 1986 was not replaced, nor did the male return, so that site remained unoccupied. Loss of site 8, however, was offset by reoccupancy of adjacent site 7 by an adult pair that successfully fledged one young. Reoccupancy of East Slope sites continued to lag with presence of pairs at only 3 sites (site 1, 39, and 49), and lone adults at 2 others (site 18 and 33). Conditions on the West Slope continued to improve with 75% (24 of 32) occupan- cy. Due to the lack of interplay between East and West Slope sites, release efforts were continued east of the Continental Divide. Reproduction Breeding behavior (production of eggs) was exhibited by 23 of the adult pairs (Table 1), none of the pairs comprised of a subadult member were successful in producing eggs. The adult pair at site 48 were just discovered in 1987 and still may have been establishing a territory. Five of the breeding pairs (sites 1, 9, 27, 39 and 42) were manipulated to receive foster young. The remaining 18 wild pairs (sites 3, 4, 5, 7, 11, 16, 25, 29, 30, 31, 34, 35, 38, 40, 41, 43, 44, 45 and 49) were permitted to incubated and hatch their own eggs without intervention. One pair (site 41) continued for the fourth consecutive year to fail to hatch their eggs. While eggshell thinning is suspected, the eyrie ledge was inaccessible and shell fragments were not obtained. The 17 successful pairs produced at least 49 young (2.88 young/successful pair) of which 39 fledged (2.29 young/successful pair). Thus, a remarkable productivity of 1.77 young fledged per wild breeding pair was achieved in 1987. This is the third consecutive year that wild productivity exceeded the productivity of 1.25 which is generally considered necessary to maintain population stability. The 5 sites that were fostered (sites 1, 9, 27, 39 and 42) were manipulated for a variety of reasons. Site 1 was fostered in order to accelerate the reproductive process and thus avoid conflict with rock climbers since the eyrie ledge was adjacent to a popular climbing route. Fostering succeeded in providing young 2 weeks older than would have been produced naturally. Rock climbers were observed on the eyrie ledge a week after the young fledged. Pairs at sites 27 and 39 were fostered because of extreme eggshell thinning experienced by the females the previous year. Sites 9 and 42 were fostered to augment wild production at Dinosaur National Monument. A total of 17 eggs were removed (average clutch size of 3.4) of which 12 were successfully hatched at the Peregrine Fund's Boise facilities. In all, 17 captive hatched young were returned (brood size of 3.4 young /pair) of wh ich 16 fledged for a success of 3.2 young/pair. When the productivity of the wild pairs is combined with the productivity of fostered pairs, a total of 66 young were produced of which 55 fledged for an overall productivity of 2.04 young fledged per pair. Eggshell Condition Eggshells representing 35 eggs were collected in 1987 at 16 eyries (sites 1, 3, 15,25,27,29,30,31,34,35,40,42, and 44).· Complete first clutches 7,9, �15 were collected from five sites (1, 9,27,39 and 42) in the course of fostering activities. Whole, nonviable single eggs were collected from 4 eyries (3, 30, 35, and 40), and shell fragments represent i ng 14 add it i ona 1 eggs were encountered at 9 eyries (7, 16, 25, 29, 30, 31, 35, 40 and 44). In all cases, the fragments were from eggs that hatched young. Due to variability of thickness measurements taken from elsewhere on the egg, only measurements obtained from the waist of eggs were utilized for comparison. Thus, only 22 whole eggs originating from the above 9 eyries were suitable for shell thickness (with shell membrane) comparison. The 22 eggshell measurements averaged 0.304mm which is 15.3% thinner than pre DDT era eggs (Table 3). This average is the th i nnes t encountered since 1979. The average was reduced by several eggs that were among the thinnest eggs ever encountered in the wild in Colorado. An egg each from sites 30 and 39 measured -27.9% thin (0.259mm), and if fragments are at all representative, the second egg from site 30 was critically thin (-28.4%, 0.257mm) as were eggs at sites 31 (-22.6%, 0.278mm) and 7 (-27.9%, 0.259mm). Further cause for concern is that 7 eggs representing 4 females (sites 3, 30, 39, and 42) were thinned more than 18%, a level generally associated with declining population. In addition to reduced shell thickness, wide thickness variability was encountered within clutches. The new female breeding at site 1 produced eggs that varied from -5.6% (0.339mm) which is nearly normal, to -14.2% (0.308mm) thin. The pair as site 27 produced eggs ranging from normal (-4.7%, 0.342mm) to critical (-17.5%, 0.296mm). The great shell thickness variability within clutches was also observed at 4 sites (16, 30, 31, and 35) in 1986. Interestingly, the eggshell thinning does not appear to be correlated with pes t ic tde residues in the egg contents. Organochlorine Whole eggs colleceted organochlorine residue in 1986 analysis. Photographic and Residues 1987 in Eggs have Documentation not yet been submitted for of Wild Adults Due to time constraints, photographic identification of individual breeding adults was not as detailed as previous years. In 1987, useable photographs were obtained of 10 individuals at 7 territories. From photographs obtained of the male at site 27, it appears that he is the same male that was present at site 9 in 1983. Hacking Efforts Hack sites were activated at 5 locations along the East Slope in 1987 (Table 4). In all, 20 falcons were released from the hack sites and 19 (86%) survived to independence. In addition, a subadult female frequented one hack site and an adult pair interacted with young at another site. Return of Released Falcons In all, 31 of 47 falcons were optically checked for the presence of bands or color markers. Eighteen were unbanded and 13 were determined to have been released either through fostering or hacking. Hence 42% of the falcons that �16 were examined originated from release efforts. LITERATURE CITED Craig, G.R. and J.H. Enderson. 1981. Nesting performance of peregrine falcons in Colorado. Job Prog. Rep., Colo. Div. Wildl. Res. Rep., Jan., pp 13-23. Prepared by: C. R~ Gerald R. cralg Wildlife Research r C �17 Table 1. Site Male Age Female 1 3 4 5 7 9 11 16 25 27 29 30 31 33 34 35 36 38 39 40 41 42 43 44 45 46 47 48 49 A A A A A A A A A A A A A A A I I A A SL* A A Total A A A A A A A A A A A A A Summary of 1987 Peregrine Production. 1st clutch Eggs 2nd clutch 3 3 4 3+ 1+ 4 2+ 4 4 4 3+ 3 Young Hatched 3 2 4 3 1 3 2 4 4 Young Fledged 3 1 0 3 1 3 1 3 4 3 4 3 1 4 2 3 1 4 3 4 3 3 3 2 3+ 3 3 3? 4 3+ 4 3+ 1+ 3 4 3 1 4 1 0 4 3 3 2 3+ 3 3 ? 0 0 78+ 49 A A A A A A A A A A A A A I A A A A A A A A A A 4 1 0 4 Total Sites Occupied: 29 Total Adult Pairs: 24 Total Breeding Pairs: 23 Total Successful Pairs: 22 Total Young Produced : 66 Total Young Fledged: 55 Average Fledged Brood Size: 2.5** Young Fledged Per Total Wild Pair: 2.04*** Young Fledged ~er Total Unmani~ulated Pairs: 1.77 * Young Fostered Approx 1 mile into Utah, so not considered here. ** Total Fledglings divided by Total Successful Pairs. *** Total Fledglings divided by Total Pairs. 17 55 �18 Table 2. Year Site 1 2 3 4 5 6 7 8 9 11 13 14 16 23 % Occ. 72 73 p P P P P P P P A P P P P P P P P P P P 74 75 Occupancy of Historic Sites 1972-73 76 77 78 79 80 81 82 83 84 85 t~ 86 87 P P P P P P P P P M M P P P P P P P P P P P P P P P M M P P P P P P P P P P P t~ M F P F P P P P P P P P P P P P F P P P P P .p P P P P P 50 57 A A F A 93 79 P = pa ir 64 43 P P 36 P 36 29 A = lone adult Table 3. P 36 P 36 M 36 29 29 M= lone male 21 21 F= lone female Cumulative Eggshell Thicknesses Year Average n 1980 .306 17 11 (65%) 9 (53%) 7 (41%) -14.8 1981 .320 22 10 (46%) 8 (36%) 4 (18%) -10.9 1982 .325 26 9 (35%) 5 (19%) 2 (8%) -9.5 1983 .319 27 10 (37%) 7 (26%) 3 (11%) -11.1 1984 .319 30 13 (43%) 8 (27%) 1 (3%) -11.1 1985 .312 22 12 (55%) 8 (36%) 2 (9%) -13.4 1986 .312 28 16 (57%) 14 (50%) 5 (18%) -13 .1 1987 .304 21 13 (62%) 12 (57%) 7 (33%) -15.3 * Measurements Number of eggs thinner than -13% -18% -15% of whole, first clutch eggs. % thinner than pre DDT era eggs �19 Table 4. Site Responsible Agency 1987 Hack Site Summary Young Into Site Young Released Young Reaching Independence Deer Mtn. NPS 5 5 5 Beaver Creek BLM 5 5 5 Big Hole BLM 4 4 3 Twin Mtn BLM 4 4 2 Adobe Creek FS 4 4 4 22 22 TOTAL Young placed per 5 ite ................... 4.4 Young released per site .•••••••••.••.•.• 4.4 Young achieving independence per site •.. 3.8 -19 ��'/ 21 FEDERAL ... .. :-~.~. i! .. ~ <~ e '''__:,', State ; AID JOB PROGRESS REPORT of Colorado ';~" Project No. 86-039-01 Work Plan Job ~Perlod Covered: 1 ., 1 1 January 1986 thro~gh 01 June 1981 ., .... " Personnel: D. Bolster, Univ. Colorado; G. C. Miller, __Colorado Division -' of Wildlife ',. ABSTRACT .. '; .... -:.," . '1 The irstudies and activities reported herein relate to the various <. objectives of the Work Plan 86-039-01 (Appendix A), ~nitiated in 1985 and ..•. 1986. Two projects·, Habitat Changes in Colorado and Habitat Conservation .: Program, Planning, were completed. A reorganization of DOW's Habitat Resources Section (Management Team decision, 1 April 1987) eliminated the posi tion responsible for 5 projects. Status, budget and personnel .•. allocations, needs for completion, and required actions to continue or ": .. ..... : terminate the 5 projects are provided. Briefly, personnel and budget needs, and required actions are: ~,:', ..... ,,' " .' :,' .: .....; ECOSYSTEM MANAGEMENT 1. Managing Forested Lands for Wildlife a. Training/Implementation.0.25 FTE (12.5 wk.); $3,500 operating; Go/No go Management Team decision. b. Evaluation (West Antelope ~imber Sale). - 0.2 FTE (10 wk.), volunteer (4 mo.); $I,200/yr. operating; Go/No go decision by-Habitat, .'_Terrestrial, -·Aquatic, SW Region needed; inform other cooperators (BLM, ..;FS). .... /. ·.c: . ,... 2. Managing Rangelands for Wildlife.-Inform that CDOW involvement has been terminated. c~op~ratin~ agencies ., .; " ·3. Standards and guidelines for low-elevation riparian zones a. Inventory/Setting ·of Objectives .. - 0.25 FTE (12.5 wk.); $7,500 operating; Go/No go decision by Mangement Team. . b. Meadow complexes. standards. - 0.05 FTE (2.,5wk.); $.3,400 operating and travel from Jul 87 .;Sep 88; Go/No go decision by Haoitat, . Terrestrial Resources, NE Region;, inform CU Boulder, Denver Audubon of . decision. 'to " HABITAT INVESTIGATIONS ,;' ..... ,.,'.:';:; ~~, .,".:~ ~\.)?' ... ',' " ," .•.. ��23 Page 2. ECOSYSTEM MANAGEMENT Gary C. Miller The Colorado Division of Wildlife is statutorily obligated to protect, preserve, enhance, and manage all species of wildlife and their environments in Colorado (C.R.S. 33-1-101, 33-8-102, 39-22-701), a process known as ecosystem management (Graul and Miller 1984). The process differs from traditional single-species management in that the minimum needs of ~ll species are addressed prior to optimizing conditions for any single or group of species of particular concern (Graul and Miller 1984,_ Hoover and Wills 1984). Although the conceptual framework of ecosystem management processes is in place and a methodology for implememtntation has been proposed, (Hoover and Wills 1984), on-ground implementation has just begun and evaluation has yet to occur. Implementation and evaluation of the processes within the Colorado Division of Wildlife is the goal of this project. 1. tlan~ging Forested Lands for Wildlif~tlELNl Ob~£tives (from Work Plan 86-039-01): 1. Ensure that all permanent CDOW personnel involved in wildlife habitat management activities are trained in the application of MFLW to a level of expertise comparable with their responsibilities. 2. Develop or modify a computer program, compatible with CDOW's WANG system, that automates the process described in MFLW 3. Monitor USDA-Forest Service implementation of MFLW and ensure linkage between that implementation.and CDOW objectives. St~t~~: Approximately 200 DOW employees received initial training in the MFLW process in 1985 and 1986 (Miller 1986). Approximately 60 DOW personnel have received "hands on" training in the use of the automated program on FS Data General computers. A WANG-compatible program became available in May 1987--0 DOW personnel have received training in its application. Training requests are outstanding from approximately 20 regional personnel in NE, SE, and SW regions. To perform any but a basic emphasis species (usually deer and elk) analysis of proposals requires a computer and program--the latter was unavailable until recently. Additionally, treatment of requests for input to FS treatment proposals has been uneven--significant numbers of both FS and DOW personnel appear to feel the MFLW process is the full extent of DOW's input. Consequently, many proposal are evaluated with only an emphasis species objective. Data are unavailable at present to evaluate the relative amounts of FS lands treated under the Emphasis, Ecological Indicator, or Combined alternatives. As of 31 December 1985, 360 mi2 of FS lands were being managed under the MFLW process. The amount of land has increased substantially since then--DOW has no centralized means for tracking areas, amounts of land, or management objectives for those management units. Table 1 . demonstrates the complexity of tracking implementation areas--using only Arapaho-Roosevelt N.F as an example. Since many implementation areas cross District and Area, less frequently Regional, boundaries, monitoring implementation and DOW input to the scoping and decision documents will be complex, but probably could be done at Regional level with either dBase III or Lotus 1-?-~ snf~w~rA �24 Table 1. Management units tentatively scheduled for implementation ,. " M'£!l~g_i.ng Forested Lands for Wildlife on Arapaho-Roosevelt National Forest, Colorado. of ~ =========================================================================== Ranger District . : .. I i PreScoping Scoping Document Decision Document Mammoth/Pisgah Lump Gulch N. Boulder Middle St. Vrain N. St. Vrain Sugarloaf Caribou James Ck. 85-86 85-86 86-87 86-87 87-88 88-89 89-90 90-91 85-86 85-86 86-87 87-88 88-89 89-90 90-91 91-92 85-86 86-87 ,87-88 '88-89 89-90 90-91 91-92 92-93 REDFEATHER Long Draw George Ck. Many Thunders Lower Green Ridge Brown Ck. Laramie R. Panhandle Roach Poudre Canyon Eaton Timberline 84-85 85-86 85-86 85-86 ? 85-86 85-86 86-87 86-87 86-87 87-88 85-86 85-86 85-86 85-86 86-87 86-87 86-87 87-88 87-88 87-88 88-89 85-86 85-86 86-87 86-87 87-88 87-88 87-88 88-89 88-89 88-89 89-90 ESTES POUDRE Buckhorn Poudre Wilderness Poverty Twin Sisters, Comanche Ski Canyon Cedar Park Elk Ridge Pingree Crozier Lion Gulch 85-86 85-86 86-87 86-87 86·-87 87"'"88 88-89 89-90 90-91 91-92 85-86 86-87 86-87 87-88 87-88 88-89 89-90 90-91 91-92 92-93 86-87 86-87 87-88 87-88 88-89 89-90 90-91 91-92 92-93 93-94 86-87 86-87 87-88 88-89 89-90 90-91 91-92 86-87 87-88 88-89 89-90 90-91 91-92 92-93 87-88 88-89 89-90 90-91 91-92 92-93 93-94 85-86 86-87 86-87 87-88 88-89 89-90 90-91 85-86 87-88 87-88 88-89 89-90 90-91 91-92 86-87 87-88 88-89 89-90 90-91 91-92 92--93 BOULDER " '''., Area I Devil's Canyon Mill Ck. W. Chicago Ck. Leavenworth Loch Lomond Blackhawk Pk. St. Hwys. 5 & 103 Corr. CLEAR CREEK SULPHUR Stillwater-Supply Corona Doe Meadow Beaver-Muddy Elk Ck. Crooked Ck. Buffalo Park Obj. Input E I C Date By �25 Needs: "Hands on " training of operations and regional staff pe'rson~el-'On the new WANG-compatible MFLW process is needed if DOW is to play an active role in habitat management activities on FS lands. A reasonable amount of standardization among Regions for monitoring MFLW implementation is needed to assess how much effort DOW does and should expend in the MFLW process. Efficiencies of operations personnel's time should be realized with, at a minimum, a "checklist" approach to responses (Fig. 1) or, preferably, a modification of the automated program to allow "What if" analyses of DOW alternatives. These needs will become more critical in several years as current Forest Plans come into the 10-year public review process. Bugget and ~ersonn~l_AllQ~~tiQn~: Annual allocations under the "039," cost-center in FY 86 and 87 were approximately .25 FTE and $3,500 operating and travel for program development and training. Reggired_Action~: Go/No-go Management Team decision to continue DOW involvement with MFLW process. If "Go," resolution of responsibilities and budget allocations (budget for FY 88 was included in the Habitat Conservation program plans) ._. b. Evaluation--West AnteloEe Timber Sale The Antelope Creek area near Gunnison provides habitat for about 400 deer and the same number of elk (Anon . , Importance of Antelope Creek for elk and deer as effected (sic) by proposed logging operations. Report dated 02/15/85. CDOW Gunnison Area Office files). Results of reconnaissance between December 1986 and March 1987 indicated that significant amounts of both hunting and nonconsumptive recreation occur on and near the area. A proposal by USDA-FS to harvest timber from the area following the guidelines of tlgngging Forested Lands for Wildlife presented an opportunity to compare results of the West Antelope Timber Sale with the results predicted by the MFLW process. This was to be a co-operative study among USDA-FS, USDI-BLM, DOW--SW Region, and DOW-Habitat Resources. QQjectiY~ (from Work Plan 86-039-01): 3. Monitor OSDA-Forest Serviced implementation of MFLW and; to the extent permitted by OSDA-FS statutes and regulations, ensure linkage between that implementation and CSOW objectiv~s: ~1atu~: Planning began in late 1986. There were to be 4 major data-collection years between 1987 and 1994. Following inter-agency discussions a draft MOO was prepared and sent for review (Appendix B). Literature was reviewed prior to final study plan development (Appendix B). Fourteen elk were trapped and fitted with radio transmitters in 1987. Field work should have begun in Spring 1987--a potential volunteer workei from the Gunnison area had been contacted. Involvement of the Habitat Resources researcher terminated in April 1987. tl~~g~: Completion of study plan and MOO. Data collection system must be standardized. Personnel needs must be accommodated (quite possibly the volunteer is no longer available). �26 \' -----------------------RESPONsEs-rO-usDA=FS-PROPOSALS--------------------DOW INPUT WORKSHEET Date: Completed by: (Name) DOW Region: _ DOW Area: Nat. Forest: Management (Title) DOW District: _ Ranger District: Unit: Impl. Area: _ Forest Plan Presc. : Scoping Doc. Pre-scoping Decision (Date) (Date) (Date) This proposal should be designed following the Managing for Wildlife process Concur / Do Not Concur MANAGING FORESTED STEP l-Identify, STEP 2-Select LANDS FOR WILDLIFE Describe Wildlife Management Habitat (FS Recommendation) Goal Lands Area: (Concur / Do Not Concur--Attach.l) (Emphasis, Spp. Rich., (Concur / Do Not Concur--Attach. Habitat STEP 5-Prescribe, Forested PROCESS STEP 3-Selection of Species: Forest Service Recommends STEP 4-Develop _ DOW Recommends Combination) 2) (Attach. 3). (Concur / Do Not Concur--Attach. 4) Objectives: Schedule Treatments: (Concur / Do Not Concur--Attach. OTHER MANAGEMENT CRITERIA FOR IMPLEMENTATION Configurations, Special Concerns): (Roads, Dead Trees, Stand Approved: (Regional Manager/State Fig. 1. Draft "checklist" treatment proposals. Wildlife Manager) form for DOW responses to USDA-Forest Date Service _ 5) �27 ~ggget_2ng_fgK~Qnngl_hlloc~tiQn: Contributions from the 01-039-01 cost center were to be 0.2 FTE and $1,200/yr operating and travel, plus providing a 4-wheel-drive vehicle for use by the volunteer lab~rer. BgggiKed_Ag1iQn: The project was originally supported by the Habitat Resources and Terrestrial Resources Sections and the SW Region. In addition, the project impinges upon the objectives of all 4 major programs of DOW. Personnel of Habitat Resource, Terrestrial Resources, SW Region, and Aquatic Resources should consult to decide the future of the project. The other cooperating agencies should be advised of the decision. QQjggtivg: Provide a system similar to ~an£ging_Forested Wildlife for Colorado's rangelands (Appendix C). L2nds for St21g~: A decision was made by the Executive Steering committee to scale down the objectives, concentrating on one ecosystem as a prototype. Subsequent meetings of the Technical Steering committee resulted in selection of one of the "sagebrush steppe" areas near Gunnison for development of the prototype system. Decisions were made to eliminate several chapters contained in the original proposal, contract the first chapter, and contract drafts of the wildlife species requirements chapter (to be subsequently reviewed and modified by agency personnel with appropriate expertise). An April 1 Management Team decision reportedly was to terminate CDOW involvement in the project. R~ggir~Q_AQtion: Notification to cooperating involvement in the project is terminated. 3. Standards and Guidelines for Low-Elevation agencies Riparian that CDOW's Zones. Objgctiyg: Develop standards and guidelines for low-elevation riparian zones based on ecosystem management principles~ Put ecosystem nanagement principles into practice through Colorado Di"vision of ~ildlife habitat management activities. £i2ig2: The standards and guidelines provided in Miller (1986. labitat Investigations Report. 66p.) were slightly modified, primarily lith respect to the method by which habitats on DOW properties are mapped Ind inventoried. A sample of the modified technique (for the Dodd 3ridge, Brush, Cottonwood SWA's) is provided (Appendix D). A draft lanuscript (co-authored with W. Snyder), incorporating this work with ~hat from project N-4-1-R, documents the rates and direction of changes _n abilities of Colorado's low-elevation riparian zones to sustain listorical levels of species richness. Recommendations for acquisition ,bjectives along various stream segments were made (background data from lildlife 21 report). A pamphlet (Hiller, Sherman, and Houston. 1986. ;ottonwood riparian zones of Colorado. 10pp.) (Appendix E) was produced �28 for, and presentations made at, the Colorado Water Workshop in Gunnison, July-Aug 1986, with approximately 40 attendees and 100 pamphlet recipients. Additionally, a project to map habitats on major DOW properties was conducted under a contract with Colorado State Forest Service (Bill Olmstead administered). Those maps that were completed are currently at the CSFS photo-interpretation shop in Fort Collins and may be obtained by appropriate personnel. ~ggg2: Standards and guidelines need to be drafted into an integrated package, including monitoring system and objectives, distributed for review, and modified as needed. Once in appropriate format (Administrative Directive, Instructional Memo, SOP, etc.), acceptance or rejection by Management Team. ~ggggt_~nQ_fgK2Qnngl_AllQQ~tion: .3 FTE/yr, plus $7500/yr. operating and travel (graduate study--intern). Reggireg_~QtiQn: Although this activity was initially included in the Habitat Conservation plans, it carried major implications to the Hunting, Species Conservation, and Watchable Wildlife programs, as well as being a large part of the basis for the proposed expenditures shown in Wildlife 21. As such, a Go/No go Management Team decision appears appropriate, with funding and responsibility decisions accompanying a "Go" decision. Objgctiyg: Develop appropriate ecological indicator species list and species requirements data for riparian meadow complexes. Statu£ (Prepared by D. Bolster and G. C. Miller): A project to ascertain habitat characteristics and develop management standards and guidelines for riparian and adjacent upland meadow complexes in northeastern Colorado was begun in July 1986 and is continuing to the present (Appendix F). Field work was performed by David Bolster, Univ. of Colorado graduate student (M.S. candidate) and CDOW intern. 3urveys of habitats and avian communities were conducted in a variety of riparian and upland meadows of the South Platte River valley in Weld and Logan counties, northeastern Colorado, between July and September 1986. Three general habitat categories--undisturbed grassy meadows, rangeland, ~nd cropland--were surveyed and a profile of avian communities for each type was compiled. Initial reconnaissance was performed by selecting ~pparently representative patches of the various habitats and walking "zig-zag" patterns to cover most of the habitat patch. Observation- of jirds were made with 10-power Minolta binoculars. 3pecies most common in the more limited habitat types were: Horned lark (~K~mQ2hil£ £12~2t£i2 ), lark bunting (Q£1£mQ22i~£ m~1£nQQQ£~2)' western dngbird (IYK£nng2 ~~£tiQ9.1i§.), and barn swallow (tlinmQQ KgstiQ£)· 3wainson's hawk (~gteQ_2~9.in§'Qni), American kestrel (E9.1QQ_§'E9.K~~£ig§.), lnd brown-headed cowbird (tlQ1Qth£g§._at~£) were not common but present in 111 habitat types. Upland sandpipers (~9.Kt£9.mi£_longiQ9.gg9.),listed as a 3pecies of Special Concern by the Colorado Division of Wildlife (Webb 1985) was locally common in lightly and moderately-grazed rangelands and 1ewly-mowed alfalfa (tl~giQ9.gQ_§'9.tiy9.) fields, especially those reported )y landowners to be infested with alfalfa weevils (Family: ~urculionidae). Bobolinks (DQliQhQnYZ_QKY~iYQKQg§) and grasshopper ;parrows (AmmQQK£mg§_§av£nn£Kgm) appeared highly localized and restricted �29 to very few habitat types (Table 2). Data from the initial surveys and recQnnaissance are being analyzed and a study plan is being developed for the'period 1987-1988. Table 2. Summary of potential indicator species observations and·habitat associations in northeastern Colorado, July - September 1986. --------------------------------------------------------------------------------------------------------------------------------------------------NO. NO. SPECIES STUDY SITE HABITAT OBSERVATIONS INDIVIDUALS NO. TYPE a/ Upland sandpiper Bobolink 39 79 9 RH,RM,RL A,AC,S,C, BG,OF Triangle 15 49 5 RH,RM,RL A,AC SUBTOTAL 54 128 9 6 9 3 0 0 0 9 12 3 RM,RL,OF Triangle 7 15 2 RM,RL SUBTOTAL 16 27 3 Crook Crook Triangle 3rasshopper sparrow Crook RM,RL,OF =========================================================================== ]./Habitat types-RH,RM,RL - heavily, moderately, and lightly grazed ~angelands, respectively; A-alfalfa; AC-cut and baled alfalfa; 3-cultivated sunflower (B~li~nthg~ sp.); C-corn (~~~ m~Y2); BG-bare 5"round; OF-old field, "go-back" �30 The project is a cooperative venture among CU Boulder (providing support 9 mo./yr. with a teaching assistantship), CDOW Northeast Region (provision of living quarters and miscellaneous support by field personnel), and CDOW Habitat Resources Section (provision of vehicle, operating expense and camp-rate per diem). Total cost to DOW is estimated at $5,000 for the entire study. As of 1 July 1987, approximately $1,600 will have been co~nitted to the project by Habitat Resources. A total of $3400, then, will be needed over 2 fiscal years to see the project to completion (field work ends in September 1988). Other entities expressing interest in the project are: DOW Terrestrial Resources Section and Denver Audubon Society. ~~~g~g_AQ1iQn.- Parties of interest need to decide if and how to continue with the project past 1 July 87. These include: ~avid C. Bolster, CU graduate student, and Alex Cruse, CU Dept. EOPO Biology, faculty advisor. There are recent indications that, at least in certain areas of Colorado's low-elevation riparian zones, significant losses of trees may occur (Miller 1986, Miller and Snyder ms. in prep.). In addition, Miller (1986) found that enhancing vertical heterogeneity of these areas could benefit a number of species with little risk to ecological indicator species. One strategy to address both of these management concerns is to explore the potential of trees and shrubs, native to North America but not currently found in Colorado's low-elevation riparian zones, to become established in selected areas. St~tu.§.:Bre ;Jp tbil1~-q:e~;d no pi occed frO'rJpc~ design and study area selection phase.$~0Plani~o contract with Colorado State Forest Service for propogation of seedlings in spring of 1987 were not initiated. ~ggg~1_~nd_r~K'§'2nn~1_A112£ati2n: 0.16 FTE (8 wk.) researcher, volunteer/intern (3 mo.), property tech. 4 da.; $10,000 operating in fear #1, $3,000/yr. thereafter. ReggiK~g_A£tiQn: The project impacted primarily NE, SE, and :entral regions, CDOW, and Habitat Resources Section. In addition, the ?roject was designed to complement an existing Terrestrial Resources cesearch project by Warren Snyder regarding propogation of cottonwood ~rees with stem cuttings. A Go/No go decision by the impacted )rganizational units is needed, with the decision forwarded to field ?ersonnel. 5. H~Qit~tLR~31_E.§.1~1~_Ey~lg~tion_§y.§.t~m.-Initial drafts on file, Habitat tesources Section. :>. H3Qi131_~hQng~.§._1.n_Col.Qrad.Q. -Project completed, (Appendix G). 7. H~Qit31_~2.n'§'~KY3112.n_rK2gK3m_r13n.§..-Project completed. :ile, Habitat Resources Section. PREPARED __:~~_::.....:bJ~~:_\~~~L_' BY: _~O__:_~.E::...!Y'~\...;f.-1 I _ Documents on �31 APPENDIX A WORK PLAN State of Project 'Nork Plan Job QQ.1Q.K£gQ. §_§=~39=~1 ____ 1 ____ 1 _ _ The Colorado Division of Wildlife is statutorily obligated to protect, preserve, enhance, and manage all species of wildlife and their environments in Colorado (C.R.S. 33-1-101, 33-8-~02, 39-22-701), a process known as ecosystem management (Graul and Miller 1984). The process differs from traditional single-species management in that the ninimum needs of all ~pecies ar~ addressed prior to optimizing conditions for any single or group of species of particular concern (Graul and 1iller 1984, Hoover and Wills 1984). Although the conceptual framework Jf ecosystem management is in place and a methodology for implementation ias been proposed (Hoover and Wills 1984), on-ground implementation and ~valuation of the process has yet to occur. Implementation and ~valuation of ecosystem management processes within the colorado Division )f Wildlife is the goal of this project. 1. Ensure that all permanent CDOW personnel involved in wildlife 1abitat management activities are trained in the application of Managing Torested Lands for Wildlife (MFLW) (Hoover and Wills 1984) to a level of ~xpertise comparable with their responsibilities. Such training may Lnclude introductory classes, "hands on" woz ks hop s , or implementation on :::DOW properties. 2. Develop or modify a computer program, compatible with CDOW's 1ANG system, that automates the process described in MFLW. 3. Moni-tor USDA-Forest Service implementation of MFLW and, to the ;xtent permitted by statutes and regulations, ensure linkage between that ~mplementation and CDOW objectives. 4. Assist operations personnel in each region in the implementation ~o ecosystem management approaches on specific CDOW properties--2 :egions/year. 5. Assist in the development of appropriate ecological ;pecies lists for various ecosystems within Colorado. ;ary C. Miller 'acant, interns or volunteers 1.0 FTE 0.25 FTE/yr. indicator �32 ~ggmQni_QQ~t: $61,261/ yr. (01~ Personal Services Title Name Rate Period Gary C. Miller Wildlife Researcher Vacant Interns, Volunteers C 1.0 yr. Total $43,211 $43,211 -0- 0.25 -0- =========================================================================== (21) Operating Supplies and Services Item Number Uni t Cost- Total 30,000 mi. Vehicles 3raphics, drafting 5 hr. Photographic supplies, processing var. :omputer software var. ~aps, aerial photographs var. 3mall equipment var. $0. 22/mi $6,600 $40/hr 200 var. 200 var. 600 var. 600 var. -~@Q $8,500 Total --------------------------------------------------------------------------------------------------------------------------------------------------(28) Travel :~ame Days }ary C. Miller Intern/volunteer 60 65 Rate Total $31. 50/da. $20/da 'I'ota L $2,250 _Ld00 $3,550 (31) Capital Expenditures -------------------------------------------------------------------------- ~amper shell for pickup \ir condo for camp trailer ~ANG PC computer, peripherals 500 600 '4,900 $6,000 $ Total ~raul, W.D. and G.C. Miller. 1984. Strengthening lpproaches. Wildl. Soc. Bull. 12: 282-289. ecosystem management loover, R.L. and D.L. Wills. 1984. Managing forested lands for lildlife. USDA Forest Service and Colorado Division of Wildlife, )enver. 459 pp. �l' a Le : Mr'LW3 33 APPENDIX MEMORANDUM B OF UNDERSTANDING This agreement, made and entered into this day of , 1981, by and between the Southwest Region and Habitat Resources Section, Colorado Division of Wildlife, and the United States Department of Agriculture, Forest Service, Grand Mesa, Uncompahgre, and Gunnison National Forests and the United States Department of the Interior, Bureau of Land Management, District. This Agreement shall become effective when all parties have signed it and remain in effect until terminated in writing by one of the parties with 30 days written notice of intent to terminate to the other party. WITNESSETH: WHEREAS, the USDA-Forest Service (hereinafter referred to as Forest Service) is organized and maintained to provide for the administration, protection, and improvement of National Forest lands within the Grand Mesa, Uncompahgre, Gunnison National Forest for the multiple benefits of timber, range, wildlife, recreation, and water, AND WHEREAS, the USDI-Bureau of Land Management (hereinafter referred to as BLM) is organized and maintained to provide for the administration, protection, and improvement of BLM lands within the for the multiple benefits of timber, range, wildlife, recreation, and water, AND WHEREAS, the Colorado Division of Wildlife, Southwest Region (hereinafter referred to as Division of Wildlife), is organized and maintained to protect, preserve, enhance, and manage the wildlife of Colorado and their ~nvironment for the use, benefit, and enjoyment of the people of the state and its visitors, AND NHEREAS, participation by the Forest Service, BLM, and Division of ~ildlife in cooperatively evaluating the effects upon wildlife of the Nest Antelope Timber Harvest is within each party's mission. ~mw, THEREFORE, in consideration of the above premises, and in the interest of attainment of common objectives, to the mutual benefit of the ?arties involved and the citizens of Colorado and the Nation, the parties iereby agree to cooperate in the evaluation of the effects upon wildlife )f the West Antelope Timber Harvest described in Appendix A as follows: \. THE FOREST SERVICE WILL: 1. Provide the needed radio transmitters, as shown in Appendix )r their cost-equivalent in goods and services needed for the ~ccomplishment of the objectives described in Appendix A. 2.Provide, install, and maintain at least 2 traffic Locations designated by the Division of Wildlife. counters A, at 3. Design and implement the west Antelope Timber Harvest in lccordance with the guidelines established in the book, ~~n~ging_EQ~~§teg_ !~ng~_fQK Wilglife, Hoover and Wills, eds. (1985). 4. Provide % of fixed-wing aircraft flight time needed lccomplishment of the objectives described in Appendix A. _- 5. Provide compensation :>imilar individual to collect for a university graduate necessary field: data. student or for the �34 B. THE BLM WILL: , 1. Provide a 4-wheel drive vehicle for a graduate student or similar individual to be used in data collection and other duties associated with the evaluation as stipulated in Appendix A, and in no case to exceed months/year. 2. Provide living quarters for a graduate student or similar individual to be used while during periods of data collection and related activities in the Gunnison area. 3. Provide Division of Wildlife personnel access and permission to carry out wildlife baiting, trapping, and marking activities upon their lands as needed for the accomplishment of the objectives described in Appendix A. C. THE DIVISION OF WILDLIFE WILL: 1. Provide a scientifically sound design for a quantifiable evaluation of the effects upon wildlife of the West Antelope Timber Harvest. 2. Capture, process, and fit with radio transmitters or other appropriate markings any animals needed for the accomplishment of objectives described in Appendix A. 3. Provide radio receivers or their cost-equivalent in goods and services needed for the accomplishment of the objectives described in Appendix A. 4. Provide snowmobile(s), operating and maintenance expenditures. 5. Provide the personnel and living expenses (per diem) needed to oversee the evaluation, and conduct on-gro~nd data collection except as stipulated in A. above. 6. Perform and otherwise. or oversee data compilation and analysis, computerized 7. Provide written reports of progress no less than annually, ~ritten final report no later than 1 year following the termination data collection. and a of 8. Acknowledge Forest Service's and BLM's contributions in all ?ublications resulting from the evaluation, and provide copies of all 3uch publications to Forest Service and BLM. �Memorandum of Understanding IT IS MUTUALLY p.3. 35 AGREED: < 1. In connection with the performance of work under this Memorandum of Understanding, all parties agree not tho discriminate against any employee or applicant for employment because of race, religion, color, or national origin. The aforesaid provision shall include, but not be limited to, the following: employment, upgrading, demotion, or transfer; rates of payor other forms of compensation; and selection for training, including apprenticeship. The Division of Wildlife further in all subcontracts hereunder. agrees to insert the foregoing provision. (Date) Supervisor, Grand Mesa, Uncompahgre, Robert K. Towry Regional Manager, Southwest Bruce McCloskey State Wildlife Manager, Region, and Gunnison Colorado Habitat Resources, National Forests (Date) Division of Wildlife (Date) Colorado Division of Wildlife �36 ECOSYSTEM MANAGEMENT MANAGING FORESTED LANDS FOR WILDLIFE EVALUATION (W. ANTELOPE TIMBER SALE) STUDY PLAN Project: ~I~~~~~~~l _ Work Plan: _1_ Job (Objective #):_~L_~ BAQKgRQQMQ_ANQ_~Q§TIFIQAIIQN: The Colorado Division of Wildlife is statutorily obligated to protect, preserve, enhance, and manage all species of wildlife and their environments in Colorado (C.R.S. 33-1-101, 38-8-102, 39-22-701), and the USDA Forest Service is likewise obligated to maintain habitats for viable populations of all native and desirable non-native vertebrates, as well as for management indicator (National Forest Management Act of 1976, 36 CFR 219, §gg ~lso Salwasser and Tappeiner 1981, Salwasser et al. 1982). Although the conceptual framework of this process, called ecosystem or ~ommunity management, is in place (Graul and Miller 1984), and a nethodology (Hoover and Willd 1984) has been implemented (Miller 1985. :010. Fed. Aid Job Prog. Rep. 86-039-001. Colo. Div. Wildl., Denver. [64 pp.]), evaluation of the process has yet to occur. Such evaluation --that is, how closely do MFLW "predictions" match wildlife community and ~abitat responses to treatment--is the objective of this study. Implementation of a timber harvest (sale) on the Taylor River Ranger )istrict, Grand Mesa, Uncompahgre, and Gunnison National Forests (GMUG ~F)has been designed in accordance with certain processes described in 1~naging_Iore§ted L~nd2-ior Wildlife (hereafter referred to as MFLW) (Hoover and Wills 1984) (letters "and memoranda provided in Appendix I, lee al§Q Decision Notice of 30 Jun 1986 (GMUG NF) and Envoronmental \ssessment, 20 Jun 1986, GMUG NF).). Treatments may impinge upon big 5ame (deer and elk) wintering range, elk calving areas, and wildlife ~ommunity dynamics. Division of Widlife, USDA-Forest Service, and JSDI-Bureau of Land Management representatives are in agreement that such 3valuation is· needed. 2~JECTIYE§ (See Habitat Investigation Work Plan, Proj. 86-039-01 and ~7-039-001) #3-Monitor implementation of MFLW and ensure linkage between ;uch implementation and CDOW objectives £ng #5-Develop appropriate !cological indicator species lists for vaious ecosystems within Colorado. Subobjectives: a. Measure habitat characteristics and responses of management ndicator (emphasis) species pre- and post-treatment (Habitat :haracteristics landscape use by species). b. Measure habitat characteristics and community response of 'cological indicator species pre- and post-treatment c. compare values ascertained post-treatment with those redicted by the MFLW process using pre-treatment values. = �Appendix B (cont'd) 37 LITERATURE CITED Graul, W.D. and G.C. Miller. 1984. Strengthening approaches. Wildl. Soc. Biull. 12: 282-289. ecosystem management Hoover, R.L. and D.L. Wills, ~ds. 1984. Managing forested lands for wildlife. Colo. Div. Wildl. and USDA Forest Service, Denver. 459 pp. Lyon, L.J. and cover. Salwasser, integrated 1979. Habitat effectiveness J. For. 77 (10): 658-660. for elk as influenced H. and J.C. Tappeiner II. 1981. An ecosystem timber and wildlife habitat management. by roads approach to Salwasser, H., I.D. Luman and D. Duff. 1982. Integrating wildlife and fish into public land forest management. Proc. Western Assoc. Fish Wildl. Agencies, 62: 293-296. Wright, K.L. 1983. Elk movements, habitat use, and the effects of hunting activity on elk behavior near Gunnison, Colorado. --M.S. Thesis, Colorado State Univ., Fort Collins~ 206 pp. Young, S. 1982. Migration patterns of the upper Gunnison River elk herd. M.S. Thesis, Colorado State Univ., Fort Collins. 104 pp. ��39 Prospectus, APPENDIX C Managing Rangelands EXECUTIVE for Wildlife SUMMARY ,. Production of a book (working title Managing GrasSiands and Shrublands for Wildlife) (MGSW) patterned after the CDOW-USDA-Forest Service publication, Managing Forested Land~ for Wildlife, is proposed. Potential cooperating agencies are: Colorado Division of Wildlife, USDA-Forest Service, USDA-Soil conservation Service, and USDI-Bureau of Land Management. Need and potential demand for such a publication are based upon: 1) high priority of non-forested rangelands for wildlife (including deer and elk winter range, and habitat for several threatened and endangered grouse species)--two of three CDOW highest-priorit~ terrestrial habitats include these non-forested areas; 2) ~he largest proportion of Colorado (about 48%) that is non-forested rangeland; and 3) the apparent high demand for the . precursor, Managing Forested Lands for Wildlife (85% of first printing gone within six months). Production of MGSW will yield increased efficiencies in cross-discipline and interagency communication, which should result in higher levels of wildlife outputs. A si"ngle. computer-compatible met.hod of predicting and assessing the effects upon wildlife of various livestock and range improvement treatments . will be designed to improve wildlife habitat. The project can be completed by 1 November 1988, given a t October 1986 starting date. Approximately $120.000 will be the cost exclusive of agency personnel's time (191 person weeks). Total cost ($279,000) is less than the cost of Managing Forested Lands for Wildlife. Several funding alternatives are presented--the recommended one calls for $8,333 and $23,333 from each of~ three (3) agencies (BLM. Dm~ .•FS) in FY '86 and '87, with DOW "fronting" an additional $25,000 for printing costs (recovered in part from book sales). SCS contributions are primarily through chapter co-authorships, technical review, and membership on the Exec~tive and Technical Steering Committees. Gary C. Miller. Colorado Division of Wildlife Dale ~~ills, USDA-Forest Service Don Gillaspie. USDA-Soil Conservation Service Lee Upham, USDI-Bureau of Land Management �40 PROPOSAL MANAGING GRASSLANDS AND SHRUBLANDS FOR WILDLIFE An interagency publication of the Colorado Division of Wildlife, USDA-Forest Service, USDA-Soil Conservation Service, and USDI-Bureau of Land Management I. NeAd. Purpose. Scope. A joint publication of the Colorado Division of Wildlife and USDA-Forest Service, Managing Forested Lands for Wildlife (MFLW) (Hoover and Wills, 1984), became available in early 1985. The ecosystem management concepts outlined in that book were quickly embraced by the Rockj Mountain Region, USDA-Forest Service (Torrence letter, dated February 11, 1985) and the Co10~ado Division of Wildlife (Ruch letter, dated February 27, 1985). By July 1 all but approximately 250 of the initial printing of 1596 books had been distributed to resource managers, libraries, other agencies, conservation organizations, and universities throughout the United States. Demand for the book has remained high; it is being considered for use as a textbook in at least one Colorado university; attendance at training sessions in the applications of the book are increasing, indicating acceptance by field-based managers. A similar demand seems to exist for a companion publication directed toward managing non-forested ecosystems for wildlife--the ecosystem management concepts of MFLW should have applicability for these lands that are primarily used for grazing. The companion publication, tentatively entitled Managing Grasslands and Shrublands for Wildlife (MGSW), will provide a medium where four cooperating agencies mutually agree on a process for managing wildlife habitats-in grassland and shrubland ecosystems. A single method of describing wildlife habitat conditions on these lands, as well as a single, computer-compatible method of predicting and assessing the impacts upon wildlife of various· livestock grazing and range management treatments will be employed by all cooperating agencies. The results should be increased efficiencies in cross-discipline and interagency communication, reductions in duplications of effort, with a resultant increase in money and manpower available to design and implement wildlife nabitat and rangeland improvements. The proposed publication will provide a valuable reference document with practical application that personnel of the four agencies can readily access. Much of the information that will be contained in the publication is not currently available in a condensed form--much of it may not be available in published literature, but will be a result of many people drawing from personal expertise, resulting from years of practical experience. The scope of MGSW will be the non-forested ecosystems of Colorado, although those ecosystems (and, hence, the book's applicability) extend to other states of the western Great Plains, the intermountain and Great Basin areas, and the Southwest. Preliminary analysis indicates that 11 ecosystems may be addressed (reference Natural Vegetation of �41 Colorado, 1972, USDA-SCS), comprising roughly 48% of the State. Format will parallel MFLW (Attachment #1). Croplands will not be addressed--a recent publication of the Great Plains Agricultural Council addressed ", those lands. II. Methods. Production organization: Technical Reviewers of MGSW will be accomplished Executive Steering Committee Technical Steering Committee with an interagency ~ Editor Authors A. Executive Steering Committee - comprised of upper-level management representative from each participating agency (CDOW, USDA-FS, USDA-SCS, USDI-BLM). Primary purposes are to formalize agency commitments of resources and resolve major interagency "glitches" (resulting from possible policy differences). G. Hetzel (FS), E.Prenzlow (CDOW), C. Roberts (BLM) , and D. Gillaspie (SCS) may be appropriate members. B. Technical Steering Committee -comprised of one representative from each participating agency. Handle normal interagency coordination, provide direction to editor, resolve editor-author conflicts .. D. Wilis (FS), L. Upham (BLM) , E. Mustard (SCS), and G. Miller or'D. Smith (CDOW) may be appropriate members. C. Editor - one person, possibly under contract, mutually acceptable to all agency representatives, provides editorial continuity and direction to authors, oversees technical review and normal operations of book production, identifies author~reviewer conflicts. One alternative considered was to use multiple editors from the agencies--such an alternative would be very cumbersome, however. D. Authors - ag~ncy personnel; agency representation. E. Technical Reviewers - experts in various disciplines who critically evaluate manuscripts for technical accuracy and completeness. multi-authored :1 ' chapters :. ~ . for diverse �42 01 Oct 86 86 Draft Ch. 2 Ch. ~3 Wor-!-::shop Proposal Go-No Go by' Agencies Begin Ch. 2 F:esults CII. 3 l'-lorkshop Go-I'Jo Go by 01 1'1.::\ r 01 Jan uw \). 01 e-o 00 Draft \ ••}\.J .. Pro i ec ted Begin May 88 8 .. Final book Book Ch. 8) Ch. 8 Costs - over tvJO f isea 1 years (three Tech. q<= , ...J Steering Com. 3) Typing Services 4) Travel (workshop~ s . 25 15 Contract :2) Design 3~ Artwork, 4) Ch. for Book on-line~ di·:;tt-ibution DO!;)) Pers. 12~OOO vJks. $ ~ 28,800 layo~t) graphics 1~ c on t rac t; SUBTOTPiL _~, 28,000 10 8,000 20,000·. 8~(lO(l 56 -rs , 35(i 15'-:; , ooo 40,000 40~OOO 40~000 10,000 10,000 70,000 5,000 95,000 ()(1(i 25,000 103~OOO 8~550 20,000 (4-color Tot.al $ 11 a , ooo Editor Center copy 74~:200 i i 3, 65() 1) .. FY 88 F'prs. vJks. Authors 01 Nov 88 Camera-ready draft, F''(87 1) (e;-:. 1~ 4-7 Ch. __ -_-_---_ _-_--_--_---_ __ ---_ Ch. Draft Agencies-~: Begin _--_. 01 Jun 87 01 Feb 87 01 Dec . ':':,' �43 P{ 87 Total $ '$ c . G!E.'Q.lt~L E:':"R§£'!..c_:j_j,j;h.~[~ Printing @ $::'5!copy x 10eO copies i.L ,\ 2) ..,., ..:.. } Inb::TagEllcy cGlltribLtted Pel~s. and Tr··.:;\vel Line It8i1 E::...tdgeting of Fur-Ids 25, ooo :25,000 113 , .:S::5(1 45!1:3.S-{) i59,(:(X) ::5 !I C(x) 70,OC(1 95,CX):) Pr irrt irio GF.f.:tID TOTAL .....• c vi. $140, ~() 25l1C(x) $279. (leO \.jariou·::; +undanq opt.i.oos are pre:..;;.Enb:::din Attachrnent #2. Ti-'e t-ecorflm.... =-nded opt.i.oo calls fot- ·~:8~..::•.::•.::. and $:23,.'::.'::.'::. from each of three (3) age=.""1c i2::; CEU1, C["O;') , FS) in FY' ..87 and FY . El'.=:l, t-e'"::5pE-C ti ve I y . In add.it.i.on , cro!j ~'-Jill provi.oe "up +ront;" capital of $::'5,CX:x)to cover printing cO':5t'::;-part of thc)"3e co::;ts ·,...I0•... .rl d be n:::-covered from book sales to agenci2::; and ott-;ers (Option #3, {4tt.~.ch-nent #2). t:::"'t.rrriP,anl - Costs of producing 1·1ana.qinq Forest.f="'d Land,::; for Wild life !.-'Jere poor Iv tr'acked, but may i-'.a\!e approached $331,(XX), including agency per~.onnel 's titTle (Attach-rent #3). The grand total Estimate of $279,<Xx) for production of f'1!~"Sl.!j,therefore, appears t-eascna.ble. Cccperating agencies will need to provide perserlnel cOTlillibnent.s of 191, persrrl-tr.eeks (""j\/ei-the 2.1-~:leat- life CJf ti':e pr-cJject. Except; for initia.I'printing cost,s, eqLlal cormitmerrt of' 1··-e--:5L.iLtrces b':l th,~ (~:::)contribLrting cigericiE-S (FS~ EU1, ceo!)) 1.S reCCJfiff'end:=.-d. Applic.:3.tion Df [13:::1,; cou Id increa·:=..e ~'Jildlif2 outputs on t.he a.pprm:imately i?!: of Color-ado lC:"\.'1d~. ~~Jh.ich·:3rE both non-forested arid putil Lc Iv-cx-ried and th? ::::,:Si~ t.ha.t CiTe pri'v"a1:el'/ contt-olleci-grasslands and shrublarlds wl-ticl! at-e oft.c"'i' crucial habi t,-;ts of d8er and el k (~.-..,tinterr-a\nge), and se\feral t-d..gt-rpj-- iot- i t.'l =;] rc::_t5e ':5~-;.~2;:: iE':-=. (f~J,3.rne , ·a':3 \rJe 11 ,3.S threEt, t.E=r'led ,=-I'id s:iidEtl"lg2t-ed sp2cies) . pt-'3.lr-l2 ·3.i2 t~~i'C3 o+ th:~ t:.hr22 1··I.i.qre:=.:.t-pi-ic:)t-.i.t·~./ tet-~-Estri-=f,l hc·lbi.t3.ts -fc)t~~J.i lef 1.i,-h: in CoLor'ado , t'+-OCJ\/Et-, F:. L. culd D. L. Vji 11 s; , E:,"C.is", 192·~l. D.i\/ision of (..!jildlHe, L,jU.~ilif..§_~ Colorado Color-ado, 45'7 p , r..t=\i'"2~~9JnCl t.f~!::?:;t::§9_!::.§1I1d·::;........:f_Qc.... U3DA-Fore':5t SE-r.'ice, [}-2nver, �44 CUILIl\E :::h. 1 :h. 2 - n",Iff}!'"?; di':;b-ibution rr.::'<.p;vE:'qetz,tion, clim:::lfic ECCt5vstem L\?sct-ipti.c~IS cond i tions; soi 1s; special cornponE:f"lts; vJi LdLi, ofe a·:;::,i.JCiatio.'O·5; s:-?ral stages and asso:: Lated ~·Ji1d 1i fe va 1Ll2:; • Es timate 11 ecosystems. :h. 3 ~.:!:.}.dlife SFECies F:equirernents - picture; binomi-cil; distri!:~_(tion map; ecoS"-Y':;b:_4TiS; habitat r'equin?ffi'"<::2nts (r·epn:;..--:iu.ction, fec-?ding, cover , sp.:3.ce, sr-ec:ial features); mi.n.im.un viable pop ..... llations; iTk3.ti~ices. Grazinq :h. 5 :h. '...J ~h -7 _,I Tr-eabneryts and Effects ~~:anqe Ir~rOVE":,>oiE.~nt F'racticf2":; VJi ld Ii fe. - b':l ec::cy::~.ystem, including effects on L t n I :h..8 Sf?t tine( Go.:3.l sand ?~pol;hcatiCln:; land::: .• - brJD Obj e-cti \./es - simi Ia~- to applicat.i.ons, Su.ggested _R~.Q... DlE' iYFLi;J. few' public lands, Dlle for- pt"-ivate At;}E?rlC'i (m=..;·:.) tGt§'nti9..Lf!:.\.tt~Jr(~l (A:::Js!!_c;:LAfflliati..flll) o 1 2 (i 1 ecosvs 3 3 B" Jc~r-·inst.orl (FS), L. J LlrgE:flS (~~:.:S) ..::. j-' tE11lS) r, Danna (Etl1), ~) (t;.) spp; / ~~Jkshop) 4 . ~··jhitt(:?kind Zj_t~r-oth (f=:::3), l3. I)Jl!J), L ~"J . :::~-'1 (F'~;), \~ roo.. -:!' ,_: " \:~/;ji~l'- ( C ..E. II ~:chi:=lIT).:iJl!f L_. a LU-'gE-1i::} (::';1:;3), ~<inch (PJ'i) C3.r-:p=:rl tE'j"- ( COVJ) , ~I F~ Strol:el II (FS)!, C" 1 t.. \..J ; i- .1.\".1 -S • C:JUDtzi E;n",icJer (F'S), (CO;))!I E. Mu~.t6rd jvj. (SCS), B1'/in,!e~- (BU"1) 7 4 3 Capp (FS) ~ tlcl.lf=~:;pin (FS), Lipsconb Bridges CEL"1), 1"1ighton (FS) 8 24 4 \..:c\PP ~FS) ~ Li pscomb (DOl'J), Upham ( HJ'I) , ( pub l i.c bv F~l, EU'1; r.:Jt"""ivateby 9:'"S) mQ..tJ) , �45 0, P·_; r WI Total Line Item Budget I\Iea-:ls $1 =~)~000 $. :::;~),C(X) each PLTEFNATI \"ES 1. Equal <::;:.'5"/.) split ,::·uilClrig 4 cooperating agencies~ e<3.chreceiving books for ar;l2ncy use. '$ 6~:L"5-() each _ 2. Equal (33';·~)sp 1it ,3mongEU1, FS~ DC);JSCS corltr-.itutio.'l prim,-3.rily tht-c;(..tgh s tet.:?t-· ing CDT.T,i t tee-::;, au thorshi j:J'S, tEdTlical review (because of e:-:tn=iT!e budget ~"'r:::~:lT ic tic::ns) ~ach agency receivirt<;i books for agency' use. d: Equal (33. 3':!~) spl it atTD'!;:;:) FS ~ EU1 ~ [O!J for all e:-q:€flses, e:-:cluding $ZS~O()() pr irrt.inq , t.'Jhich iAA.Juld in i tia 11y be borne by CDO.!J. (.:Ic:~€'~nciE:.Y;:; conb-ibute through j:..1Lln:ha<:=...e of books, for theituse--D]i,'j h:3f",dlE:;s =,ah=-::;~ dis-. tribution of complimentary copie-:;. '$ ." ~Io.. ~-:: 0 '_,' , ,_10_10_1 each 8, ..:: ..:: ..::.each $. 4()=,(:(X) each $ 23,::::33 FS $ 23 ~..:: •.:: •.::.[-tJ1 $ 4B~~4 OO;.J lFigun?<..::; aSSUJT€~igE..:.rICies ccrit.r i.b... lte typing servicEs-if not, total budget. m:.->eds inc reas.e by $12 ~COO (F'{ 80S) and $-8,000 (FY 87)· 'for con tr Cl.Ct typing. .•..... . �46 ATIACH"Hff 3 Est. Personnel Ta'::l: R. Hoover R. Rltterf Chapt; , 1 ~·~]"~<6$i. editor* Chapt; , L 8:: .:3.ppenc.1i>~* . Chapt , .-: ..:.. Chapt. 3* ''0,) -:;~.t::: ..:..!! ,_c.;:.._. Chapt. B. Gillam R. TOIt.JeI·~y' 11-FS Detaile~-s 1o-rxn Det.:3.i 1Er-·::;· F:. Kufeld C. t-'fcl~r,ich D. \.lJills R. Mc.....••.:we B~' tviel ton D. Pfankuch J. Li psconb J. Capp S. M?3.1e~y' vJ .. 2.=:.ndfot-t Gal13.her w. I.::, Chapt., :~5,(}{)() ., editor- ...J~ 8< appd.* i.. w 6 :5~(}{):) 8 s.voo so.coo 12~t.C!{) 2;J),OOO - -::~) ~000 11,::;(:() s, E:di tot- 1':/, Chapt. Chapt. 7 ,8 '':', r·~.......... ~- 7~8 ,_:,', .-:-=: ,-,,-v-, .._._, !l .•_,,_r,_, assist. 3 s, append.i Y. :-:>l( append i.xx Chapt. 7 Chapt., C) w In b-oduction;¥. 1S-3() 4 8-16 6-10 9JBTOTPL 2?iLAF:IES: EstifTBted Typing Des.l.gn Center-layout 20,000 :5,C)():) Art ~.l.AJr-k F't-inting 3(J,()():) ~'5,(:(lO ::0:,000 30,000 35~C(X) :~:5,OOO 2;~)~OOO 50,0(:0 61 ~500 6,::::'()O 1 ~500 2;.0,000 ::..(),C(X) +r i.. w ,-"I t-='t-'''-n 3 4 c:- Chapt. Chapt. Chapt. 40,000 40,000 ":"1'::,,_1 Men:. Cost ::::.s~5C() 6-'E~ '-;'9.< Chapt; , 4 Chapt. 5(>-8() Cost 2,5<):) i ,:7C(> 23,400 17,400 47~80(l 4,6~)(i _5,8:)0 1,9X) 10,100 10,1(:(> 2~500 7,700 ::0,000 5,7(x) $·143,tJ.)() $Z:S , ooo 2(l~020 20~(l=~) ·15,Lj.{X) 9 ~tJ.)O C051: 8~OOO $ 93,C(¥.) F'e.-scJ.itiel CG5t::; l!··~0?n2 (:;:~3t:i.iT:2:(:'2(::l [J'/ H::-f3,fET and ("h.Ils aftett3.1king tirTK? '=lctu.=tll",-.' '3p==nt It-:orking on the pub l Lce t i.on . Tha:::,::?individuals ~'Jith the aut.ho~-s, for the ~',ork.ing en the Steering f-l'':O'./ET ha.d kept c.i i:c:r 2:::.ti.fTk3.t.ing the· ott-iErs. part: , log �47 APPENDIX D Description of Management Areas Dodd Bridge, Brush, and Cottonwood SWA's C H GR S - tJG - D NV L R - COTTONWOOD HAYMEADOW GRASSLAND SHRUBS AGRICULTURE DEVELOPED NON-VEGETATED LAKES RIVERINE SIZE CLASSES I - under 6" dbh 2" - 6 - 16" dbh 3 - 16 - 3O"dbh 4 - over 30· dbh CROWN DENSITY A - 10 - 35% B - 35 - 55 "I. e - 55 - 100°J(. Fig. 1. Vegetation map, Cottonwood State Wildlife Area, from 1979 aerial photograph. �.j::'- 00 C H GR 5 AG o NV l R - COTTONWOOD HAYMEADOW - GRASSLAND - SHRUBS - AGRICULTURE - DEVELOPED - NON-VEGETATED - LAKE - RIVERINE SIZE CLASSES I - under 6' dbh 2 - 6 - 16' dbh :3 - 16 - 30' dbh 4 - over 30' dbh CROWN DENSITY A - 10 - 35,},. B - 35 - 55 % C - 55 - 100·" Fig. 2. Vegetation map, Dodd Bridge State Wildlife Ar~a, from 1979 aerial photograph , ,/ �4..IWW 49 Page S PLATTE COTTONWD H GR 1 BERRY r:: -e.....J .•• .:;, 10·~.(I 164.5 r' . C2f~UE C:28 CJ~BUE ·:2C ~::2ClJE .-":-,,"\ I~·~':-I 21. 7 18.0 64.8 14.5 2.9 4.6 - :::::5.4 ...•.• &~ •• 8.2 .~C:.8 L::. :J .34. '=J ~ .~.7 .-:::.. .. -I ,-~ r ", J"t.l\,'·1 . .,.. ...• ._;. I :::5. ~ .,0 c• •• ;:, ::::.::. .• 7 .L ::~ oS -3 ~:~• :3 ~j • 1.s • 7 , i \).~ ..~) ·lS. i .:".i- .~~ •. c.sc C:3CUC: C4A C.:'\·AUE C4B C4BL.:E C4C C4CUE R NV L 69.3 66. 1 89.7 102. 1 C.::::(~U~ C3B '::~~.8:_.~E 111. 4 28.2 CHARTIER 24.0 Q PJ AG D C1A CH\UE CIB CIBUE CIC CICUE C2A DODD BR BRUSH " '1' 14.5 7 .. ~. 1..2 41. I) 8 '') • .L- 10.9 42.0 96.7 ,.'?' ,..... ":_.j • .::.. 8.9 8.3 vi TOTAL C"C'I""'\ C'" ..J..J..:,. • ..J -:0'1"""'\ •• ·_..,..:..4 61. 9 485.9 658.6 -", .. ,...•... 64.8 , .. .. -- _;_. ," ':~. �50 Page HALE PO S REPUS BONNY J MARTIN GR 332.6 S PJ 21.7 66.1 543.9 17.6 494.7 3088.8 725.1 603.3 10473.1 3217.5 180.6 176.6 177.4 AG 44.9 614.1 199.3 178. 1 740.6 46.7 46.2 116.9 2.0 13.7 28.2 7.9 46.5 15. 1 19.0 20.8 -9.9 10.8 18.9 192.4 21.9 .' H D . 1 CIA CIAUE C'_B CIBUE C1C C1CUE ['2A C2AUE C2B C2BUE C2C C2CUE I""'~~"\ ~.':t"'1 "? ..., . .a:.,. • "'- 18.3 79.4 • ,1 .1. • ..., :-':::4. (~ ::5 ...l •• RIO GR 407.7 21.9 90.5 i.'J. • '"' "J 7 15.5 54.6 9.5 46.3 4·3.4 4c)·.:l. ~) . .:.•• t:$ PURGATOR ,,-..-, ..!..:. " • r= .J. 1""'\ I.;) "'7 I C3AUE .'-.'7~ ._, ..,:-.0 2~:'.9 81.3 ~,",:.C" 4-J • ...:.. ~. ';'7.5 95.9 21.2 47.8 21.8 64.8 1.8 10.1 83.0 847.1 942.8 C::::'BUE C3C- 45.8 2.8 C3CUE C4A C4AUE cos C4BUE C4C C4CUE R NV L -. ...::. • C" ....J 13.8 6.8 1724.7 28.4 1277.0 6954.0 22214.7 ..,.. c: W TOTAL ,..J 186.0 824.8 5444.4 4.7 .J. 544.3 '" • .:". ...... .;. ":, ::.~.:;':~ .;,~.j~~~;~I···<;\\4;.i'-l·;';~~:.:;. : J • ;;, ..~.:,:t~t �erIe: DOW 51 Page 1 ROCKY FD LUSK DOLOR R ESC ALA 118. I) 31.9 38.0 254.9 56.0 57.4 3014.6 AG D CtA CIAUE CIB C1BUE e1C CICUE 111.3 6.7 1.0 204.1 1. 1 4. 1 37.1 60.2 413.5 570.2 79.6 C2PI 6.4 H GR S PJ 29.9 WALKER 1.1 70.4 251.8 3C)3.2' 58.8 15.6 34.7 108.0 3.1 13.8 C' ,J./ - 44.3 '-.""':'" • ...J ..;;..~ 70.7 37.8 C' C2~ilJE C28 1·.L • ., -r C2EiUE C2C (:2CUE C3i~ 24.1 14.2 ~S4. 8 26~2 12.1 l'0 • ~, ..::. '!::" ~-' .• , -7 30. ,:;. 1. i~ C3AL;E C38 39.0 i s. 1 24.6 1(;.8 9.5 9c).3 34.2 C:38UE C3C C:":;CUE C4A C4AUE 1.5 4. 1 C4B C4BUE C4C C4CUE R NV L 8.9 6.0 20.2 1.1 21.2 2.9 4.6 1.9 ~'J TOTAL . 120.3 29.9 115.2 350.0 631.7 1478.2 3665.1 755.0 ;", , . r, ...... i:I~ �52 Pagl! 1 COTTGN~ilOD SOUTH PLATTE STAND I TYPE ~CRES TYPE 1 C2AUE 6R 3.3 15.5 32.1 14.9 8.1 R C3B C3AUE 2 3 4 5 6 7 H GR L ACRES TYPE ACRES .8 R CIA S CIA 10.9 9.6 24 12.1 5.3 AS '33.5 47.1 .5 3 .1.8 40.7 36 2.2 16.7 7.3 7.4 H H C3A H C2B 39 50.7 36.3 13.2 .9 ~.3 C3A CIA C2B C3C C3A 10.:3 AS 7.4 C:~~E 1S ..2 12 ~.6 10.3 .., C1~ AS R C3~ C3i1 C3A C3~ 15 111 17 18 TYPE H 8 13 ~4 ACRES SR 9 10 11 12 R BRUSH BERRY C3~ H C"'''c 1:-t'.; ••• !'lli ~.'" C2ri en . •• R L 3.2 b 55.4 2.9 12.3 .s .4 C3~ j*'~.' I ..,.,;11 t9 ;:: [23 ",' 21 ... C::U£ b.7 j"'A,\ C2t) !:.1 • 7 2.6 •& B.l C3C 5.b ., " 1...1.. 8.3 3.4 "T " 1.1.. ~~n w~,:o b.~ C3B 17.5 C3B 22 L AS .8 4;).8 C3C 2.9 .9 27 28 29 BR GR 13 1·;.8 2.4 30 AS 5R •• 'J ..,., 23 C33 C3B 24 25 •• I.. 26 31 32 33 34 35 36 ." 110 6R ~'n C3AUE L L H sa 59.9 C3~ ~5.8 1.6 H L C2A C3~ C3B C2A L 37 38 39 40 H C38 R C3B C4A C28 C3A SR 41 42 43 44 45 TOTAL 552.5 32.2 61.9 .6 2.9 19.9 1.8 40.2 .7 4.7 53.1 4.9 3.S 5.7 18.2 2.4 1.3 .8 5.2 3.8 1.2 8.3 485.9 �53 Piqe CHARTIER DODD BRIDGE STAND I TYPE 1 R 2 3 C3AUE C2BUE 4 5 0 7 8 9 H C4B C2A C3A H I' .1 C3A C2A C33 12 ~V 13 t4 C2BU£ II) 15 16 17 18 19 2') 21 22 23 24 25 26 27 28 29 30 31 32 33 34 3S 36 37 38 39 40 N~ ACRES 94.0 10.4 8.3 3.4 1.2 30.4 27.0 12.9 12.1 8.3 ~ 1.0 I". 4.7 C3~ [3aUE u.s C3A~E H H C3AUE SR ~w as. ·1 7.~ 2.b 1 C3A· 6R C3B C~ NV C38 GR C3A MV C2B 41 H'J 42 43 44 4S R C3AUE 04.8 n.B 3.4 27.5 3.5 7.5 29.3 37.4 14.B 17.4 9.1 23.9 9.2 6.3 2.9 34.9 1.S 2.1 5.6 NV AS 4.1 1.1 15.7 It sa ACRES 16.'1 ".I.;), C33 C3BUE TYPE 11.5 CZA BR H 1 658.6 64.S ��APPENDIX E COTTONWOOD 55 RIPARIAN ZONES OF COLORADO· .. , ': :-~~.!-.::--. Colorado Division of Wildlife ��57 "To keep every cog and wheel is the first precaution of intelligent tinkering." Aldo Leopold - A Sand County Almanac COTTONWOOD RIPARIAN ZONES of COLORADO Gary C. Miller Richard D. Sherman Jim Houston Colorado Division of Wildlife Colorado Water Workshop Gunnison Colorado 30 July 1986 �58 I. INTRODUCTION The word "riparian" (from the Latin ~iE~ri~~, meaning the bank of a stream) seems to have virtually exploded into the western lexicon over the past 15 years. That riparian zones carry high public interest, whether from the standpoint of flood control, recreation, aesthetics, beef production, or wildlife, is evidenced by the many recent federal, and state regulations and local ordinances (such as in Boulder and Greenwood Village) governing their use (see Johnson et al. 1985). The term carries various connotations in different fields of endeavor, however. The objective of this paper and the accompanying discussions, then, is to increase the effectiveness of communication between practicioners of diverse disciplines ...the following pages summarize what the Colorado Division of Wildlife and others have learned in recent years regarding certain of these areas--cottonwood riparian zones-- which are so vital to the state's wildlife. Riparian zones, in the strict sense, are vegetation complexes which exist because of surface or subsurface inputs of water associated with permanent or ephemeral streams. In Colorado, the ones of greatest direct significance to the state's overall wildlife species richness, and the ones receiving greatest attention in recent years, are those forested with cottonwoods and willows. That is not to imply a lesser importance of those riparian zones of grasslands, shrublands, or coniferous forests--after all, the cottonwood riparian zones depend upon the proper functioning of those others for the needed inputs of water. Acknowledgements.-Division of Wildlife research reported herein was supported in large part by voluntary contributions of Colorado taxpayers (Nongame Checkoff Fund) combined with excise taxes collected on hunting equipment (Pittman-Robertson funds). W. D. Graul supplied some of his unpublished data and reports. �59 II. ECOLOGY .Many of Colorado's most important riparian zones are best recognized by their cottonwood (Populus spp.) and willow (Salix spp.) trees. These trees, and accompanying grasses, "weeds", and shrubs comprise a vegetation aS~Q£iation which supports the most diverse and productive natural commgnity of living organisms in the state. The non-living features of riparian areas (water, soils, climate, atmosphere, and history) combine with this community to form the cottonwood-willow riparian ecosystem. Classification of cottonwoods is not easy. The species belongs to the Salicacea~ or willow family--beyond that, it is difficult to find agreement among experts as to classification. Plains cottonwood is afforded full species status (P. ~argentii) in some references, while others classify it as a variant of the eastern or common cottonwood (P. deltoides var. occidentalis). Likewise, confusion over the "correct" classification of the Rio Grande cottonwood of the western slope exists ...they may be P. wisliz~ni, or P. fremontii, or P. fremontii var. ~isliz~ni, again depending upon the authority consulted. Other cottonwoods, named Fremont cottonwood and lanceleaf cottonwood are reported to occur in the state. The latter species may in fact, be a hybrid of plains and narrowleaf cottonwoods (P. angustifolia). Three cottonwood-willow vegetation associations may be said to occur in Colorado's forested riparian zones (Fig. 1). Plains cottonwoods dominate lower reaches (generally below approximately' 5,000 feet elevation) "of the South Platte, Republican, and Arkansas drainages in eastern Colorado , while Rio Grande cottonwoods are the dominant feature of wooded riparian areas along the lower portions of the Yampa, White, Colorado, and"Gunnison rivers (scattered stands of Rio Grande cottonwoods also occur along the Rio Grande River). Narrowleaf cottonwoods are sympatric (co-exist) with the "broad-leafed" cottonwoods in a tran-sitional zone which may occur between roughly 5,000 and 6,000 feet elevation. Most cottonwood - willow associations of the riparian zones above 6,000 feet (but only occasionally exceeding 8,500 feet) are dominated by narrowleaf cottonwoods. Cottonwoods are tall (often 50, occasionally to 90 £eet at maturity), large (occasionally to 6 feet in diameter) trees of stout, spreading limbs. In Colorado, they often occur in almost pure, evenaged stands where water table depth does not exceed 12 feet. They are an early-successional species, which do not reprodUce well in deep shade. Many seeds are produced each year and dense "nurseries" of seedlings may become established where conditions are suitable. As the stand of trees becomes older, it thins naturally until the mature cottonwood stand becomes open and savannah-like (perhaps 10 trees/acre). Under natural circumstances, a new cottonwood stand will not become established until the old stand is fairly broken up, perhaps at 50 years of age, although trees may live to 80 or 90 years under very favorable conditions (Sudworth, 1934; Read, 1958). �60 1, , \ , I I ,,L , •• ••• Pueblo 86fo:L: City I , I I I I , 7500' -- Plains ••• Rio Grande Cottonwood CottonwOOd _ Norrowleof Cottonwood LCOLORADO -- ----- - --- Fig. 1 Distribution major Colorado ---- ---- ----- of cottonwood-willow drainages. - vegetation - ----........_---- associations along . , �61 • The cottonwood riparian zones are vegetated with much more than cottonwoods and willows. In study areas along the lower reaches of the South Platte and Arkansas rivers, trees such as red ash (Fraxinus ~~nnsyslvanica var. suhintggmn~), russian olive (Elaeagn~§ angustifolia), and tamarisk (Tam~~ix gl~ca) are common in places. Shrub cover, primarily snowberry (SYmphoricarpos sp.), is highly variable (ranging from 0.3 to over 60% cover), and most of that cover (generally more than 90%) occurs within 20 inches of the ground. Grass and forb species most commonly occurring along the lower South Platte are wheatgrasses (Agropyron spp.), cheatgrass (~~omus tectQrum), switchgrass (Panicum yirgatum), Prairie cordgrass (~artina pectinata), and a tall "weed" called Cardaria (Cardaria sp.) The cottonwood riparian zones are much more than a homogenous forest of tall trees along a streambank. Within the forested areas, discrete stands forming varying combinations of tree density and age can be identified. Non-forested areas, such as sub-irrigated-meadows, sand bars, and sloughs, are also important to the maintenance of the ecosystem. The Colorado Division of Wildlife, in conjunction with the Colorado State Forest Service, has developed a method to describe these cottonwood riparian zones using over 20 structural classifications (Miller, 1985) (Fig.2). COTTONWOOD STATE WILDLIFE AREA VEGETATION MAP C H GR S fC, - COTTONWOOD HAYMEADOW GRASSLAND SHRUBS AGRICUL HiRE o - DEVELOPED NV - NON-VEGETATED L - LAKES R - RIVERINE SIZE CLASSES I - under 6· dbh 2 - 6 - 16· dbh 3 - 16- 30· dbh 4 - ever 30· dbh CROWN DENSITY A - 10 - 35% B - 35 - 55 C - 55 - 100% ·1. .1 Fig. 2. Example of a cottonwood riparian zone mapped with the classification system used in habitat management by the Colorado Division of Wildlife. �62 • The cottonwood riparian zones of Colorado are undergoing major changes in many areas, especially along the lower South Platte and Arkansas Rivers. For reasons as yet unclear, new cottonwood stands are not developing fast enough to replace those being lost due to old age. There is much seedling "nursery" establishment, but young trees are not surviving into the older age classes, as demonstrated by Miller's (1984) study of great blue heron nest sites (Fig.3). In fact, Miller (1985) has. demonstrated that up to 75% of the cottonwood forests on his study areas near Fort Morgan may be treeless by the year 2000. Many hypotheses have been advanced as to the cause of these changes--grazing by domestic cattle and wildlife, changes in water regimes, lack of periodic flooding, and soil accretion on floodplains--but none have yet held up to serious analysis and the cause remains a mystery. - o o ~ ACTIVE COLONY INACTIVE COLONY ALTERNATE STANDS (1,104) STANDS (1,062) STANDS (4,807) 50 C/) w w ex: •.... u.. 40 0 •.... z w 30 27 0 ex: w o, 20 10 DIAMETER BREAST HEIGHT (mm) Fig. 3. Size (age) distributions of cottonwoods discovered during statewide great blue heron study. Note the very high proportion of seedlings (0-25 mm), but the extremely low percentages of trees in the 26-150 mm intervals. Obviously, as the larger size classes die out, there will not be enough trees to replace them (from Miller 1984). �63 WILDLIFE • Although cottonwood riparian zones make up only an estimated 0.2% of Colorado's area, 264 (43.4%) of the state's 608 terrestrial vertebrates (reptiles, amphibians, birds, mammals) have been recorded there (Graul and Svoboda, unpubl. ms. in Miller, 1985). A study of the lower South Platte River showed that 209 (34.4%) occurred in that thin habitat corridor. Approximately 50% of all bird species in the state occur there (Bottorff, 1974). Although some of the species found in riparian zones ffiQY be able to exist in other areas, a number of species appear to be closely linked to cottonwood riparian areas for at least part of their annual life cycle. Among the more familiar wildlife species which are clearly linked to the lower-elevation cottonwood riparian zones are: White-tailed deer, wood duck (both fairly recent arrivals), mink, bald eagle, great blue heron, Rio Grande wild turkey, and northern bobwhite. Other species, not as well-known but perhaps no less important to the health of the ecosystem, are red bat, spiny softshell turtle, yellow-billed cuckoo, and upland sandpiper. Our knowledge is not nearly so complete for the narrowleaf cottonwood riparian zones, although we do know of their great importance as elk and deer wintering range, sage grouse brood-rearing areas, great blue heron nesting sites, and bald eagle winter roosts. Beaver, goshawk, sharp-shinned hawk, willow flycatcher, yellow-bellied sapsucker, warbling vireo, and 3 species of warbler may, in some cases , be among species closely associated with these areas. Within the riparian zones, animals do not do equally well in all of the different vegetation categories described. Upland sandpipers depend almost exclusively upon the sub-irrigated meadows. Wood ducks must have the mature to overmature cottonwood stands for their nesting cavities as well as sloughs for rearing of young. Yel19w-billed cuckoos generally nest in shrubs or small trees fairly close to the ground. Wild turkeys need mature cottonwoods with little understory for roosting sites, but dense understory areas for nesting and areas of younger (shorter) trees for brood-rearing areas. Warblers are classic examples of species which are specialists in using particular habitats--some forage in dense understory shrubs while others use the outer portions of the uppermost forest canopy, and still others forage on branches close to the trunk of the tree. �64 III. MANAGEMENT • Knowledge of the vegetation associations and the wildlife of the cottonwood riparian zones allow us to manage these areas to benefit some wildlife species while eliminating none--following Leopold's first precaution of "intelligent tinkering." Habitat management activities by· the Colorado Division of Wildlife follow precepts laid out in a book, published jointly by the Division of Wildlife and the USDA-Forest Service, titled Managing Forested Land2 for Wildlife (Hoover and Wills 1984). Although that book was aimed primarily at coniferous forests, it provides a system by which certain species' habitats can be optimized without violating critical levels of other species (as a general rule, one does not improve conditions for certain species without having some sort of detrimental effect upon others). With few differences, the conceptual framework for the process was described as an ecosystem management approach by Graul and Miller (1984). At the heart of the system is the need to identify and address the needs of ecological indicator species, also known as stenotopic species (steno narrow). These are species which have very stringent life requirements, or narrow tolerances, such as the upland sandpipers or wood ducks mentioned in the previous section. Only after the needs of the ecological indicators are assured do activities designed to optimize conditions for emphasis species take place. Emphasis species are those which carry high public interest regardless of their ecological role--these are often threatened, endangered, or game species. In some cases, lists of ecological indicators and emphasis species may overlap substantially--of 10 ecological indicators identified by Miller (1985) for his South Platte study areas, 6 were also potential emphasis species. = The process is graphically illustrated in Figure 4. Ten possible habitat conditions are represented along the X-axis. The bars (A through D) indicate conditions in which each of the 4 species of the Y-axis can exist. By creating conditions in which species A and B mgst exist, we also have conditions in which species C and D can exist. Conversely, if we manage for species C and D, we have an 80% chance of failing to provide for species A and B--and failing to meet our primary objective. I . D w C o w 8 (J) 0.. (J) A 1 2 3 4 5 6 7 8 9 10 HABITAT CONDITIONS IN WHICH SPECIES OCCUR Fig. 4. A simplified ecosystem with 4 species and 10 habitat conditions. Horizontal bars indicate conditions suitable each species (after Graul and Miller 1984). for �65 f In certain circumstances, an area may be managed with a ~ingle-§Eecie§ management approach (as opposed to an ecosystem management approach). This approach seeks to maximize habitat conditions for a single species, and therefore carries the greatest risk to other species. Hence, this approach is generally used only to meet particular species objectives--management for a threatened or endangered species, for instance, or, more commonly, to provide critical winter range for deer or elk. The single-species approach is rarely applied to the low-elevation cottonwood riparian zones. Even where this approach must be used, care must be taken to assure that other species' requirements are met elsewhere--to at least maintain "between-habitats" or "geographic area" species diversity (Samson and Knopf 1982) ..- Whichever approach is used, vegetation is managed to change its structure, and its value as wildlife habitat. Knowledge of the vegetational structure of the area (Fig. 2) is coupled with knowledge of ecological indicator and emphasis species' requirements to-develop habitat management standards. Examples of such standards developed for low-elevation riparian areas in eastern Colorado are given in Table 1. Table 1. Example of ecosystem management standards for low-elevation cottonwood-riparian areas in eastern Colorado (adapted from Miller 1985). Step 4. a). Main~enance of at least 20% of the area in forest, structured to provide sustained yield of 20 habitat types, and arranged in forested patches of at least 25 acres. b). Maintenance of mesic haymeadows of 400 acres in area, or clusters of smaller meadows (each at least 25 acres in area) which total 400 acres within 15 mile radius. c). Major alterations (type-conversions) in habitat types Cl, C2, C4, or S. are not permitted Step 5. b) Cottonwood stand establishment to take place in existing C3A, C3B, or H habitat types. Step 6. Future habitat acquisitions to evaluate inclusion of xeric or upland shrub habitat types ...to serve as refugia during periods of deep snow accumulations or flooding. �66 . The preceding standards were developed for areas controlled by the Colorado Division of Wildlife, with wildlife benefits the sole purpose. !n many areas, however, wildlife benefits may be only one of several values for which a cottonwood riparian zone is managed. In such cases, standards may be quite different. In any case, the techniques used to attain the desired habitat configuration may include: "Doing nothing," burning, mowing, cutting, plowing or rototilling, planting food or cover-producing plants, irrigating, periodic grazing or elimination of same. LITERATURE Bottorff, R. L. 1974. Cottonwood Birds 28(6): 975-979. CITED habitat for birds Graul, W. D. and G. C. Miller. 1984. Strengthening approaches. Wildl. Soc. Bull. 12(3): 282-289. in Colorado. ecosystem Am. management Hoover, R. L. and D. L. Wills, eds. 1984. Managing forested lands for wildlife. Colo. Div. Wildl. in cooperation with USDA Forest Serv.,Rocky Mountain Region, Denver, Colorado. 459 p. Johnson, R. R., C. D. Ziebell, D. R. Patton, P. F. Ffolliott, and R. H. Hamre, tech. coords. 1985. Riparian ecosystems and their management: Reconciling conflicting uses. First North American Riparian Conference. USDA Forest Servo Gen. Tech. Rep. RM-120. 523 p. Miller, G. C. 1984. Population surveys of selected bird and mammal species in Colorado. Job Prog. Rep. N-1-R(W-136-R). Colorado Div. Wildl., Denver. Miller, G. C. 1985. Habitat investigations report. Job Prog. Rep., 86-039-01. Colorado Div. Wildl., Denver. [64p] Read, R. A. 1958. Silvical characteristics of plains cottonwood. USDA Forest Service, Rocky Mountain Forest and Range Exp. Sta. Paper no. 33. 18p. Samson, F. B. and F. L. Knopf. 1982. In search of a diversity ethic for wildlife management. Trans. N. Am. Wildl. and Nat. Resour. Conf. 47:421-431. Sudworth, G. B. 1934. Poplars, principal tree willows and walnuts Rocky Mountain region. USDA Tech. Bull. 420. 112p. of the �67 c IV. SELECTED REFERENCES Graul, W.D. and S.J. Bissell tech. coords. 1978. Lowland river and stream habitat in Colorado: A symposium. Univ. Northern Colorado, Greeley, 4-5 October, 1978. 195p. Great Plains Agricultural Council, Forestry Committee. 1979. Riparian .. and wetland habitats of the Great Plains. Proc. of the 31st Annual Meeting. Great Plains Agricultural Council Publ. No. 91. Johnson, R.R. and D.A. Jones, tech. coords. 1977. Importance, preservation and management of riparian habitat: A symposium. USDA Gen. Tech. Rep. RM-43. 217p. Madson, C. and L. Lahman. The death of a river. (non-technical article) Audubon 84(3):70-85. Swanson, G.A. 1979. The mitigation symposium: A national workshop mitigating losses of fish and wildlife habitats. USDA For. Servo Gen. Tech. Rep. RM-65. USDI Fish and Wildlife Special Research Prairie Wildlife Service. Report. Research on 1981. The Platte River ecology study. U.S. Fish and Wildlife Service, Northern Center, Jamestown, North Dakota. 187p. ��69 APPENDIX F A Report on Research Done on the Upland Sandpiper (Barlramia longicauda) in northeastern Colorado during the Summers of 1986 and 1987 by David C. Bolster" Department of EPO Biology P. O. Box B - 334 Univer sitv of Colorado Boulder, Colorado 80309 "In cooperation with the Colorado Division of \\' ildlif e . �70 This report summarizes the work done on . Upland Sandpipers (Bartramia t"\\7(1 populations of longicauda) in northeastern Colorado during the latter half of the summer of 1986, and throughout summer the of 1987. The main aim of the research was to document the status of these populations. the habitat and to try to identify what it was about available to and used by these birds which allowed them to remain so healthy and viable relative to the rest of the Upland Sandpipers in the state.' This work constitutes the field wort habitat I anticipate approximately doing for my Masters half of Degree on the My field use and breeding biology of Upland Sandpipers. work during the 1987 season was supported, . both financially and INTRODUCTION In 19135. the Nongame Advisory Council of the Colorado Division of \\~ildlife ranked the Upland Sandpiper "Species of Special Concern" (Winternitz ~1 (tied.! out of 4(l·avian and Crumpacker 190)). They cited "historic population decline due to tallgr ass prairie habitat loss" as the apparent reason for this ranting, although they indicated that this cause and effect had not been proven. species has a small and declining population there is inadequate information, They noted that the in Colorado, about which and they considered it to have special ecological value as an indicator species. To date. no stud)' has ever been published on this species in Colorado. This preliminary report wil! indicate the work done so far in investigating use patterns of the Upland Sandpiper 1 in northeastern the habitat Colorado. as �71 well as the intended direction of research, to be conducted summer of 1988. Any and all suggestions in the on how this study can be improved would be appreciated. STUDY AREAS The habitat in Logan, Sedgewick, South Platte river valley in northeastern cropland. rangeland, woodlands scattered and Weld Counties in the Colorado is a mosaic of towns, and a narrow strip of riparian along the South Platte River. Upland Sandpipers spend the summer in open areas composed .in mixed amounts of range and cropland. although I infrequently isolated farmhouses. saw them on grassy "lawns" around I never observed Upland Sandpipers on mudflats along the shore of either the river. or any of the ponds or reservoirs, in my study areas, which is corisistant with the published accounts of their behavior and natural history (Bailey and Niedrach 1965, Bo",-en-1976, Buss and Hawkins 1939. Higgins and Kirsch 1975. \fhite 1983, among others). In the Crook -jule sb urg Reservoir area, four parallel farm roads leading due north from the edge of the South Plane River were chosen as replicate transects through the area where. in 1986, preliminary observations Sandpipers. Because many of the pastures spanned 1\\"0 of Upland and corn fields in the area entire mile square sections, each road chosen was at least miles apart, thereby types. had located the largest numbers avoiding duplicate sampling of some habitat The sampling area along each of the four roads extended approximately 2.5 miles (4.5 kilometer s i north from the edge of the 2 �72 riparian wood edge along the river: thus in some cases the transects actually began some distance from the river's edge. In Weld County. my sampling was conducted 49), beginning along Kersey Road (\'reld County Route 1 mile north of Interstate Hudson, and extending approximately tracts 76, east of the town of 15 miles north to the railroad 1 mile south of Colorado Highway 34. west pf the town of Kersey, DATA COLLECTION Observations were made at .25 mile \,45 tm', intervals along the roads. starting 0.1 mile from the riparian wood edge. I'pon stopping, the general habitat category of the fields on each side of the road was recorded. Habitat categories were as follows: bare ground (bg I; heavily grazed range land (rh): moderately . grazed range land (r m '1; lightly grazed range land trl): old fields, often partially overgrown with weeds {of); densely growing hay fields (na): rows of planted corn or sorghum. shorter than 28 cm (cs): rows of corn or sorghum, taller than.28 cm (ct l; freshly cut alfalfa fields. where the alfalfa was still lying in parallel rows drying (ac i: recently cut alfalfa fields. where the dry rows had been baled but not yet collected (ab); short alfalfa, less -than 28 ern (as); and tall alfalfa, taller than 28 cm (at). The short-tall cutoff of 28 em (11 inches) was chosen because that is the height of an alert, heads up Upland Sandpiper. Vegetation taller than 28 em would be too tall for the birds to effectively for danger. ... ) scan �73 At each stop the fields on both sides of the road were scanned for approximately two minutes using l Ox binoculars, in an attempt to locate any Upland Sandpipers recorded. present. The nu mber observed was Then a 100 meter by 50 meter rectangular axis perpendicular (long to the road) was walked through the fields on each side of the road. This transect-flush method was used to increase the Iikelihood of finding Upland Sandpipers vegetation, transect in dense or to flush any birds which, because of their cryptic coloration, tiad gone unobserved during the roadside scans. The nu mber of birds seen within approximately the transect 75 m (visual estimate) route. and the number of sandpipers during this process, were recorded separ atelv. behavior of the bird when it was first observed of heard but not seen In addition. the (i.e. feeding, resting, preening. on alert. etc.l v•.as also noted. Fer those birds which were observed in some stationary location ii.e. not already running awayi a detailed habitat analysis was made of the precise site where they were first seen. In some cas~s this was done right away; in others the site 'was marked or otherwise "\\"i111 48 identified, and the analysis was done later on. but always hI'S. of the observation. following measurements: vegetation observation; These habitat analyses included the the range (minimum in a 3 m diameter circle centered the average vegetation was grass or other monocots: the percentage litter. on the point of height within that circle; the percent cover in that circle; the percentage percentage and maximum) of the of that vegetation which was Forbs: the which was shrubs; and the percentage Then, a vegetation which profile was measured of twigs and leaf along a randomly :. �74 oriented of the 3 m circle: diameter the line, the vegetation height was recorded, (grass, forb, shrub, etc.). nearest emergent at every rock, fencepost, randomly chosen location within and again in the afternoon. around made daily between from about (usually around in 1986 indicated . spend the hottest shade of a dense clu mp of grass or shrub. behavior 1986 also indicated although precise part of the day loafing, thev'. appear to engage it was not clear whether IOC:llicJ11s throughout at dusk almost always . morning. It seems unlikely great distance birds I can not be certain areas where behavior Preening is the only active this time .. Observations . stop moving around Fields having in dust. in those sandpipers in in them the following individually ofthe marking roadside day, and on morning surveys. the been observed, lite good habitat 5 when I spent time either that they might be copulating or in areas that looted in the meager of this. birds had recently indicated Sandpipers that the birds would fly or walk any portion was not doing the regular usually had sandpipers in the dark. but without During the middle . 10:30 A:\.I), or not the birds remained the night. - of the day. that Lpland in during that the sandpipers dawn 3P:\1 until dusk (usually 7:45 Pxl), to coincide with the cooler pans Preliminarv . observations them patch, no less than the bird was observed. 6 A:\J) and mid-morning (approximately point was the same habitat were to the in full at some other but no more than 50 m from where surveys along of plant type in meters or other vantage This process was then repeated These roadside regardless Finally, the distance recorded. 10m 10 em interval especially in if their or nesting for sandpipers. I nearby, These sites �75 were not necessarily along my regular roadside routes: were composed of uncommon habitat or vegetation usually they types not found on my study areas. Only rarely did Ifind Upland Sandpipers of these sites. Thus some of my observations on an)! on behavior, habitat usage, etc. were not actually collected along the -4 roads mentioned previously. Samples of all the dominant plant species in the different habitat types were collected and pressed for future identification: . facilitate this identification, To multiple specimens were collected, especially of different phases of f lowering and/or fruiting. Similar looting plants were often collected more than once, to insure against lurn ping or! sister ""'PC1'e" apns~ were also ';"~I,l_! ~.L~•• ~!.'_ ~. Photogr 1 l • "'t' ,~. _~\...tal-en of l all 1 characteristic habitat types. RESULTS A total of 380 observations during my roadside surveys. nu mber of observations of Upland Sandpipers were made The Following table (Table 1) 5hO\\'5 the made on those surveys. Observations are broken down according to habitat type, and by road, for the whole of 1987, Seasonal totals for each road and habitat type are also given. Note that the observation period was divided into 3 roughly equal periods; from early May through and 22July until 18 August. 14 June, 14 June through 21 July, I believe these time blocks correspond to the on-nest phase, the chick rearing phase, and the pre-migratory phase of the Upland Sandpiper's breeding season in 1987. �76 . TABLE! - Cumulative Road 85 Phase Habitat Bare Grnd Range Hvv Range Med Range Lt Old Field J roadside data. 1987. survey Ker sev Road 89 Road 93 2 3 1 2 j" 0 7 1 11 0 0 0 27 0 6 0 2 4 0 44 Road 97 "') 1 2 0 3 0 0 1 0 1 0 7 4 0 4 1 14 .., o 18 10 4 5 7 9 9 1 2 9 9 4 9 8 107 8 0 1 18 2 1 0 1 14 O· 0 0 0 0 46 .., 3 2 0 ":> . I 2 Corn reu 2 0 0 0 0 o 0 3 0 0 2 6 2 -0 0 0 16 0 0 61 0 0 0 0 0 5 4 o 0 0 70 0 4 0 0 0 0 5 0 2 .? 1_ 0 O· 0 0 26 o 0 0 0 0 0 0 0 0 0 0 n 0 0 0 0 0 0 2 3 2 5 0 0 0 0 0 0 0 13 0 0 -1 0 0 0 0 0 0 0 0 5 6 1 0 0 0 0 0 0 10 0 0 Cern Short Sum 2 Hay FieJd ., 1 ...• .) CUl AII' 0 Baled All' 0 0 0 0 0 0 Short _~.lf 5 .., .) Tall 26 :~?: _,;If 0 0 0 0 0 n 0 0 0 0 0 n I) [) 0 0 Phase Totals Road Totals 11 34 75 120 The number during the entire distinguishable. 6 39 18 35 25 7 63 of birds counted 67 37 30 26 using a non-parametric 37 93 on the different year was determined 6 13 18 habitat to be statistically equivalent to single types 380 .. �77 classification anova, the Kruskal- Wallis test (H Partitioning the summer into the 3 nesting-phase = 26.652, p < .01). time-blocks, the birds' use of the different habitat types was still statistically distinguishable (May-June at the .05 level. june-july and JUly- August at the .0 I level). The habitat types themselves Upland Sandpipers. comparison were used differentially according to a non-parametric by multiple test (Sokal and Rohlf 1981). In the first third of the nesting cycle. lightly grazed rangeland was used most often by the birds, followed by medium rangeland, and then heavy rangeland short corn (used equally often). and Bare ground. old fields. hayf'ie lds. tall corn, and all 4 types of alfalfa fields were hardly ever used by the birds. and were statistically similar to that observed indistinguishable. This pattern was in the latter 2/3 of the nesting season, although the Upland Sandpipers did appear to use significantly more . alf alfu fields-in late summer (pers obs.). Only 36 of the 380 (9.4 ';0) Upland Sandpipers observed the roadside surveys. were heard but not actually seen. during In only 3 of the 180 cells in the table above (habitat by road by nesting phase) was the number of birds heard larger than the number seen (in short corn, Road 93. in May-June; light range. Road 93. in july-August: in old fields, Road 97, also in July-August). The number seen was equalled by the number heard in only 5 of those cells. The vast majority of observations required during my transect-flush walks. and the birds to actually be flushed �78 DISCUSSION Although the habitat in the Kersey Road area appeared along its entire Sandpipers IS mile length. only 2 small colonies of Upland were ever located there. and they were close enough (approximately appropriately were observed 1 mile) to each other that they might more be called only one population. fairly consistently parallel roads. These transects In the Crook area. birds from end to end of each of the 4 . were not extended from the river because of the topography '. similar any further north of the South Platte River valley: approximately 2 miles north. of the river the land slopes quietly upward, rising roughly 18 meters in half a kilometer or less. Although this is no barrier to the sandpipers, upper fields, they quickly decreased and they did use these in abundance north of the rise. and were usually absent from the fields more than 4 miles from the river. It should be noted that, by that point, the land was rarely irr igate d. and the majority of the fields were dry range. This apparent affinity for the lower. more productive in the Crook area in particular, makes sense evolutionarily habitat types are analyzed individually. habitats. when the These birds are adapted to the Great Plains. and thus it makes sense that they would spend their time in moderately vegetated grassland, Light and Medium (grazed) Rangeland. cornfields through. are structurally such as those I categorized In the early summer. (short) similar to grasslands Because of the moisture and nutritious yet easier to move young plants growing there, they were always teeming with grasshoppers . other insects. and therefore . and . probably ideal foraging sites. Short or �79 recently cut (+ /- baled) alfalfa fields, again presumably the moisture and insect rich vegetation, also were attractive birds, especially in mid- and late summer. alfalfa was growing extremely was cut the sandpipers insects. because of During this period, the rapidly, and every few weeks when it moved in in droves to gobble up the exposed Although some of these habitats have relatively nu mbers of observations to the low associated with them. that is misleading. because of their very limited availability in space and/or time. Although I have not done the analyses. Ifeel confident that these habitats will prove to be important when corrections-are done for space and time. In addition 10 the sandpipers using field types which were similar to the grasslands they evolved in. they tended to avoid habitat types, such as tall cornfields. tall alfalfa. old "abnormal" fields. and bare ground. Bare dirt fields were used infrequently. often in the early to mid morning s ' and may have been an effective means of warning up after a foraging bout in the dew-covered Dense vegetation probably grass. (i.e. tall corn, alfalfa, and weedy old fields i is difficult for them to get through. and more importantly, impossible to see around in. I defined "tal!" as greater the height of an Upland Sandpiper . observations standing fully erect. in 1986 and 1987, I never once observed dense vegetation than 28 em. In all of my these birds in taller than they were. One habitat type barely used by the Upland Sandpipers Colorado, but frequently referred to in the literature, in was havf'ields. In my study areas. hayfields were never used by the birds once the gr ass got thick and tall. Prior to this. i.e. in the early summer. 10 when �80 these fields were shoruer) and patchy, they were used, sometimes extensively. However, during this period I considered structurally more similar to lightly grazed rangeland in the vicinity, and so categorized sandpipers' them. Because I am attempting categories to look at the response to differing patterns of vegetation well as actual use of that vegetation is to be expected. them to be structure as by humans. some overlap in I think it most appropriate to think of those areas I labelled "hayfields" as lying at the end of the heavymedium-lightly grazed rangeland continuum. It may' be that. because of the density of the haygr ass, Upland Sandpipers in Colorado prefer not to use it, possibly because they are too vulncr able to predation from coyotes, feral cats, and weasels. This still does not explain why they would be so strongly opposed to using it in Colorado, when they do in other parts of the country (see Ailes 1976; Bowen 1976; Buss and Hawkins 1939; Dorio and Grewe 1979; Orr 1943: and White 1983; among others) which presumably of the same terrestrial have many predators. ANTICIPATED RESEARCH FOR 1988 During the summer of 1988 I intend to continue the roadside habitat surveys in the same manner they were conducted during 1987. My major new approach to the study of Upland Sandpipers' habitat use will be to capture and radio tag approximately birds, and intensively throughout 12 adult monitor their daily and season movements the study areas. This technique seems to be ideally suited for such a study, because of the cryptic nature of the birds, ] 1 �81 and because of the wealth of precise data which can be obtained. Roosting birds, and later on in the season nesting birds, will be located by spotlight at night, and captured using a long handled "dip net" as described in Ballard and Bowen 1973. Birds will be banded with a U.S. Fish and Wildlife Service numbered weighed. and then individually aluminum band, color marked with plastic leg bands. They will also be measured, their fat load assessed, and checked for external parasites. transmitter Birds to be radio tagged will have a small attached to the basal portion of the shafts of their rectrices (tail feathers) with epoxy and/or thread, as -described in Kenward (1987). This method of attachment has been chosen because of the minimal impact it will have on the birds flight. and because these radios will be molted off prior to their long migration south in the fall. In the only other published account of radio tracking with this species (Ailes and Toepfer 1977),2 birds were mounted with backpack -style harnesses, 23 days, respectively. approximately 3.5 and monitored for 14 and Although those radio packages only weighed % of the birds' body weights. and they appeared to cause the birds no discomfort during the study, I believe that that weight may have been a significant impediment The transmitters I anticipate using will be roughly half the weight of the ones used by Ailes and Toepfer. approximately during migration. In addition. they should last 100 days, as opposed to Ailes and Toepfer's, which had a 40 day life expectancy. 12 �32 REFERENCES . -.•. . Ailes, 1. 1976. Ecology of the Upland Sandpiper in Central Wisconsin. :M.s. Thesis. University of Wisconsin, Steven's Point. Bailey, A. 1-1.and R. j. Niedrach. 1965. Birds of Colorado. Museum of Natural History. Ballard. \\7. B. and D. E. Bowen. Jr. 1973. Capture L pland Plover. Inland Bird Banding Denver Methods for the News 45: 132-135. Bowen. D. E. Jr. 1976. Colonialitv, Reproductive Success, and Habitat lnteractions of the Upland Sandpiper (Bartrami.~ longicaud'll. Ph. D. Dissertation. Kansas State University: Manhattan. Kansas. Buss, I. O. and .A. S. Hawkins. 1939. The Upland Plover at Faville Grove, \'\'isconsin. \\~ilson Bull. 51( 4): 202-220. Dor io. ~1. C. and A. H. Grewe. 1979. Nesting -- and Brood Rearing Habitat of the Upland Sandpiper. J Minn. Acad. Sci. 4S( 1): 8 -] 1. Higgins. K. F. and L 1V1. Kirsch. 1975. Some Aspects of the Breeding Biology of the Upland Sandpiper in North Dakota. \,'ilson Bull. 87i.lj: 96-102. Kenward, R. 1987. 'Wildlife Radio Tagging: Equipment. Field Techniques and Data Analysis. Academic Press, New York. Orr, E. 1943. The Bartr amian Sandpiper Bird Life 13:38-40. in Northeast Sokal. R. R. and F. J. Rohlf. 198 I. Biometry. Freeman and Co., New York. 2nd edition. White, R. P. 1983. Distribution and Habitat Preference Sandpiper (Bartramia longicauda) in Wisconsin. 37( 1): 16-22. Iowa. Iowa \\7. H. of the Upland Amer. Birds �83 APPENDIX HABITAT G CHANGES IN COLORADO Gary C. Miller Wild animal habitats are defined by the creatures which inhabit them. Since populations of those creatures are often quite mobile, wildlife habitats are likewise dynamic, and therefore unrealistic to measure. Fortunately, the vegetation associations of Colorado are somewhat easier to identify and measure, and often those associations carry a high likelihood of being habitat for particular species. Thus, we know that low-elevation (cottonwood) riparian associations are-potential habitat for 264 (43.4%) of the state's 608 terrestrial vertebrates (reptiles, amphibians, birds, and mammals) (Graul and Svoboda, unpubl. ms. !.!2 Miller, 1985, Job Prog. Rep. 86-039-01, Colo. Div. Wildl.). We also know that the habitats of greater and lesser prairie-chickens, and propably upland sandpipers are almost exclusively in the sandsage-bluestem habitat association (the association is also of prime importance to eastern plains deer herds). Pronghorn, swift fox and mountain plover habitat is probably in the grama-buffalograss association. Winter ranges of elk and "high country" mule deer have a high probability of being in the pinyon-juniper or mountain grasslands and shrublands. The latter association also serves as the principle habitat of sage grouse and Columbian sharp-tailed grouse. Thus, in order to assess the condition and trend of wildlife habitats in Colorado, we track the status of various vegetation associations. f METHODS The historical area of vegetation associations ("habitat") reported was gen~rally from the classification system used by Kuchl~r (1964, Am. Geogr. Soc~ Spec. Publ. 36) and the computations from CDOW's PNVPOLY computer program, on file in the Habitat Resources Section. At variance from those computations are the low-elevation riparian areas which combined Kuchler's (1964) "northern floodplain forest" classification with subsequent studies by the Colorado Division of Wildlife (Miller. 1983, Job Pr o q , Rep. N-4-R-i and Snyder. 1984, Job Prog. Rep. N-4-R-2). Area thus computed for l~w-elevation riparian (minus the area o~ "northern floodplain forest) was subtracted from the area computed with PNVPOLY for grama-buffalograss. Changes in habitats that had occurred in Colorado by the mid-l~60's were computed from the 1967-based Conservation Needs Inventory (USDA 1971) as perfor-med by Klopatek et a l , (197S', En v i r nn , Conserv. 6(3):191-200), but adjusted to the historical areas previously computed. Data were cross-checked against those of Miller and Choate (1964, US For. Servo Res. Bull. INT-3) for forested areas. Low-elevation riparian status in the mid 1960's was estimated from a mid-range of values for the early-1940's through late 1970's interval (Miller 1983). Current status of low-elevation riparian areas was computed from recent Colorado Division of Wildlife studies (Miller _1983, Snyder 1984, Miller 1985, Job Prog. Rep. 86-039-01). For all other habitat types, status was estimated from county data of the 1982 Census of Agriculture (Vol. 1, Part 6, U.S. Dep. Commerce, Bur. Census, AC82-A-6). For counties �84 containing a habitat type of interest, the county area ascertained by PNVPOLY was divided by the area of the county reported in the census, yielding a "not-r~ported factor", typically, of slightly over 1. County acreages reported for "man-made systems" (cropland, roads, habitation, impoundments) were tabulated, multiplied by the not-reported factor, then subtracted proportionately from the areas of habitat types within the county. RESULTS AND RECOMMENDATIONS As long ago as 1967, nearly 25% of Colorado's natural vegetation ass 0 cia t ion s ("h abit atty pes") had bee n los t too the r -it s es ITab Ie 1). 0f particular concern in recent years is the reduction of low-elevation riparian habitats, the most important in the state for overall species richness, at the rate of approximately 1% los5 every 3-4 years. However, due to the rapidly-aging condition of cottonwood stands within this habitat, the loss will greatly accelerate within the next 15 years--as much as 75% of areas currently forested will be virtually treeless by the year 2000 without corrective action (Miller 1985). Fully 10% of the species of this system may be expected to be lost in such a circumstance. Since no low-elevation ripari~n areas are protected by other agencies, CDOW will need to acquire some amount of control over approximately 40,000 acres to meet minimum viable population standards for the 264 wildlife species found there, to address expected increases in non-consumptive use and other recreational demands (currently nearly 50% of property use), as well as increased demands from consumptive users (hunter densities on current CDOW properties can be expected to increase substantially with the current trend of private "low-hunter-density" hunting clubs). Best cost estimates availabl~ at this time indicate that: a. -Maintenance costs of the additional properties will approximate 515K/yr cumulative, or 51.6 million b. -Habitat improvement due to losses of cottonwoods could approximate $1,050,000 OR currently-owned properties (treat 10,500 acres with 20 trees/ac.). c. -Habitat improvement due to losses of cottonwoods on newly-acquired properties could approximate 52,650,000. d. -The $3.7 million of improvements on the total low-elevation areas amounts to approximately 51.50/ ac/yr, using a conservativ~ 50-year estimate of project longevity. -Special habitat improvements in the reservoir-riparian system benefitting 10 species with specialized requirements and exhibiting high public interest (colonially-nesting waterbirds) will cost $5 million. §~~~~~gg:~!yg~!gm_Q[~irig has demonstrated the greatest loss (50%) of all natural habitats in Colorado ITable I}, and ranks highest of all non-forested types in species richness; In Colorado, the highest densities of ring-necked pheasants, greater and lesser prairie-chicken~ occur in the areas that were at least originally vegetated with this association. The type also demonstrates great importance to the eastern plains deer herds, upland sandpipers, scaled quail, and appears to be of importance to bobwhite as secondary nesting cover (various CDOW reports). CDOW controls the only significant amount of this type in public ownership primarily for wildlife benefits--approximately 5,000 acres. �85 CDOW should acquire some control over at least 20,000 acres of this type (for management core areas) by the year 2000 in order to meet targets of de-listing at least 2 threatened or endangered species, provide for expected increases in demand for non-consumptive wildlife uses, and to accommodate consumptive users of a decreasing habitat base. Additional considerations are: a. -Habitat improvement costs will vary gre~tly in this type--approximately 40% of the area can be expected to require perrennial cover establishment. Total improvement costs of $1 million amounts to about tl/ac/yr over a 50-year project longevity period. b. -Habitat maintenance will approximate 51.501ac/yr, or 5310,000 over the 14-year period. Date: September 1986 �86 Table 1. Vegetation (habitat) changes in Colorado--from conditions to most recent estimates. Supportive data Investigations office, Colorado Division of Wildlife. historical on file in Habitat =========================================================================== Vegetation Type mid-1960's Recent Historical mi2 (~~) mi2 mi2 Change (7.) --------------------------------------------------------------------------- Low-elev. riparian Sandsage-bluestem grama Pinion-juniper/mt. mahogany-oak scrub Grama-buffalograss Mt. grass-shrubs(5) Mt. forests(4) Desert Alpine TOTALS * - Project terminated 210 3,660 ( 0.2) 3.5) 171 1,798 179 1,918 -19 -50 12,163 (11.7) 9,905 JO,685 -12 40,735 13,982 27,794 67 5,263 (39.1) (13.4) (26.7> 23,913 11,619 24,805 64 5,237 20,956 -49 -17 -11 - 4 -tr 103,874 prior ( 0.1) ( 5.0) (100) to completion 77,640 * * * '+ (75) 72,657 (70) (estimated) -30 of analyses. bQ~:~lgY~!iQD_[iQ~[i~D: Data represent minimum estimates based only upon 318 river miles of the South Platte and Arkansas Rivers--upper reaches of these rivers, as well as Republican, Arikaree, Purgatoire, and smaller eastern streams not included due to extremely narrow configuration and limitations of remote sensing. §~U~~~qg:~l~!~t!m:g~!!!:This category combines sandsage-bluestem praIrIe and bluestem-grama types of Kuchler (1964). Of the 2, bluestem-grama comprised the smaller amount, only 112 mi2 historically, although by the mid-1960's 54% had been lost. Bluestem-grama was probably important to prairie grouse historically. ~t~_g[!~~:~t[~~~: Table combines 5 types identified by Kuchler (19641--ranging from fescue-mt. muhly (41% loss) and sagebrush (177. loss) to Great Basin sagebrush (1.51. loss). steppe ~t~_£Q[~~t~: Table combines 4 types identified by Kuchler (1964). Killer and Choate (1964, U.S. For. Servo Res. Bull INT-3) computed the area of these forests (commercial) as 19,063 mi2 (18.41.), in the early-mid 1960's, ~~. Klopatek's computation of 24,343 mi2. EiUyQQ:i~~i~~cL~t~_@~QQg!rrz:Q~t_~~[~~:: These types have been combined due to their general importance as winter range for deer and elk. These figures compare favorably with Miller and Choate's (1964) computation ~f 13,090 mi2 for these "non-commercial" forested lands, and figures from Kl op a t e k s t al. "s data set for Colorado (13,103 m i Z) , � https://cpw.cvlcollections.org/files/original/22295f9a356e1d37cf0d7e55da58c2b6.pdf c0c56254215fc381d7f4e77def7ef2d7 PDF Text Text 1 Colorado Division of Wildlife Wildlife Research Report April 1987 JOB PROGRESS REPORT State of Colorado ---------------------------------- Project 01-03-045 CW-88-R) Hork Plan 1 Job Title: 14 ----- Ecological Studies of the Flightless Period of Ducks in Colorado Period Covered: Author: : Job Avian Research 01 January 1985 through 31 December 1986 Michael R. Szymczak Personnel: C. Braun, J. Corey, J. Ringelman, S. Steinert, and M. Szymczak, Colorado Division of Wildlife ABSTRACT Comparative weight of Gadwall CAnas strepera) captured and measured in North Park indicated that the rate of weight loss in birds that were handled was greater than in the population as a whole. Atypical weight loss was immediate and lessened over time. Attempts to define and compare weight dynamic curves for gadwall during the flightless period met procedural problems but 14 of 20 location/sex/year class populations exhibited weight loss during the 20-day period when Primary lengths grew from 15-25 mm to 115-125 mm. Daily energy use of birds on metabolic trials was 153 Kcal/kg metabolic weight during non-growth periods indicating additional energy is not required for primary feather growth. Estimated survival through the flightless period of 7 female mallards CAnas platyrhynchos) was 0.57 ± 0.36% while 11 female gadwalls survived at an estimated rate of 0.73 ± 0.26%. X ��3 ECOLOGICAL STUDIES OF THE FLIGHTLESS PERIOD OF DUCKS IN COLORADO Michael R. Szymczak James K. Ringelman P. N. OBJECTIVES 1. Document the species and sex composition and seasonal abundance of ducks molting on selected wetlands in North Park. 2. Identify physical and biological characteristics of wetlands used by molting ducks. 3. Investigate spatial and temporal differences in use of wetlands by molting ducks. 4. Determine behavioral time budgets and net energy balance of molting ducks. 5. Investigate differences in duration of the flightless period of selected species of ducks in relation to sex, body condition, habitat quality, and time of molt. 6. Determine the survival rate of molting ducks in relation to sex, body condition, habitat quality, and time of molt. 7. Identify and quantify duck molting wetlands in Colorado. SEGMENT OBJECTIVES 1. Identify physical and biological characteristics of wetlands used by molting ducks. 2. Investigate spatial and temporal differences in use of wetlands by molting ducks. 3. Determine behavior time budgets and net energy balance of molting ducks. 4. Determine the survival rate of molting ducks in relation to sex, body condition, habitat quality, and time of molt. 5. Compile and analyze data and prepare progress report. METHODS Physical and Biological Characteristics of Wetlands Mapping of vegetation on 76 Pond, Elk Pond, Case #2, School Section, and Home Pond was completed and plant species in vegetation complexes at South Allard Pond and Wattenberg's were identified. Distances were measured between landmarks identifiable on aerial photos of wetlands to correct scales on those photos. �4 Gadwall Condition/Energy Balance Adult gadwall trapped in conjunction with a banding operation were weighed and measured to classify stage of molt and relate molt stage to physical condition. The lengths of primaries I, ~ and X were measured but, in analysis to date, only P X has been used in establishing molt stage. Measurements of girth and body length (Szymczak and Ringelman 1983) were taken to use in condition index estimation (Szymczak and Ringelman 1984). If a bird was subsequently recaptured, only weight, girth, and primary measurements were recorded. Birds were captured at Walden Reservoir (6, 8, and 16 Aug; 9, 12, and 17 Sep), MacFarlane Reservoir (7 and 15 Aug, 11 Sep), Lake John Annex (18 Sep) and Pole Mountain Reservoir (19 Sep). Metabolic Feeding Trials-Gadwall On 1 July 1986, 8 gadwalls, 4 males and 4 females, were placed in individual digestion cages inside a wooden shed. The birds had hatched from eggs taken from the wild in July 1985. Birds were provided with feed (lay pellets - 20% protein, 2% fat, and 10.5% fiber) and water ad libitum until they were acclimated to the cages. All males were removed after they failed to eat and one female was removed after injury. Trials were initiated on 21 July and 2 trials were conducted weekly with one exception until termination on 15 October. Each trial began between 0830 and 0930, ran for 24 hours, and was followed directly by a second trial. For each trial, each bird was provided with 200 gm (wet wt.) lay pellets. At the conclusion of the trial all remaining food and accumulated feces was collected in individual aluminum pans. Birds were weighed at the beginning of the first trial of the week and at the conclusion of the second trial. The stage of primary molt was determined at the start of each 2-trial series. Ambient temperatures (max-min) were recorded daily. Collected food and feces, and a weekly sample of the lay pellets were dried in a convection oven at 100 C for at least 48 hours, and then weighed. Feces were ground in a Wiley Mill and analyzed in duplicate for caloric content through bomb calorimetry. The mean of the 2 caloric values was selected as the true caloric content of the sample. The caloric value of the lay pellets was also determined. To determine existence energy (Kcal/kg metabolic weight) for each trial, food and feces weight values were converted to caloric values and bird weights were converted to metabolic weight (weight in Kg·72). Trial weights used in calculations were the mean of the starting and ending weights of the 2 successive trials. Survival Radio transmitters were attached to 7 flightless adult female mallards and 12 flightless or near flightless adult female gadwalls in September 1985 to specifically monitor survival during the flightless period. One gadwall (#578) apparently left the study area before it became flightless reducing the gadwall sample to 11. Birds were monitored almost daily between the time of radio attachment and the estimated time they reached flight capability, and periodically after that time until 13 November or until signal loss. �5 The estimated number of days (exposure days) the birds were considered flightless was calculated by subtracting the length (mm) of primary! at capture from the estimated length at which the birds were assumed capable of flight (mallards = 131 mm, gadwall = 139 mm) and dividing that by the daily feather growth rate (mallards = 5.21 mm, gadwalls = 4.69 mm). Mallard growth rates and involuntary flight capability estimates were obtained from Owen (1979). Energy Budgets of Flightless Gadwall - Effects of Handling Birds recaptured and measured one or more times within the same flightless period should present an accurate measure of weight change during that period. However, examining weight changes in birds recaptured within the approximate 20 days required for primary to grow for 20 to 120 mm showed that weight loss (gm/day) may have exceeded that occurring in the wild population. Weight changes were variable, ranging from +2 to -50 gm/day, indicating varying response to handling. Trapped male and female gadwall held in captivity without food lost an average of 38 gm/day and 55 gm/day, respectively for the first 2 days of captivity (M. Szymczak, unpubl. data) showing that a 50 gm/day weight loss is possible. X To evaluate the affect of handling on weight dynamics, weights of birds captured at Walden Reservoir in 1982 were examined. The weights of birds subsequently recaptured were compared to the weights of birds in the general population by molt stage (Table 1). The weights of the birds at recapture were then compared with weights obtained from the wild population. Birds were assigned by sex to 4 molt stage categories (Table 1). Birds with primary ! greater than 150 mm at the time of the original capture were not included. The sample of subsequent recaptured birds allowed original weight comparisons in 7 of 8 molt class/sex categories. In 6 of the 7 classes, mean weights of sample birds were greater than the population. Generally, numerical differences were not great and there was no difference in weight between the general population and those birds that were subsequently recaptured. The second level of comparison between the sample of birds recaptured and the population resulted in 15 molt class/sex categories. Unfortunately, in most categories the number of sample birds was small hampering the ability to find statistically different mean weights even though numerical weight differences were substantial. In 11 of 15 categories the mean weight of sample birds was less than the mean weight of the population. Mean weights were significantly less than population weights (p <0.05) in 4 categories and approached significance (R <0.10) in 2 others (Table 1). Generally, the shorter the period between the original capture and recapture the greater the difference in mean weights indicating an initial reaction to handling which lessened over time. A plot of time after capture vs. rate of weight loss of individual birds also indicated an immediate response to handling (Fig. 1). Energy Budgets of Flightless Gadwalls - Weight Dynamics Body weights by stage of primary growth have been plotted annually by 30 m (~5 days) increments since 1982. Generally, a significant loss in weight has been recorded for the mid-molt period (Szymczak and Ringelman 1986). Following the �0\ Table 1. Comparative weights, by molt stage, of gadwa11s captured more than once during the flightless period with those birds captured only once. x Stage No molt 0-50 mm 51-100 101-150 Reca]2 Stage X weight PO]2u1ation Sam]21e birds +1.29 0-50 mm 51-100 mm 101-150 mm 734.62 (91) 708.75 (48) 698.00 (20) 685.00 (4) 717.50 (4) 700.00 (2) -6.75* +1.23 +0.29 813.33 (3) -0.16 0-50 mm 101-150 mm 829.81 (103) 795.98 (117) 785.00 (2) 850.00 (1) -5.40** +6.79 734.62 (91) 736.88 (16) +0.31 0-50 mm 51-100 mm 101-150 mm 734.62 (91) 708.75 (48) 698.00 (20) 682.00 (5) 680.00 (8) 693.33 (3) -7.16** -4.06*** -0.67 M 829.81 (103) 839.52 (21) +1.17 0-50 mm 51-100 mm 101-150 mm 829.81 (103) 802.55 (110) 795.98 (117) 840.00 (3) 764.62 (13) 766.67 (6) +1.23 -4.73** -3.68*** F 708.75 (48) 760.00 (2) +7.23 51-100 mm 101-150 mm 708.75 (48) 698.00 (20) 690.00 (1) 770.00 (1) -2.65 +10.32 M 802.55 (110) 807.89 (19) +0.67 51-100 mm 101-150 mm 802.55 (110) 795.98 (117) 706.00 (5) 769.33 (15) -12.03* -3.35** M 795.98 (117) 822.50 (4) +3.33** 101-150 mm 795.98 (117) 795.00 (4) -0.12 wei8ht Sam]21e birds Sex PO]2u1ation F 711.84 (38) 721.00 (10) M 814.67 (15) F *p < 0.01. **P<0.05. ***p < 0.10. % diff. % diff. �7 10 + A o + A A A A A A B A B B A B B -10 + C C A A A B A B B B C A A A B -20 B A A A D B A A A A A A B C A B A A A A A B A A A B C A A A + ~ Cl .•..•.• '"'"0 H E-< ::r:: A= 1 2 C 3 D = 4 C-' H gl -~:t) + -40 + A -50 + A B A A Observation Observations Observations Observations ---+----------+----------+----------+----------+----------+----------+-c) 5 10 15 20 25 30 DAYS ELAPSED Fig. 1. Daily rates of weight (gm) change of gadwalls for specific recapture intervals during the flightless period when Primary X lengths are >19 and <120 mm. �8 1985 banding season, 4 years of measurements data were pooled by sex in an attempt to define weight loss curves. The first test of the pooled data using 16 stages (pre-flightless + 10 mm increments) of feather growth showed that significant variation occurred in weight by stage, location, and year (p <0.001). Attempts to combine individual year-location groups whose-weight stages were not significantly different were abandoned since the shape/slope of the growth curve could be the same even though weights were different. Data from either end of the weight/stage curves were deleted. Birds that were in the pre-flightless stage (no molt) were eliminated since the actual number of days until feather drop was unknown. Birds whose primary X lengths were greater than 120 mm were also deleted from the data set, since weights of flightless gadwall generally reach a low point at 120 mm. Within year recaptures were eliminated because of suspected handling effects. A regression analyses showed significant weight differences (p <0.05, negative slopes) for males at Walden Reservoir in 1982 and 1985, for-females at Walden Reservoir in 1982, for males at MacFarlane Reservoir in 1983, 1984, and 1985 and females in 1985, for males and females at Lake John Annex in 1983, and for males at Pole Mountain Reservoir in 1983. However, average estimated weight loss was small, 0.5 gm/day in 8 of 10 location/sex/year classes. Further refinement was accomplished by examining plots of weight/stage data. Weight loss generally begins within the 3-day period when primary length is in the 15-25 mm range and reaches a low point ~20 days later when primary X is in the 115-125 mm range. A comparison of weights of birds between these 2 stages indicated weight loss in 14 of 20 location/sex/year classes (Table 2). Weight differences were far more substantial than in the above comparison, thus projecting significant weight loss/day. However, smaller samples reduced the ability to detect statistically significant differences. X Energy Budgets of Flightless Gadwalls - Metabolic Trials Trials were conducted for 3 weeks before the first bird (#3) lost its' remiges. Bird #7 did not lose its' remiges until the 7th week of the trial and bird #1 underwent an atypical molt cycle and was not used in the analyses. Weights of experimental birds increased during the trial period (Fig. Birds were lighter in the pre-molt stage (P <0.0001) than during molt post-molt stages. However, weight also differed by trial, increasing progressed (Fig. 2), regardless of stage, masking possible molt stage 2). and as time effects. Food consumption was greater during the post-molt period than during the molt (p = 0.07) although, as with body weight, there was also a difference by trial. Energy density of excreta did not differ by molt stage (R>0.12) nor did assimilated energy (p = 0.10) when expressed as Kcal/gm metabolic weight, indicating the efficiency of food use remained the same throughout the different molt stages. However, energy density of excreta did differ among trials (R<O.OOOl) suggesting a pen effect. Daily energy use per bird during non-molt periods was about 185 Kcal/kg metabolic weight or about 2.5 times the estimated basal metabolic rate (BMR = 75 * wt·72). During the period of primary feather growth, estimated energy �Table 2. Weight (gm) differences of flightless gadwall at 2 primary are in parentheses. X growth (mm) stages. Wei~ht at: Year Location 1982 Walden Res. MacFarlane Res. 1983 Walden Res. MacFarlane Res. L. John Annex Pole Mountain 1984 Walden Res. MacFarlane Res. L. John Annex Pole Mountain 1985 Walden Res. MacFarlane Res. Sex 15-25 115-25 F M F 720.45 (22) 830.00 (21) 776.67 (3) 680.00 (5) 799.55 (22) 680.00 (2) F M F M M F 714.12 803.00 656.25 742.50 860.91 730.00 (17) (10) (8) (4) (11) (1) 686.67 758.00 716.67 727.50 757.14 785.00 F M F M F M F 730.00 826.67 714.57 873.33 640.00 880.00 710.00 (6) (6) (11) (3) (1) (4) (1) F M F M 757.00 879.97 743.39 821.50 (11) (9) (4) (4) Difference Sample sizes Est. ave. wt. chan~e/da~ -40.45 -30.45* -96.67** -1.90 -1.51 -4.53 (3) (5) (3) (4) (7) (2) -27.45 -45.00 +60.42* -15.00 -103.77*** +55.00 -1.29 -2.23 +2.83 -0.74 -5.15 +2.58 739.21 837.86 752.43 839.56 735.00 826.67 700.00 (17) (14) (18) (22) (2) (9) (1) +9.21 +11.91 +37.86* -33.77 +95.00 -53.33 0 +0.43 +0.55 +1. 78 -1.67 +4.46 -2.65 733.38 821. 95 683.94 765.81 (19) (12) (4) (22) -23.62 -58.02* -59.45* -55.69* -1.11 -2.88 -2.79 -2.71 *p < 0.10. **P<0.05. ***P < 0.01. \.0 �t-' a WEIGHT _----""1....J ~~~~~~, .•..•. ..... •• •.- .- # " -_- -~ _ ~ _ ~ __ - 600 ~;-~-------------~I ~- ••.... I 200 ~ . I- OJ ENERGY I I I I I I I 150 W ~ CJ) ~ 100 <, '--------'" co 0 --BIRD 50 --_. -~ • +- -- -- -< 400 .•.•,/ .....- _.,'" .......... o ~ ", ", ", ", 0 'I CO 3 200 BIRD 7 PRIMARY MOLT 21 29 JUL Fig. 2. molt. 6 19 AUG 26 2 9 15 SEPT 23 30 7 15 OCT Energy and weight dynamics of captive gadwalls before, during, and after primary feather �11 use was actually less per day (153 Kcal/kg met. wt.) than during non-molt periods (Fig. 2). Energy use coefficient (% ingested energy not excreted) was about 30% during he trials. The results of these trials were basically inconclusive. The inability of the captive birds to adapt to the small digestion cages resulted in the removal of 5 of the 8 original test birds. A 6th bird underwent an atypical molt cycle reducing the number of experimental birds to 2. Throughout the test environmental factors masked the effects of the molt cycle on different parameters. The results of the experiment suggest that primary feather growth has little effect on nutrient dynamics in adult gadwalls as suggested by Ankney (1979) for waterfowl and Murphy and King (1984) for Passeriformes, specifically white-crowned sparrows (Zonotrichia leucophreys). The results do little to explain the significant reduction in weight during primary molt experienced by free-ranging adult gadwalls in North Park. Survival Four of 7 instrumented female mallards survived through the flightless period (Table 3). One bird at MacFarlane Reservoir was killed by a predator shortly after marking, another died of starvation at Lake John Annex, and a third was shot by a hunter on Walden Reservoir. Estimated survival was 0.57 ±0.36 for the flightless period. Only 3 mallards remained alive and/or were flightless going into the hunting season. All were on Walden Reservoir which was subject to heavy hunting pressure on 5 and 6 October 1985. Two of the birds, of which 1 was shot, remained on the main portion of the lake in open water areas. The third bird remained on the north pond in a dense bulbush (Scirpus spp.) stand, before moving to the Illinois River in the late stages of feather growth on 8 October. A simplistic estimate of hunting season survival for these birds was 0.67±0.53. However, since the surviving birds attained flight before the end of the hunting season the length of exposure should be considered. Eight of 11 instrumented female gadwalls survived through the flightless period (Table 3). One bird was taken by an avian predator at Pole Mountain Reservoir and 2 birds were shot by hunters, one each at Walden Reservoir and Lake John Annex. Estimated survival was 0.73± 0.26 for the flightless period. Ten gadwalls remained alive and were flightless when the hunting season began. One bird each was at MacFarlane Reservoir, an area not hunted, and Lake John Annex, while the other birds were on Walden Reservoir. All but 2 of these birds survived the hunting season for a survival rate of 0.80± 0.25. Survival for birds on Walden Reservoir during the hunting season was 0.88± 0.21. All but one bird on Walden Reservoir was estimated to be flightless during the first 2 weekends of the hunting season and that birds (#623) had just reached flight capability at the beginning of the second weekend. Analysis of the data indicate substantial mortality of late-molting female mallards on North Park wetlands. Unfortunately sample sizes were small and geographically they were not distributed in the same fashion as the population. Annual survival of North Park adult females during 1971-80 was 59.8% (Szymczak 1986). Data accumulated through wetland counts, monitoring instrumented birds, and trapping indicated late-molting mallard females �f-' N Table 3. Molt locations, characteristics, the flightless period in North Park 1985. Bird no. and chronology K S2ecies Location Mallard MacFarlane Res. Walden Res. Walden Res. Walden Res. L. John Annex L. John Annex Pole Mt. 622 570 576a 582 630b 629 621 889 1063 1086 1080 844 945 MacFarlane Res. Walden Res. Walden Res. Walden Res. Walden Res. Walden Res. Walden Res. Walden Res. Walden Res. L. John Annex Pole Mt. Pole Mt. 568 569a 623 572 573 624 580 581 571c 579a 578d 625 729 689 585 710 681 690 750 700 739 709 685 770 Gadwall aShot bDied cBird dBird of events for instrumented Wt. -- primary length (mm) 28 0 0 19 52 70 73 0 15 10 0 15 0 13 0 No molt 5 No molt 37 Ca2tured female mallards and gadwalls monitored Date Est. capable of flight Est. time of death 12 17 17 17 18 18 19 Sep Sep Sep Sep Sep Sep Sep 2 12 12 9 3 30 30 Oct Oct Oct Oct Oct Sep Sep 12 13 13 17 17 17 17 17 17 18 19 19 Sep Sep Sep Sep Sep Sep Sep Sep Sep Sep Sep Sep 12 10 11 17 14 17 14 17 Oct Oct Oct Oct Oct Oct Oct Oct ----6 Oct ------- 17 Oct 5 Oct --- 11 Oct by hunters. of starvation; primary feather growth retarded. captured prior to primary feather drop. captured prior to primary feather drop; departed area prior to wing molt. Last contact -- 26 Sep -- 19 Oct -- 6 Oct 8 Nov -- 5 Oct -- -- 1 Oct 24 Oct 4 Nov 7 Nov -- 14 22 22 13 4 31 28 Oct Oct Oct Nov Nov Oct Oct -- for survival during EX20sure daxs PostFlightless flightless 14 25 18 22 17 12 11 30 24 28 30 27 30 27 30 30 16 26 Sep -- 11 7 30 24 28 27 3 5 8 27 21 14 �13 primarily use smaller wetlands that support emergent vegetation. Walden and MacFarlane reservoirs are not prime molting areas for female mallards (M. Szymczak, unpubl. data). However, monitoring birds in these areas, particuarly Walden Reservoir, gives an indication of mortality where molting populations and extensive hunting occur on the same areas. In contrast, instrumented female gadwalls were distributed geographically similar to the molting population, although MacFarlane Reservoir should have been represented by a few more birds and some late molters use small pond areas. Adult female gadwalls banded on molting areas have an annual survival rate of about 59% (M. Szymczak, unpubl. data). Mortality of flightless females do not reflect mortality of the entire molting population, particularly males. Males and unsuccessful females of both species complete the molt cycle earlier in the summer and thus are not subject to hunting mortality during the flightless period. However, females marked in this study represents the productive segment of the population. Most likely these birds have successfully completed the nesting and brood rearing cycle. Additional survival analysis will be conducted using information from birds instrumented in other years with an analysis program that will consider exposure days, thus allowing a calculation of a daily survival rate. LITERATURE CITED Ankney, C. D. 1979. Does the wing molt cause nutritional stress in lesser snow geese? Auk 96:68-72. Murphy, M. E., and J. R. King. 1984. Sulfur amino acid nutrition during molt in the white-crowned sparrow. 2. Nitrogen and sulfur balance in birds fed graded levels of the sulfur-containing amino acids. Condor 86:324-332. Owen, M. 1979. The duration of the flightless period in free-living mallards. Bird Study 26:267-269. Szymczak, M. R. 1986. Characteristics of duck populations in the intermountain parks of Colorado. Colorado Div. Wildl. Tech. Publ. 35. 88 pp. , and J. K. Ringe1man. ------~~ period of ducks in Colorado. Aid Wildl. Rest. Oct. 1983. Ecological studies of the flightless Job Prog. Rep., Colorado Div. Wild1., Fed. Pp. 13-31. , and 1984. Ecological studies of the flightless period -----o~f~d-u~cks in Colorado. Job Prog. Rep., Colorado Div. Wi1dl. Fed. Aid Wildl. Rest. Oct. Pp. 13-31. __________ , and 1986. Ecological studies of the flightless period of ducks in Colorado. Job Prog. Rep., Colorado Div. Wild1. Fed. Aid Wildl. Rest. Apr. Pp. 13-30. �14 Prepared by ~ Ie J; T Michael R. SZymcz~ Wildlife Researcher C �Colorado Division ot Wildlite Wildlife Research Report April 1987 1) JOB PROGRESS REPORT State of Colorado --~~~~~---01-03-045 (W-88-R) Project Work Plan 1 Job Title: : Job Avian Research __ 15 .::.::..._ Development and Use of a Physiological Condition Index for Monitoring Wintering Mallard Nutrient Reserves Period Covered: Author: ------ __ ---- 01 July 1985 through 31 December 1986 J. K. Ringelman Personnel: J. F. Corey, J. K. Ringelman, M. R. Szymczak, Colorado Division of Wildlife; D. R. Anderson, Colorado Cooperative Fish and Wildlife Resea!ch Unit; D. C. DeLong, Colorado State University ABSTRACT Mallards (Anas platyrhynchos) were trapped at Bonny Reservoir during early December, mid-January, and late February. Body condition indices ranged from 0.142 to 0.179 during winter 1985-86. Female mallards of both ages were in highest condition during January, but males did not change in condition over time. Within trapping periods, males were in lower condition than females during January, while no differences existed among age-sex classes in either December or February. In December 1986, all age-sex classes were in good condition, except adult males had lower relative reserves than the other age-sex classes. Body mass of juvenile female mallards at Bonny Reservoir during 1977-84 was positively associated with mean maximum December temperatures and hectares of corn. Juvenile maLe body mass was also positively associated with,.hectares of corn. Among'adults~ males were positively associated with mean minimum December temperatures and negatively related to the number of days with December minimum temperatures below 0 C. Adult female body mass did not correlate with any variables. The population size of mallards overwintering at Bonny Reservoir was positively correlated with maximum and minimum temperatures, and negatively associated with December snowfall. The distribution of recoveries prior to legalization of hunting below Bonny Dam, compared to recoveries after hunting was permitted, did not support the hypothesis that mallards originally banded at Bonny selected a new terminal wintering area. In fact, because of higher harvest rates as a result of legalizing hunting below the dam, relatively more banded birds were reported from the Bonny degree block during the latter period. �The thermoregulation sub-model of a mallard winter energetics program was developed and tested. The sub-model integrates winter temperature records with body mass data to compute acclimation temperatures, lower critical temperatures, and total energy required for thermoregulation. Results from this sub-model indicate that lighter mallards incur higher total thermoregulatory costs when wintering in a cold climate. �17 DEVELOPMENT AND USE OF A PHYSIOLOGICAL CONDITION INDEX FOR MONITORING WINTERING MALLARD NUTRIENT RESERVES James K. Ringe1man Michael R. Szymczak Only recently have waterfowl biologists come to appreciate the role of stored nutrients in the life histories of ducks and geese. Earliest evidence of the importance of nutrient reserves was obtained from studies on arctic-nesting snow geese (Chen caeru1escens); (Ankney 1974, Ankney and MacInnes 1978), in which it was demonstrated that the size of reserves prior to breeding was directly correlated with the number of eggs in the clutch. The same relationships were later demonstrated for Canada geese (Branta canadensis); (Raveling 1979). Nutrient reserves important for maintaining physiological condition are not only obtained in migratory habitat during the spring (Raveling 1979), but may also be accumulated on wintering grounds (MacInnes et a1. 1974). Current studies on arctic-nesting geese have progressed beyond the point of demonstrating the importance of nutrient reserves to investigations addressing when and how these reserves are obtained. The role of nutrient reserves during the breeding period of ducks is less evident than in geese. Bengston (1971) provided field data to support Lack's (1967) hypothesis that the clutch size of ducks is related to food availability on the breeding grounds, yet Milne (1976) demonstrated that the weight of female Common eiders (Somateria mo1lissima) in winter was directly related to clutch size the following spring. The apparent discrepancy in these results was resolved by more detailed nutritional studies: clutch size in eiders is limited in part by the size of stored protein reserved (Korschgen 1977) whereas many dabbling ducks wait until arrival on the breeding ground to obtain protein needed for egg production (Krapu 1974, 1981; Drobney 1977; Reinecke 1977). For the mallard, there remains a common link between nutrients obtained in winter and spring and subsequent reproductive success: fat reserves provide the "fuel" that enables mallards and similar species to forage for the higher protein but lower energy animal foods (Krapu 1981). If the mallard population is regulated by limitations imposed during winter, as has been suggested for most temperate bird populations (Fretwell 1972), such regulation may also result from increased winter mortality. Most reports of waterfowl mortality during winter cite severe cold and/or snow as the causative factors (Pearson 1934, Gromme 1936, Trautman et a1. 1939). Fat, the most labile endogenous reserve, serves as the principal energy source used by waterfowl to survive severe winter weather. In all likelihood, winter fat reserves of mallards serve dual functions of increasing winter survival probabilities as well as providing nutrients and energy essential for reproduction. The questions of foremost interest are (1) what minimum level of winter fat reserves are necessary to optimize both survival and reproduction, (2) what role does weather-induced stress play in the dynamics of fat reserves, and (3) can conditions that are detrimental to the . acquisition of fat reserves cause a shift in the winter distribution of mallards? �18 The latter 2 questions were the focus of much of this segment's work, using data obtained from mallards captured at Bonny Reservoir. This reservoir was selected for study because it typically supports the largest winter concentration of mallards in Colorado, with a recent high population of 67,700 in 1980 and a low of 6,100 in January 1985. Research personnel have banded mallards at Bonny since 1963, thus this population contains one of the largest samples of banded birds in the Central Flyway. A dramatic decline in the Bonny mallard population began in 1982 (Jan inventories: 58,350 [1982]; 12,500 [1983]; 18,000 [1984]; 6,100 [1985]). Concurrent with this decline was a change in hunting regulations, beginning in the 1980-81 season, which allowed hunting from "controlled" blinds along the toe drain and outflow stream below the dam. This area was deemed an "experimental hunting area", the "experiment" intended to determine if more ducks were harvested as a result of the regulations change. However, the action was not an experiment in the scientific sense; no pre-treatment data were obtained, no control area was designated, nor was a formal hypothesis proposed. In addition, several other factors including relatively severe winter weather, a reduction in the area of corn grown in the vicinity, and a substantial decline in the Central Flyway mallard population have also occurred since 1982. These changes may also be contributing to the decline in the Bonny population. P. N. OBJECTIVES The objectives of this study are to: 1. Develop a physiological condition index to accurately assess nutrient reserve levels of wintering mallards, 2. Determine if differences exist in the physiological condition of mallards wintering in several areas of northeastern Colorado, 3. Document temporal changes in mallard physiological condition on a major Colorado wintering area, and 4. Relate spatial and temporal differences in mallard condition, if such differences exist, to weather and the availability of waste cereal grain. SEGMENT OBJECTIVES 1. Capture, weight, measure, and derive a condition index for at least 40 mallards of each age-sex class during early, mid, and late winter at Bonny Reservoir. 2. Quantify temporal and age-sex specific differences in the condition index. 3. Tabulate and analyze banding data to test the hypothesis that changes in philopatry are responsible for the observed decline in winter mallard populations at Bonny Reservoir. 4. Begin development of a computer simulation program to model the role of weather in the body condition dynamics of wintering mallards. �19 METHODS Field Procedures Salt Plains bait traps (Szymczak and Corey 1976) were used to capture mallards at Bonny Reservoir on 2-6 December 1985, 19-24 January 1986, 25 February-l March 1986, and 4-6 December 1986. Mallards were held 4-12 hours to allow digestion of ingested corn prior to being weighed to the nearest gram. Wing length was measured to the nearest millimeter. Sex-specific condition index equations (Ringelman and Szymczak 1985) were used to estimate total and relative fat reserves. Conventional analysis of variance procedures were used to compare mean condition index values by time, age, and sex. Multiple Regression Analysis Mallards have been weighed during winter banding operations at Bonny since 1977. These data were included in an analysis designed to investigate the relationships between mallard body mass and January population size at Bonny vs. weather, food, and population parameters. With body mass and population size as dependent variables, a multiple regression analysis was performed using 8 independent variables (Table 1). The objective was to determine which factors accounted for most of the variation in either body mass or population size. Hunting regime was not considered as an independent variable, primarily because it represented discrete rather than continuous data. Ten years of data were used, 6 of which included years when no hunting was allowed directly below the dam. Thus, the data are weighted to the "no-hunting" regime, and hunting may be adding "noise" to the analysis. Table 1. Variables used in multiple regression analyses. Variable Significance Dec snowfall Snow limits food availability Mean maximum Dec temperature Mean minimum Dec temperature Low temperatures freeze water areas and increase energy demands Number of days with temperature Availability of open water > a C Area of corn in Yuma and Kit Carson counties Corn is the primary food item in the mallard diet Breeding mallard population Population size at Bonny Reservoir may depend on breeding population size Central Flyway age ratio in harvest Fall flight estimate Bonny Reservoir population size may be a function of recruitment Body mass Dependent variable Jan population size at Bonny Reservoir Dependent variable �20 Philopatry Analysis The purpose of this analysis was to test the hypothesis that hunting below the dam excludes birds from one of the few open water areas available during severe weather. This precludes birds from obtaining supplementary foods (especially invertebrates) essential for complementing a corn diet, which is nutritionally deficient. It also places birds in a negative energy balance, since they are forced to roost in colder, wind-blown habitats. This energy stress causes those birds with the lowest energy reserves to leave Bonny. These birds are typically immature males and females (J. K. Ringelman, unpubl. data). The net result of forcing birds from an energetically favorable environment is a break in the homing tradition to the old terminal wintering area. During banding operations, previously banded birds are often recaptured. In this case, their probability of recapture is a function of both homing rate (philopatry) to the terminal wintering area and survival rate. Recapture probabilities can be estimated using mark-recapture survival models (Jolly 1965, Seber 1965). Survival rates can be independently estimated by maximum-likelihood techniques (Brownie et al. 1978) using recoveries of banded birds. Theoretically, the contrast between the rates derived using these 2 methods will be an estimate of "homing rate". A comparison of homing rates between years in which hunting was not allowed below the dam with years where hunting was permitted serves as a test of the effect. of a disturbance (hunting) on philopatry to a wintering area. A second means to assess changes in philopatry is to examine the distribution of recoveries of mallards banded at Bonny Reservoir. Recoveries for the pre-hunting period (1963-64 through 1975-76) were compared to the hunting period (1981-82 through 1984-85) to detect any changes that may have occurred in the distribution of harvest of banded birds. Simulation Model Modeling has become a powerful tool now widely used in vertebrate biology (Verner et al. 1986). In waterfowl biology, models have been used to guide management of nesting habitat (Johnson et al. 1986), understand breeding bioenergetics (Owen and Reinecke 1979), and direct future research (Ringelman and Longcore 1980). The advantages of attempting to model a biological system are (1) the modeling process requires a detailed assessment of the state of our knowledge and a thorough understanding of inter-relationships, (2) assumptions are stated explicitly, thus allowing them to be isolated and tested, (3) simulations involving manipulations of several variables simultaneously can be quickly and easily performed, and (4) sensitivity analyses can be performed to evaluate the relative "importance" of model components in terms of how they change the dependent variables of interest. Model building becomes a deductive-inductive process, since models suggest questions to ask of nature and nature provides answers that further shape the models (Stormer and Johnson 1986). An overview of the model being developed is depicted in Fig. 1. Weather conditions (temperature regimes, snowfall, wind) drive the energy submodels (behavior, thermoregulation, feeding) which in turn regulate energy balance and subsequent nutrient reserve dynamics. Weather records for both Bonny �21 Weather (max. temp., min. temp., snowfall, wind) Body Mass (randomly generated at t = 1 ) Population = 100 birds , Behavioral Energy Basal Metabolic Rate (activity budgets and multiples of basal metabolic rate) (standard mass-based equation) Net Energy Balance (sum of behavior, thermo, and feeding) - ~ - Thermoregulatory Energy (acclimitization, hours below LeT, convective heat loss) It Reduce or increase fat and protein reserves based on energy balance Food Energy It Add or subtract body mass based on changes in nutrient reserves. (corn consumption corrected for snow, post-harvest treat.) Fig 1. Flow diagram of the winter energetics for mallards wintering in eastern Colorado. model being developed �22 Reservoir and other mallard wintering areas were used for initial weather input. Sensitivity analyses will be used to explore the impact of weather on mallard energy dynamics. In this instance, weather input will be modified to simulate heavy snowfall, prolonged cold, etc. The simulation begins with a population of 100 mallards on 1 December. Body mass for each bird in the population is assigned by a random number generator. The mean and standard deviation of the population is set to approximate the distribution of body mass values in a wild population. Basal metabolic rate is expressed as 75 (massO•72) (after Smith and Prince [1973] and Wooley and Owen [1977]) for each bird. A net energy balance is then calculated for each individual based upon energy demands and acquisition from the 3 sub-models. Body composition is calculated from regression equations relating nutrient reserves to body mass (J. K. Ringe1man, unpub1. data, Whyte and Bolen 1984, Whyte et a1. 1986), then fat and protein reserves are "catabolized" or "anabo1ized" according to energy use. The net impact of changes in body constituents is reflected in changes in mass for each bird. These new values for body mass are then cycled back to the beginning of the model for day t + 1. The model simulates a 90-day winter from 1 December to 28 February. The behavior sub-model uses time budget information (e.g., Jorde 1981; J. K. Ringe1man, unpub1. data) to assign a daily behavioral energy budget equal to the sum of multiples of basal metabolic rate for each behavior (Wooley and Owen 1977, Owen and Reinecke 1979). Changes in behavior as a function of weather will be incorporated into the sub-model (Jorde et a1. 1984). Basal metabolic rates for each bird, and therefore behavioral energy, change on a daily basis. Thermoregulatory energy is determined by summing convective heat loss (Jorde 1986) and hours below lower critical temperature (Smith and Prince 1973, Prince 1979) corrected for seasonal acclimatization period, using actual temperature data beginning on 17 November for each simulation. Corn is considered the sole food resource in the food energy sub-model. Corn consumption is modeled using functional response equations that relate feeding rate to snowfall and post-harvest treatment (J. K. Ringe1man, unpub1. data). Available cornfields are initialized at the beginning of the simulation, post-harvest treatments are described, and corn is "depleted" as winter progresses. Core areas and feeding arenas will be designated in a manner similar to another waterfowl simulation model, REFMOD (Frederick et a1. 1983). After verifying the model's performance, sensitivity analyses will be conducted to assess the impacts of severe weather and/or food abundance and availability. Bounds of the parameters used in the sensitivity analysis will reflect current or anticipated conditions on the high plains of Colorado during winter. RESULTS AND DISCUSSION Trapping Effort Nearly 10 cm of snow was present on the ground when trapping commenced in December 1985. Cold temperatures and high winds produced wind chills of -43 C �23 which caused the reservoir to freeze completely on the night of 1 December. Cold weather prevailed during the December trapping period. In an effort to increase the probability of recapturing females during subsequent trapping periods, 99 adult female and 100 juvenile female mallards were captured, weighed, measured, and banded. Regular quotas were maintained for males. High water releases from the reservoir during January 1986 made trapping below the dam difficult, but daytime temperatures near 15 C and overnight lows at or above freezing provided ample alternate trap sites. Snow cover was non-existent during this trapping period. During February 1986 trapping, temperatures were mild, the entire reservoir shoreline was open and birds were captured along the north shoreline of the reservoir. The reservoir water level was atypically low in early December 1986. Sheltered bays along the north side of the reservoir would freeze during nighttime hours as temperatures fell to -5 C. Birds were trapped along the east side of the north cove in the northeast corner of the reservoir. Condition Indices Mean condition indices (CI) ranged from 0.142 to 0.179 during winter 1985-86 (Table 2). Both adult and juvenile females were in higher condition during January than during either December or February. Condition indices for adult and juvenile males, however, did not change during the winter. Across all age-sex classes within trapping period, no differences existed in the values of CI during December of February. In January, adult males = juvenile males < juvenile females = adult females. Extremely mild January weather apparently enabled males to maintain fat reserves through mid-winter, when in normal winters condition decreases significantly. The mean CI for adult males in January was relatively low compared to data for adult males at other locations or years during the same month (Table 3). Females were actually in better condition during January than during early or late winter. Table 2. Condition indices of mallards trapped during winter 1985-86 at Bonny Reservoir, Colorado. Within rows, means with the same letter do not differ by capture date (P>0.05). Age-Sex class Ad F Ad M Juv F Juv M x 0.157a 0.159a 0.143a 0.142a Dec SE 0.005 0.006 0.005 0.008 N 99 39 100 37 Capture date Jan SE x 0.179b 0.149a O.l72b 0.154a 0.005 0.004 0.006 0.006 N 38 62 38 49 x 0.157a 0.147a 0.146a 0.144a Feb SE O.OOp 0.006 0.007 0.006 N 40 42 40 42 �24 Condition indices for adult male mallards captured at 4 Colorado Table 3. locations during January 1983, 1984, and 1985. Within columns, means with the same letter do not differ (p> 0.05). At Bonny Reservoir mallards captured during 1983 had a greater cI than during 1984 and 1985 (p < 0.05). Location 1983 Year 1984 1985 Bonny Kodak Ponds Valmont Reservoir Segelke 0.173ab 0.168a 0.195c 0.188bc 0.14la 0.179b 0.143a 0.149a 0.156a 0.147a 0.149a 0.182b In December 1986, mallards were generally at a high level of condition (Table 4), except that adult males had lower relative reserves than other age-sex classes. Record warm weather preceded this trapping period. Thus, low thermoregulatory costs coupled with abundant food created optimum conditions for building and maintaining nutrient reserves. Table 4. Condition indices of mallards trapped during December 1986 at Bonny Reservoir, Colorado. Means with the same letter do not differ (p > 0.05) • Age-Sex class Ad F Ad M Juv F Juv M Condition index SE x 0.007 0.007 0.007 0.006 0.2l5a 0.175b 0.204a 0.198a Multiple Regression Analysis Among juvenile females during 1977-84, variation in body mass was positively associated with mean maximum December temperatures and hectares of corn. These 2 variable accounted for 85% of the variation in body mass (p = 0.059). Among adult males, mass was correlated positively with mean minimum December temperatures and associated negatively with the number of days with December minimum temperatures below 0 C. These variables accounted for 95% of the variation in adult male mass (R = 0.009). Bod mass of juvenile males was positively correlated with hectares of corn (R = 76%, P = 0.023). Adult female mass did not correlate with any variables (R> 0.(5). 2 This analysis suggests that adult males, the dominant individuals, tend to alter their energy balance (and therefore body mass) in response to energy costs associated with thermoregulation (low temperatures). Subordinate juveniles, however, appear to respond mostly to food availability (energy acquisition), and energy balance changes accordingly. It would be expected that subordinate birds would be at a competitive disadvantage under a regime of limited food availability. N 40 40 38 41 �25 The number of mallards wintering at Bonny during January was positively associated with both the mean maximum and mean minimum temperatures, and negatively associated with December snowfall. Thus, December snowfall and temperatures were of over-riding importance, accounting for 82% of the variation in mallard numbers (P = 0.027). Area of corn or flyway population size had little to do with population variation. Phi10patry Analysis The distributions of harvest for normal, wild mallards banded post-season at Bonny Reservoir and recovered elsewhere were plotted for 2 time periods: 1963-64 through 1975-76 (pre-hunting below the dam), and 1981-82 through 1984-85 (hunting below the dam). Only adult males had a sufficient number of recoveries in both periods to draw valid comparisons. During the pre-hunting period, 102 of 281 recoveries (36.3%) occurred in the degree block including Bonny Reservoir (Fig. 2). A higher proportion (!2 = 4.94, df = 1, R <0.05) of recoveries (87/197 = 44.2%) occurred in the same degree block during the hunting period (Fig. 3). This result is counter to the direction of expected change. The analysis underscores the extent to which a change in hunting pressure can alter recovery rates of banded birds and the harvest distributions which rely on reports of recoveries. Hunting below the dam appears to have sufficiently increased harvest rate, resulting in a change in the apparent harvest distribution. This change may have effectively masked any subtle changes in phi10patry that occurred as a result of changes in hunting regulations. Maximum likelihood banding analysis models (Brownie et a1. 1978) were used to estimate survival rates of mallards. During 1965-84, 10,634 male and 8,169 female mallards were banded at Bonny Reservoir. The Jolly-Seber survival rate model was used to analyze data on recaptures of 962 male mallards which fell into 272 recapture patterns, as well as 10,935 males that were banded but not recaptured. A similar analysis was performed on female mallards with 252 recaptures, 113 recapture patterns, and 8,273 birds that were banded but not recaptured. Unfortunately, adults and subadu1ts were pooled for these analyses which may have been appropriate for calculating survival estimates using recoveries (Hopper et a1. 1978) but was not appropriate for comparing recapture patterns with recoveries. Hopper et a1. (1978) found contrasts in recovery distributions of mallards banded as adults and subadu1ts in eastern Colorado. The data will be re-ana1yzed using only the adult segment of the population. Simulation Model Work on simulation modeling during this reporting period focused on outlining the overall model flow, development of the thermoregulation sub-model, and obtaining and coding weather data. Coding for the thermoregulation sub-model is essentially complete (Appendix 1), and has been compiled and tested for accuracy. The program begins by initializing starting body masses and temperature maximums and minimums beginning on 17 November. The first 2 week period of temperature regimes was used to acclimatize birds and adjust lower critical temperatures on a daily basis, following equations used by Owen and Reinecke (1979). Basal metabolic rates are calculated for each bird. Hourly temperatures are computed based on daily maximum and minimum temperatures using the program developed by Parton and Logan (1981). �26 Fig. 2. Distribution of harvest for adult male mallards banded at Bonny Reservoir during which time hunting was not allowed below the dam (1963-76). , �27 Fig. 3. Distribution of harvest for adult male mallards banded at Bonny Reservoir during which time hunting was allowed below the dam (1981-85). �28 Time below lower critical temperature per day is calculated for each bird, then converted to an estimate of energy needed for thermoregulation. Results of a trial run of the thermoregulation sub-model are presented (Table 5) for a hypothetical population of 12 birds. Body mass remains the same throughout the winter in this example, since no attempt has yet been made to decriment mass for energy use. Resting metabolic rate (RMR), maximum (MAXT) and minimum (MINT) daily temperatures, average temperature of acclimation (TACCL), lower critical temperature (LCT), and energy needed for thermoregulation (in kcal, HEAT) are shown for days 1 and 2. Particularly on day 2, thermoregulatory costs become appreciable. Lighter birds, because of their higher lower critical temperatures which puts them below their thermoneutra1 zone more frequently during the day, experience higher total thermoregulatory costs. The sum of thermoregulatory energy for winter 1982-83 (HEATNRG, Table 6) was 45.6% greater for an 800 gm bird than for a 1300 gm mallard. Example output (Colorado, winter 1982-83) from the thermoregulation Table 5. sub-model. Abbreviations are: RMR (resting metabolic rate), MAXT (daily high temperature), MINT (daily low temperature), TACCL (acclimation temperature), LCT (lower critical temperature), and HEAT (total thermoregulatory costs in kcal/day) • Mass Day 1 0.800 0.850 0.900 0.950 1.000 1.050 1.100 1.150 1.200 1.250 1.300 0.999 Day 2 0.800 0.850 0.900 0.950 1.000 1.050 1.100 1.150 1.200 1.250 1.300 0.999 Temperature and thermoregulatory data MAXT MINT TACCL RMR LCT HEAT 80.3 82.9 85.5 88.1 90.6 74.9 15.0 15.0 15.0 15.0 15.0 15.0 15.0 15.0 15.0 15.0 15.0 15.0 -5.0 -5.0 -5.0 -5.0 -5.0 -5.0 -5.0 -5.0 -5.0 -5.0 -5.0 -5.0 -0.5 -0.5 -0.5 -0.5 -0.5 -0.5 -0.5 -0.5 -0.5 -0.5 -0.5 -0.5 -2.2 -2.5 -2.9 -3.3 -3.6 -4.0 -4.4 -4.7 -5.0 -5.4 -5.7 -3.6 0.2 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 63.9 66.7 69.5 72.3 75.0 77.7 80.3 82.9 85.5 88.1 90.6 74.9 -2.2 -2.2 -2.2 -2.2 -2.2 -2.2 -2.2 -2.2 -2.2 -2.2 -2.2 -2.2 -16.7 -16.7 -16.7 -16.7 -16.7 -16.7 -16.7 -16.7 -16.7 -16.7 -16.7 -16.7 0.2 0.2 0.2 0.2 0.2 0.2 0.2 0.2 0.2 0.2 0.2 0.2 -1. 7 -2.1 -2.4 -2.8 -3.2 -3.5 -3.9 -4.2 -4.6 -4.9 -5.2 -3.2 20.3 20.3 20.2 20.0 19.8 19.6 19.4 19.1 18.7 18.4 18.0 19.8 63.9 66.7 69.5 72.3 75.0 77. i �29 Table 6. Output of mean values (abbreviations in Table 5) and total winter thermoregulatory energy (HEATNRG) using weather data from Bonny Reservoir, 1982-83. Mass RMR Averages for the winter period MAXT MINT LCT TACCL 0.800 0.850 0.900 0.950 1.000 1.050 1.100 1.150 1.200 1.250 1.300 0.999 63.9 66.7 69.5 72.3 75.0 77.7 80.3 82.9 85.5 88.1 90.6 74.9 10.6 10.6 10.6 10.6 10.6 10.6 10.6 10.6 10.6 10.6 10.6 10.6 -7.2 -7.2 -7.2 -7.2 -7.2 -7.2 -7.2 -7.2 -7.2 -7.2 -7.2 -7.2 1.2 1.2 1.2 1.2 1.2 1.2 1.2 1.2 1.2 1.2 1.2 1.2 -1.1 -1.5 -1.8 -2.2 -2.6 -2.9 -3.3 -3.6 -4.0 -4.3 -4.6 -2.6 HEAT 6.1 6.0 5.8 5.6 5.4 5.2 5.0 4.8 4.6 4.4 4.2 5.4 HEATNRG 549.9 536.8 522.2· 506.5 489.5 471.8 453.4 434.7 415.7 396.5 377 .6 489.8 These preliminary results suggest that thermoregulatory costs for mallards at Bonny Reservoir may be considerable, particularly for mallards of low body mass. 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Body weights and carcass composition of the common eider. Wildfowl 27:115-122. Owen, R. B., Jr., and K. J. Reinecke. 1979. Bioenergetics of breeding dabbling ducks. Pages 71-93 in T. A. Bookhout, ed., Waterfowl and wet1ands--an integrated review. La Crose Printing Co., La Crosse, Wisc. Parton, W. J., and J. A. Logan. 1981. A model for diurnal variation in soil and air temperature. Agric. Metero. 23:205-216. �31 Pearson, T. G. 1934. Feeding wild ducks in a crisis. Bird-Lore 36:143. Prince, H. H. 1979. Bioenergetics of post-breeding dabbling ducks. Pages 103-117 in T. A. Bookhout, ed., Waterfowl and wetlands--an integrated review. -raCrosse Printing Co., LaCrosse, Wisc. Raveling, D. G. 1979. The annual cycle of body composition of Canada geese with special reference to control of reproduction. Auk 96:234-252. Reinecke, K. J. 1977. The importance of freshwater invertebrates and female energy reserves for black ducks breeding in Maine. Ph.D. Thesis, Univ. Maine, Orono. 113 pp. Ringelman, J. K., and J. R. Longcore. 1980. Computer simulation models as tools for identifying research needs: a black duck population model. Trans. Northeast Sect. The Wildl. Soc. 37:182-193. ____~----, and M. R. Szymczak. 1985. A physiological condition index for wintering mallards. J. Wildl. Manage. 49:564-568. Seber, G. A. F. 52:249-259. 1965. A note on the multiple-recapture census. Biometrika Smith, L. G., and H. H. Prince. 1973. The fasting metabolism of subadult mallards acclimatized to low ambient temperatures. Condor 75:330-335. Stormer, F. A., and D. H. Johnson. 1986. Introduction: biometric approaches to modeling. Pages 149-169 in J. Verner et al. eds., Wildlife 2000: Modeling habitat relationships of terrestrial vertebrates. Univ. Wisconsin Press, Madison, Wisc. Szymczak, M. R. and J. F. Corey. 1976. Construction and use of the Salt Plains duck trap in Colorado. Colorado Div. Wildl., Div. Rep. 6. 13 pp. Trautman, M. B., W. D. Bills, and E. R. Wickliff. 1939. Winter losses from starvation and exposure of waterfowl and upland game birds in Ohio and other northern states. Wilson Bull. 51:86-104. Verner, J., M. L. Morrison, and C. J. Ralph, eds. 1986. Wildlife 2000: Modeling habitat relationships of terrestrial vertebrates. Univ. Wisconsin Press, Madison, Wisc. Whyte, R. J., G. A. Baldassaree, and E. G. Bolen. 1986. Winter condition of mallards on the southern High Plains of Texas. J. Wildl. Manage. 50:52-57. , and E. G. ---------condition indices Bolen. 1984. Variation in winter fat depots and of mallards. J. Wildl. Manage. 48:1370-1373. Wooley, J. B., Jr., and R. B. Owen, Jr. 1977. Metabolic rates and heart ratemetabolism relationships in the black duck (Anas rubripes). Compo Biochem. Physiol. 57:363-367. �32 Prepared bY~1(.~ JacRingean Wildlife Researcher �33 APPENDIX PROGRAM 1 BIM INTEGER DATE,DUCKS,SEX REAL BoDYWT(100) ,RMR(100) ,METBWT(100) ,LCT(100) ,TACCL(100) +,T(24) ,HEAT(lOO) REAL MAXTEMC-13:100) ,MINTEM(-13:100) ,AVETEM(-13:100) REAL NDAY REAL TLCT(12) ,THEAT(12) ,XLCT(12) ,XHEAT(12) DATA TMAXT,TMINT,TTACCL/O.,O. ,0.1 OPEN (UNIT=2, FILE='COL081.DAT') OPEN (UNIT=1, FILE='WGTS.DAT') DUCKS = SUM = 0. SEX = 1 WRITE(4,15) 15 FoRMATC'l'I!15x, '****** COLORADO, WINTER 1982-83 ******'///) ° TLCT(IN)=O. THEAT(IN)=O. 13 CONTINUE READ IN BODY WEIGHT DATA DO 100, N = 1,12 READ(1,1,END=901) BODYWTCN) DUCKS = DUCKS + 1 1 FORMAT (F4.3) 100 CONTINUE READ IN TEMPERATURE DATA 901 DO 101. M = -13,90 READ (2,2) MAXTEM(M) ,MINTEM(M) 2 FORMAT (6X,2F3.0) ~ CONVERT FARENHEIT TEMPERATURES TO CELCIUB MAXTEM(M) - 5.0 I 9.0 * (MAXTEM(M) - 32.0) MINTEM(M) = 5.0 / 9.0 * (MINTEM(M) - 32.0) AVETEM(M) = (MAXTEM(M) + MINTEMCM» / 2.0 IF (M .GE. 1) THEN TMAXT = TMAXT + MAXTEM(M) TMINT = TMINT + MINTEM(M) END IF 101 CONTINUE C CALCULATE 902 DO 102! L TEMPERATURE = OF ACCLIMATION 1,M DO 103, K - -14,-1,1 SUM = SUM + AVETEM(L+K) 103 CONTINUE TACCL(L) = SUM / 14.0 TTACCL = TTACCL + TACCL(L) SUM = O. 102 CONTINUE C MAIN LOOP C C CALL SUBROUTINES DUCKS = NUMBER OF INDIVIDUALS IN THE POPULATION DATE = NUMBER OF DAYS SINCE 1 DECEMBER C CALCULATE JULIAN DATE �34 NDAY - REAL(DATE) + 334. NDAY - REAL(DATE) - 31. TMX - MAXTEMCDATE) TMN - MINTEM(DATE) c CALL BASAL(DUCKS,BODYWT,METBWT~RMR) CALL LCTEMP(DUCKS,DATE,METBWT,MAXTEM,MINTEM,AVETEM,LCT, -l- l" ACC:l_ 'J -rL..c~-r) CALL TEMPSCTMX?TMN,NDAY,T) CALL HETREG(DUCKS,SEX,METBWT,LCT,T,HEAT,THEAT) L, V,JF~I ;;,,1[';: (/j, 'J 40:t)l)(lTE WRITE(4,403) BODYWT(K) ,RMR(K) ,MAXTEM(DATE) , MINTEM(DATE) ,TACCL(DATE) ,LCT(K) ,HEAT(K) + :I.();S TE I Tt:: (,;;.'J 4(2) corrr I i\IUE 10;5 Cot\IT I!\!UE 401 FORMAT(!!T13, 'TEMPERATURE AND THERMOREGULATORY DATA FOR DAY', -1-13//) 402 FORMAT (T9, 'WEIGHT',5X, 'RMR',5X,'MAXT',4X, 'MINT',5X, 'TACCL', +5X, 'LeT' ,f.~X, 'HE{-~T' /') 403 FORMAT(T9,F5.3,4X,F5.1,4X,F5.1,3X,F5.1,5X,F4.1!4X,F5.1,3X,F3.1) !/JF;: I -rE~ ( l+ \!,IF': ITE " l+() 4. ) 40:3;' XMAXT = TMAXT/90. XMINT = TMINT/90. XTACCL = TTACCL/90. DO 1. 2? 1\11",1 = 1, i 2 XHEAT(NN) = THEAT(NN) 190. XLCT(NN) = TLCT(NN)!90. WRITEC4,4(6) BODYWTCNN) ,RMR(NN) ,XMAXT,XI1INT,XTACCL,XLCT(NN), + XHEAT(NN) ,THEATCNN) 404 405 + 406 (.:1. , FORMATe/I/,T??, '*** AVERAGES FOR THE WINTER ?ERIOD ***'/!) FORMAT(T9, 'WEIGHT',5X, 'RMR',4X, 'MAXT·~4X. 'MINT'~~X. 'TACCL', 51, 'LCT' ,4X, 'HEAT',4X, 'HEATNRG'/) FORMAT(T9,F5.3,4X,F5.1,4X,F4.1,3X,F5.1,5X,F4.1,4X,F5.1.3/, SUBROUTINE j,OO BASAL (DUCKS~80DYWT,METBWT,RMR) REAL BODYWT(100) ,RMR(lOO) ,MET8WT(100) DO 100, II = 1,DUCKS MET8WT(!I) = (BODYWT(II»**.72 RMR(II) = 75.0 * METBWT(II) currrIl\!UE: I~ SUBROUTINE LCTEMP(DUCKS,DATE,METBWT,MAXTEM,MINTEM,AVETEM, "!-L_C:''- ~ -r{~C:C:L '.1 ~rL[~~r) INTEGER DATE,DUCKS REAL TACCL(100) .LCTCi.OO).METBWT(100) ,TLCT(12) DO 20, ] = 1,DUCKS LCTe]) = (.67 * TACCLCDATE) - 3.3) + (-:1.0.0* METBWT(J) TLCT(J) = TLCT(J) + LCT(J) 20 COt'H INUE::~ :l 00 F':ETUF\i\~ + 10.0) �SUBROUTINE 35 TEMPSITMX,TMNjNDAY,T) DATA A,B1C,APHl/l.52,2.00,-O.18.0.698! THIS SUBROUTINE CALCULATES AIR TEMPERATURES BY HOURLY BASED ON A MODEL BY PARTON AND LOGAN (1981, AGRIC. METEOR. 23) TMX = MAXIMUM TEMPERATURE TMN = MINIMUM TEMPERATURE T(I) = TEMPERATURE AT SPECIFIED I....!r» I lr.:·, A - TIME LAG IN MAXIMUM TEMPERATURE AFTER NOON B = COEFFICIENT THAT CONTROLS TEMPERATURE DECREASE AT NIGHT C = TIME LAG FOR THE MINIMUM TEMPERATURE AFTER SUNRISE MDAY = JULIAN DATE APHI = LATITUDE (RADIANS) **NOTE** -- APHI = CLATITUDE/360)*2*PI CALCULATE DAY LENGTH AND NIGHT LENGTH ADELT=.4014*SIN(6.28*(NDAV-77.)!365.l TEM1=1.-(-TAN(APHI)*(ADELT»**2 : 1'__"_1: C I: r-> I••~. L \. TEM2=(-TANCAPHI)*TAN(ADELT» AHDU=ATAN2(TEM1,TEM2) ADY=(AHOU/3.14)*24 DD 4 C I I:::: 1 'j 21.1· DETERMINE IF THE HOUR IS DURING THE DAY OR NIGHT E:)3::::12. /2. ··-{Y()"-/ +C BE:::: 12.+t:IDV/2. IFCBT.GE.BB.AND.BT.LE.BEl GO TO 3 CALCULATE TEMPERATURE FOR A NIGHTTIME IF(BT.GT.BElBBD=BT-BE IF(BT.LT.BB)BBD=(24-BE)+BT HOUR TSN=(TMX-TMN)*SIN«3.14*DDY)/(ADY+2*A»+TMN TCI)=TMN+(TSN-TMN)*EXP(-B*BBD/ANI) so TO 4CALCULATE TEMPERATURE FOR A DAYTIME HOUR C T(I)=(TMX-TMN'*SIN«3.14*BBD)!(ADY+2*A»+TMN END SUBROUTINE HETPEG (DUCKS,SEX,METBWT,LCT.T,HEAT~THEAT) REAL METBWT(lOO) ,LCT(lOO) ,HEAT(100) ,T(24) ,THEAT(12) INTEGER DUCKS,SEX DO 10, KK = 1,DUCKS DD 1.i ~ LL ".::: j 'J :::4 IF (TCLL) .GE. LCT(KK» GO TO 11 IF (SEX .EO. 1) THEN HEAT(KK) - (.12*(LCT(KKl-T(LL»*METBWT(KK»+HEAT(KK) ELSE HEAT(KK) - (.11*CLCT(KK)-T(LL»*METBWT(KK»+HEATCKK) EI\W IF" CONT I,,!ut:: THEAT(KK) = THEAT(KK) 10 CiJNTINUE i 1. 1 J. EI\ID + HEATCKK) ��37 Colorado Division of Wildlife Wildlife Research Report April 1987 JOB PROGRESS REPORT State of Colorado ------------------------------------ Project 01-03-045 (W-88-4) Work Plan 1 Job Title: 16 --~-- Field-feeding Ecology of Mallard Ducks Period Covered: Author: : Job Avian Research 01 July 1985 through 31 December 1986 James K. Ringelman Personnel: J. F. Corey, J. K. Ringelman, M. R. Szymczak, Colorado Division of Wildlife ABSTRACT Feeding trials using pen-reared mallards (Anas platychynchos) were conducted in standing corn stubble, disced stubble, and snow-covered stubble during January-March 1986. Mallard feeding rates on corn kernals varied by stubble treatment and corn density, which ranged from 100 to 1200 kg/ha. Feeding rates also differed by individual birds across all corn densities. Search time over and above handling time decreased feeding efficiency by 43% at normal waste corn densities, but mallards were still able to acquire up to 9 g corn/minute in standing stubble. A 2 cm snowfall further reduced feeding rates by 74%, decreasing intake to about 2 g/minute. Discing stubble containing only kernals caused food consumption rates to decrease to fractions of a gram/minute. ��39 FIELD-FEEDING ECOLOGY OF MALLARD DUCKS James K. Ringelman Man has drastically altered waterfowl habitat during the last 100 years. Whereas widespread drainage of inland wetlands and alteration of coastal areas have had detrimental effects, water development projects and irrigated agriculture have created new wintering and migration habitat. In Colorado, relatively few waterfowl were present in winter until irrigated cereal grain crops were introduced in the mid-1800's (Steinel 1926:174). Within a decade, waterfowl were being "short-stopped" along their traditional migratory routes, attracted by newly-created reservoirs and abundant waste cereal grain (Buller 1975). Wintering waterfowl, particularly the mallard, became so plentiful in the 1940's that a special season was held on mallards in Colorado during 1942-43 to help alleviate damage to grain crops (Wagar 1946). Mallards remain the most common wintering duck species today and accounted for 65% of the total duck harvest in 1981 (Colo. Div. Wildl. 1981). D Wintering mallards, like all inland wintering waterfowl populations, are highly depending on waste products of agriculture for food (Girard 1941, Reed 1971, Sugden et al. 1974). In Colorado and adjacent states to the east and south, waste corn comprises from 50 to 90% of the winter diet (Gordon 1981, Jorde 1981, Baldeassaree and Bolen 1984). Despite the importance of corn to waterfowl, little is known of the factors that regulate its use by ducks. Harvester efficiency, slope of the land, and corn moisture content all affect the amount of corn remaining in the field after harvest (Baldassarre et al. 1983). This "wastage", typically 3-5% of the pre-harvest corn crop, contributes the bulk of cereal grains necessary to sustain waterfowl throughout the winter. The amount of this wastage used by ducks varies depending on post-harvest treatments including discing, plowing, burning, and cattle grazing (Jorde 1981, Baldassarre and Bolen 1984). Economic forecasters predict radical changes in eastern Colorado irrigated agriculture within the next 7-17 years. Four factors will play important roles in this transformation: (1) depletion of ground water aquifers, (2) rising energy costs and related impacts on the cost of operating pumps, (3) conversion of farmland to urban use, especially along the Front Range, and (4) corn prices. Nearly 13% of all Colorado cropland is planted to irrigated corn and it ranks second behind wheat in crop value (Colo. Dep. Agric. 1982). However, under scenarios of rising energy costs and groundwater depletion, corn is the first irrigated crop to become economically unfeasible (Sharp 1979). The break-even point is when electricity to power pumps reaches $0.12/kwh (Oamek 1981:64), and some scenarios predict the disappearance of irrigated corn from the region overlying the Ogallala Aquifer by 1990 (Young et al. 1982:129). Front Range area croplands, which benefit from gravity flow irrigation water transported by canals, are under increasing pressure from urban growth. In the last 2 decades, Colorado has seen over 1.3 million acres converted from agricultural use (Colo. Dep. Agric. 1980). Much of this development occurs on irrigated land (U.S. Dep. Agric. 1978). The key to preserving wintering waterfowl and subsequent harvest opportunity is the management of water areas coupled with more intensive land management practices, particularly in relation to cornfields on Division of Wildlife and �40 private lands. Based on historical evidence and knowledge of wintering waterfowl food habits, it is believed that loss of cornfields will result in major shifts of birds out of an area. In the case of the Front Range, this would result in severe loss of hunting opportunity where demand is greatest. This would also be true elsewhere in eastern Colorado where hunting pressure and demand is somewhat less pronounced, but nevertheless important. So little is kno~m of how mallards use cornfields and waste grain that it is not possible to determine the area of cornfields needed to sustain a given mallard population. In the past, such information was unneeded because of abundant corn crops. However, with the expected declines in irrigated agriculture in the near future, these data will be necessary to maintain wintering waterfowl. Past field-feeding studies have focused on the timing of flights and the relationships between flight times and weather (Hochbaum 1955, Swinebroad 1956, Bossenmaier and Marshall 1958, Winner 1959, Gordon 1981, Jorde 1981), or on ways to alleviate crop damage by ducks (Gollop 1950, Hammond 1952, MacLennan 1973). Only recently have studies documented the potential value of post-harvest treatments in making fields more attractive to ducks. Jorde (1981:50) described a commensal relationship between field-feeding mallards and cattle which exposed corn in times of heavy snowfall. Baldassarre et al. (1983) reported that treatments such as burning of discing of fields with moderate amounts of waste corn made these fields more attractive to ducks than untreated cornfields of standing stubble with waste corn densities many times greater. This suggests an "optimal foraging" strategy of field-feeding ducks in which feeding rate is balanced with other considerations such as search time, handling time, and distance to field. A large body of information on optimal foraging in birds exists in the ecological literature, but thus far these findings have not been applied to the management of field-feeding ducks. Efforts must be made to evaluate methods of making waste corn more available to waterfowl, thereby optimizing feeding rates and winter physiological condition. P. N. OBJECTIVES 1. Identify the relationships between feeding rates of mallards and waste corn density, group foraging size, sex, and age of duck. 2. Relate post-harvest cornfield treatments to foraging efficiency. 3. Document the foraging habitat requirements of wintering mallards. 4. Develop a model for post-harvest management of cornfields for wintering mallards. SEGMENT OBJECTIVES 1. Contact local farmers to arrange a cornfield lease. ears from 1 ha of corn during late summer. Hand pick and remove 2. In fall 1986, "harvest" the cornfield plot using standard techniques leaving canes, leaf litter, and stubble in place. �41 3. During January-March 1986, quantify the feeding rates of mallards as a function of kernal density in standing stubble. 4. During January-March 1986, quantify the effect of snowfall on feeding rates in stubble as a function of snow depth and kernal density. 5. During March-April 1986, quantify feeding rates of mallards on disced stubble plots that were pre-dosed with kernals at several densities. METHODS Farmers in the Fort Collins area were asked if they would lease their cornfield for field-feeding experiments. Eldon Edwards, a farmer 1 km northeast of Wellington, agreed to the use of his field. Whole ears were hand-picked from an area about 8 x 500 m in area during early September 1986. Corn was harvested in this field in mid-September, and permanent plot markers were established shortly thereafter. An electric fence was erected to exclude cattle from the plots. Panels measuring 1.2 x 2.0 m were used to enclose square plots 7.3 m on a side. Whole kernal corn was broadcast uniformly within the 53.3-m2 enclosure using a hand operated seed spreader. Mallards used in the experiments were deprived of food 24 hours before a trial. Eight (occasionally 9) birds of each sex were weighed to the nearest gram on an electronic digital balance immediately before each trial. Mallards were released en masse into the enclosure, allowed to feed for precisely 5 minutes, then rounded up and reweighed. The amount of corn ingested was assumed to equal the difference between starting and ending body mass. Feeding rates in standing stubble were measured at initial densities of 100, 200, 400, 800, and 1200 kg/ha. Since mallards depleted a significant proportion of available corn, particularly at lower initial densities, an "ending density" was calculated by subtracting the total corn consumption for the trial from the initial dosage. An "average density" was then computed as the mean of the initial and ending densities. Average density was used to test for differences in feeding rates by density and treatment. Birds with feeding rates above the median were used to quantify maximum feeding rates. Six standing stubble plots were pre-dosed at densities ranging from 284 to 1094 kg/ha, then disced on 18 March 1986. Enclosures were re-erected on the original plots, and feeding trials were conducted in the standard manner. The effect of 2-2.5 cm of fresh snow on the feeding rates of mallards in standing stubble was evaluated on 6 and 11 February 1986. Plots were pre-dosed at 1200 kg/ha (6 Feb) and 400 kg/ha (11 Feb). Feeding trials were then performed as previously described. Twelve feeding trials were conducted in standing stubble. A wide variation in feeding rate within density, reflecting differential foraging success among individual birds, was apparent in every trial. In considering birds only above the median, variation was greatest at corn densities from 200 to 800 kg/ha in standing stubble (Fig. 1). In contrast, variation in feeding rates in disced stubble (Fig. 2) and snow (Fig. 3) was highest at densities above �42 20 ~ z .- 15 ~ ......•• CJ 0 "'" § w le::( a: 10 8 Q 0 0 Cl W W 0 5 @ § § § 0 ~ 0 ~ 0 9 ! ~ 0 @ CJ Z I..L. @ ~ § 8 0 0 ~ ~ 0 ~ 0 0 0 0 200 MEAN 600 CORN DENSITY 1000 (KG/HA) Fig. 1. Variability in feeding rates of mallards in standing stubble dosed with a known density of corn kernals. Only ducks above the median feeding rate in each trial are depicted. �43 20 ~ z - 15 ~ .•••...•• C} "'"" W t- « a: 10 C) z o w W LL 5 0 § 00 0 00 e ~ 0 0 ®> ® 200 MEAN 600 CORN DENSITY § 1000 (KG/HA) Fig. 2. Variability in feeding rates of mallards in disced stubble pre~dosed with a known density of corn kernals prior to discing. Only ducks above the median feeding rate in each trial are depicted. �44 20 ~ z - 15 ~ •..•..••• C) ....._, W J- < .0: 10 C) Z 0 W W u, @ 5 0 i 600 200 MEAN Fig. 3. Variability by 2-2.5 cm of snow. are depicted. CORN DENSITY 1000 (KG/HA) in feeding rates of mallards in standing stubble covered Only ducks above the median feeding rate in each trial �45 1000 kg/ha. When averaged across all densities, feeding rates differed among birds (p = 0.005; Table 1). The feeding rate of the highest bird (80) was 90% greater-than that of the lowest bird (10). Trends in individual feeding rates by sex were not apparent. Table 1. Feeding rate of individual mallards in standing stubble, averaged across all corn densities and trials. Bird 1/ 80 4 6 82 7 77 5 2 11 3 1 12 78 8 9 10 Mean feeding rate (g/min)a 7.84 7.55 7.51 7.41 7.30 7.18 6.93 6.92 6.87 6.03 5.55 5.46 5.28 4.90 4.39 4.13 Sex a a,b a,b a,b a,b a,b a,b,c a,b,c a,b,c a,b,c,d a,b,c,d a,b,c,d a,b,c,d b,c,d c,d d F M N F M F M F M F F M F M M M a~1eans with the same letter do not differ (p> 0.05). Six feeding trials were conducted in disced corn stubble. No differences in the performance of individual birds were detected in these trials (p = 0.73). Since only 2 trials were conducted in snow, no comparisons in individual foraging rates could be made for this treatment. Mallards were efficient foragers in standing corn stubble, averaging about 9 gms of corn kerna1s/minute (corrected for average corn density) at densities above 800 kg/ha (Fig. 4). The maximum rate of change in the functional response curve occurred at about 200 kg/ha. Means of feeding rates within average density separated poorly except at densities below 160 kg/ha. Discing corn stubble drastically decreased corn kerna1 availability. Feeding rates declined to an average high of about 2 g/min at initial densities of nearly 1100 kg/ha (Fig. 5). Snowfall of 2-2.5 cm had a similar effect, reducing feeding rates to 4.86 g/min at an initial density of 1167 kg/ha, and to 1.90 g/min at an initial density of 389 kg/ha (Fig. 6). �46 8 a • ,..... Z - 6 .b ~ .•••...•• o • be c • '-" • cd W .cd •••• « a: o z 4 (:) W W U. 2 200 600 1000 MEAN CORN DENSITY (KG/HA) Fig. 4. Mean feeding rates of mallards in standing stubble as a function of average kernal density. Means with the same letter do not differ (f >0.05). The functional response curve was fitted by hand. �47 ,...... z ~ 6 .....•. C) ~ w 4 I- « • a: {!J a 2 z - c 0 c c •• W W u, 200 • 600 1000 MEAN CORN DENSITY (KG/HA) Fig. 5. Mean feeding rates of mallards in disced stubble as a function of average kernal density. Means with the same letter do not differ (~ >0.05). The functional response curve was fitted by hand. �48 '"z - 6 ~ .•..•.•• • (!J ~ w a 4 I- « a: (!J • 2 b Z 0 W W IJ.. 200 600 1000 MEAN CORN DENSITY (KG/HA) Fig. 6. of snow. Mean feeding rates of mallards in standing stubble covered by 2-2.5 cm Heans with the same letter do not differ (~>0.05). �49 DISCUSSION Optional foraging theory (Pyke et al. 1977), when applied to field-feeding mallards, can be helpful in understanding the behavior of feeding birds and making management decisions about post-harvest treatments. Although handling time can be an important factor in the selection of barley and wheat strains (Clark et al. 1986), data on feeding rates for corn kernals on dirt substrate indicate that handling time is minimal (Colo. Div. Wildl., unpubl. data). The feeding rate in standing stubble was 33% lower at 1200 kg/ha than the feeding rate on dirt at the same corn density (Table 2), suggesting that about one-third again as much time is needed to forage in standing stubble owing to increased search time. In the range of normal waste corn densities (200 to 400 kg/ha) (Baldessarre and Bolen 1984), search time resulted in a 43% decrease in feeding efficiency: Since mallards consume from 34 to 64 g of corn/feeding bout (Whyte and Bolen 1985), minimum feeding time in stubble would be about 4.7-8.9 minutes. Table 2. Estimated foraging rates of mallards by density of corn kernals and treatment. Density (kg/ha) 100 200 300 400 600 800 1000 1200 Bare ground 10.3 11.8 12.7 13.3 13.8 13.9 13.9 14.0 Foraging rate (g/min) by treatment Stubble Disced stubble 4.7 6.3 7.2 7.8 8.6 9.0 9.3 9.4 0.1 0.1 0.3 0.7 1.7 Snow 2.0 5.0 Assuming 2 field-feeding flights/day, total daily feeding time could be less than 20 minutes (not including flight time). Thus, under normal conditions, mallards on high plains wintering areas can afford to adopt a strategy of time minimization (Pyke et al. 1977), thereby conserving energy and affording time for courtship as well as other, less demanding behaviors. The ease and efficiency with which mallards forage in standing stubble changes dramatically with only a 2-cm snowfall (Table 2). At corn densities of 400 kg/ha, feeding rates in stubble decrease 74% from 7.8 g/min to 2.0 g/min: Given that the basal metabolic rate (BMR) of a mallard can be modeled allometrically as 75 x mass 0.72 (Owen and Reinecke 1979), then a 1.2 kg mallard would have a BMR of 86 kcal/day. Daily existence energy costs could exceed 3.5 times BMR in severe winter weather (J. K. Ringelman, unpubl. data), thereby bringing total daily energy requirements (DEE) to 300 kcal/day •. At an estimated metabolizable energy of 3.5 kcal/g corn and a maximum feeding rate of 2 g/min, nearly 45 minutes would be required to meet DEE. During periods of heavy snowfall (>10 cm), and severe cold, mallards may forage throughout the day (J. K. Ringelman, unpubl. data), suggesting that feeding rates fall �50 far below 2 g/min and that several hours are needed to attain or approach DEE requirements. Discing reduces the abundance of available kernals by 93% (Baldessarre et al. 1983), therefore, the 300 kg/ha density of kernals prior to discing was reduced to about 20 kg/ha for the low density trial in disced stubble. At this density, birds acquired less than 1 g of corn during the 5-minute trial (Table 2). Baldassarre et al. (1983) suggested that disced cornfields were often preferred by field-feeding mallards because litter was largely eliminated and the discs shattered whole ears making more kernals available. Since whole ears were not introduced into feeding trials this year, the overall effect of discing on mallard feeding rates cannot be quantified. Feeding trials next segment will emphasize feeding responses to whole ears. All feeding rate curves will be described mathematically. A computer model will be developed to explore changes in feeding rates and net energy acquisition by treatments. This model will allow estimation of winter carrying capacities based on average waste corn density, post-harvest treatments, and snowfall. LITERATURE CITED Baldassarre, G. A., and E. G. Bolen. 1984. Field-feeding ecology of waterfowl wintering on the southern high plains of Texas. J. Wildld. Manage. 48:63-71. ______ ~--' R. J. Whyte, E. E. Quinlan, and E. G. Bolen. 1983. Dynamics and quality of waste corn available to postbreeding waterfowl in Texas. Wildl. Soc. Bull. 11:25-31. Bossenmaier, E. F., and W. H. Marshall. 1958. Field-feeding by waterfowl in southwestern Manitoba. Wildl. Monogr. 1. 32 pp. Buller, R. J. 1975. Redistribution of waterfowl: influence of water, protection, and feed. Proc. Int. Waterfowl Symp. 1:143-154. Clark, R. G., H. Greenwood, and L. G. Sugden. 1986. Influence of grain characteristics on optimal diet of field-feeding mallards. J. Appl. Ecol. 23:763-771. Colorado Department of Agriculture. 1980. Agricultural land conversion in Colorado. Resour. Analysis Sect., Colorado Dep. Agric., Denver. 8 pp. 1982. Colorado agricultural statistics. Rep. Serv., Denver. Bull. 1-82. 94 pp. Colo. Crop and Livestock Colorado Division of Wildlife. 1981. Colorado small game, furbearer and varmint harvest, 1981. Colorado Div. Wildl., Denver. 228 pp. Girard, G. L. 1941. The mallard: Wildl. Manage. 5:233-259. its management in western Montana. J. �51 Gollop, B. J. 1950. Report on investigation of damage to cereal crops by ducks in the prairie provinces. Unpub1. Rep., Can. Wi1d1. Serv., Ottawa. 11 pp. Gordon, D. H. 1981. Condition, feeding ecology, and behavior of mallards wintering in northcentral Oklahoma. M.S. Thesis, Oklahoma State Univ., Stillwater. 68 pp. Hammond, M. C. 1952. Waterfowl damage and control measures, Lower Souris Refuge and vicinity. Unpub1. Rep., U.S. Dep. Inter., Fish and Wi1d1. Serv., Washington, D.C. 5 pp. Hochbaum, H. A. 1955. Travels and traditions of waterfowl. Press, Minneapolis. 301 pp. Univ. Minnesota Jorde, D. G. 1981. Winter and spring staging ecology of mallards in southcentral Nebraska. M.S. Thesis, Univ. North Dakota, Grand Forks. 116 pp. MacLennan, R. 1973. A study of waterfowl crop depredation in Saskatchewan. Can. Wi1d1. Serv., Saskatoon, Saskatchewan. Wi1d1. Rep. 2. 38 pp. Oamek, G. E. 1981. Economic adjustments to rising energy costs: the case for pump irrigators. M.S. Thesis, Colorado State Univ., Fort Collins. 90 pp. Owen, R. B., Jr., and K. J. Reinecke. 1979. Bioenergetics of breeding dabbling ducks. Pages 71-93 in T. A. Bookhout, ed. Waterfowl and wet1ands--an integrated review. La Cross Printing Co., La Crosse, WI. Pyke, G. H., H. R. Pulliam, and E. L. Charnov. 1977. Optimal foraging theory: a selective review of theory and tests. Quart. Rev. Bio1. 52:137-154. Reed, L. W. 1971. Use of western Lake Erie by migratory and wintering waterfowl. M.S. Thesis, Michigan State Univ., East Lansing. 71 pp. Sharp, R. L. 1979. Economic adjustments to increasing energy costs for pump irrigation in northeastern Colorado. M.S. Thesis, Colorado State Univ., Fort Collins. 72 pp. Steinel, A. T. 1926. History of agriculture in Colorado. Fort Collins, Colo. 659 pp. State Agric. Co11., Sugden, L. G., W. J. Thurlow, R. D. Harris, and K. Vermeer. 1974. Investigations of mallards overwintering at Calgary, Alberta. Can. Field-Nat. 88:303-311. Swinebroad, J. 1956. Some aspects of the role of weather in bird migration. Ph.D. Diss., Ohio State Univ., Columbus. 315 pp. U.S. Department of Agriculture. 1978. Urbanization of rural lands in the northern Colorado Front Range. U.S. Dep. Agric., Washington, D.C. 23 pp. Wagar, J. V. K. 1946. Colorado's duck-damage, grain-crop problem. North Am. Wi1dl. Conf. 11:156-162. Proc. �52 Whyte, R. J., and E. G. Bolen. 1985. Corn consumption by wintering mallards during morning field-flights. Prairie Nat. 17:71-78. Winner, R. W. 1959. Field-feeding periodicity of black and mallard ducks. Wi1d1. Manage. 23:197-202. Young, R. A., L. R. Conklin, R. A. Longenbaugh, and R. L. Gardner. 1982. Energy and water scarcity and the irrigated agricultural economy of the Colorado high plains: direct ecnomic and hydrologic impact forecasts (1979-2020). Colorado Water Resour. Res. lnst., Colorado State Univ., Fort Collins. 362 pp. Prepared by ~ It'~ Ja K. Ringelm Wildlife Researcher C J �53 Colorado Division of Wildlife Wildlife Research Report Avril 1987 JOB PROGRESS REPORT·· State of Colorado Project 01-03-045 Work Plan 1 Job Title: 17 --- Habitat Use of Wintering Mallards Near the Front Range of Colorado Period Covered: Author: : Job Avian Research 01 July 1985 through 31 December 1986 Michael R. Szymczak Personnel: J. Barnett, J. Corey, C. Crawford, J. Ringelman, L. Rogstad, W. Russell, and M. Szymczak, J. alterman, G. Skiba, Colorado Division of Wildlife ABSTRACT Some mallards (Anas platyrhynchos) radio-marked at Chestnut Slough in December 1985 periodically used Kodak Ponds, north of the proposed study area. Therefore, the study area bordered by Interstate Highway 25 on the west, State Highway 34 on the north, State Highway 66 on the south and Milton and Latham reservoirs on the east with Chestnut Slough as the epicenter was expanded northward to include Kodak Ponds and the Poudre River drainage south of State Highway 392. Seventeen of 25 birds marked a Chestnut Slough on 3 December 1986 generally remained in the study area during December. There was considerable use of a non-hunted portion of the Big Thompson River during daylight hours by birds using Chestnut Slough as a nocturnal roost; movement was not dependent upon hunting activity at Chestnut Slough itself. Weight loss of captive mallard groups fed only yellow corn or yellow corn plus 1 snail (Physa sp., 0.03 g) twice a week during a 6-week period was not significant whereas groups fed yellow corn supplemented with 4 (0.11 g) and 8 (0.22 g) snails respectively, twice a week lost weight (p < 0.01 and P < 0.05). However, birds fed hen lay pellets during the same period gained weight (p < 0.01). Weight loss in all groups (pooled) fed corn was significant (~< 0:-01). ��55 HABITAT USE OF WINTERING MALLARDS NEAR THE FRONT RANGE OF COLORADO Michael R. Szymczak INTRODUCTION Irrigated agriculture along the Front Range of northcentral Colorado has created a diversity of wetland habitats ranging from large reservoirs to small irrigation seep ditches. Riparian areas compliment these man-made wetlands, and together with abundant cereal grain crops, provide attractive habitats for wintering waterfowl. Mallards that winter along the Front Range originate from northern breeding grounds and mountain park production areas in Colorado. Traditionally, the Front Range region (Fort Collins-Greeley-Denver) has supported about 25% of the post-hunting season mallard population in the South Platte Valley. However, this percentage has been declining in recent years (Colo. Div. Wildl., unpubl. data). In the late 1950's, a period of high continental mallard populations, and even in the early 1960's when mallard populations were at a low level, the Front Range region wintered about 75,000 mallards. January inventories from 1979 to 1983 indicated an average of 30,000 mallards in this area. The Front Range region has undergone considerable change in recent decades. Sprinkler irrigation systems have become widespread, resulting in increases in the area and yield of small grain crops, particularly corn (Buller 1975). Whereas irrigation has made land more productive, urbanization has resulted in the loss of over 400,000 ha of prime agricultural land (Colo. Dep. Agric. 1980). Late fall and winter duck distribution has changed in response to these factors. These changes reflect the suitability of wetland and upland waterfowl habitats for meeting the complex requirements of wintering ducks. Most apparent among the needs of wintering mallards are waste cereal grains, particularly corn (Gordon 1977, Jorde 1981). Corn is a high energy, high carbohydrate food, yet is nutritionally unbalanced (lacking calcium and the amino acids lysine, methionine, and tryptophan) (Baldassarre et al. 1983). Thus, mallards must supplement their corn diets with other foods, notably aquatic invertebrates (Jorde et al. 1983). These invertebrates occur in low densities in wetlands with specific chemical and biotic characteristics (White 1982). The physical properties of wetlands are also important. Large reservoirs-are used for roosting and as an arena for courtship and pairing. By concentrating birds, they may also synchronize physiological changes (such as ovary recrudescence preparatory to breeding) or serve as information centers for feeding flights (Weatherhead 1983). Small wetlands are no less valuable, providing areas for pair isolation and usually a more favorable thermal environment (Jorde et al. 1984). Superimposed on the requirements of mallards during winter are human and weather-induced constraints that limit availability of wetlands. Hunting may dramatically affect distribution of mallards during fall and mid-winter, yet �56 hunting has barely been considered (Kirby et ale 1976) as an ecological component that shapes habitat use by mallards. Sub-freezing temperatures make many wetlands unsuitable during winter. The few open water areas that remain are often heavily hunted. Thus, mallards in northeastern Colorado, a region with seemingly abundant wetlands during fall, have to cope with limited habitat when weather and hunting interact to severely limit wetland availability. Suboptimal wetland habitat is believed to influence winter survival rates of mallards (Hepp et ale 1985) and regulate winter body condition (White 1982), which may be linked to reproductive performance (Krapu 1981). Wintering mallards must satisfy demands of courtship and palrlng, pre-basic molt (females), lipid deposition, and thermoregulation, all within a regime of changing availability of secure, ice-free wetlands. As a result of these physiological, behavioral, and physical constraints, mallards may use only a small proportion of the available wetlands during winter. These relatively few wetlands, whose characteristics and biological significance are largely unknown, are the key to attracting and maintaining wintering mallard populations. The strategic plan of the Colorado Division of Wildlife specifies an objective of increasing duck harvest by 18% in the next 5 years. Habitat, through improvement, lease, purchase, or preservation, has been identified as the vehicle to accomplish this and other objectives. Yet our knowledge of the habitat requirements of wintering mallards is incomplete. A need exists to identify the wetland habitat requirements of mallards, the role of these wetlands in fulfilling mallard nutritional, energetic, and behavioral demands, and the interactions of weather and hunting in modifying the availability of habitat. P.N. OBJECTIVES 1. Relate mallard habitat use to the physical characteristics of wetlands, aquatic and upland plant communities, wetland macroinvertebrate populations, weather, and hunting regimes. 2. Characterize mallard use of wetlands through time budget techniques. 3. Document the composition of the wetland community within the study area, and relate wetland use by mallards to availability. SEGMENT OBJECTIVES 1. Conduct a comprehensive literature search on habitat requirements, food habitats, local movements, condition dynamiCS, and behavior of mallards during winter. 2a. Examine aerial imagery, harvest data, winter waterfowl surveys, and wetland inventory data bases to identify potential study areas. Contact District and Area Wildlife Managers and Biologists with jurisdiction in potential study areas. �57 2b. Survey potential study areas during early, mid-, and late winter to evaluate the suitability of each site. Prime considerations are road systems, access to private land, existence of sizeable mallard populations, diversity of hunting regimes practiced in the area, terrain favorable to a biotelemetry study, and proximity of one or more weather stations. 2c. Select a study area and obtain a detailed set of aerial photographs and maps. 2d. During December, capture and radiomark 6 mallards (3 females, 3 males) to ascertain fidelity to the study area, frequency of movements, and suitability of transmitter design. Refine tracking methodology. 3a. Capture and radiomark a minimum of 20 mallards (sample divided equally by age and sex) during early December. 3b. Monitor the movements of instrumented mallards using aerial and vehicle-mounted tracking systems. Alternate tracking days between (1) relocating each bird as many times as possible during 1 day and (2) intensive tracking of 1 or 2 birds during 1 day. Rotate tracking periods over a 24-hour period, unless habits of birds indicate nocturnal and/or diurnal tracking is unnecessary. 3c. Conduct aerial tracking flights once a week, or more frequently if weather conditions or bird movements warrant. METHODS Selection of Pilot Study Area Winter mallard distribution and band recovery data were examined in conjunction with different wetland complexes along the Front Range. In recent years, mallards in late winter have been concentrated in 3 areas centered around wetlands that have open water regardless of weather conditions. The central wetlands for these areas are Va1mont Reservoir near Boulder, Chestnut Slough near Milliken, and Kodak Ponds near Windsor. Va1mont, a power plant cooling reservoir has numerous small ponds and some large reservoir roosts in the vicinity, but only a small amount of riverine habitat. The number of birds using Va1mont as a winter roost in recent years has been decreasing. Chestnut Slough is an off-channel slough located adjacent to riverine (South Platte River) habitat. There are additional warm winter sloughs, some small ponds, and 2 large reservoirs (Milton and Latham) that are used as winter roosts in the area. One of the Kodak Ponds is along the course of a warm water seep ditch while the other is the result of groundwater seepage from agricultural areas. Kodak Ponds are within a large hunting closure that includes riverine (Poudre River) habitat and 2 small ponds. In the Kodak vicinity are New Windsor Reservoir and Woods Lake which serve as major winter roosts. �58 Valmont Reservoir was rejected as the pilot study epicenter because of increasing urbanization in the area, decreasing mallard population in recent years, and lack of riverine habitat. Kodak Ponds were rejected primarily because they were in a large hunting closure that contained a variety of habitats, including agricultural feeding areas. Thus, sample birds could spend the entire winter within the closure negating the possibility of evaluating the effect of hunting regimes. Chestnut Slough was selected as the epicenter because the variety of habitat and potential variety of hunting regimes on both private and public land in the vicinity. Preliminary study area boundaries were Interstate Highway 25 on the west, State Highway 34 on the north, State Highway 66 and on eastward extension on the south, and Milton and Lower Latham reservoirs on the east. The preliminary boundaries were considered logical because of topographic changes that occurred to the north (winter wheat uplands between the Big Thompson-South Platte, and Poudre River basins) east (grasslands/winter wheat uplands eat of Milton and Latham), and logistical considerations in reference to the size of the study area to the west and south. In addition, band recovery and recapture data of mallards banded post-season at Valmont, Chestnut Slough, and Kodak Ponds indicated limited interchange between the 3 locations with few recoveries east of Greeley. Radio Application Mallards were captured in Salt Plains bait traps (Szymczak and Corey 1976) at Chestnut Slough, 2 miles east and 1 mile south of Milliken, Colorado on 8 December 1985 and 3 December 1986. In 1985, 8 birds (Table 1) were fitted with 19-9 transmitters mounted with a back-pack harness (Dwyer 1972). In 1986, a 24-g transmitter in a back-pack harness was attached to 25 birds (Table 2). All birds were released at Chestnut Slough. Table 1. Physical characteristics, band number, and radio frequency of mallards instrumented at Chestnut Slough, 8 December 1985. Wing length (mm) Condition a index Radio frequency Age/sex Band # Weight (g) AM 1397-16612 -16615 -16616 1304 1442 1223 290 312 292 26.23 27.40 22.43 150.261 150.450 150.820 AF 1397-16605 -16610 -16619 1241 1104 945 280 282 279 30.91 23.68 15.34 150.860 150.931 150.840 IF 1397-16604 -16613 1032 1023 283 275 19.53 20.92 150.280 150.920 aBody fat/fat-free body weight (Ringelman and Szymczak 1985). �59 Table 2. Physical characterisitics, band number, and radio frequency of mallards instrumented at Chestnut Slough, 3 December 1986. Wing length (mm) Conditiona index Radio frequenc! Age/sex Band II Weight (g) .AM 1397-17508 -17509 -17510 -17511 -17515 1054 1068 1236 1173 1208 301 308 300 296 309 12.68 12.06 21.45 19.42 18.53 150.837 151.020 151.126 150.936 151.063 AF 1397-17518 -17521 -17522 -17530 -17538 -17539 -17540 -17541 1140 1021 1111 972 1076 999 1082 1110 281 275 283 281 281 289 284 275 25.78 20.81 23.82 16.51 22.43 16.24 22.05 25.66 151.111 151.189 151.099 150.862 150.946 150.887 151.249 151. 202 1M 1397-17504 -17505 -17506 -17533 -17536 1205 1116 1214 1226 1210 296 287 291 297 285 20.86 18.52 22.23 21.59 23.25 151.174 151.155 151. 714 151.232 151. 732 IF 1397-17519 -17520 -17523 -17524 -17527 -17529 1033 1202 873 985 288 289 282 277 266 271 281 18.42 26.99 10.02 18.24 150.816 150.916 151.142 150.979 150.963 150.995 150.900 896 1060 14.18 21.55 aBody fat/fat-free body weight (Ringe1man and Szymczak 1985). Radio Monitoring Radio signals were received through 4 different antenna systems. A truck mounted 3-e1ement precision directional antenna that enabled precise triangulation was used predominantly. A truck mounted 14-e1ement and a hand held 3-e1ement antenna were also used on the ground whereas aerial tracking was accomplished with a 2-e1ement system (Dodge 1985) using technqiues described by Gilmer et a1. (1981). In winter 1985-86, the majority of the proposed study area was traversed 7 times including 2 aerial flights during which the location of radio-marked birds was recorded. Duck concentration areas outside the proposed study area, particularly to the north, were routinely checked. In addition, daily monitoring of birds at Chestnut Slough was initiated on 14 January 1986 and �60 terminated on 26 February 1986. Inadequate radio signal-strength made finding marked birds away from waterfowl concentration areas difficult, particularly from the ground. Therefore, the routine for ground monitoring was to check known concentration areas for radio-marked ducks. In December 1986, monitoring of radio-marked ducks began on 8 December and continued daily through the month, except during 3 days. Most monitoring centered around Chestnut Slough with the intent of recording the response of birds using the area to hunting and weather changes. A beacon transmitter was placed near Chestnut Slough and permanent telemetry stations were established to allow precise plotting of bird locations in the vicinity. Bird locations throughout the study area were recorded by a Universal Transverse Mercator grid. To measure the responses of radio-marked ducks to hunting on Chestnut Slough, birds present in early morning were monitored continuously until they left the area. Disturbance associated with hunting activity (shots fired, driving, walking, etc.) was also recorded as were weather conditions. Early morning movement by birds around Chestnut Slough on days prior to expected hunts were also recorded. Micronutrient Study Yellow corn, the assumed predominant item in the diet of mallards wintering along the Front Range does not provide nutrients adequate for winter maintenance (Baldassare et ale 1983, Jorde et ale 1983). Corn is particularly deficient in calcium and the amino acids lysine, tryptophan, and methionine (Baldassare et ale 1983). Mallards wintering along the Front Range are thought to supplement their corn diet by ingesting invertebrates obtained from small wetlands. Pond snails (Physa sp.) containing the missing nutrients (Heitmeyer 1985:292) are present in wetlands along the Front Range and are known to be ingested by ducks. Therefore, we evaluated the effects of micronutrients on weight of captive mallards. Five groups of 4 adult mallards each, 2 males and 2 females, were selected on 4 November 1986 from the captive flock and placed in pens that had shelter and open water. The birds were selected so that the mean weights of each group were about equal. The birds were kept on their normal diet, a mixture of whole kernel corn and hen lay pellets during a 2+ week acclimation period. Body condition in terms of fat stores varied when birds were placed in the pens and weight loss during the acclimation period varied from less than 1 to 28%. Treatments were randomly assigned to groups 4 days before the start of the trials with 4 groups receiving corn (experimental) only and the fifth group hen lay pellets (control). Beginning on 20 November 1986, each bird in the corn only groups 3, 4, and 5 received approximately 0.03 g (1 snail), 0.17 g (4 snails), and 0.22 g (8 snails) of whole snails respectively, injected into the esophagus using a disposable syringe. Group 1 (hen lay pellets) and 2 (corn only) were handled in a like manner, however, only fragments of the regular diet were injected into the esophagus. The birds were treated on Monday and Thursday of each week through 29 December 1987 (12 treatments). The birds were weighed during each treatment and food consumption of each group was measured (weight). The final measurements were made on 1 January 1987. �61 RESULTS AND DISCUSSION Selection of Study Area Attachment of radios on 8 December 1985 was followed immediately by a 12+ inch snowfall that evening. The snowfall did not cause movement of birds out of the general study area as 7 of 8 birds were subsequently located in mid-December. Four of the 8 birds were at Kodak Ponds at least once before the end of December. Three of the birds used Kodak Ponds and Chestnut Slough interchangeably. With the exception of Kodak Ponds and 1 record at Wood's Lake about 4 miles northeast of Kodak Ponds, no birds were found outside of the study area. Therefore the study area boundaries remained the same except for moving the north boundary to State Highway 392. Distribution of Radio-marked Ducks Locations and movements of radio-marked ducks during December 1986 have not been plotted or analyzed but some general information can be presented. Of the 25 birds marked, 17 generally remained within the study area during December, 1 bird (AF) was located periodically within the study area, 2 birds (AM, AF) left the study area immediately after marking but returned within a few days, 3 birds (AM, 1M, IF) died during December, 1 bird (IF) was not relocated, and 1 bird (AM) was not monitored because of continuous radio interference on it's frequency. Considerable interchange occurred between Chestnut Slough and a portion of the Big Thompson River about 2.5 km north of Chestnut Slough. The birds used Chestnut Slough as a nocturnal and partial daytime roost and moved to the Big Thompson River during daylight hours. Movement to the Big Thompson was not dependent on hunting activity at Chestnut Slough. Kodak Ponds, Arrowhead Lake, and Latham Reservoir were other wetlands used by radio-marked ducks during December. Micronutrient Study Weight dynamics of birds by treatment group were minimal during the 6-week treatment period (Fig. 1). Comparing starting and ending weights within groups indicated that birds fed hen lay pellets had a positive weight change while those on a corn diet receiving 4 and 8 snails lost weight (Paired t test, .!: < 0.05, Table 3). No weight change (p > 0.05) was noted in the other 2 groups. Table 3. trials. Weight change and total food consumption of ducks on micronutrient Treatment GrouE Corn Corn + 1 snail Corn + 4 snails Corn + 8 snails Hen lay pellets Mean wt (g) Starting Ending 1075 1172 1197 1234 1157 1057 1140 1123 1193 1227 Diff. P Food consumption avg g body weight -18 -32 -74 -41 +70 0.39 0.43 0.01 0.05 0.01 2.19 2.11 2.01 1.98 3.82 �0' N 1500 _------ ""0> ~ t- J: - (!) ...._------.. .._.......••. _-~-----_.a-..•..•......•......,....,.--...".--:-~------------..------,.. " d--_--______ "----_ __ ."""""", ...-.-.-.-'-'_'_'_' _._.-._ .. ._._._ .- .-.-_.- .- - .- .-...• " '." " .•.•..•.. _. - ; __ .....••.....•.•. " '..D--:.---------. ..•.. ......... ..•.., "" " .,... •..•..•..•.. '" .. 1000 ·_·-·-CORN W ---CORN + 1 SNAIL ~ 5 00 -,'•••••••• ~i~•••• 20 NOV .,i~ T'•••••• ----·CORN+4 SNAILS ............ CORN+8 SNAILS ---------- HEN LAY PEL LET S -ri••••••••.,i~••••.,i••••••••~i~ ••••~i~ •••• ~i•••••••••• i,.•••••••• ~i~•••• ~i•• 1 DEC 11 DEC 22 DEC 1 JAN DATE Fig. 1. Mean weights (g) of mallards snails and hen lay pellets. fed diets of corn, corn plus a specific number (wt.) of �63 Because corn is deficient in nutrients needed for survival, the hypothesis was that birds on a corn diet, without snail supplements would lose weight during the test period, since they would need to catabolize protein to obtain essential amino acids. Birds receiving snail supplements mayor may not lose weight depending on the quantitative nutrient composition of snails relative to the bird's requirements. Theoretically, there was the possibility of a comparative decreasing amount of weight loss with increasing micronutrient dose. That did not occur (Table 3). However, all groups on a corn diet lost weight whereas controls gained weight (Table 3). The weight change difference was as great as 140 g (Group 3 vs. Control). Possibly, supplemental nutrients were deficient at all levels. When all corn groups were combined as one treatment, they lost an average of 41 g during the period (R<O.Ol). LITERATURE CITED Baldassare, G. A., R. J. Whyte, E. E. Quinlan, and E. G. Bolen. 1983. Dynamics and quality of waste corn available to postbreeding waterfowl in Texas. Wi1d1. Soc. Bull. 11:25-31. Buller, R. J. 1975. Redistribution of waterfowl: influence of water, protection, aridfeed. Proc. Int. Waterfowl Symp. 1:143-154. Colorado Department of Agriculture. 1980. Agricultural land conversion in Colorado. Resour. Analysis Sect., Colorado Dep., Agric., Denver. 8 pp. Dodge, W. E. 1985. A telemetry antenna mount for Cessna-type aircraft-construction details. U.S.D.I., Fish and Wi1d1. Servo Res. Info. Bull. 85-126. 4 pp. Dwyer, T. J. 282-284. 1972. An adjustable radio-package for ducks. Bird-Banding 43: Gilmer, D. S., L. M. Cowardin, R. L. Duval, L. M. Mech1in, C. W. Shaiffer, and V. B. Kuech1e. 1981. Procedures for the use of aircraft in wildlife biotelemetry studies. U.S. Dep. Inter., Fish and Wi1d1. Servo Resour. Pub1. 140. 19 pp. Gordon, D. H. 1977. Condition, feeding ecology, and behavior of mallards wintering in northcentral Oklahoma. M.S. Thesis, Oklahoma State Univ., Stillwater. 68 pp. Heitmeyer, M. E. 1985. Wintering strategies of female mallards related to dynamics of lowland hardwood wetlands in the upper Mississippi De1ta._ Ph.D. Diss., Univ. Missouri, Columbia. 376 pp. Hepp, G., R. Blohm, R. Reynolds, J. Hines, and J. Nichols. 1985. Physiological condition of autumn-banded mallards and its relationship to the probability of recovery. Proc. Waterfowl in Winter Symp., 7-10 January 1985 (in press). Jorde, D. G. 1981. Winter and spring staging ecology of mallards in southcentral Nebraska. M.S. Thesis, Univ. North Dakota, Grand Forks. 116 pp. �64 , G. L. Krapu, and R. D. Crawford. 1983. Feeding ecology of J. Wi1d1. Manage. 47:1044-1053. ---:-=-mallards wintering in Nebraska. , , and M. A. Hay. 1984. Effects of weather ----::--~ on habitat selection and behavior of mallards wintering in Nebraska. Condor 86:258-265. Kirby, R. E., J. H. Reichmann, and M. E. Shough. 1976. A preliminary report on Minnesota's innovative 1973 waterfowl season. Wi1d1. Soc. Bull. 4:55-63. Krapu, G. L. 1981. Auk 98:29-38. The role of nutrient reserves in mallard reproduction. Ringe1man, J. K., and M. R. Szymczak. 1985. A physiological condition index for wintering mallards. J. Wi1d1. Manage. 49:564-568. Szymczak, M. R., and J. R. Corey. Plains duck trap in Colorado. Weatherhead, P. J. 1983. Am. Nat. 121:237-243. 1976. Construction and use of the Salt Colorado Div. Wi1d1., Div. Rep. 6. 13 pp. Two principal strategies in avian communal roosts. White, D. C. 1982. Leaf decomposition, macroinvertebrate production, and wintering ecology of mallards in Missouri lowland hardwood wetlands. M.S. Thesis, Dniv. Missouri, Columbia. 264 pp. Prepared by 7J1,.:.L}J? ~ Michael R. Szymczak Wildlife Researcher C �65 Colorado Division of Wildlife Wildlife Research Report April 198} JOB PROGRESS REPORT .. State of Colorado ------------------------------------- Project 01-03-045 (W-88-R) Work Plan 1 Job Title: __ - 18 Winter Survival and Reproductive Success of Female Mallards Period Covered: Author: : Job Avian Research 01 July 1985 through 31 December 1986 James K. Ringelman Personnel: J. F. Corey, J. K. Ringelman, M. R. Szymczak, Colorado Division of Wildlife; D. R. Anderson, Colorado Cooperative Fish and Wildlife Research Unit; C. W. Jeske, M. W. Miller, Colorado State University ABSTRACT Mallards (Anas platyrhynchos) were captured in the San Luis Valley (SLV), Colorado during February and December 1986. A condition index (CI), which provides an estimate of relative fat reserves, was calculated using wing length and body mass measurements from 166 birds. Birds captured in February had a low CI (x = 0.092). When body mass of these birds was compared to published accounts of mallard body mass from other wintering areas, SLV mallards were about 20% lighter than their counterparts in Mississippi and Texas. However, mallards captured in the SLV during December had high fat reserves (x CI = 0.179), similar to CI values computed for other Colorado mallard populations during winter. These data suggest that SLV mallards begin winter with average reserves, but either low energy intake (food availability or digestibility) or high energy demand (thermoregulatory costs, etc.) preclude the acquisition of endogenous fat reserv,es during winter •.. The effect of radio instrumentation on the weight dynamics of mallards in winter was evaluated for transmitterss attached in 2 ways: "harness radios" with PVC tubing to attach the radio in the traditional backpack style and "suture radios" surgically attached to the back of the bird using subcutaneous sutures. A non-instrumented control group of birds was also monitored. All birds lost weight until mid-winter, but birds with harnesses lost weight at a faster rate than controls during the first 2 weeks following instrumentation. Sutures used to attach suture radios eroded through the skin, resulting in loss of transmitters in 20~60 days. Behavioral responses to harness radios varied among individuals, but most birds spent an abnormally large amount of time preening around the harness and transmitter during the 2 weeks following instrumentation. Behavior became normal after this period of acclimation. Overall, no significant difference in weight change between harness and control groups was apparent over the term of the experiment. ��67 WINTER SURVIVAL AND REPRODUCTIVE SUCCESS OF FEMALE MALLARDS James K. Ringelman Michael R. Szymczak In the 9 years since Anderson and Burnham (1976) demonstrated that hunting mortality of mallards is largely compensated for by decreased natural mortality, researchers have made little progress in identifying the period in which compensatory natural mortality occurs. Since many temperate bird populations are believed to be regulated by limitations imposed during winter (Fretwell 1972), it is appropriate to consider whether the North American mallard population may also face limitations and/or substantial mortality during winter. One study has addressed the question of natural mortality among mallards in the heart of their wintering range, the Mississippi Alluvial Valley of Arkansas and Mississippi. Little natural mortality occurred among mallards in this winter habitat (U.S. Fish and Wildlife Service 1986). However, one could argue that this region has become the heart of the winter range because mortality rates are low, and that substantial (detectable) natural winter mortality would be expected to occur only in severe winter habitats characteristic of the northern periphery of the mallard's winter distribution. These northern wintering areas were created earlier in this century as a result of irrigated agriculture, which provides waste grains and was also responsible for the creation of numerous reservoirs. Among northern wintering mallards, an increase in one component of fitness (proximity to the breeding habitat leading to "first choice" of breeding territory) may be countered by a decrease in another (over-winter survival). Mortality of ducks in response to severe winter weather is not uncommon. The first published accounts of waterfowl mortality during winter (Pearson 1934, Gromme 1936, Trautman et al. 1939) cited severe cold and/or snow cover as the principal cause, as have the most recent reports on duck mortality during winter 1976-77 and 1977-78 (Kirby and Ferrigno 1981). An appropriate location for studying winter mortality should meet 3 criteria: (1) it should maintain a relative large (>10,000) population of mallards during the entire winter, (2) winter weather should be severe, with low daily mean temperatures and at least moderate snowfall, and (3) the wintering mallard population should be isolated, thereby minimizing the likelihood of "escape" from severe weather by migration to another area. Studies of winter survival should also incorporate habitat use information, since an individual bird's habitat use in response to dynamic habitat conditions will ultimately determine its survival probability. The idea that sub-optimal winter body condition of mallards adversely affects reproductive performance has become a tacitly accepted, but as yet untest_ed, hypothesis. The physiological basis for this potential relationship was descrbied by Krapu (1981), who found that mallards arrived on the breeding grounds with most of the fat needed for reproduction. Furthermore, the magnitude of fat reserves was positively correlated with clutch size. Thus, fat reserves accumulated in migratory or wintering habitat are important for reproduction. Heitmeyer and Fredrickson (1981) reported a correlation between wetland habitat conditions in bottomland-hardwood wintering habitat and mallard recruitment rates (age ratios) the following breeding season; the correlation with winter habitat was stronger than that with either Mayor July pond numbers on the breeding grounds. More recently, Barnett and Ringelman �68 (unpubl. data) found that captive female mallards in poor late-winter condition (but provided unlimited food at the date of arrival on the breeding grounds) initiated nests later than hens in good condition. Similar pen experiments, one using game farm mallards (L. Vangilder, pers. commun.) the other using black ducks (Anas rubripes) (G. Hepp, pers. commun.) are now being replicated on a larger scale. Although informative and useful as a basis to formulate hypotheses, pen studies should not be considered definitive in resolving the question of cross-seasonal relationships between winter fat reserves and mallard reproduction. Final resolution of the question can only be attained by monitoring the winter fat reserves and subsequent reproductive performance of individual hen mallards in the wild. Biotelemetry provides a tool to accomplish such a study. However, it is not feasible to instrument mallards in the heart of the wintering range and then attempt to relocate them on breeding grounds several hundred kilometers away. The only way to effectively address this question is through an intensive study of a relatively "closed" population of mallards, one in which mallards winter and breed in the same geographic area. The objectives of this study are to: 1. Measure and compare selected reproductive parameters of adult female mallards in "good" and "poor" body condition. 2. Estimate and compare winter survival of adult female mallards in "good" and "poor" body conditions. 3. Document behaviors of male and female mallards in "good" and "poor" body condition. 4. Document flock structure at daytime roosts. 5. Document the amount of time spent foraging by birds in "good" and "poor" body condition. 6. Measure basal metabolic rates of adult male and female mallards and determine the lower critical temperature. 7. Compare proximate composition of barley from fields used foraging birds with fields not used by foraging birds. REPORTING PERIOD OBJECTIVES 1. In cooperation with personnel of the Colorado Cooperative Fish and Wildlife Research Unit and the U.S. Fish and Wildlife Service, conduct field surveys to select study sites. 2. Interview and select a graduate student. 3. Capture, measure, and weigh 40 mallards of each age/sex class during late February 1986 and mid-December 1986. 4. Compare weights and condition indices of mallards from the San Luis Valley with the same statistics from other wintering mallard populations. �69 5. Investigate new methology for attaching radio transmitters to mallards. 6. Evaluate the effect of radio instrumentation on the weight dynamics of mallards during winter. METHODS Study Area The San Luis Valley (SLV) in south-central Colorado contains a mallard population uniquely suitable for addressing the study objectives. The mid-latitude location of the SLV belies the severity of its wintering habitat. Because of its high elevation (2000 m) and encirclement with mountains, January temperatures average -4 C (average minimum = -17 C), and snows and ice fog are common. Winter mallard populations during 1982-84 averaged 14,700 (Colorado Division of Wildlife, unpubl. data). Over 5,000 mallard pairs breed on the valley floor (Colorado Div. Wildl., unpubl. data) and 1,200-2,000 pairs nest in high mountain breeding habitat directly to the west (Rutherford and Hayes 1976). Most importantly, banding data suggest that many mallards that winter in the SLV remain to breed in the valley or the surrounding mountains (Hopper et ale 1975). Thus, the SLV is ideally suited for a study designed to examine both winter mortality and the relationship between winter fat reserves and reproduction, because it (1) contains a sizeable wintering and breeding population of mallards, (2) a portion of the wintering population remains to breed in the valley, (3) winter weather conditions are severe, and (4) its geography minimizes bird movements to other wintering areas. Moreover, the SLV contains 2 federal refuges, 3 state wildlife areas, and 1 Bureau of Land Management wildlife area, all dedicated to intensive waterfowl management. Accessibility to waterfowl areas, study logistics, and inter-agency cooperation are all enhanced under these circumstances. Field Procedures Salt Plains bait traps (Szymczak and Corey 1976) were used to capture mallards on Monte Vista National Wildlife Refuge in the southwestern portion of the San Luis Valley. Wing length was measured to the nearest millimeter. Body mass was determined to within 1 gm with an electronic, digital balance. Sex-specific fat estimator equations using body mass and wing length were used to derive a condition index (Ringelman and Szymczak 1985). Effect of Radio Transmitters Two methods of radio attachment, backpack harness and sutures, were evaluated. Simulated radio transmitters measuring 2.0 x 4.0 x 1.5 cm were constructed of wood. Lead shot was used to weight the transmitters to 25 + 0.5 gm. Clear polyvinylchloride (PVC) tubing with an outside diameter of 2.6 mm was used to construct a "backpack" design harness (Dwyer 1972). Knots tied in the tubing and sealed with PVC cement were used to secure the harness. A one-eighth turn suturing needle was used to make 2 subcutaneous sutures in the back to hold the suture transmitters. Eaeh suture was about 3 em long, running longitudinally in the prodorsal region approximately 1 em on either �70 side of the dorsal apterium. Three suture materials were used: (1) Vetafil, a stranded nylon material, (2) #1 Nylon monofilament, and (3) PDS, a monofilament dissolving suture material. Ten birds (5 males, 5 females) were randomly assigned to each of 3 groups (control, backpack, and suture treatments). All birds were weighed when first instrumented and at approximately 2-week intervals thereafter. The variable of interest was change in body mass over time in response to method of instrumentation and bird sex. Long-term durability of the attachment method, feather wear and skin abrasion, and behavior of instrumented birds were also noted. All birds were fed a nutritionally balanced ration ad libitum. RESULTS AND DISCUSSION After discussions with U.S. Fish and Wildlife Service Refuge personnel, the Monte Vista National Wildlife Refuge (MVNWR) was selected as the primary study site for winter mortality studies. However, bird movements were monitored on a SLV-wide basis throughout winter and spring. Clinton W. Jeske was selected as the graduate student on this project. February 1986 Trapping Trapping was conducted on 11VNWR from 20 to 25 February. The weather during this period was clear and sunny, with overnight lows of -10 to -6 C giving way to daytime highs of 12 to 18 C. Most shallow wetlands were covered with 1-2 cm of ice at daybreak, but opened by late morning each day. Condition indices were computed for 166 birds (Table 1). Adult females were in the best condition but they, like the other age-sex classes, were still in much poorer condition than mallard populations wintering elsewhere in Colorado (J. K. Ringelman, unpubl. data). Comparisons of condition were also made among mallards from the San Luis Valley and mallards in 3 major winter areas: Bonny Reservoir (Colorado), the Texas Panhandle, and the Mississippi Alluvial Valley. Since condition indices were not computed by researchers studying the out-of-state populations, comparisons were made using published studies of body mass. Table 1. Condition indices for mallards captured in the San Luis Valley during February 1986. Age-sex class x Ad female Ad male Juv female Juv male 0.102 0.096 0.087 0.082 x 0.092 Condition index SE 0.006 0.005 0.006 0.005 N 46 40 40 40 �71 Mallards in the San Luis Valley weighed less than their counterparts captured at the same time of year at other locations (Table 2). Mean body mass of birds from the heaviest population (Texas) was nearly 28% more than SLV mallards. Since most fluctuations in mallard body mass are attributable to fat deposition or depletion (Ringelman and Szymczak 1985), these data suggest that SLV mallards either are unable to deposit high fat reserves or have such high energy demands during winter that most food consumed is used for winter maintenance and activity. In either case, low pre-breeding fat reserves could have significant consequences for breeding, since mallards begin the breeding period with most of the fat they will use for reproduction (Krapu 1981). The low mallard condition indices in the SLV were particularly unexpected in 1986, since temperatures were mild in winter 1985-86 and snowfall was less than normal (M. Nail, pers. commun.). Table 2. Mean body mass (gm) of mallards from the San Luis Valley, Bonny Reservoir, Colorado (3-year Feb mean), Texas Panhandle (Whyte et al. 1986), and Mississippi Alluvial Valley (Delnicki and Reinecke 1986). Location San Luis Valley Bonny Rservoir Mississippi Allevial Valley Texas Panhandle AF Body mass by age-sex class JF AM JM 876 956 1,095 1,114 994 1,080 1,246 1,264 829 928 1,040 1,090 929 1,042 1,181 1,160 Means 907 1,002 1,140 1,157 December 1986 Trapping Cold temperatures prevailed during this trapping period, with overnight lows near -18 C and daytime highs about 2 C. Approximately 7 cm of snow was on the ground along the west side of the SLV. Trapping was conducted from 16-19 December. A population of 8,000 mallards was present on or near the trapsites, all of which were located within MVmiR. Condition indices were high for all age-sex groups. Condition varied by age-sex class (p = 0.01), with adult females in better condition than either juvenile males or juvenile females (Table 3). The mean conditiort index for December was 95% greater than the mean index for February 1986. The December condition index was similar to indices derived for other winter mallard populations elsewhere in Colorado (Ringelman and Szymczak 1985). Thus, ~LV mallards apparently begin winter with substantial fat reserves, but quickly fall into a negative energy balance in either mid or late winter. �72 Table 3. Condition indices for mallards captured in the San Luis Valley during December 1986. Age-sex class x Ad female Ad male Juv female Juv male 0.199 0.180 0.171 0.166 x 0.179 Condition index SE 0.010 0.004 0.008 O.OOS N. 18 79 30 62 Effects of Radio-marking All simulated transmitters were attached on 18 December 1985. Unfortunately, even though birds were randomly assigned to groups, the mean mass of mallards in the control group was greater than that of either treatment group on the date of instrumentation. This necessitated expressing mass change as percentage change from starting body mass. Sutures proved to be a poor method of attachment. Therefore, those birds were not used in evaluation of mass dynamics. Body mass of all birds declined during the 60-day period following instrumentation (Fig. 1). This loss may be attributable to handling at bi-weekly intervals, but more likely represents an endogenous cycle of mass loss during winter, a phenomena common among Anatidae (Reinecke et al. 1982). Weight loss among harnessed birds during the 10 days following instrumentation was 11.S%, significantly greater than the S.S% loss experienced by control birds (p = 0.003, Wilcoxan 2-sample test). Significant differences in mass loss also occurred during the first 20 days after instrumentation (control = 8.9%, harness = l4.S%, P = 0.008). However, during the period 20-34 days following instrumentation, mean body mass of harnessed birds did not change. From 34 days until the end of the experiment 118 days after instrumentation, mass dynamics of control and harnessed mallards was nearly identical. During the experiment, control birds lost an average of 172 gm, whereas mean mass of harnessed birds declined 197 gm, an insignificant difference (p > O.OS). Moreover, variation in body mass remained nearly identical within groups from start to finish (control c.v. = 9.0% start, 8.S% finish; harness c.v. = 12.7% start, 12.3% finish). Mass loss of birds with sutured radios was similar to, but not as dramatic as, the loss experienced by harnessed birds. Unfortunately, evidence of sutures "tearing" through the skin was apparent as early as 10 days after instrumentation. The tearing process consisted of a gradual "eroding" of the suture material through the skin away from either end of the radio. Healing then occurred quickly as the suture eroded through the skin, leaving no evidence of a wound. This effectively reduced the length of the subcutaneous portion of the suture from about 3 cm at installation to less than 1 cm by 40 days after instrumentation. Continued pulling at the transmitter by the bird, along with the bounce of the radio created by slack in the suture, eventually �1600 1400 -C) lI o 1200 - w ~ >- C 0 CO 1000 800--------------------~------------------------~--10 30 50 DA YS AFTER Fig. 1. Weight dynamics of mallards instrumented control group of non-instrumented birds (top). 70 90 110 INSTALLATION with back-mounted harness radios (bottom) vs. a -...J W �74 caused the sutures to erode completely through the skin and the radios to fall off. All remaining sutured radios were removed, and the suture portion of the experiment ended, on day 47. No infection from the sutures was observed in any bird. Responses of birds to the harness radios varied by individual. No differences in weight dynamics occurred between sexes. Differences in weight change caused by tight or loose fitting harnesses, as judged retrospectively by indications of skin abrasion, were also not apparent. Behaviorally, however, birds with tight fitting radios preened more than their counterparts with loose transmitters. In general, all harnessed birds spent a great deal of time preening in the region of the harness and radio during the first 2 weeks after instrumentation. Thereafter, no qualitative differences in behavior were apparent between harnessed and control groups. An increase in preening behavior of ducks for a 1-2 week period after instrumentation has been reported by several researchers (Greenwood and Sargeant 1973, Gilmer et ale 1974, Siegfried et ale 1977, Ringelman 1980), but in each case preening behavior tended to diminish with time until behavior patterns became indistinguishable from wild birds. Siegfried et ale (1977) also concluded that the physical condition of African black ducks (Anas sparsa), as indexed by changes in body mass, was hardly if at all affected. Data presented here suggest that body mass dynamics are altered during the first 20 days after instrumentation, but are not distinguishable from unmarked birds thereafter. Concurrent with this weight loss is an increase in preening behavior. These factors may alter survival probabilities of harnessed birds during the first 2 weeks following instrumentation. However, the long-term effects of instrumentation seem to be negligible, hence it can be assumed that radio instrumentation is an unbiased technique for assessing the relationship between winter body condition and reproduction. LITERATURE CITED Anderson, D. R., and K. P. Burnham. 1976. Population ecology of the mallard VI. The effect of exploitation on survival. U.S. Dep. Inter., Fish and Wildl. Servo Resour. Publ. 128. 66 pp. Delnicki, D., and K. J. Reinecke. 1986. Mid-winter food use and body weights of mallards and wood ducks in Mississippi. J. Wildl. Manage. 50:43-51. Dwyer, T. J. 282-284. 1972. An adjustable radio-package for ducks. Bird-banding 43: Fretwell, S. D. 1972. Populations in a seasonal environment. Press, Princeton, N.J. 192 pp. Princeton Univ. Gilmer, D. S., I. J. Ball, Jr., L. M. Cowardin, and J. H. Reichmann. 1974. Effects of radio packages on wild ducks. J. Wildl. Manage. 38:243-252. Greenwood, R. J., and A. B. Sargeant. 1973. Influence of radio packs on captive mallards and blue-winged teal. J. Wildl. Manage. 37:3-9. Gromme, D. J. 1936. 53:324-325. Effect of extreme cold on ducks in Milwaukee Bay. Auk �75 Heitmeyer, M. E., and L. H. Fredrickson. 1981. Do wetland conditions in the Mississippi Delta hardwoods influence mallard recruitment? Trans. No. Am. Wildl. and Nat. Resour. Conf. 41:44-57. Hopper, R. M., and A. D. Geis, J. R. Grieb, and L. Nelson, Jr. 1975. Experimental duck hunting seasons, San Luis Valley, Colorado, 1963-70. Wildl. Monogr. 46. 68 pp. Kirby, R. E., and F. Ferrigno. 1981. Management of waterfowl during severe weather. New Jersey Outdoors 8:14-16. Krapu, G. L. 1981. Auk 98:29-38. Pearson, T. G. 1934. The role of nutrient reserves in mallard reproduction. Feeding wild ducks in a crisis. Bird-Lore 36:143. Reinecke, K. J., T. L. Stone, and R. B. Owen, Jr. 1982. Seasonal carcass composition and energy balance of female black ducks in Maine. Condor 84:420-426. Ringelman, J. K. 1980. The breeding ecology of the Black Duck in southcentral Maine. Ph.D. Diss., Univ. Maine, Orono. 89 pp. _____ , and M. R. Szymczak. 1985. A physiological condition index for wintering mallards. J. Wildl. Manage. 49:564-568. Rutherford, W. H., and C. R. Hayes. 1976. Stratification as a means for improving wildlife studies. Wildl. Soc. Bull. 4:74-78. Siegfried, W. R., P. G. H. Frost, I. J. Ball, and D. F. McKinney. 1977. Effects of radio packages on African black ducks. S. African J. Wildl. Res. 7:37-40. Szymczak, M. R., and J. F. Corey. Plains duck trap in Colorado. 1976. Construction and use of the Salt Colorado Div. Wildl. Rep. 6. 13 pp. Trautman, M. B., W. D. Bills, and E. R. Wickliff. 1939. Winter losses from starvation and exposure of waterfowl and upland game birds in Ohio and other northern states. Wilson Bull. 51:86-104. u.s. Fish and Wildlife Service. 1986. Preliminary report on the evaluation of stabilized hunting regulations. U.S. Dep. Inter., Fish and Wildl. Serv., (mimeo.). Whyte, R. J., G. A. Baldassarre, and E. G. Bolen. 1986. Winter condition of mallards on the southern high plains of Texas. J. Wildl. Manage. 50:52-57. Prepared by ~z:~ JK:Ringeln Wildlife Researcher C ��77 Colorado Division of Wildlife Wildlife Research Report Ap r LL 1987 JOB PROGRESS REPORT State of Colorado Project 01-03-045 (W-37-R) Work Plan 1 Job Title: 23 Evaluation of No-till Wheat Farming Period Covered: Author: : Job Avian Research 01 January through 31 December 1986 Warren D. Snyder Personnel: Warren D. Snyder, Colorado Division of Wildlife ABSTRACT Fields and tracts were selected within eastern Colorado and height-density (HDI) samples were obtained for wheat stubble, green wheat, alfalfa, and seeded and native grasses. Wheat stubble indices within 21 biennial no-till, ecofa11ow, and conventionally fallowed fields were highly variable (range: 0.22-1.98 dm) but averaged 0.64 dm indicating generally fair nesting quality. Most of 12 tracts of green wheat and 7 tracts of alfalfa made early, rapid growth and their HDI's surpassed that of wheat stubble in mid April. Growth patterns of wheat under conventional wheat-fa1low-wheat and annual no-till were the same within samples. Native grasses provided marginal nesting cover, below that of wheat stubble in quality into late spring. Residual switchgrass provided the highest early spring HDI; inadequate samples of cool-season grasses were obtained. Tillage of wheat stubble within conventionally fallowed fields progressed ahead of average with 60% completion by 1 May. Surveys of row crop planting into ecofallowed stubble revealed that nearly all fields were planted during. the first 2.weeks in l1ay and that subsequent stubble cover conditions were extremely poor •. ��79 EVALUATION OF NO-TILL WHEAT FARMING Warren D. Snyder This study has been transferred from the Division's South Republican Wildlife Area primarily to private lands scattered over eastern Colorado because of problems previously discussed (Snyder 1987). The transfer allowed expansion of sampling to include limited evaluation of ecofa11ow and annual no-till farming systems. P. N. OBJECTIVE To test biennial, annual, and ecofa11ow wheat and wheat-row crop no-till, and reduced tillage cropping systems for increasing the quality, quantity, and security of nesting cover for ground nesting birds within eastern Colorado. SEGMENT OBJECTIVES 1. Contacts with" USDA agency and CSU Extension personnel and others within eastern Colorado to locate farmers using no-till (annual and biennial) and ecofa11ow farming systems were made. Farmers were contacted to gain access for vegetation monitoring and to determine field locations and planned treatments. 2. Fields and other vegetation tracts were randomly selected (when possible) in spring 1986 and height-density sampling, using the Robel method, was conducted. Efforts were made to distribute sampling among several farmers and locations within northeastern Colorado and to randomize tract selection when possible to obtain a general rather than specific representation of vegetation height-density conditions. A minimum of 40 measurements per transect were obtained within all samples except when wheat growth became tall. Sampled vegetation types included: a. Wheat stubble from conventionally-fallowed, biennially-cropped no-till and ecofa11ow fields was monitored in late March or April prior to major green up of volunteer vegetation. A 2nd heightdensity sample was obtained within ecofa11ow fields within 2 weeks after they were planted to a row crop in May. Approximately 5 fields per type were to be sampled. b. Green wheat, no-till planted directly into the previous year's wheat stubble, was sampled within at least 5 fields at approximately 2-week intervals in spring. The first sample was obtained to measure wheat stubble quality remaining after fall seeding. c. Green wheat, planted on a biennial rotation into 5 conventiona11yfallowed fields, was monitored at ~2-week intervals beginning in mid-April (earlier in phenologically early years). Monitoring continued until late Mayor until a height-density index exceeding 5 dm was obtained. �80 d. Native, unmanaged, ungrazed mixed-grass pra1r1e within northeastern Colorado tableland (loam soil) was monitored within at least 4 sites beginning prior to the start of major growth and continuing at monthly intervals in April, May, and June. In addition, at least 2 tracts of switchgrass and 2-tracts of seeded cool season grasses were monitored for comparison. e. Unharvested, ungrazed alfalfa or alfalfa-dominated alfalfa-grass tracts planted for wildlife nesting cover within at least 4 Division of Wildlife properties or leases in northeastern Colorado were monitored at 2-week intervals beginning in mid-April and continuing to 1 June. 3. Progression of spring tillage of wheat stubble fields under conventional summer fallow was monitored along a route between Holyoke and Fleming. Monitoring was begun 1 April and continued at 2-week intervals until about 95% or more of the tillage had been completed. 4. Progression of slot-planting of row crops within ecofa110w stubble fields was monitored at 7-10 day intervals along the Haxtun routes established in spring 1986. Monitoring began in late April. RESULTS AND DISCUSSION Cover Quality Sampling Contacts with federal and state agency personnel, and subsequently with farmers, in early 1986 revealed that numerous fields managed under the ecofa110w system were available for sampling but almost no fields managed under the biennial no-till wheat system existed. A few farmers were trying annual no-till wheat farming in extreme northeastern Colorado. The primary concentration of ecofa110w, in wheat-corn or sorghum-fallow (3-year rotations) was in western Phillips county but a few fields were scattered elsewhere (Figs. 1, 2). Randomization throughout eastern Colorado (north of the Arkansas River, Fig. 1) was not possible because of poor distribution of locations and in some cases, because of inadequate fields. Partial randomization occurred in selecting among fields farmed by individual landowners. Likewise only partial randomization was attained in selection of sample tracts for measuring other herbaceous covers. Wheat Stubb1e.--Stubb1e quality within 3 conventionally fallowed fields, 10 ecofallow fields, and 8 biennial no-till fields was sampled prior to major vegetative growth in spring 1986. Average height-density indices (HDI's)_were 0.69 dm for the conventionally-farmed fields, 0.43 dm for ecofal10w fields, and 0.94 dm for the biennial no-till fields yielding an overall mean of 0.64 dm. The wide discrepancy among fields and types (Table 1) was primarily due to location and differing soil and weather variables. The poor readings obtained in ecofa110wed fields resulted because most were in sandy soils of western Phillips County (Figs. 1, 2) where wheat growth and stubble quality are routinely below those in nearby loam soils. The biennial no-till tracts were primarily located in sites 7 and 8 (Fig. 1) where above average precipitation in 1985 had promoted excellent wheat and subsequent above average wheat stubble. Sample means from individual fields illustrate the �81 , 'I I 'I. B E R H ASO BLO 1 -1 Fig. 1. Locations within eastern Colorado where heightdensity sampling was conducted in spring 1986. �~ ... ", 00 N .~ .• I .' ~~~~~~4d~~~~~2& ~~. I ~'! .!i![4~T:tLl1~J:129j±-:;:)N.I·2t~J~r-E' r ~1· JI~ I ~ "",,-""<..~ I'-' -+_ ..t: !'.,. ::;" i• : ".: ._ ~ ~ • I I' : "1' ! . s. • L'\..,' -'-'" I) - ., .. , . . 'd _._ 1 """... "" I. ..,.,'. • r--'-r--":"'jp~iL' -r;:~i--: . . • ~.-!JW ,,{. .I.-".. . ! L,'_ ",.I~'(:'I' .,•.••·rf~'<."--- . I i .I v.. •.•• .';::i).~ \:' )1'· - . ',c'. --....:N .J- " ' = ".", :~ .. __. _==i~.. _:_ .:_:.. : , ... ' .: ;-Y.-,.; "";'.' :1; . . ,J' __. :: ::_'( • ~. ~' . , . ~. ~ '!..;;r.. • _.:;- -II~("'_<·'''''_''''' ·c _'-'.'~" . ,..•....•.. ~ ~: •• ,_~ ~~~~. '. ·;;j:·"·.N " ~. ,.:-_.<-,,::,:. ,V J> .•._-;'.-:'~~'lo __J~::::': \ . ~~ -,~~ '1'. ~ f R-' .,:.r.. 'T .. ,' ". . t. . i;;;;,. .._ I ;J-"> "'" _. • .-r ,_~ . I '- : w.,;:~ II"':"""' • , ~ ."<; P.' ,,::, . '... :'1 '.' ~ i ••...• - ,"....' ~~."oJ: ·-H . 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I:: I ,: '.: "- " ~'- ··~L~~'·i "--i ~., ~ '\:ni'.':"b'\., :Ii L,t" .t_\. h.'-,~~'i':,~[);-,)"\·.-:'\t;;:::~/.) .~--..:~".:: ;r~.;:~if·i 1 .~ ~~"~R'~-=eo,,~· ,'-, ; .," .f:)~_..~:: . "._.... :~I.~~-'-~J., 1!"~'~ :-J.i~:t:--.l. :l>'''~~:;+~~ '. "'~ r",..-s ~:. ''". ~s,_!l--l':'1" .' L'l' .•., .~~~) ~.~ \II .' _j .: . ". 1).0' ~I. ,"_ O>¥ ~.:.,.'~ .. I )"ir~ i -...-; • ~ i~ ••;:' '.! I ~"" :/ _;...,~~. .' ~ . I_.~·"I..~~=--._· " ~., ,,'-:-":~ ! .,...,. '~'.'" ::"5~ I''t'>', n~t'. -,_-; ."--' t-" t .".~ ..:"-'~;;J -»!~f-:~ -"!~ .... /'\;, . I :~I-"-,r'Z:;~'v'J .~_~.~I '.. _:'!' c-; ~_ J. • : .•.. .., . .1. ';' . . . - ~ _, c',):a.Li,~c;E 1 ~o h'iIJ~I-:,.;< .-·I_~t.::- "'~ .·T· ~p.(i, "_ ''_ "li..:tIO> .:~ -~ -t,c- 'I,J " •• : i' L-~~~~."",. ~: .....• : .-,,,,,.,, .' J ... :...• ;:::~I::'-"""l~~ ... i-..__ ';;Y'" ....,..... .--0. .-' r~~~-J ." . ,'. ""."-I _~ .:....: rt-J-""'. . ~ I ~'Wi" ~_, .' .:, ._+_. ".I' ',,'_ ~_, .'. .' - '. ~'~l+.-!·.,_r1· ·Lf~GP.:~··:·~ c{~rv;;k_~~ I 'j JI '. L . ~L~' II.'·"B~>' -;.~~=::f" ~""'-7.' -• .~C: t .,'\.1::" I;'\.. .•.•• -~"" ~ 1",l~ •.•.•.. ~, Fig. 2. Location of the Holyoke-Fleming stubble tillage route and the Haxtun ecofa11ow row-crop planting route, Phillips-Logan counties, Colorado, Spring 1986. �83 Table 1. Height-density indices (dm) of wheat stubble within fields farmed by biennial no-till, ecofallow, and conventional systems in eastern Colorado, Spring 1986. Site County a Legal description Sample size x HDI Biennial No-till Kiowa Kiowa Kiowa Kiowa Washington Washington Washington Washington Larson N. Larson SW Jacobs IFl Jacobs IF2 Wagers III Wagers IF2 Mickleson IFl Mickleson IF2 31 1 9 5 22 15 13 31 - l8S l8S 17S l7S lS IS 2S IS - 48W 49W 50W 50W 56W 56W 56W 55W 48 48 48 48 44 44 48 40 0.71 0.34 0.52 0.43 1.49 1.98 1.08 0.89 - 7N - 7N - 9N - 8N - 8N - 5N - 6N - 7N - 17S - l7S - 46W 46W 47W 47W 48W 49W 49W 45W 45W 45W 64 60 60 64 60 44 40 40 48 48 0.43 0.82 0.25 0.32 0.28 0.64 0.43 0.63 0.29 0.22 19 - ION - 43W 19 - 10N - 43W 19 - 10N - 43W 60 40 40 0.78 0.92 0.31 Ecofallow Ham IFl Ham IF2 Livingston Hofmeister III Hofmeister 112 Whittington III Whittington 112 Anderson Scherler III Scherler 112 Phillips Phillips Phillips Phillips Logan Washington Logan Phillips Kiowa Kiowa 30 30 28 24 12 11 36 31 22 15 Conventional Sand Draw III Sand Draw 112 Collins Overall Sedgwick Sedgwick Sedgwick x aSection, Township, Range. 0.64 �84 wide variability of stubble quality available to nesting pheasant hens in spring 1986. Early spring nests of radio-marked pheasant hens were not found in wheat stubble in 1980 when its HDI was exceptionally poor (0.34 dm) whereas 10 of 21 early nests were in stubble in 1979 when an excellent HDI (1.34 dm) existed (Snyder 1984). In 1981, 8 of 34 early nests were in stubble when the average HDI was 0.83 dm. Using these data for evaluating 1986 stubble quality implies that at least one-half of the wheat stubble fields with HDI's <0.5 dm probably provided poor nesting cover for hen pheasants and a few fields offered good nesting sites. The fact that stubble fields within biennial no-till fields possessed higher HDI's than those under ecofallow should not be misinterpreted as a product of treatments. Most no-till fields were first-time treatments and the stubble there had come from conventionally-fallowed fields. Several years of farming under the biennial no-till system would be needed to compare stubble quality with that in other farming systems. At present ecofallow is dominant among the new farming approaches and is the only one gaining increased acceptance as a standard practice in northeastern Colorado. This system yields 2 crops in 3 years as opposed to a crop every other year, allows crop diversity, and uses inexpensive and less frequent herbicide applications. Green ~fueat Under Conventional and Annual No-till Farming.--The warm spring in 1986 stimulated early wheat growth in most sampled fields. Poor growth, a product of poor farming practices by lessees, lack of fertilizer, and deficient soil moisture occurred in 3 sampled fields within Division of Wildlife Properties. Wheat HDI's there on 1 May average only 0.45 dm compared to 1.87 dm within 4 sampled fields on private lands (Table 2). The average HDI for green wheat surpassed that for wheat stubble in mid to late April just ahead of the primary pheasant nesting season (Fig. 3). Five fields under annual no-till winter wheat management were sampled during spring 1986. Under this farming approach herbicides with short residual effectiveness are applied after July wheat harvest and wheat is seeded directly back into the unworked stubble in September using special no-till drills. Fertilizer is applied prior to or during the seeding operation. The no-till drill knocks down most of the residual stubble leaving limited cover over winter to stop most drifting snow but that is marginal for wintering pheasants. The height-density quality of remalnlng wheat stubble and green wheat were listed separately and then combined during 1 April sampling of no-till wheat fields. Tallies where stubble was the dominant obstruction averaged 0.27 dm and those for wheat were 0.12 dm yielding a combined average of 0.18 dm. This indicates that planted no-till fields provide marginal cover in early spring. Green wheat rapidly became the dominant vegetation in subsequent sample intervals and followed the same growth curve obtained for wheat planted in conventionally-fallowed fields (Fig. 3). The residual straw, both standing and flattened, may make annual no-till fields more attractive for pheasant nesting than conventionally-fallowed fields where the soil surface is bare. Evaluations would be needed to test this hypothesis. �85 5 I I I I 4 ,~ I It' 1$ -.a 'g I!!, 10 3 I ....., I ~ ~ _- a' /' til Z til 0 I 2 I ~ ~ I .-.,.' ).0£ a "1.0£•••••• a ••••• I:' .." •••• . •a· .•. ' .-w- •• I ~ •• /- ...•.".. /I ./ 1 . ... ~.. Cool Sea!~~r~~e~_~7 .,,".,..,. - .. -- , .p.r.".~ •.... • .". ---.---~ .. .. _...-- ,.- - -------_.. :..:.- •...'L ."._.__. _. _. .- .. ,0 _ Wheat Stubble .•.- 28 MAR 3 --'"•••• •• " '" 9 15 APR Native Grasses 21 27 3 __ / •. " • _.- 9 15 MAY -> .. •.. .- 21 27 2 8 JUN Fig. 3. Comparison of height-density indices among sampled vegetative covers in relation to wheat stubble cover quality, eastern Colorado, spring 1986. .• �86 Table 2. Height-density indices (dm) of green wheat within conventional and annual no-till fields, northeastern Colorado, spring 1986. Location Apr 15 1 May 29 13714 22729 4.22 1.99 2.56 3.19 6.22 4.81 4.77 5.35 4.11 3.40 4.53 Conventional tillage Hodges Sand Draw 111 Sand Draw 112 Collins NW Bamford Duck Creek Holyoke-Heimer 0.07 0 0 0 0.31 0 0.63 0.02 0.16 0.19 0.89 0 0.37 1.84 0.45 0.77 1.53 2.42 0.10 1.75 1.38 2.96 Annual No-till Kinnie 6-1 Kinnie 30-2 Kinnie 20-3 Livingston 9 Livingston 3 0.30 0.20 0.12 0.13 0.14 0.49 0.53 0.38 0.38 1.58 3.20 3.38 1.81 1.26 3.00 2.10 0.76 field destroyed 4.73 5.68 4.93 4.33 Alfalfa.--Seven transects containing a dominance of unharvested alfalfa (some cool season grasses were present) within Division of Wildlife Properties or leases were sampled through the spring growing season (Table 3). Most stands had been planted >5 years. Inadequate precipitation and soil moisture were factors at the Duck Creek and Holyoke sites. Excellent growth and nesting cover conditions were present on the other sites and on the average, alfalfa surpassed the HDI of wheat stubble at about the same time (mid-April) as green wheat (Fig. 3). A primary weakness with alfalfa is that it does not stand up over winter to provide winter protection during snow cover or to provide early spring residual cover. Table 3. Height-density transect means (dm) of alfalfa of alfalfa-dominated mixtures within 7 northeastern Colorado sites, spring 1986. Location 15 Apr 1 May 15 May 30 May 11 Jun Sand Draw Duck Creek Holyoke property Whittington E. lVhittington W. Wisdom S. Wisdom N. 0.33 0.53 0.59 0.52 1.10 1.02 1.39 1.47 1.43 1.28 1.68 1.46 1.13 2.56 2.30 2.97 2.36 3.07 1.52 2.19 3.97 3.72 3.86 3.20 4.50 1.61 2.02 4.53 4.18 4.80 3.70 �87 Seeded and Native Grasses.--Samp1ing of seeded cool-season grasses was not sufficient to provide meaningful comparative data. A stand of smooth brome within a waterway on the Duck Creek Property provided fair to good nesting cover (Table 4). A dry1and slope on the same site, seeded to crested wheatgrass was of poor quality and an old stand (15+ years) of intermediate wheatgrass also was of little value for pheasant nesting through the spring (Table 4). Table 4. Height-density index transect means (dm) of cool season grasses, switchgrass, and native grass tracts within several ungrazed northeastern Colorado sites, spring 1986. Apr Location Vegetation Mid 1 1 May Mid Late Mid Jun 2.11 0.37 0.25 2.17 0.65 0.27 2.45 0.62 0.89 Cool-Season Grasses Duck Creek Ham Smooth brome Crested wheat Inter. wheat 0.64 1.61 0.14 0.15 Switchgrass 3.23a Sand Draw Holyoke Property Messex site S. Republican 4.85 3.10 1.81 1.31a 1.21a 1.24a (Hailed) Native Grasses Sand Draw Holyoke Property Duck Creek Ham Mixed Mixed Mixed Mixed Pr Pro Pro Pro 0.20 0.38 0.24 0.61 0.34 0.39 0.26 0.53 0.41 0.71 0.28 0.50 1.11 0.93 0.54 0.71 3.18 1.45 1.09 0.89 aA11 first measurements of switchgrass sampled only residual vegetation. Seeded stands of switchgrass provided good to excellent residual nesting ~over through spring within 4 widely distributed locations (Table 4, Fig. 3). New growth did not begin to impact height-density readings until early June, consequently only two samples were obtained. A severe hailstorm on the South Republican flattened all residual cover preventing a comparable late spring measurement. These data indicate that this species, if properly managed on a periodic basis, can provide by far the best early spring residual nesting cover where it is adapted for planting in extreme eastern Colorado. During an interval of deep (25 cm) snow in December 1985, switchgrass provided the only herbaceous night-roosting cover available to wintering pheasants attesting to its overwinter values to wildlife. �88 Height-density indices of native grasses were all rather uniformly low among 4 sampled locations in northeastern Colorado through mid-May (Table 4). May precipitation stimulated improved HDI's in late May and mid-June with cheatgrass providing the dominant visual obstructibn on most sites. None of the tracts was grazed. Native grass stands did not compare favorably with wheat stubble as nesting cover during early to mid spring intervals (Fig. 3). Time of Conventional Stubble Tillage The timing and impact of initial stubble tillage on nesting pheasants and other ground nesters at the onset of summer fallow operations was discussed by Snyder (1984). During 1979 and 1981 only 1 of 18 nests hatched successfully from wheat stubble in spring with most of the losses being attributed to tillage operations. Nest predation was a factor in several losses but if it had not occurred, all would have been lost to tillage operations anyway. A route from Holyoke to Fleming was established in spring 1986 and over 100 stubble fields were censused routinely to assess progress of stubble tillage (Fig. 2). Results revealed tillage progression was early in 1986 compared to that of the 1979-81 interval and 60% had been completed by 1 May (Fig. 4). Thus, the majority was completed prior to the primary May-June pheasant incubation interval. Time of Row Crop Planting Within Ecofallow A 2nd survey route was established within western Phillips County to identify timing and progression of row crop planting in ecofallow (herbicide treated) wheat stubble fields (Fig. 2). Most of the land was sandy and strip farmed to reduce erosion prevails. Therefore, individual strips within fields or sections were listed as not planted, planted, or in progress. The 29 April survey revealed that planting had not been initiated on any of the 145 strips or tracts within 34 larger field units along the route. By 7 May 34% of the strips were planted compared to 79% completion on 15 May. A 30 May survey revealed nearly all had been completed and most of the 5 remaining strips would probably be fallowed. Most planting occurred during the peak pheasant nesting interval in 1986. However, since most fields contained low HDI's prior to planting, nest disruption was probably not a major factor. Post-planting HDI's were obtained on 8 ecofallow fields prior to significant corn or milo emergence. Most HDI's were about zero and averaged only 0.19 dm indicating little attractive nesting cover remained after planting. SUMMARY The preceding data show that among the 4 farming systems, biennial no-till wheat farming, by leaving stubble standing undisturbed through the spring and summer has the greatest value for ground nesting wildlife. Its primary weakness lies in the height-density quality of.the stubble which in most sampled fields, was only poor to fair in 1986. Annual no-till is not expected to be adopted in the near future because of severe acreage restrictions required under federal farm programs. Economics dictate that most farmers comply with these restrictions. Wheat planted directly into stubble may �89 100 / -----",. ",.--- _" / 80 / / / I / 60 / 0 / tal tl / ..l ~ 0 u 40 fi tal hl tal III 20 / / / / / / / / .;.;/ 0 4 8 12 16 20 24 28 APRIL 2 6 10 14 18 22 26 30 MAY Fig. 4. Progression of spring tillage of wheat stubble fields along the Holyoke-Fleming survey route, spring 1986. �90 provide a better nesting environment including early spring use by mourning doves, but has the limitation of marginal over-winter value for wildlife. This approach will probably be restricted to extreme northeastern Colorado, at least in the near future, where favorable annual precipitation amounts permit its use. Gradual expansion of ecofallow into loam soils may improve stubble quality. However, slot planting of row crops during the peak of spring nesting, especially milo planting in late May will undoubtedly cause considerable nesting disruption. Most fields offer only marginal cover for nesting subsequent to planting. Although stubble tillage under conventional fallow operations was earlier than usual in 1986, it probably still impacted many early spring nesting efforts and forced renesting elsewhere. LITERATURE CITED Snyder, W. D. 1984. Ring-necked pheasant nesting ecology and wheat farming on the High Plains. J. Wildl. Manage. 48:878-888. 1987. Evaluation of no-till wheat farming. Colorado Div. Wildl. Wildl. Res. Rep. Fed. Aid Proj. 01-03-045. Apr. (In press). Prepared by ¥ 1UMMtJ £) War~D. Snyder Wildlife Researcher C �91 Colorado Division of Wildlife Wildlife Research Report April 19_8] JOB PROGRESS REPORT State of Colorado ----------------------------------- Project 01-03-045 3 Work Plan Job Title: 10 ------ Some Population Characteristics of Adult Gadwall Molting in North Park Period Covered: Author: Job Avian Research 01 April 1985 through 31 December 1986 Michael R. Szymczak Personnel: J. Corey, J. Ringelman, S. Steinert, and M. Szymczak, Colorado Division of Wildlife ABSTRACT Trapping of gadwall (Anas strepera) in North Park resulted in 648 normal wild-trapped banded birds released into the population. ��93 SOME POPULATION CHARACTERISTICS OF ADULT GADWALL MOLTING IN NORTH PARK Michael R. Szymczak P. N. OBJECTIVES 1. Document the distribution of harvest of gadwall banded as flightless young or molting adults in North Park, Colorado. 2. Estimate recovery and survival rates of adult gadwall molting in North Park, Colorado. SEGMENT OBJECTIVES 1. Trap and band 400 adult males, 350 adult females and, if available, up to 100 local gadwall on molting and brood rearing areas in North Park (1985). 2. Submit banding schedules and recapture reports to the U.S. Fish and Wildlife Service's Bird Banding Laboratory. File return information at the Colorado Division of Wildlife Research Center (1985). 3. Obtain computer tapes containing: (1) all records of gadwall banded in North Park through September 1985 and (2) all records of recovery, recapture, or other types of encounters through April 1986 of gadwall banded in North Park from the U.S. Fish and Wildlife Service's Bird Banding Laboratory. 4. Compile a complete listing of North Park gadwall returns (banded birds recaptured within the same la-minute latitude-longitude grid of banding) from CDOW banding files. 5. Analyze recovery, recapture, and return data to document: (1) the geographic and temporal distribution of recovery, (2) homing rates to molting areas, and (3) recovery and survival rates. 6. Prepare final report as a technical publication. METHODS AND MATERIALS Birds were captured from an air-thrust boat at night using a hand-held 12-volt landing light and a long-handled net. Flightless adults captured were weighed and measured prior to being banded and released in conjunction with a study of the flightless period in ducks (Szymczak and Ringelman, 1984). The band numbers of birds captured that had been previously banded were recorded. Banding schedules and recapture reports were prepared and submitted to the U.S. Fish and Wildlife Service's Bird Banding Laboratory. Information on returning birds recaptured in the same la-minute grid of banding were filed at the Colorado Division of Wildlife Research Center. Following the 1985-86 (31 Aug - 1 Sep) recovery year, a banding tape listing all gadwall banded in North Park from 1971 through 1985 and a companion �94 recovery tape listing all encounters (recoveries, recaptures) of these birds through the 1985-86 recovery year were requested from the u.s. Fish and Wildlife Service's Bird Banding Laboratory. Returns, birds recaptured in the same lO-minute latitude-longitude banding, were compiled through the 1984 summer recovery year. grid of RESULTS Banding Trapping resulted in 329 adult males, 285 adult females, 16 local males, and 18 local females being released as newly banded birds useable in future analyses (Table 1). In addition, bands were replaced on 5 adult males and 8 adult females. Radio transmitters were attached to 12 additional females to study short-term survival of flightless adults as part of another study. The total number of adults released was greater than banded in 1984, but was short of established sex quotas. Analysis The analysis was not initiated during this reporting period. LITERATURE CITED Szymczak, M., and J. Ringelman. 1984. Ecological studies of the flightless period of ducks in Colorado. Colorado Div. Wildl., Wildl. Res. Rep. Oct. Pp. 13-31. Prepared by ~1;;J_J??.4 010~ MiC ael R. Szymczak Wildlife Researcher C �Table 1. Gadwa11s banded in North Park by location, August through September 1985. Location Adult males Walden Reservoir MacFarlane Reservoir Pole Mountain Reservoir Lake John Annex 215 (4)a 116 (l)a 3 Totals 334 (5)a ° Age and sex Adult females Local males 231 52 19 3 (4,8)b (l,l)b (3,2)b (O,l)b 305 (8,12)b Local females 4 3 8 1 6 1 8 3 16 18 Totals 456 172 38 7 (8,8)b (2,1)b (3,2)b (O,l)b 673 (13,12)b aNumber of birds included in the total whose bands were replaced. bNumber of birds included in the total whose bands were replaced (1st # in parentheses) to which radio transmitters were attached (2nd # in parentheses). ~ L11 �� https://cpw.cvlcollections.org/files/original/76ced8ce22a8f59100a7019976b28cb7.pdf 7e7d9786708ee52d200119f3c4ae19c3 PDF Text Text 97 Colorado Division of Wildlife Wildlife Research Report April 198] JOB PROGRESS REPORT State of Colorado ----~~--------------------------01-03-045 (W-37-R) Project Work Plan 3 Job Title: l3(b) ----~~- Responses of Sage Grouse to Vegetation Ferilization Period Covered: Author: : Job Avian Research 01 January through 31 December 1986 Orrin Myers Personnel: C. Braun, L. Carpenter, J. Corey, Colorado Division of Wildlife; W. Brown, O. Myers, G. White, Colorado State University; C. Cesar, L. Upham, U.s. Bureau of Land Management ABSTRACT The response by sage grouse (Centrocercus urophasianus) to application of nitrogen fertilizer to sagebrush-dominated rangeland was studied in North Park, Jackson County, Colorado. Ammonium nitrate fertilizer was applied to 330 ha of sagebrush in Fall 1985 at a rate of 112 kg-N/ha. Seasonal collections of foliage from Artemisia tridentata wyomingensis (ATW) and A. t. vaseyana (ATV) were made to document chemical responses to fertilization~ Information on vegetation growth response also was collected. Diameters of sagebrush plants were not affected by fertilization but plant heights increased 18% overall. The average leader length of fertilized sagebrush plants was 2.1 times the length of leaders from plants on control plots. ��99 ..' RESPONSES OF?SAGE GROUSE TO VEGETATION FERTILIZATION Orrin Myers PROGRAM NARRATIVE OBJECTIVE The project is part of a 2-phased study to (1) collect baseline information on sage grouse winter distribution and feeding ecology in areas to be impacted by coal mining in North Park, Colorado, and (2) evaluate whether nitrogen fertilizer, when applied to sage grouse winter habitat, can be used as a tool to mitigate reduction in habitat available to sage grouse. The focus of this portion of the project is on phase 2. SEGMENT OBJECTIVES 1. Document the chemical and growth response of sagebrush to nitrogen fertilizer, 2. Evaluate feeding preferences of sage grouse for fertilized and unfertilized sagebrush subspecies, 3. Estimate the digestibility of fertilized and unfertilized sagebrush, and, 4. Monitor the reproductive parameters of radio-marked sage grouse to learn if sage grouse reproductive success is affected by fertilizer treatment. DESCRIPTION OF STUDY AREA The study area is North Park, Jackson County, Colorado. The Park is an intermountain basin at an elevation of about 2500 m. It is drained to the northwest by the North Platte River, which is fed by many smaller streams. The topography is flat to rolling with numerous ridges and benches. The climate is cold and dry with an average annual frost-free period of 46 days. Sagebrush-dominated grasslands cover upland sites in the Park; grasses and sedges occur in native and irrigated meadows that border drainages. Artemisia tridentata is the dominant sagebrush species and includes 2 subspecies: A. t. wyomingensis and A. t. vaseyana. Other species of sagebrush occurring with limited distribution-in North Park are A. longiloba, ~. cana, and A. argillosa. METHODS In October-November 1985 ammonium nitrate fertilizer (33.5% nitrogen) was applied at a rate of 112 kg-N/ha to a total of 330 ha of sagebrush rangeland. Fertilizer was applied with mechanical spreaders at 3 areas in North Park representing occasional (Area C), regular (Area A), and intensive (Area B) sage grouse use (Schoenberg 1982). Within each Area, 11 20-ha experimental blocks were selected in random locations. Each block was divided in half with the northern or southern 10 ha randomly assigned as fertilized or control and treated accordingly. �100 Sagebrush Foliage Collec~ons Sagebrush foliage samples were collected from study blocks in May, August, and November-December. Samples of ATW and ATV were clipped from the control and fertilized halves of randomly selected blocks. During the May and August collections 10 blocks were sampled with 10 plants of each subspecies being clipped from the control and fertilized halves of each block. Individual plants clipped were those closest to random points along pace transects parallel to the long axis of each block. All samples were placed in zip lock bags, labeled, sealed, and placed in frozen storage at the CDOW Research Center, Fort Collins. The November-December foliage collections differed in that fewer plants were clipped from each block, however more study blocks were sampled so the total number of plants clipped remained about the same. Four plants per treatment of each subspecies were clipped from a total of 24 study blocks evenly distributed across the 3 main study areas. Measurement of Sagebrush Growth Response A total of 12 randomly-selected blocks was sampled in October. Blocks were selected to stratify samples evenly across all 3 Areas (A, B, and C). At 15 points along pace transects, distance to nearest sagebrush plant, 2 perpendicular diameters of this plant, its height, and the distance to its nearest neighbor were measured. Each half of the experimental block (fertilized and control) was measured. Information on the current year's growth increment was gathered by measuring 3 plant leaders from 15 plants on each half of 1 experimental block in each Area. RESULTS Diameters of sagebrush plants measured on the control and fertilized halves of each block did not differ between treatments (Table 1). Height of fertilized plants was greater than controls within Area A and Area B but not in C. In each of the areas the average leader length was greater for fertilized plants than for control plants (Table 2). The ratio between leader length and plant height (LHRATIO) was examined as a means of standardizing plant response to plant size. On a gross scale this may minimize effects due to subspecies status. There were no differences between areas (p =-0.35 in LHRATIO), but LHRATIO for plants from fertilized plots was greater than for control plants (P = 0.0001). Due to varying subspecies composition of blocks, soils, and topography, block effects and area-treatment interaction effects were sometimes significant. Within block 38 in Area B for example, plant height was greater among control plants (56.3 cm) than among those that were fertilized (35.0 cm). This finding was a result of a preponderance of ATV plants being measured in the control plot, which was on a north-facing slope, and a preponderance of ATW plants being measured on the fertilized portion of the block, which was mainly on the hilltop. �101 Table 1. Selected measurements of Artemisia tridentata plants from 4 experimental blocks in each of 3 study areas in North Park, Colorado, October 1986. Parameter Treatment A ~N - 60) Diameter 1 (cm) Control K SD Fertilized X SD Diameter 2 (cm) Control x SD Fertilized x SD Height (cm) Control X SD Fertilized x SD ap < B ~N - 60~ Area C ~N - 60~ All (N - 180) 34.4 21.8 45.9 24.6 31.2 21.6 37.2 23.4 36.9 18.7 43.6 27.2 39.2 22.7 39.9 23.2 25.8 17.6 32.5 18.0 23.0 16.8 27.2 17 .8 28.3 16.5 33.0 21.8 27.9 16.8 29.7 18.6 14.6 9.6 32.1 19.4 21.6 11.9 22.8 15.9 20.4a 8.8 32.3 16.7 28.0a 12.4 26.9 13.9 0.01 that fertilized equal to control. Table 2. Average leader length (mm) from Artemisia tridentata plants on fertilized and control plots in North Park, Colorado, October 1986. Area Treatment N x A Control Fertilized 15 15 20 79 10.3 41.0 0.0001 B Control Fertilized 15 15 33 56 21.3 25.4 0.002 C Control Fertilized 15 15 23 62 8.3 25.0 0.01 All areas Control Fertilized 45 45 25 66 15.2 32.2 N.T.a aN.T. = not tested. SD P > (t) �102 LITERATURE CITED Schoenberg, T. J. 1982. Sage grouse movements and habitat selection in North Park, Colorado. M.S. Thesis, Colorado State Univ., Fort Collins. 86 pp. Prepared by ~ __~ __ =-_~ __~_.+~~-~ Orrin B. MY;?;; Graduate Research Assistant Approved by i!tJ L. ~ _ ~C~l-a~i~t~E~.~B~r~a-u~n~~~~---------Wildlife Research Leader �Colorado Division of Wildlife Wildlife Research Report April 1987 -~ JOB PROGRESS REPORT State of Colorado --~~~--------------------------- Project 01-03-045(W-37-R) Job 3 Work Plan Job Title: 15 ------ Sage Grouse Distribution and Habitat Use in the Gunnison Basin Period Covered: Author: Avian Reseqrch 01 July 1985 through 31 December 1986 Jerry Hupp Personnel: C. Braun, C. Coghill, B. Groshek, J. Houston, T. Henry, P. Mason, J. Olterman, T. Sherrill, Colorado Division of Wildlife; J. Hupp, R. Ryder, Colorado State University; M. Blymeyer, J. Capodice, U.S. Bureau of Land Management; C. Molitoris, W. Shuster, B. Wallis, U.S. Forest Service ABSTRACT Sage grouse (Centrocercus urophasianus) winter distribution, foraging ecology, and habitat use were studied during January-March 1986. widely distributed across the Gunnison Basin. Flocks were In 3 winters of study, no evidence of a single or small number of concentrated winter areas was obtained. Sage grouse use sites were located on 52 of 120 l-km diameter randomly-located survey plots in 1986. Most (61%) of the 83 winter feeding sites located were in sagebrush (Artemisia spp.) stands in drainages and on 0 6-15 slopes with southwest aspects. Use of 0-50 topographically low sites in 1986 (11% of feeding sites) was less than in 1985 (37% of feeding sites). aspects. Only 4% of 1986 feeding sites occurred on slopes with northeast Sagebrush height in relation to snow depth likely affects sage grouse distribution among terrain categories. Shrub structure differences �104 were most apparent between sage grouse feeding and random sites on xeric terrains (>5 0 southwest, 0-50 high) and least apparent in drainage sites. These were no differences in crude protein or monoterpene content among randomly-located sites in different terrains. Crude protein and monoterpene content was similar between browsed and unbrowsed mountain.big sagebrush (Artemisia tridentata vaseyana) plants at sage grouse feeding sites. Early spring lipid reserves of adult male sage grouse in Gunnison County were similar betwen 1985 (4.1%) and 1986 (4.2%). Lipid reserves of adult males in Jackson County were lower in 1986 (3.6%) than during 1985 (5.5%) due to severe early winter conditions. Differences in male strutting display rates were observed between Jackson and Gunnison County males but could not be attributed to unequal lipid reserves. �105 SAGE GROUSE DISTRIBUTION AND HABITAT USE IN THE GUNNISON BASIN Jerry Hupp P. N. OBJECTIVES The primary objectives of this study are to: (1) evaluate winter and spring distribution of sage grouse in the Gunnison Basin, Colorado; (2) describe topographic, vegetational, and shrub structural characteristics at sage grouse feeding sites; (3) compare spring lipid reserves of male sage grouse in Jackson and Gunnison counties; and (4) develop a sage grouse management plan for the Gunnison Basin. SEGMENT OBJECTIVES 1. Compare snow depths and winter sagebrush availability among years and terrain categories. 2. Evaluate sage grouse winter distribution in the Gunnison Basin. 3. Describe topographical characteristics of winter use sites. 4. Describe shrub species composition and structure at winter and spring use sites. Evaluate chemical composition of sagebrush plants at random locations and sage grouse winter feeding sites. 5. Locate previously undiscovered leks in the Gunnison Basin. Survey sage grouse lek attendance. 6. Evaluate spring lipid reserves of adult male sage grouse collected in Gunnison and Jackson counties. Test whether observed differences in courtship behavior between populations can be attributed to differences in energetic reserves. 7. Evaluate and analyze data. Prepare progress reports. Present findings at professional meetings. �106 DESCRIPTION OF STUDY AREA The study area in Gunnison and Saguache counties has previously been described by Hunter and Spears (197S) and Hupp (1984) and will be described in the final report. METHODS Snow Depth and Sagebrush Availability I compared snow depth among years and terrain categories (Table 1) in the Gunnison Basin. In 1985, snow depth was measured along a series of randomly located transects in Chance and Graflin gulches. Snow depth as evaluated in these areas because conditions appeared representative of snow'cover in the Gunnison Basin, and because access to a variety of topographical types and elevations was feasible. In 1985 and 1986, measures of snow depth were obtained at S points spaced at SO-m intervals along transects in each terrain category present in randomly located l-km diameter survey plots. In each year of the study, I estimated mid-winter availability of exposed sagebrush along aerial transects between 24 January and 24 February. Approximately 416 km of transects were flown each year via fixed-wing aircraft at approximately lSO-200 m elevation. I observed sagebrush through a 2S-cm 2 Air speed was approximately 130 km/hr. viewing square marked at shoulder level on the passenger window of the cabin. At IS-second intervals, I recorded the presence or absence of exposed sagebrush in the viewing square. Exposure was "excellent" if sagebrush comprised >SO% of the viewing square, "moderate" if sagebrush was exposed but comprised <SO% of the viewing square, and "unavailable" if no sagebrush crowns were observed in the square. A second observer was responsible for keeping the pilot informed of the aircraft's position relative to transect lines. Transects were marked on 7.S minute topographic maps to facilitate correct orientation of the aircraft. �107 Table 1. Categories or:~·slopeand aspect used to classify terrain in randomly selected sage grouse survey plots, Gunnison Basin, Colorado. Terrain category Description 0-50 slope, high Areas of <50 slope; broad (>100 m) mesa or ridge topography tops topographically higher than surrounding terrain. 0-5 o slope, low topography Areas of <50 slope; broad (>100 m) flood plains and stream drainages adjacent to areas of higher topography; terraces that occur on slopes. Drainages Narrow «100 m) flood plains of permanent or intermittent streams; shallow, eroded gulches on slopes. 6-15 o slope, north- east aspect 6-15 o slope, south- west aspect > 15 o slope, north or east aspect >15 o slope, south or west aspect 0 Slopes between 6 and 15 with primarily north or east aspects.a Slopes between 6 and 150 with primarily south or west aspects. b Slopes of >150 with primarily north or east aspects. a South of >150 with primarily south or west aspects. b aAspects with bearings of 316 to 1350• bAspects with bearings of 136 to 3150• �108 Winter Distribution I evaluated sage grouse distribution among 8 regions of the Gunnison Basin (Fig. 1). Ground searches for use sites were conducted in each region between 7 January and 31 March 1984, 3 January to 15 March 1985, and 7 January to 11 March 1986. The slightly longer winter survey period in 1984 was due to severe winter conditions that existed during that year. Use sites were located from flock sightings as well as tracks in snow. Track observations included snow roosts and feeding sites. The latter were distinguished by the meandering path of tracks and the presence of clipped leaves on sagebrush plants. Track observations indicated that sage grouse usually landed, foraged in a localized area, then flushed. Therefore feeding sites were discrete and it was usually possible to identify the extent of most use sites. Track observation was a valid criterion of use because snow cover was present throughout each winter period. Two different approaches were used to distribute search effort in each region of the Gunnison Basin. In 1984, severe winter conditions restricted access to many areas of the Basin. Accessible areas within each region were selected and systematically searched for sage grouse use sites via snowmobile, snowshoes, or skis. topographic maps. Approximate boundaries of use search areas were marked on Improved access in 1985 and 1986, allowed use of a different approach that insured unbiased sampling of sage grouse winter distribution. Random points were chosen in each region, and 1-km diameter circular plots were established around each point. drawn on 7.5 minute topographic maps. Boundaries of plots were Circular plots were visited once each winter and searched on foot for use sites. Plots were located in the field by comparing features illustrated on topographic maps with terrain characteristics in the area to be surveyed. Because of the rugged topography �~ ":"> A\ MciNTOSH 012345 KILOMETERS STEUBEN Fig. 1. Regions !"I of the Gunnison Basin, Colorado. f-' o '" �110 of the Gunnison Basin, boundaries of survey sites were easily discerned. sage grouse use sites observed within a survey plot were recorded. effort was approximately equal in each plot. All Search The intent of this approach was not to locate all sage grouse use sites within a survey area, but rather to equally distribute sampling effort among all regions of the Gunnison Basin. A plot was considered "occupied" by sage grouse if at least 1 use site was observed within its boundaries. I used chi-square goodness-of-fit tests to compare the proportion of occupied random plots within a region to an expected proportion that would have occurred if sage grouse had been randomly distributed across the Gunnison Basin during winter months. Distribution Among Topographic Categories Topographic features were recorded at sage grouse feeding sites during each of the 3 winters. Because winter search efforts in 1985 and 1986 were within discrete random plots, it was possible to determine whether sage grouse feeding sites were proportionately distributed among topographic features. Terrain within each random survey plot was classified by slope and aspect (Table 1). Where necessary I used a compass and abney level to verify slope aspect and steepness. Boundaries of terrain categories within random plots were drawn on topographic maps and each topographic type was searched on foot. Ideally, search effort and the likelihood of detecting use sites would have been independently assessed for each terrain category. to do so via line transects were made in 1985. Initial attempts However, due to the high degree of interspersion of terrain categories maintenance of straight lines was not feasible. Also, use sites in 1 terrain category were often located when an observer was in a different terrain type. Therefore, it was not possible to measure search effort independently for each topographic category. Instead, each survey plot was searched such that all terrain �III categories were sampled. At each sage grouse feeding site, terrain features were classified to 1 of the 7 possible categories. feeding sites overlapped 2 terrain categories. On a few occasions, large When this occurred, the observation was assigned to the terrain in which most foraging activity appeared to take place. Elevation was also recorded at feeaing sites. Because feeding sites were discrete, it was possible to observe >1 foraging area within a plot. To improve independence of observations, a feeding site was not included in the analysis if it was close «200 m) to a feeding site in the same plot that had been observed earlier in the survey. Discovery of adjacent, discrete feeding sites rarely occurred. Observed use of each terrain category in 1985 and 1986 was compared to expected use by chi-square goodness-of-fit tests. elevation categories «2,460 Distribution among 3 m, 2,461-2,615 m, >2,615 m) was also evaluated. Expected use was based on a uniform distribution of feeding activity across all terrain and elevation categories within survey plots where feeding activity was observed. Terrain categories in unused survey plots were not considered in calculation of availability. Availability of terrain types was estimated from topographic maps with a dot-grid area estimator. Confidence limits (95%) were calculated for the observed proportions of total use within each terrain. Confidence intervals were compared to expected proportions to evaluate whether observed use of a terrain category differed from expected use (Byers et ale 1948). Shrub Structure Shrub structure was measured at a sample of sage grouse feeding sites during each winter. Sites selected for measurement of shrub structure were those where approximate boundaries of the use area could be discerned from tracks or fecal droppings, and where browsed leaves were apparent on sagebrush �112 shrubs. Shrub structure -;as not measured at sites where sage grouse tracks were partially obscured by melted or windblown snow, or where feeding activity was not apparent. vinyl flagging. The approximate center of the use area was marked with I returned to marked sites and measured shrub structure following the winter period. A l5-m tape was stretched in direction over the marked center of the foraging area. a north-south Species of each shrub beneath the transect was recorded and sagebrush was classified as living or dead. Dead shrubs had living foliage on 20% of stems. individual shrubs beneath the tape were measured. Canopy dimensions of Individual shrub measurements included (1) length of transect intercept, (2) longest axis of crown, (3) crown width perpendicular to the longest axis, and (4) height from ground level to the tallest stem (excluding inflorescenses). Shrubs other than sagebrush were not recorded if they were less than 20 cm tall. Live sagebrush plants within 0.5 m of the l5-m transect were counted to obtain an estimate of density. Shrub dimensions and density were also measured along a 2nd, l5-m transect that crossed the center of the 1st line at a perpendicular angle. Shrub structure was also measured at 100 random sites in the Gunnison Basin. Random sites were stratified with 20 locations sampled in each of 5 terrain categories (Table 1). However, no distinction was made between 6-150 and >15 0 slopes; random sampling occurred on all slopes> 50 • Sites were selected within a random sample of the same l-km diameter survey plots used to evaluate winter distribution and habitat use. stratified among all regions of the Gunnison Basin. Survey plots were Within a survey plot I located the approximate center of the terrain category to be sampled, then walked 200 steps in a randomly selected direction to locate the site where shrub structure was to be measured. If the line of travel took me out of the �113 selected terrain, I choose another random direction and continued walking for 200 steps. In drainages, travel was either 200 steps upstream or downstream from the center. I rejected sites if no sagebrush was present and replacement sites were selected. Two, l5-m transects were oriented in north-south and east-west directions at random sites. Measurement of shrub structure and density at random sites followed the same procedures used at sage grouse feeding sites. Summary statistics were calculated for each use and random plot. These included (1) the total length of live big sagebrush canopy that was intercepted by transects (canopy cover), (2) the mean height of big sagebrush plants beneath the transects (mean height), (3) the standard deviation of big sagebrush plant height as a proportion of mean height (coefficient of variation for height), (4) the mean combined measures of crown length, width, and height (plant size), (5) the standard deviation of plant size as a proportion of the mean (coefficient of variation for plant size), and (6) the number of live sagebrush plants per m2 (density). I calculated correlation coefficients among variables and selected variables for analysis that were not highly correlated with each other. I used analysis of variance procedures to compare differences within each of these variables among terrain categories and between random and use sites. Kolomogorav-Smirnov All data were evaluated by distribution tests to insure that assumptions of normality were met (Conover 1980:357). Sagebrush Chemical Analysis Sagebrush foliage was collected at winter feeding sites in 1985 and 1986 for analysis of crude protein and monoterpene content. were located while searching l-km diameter survey plots. Winter feeding sites Sagebrush foliage was collected at winter feeding sites at the time of their discovery. Because �114 collection of foliage at a feeding site usually required 2 hours, it was not feasible to obtain sagebrush leaf samples at sites discovered late in the day. Sites not included in the analysis of sagebrush chemical content were marked for measurement of shrub structure. As with feeding sites where only shrub structure was measured, sagebrush foliage was collected only if approximate boundaries of the use area could be discerned. Foliage samples and shrub structure measurements were obtained along 2 l5-m transects near the center of the use area. often narrow «5 plants (~>10) Sage grouse feeding sites in the Gunnison Basin were m) and linear. To obtain an adequate sample of browsed at a feeding site, it was necessary to sample sagebrush plants along the path of feeding activity. Therefore, 1 transect was oriented along the axis the birds traveled while feeding. the 1st at a perpendicular angle. The 2nd line crossed the center of Snow cover beneath transects was removed, and shrub canopy dimensions and density were measured. Sagebrush plants beneath each transect were examined for evidence of feeding activity. Sage grouse feed only on the leaves of sagebrush and do not consume woody material. Browsed plants were distinguished by the dark, exposed mesophyll of clipped leaves. Plants were considered browsed if ~2 leaves had been clipped by sage grouse. Browsed and unbrowsed plants were marked with different colored vinyl ribbon. Leafy stems were collected from several areas of the crown of each plant. Samples were not obtained from plants that were totally covered by snow because these were not available to sage grouse. Each sample was individually sealed in a plastic bag and frozen. Sagebrush samples were also obtained at 32 randomly located sites during March and April 1986. Sagebrush was collected from 8 sites in each of 4 0 terrain categories (0-50 low, 0-5 sites were selected in the >5 o 0 high, drainage, >5 southwest). No northeast category because these areas were �115 usually snow-covered and not available to sage grouse. Sites were located in the same l-km plots used to evaluate winter distribution and habitat use. Selection of random sites within survey plots followed the same procedures used to select random sites for shrub structure measurements. Sagebrush foliage and shrub measurements were obtained along 2 l5-m transects oriented in cardinal directions. Foliage samples were individually bagged and frozen. I compared chemical content of pooled samples of browsed and unbrowsed plants at 1986 feeding sites. A 15-30 gm pooled sample of leaves was prepared from each group of browsed and unbrowsed plants for each sagebrush species within a feeding site. Equal amounts of leaves (1 gm minimum) were picked from individual plants so that all plants were equally represented in pooled samples. Pooled samples of leaves were also prepared from each random site. Samples were homogenized by grinding leaves in a mortar. Liquid nitrogen was added to the mortar immediately prior to grinding to facilitate fragmentation and to prevent loss of monoterpenes. Monoterpenes were extracted from samples by continuously dripping diethyl ether through 5-gm subsamples for 6 hours in a soxhlet apparatus. flasks and saved. The extract was captured in A carvone standard (0.625 ug/ul) was added and the volume increased to 100 ml by adding diethyl ether. Approximately 20 ml subsamples were retained for analysis. I used a Perkin-Elmer gas chromatograph with a hydrogen flame ionization detector to separate and quantify monoterpenes (Table 2). ul) were carried through a 30 m tubular column. identified via mass spectrum analysis. Extract samples (4 Individual monoterpenes were Identification was made by comparison to published spectral patterns (Epstein et ale 1976, Heller and Milne 1978). �116 Table 2. Gas chromatograph programming specifications used to separate and quantify monoterpenes in Artemisia tridentata foliage from the Gunnison Basin, Colorado. Carrier gas Helium Flow rate 1.5 m1/min (approx) Initial temperature 70 C Initial rate of temperature increase 2 C/min Final temperature 130 C Final rate of temperature increase 20 C/min Post temperature 300 C Approximately 5 gms of homogenized leaves were retained from pooled samples for analysis of crude protein in random site, browsed, and unbrowsed plants. Samples were freeze dried to remove moisture and dry matter percentages were determined. Nitrogen content was evaluated by microkje1dah1 procedures (Horowitz 1980). Nitrogen content was mUltiplied by 6.25 to obtain an estimate of crude protein. Comparisons of sagebrush chemical content were made between browsed and unbrowsed plants at feeding sites, and among random sites in different terrain categories via analysis of variance procedures. Sage Grouse Lipid Analysis Adult male sage grouse were collected annually on leks in Jackson County during 2 sampling periods (early and late season courtship) between 1983 and 1985. The early season sampling period occurred during the first 2 weeks that males consistently displayed on leks, and was usually between 1 and 15 April. Severe winter weather delayed courtship behavior in 1984; males were not �117 collected until the final 2 weeks of April. was in mid- to late May. The late season sampling period In Gunnison County, early and late season collections of adult males were made in 1984 and 1985. In 1986 adult males in Gunnison and Jackson counties were collected only during the early part of the breeding season. Between 8 and 10 males were collected in each sampling period during all years in both populations. Males were weighed immediately upon collection, then sealed and frozen in plastic bags. Males were thawed and feathers were removed in preparation for carcass analysis. The head, tarsi, and wings distally from the carpals were removed and discarded. weighed. Breast muscles on 1 side of the body were excised and Internal organs were also removed and weighed or measured. contents were removed, weighed, and discarded. Gut Muscles and internal organs were returned to the carcass. The carcass was refrozen then homogenized 6-8X in a commercial meat grinder. The homogenate was spread on aluminum foil, dried at 80 C for 24 hours then further homogenized in a high speed Wiley Mill. Lipids were extractd from 10-gm subsamples of homogenate with diethyl ether for 6 hours. Weight of lipids in the dried homogenate was determined and the percentage of lipids as live body weight calculated. Comparison of Adult Male Courtship Behavior Between Populations Courtship behavior of adult male sage grouse in Gunnison and Jackson counties was compared in 1986 to test the relationship between strutting display rates and lipid reserve size. for study. I selected 3 leks in each population Leks were accessible by vehicle and were attended by similar numbers of males. Courtship behavior was observed during 5 mornings on each lek between 9 April and 6 May. All observations were made during mornings when low temperatures ranged between -8 and 2 C, winds were less than 16 kmihour, and there was no precipitation. Observers arrived 1.75-2.0 hours prior to sunrise to record time of 1st display. �118 Evaluation of courtship behavior began as soon as light was adequate to distinguish males from females (usually about 0.75 hr before sunrise). Numbers of females and adult male sage grouse present on leks were recorded. Lek counts were repeated at approximately 20-minute intervals. Adult males were randomly selected for evaluation of strutting display rates. Adults were distinguished from yearlings by their large size and distinctive nuptual plumage development. Selected males were observed for 5 minutes and the number of strutting displays observed during that time was recorded. Time that males were involved in territorial disputes (chases, wing thrashes, and face-offs) during the 5 minutes of observation was recorded. I also recorded time birds spent in crouched positions in response to potential avian predators and common ravens (Corvus corax). Number of displays/minute was calculated for each male. Display rates were corrected for time of disruption due to territorial disputes or avian predator disturbance. Males for which disruption time exceeded 2 minutes were not used for comparison of display rates between populations. Distance between a randomly selected male and the nearest female during the observation period was recorded as either <10 m, 10-30 m, or >30 m. «20 Small em-high), orange-flagged stakes were placed at 10 m-intervals on leks to facilitate distance estimation. Evaluation of display rates of randomly selected males continued until the number of males remaining on the lek was <25% of the maximum number of males observed during the morning survey period. Time at which 75% of the males had abandoned the lek was recorded. Lek Surveys Counts of sage grouse attending 19 leks in the Gunnison Basin were made between 27 March and 21 May in cooperation with local District Wildlife Managers. Lek surveys were usually conducted between 0.5 hours before and 1.0 hours after sunrise. �119 RESULTS Snow Depth Variation Among Winters and Terrains Winter conditions were severe in 1984 (Table 3). Between January and March, mean low temperatures were 3-8 C below the 3D-year average. cover persisted into April. Heavy snow Winter temperatures were similar to or slightly above normal in 1985 and 1986 and snow depths were less than in 1984. Ground transect snow measurements reflected differences in winter severity. February snow depth measurements were obtained in 3 terrain categories during each winter. In 1984, snow depths were approximately 1.5X greater than during 1985 and 1986 (Table 4). of 7 terrain categories in 1985 and 1986. Snow depth was measured in each In both years snow depths increased between January and February, and decreased in March (Fig. 2). deepest in drainages and on 6-150 and >150 northeast aspects. 0 shallow on 6-15 Snow was Snow was and >150 southwest slopes while depths were intermediate on 0-50 high and 0-50 low sites. Aerial Transect Estimates of Sagebrush Exposure Winter severity dramatically affected mid-winter sagebrush exposure in the Gunnison Basin. (Table 5). Deep snows greatly reduced sagebrush availability in 1984 Exposed sagebrush was present in only 7% of the areas surveyed during aerial transects. «50%) I considered sagebrush exposure to be moderate at all sites where foliage was available. Sagebrush was much more available during the more moderate winters of 1985 and 1986. Exposed sagebrush was available in approximately 80% of the areas surveyed along aerial transects during those years. I considered sagebrush exposure to be excellent (>50%) at 11 and 24% of the sites where sagebrush crowns were visible in 1985 and 1986 respectively. �t-' N Table 3. Departure of mean winter (Nov-Mar) temperatures from 30-year (1951-80) averages and monthly snowfall totals in Gunnison County, Colorado 1983-86. Data were obtained from National Oceanic and Atmospheric Administration monthly Climatological Data for Colorado. Departure of mean temperature from 1951-80 average Mean Nov Dec Snowfall (cm) Jan Feb Mar departure Nov Dec Jan Feb Mar Totals 1983-84 -0.2 1.7 -8.1 -6.0 -2.9 -3.1 37.6 96.0 6.4 22.9 22.9 186 1984-85 0.7 3.7 2.0 -0.7 2.9 1.7 6.4 19.1 29.2 2.5 39.4 97 1985-86 1.2 -1.2 -0.2 6.7 5.2 2.3 20.3 26.7 6.4 34.3 2.5 90 o �121 Table 4. Mean February snow depths in 3 terrain categories in the Gunnison Basin 1984-86. Snow depths in 1984 were from randomly located transects in Chance and Graflin gulches. Snow depths in 1985 and 1986 were from ground transects in randomly located l-km diameter survey plots throughout the Gunnison Basin. 1984 Terraina 1984 1986 cm N cm N cm 1! Drainage 65 108 39 244 38 169 6-15°, southwest aspect 39 56 18 230 28 139 6-15°, northeast aspect 57 132 37 229 46 146 aDescription of terrain categories are in Table 1. �122 SNOW DEPTHS -1986 50 - - ) 1 5 NORTHEAST 40 ~ 6-15 o J: f- NORTHEAST 30 a. UJ c $ o z tJ) 0-5 LOW 20 , DRAINAGE 0-5 HIGH 10 6-15 ) 15 SOUTHWEST o FEB JAN 1985 _ - DRAINAGE 40 •....•.... 6-15 NORTHEAST /-----~~~---------. ) 15 NORTHEAST " .._ _ ..-.._ _ ..-.._ _ _ .•.•. o f- a. MAR SNOW DEPTHS ~ J: SOUTHWEST .. ../"'_ .. .. .. .. ,,/ 30 .._ .._ .._ ....:.'4-.. ,,,,,,,,, ._ ••• 0 - 5 LOW 0 UJ o 0-5 HIGH $ 20 o z so 10 ) 15 SOUTHWEST o JAN Fig. 2. Mean winter 1985-86. FEB snow depths in the Gunnison MAR Basin, Colorado, �123 Table 5. Mid-winter availability of sagebrush in 8 regions of the Gunnison Basin, 1984-86. Availability was estimated during aerial transects. Exposed sagebrush in mid-winter (%) Regiona 1984 1985 1986 6.7 81.0 69.2 11.7 84.1 70.7 3.5 84.2 80.0 72.5 84.8 7.9 86.3 89.7 Beaver 11.7 95.4 92.2 Chance 11.8 84.7 79.7 Doyleville 7.6 81.0 57.7 Average 6.7 83.7 78.0 Tomichi Parlin McIntosh Ob Steuben Sapinero ~egion boundaries are delinieated in Fig. 1. bSevere air turbulence may have affected sagebrush exposure estimates in the Steuben region. �124 Winter Distributi~n -~ I observed 95 sage grouse winter use sites in 1984. Use sites were present throughout the Basin (Fig. 3), although some areas appeared to be used more heavily than others. A high percentage (55%) of use sites occurred south of Gunnison between South Willow Creek east to Graflin Gulch. This included the Sugar, Pole, Camp, South Beaver, Willow, and Gold Basin Creek drainages, as well as the Stubbs, Chance, and Sage Hen Gulch areas. I also observed use of areas east of Gunnison between Signal Peak and Cabin Creek, as well as in the vicinity of North Parlin Flats. Scattered use sites occurred near the north end of Sapinero Mesa, lola Access Area, East Antelope Creek, and Woods Gulch. Helicopter crews involved in aerial feeding of elk herds observed scattered flocks along the rims of mesas north of Blue Mesa Reservoir. I observed little evidence of sage grouse use in areas south of Parlin and Doyleville and east of Colorado Highway 114. Although approximate boundaries of areas searched in 1984 were marked on topographic maps, I cannot be assured that search effort was uniform among all survey areas. Therefore, statistical analyses of distribution of use sites among regions would not be valid and were not conducted. Ground searches for use sites were conducted in 142 and 120 l-km diameter survey plots in 1985 (Fig. 4) and 1986 (Fig. 5), respectively. The smaller 1986 sample was due to heavy, wet snow conditions in late February that prevented use of snow machines and limited access to some plots. Sage grouse use sites were observed on 51 and 52 plots in 1985 and 1986, respectively. In both years the distribution of used plots among regions did not differ from a distribution that would occur if sage grouse randomly occupied the Gunnison Basin (R> 0.10, Table 6). Preference for one or more regions of the Gunnison Basin was not apparent in 1985 and 1986. This suggests that areas of concentrated winter use did not exist during those winters. �~ A\ 012345 KILOMETERS I PI I~l •••••• •• • ~ ..o ! ,;; Fig. 3. Sage grouse winter use sites in the Gunnison locations represent >1 use site. Basin, Colorado, 1984. Some •.... N lJl �~ N 0'\ ~ ~ 012345 KILOMETERS 0 • 0 ~\~ (,0 ,J' 7 I 0 0 0 0 ~ • • 0 Ooyle':.l"e o f o .. I/) Fig. 4. Sage grouse winter survey circles represent used plots. plots in the Gunnison Basin, Colorado, 1985. Darkened , \{! �~ A\ \. 0\:\ 01 21--n sf KILOMETERS o o 7 • ~ • 0 o •• • o 0 ~ . • • 0 Parlin O. ~ •~ 0 - -0 0 Dayle~lIe. o ~ Fig. 5. Sage grouse winter survey plots in the Gunnison circles represent used plots. Basin, Colorado, 1986. Darkened I-' N -...j �128 Table 6. Observations of sage grouse use in randomly selected 1-km diameter survey plots in the Gunnison Basin, January-March 1985, 1986. Expected number N random !! plots of plots with plots with use sites use sites 1985 1986 1985 1986 1985 1986 Tomichi 11 9 6 4 4 4 Parlin 20 16 4 10 7 7 McIntosh 10 8 2 4 4 3 Steuben 10 10 0 4 3 4 Sapinero 17 14 6 5 6 6 Beaver 27 22 11 10 10 10 Chance 27 22 11 9 10 10 Doy1evi11e 20 19 11 6 7 8 142 120 51 52 51 52 Re~iona Totals 1985 X2 = 8.78 with 7 d.f., 0.25> P > o.r. 1986 X2 ~egion 2.10 with 7 d. f , , P>0.95. boundaries are delineated in Fig. 1. �129 Distribution of Feeding Sites Among Topographic Categories In 1984, topography was classified at 91 sage grouse feeding sites (Table 7). A high percentage (73%) occurred in drainages (35%) or 6-15 with southwest aspects (38%). 0 low, 0-5 o slopes Fewer feeding sites were observed on 0-50 high, and 6-150 northeast slopes (Table 7). Due to difficult o access I made few attempts to survey use of steep (>15 ) slopes, therefore, use of these sites in 1984 is not known. I observed 74 winter feeding sites within 51 used survey plots in 1985 and 83 feeding sites within 52 used survey plots in 1986. Feeding activity was not proportionally distributed among terrain categories in either year (~< 0.001, Table 7). A high percentage (23-33% of feeding sites) of observations occurred in drainage sites in both 1985 and 1986, even though these areas comprised only 3-4% of the survey plots. A high percentage (22-28%) of sage grouse feeding sites also occurred on 6-150 southwest slopes in each year, Use of 0-50 low sites was although use was proportionate to availability. greater (37% of feeding sites) in 1985 than in 1986 (11% of feeding sites). 0 Feeding activity (11-16% of feeding sites) on 0-5 proportional to availability. high sites was I observed little use (1% of feeding sites) on >150 southwest slopes in 1985 while use of these sites was slightly greater (15% of feeding sites) in 1986. In both years I observed little (1-2%) feeding activity on 6-150 and >150 slopes with northeast aspects even though these terrains comprised 30-31% of the survey plots. Flock sightings and track observations were used to locate feeding sites. This analysis of sage grouse distribution among topographic features is not accurate if these 2 criteria do not provide similar estimates of use among terrain features. Dissimilarities could occur if more rapid melting of snow on southwest aspects caused tracks to be less visible than tracks in more sheltered sites such as drainages or northeast slopes. would not be subject to this potential bias. Flock observations I compared the distributions �I-' l;.) Table 7. Distribution parentheses. of sage grouse winter Confidence limits feeding (95%) of observed Observed 1984a Terrain sites among terrain categories proportions were estimated in the Gunnison following use Basin, 1984-86. Proportions of annual totals are in Byers et a1. (1984). 95% confidence limit EXl2ected use c 1985 1986 1985 1986 1985 1986 10 (0.110) 27 (0.365) 9 (0.108) 0.214::.f.1::. 0.516 0.016::.f.1::. 0.200 17.2 (0.232) 15.9'(0.191) 10 (0.110) 12 (0.162) 9 (0.108) 0.047::.f.2::. 0.277 0.016::.f.2::. 0.200 14.1 (0.190) 13.0 (0.157) 32 (0.352) 17 (0.230)d 27 (0.325)d 0.098::.f.3::' 0.362 0.187::,f.3::' 0.463 3.0 (0.040) 3.9 (0.047) southwest 35 (0.385) 16 (0.216) 23 (0.277) 0.087::'~::' 0.345 0.145:::~::' 0.409 12.7 (0.172) 16.6 (0.200) 6-150 northeast 4 (0.044) 0::.f.5::' 0.051 0::.f.5::' o. 044 19.5 (0.264) 15.8 (0.190) O::'~::'0.051 0.041::'~::' 0.249 3.4 (0.046) 9.0 (0.1()8) 0::.f.7::. 0.069 4.1 (0.055) 8.9 (0.107) 0-50 low 0 0-5 high Drainage 0 6-15 1 (0.014)e 0 southwestb 1 (0.014) 0 northeastb o > 15 > 15 aBased b on non-random systematic Steep slopes not surveyed cExpected surveys 1 (0.012)e 12 (0.145) (O.O)e 2 (0.024)e between 7 January Observed eObserved Availability not estimated. for use in 1984 due to deep snow conditions. use derived from the proportional use greater than expected availability grouse. d and 31 March. I~! use less than expected . use (f.< 0.05). use (f.<0.05). of terrain categories within randomly selected 1-km diameter survey plots used by sage o �131 of track and flock sightings among terrains via chi-square contingency table analysis. This test determines whether the~estimated proportional use of any terrain category differed between track and flock observations. 1985 and 1986 data for this analysis. I combined There was no difference in topographic distributions of track and flock sightings (0.25> R. > 0.10, Table 8). I concluded that each sighting criteria provided similar estimates of topographic use, and that combining track and flock observations for analysis of topographic distribution was valid. Table 8. Numbers of sage grouse flock vs. track observations within terrain categories of 1-km diameter plots surveyed between January and March, 1985-86. Values in parentheses are expected numbers that would occur if the distribution of flock and track sightings were similar across terrain categories. a Terrain 0 Observation 0-5 0-50 criteria Low High 13 5 Flocks (12.3) Tracks 23 (23.6) X2 ( 7.2) 16 (13.7) 6-150 southwest Drainage 11 16 (13.4) (15.1) 33 23 (28.9) (25.6) Other b Totals 9 (5.8) 8 (11.1) 6.2 with 4 d.£., 0.25> P> 0.10c aTerrain categories are described in Table 1. boo 6-15 northeast, >15 southwest, and >15 0 northeast observations were combined due to small numbers of observations. c Flock and track observations are similarly distributed among terrain categories. 54 103 �132 Distribution Among Elevation Categories I compared the distributions of 1985 and 1986 sage grouse feeding sites among 3 elevations zones. In each winter sage grouse were not proportionately distributed among elevation categories (~<0.001), less than expected. use of sites >2,615 m was However, these findings should not be interpreted as evidence that sage grouse avoid sagebrush stands >2,615 m. Topograpic catgories were not proportionately distributed among elevation zones (~ 0.001), 0-5 o elevations. availability. < low sites occurred less frequently above 2,615 m than at lower I eliminated observations in 0-5 o low categories and adjusted I considered the elevation distribution of remaining observations in other terrains and found that distribution among elevation zones was proportional to availability (~>0.25) in each year. I conclude that, within the areas surveyed, there is no evidence that sage grouse were not proportionately distributed among elevation zones. Sagebrush Structure Selection of Variables for Analysis.--I calculated 6 summary variables-for each random and sage grouse feeding site. Plant size and the coefficient of variation for plant size were highly correlated (E>0.7) with either mean height or the coefficient of variation for height (Table 9). Analysis of individual structural measures other than height or the coefficient of variation for height would not have yielded additional information regarding structural characteristics of use and random sites. variables selected for analysis were: Therefore, the 4 (1) canopy cover, (2) mean height, (3) coefficient of variation for height, and (4) density. measured different components of sagebrush stands. of overhead cover provided by sagebrush. Each of these variables Canopy cover was a measure Mean plant height was a measure of average plant size at use and random sites. The coefficient of variation for �133 height indexed the variation in sagebrush plant size relative to average height. Density was a measure of sagebrush plant spacing. Table 9. Correlation coefficients (E), examining relationships among pairs of big sagebrush structural variables measured at sage grouse winter feeding and random sites in the Gunnison Basin, Colorado, 1985-86. Coefficient Category Canopy Coefficient Mean of variation Plant of variation height for height size for size 0.56 0.12 0.33 0.04 0.05 -0.02 0.77 -0.07 -0.22 -0.09 0.66 -0.04 -0.14 -0.37 Mean height Density Coefficient of variation for height Plant size Coefficient of variation for size -0.05 Random Site Variation Among Terrains.--Shrub species composition and structural differences were apparent among randomly selected sites in different terrains (Table 10). canopy cover) most sites. Big sagebrush dominated (comprised >50% of However, black sagebrush (~. nova) was abundant (present at 13 of 20 random sites) on xeric 0-5 0 high sites. Black 0 sagebrush was the dominant sagebrush species at 9 of 13 random, 0-5 high �Table 10. Shrub species composition and big sagebrush structural characteristics among random sites in 5 terrain categories (N Variables = 20 sites/terrain), Gunnison Basin, Colorado, 1985. 0-50 0-50 low high Standard errors are in parentheses. 50 Drainage southwest f-' W 50 northeast F P a Number of sites with black sagebrush 1 13 0 8 9 , !~! Number of sites with rabbit brush Canopy, cm Mean height, cm 6 2 16 1 7 855 367 959 478 636 (47) (54) (83) (66) (82) 43.3 31.6 52.8 28.8 41.5 (2.9) (2.1) (1.9) (2.5) (3.5) 29.6 27.6 31.2 29.9 27.8 (1.5) (2.8) (1.6) (2.2) (2.1) 1.3 2.0 1.3 1.4 1.4 (0.08) (0.20) (0.10) (0.18) (0.11) 13.3 0.001 13.4 0.001 Coefficient of variation for height, % Density, plants/m a 2 Probability that structural variable is similar among all terrain categories. . 0.53 0.71 3.85 0.006 ~ �135 sites where it was present and northeast sites. 0 and was also present It was rarely present southwest on some xeric >6 in mesic 0-50 low and drainage terrains. Rabbitbrush (Chrysothamnus spp.) was also not evenly distributed terrains (Table 10). Rabbitbrush drainage sites (16 of 20 sites). low and >60 northeast >60 southwest plants >20 cm high were common in the mesic Rabbitbrush slopes. structural variables 0.71, Table 10). This indicates height was similar among terrains. height, and sagebrush density plants in drainages and canopy cover was greater of sagebrush high sites. shorter 0 southwest >60 northeast drainages. sites. slopes. sites was low relative Big sagebrush 0 (~<0.006, sites were similar was shortest Sagebrush and >60 southwest density sites. in other terrains on 0-50 sites were on 0-50 canopy cover at 0-50 Sagebrush to sagebrush on >60 southest Canopy cover in these terrains was low relative terrains. to stands, while canopy cover on to drainages. 0 was higher on 0-5 = Table 10). to density northeast Big sagebrush to drainage (~ site canopy cover, mean was low relative low and >6 relative sites, but taller than sagebrush and similar 0-50 low and >60 northeast o terrains of other than 0-50 low. than in all terrains plants on 0-5 low sites was vigorous Random were taller than those growing than plants in drainage high sites and >6 in plant height varied among terrains plants in drainages Sagebrush only the coefficient sites in different the variation average Density analyzed, for height was similar among random Sagebrush on some 0-50 was also present sites, but rarely grew on the xeric 0-50 high and Of the 4 sagebrush variation among densities density 0 and 0-5 in in high to that in most other high sites than in drainages �136 Comparison of Transect Orientation at Feeding Sites.--Shrub structure was measured at feeding sites where foliage was collected for chemical analysis as well as at sites where collection of foliage did not occur. obtained after the winter season for the latter group. orientation differed between the 2 groups of samples. Measurements were Line transect Transects were oriented along and perpendicular to the direction of sage grouse feeding at the foliage collection sites, but were aligned in cardinal directions at sites where only shrub measurements were made. Differences in sampling of shrubs could result in different estimates of shrub structure. If transect orientation affected shrub structure estimates, then comparison of feeding site shrub structure at foliage collection sites to random site structure would be incorrect because any observed differences could be due to different sampling procedures. If transect orientation did not affect structure estimates, then shrub measurements from all feeding sites could be combined and collectively analyzed regardless of transect orientation. This would be advantageous because sample size would increase. To evaluate whether structure estimates varied between transect alignment methods, I used both approaches to obtain replicate measurements of structure at 9 winter feeding sites where foliage was collected for chemical analysis. Estimates of sagebrush structure were compared between feeding direction and cardinal direction with paired! tests. Mean structural measures for the 9 sites were comparable between transect alignment methods (Table 11). I also compared variation in height of individual big sagebrush plants (~ = 372 plants) among sites and between transect orientation methods with a 2-way analysis of variance. Site effects were significant (F = 20.0, ~ <0.001) while transect alignment effects were not (I = 0.001, ~ = 0.975). This suggests that shrub height was much more variable among sites than between �137 methods. Different transect orientation approaches did not affect estimates of shrub structure. This is probably because the area available for sampling was a small 15-m diameter circle. was usually homogenous. Shrub structure within these small areas Therefore, I used structural data obtained at all feeding sites (regardless of transect orientation) in comparisons of winter feeding and random sites. Table 11. Sagebrush structural characteristics obtained by 2 methods of line transect orientation (feeding direction and cardinal direction). Replicate measures of sagebrush were obtained using both transect orientation methods at 9 winter feeding sites. Transect orientation Feeding direction Structural variable Canopy, cm x 1,351 Mean height, cm SE 117.6 Cardinal direction x 1,243 SE t P 88.4 1.5 0.17 42.5 2.4 43.0 2.8 0.30 0.78 35.7 0.02 34.9 0.03 0.58 0.58 1.8 0.29 1.8 0.30 0.18 0.87 Coefficient of variation-height, % Density, plants/m 2 Differences in Feeding Site Shrub Structure Between Years.--I measured shrub structure at 18, 37, and 50 winter use sites in 1984, 1985, and 1986, respectively. Because sites searched in 1984 were not randomly selected, I cannot assume that shrub structure at located feeding sites was representative of feeding sites within the Basin during that winter. This invalidates �138 statistical comparison of 1984 winter feeding sites with random and feeding sites in other winters. However, structural measures for 1984 feeding sites are presented for qualitative comparison with other winters (Table 12). During analysis of shrub structure I concentrated on data gathered at feeding sites within random plots in 1985 and 1986. I evaluated differences in structural characteristics of winter feeding sites between 1985 and 1986 (Table 12) within terrain categories where observations were available for both years. Statistical comparisons were only made between sites within a similar terrain category. Of the 16 comparisons (4 structural variables x 4 terrain categories) only 1 showed a significant difference between years; sagebrush density in drainage feeding sites was higher in 1985 than in 1986. Because differences between years were minimal, I pooled 1985 and 1986 feeding site shrub structure data for comparison with random sites. Comparison of Winter Feeding and Random Sites.--Sagebrush structure at random sites differed among topographic categories (Table 13). characteristics at sage grouse feeding sites (~ = 87, 1985 and 1986 sites pooled) also varied among terrain categories (Table 13). was greatest at drainage and >50 northeast sites. 0 low sites had higher density than sites on >5 Structural Mean plant height Feeding sites in 0-50 southwest sites. Because of structural variation associated with topographic features, feeding sites were only compared to random sites within the same terrain category (e.g., drainage use sites vs. drainage random sites). Differences between feeding and random sites were most pronounced in the xeric >50 southwest and 0-50 high terrains (Table 13). Within these categories, sage grouse used sites with greater canopy cover and taller plants than occured at random sites. Density and coefficient of variation for height �139 Table 12. 1984-86. Big sagebrush structure at sage grouse winter feeding sites in the Gunnison Basin, Colorado, Standard errors are in parentheses. Terrain categor! Variable 0-50 0-50 low high a a Year >50 Drainage southwest N sites 1984 8 10 1985 20 4 6 7 1986 5 6 15 21 Canopy, cm a 1984 a 1,057 (76) 631 (176) 827 (123) 1985 1,090 (62) 928 (160) 1,276 (121) 1986 1,029 (129) 989 (127) 1,066 (69) 0.67 0.77 pb 0.13 1,066 (89) 0.18 Mean height, cm a 1984 a 64 (5.8) 33 (5.5) 1985 43.5 (1.9) 45.7 (5.7) 53 (2.0) 41 (2.7) 1986 45.7 (3.1) 43.5 (4.6) 54 (4.1) 42 (2.0) pb 0.61 0.77 0.89 0.94 Coefficient of variation for height, % a 1984 a 38 (0.02) 31 (0.03) 1985 31.5 (0.01) 36.4 (0.04) 29.7 (0.01) 31.3 (0.01) 1986 27.2 (0.02) 33.1 (0.03) 3i.5 (0.02) 33.0 pb 0.11 Density, p1ants/m 1984 0.51 0.67 0.53 1.1 (0.07) 1.2 (0.25) 2 a a 1985 1.9 (0.14) 1.3 (0.23) 1.8 (0.16) 1.1 (0.15) 1986 1.9 (0.39) 1.4 (0.18) 1.3 (0.08) 1.2 (0.15) pb 0.98 0.68 0.006 0.25 aNo measurements obtained at use sites in this terrain in 1984. b Probability that no difference exists between 1985 and 1986 measures. �140 Table 13. Sagebrush struc6tra1 characteristics at 1985 and 1986 sage grouse winter f eed.Lng sites and random locations in the Gunnison Basin, Colorado. Coefficient Mean of variation N Canopy height height Density Elots (cm) (cm) (%) (Elants/m2) Random 20 855 43.3 29.6 1.3 Feeding 25 1,078 43.9 31.0 1.8 Terrain categor:z: 0-50 low F 8.8 0.03 0.34 5.5 pa 0.005 0.86 0.56 0.02 0 0-5 high Random 20 367 31.6 27.6 2.0 Feeding 10 965 44.4 34.5 1.4 10.9 2.7 3.9 0.11 0.06 F 34.6 P <0.001 0.003 Drainage Random 20 959 52.8 31.2 1.3 Feeding 21 1,127 54.1 31.0 1.5 F 2.7 0.13 0.01 2.9 P 0.11 0.73 0.94 0.10 >50 southwest Random 20 478 28.8 29.9 1.4 Feeding 28 1,007 41.1 32.6 1.4 25.1 20.0 1.5 0.5 <0.001 <0.001 0.23 0.5 F P >5 0 northeast b Random Feeding aprobabi1ity 20 636 41.5 27.8 1.4 3 915 51.6 28 1.0 that use and random site measures are similar. bNo statistical comparison due to small feeding site sample. �141 did not differ between feeding and random sites. In the 0-50 low terrain, differences between use and random sites were apparent. Canopy cover and density were greater at feeding than at random sites but there were no differences between mean height and the coefficient of variation for height. Use site structure was almost identical to random site structure in the mesic drainage category. 0 Comparison of use and random sites in the >5 northeast terrain was not possible due to the inadequate sample (~ = 3) of feeding sites. I observed black sagebrush at only 8% of 87 winter sites while it was present at 31% of 100 random sites. Within sites where it occurred, black sagebrush canopy intercept was similar (p and random sites (~ = 382 cm). random sites. = 0.14) between feeding (x = 223 cm) Rabbitbrush occurred at 32 feeding and 38 Rabbitbrush canopy cover within sites where it was present was also similar (~ = 0.10) between feeding (x = 66 cm) and random sites (x = 101 cm) • Chemical Variation Among Random Foliage Collection Sites I collected sagebrush plants at 8 random sites in each of 4 terrain categories. Big sagebrush plants were present at each site (~ = 21 Black sagebrush was present at 7 sites (~ = 20 plants/site, SE 1.1). plants/site, SE 2.6) and was also sampled. Big sagebrush plants in drainages were taller than plants in other terrains (Table 14). This reflects variation in plant growth and vigor that occurred among terrains. Despite differences in site conditions and sagebrush growth forms, there were no differences in crude protein or total mono terpene content among big sagebrush plants that grew in different terrains (~> 0.05, Table 14). There were also no topographically-related differences in the 6 individual monoterpenes that comprised >90% of the total monoterpene content (p > 0.05, Table 14). Crude protein and total monoterpene content were highly variable �142 --~.., Table 14. Mean height, crude protein total and individual mono terpene contents of big sagebrush leaves collectd at 32 random sites in 4 terrain categories 1986. (8 sites/terrain), Gunnison Basin, Colorado, March-April Values other than mean height are expressed as percentage of dry matter. Standard errors are in parentheses. 0-5 0-50 low hi~h 0 Cate~orl Mean height, cm Crude protein Total monoterpenes >50 Draina~e southwest 35.3 36.4 48.2 37.8 (2.9) (3.2) (3.1) (3.5) 19.7 22.2 21.6 21.5 (1.0) (1.5) (1.6) (0.9) 2.08 1.86 2.23 2.06 (0.23) (0.25) (0.17) (0.14) 0.16 0.15 0.15 0.15 (0.02) (0.03) (0.01) (0.01) 0.09 0.10 0.08 0.12 (0.0l) (0.02) (0.01) (0.0l) 0.46 0.40 0.48 0.48 (0.06) (0.06) (0.03) (0.04) 1.03 0.95 0.96 1.01 (0.11) (0.12) (0.06) (0.05) 0.10 0.08 0.10 0.10 (0.0l) (0.0l) 0.05 0.05 F P 0.73 0.54 0.60 0.62 0.06 0.98 1.22 0.32 0.60 0.62 0.19 0.91 0.59 0.63 2.98 0.05 Individual monterpenes Camphene Arthole 1,8 Cineole Camphor Unknown A (retention time, 10.2 min) Unknown B (0.02) 0.05 (0.008) 0.03 (retention time, 28 min) (0.003) (0.007) (0.004) (0.005) �143 among sites within a single terrain. There was no correlation (p >0.05) between mean plant height and levels of crude protein (~ = -0.21) and total monoterpene (~ = 0.26) in big sagebrush plants sampled at random sites. Black sagebrush crude protein (~= 18.1%) and total mono terpene content (x 1.08) were lower than big sagebrush protein (x = 21.3%) and total monoterpene content (x = 2.15%) at 7 sites where each was found. I did not attempt to evaluate black sagebrush chemical differences among terrains due to small sample sizes in some categories. Chemical Differences Between Browsed and Unbrowsed Sagebrush Plants Sagebrush foliage samples were collected from 20 winter feeding sites in 1986. sites. Big sagebrush dominated (>50% of total canopy cover) all feeding Black sagebrush was present at 1 feeding site. (x = 15 plants/site, SE A total of 300 browsed 1.0) and 285 unbrowsed (~ = 14.3 plants/site, SE = 1.5) big sagebrush plants was collected from feeding sites. Total monoterpene content and 6 individual monoterpenes were similar (p > 0.05) between browsed and unbrowsed big sagebrush plants at winter feedingsites (Table 15). samples. Monoterpenes averaged 2.0% of dry matter in each group of Total monoterpene content at feeding sites was similar to values observed at random foliage collection sites (Table 14). Crude protein content was also similar (p = 0.80) between browsed (x = 20.6%) and unbrowsed (~ = 20.3%) big sagebrush plants at winter feeding sites (Table 15). Crude protein content of big sagebrush plants at feeding sites was similar to values observed at random foliage collection sites. �144 Table 15. Crude protein, and total and individual monoterpene contents of leaves of browsed and unbrowsed big sagebrush plants at sage grouse winter feeding sites, (~= 20) Gunnison Basin Colorado, January-March 1986. errors are in parentheses. Standard All values are percentage of dry matter. Category Crude protein Total monoterpenes Browsed Unbrowsed 20.6 20.2 (1.09) (1.07) 2.00 2.01 (0.10) (0.12) 0.14 0.15 (0.01) (0.01) 0.08 0.09 (0.009) (0.01) 0.41 0.41 (0.02) (0.03) 1.01 1.05 (0.06) (0.06) 0.10 0.10 (0.01) (0.01) 0.04 0.05 F P 0.06 0.80 0.001 0.97 0.22 0.64 0.04 0.83 0.006 0.94 0.17 0.68 0.002 0.96 0.37 0.55 Individual monoterpenes Camphene Arrho1e 1,8 Cineole Camphor Unknown A (retention time, 10.2 min) Unknown B (retention time, 28.0 min) (0.002) (0.003) �145 Male Lipid Reserves Early courtship period lipid reserves of adult male sage grouse were lower following severe winters than following normal or mild winters. Jackson County experienced heavy snow accumulations in 1984 and 1986 (Table 16). winters of 1983 and 1985 were less severe. The Early spring lipid reserves of adult males in Jackson County were low (~ = 2.3-3.6% of live weight) following the severe 1984 and 1986 winters, but were higher (~ = 5.5% of live weight) following the milder 1983 and 1985 winters (Table 16). Gunnison County also experienced a severe winter in 1984. Early spring lipid reserves were low (~ = 2.4% of live weight) in 1984 relative to lipid reserves (x = 4.1-4.2%) following the mild 1985 and 1986 winters. During mild winters, adult male sage grouse in Gunnison County did not accumulate as large of fat reserves as males in Jackson County. Comparison of Male Courtship Displays A total of 23.6 hours (284 5-min observations) was spent measuring display rates of adult male sage grouse in Gunnison and Jackson counties. Mean display rates were calculated for each lek-female distance combination within a population (Table 17). Hotellings!2 was used to test the hypothesis that mean display rates across lek-female distance combinations differed among populations. Hotellings!2 is a multivariate statistical procedure that simultaneously tests differences between populations across several variables (Norusis 1986:117). During morning display periods, adult male sage grouse in the Gunnison Basin displayed less rapidly than males in Jackson County (Hotellings !2 64.2, P = 0.023). Males in Gunnison County spent longer periods sitting passively on leks without displaying. Strutting display rates were not evaluated during evening periods, however, leks in both populations were �t-' .j:-- Table 16. c- Total winter (Nov-Mar) snowfall, lipid reserves, and body and breast muscle weights of adult male sage grouse in Gunnison and Jackson counties, Colorado, during early courtship periods (Apr), 1983-1986. 1983a b Snowfall, cm 1984 1985 1986 Jackson Jackson Gunnison Jackson Gunnison 124 163 186 102 97 Jackson 166c Gunnison 90 Lipids, % of live weight Live weight, gm M. pectoralis, gm N a 5.5 2.3 2.4 5.5 3.6 4.2 3,195 2,916 2,104 2,929 2,146 3,005 2,200 265 244 174 246 183 266 194 9 10 10 10 10 10 10 No males were collected in Gunnison County in 1983. b Data are from National Oceanic and Atmospheric Administration Monthly Climatological Data for Colorado, 1983-1986. c 4.1 Does not include snowfall accumulations in December 1985. �147 Table 17. Mean number of strutting displays/minute of adult male sage grouse in Jackson and Gunnison counties, April-May 1986. Mean display rates are presented for each 1ek-fema1e distance combination within a population. Female distance (m) is the closest distance that a female approached the male under observation. Standard errors and number of 5-minute observation periods are in parenthesis for each mean. Comparison of means between populations was by Hote11ings !2. Gunnison County Lek Jackson County Lek Ohio Ridge Creek road Deer Female S. distance Parlin Chance 4.93 5.22 5.40 5.73 6.04 6.97 (0.36) (0.80) (0.59) (0.57) (0.44) (0.15) (N = 24) (N = 9) (N = 14) (N = 18) 4.20 3.97 4.55 3.67 3.30 5.81 (0.40) (1.02) (0.94) (0.96) (1.19) (0.57) (N = 2) (N = 6) (N = 7) (N = 5) 0.85 1.73 1.42 1.69 2.43 2.31 (0.30) (0.25) (0.30) (0.30) (0.57) (0.48) (N = 20) (N = 42) (N = 39) (N = 18) <10 m 10-30 m >30 m (N = 7) (N = 4) (N = 15) Hote1lings !2 = 64.2, P 0.023 Railroad Creek (N = 20) (N = 9) (N = 25) �148 periodically checked for-displaying males during evening hours. Sage grouse in Jackson County were observed displaying on 7 of 7 occasions when evening display was evaluated on S leks. Numbers of males seen during evening display approached numbers observed on leks during morning hours. In Gunnison County, males were observed on only 2 of 9 occasions when evening courtship was evaluated at S leks. In each instance only 2 males were displaying on leks where between 36 and Sl males displayed during morning periods. Variance of mean peak daily attendance was compared among leks within populations via Bartlett's test for homogeneity (Zar 1984:181). among leks was similar (R > O.OS) within populations (Table 18). Variance Pooled variances (~2~) were calculated for each population and comparison among populations made using an F test. at Gunnison Basin leks (! = There was more variation in male attendance 9.44, P<0.02S). Lek Surveys Males and female attendance was evaluated at 19 leks. Although females were observed on leks throughout the breeding season, most mating appeared to occur during the second week of April (Table 19). This is consistent with observations on leks where intensive surveys of male display activities were conducted. (Table 20). Mean, peak male attendance was down slightly from previous years This was primarily due to lower counts at South Parlin, Chance Gulch, and Mashburn 1 leks. Because the validity of lek counts as a population index has not been demonstrated, the slight decline in lek attendance should not be interpreted as a population decline. �149 Table 18. Mean peak daily attendance of male and female sage grouse at 3 leks in Gunnison and Jackson counties where courtship display behavior was evaluated, April-May 1986. County Surveys were made during 5 mornings on each 1ek. Females Males 2 x s2 Range x Ohio Creek 17.8 20.7 11-23 1.8 9.2 0-7 Chance 20.6 29.8 12-26 2.8 13.0 0-9 South Parlin 19.8 13.7 14-24 2.4 6.8 0-6 Lek a RanEie Gunnison Sp2 21.4 Jackson Railroad 27.0 0.5 26-28 7.6 46.3 2-19 Ridge Road 17.0 2.0 15-19 7.0 50.0 0-16 Deer Creek 19.6 4.3 16-21 4.8 26.7 0-13 Sp2 2.27 DISCUSSION Winter Distribution Sage grouse were widely distributed throughout the Gunnison Basin during the mild 1985 and 1986 winters. Sage grouse were also observed throughout the Basin during the severe 1984 winter, although an extensive area south of Gunnison appeared to be used more heavily than other regions. data in 1984 may be biased due to lack of random sampling. Distribution There is no evidence that sage grouse in the Gunnison Basin primarily winter in a single or small number of geographic locations. The rugged terrain of the Gunnison �150 Table 19. Number of individuals and dates of peak sage grouse attendance at Gunnison Basin leks, 1986. Males Lek Gold Basin N Females Date 0 N Date 0 36 11 Apr 9 09 Apr 4 17 Apr 1 12 Apr Needle Creek 11 16 Apr 0 South Parlin 24 09 Apr 9 04 Apr North Parlin 20 17 Apr 11 02 May Allen 55 22 Apr 6 22 Apr Ohio Creek 23 05 May 7 11 Apr Mashburn 1 24 06 Apr 19 27 Mar Mashburn 2 17 22 Apr 4 22 Apr Sapinero 1 25 02 May 1 02 May Sapinero 2 0 Sapinero 3 24a 07 May 4 28 Apr Razor Creek 57 30 Apr 18 11 Apr Antelope 51 27 Apr 4 10 Apr Signal Peak 4 13 Apr 1 13 Apr Upper Six-mile 3 19 Apr 0 Kezar Basin 15b 12 May 0 Sugar Creek 5 08 May 0 Strattman 12 06 Apr 4 Monson Gulch 17 07 May 0 5 13 Apr 0 Chance Gulch Woods Gulch Lost Canyon 0 aForty-nine sage grouse of unknown sex flushed from lek on 14 May. bFifty sage grouse of unknown sex flushed from 1ek on 06 May. 06 Apr �151 _-:;. •.. Trends in peak 1ek attendance of male sage grouse in the Gunnison Basin 1980-86. Table 20. Peak male attendance 1980 Lek Gold Basin 0 1981 NCa 1982 1983 1984 1985 1986 0 NC 7 0 0 15 28 21 38 4 53 36 7 10 15 8 5 2 4 Needle Creek 36 27 18 14 9 9 11 Parlinb 43 72 47 88 66 94 44 Allen 46 16 27 51 37 33 55 112 53 38 82 18 32 23 68 60 NC NC 17 41 49 43 48 39 NC 19 12 57 Antelope 0 63 52 62 53 46 51 Blue Mesa 0 2 NC NC NC NC NC 7 10 NC NC 4 5 NC 0 NC Signal Peak 11 3 4 4 Lost Canyon 5 NC 12 5 Mashburn 1 22 54 24 Mashburn 2 43 25 17 Kezar Basin 12 15 Sugar Creek 5 5 Chance Gulch Woods Gulch Ohio Creek Sapinero c Razor Creek Upper Six-mile McCabe Strattmann 12 Monson 17 ~ 33.6 aNC = 37.9 26.4 No count. blnc1udes North, South, and Upper South Parlin leks. clnc1udes Sapinero 1, 2, and 3 leks. 34.0 23.3 25.5 22.7 �152 Basin likely contributed~to widespread winter distribution. Snow depths and sagebrush structure vary among topographic features (Fig. 6). Interspersion of diverse terrains insures that even in severe winters small areas of sagebrush habitat are available in most regions of the Gunnison Basin. Habitat managers cannot concentrate winter habitat management efforts in one or a small number of major sage grouse winter areas in the Gunnison Basin. Sage grouse are widely distributed throughout the region during winter months and impacts of proposed sagebrush treatments on winter habitat should be considered regardless of their location within the Gunnison Basin. Winter Topographic Associations Topographic distribution of sage grouse feeding activity was affected by shrub structure and snow depth. Drainages were heavily used in both winters even though these sites comprised a small percentage of the Gunnison Basin. Big sagebrush plants in drainages were vigorous and taller than plants in other terrain categories. Exposed sagebrush was available in drainages even though snow was deeper than in some other c&~egori2s (Fig. 6). During the severe 1984 winter much of the exposed sagebrush used by sage grouse during January and February occurred in drainages. Treatment of sagebrush stands in drainages would have a disproportionately severe impact on availability of sage grouse winter habitat. To insure availability of winter habitat, there should be no removal of sagebrush in drainages. Although a high proportion of sage grouse feeding occurred on 6-150 southwest slopes, use of this terrain was not disproportionate to availability. While use was not disproportionately high, the large percentage of feeding sites that occurred on southwest slopes suggests that these sites provide important winter habitat. Although sagebrush plants on these sites are short, snow depth is shallow and exposed plants are usually available. �60 ~ MEAN SAGEBRUSH HEIGHT AT RANDOM SITES _ 1985 FEBRUARY MEAN SNOW DEPTH _ 1986 FEBRUARY MEAN SNOW DEPTH 53 50 40 :E o 30 20 10 o DRAINAGE 0-5 LOW o >5 NE Fig. 6. Mean sagebrush height and February in the Gunnison Basin, Colorado. o ,0-5 HIGH o o >5 SW snow depths among terrain categories ~ V1 W �154 Southwest slopes >15 o were rarely used in 1985 (1% of feeding sites) but were more heavily used in 1986 (15% of feeding sites). 0 observations) use of >15 Most (7 of 12 southwest slopes in 1986 occurred during a 3-day period following a severe snow in mid-February. During the storm, wet snow became lodged in crowns of sagebrush plants in most terrains categories and made access to foliage difficult. Sagebrush plants on southwest slopes appeared to become snow-free more quickly than plants in other terrains and sage grouse may have temporarily exploited these sites because of more readily available forage. No sagebrush treatment should occur on slopes with southwest aspects due to the effects on availability of sage grouse winter habitat and because livestock forage response would likely be minimal due to xeric conditions. Use of 0-50 low sites was greater in 1985 than in 1986. reflected differences in snow depths between years. on survey plots was lower in 1986 (~ = This probably Mean January snow depth 17.5 cm) than in 1986 (x = 32.9 cm). Many (48%) observations in the 0-50 low category in 1985 occurred in January 0 prior to heavier mid-winter snow accumulations. Use of 0-5 low sites was reduced when snow depths exceeded approximately 30 cm in February 1985 and throughout winter 1986. 0 Sagebrush stands on 0-5 low sites consisted of slightly shorter plants that had a more open canopy than stands in drainages. Snow cover >30 cm covers a higher proportion of sagebrush structure on 0-5 low sites than at drainage sites. o Moderate or heavy snow depths reduced foliage availability for feeding, increased exposure of foraging birds, and made these areas less favorable than drainages. This was especially apparent in 1984 when little feeding activity occurred on 0-50 low sites. 0 Only 13% of feeding sites were observed on 0-5 high sites. Average height of big sagebrush plants at these sites was shorter than at the more mesic drainage sites and canopies were more open. Snow depths were greater �155 than on southwest aspects. big sagebrush on 0-5 o Black sagebrush was frequently interspersed with high sites. Because of its short height, black sagebrush was usually completely covered even by shallow snow depths of 20-30 cm. Small plant size, moderate snow depths, and greater exposure to wind may make 0-50 high sites less favorable as winter habitat. Treatment of 0-50 high sites would likely have less impact on sage grouse habitat than treatment in more heavily used terrain categories such as drainages. However, due to the xeric conditions on many of these sites, post-treatment forage response would be minimal, therefore sagebrush removal should not be attempted on most 0-50 high sites. I observed only 3 feeding sites on slopes with northeast aspects, even though these terrains comprised a large percentage of the Gunnison Basin. Snow depths were greater on northeast slopes than in most other terrains. Also, big sagebrush plants were smaller and canopy closure less than in drainages. Most sagebrush foliage on northeast slopes was covered as a result of deep snows and smaller plant size. Limited treatment of sagebrush stands on northeast slopes would likely have minimal impact on sage grouse winter habitat. Effects on other seasonal habitats should be considered. Shrub Structure Characteristics of Use Sites Differences in structural characteristics of big sagebrush plants were not apparent between feeding and random sites within drainages. Canopy cover, mean plant height, variation in mean plant height relative to the mean, and density were similar between samples. This indicates that structural measures of big sagebrush would be of little value in differentiating areas that are potentially suitable as sage grouse feeding sites within drainages. Differences in shrub structure were more apparent on xeric >5 and 0-50 high sites. o southwest Canopy cover and plant height were greater at feeding �156 sites than at random locations. Sites used by foraging sage grouse in these terrains were often sites that received greater moisture than surrounding areas and, as a result had more vigorous sagebrush. These sites included shallow depressions, areas beneath rock outcrops or snow cornices, and lower slopes. While shrub measurements are potentially useful to identify areas that may be favorable foraging sites within these terrains, they are probably not necessary. Because of the xeric nature of 0-50 high and >50 southwest sites, sagebrush plants are small and sagebrush control is not likely to occur on these sites. There is not sufficient fuel to sustain prescribed fires on these terrains and grass response following herbicide spraying would be minimal and would not justify the costs of treatment. Therefore, there is little management need to identify structurally suitable foraging sites on 0-50 high sites and >50 southwest slopes. Application of structural measurements to identify potentially suitable winter feeding sites in 0-50 low areas is also questionable. cover and density varied between feeding and random sites. Only canopy Mean height and variation in height relative to the mean were similar between random and feeding sites. Most (56%) measurements of feeding sites in this terrain were obtained in January 1985 when use was highest due to shallow snow cover. These measurements may not be similar to values that would be obtained at a sample of sites measured during periods of deeper snow. Sagebrush availability and sage grouse use of 0-50 low sites varies with snow depth. It would not be feasible to obtain an adequate description of structural characteristics of sage grouse feeding sites within the 0-5 o low terrain 0 that could consistently be applied to identify potential 0-5 sites among winters of differing severity. low feeding �157 Application of shrub structure criteria to identify areas of potential winter feeding activity is not necessary. These criteria are not adequate to identify winter habitat in some terrains (drainages and 0-50 low). In other terrains (>50 southwest, 0-50 high) distinctive structural characteristics of feeding sites exist, however the xeric nature of these terrains makes their treatment unlikely. Managers should rely more heavily on topographic criteria to evaluate potential impacts of proposed treatments on sage grouse winter habitat in the Gunnison Basin. Sage Grouse Forage Selection Although site conditions and big sagebrush shrub structure varied among topographic categories, there were no differences in big sagebrush crude protein and monoterpene content among random sites in the 4 terrains evaluated. Chemical composition was highly variable among sites within a single terrain. This indicates that topographic associations have little influence on sagebrush chemical content. Distribution of sage grouse feeding activity among topographic categories cannot be attributed to selection of terrains with favorable sagebrush chemical composition. Remington and Braun (1985) observed that sage grouse that foraged within stands of Wyoming big sagebrush (!. £. plants with high crude protein content. wyomingensis), selectively fed on Selection of high crude protein plants within winter feeding sites in the Gunnison Basin was not apparent. Protein levels of both browsed (~= 20.6%) and unbrowsed (~ = 20.2%) mountain big sagebrush plants in the Gunnison Basin were high relative to protein content in Wyoming (K = 14.1%) and mountain (~ = 10.8%) big sagebrush plants in Jackson County observed by Remington and Braun. Monoterpene levels also did not differ between browsed and unbrowsed big sagebrush plants in the Gunnison Basin. Total monoterpene content of mountain �158 big sagebrush at feeding sites in the Gunnison Basin was lower than monoterpene content observed within this subspecies in Jackson County, by Remington and Braun (1985). However, differences in gas chromatograph equipment and procedures make direct comparison difficult. Sage Grouse Lipid Reserves Following mild winters (1983, 1985) adult male sage grouse in Jackson County entered the courtship period with lipid reserves that comprised approximately 5.5% of live weight. In Gunnison County, mean lipid reserves of adult males were 4.1-4.2% of live weight following mild winters (1985, 1986). Lower lipid reserves of Gunnison County males may be due to the smaller physical structure of birds in this population previously reported by Hupp (1984). Small body size may result in greater energetic investment in thermoregulation during winter. Increased thermoregulatory costs could limit lipid deposition. Winter severity affected sagebrush exposure and availability and influenced early spring lipid reserves in both Jackson and Gunnison counties. In Gunnison County, exposed sagebrush was available in only 7% of the Basin during the severe winter of 1984. crowns were widely scattered. Where sagebrush was available, exposed Sage grouse access to forage resources was restricted and, within feeding sites, birds had to expend additional locomotion costs when traveling through deep, soft snow to reach scattered sagebrush crowns. Conditions were similar in Jackson County. persisted through April in each study region. Deep snow cover Reduced availability of forage resources and greater energetic investment in locomotion limited lipid deposition in each population in 1984. In 1986, heavy snows accumulated in Jackson County during November and December but dissipated during February and March. Early winter snow accumulations apparently limited lipid deposition in �159 this population. -~ Fat reserves of adult male sage grouse in Jackson County in 1986 were lower than reserves accumulated during mild winters but higher than reserves deposited during the severe 1984 winter. Comparison of Male Sage Grouse Courtship Behavior Adult male sage grouse mobilize lipid reserves during spring courtship (Hupp 1984). Lipid reserves may be necessary to meet energetic costs of spring courtship, and males may adjust courtship behavior dependent upon size of lipid reserves. Following the severe 1984 winter, lipid reserves in Jackson and Gunnison county male sage grouse were low. Males in each population were often observed passively sitting on leks without displaying. Numbers of males observed on Jackson County leks was lower in 1984 (x = 21.2 males/lek) than during the previous 5 years (K = 39.4-43.5 males/lek). Lek counts also declined in Gunnison County although not as severely as in Jackson County. Reduced lek attendance may have been due to severe winter mortality. Reduced display intensity may have been the result of persistent snows near leks that covered sagebrush foliage and reduced availability of exogenous sources of energy and nutrients. However an alternative hypothesis is that males reduced energetic investment in courtship due to low endogenous reserves. Males may have displayed more slowly and attended leks less frequently to conserve their lipid reserves. A relationship between endogenous reserves and courtship behavior may therefore exist. Further evidence to support this hypotheses was observed in 1985. Despite similarities in winter conditions, males in the Gunnison Basin entered the display period with lower lipid reserves than males in Jackson County. Behavior of males in Gunnison County appeared to be similar to that observed in 1984; males often sat passively on leks and did not display. In addition, �160 males in Gunnison County did not roost on leks at night, but instead were usually observed roosting in heavy sagebrush cover near strutting grounds. Jackson County males typically roost on the exposed display areas. Due to lower lipid reserves and smaller body size, males in Gunnison County may have roosted under sagebrush canopy to reduce night-time costs of thermoregulation. Comparisons of courtship display were made between populations in 1986 to identify differences in rates of display and whether those differences could be attributed to unequal lipid reserves. The strutting display is the predominant courtship behavior of males on leks. Faster rates of display in a population are likely indicative of greater energetic investment. Differences in display rates were observed. Strutting display rates of males in Gunnison County were slower than display rates of males in Jackson County. Also, males in Jackson County were observed displaying on leks during evening hours, while males in Gunnison county were not. Slower display rates and lack of evening display indicate that energetic investment in courtship behavior was lower among adult males in Gunnison County. Male attendance on Gunnison County leks was highly variable among mornings, while numbers of males on Jackson County leks changed little from one survey period to the next. Higher variance in attendance may indicate that Gunnison County males shift display activities among leks and are not as strongly affiliated with a single display area as Jackson County males. However, higher variance may be indicative of reduced energetic investment in display behavior among Gunnison County males. Some males in Gunnison County may not display during some mornings, or may display only for brief periods. Although slower display rates, lack of evening display, and higher variance in 1ek attendance suggest that male energetic investment in display is lower in Gunnison County, the differences in courtship behavior between �161 populations cannot be attributed to unequal lipid reserves. Due to severe early winter conditions in 1985-86, adult male sage grouse in Jackson County entered the display season with low lipid reserves. County males (~= Lipid reserves of Jackson 3.6% of live weight) were similar to reserves of adult males in Gunnison County (~= 4.1% of live weight). These results fail to support the hypothesis that male display behavior is affected by size of lipid reserves. However, tests of the relationship between lipid reserves and courtship behavior would be more conclusive through experimental manipulation of fat reserves in a sample of adult males, or comparison of display behavior within a single population among years when early spring lipid reserves vary. LITERATURE CITED Byers, C. R., R. K. Steinhorst, and P. R. Krausman. 1984. Clarification of a technique for analysis of utilization-availability data. J. Wildl. Manage. 48:1050-1053. Conover, W. J. 1980. Practical nonparametric statistics. Sons, New York, N.Y. John Wiley and 493 pp. Epstein, W. M., L. R. McGee, C. D. Poulter, and L. L. Marsh. 1976. Mass spectral data for gas chromatograph - mass spectral identification of some irregular monoterpenes. J. Chem. and Eng. Data 21:500-502. Heller, S. R., and G. W. A. Milne. base. U.S. Dep. Comm., Washington, D.C. Horwitz, W. Chem. Editor. 1980. Washington, D.C. Colorado. 1984. Basin. 107-128. 1978. EPA/EIH mass spectral data 988 pp. Official methods of analysis. Assoc. Off. Anal. 1018 pp. Hunter, W. R., and C. F. Spears. Hupp, J. Editors. 1975. Soil survey of the Gunnison area, U.S. Govt. Print. Off., Washington, D.C. 85 pp. Sage grouse distribution and habitat use in the Gunnison Job Prog. Rep., Colorado Div. Wildl. Res. Rep. April 1984. Pp. �162 Norusis, M. J. 1986. SPSS/PC+ advanced statistics. SPSS Inc., Chicago, Ill. 230 pp. Remington, T. E., and C. E. Braun. winter, North Park, Colorado. Zar, J. H. 1984. Cliffs, N. J. Prepared BYJer~ 1985. Sage grouse food selection in J. Wildl. Manage. 49:1055-1061. Biostatistical analysis. 718 pp. &= Graduate Research Assistant Approved By ~--:~----..,~~=~_Z_'-'--~ _ Clait E. Braun Wildlife Research Leader 2nd ed. Prentice-Hall, Englewood �163 Colorado Division of Wildlife Wildlife Research Report April 1~87 JOB PROGRESS REPORT State of Colorado Project 01-03-045 (W-37-R) 3 Work Plan Job Title: 17 ------ Response of Selected Avifauna to Prescribed Burning in the Big Sagebrush Type Period Covered: Author: : Job Avian Resea~ch 01 July through 31 December 1986 Ernesto J. Hernandez Personnel: C. Braun, Colorado Division of Wildlife; E. Hernandez, Colorado State University ABSTRACT Preparations for this study were initiated in July 1986 with advertisement for and selection of a graduate research assistant. Consultations were held with personnel of the Bureau of Land Management and delay in burning until 1987 was requested. Treatment (Deer Creek) and control (Railroad) leks were selected. Review of literature was initiated and study design was discussed with D. C. Bowden and personnel of the Colorado Division of Wildlife and the Bureau of Land Management. Despite effects on study design, a portion of the treatment area east-southeast of the Deer Creek Lek was burned in September-October 1986. ��165 RESPONSE OF SELECTED AVIFAUNA TO PRESCRIBED BURNING IN THE BIG SAGEBRUSH TYPE Ernesto J. Hernandez P. N. OBJECTIVES The primary objectives of this study are to: (1) describe pre- and posttreatment (burning) patterns of seasonal habitat use by sage grouse in North Park, Colorado; (2) describe pre- and post-treatment sagebrush structure and vegetation composition at observed sage grouse use sites; (3) compare pre- and post-treatment frequency of strutting displays by male sage grouse; and (4) compare relative abundance, species composition, and species richness of passerine birds of the big sagebrush type on burned vs. unburned areas. SEGMENT OBJECTIVES 1. Review available literature concerning sage grouse and the effects of prescribed burning in sagebrush rangelands. 2. Prepare a research proposal. 3. Select treatment and control sites in North Park, Colorado. STUDY AREA Selection of study areas in North Park, Colorado (Fig. 1) was done in consultation with Clait E. Braun, Colorado Division of Wildlife. A description of these areas will be included in the final report. Ernes Graduate Research Assistant Approved by t&Lr &awu) Clait E. Braun ' Wildlife Research Leader �166 .• TR.EA"TMEN1 • CoNTRol-- 1-~~ LE 'K.. _./ -: ./ /' 4i. I) Col.i-i NoS ~N\t~ Q.O '-0 RA vO Fig. 1. Study areas selected for sagebrush burning Park, Jackson county, Colorado. treatments in North �~u~urauu ULVLbLUll UL ..LVI WLLULLLe Wildlife Research Report April 198] JOB PROGRESS REPORT State of Colorado --~~~~~------------------------- Project 01-03-045 (W-37-R) 8 Work Plan Job Title: 5 --- Population Inventory and Habitat Use by Lesser Prairie-chickens in Southeast Colorado Period Covered: Author: : Job Avian Research 01 January through 31 December 1986 Kenneth M. Giesen Personnel: Beth Dillon, Ken Giesen, Gwen Kittel, Jennie Slater, Chuck Wagner, Bryant Will, Colorado Division of Wildlife ABSTRACT Approximately 41.4 km2 of rangeland encompassing pasture 1AE of the Comanche National Grasslands was selected as the study site for lesser prairie-chicken (Tympanuchus pallidicinctus) habitat use and home range information. Eighty random vegetation transects were measured on this area to document residual vegetative structure and species composition. Surveys in southeast Colorado indicated that 39 of 48 lesser prairie-chicken lek sites were active in 1986 with an average of 8.4 birds/lek in Baca County. Thirty-two lesser prairiechickens were trapped on leks and banded with 22 being fitted with miniature radio transmitters. Only 1 of 13 radio-marked hens was known to be successful in hatching a clutch of eggs. Nests contained an average clutch of 11.2 eggs and were at sites having greater than average vegetative heights. The median spring-autumn home range of 11 grouse was 217 ha with females having larger home ranges than males. Preliminary analysis of microhabitat suggests no structural differences between random sites and sites used by lesser prairiechickens •. ��169 POPULATION INVENTORY AND HABITAT USE BY LESSER PRAIRIE-CHICKENS IN SOUTHEAST COLORADO Kenneth M. Giesen Both distribution and populations of lesser prairie-chickens in North America have decreased by >90% from historic levels of the 1800's (Taylor and Guthery 1980). Although the exact historic distribution of lesser prairie-chickens is unknown, early reports (Bendire 1892, Judd 1905, Bent 1932, Baker 1953, Sands 1978) suggested they were abundant and widely distributed throughout their range. Although Aldrich (1963) indicated lesser prairie-chickens historically inhabited about 360,000 km2 in 5 states, recent estimates suggest a current population of 50,000 birds existing on 125,000 km2 (Crawford 1980, Taylor and Guthery 1980, Johnsgard 1983). Although evidence suggests lesser prairie-chickens were historically peripheral in Colorado, they were thought to be common to abundant in 6 southeastern counties (Baca, Prowers, Bent, Kiowa, Lincoln, and Cheyenne), and peripheral in adjacent counties (Loeffler 1983). Recent surveys have documented breeding populations in Baca, Prowers, and Kiowa counties (Hoffman 1963, Loeffler 1983, Rash 1985). The lesser prairie-chicken is currently classified as a threatened species in Colorado with an estimated population of <800 birds. P. N. OBJECTIVES The objectives of this study are to evaluate lek surveys as indices to population trends, ascertain the accuracy of aerial and ground surveys in detecting leks, describe the seasonal floristic and structural characteristics of lesser prairie-chicken habitats in southeast Colorado, and contribute to preparation of a recovery plan for lesser prairie-chickens in Colorado. Segment Objectives 1. Review pertinent literature applicable to the objectives of this study. 2. Complete study plan. 3. Select a primary study are of 16 sections (41.4 km2) for telemetry studies, based on lesser prairie-chicken densities, distribution of leks, land use, and habitat type. 4a. Locate all active leks within the primary study area and obtain at least 1 count/week (Mar-May) of all males and females on each lek. 4b. Survey all historic leks in Baca County and obtain at least 1 count of males on each active lek. 5. Trap and band lesser prairie-chickens on active leks within the primary study area. At least 20 will be marked with miniature radio transmitters to facilitate their periodic location. �170 6. Locate lesser prairie~chicken nests by following radio-marked hens. Record clutch size, incubation period, and nest fate. 7. Locate all radio-marked birds weekly for estimates of movement and home range. 8. Measure vegetative cover at grouse use sites and random sites. canopy-cover of shrubs, forbs, and grasses will be recorded. 9. Compile data, analyze results, and prepare annual progress report. Height and METHODS Field surveys were conducted on the Comanche National Grasslands and adjacent areas in Baca county from March through June using binoculars, a parabolic microphone listening device, and a trained pointing dog to locate active lesser prairie-chicken leks. Active leks were visited within 2 hours of sunrise to count grouse and classify them to sex. Cannon-nets and walk-in funnel traps (Giesen et al. 1982) were used to capture grouse on leks. Each captured grouse was marked with a numbered aluminum band and a unique combination of colored plastic bandettes. Miniature solar- or batterypowered transmitters (weight 18-24 gms) were attached to all captured females and selected males using a poncho (Amstrup 1980). Radio-marked birds were located using a portable receiver and a hand-held yagi antenna. Vegetative structure and species composition were measured using line intercept of canopy cover (Canfield 1941) and a range pole (Robel et al. 1970). The minimum convex polygon method (Mohr 1947) was used to calculate home ranges. Description of Study Area Approximately 41.4 km2 in and adjacent to pasture lAE of the Comanche National Grasslands was selected as the primary study area for habitat use and home range information. Included are sections 21-28, and 33-36, T34S, R44W, and sections 1-4, T35S, R44w. Approximately 90% of the area is rangeland with the remainder being dryland cropland. A series of 80 random vegetative transects was measured from March through May to quantify residual vegetation on the area. Preliminary analysis indicates a mean density of 3235 sandsage (Artemisia filifolia) plants/ha. The mean heights of shrubs, forbs, bunch grasses and short grasses were 35.2, 4.4, 16.3, and 5.9 cm, respectively. Bare ground comprised 73.3% of the canopy cover. A complete vegetative description of the area will be included in later progress reports after additional sampling and analysis of results. RESULTS AND DISCUSSION Lek Counts A total of 304 counts was obtained from 39 active leks in southeast Colorado between 5 March and 27 June 1986. A total of 292 counts of 31 active leks was obtained for Baca county on or near the study area; 9 additional historic leks were surveyed and found to be inactive. A minimum of 197 males, 22 females, and 261 total birds was counted (Table 1) resulting in an average of 8.4 �171 birds/1ek. Summaries of 1ek counts since 1977 indicate the dynamic nature of 1ek sites with many leks becoming extinct and additional leks becoming established (Table 2). Although efforts to survey and count leks were not constant among years, it appears that lesser prairie-chicken populations are relatively stable. Table 1. Lesser prairie-chicken 1ek count data, Baca County, 1986. N Lek 2 3 4 5 6 7 12 14 17 18 23 25 28 30 31 counts 47 11 12 46 9 7 3 5 4 8 2 3 21 2 5 35 36 37 38 39 2 3 41 42 1 43 2 46 1 Total Average 17 18 17 15 1 3 8 5 7 86-1 86-2 86-3 OK-1 OK-2 05 05 05 07 06 08 21 09 15 07 17 15 08 1 33 40 Count period 32 17 10 7 7 292 12 08 09 07 31 17 11 11 10 08 Mar Mar Mar Mar Mar Mar Mar Mar Mar Mar Apr Mar Mar 17 Apr Mar Apr Mar 17 Mar Mar Mar Mar 01 Mar 17 Mar Apr Apr Mar Mar - 27 Jun - 23 May - 23 May 27 Jun - 21 May - 20 May - 09 May - 09 May - 01 May - 21 May - 01 Jun - 01 May - 18 Jun Apr - 01 Jun - 21 May - 01 Jun - 02 Apr Apr - 06 May - 21 May - 09 May - 13 May Jun - 09 May Apr - 27 Jun - 27 Jun - 27 Jun - 20 May - 20 Hay Males 16 2 o 17 6 9 5 High count Females Totals 5 19 o 3 1 o 5 1 19 8 o 10 o 10 4 o 8 10 2 14 2 2 o o 2 4 1 14 o 1 8 7 10 12 o 2 o 10 o 12 2 11 8 3 o 5 1 15 6 7 9 11 197 7.6 o o o 1 o o o o 1 o o o o 4 1 1 o o 22 2.2 5 13 1 14 2 11 8 3 10 5 1 19 8 8 10 15 261 8.4 �172 Table 2. Lek 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 86-1 86-2 86-3 High counts of male lesser prairie-chicken, Baca County, 1977-86. 1977 4 18 8 9 24 22 15 1 0 0 7 0 7 17 17 16 2 3 3 5 __b 1978 1979 1980 5 14 6 8 NC 15 11 NC NC NC NC 5 13 11 11 0 4 14 NC 0 4 13 16 6 NCa 10 2 7 12 15 11 0 0 0 NC 7 12 10 12 NC 3 NC NC 2 1 6 17 NC 0 1 17 12 15 14 11 0 0 0 0 17 8 9 9 0 0 9 0 0 0 4 19 0 3 9 7 7 aNo count. bLek not yet located. Year 1981 1982 0 6 15 7 16 14 6 0 0 0 0 20 8 8 0 0 0 9 0 0 0 3 30 0 6 4 4 7 4 18 14 6 8 6 0 3 12 7 29 19 9 0 0 0 0 14 7 6 0 0 0 10 0 0 0 0 33 0 11 2 4 4 0 10 17 0 6 0 10 12 1983 1984 1985 1986 0 12 11 9 26 14 8 0 0 0 0 16 5 12 0 0 0 9 0 0 0 0 23 0 3 2 3 4 0 0 13 0 0 0 16 7 12 7 10 4 0 9 7 7 23 17 7 0 0 0 0 18 0 11 .. 0 0 2 5 0 0 0 0 21 0 2 3 0 0 0 6 11 0 0 0 21 0 19 8 18 5 3 14 0 7 7 0 18 6 5 0 0 0 0 0 0 11 0 0 5 9 0 0 0 0 16 0 '0 0 4 10 2 2 7 5 0 0 13 0 12 3 9 7 7 7 7 3 4 4 NC 16 2 0 17 6 9 NC NC NC NC 5 0 10 NC NC 4 10 NC NC 0 NC 2 NC 2 0 0 12 0 0 2 0 0 0 10 0 12 2 11 8 3 0 5 0 0 1 15 6 7 �173 Analysis 4l.4-km2 breeding breeding of lek surveys -and lek count data (1980-86, Table 3) on the study area indicates a strong positive correlation between male density and number of active leks (r = 0.90) but not between male density and average lek size (r = 6:21). Table 3. Lek count trends of lesser prairie-chickens on a 4l.4-km2 (16 2 mi ) study area, Baca County, Colorado, 1980-86. Year N leks N males x males/lek Breeding density (males/km2) 1980 1981 1982 1983 1984 1985 1986 6 6 6 6 4 5 7 59 55 59 65 46 46 75 9.8 9.2 9.8 10.8 ll.5 9.2 10.7 1.42 1.33 1.42 1.57 1.11 1.11 1.81 A mlnlmum of 76 females visitations to leks was recorded between 20 March and 21 May. Most (N = 65, 85.5%) occurred between 26 March and 11 April which likely represented the peak of mating in Baca county. Trapping and Banding A total of 32 lesser prairie-chickens (19 males, 13 females) was trapped on 5 of 7 leks on the primary study area. Design problems with the walk-in funnel traps were not corrected until after the peak of hen attendance which resulted in relatively few females being captured. Radio transmitters were placed on 13 female and 9 male prair~e-chickens to facilitate their future location. At least 10 radio-marked birds (5 males, 5 females) were killed by predators within 60 days of being trapped and another 5 weren't located after 1 June because of radio failure, mortaility, or possible long range dispersal from the study area. Yearlings comprised 9 of 19 males (47.4%) and 7 of 13 females (53.0%) captured. Adult males weighted the most (x = 768 gms) with weights of yearling males, adult females, and yearling females being similar, 716, 722, and 719 gms, respectively. Nesting Of 13 radio-marked hens, 5 were killed by predators prior to or during egg laying (nests were not located), 3 hens had malfunctioning transmitters or could not be located after 3 weeks, 1 hen abandoned her nest, 3 nests were lost to predation during incubation, and 1 hen successfully hatched her clutch. Four complete clutches examined contained 10, 10, 11, and 14 eggs, respectively. �174 Distance to nest from le~ of banding averaged 2.5 km (N = 6, range 0.9-4.8 km). During egg laying and incubation females generally restricted their daily movements within a 500-m radius of the nest. Nests were generally placed under a sandsage shrub or bunchgrass (sand dropseed or sideoats grama) which was taller (average height = 48.5 cm) than surrounding vegetation (33.3 cm). Canopy cover of grasses within 5 m of nests averaged 12.7%. Home Range Home range size was measured for 11 lesser prairie-chickens (5 males, 6 females) which were observed at least 6 times after capture. Home range varied among individuals (range 27-3787 ha, median = 217 ha) with females tending to have larger home ranges than males (Table 4). The largest home range was recorded for an adult female which continued to move long distances all summer after her nest was depredated. Table 4. Band If 301 306 308 315 317 318 319 320 323 324 327 Home range of lesser prairie-chickens, Baca County, Colorado, 1986. Age Sex 2+ 2+ 2+ 1+ 2+ 2+ 2+ 2+ 2+ 1+ 1+ M M M F F M F F F M F Time interval Home range (ha) - 27 112 217 278 3784 247 794 89 104 117 229 21 17 17 17 17 18 25 17 17 20 26 Apr Apr Apr Apr Apr Apr Apr Apr Apr Apr Apr 14 25 06 04 08 07 08 02 27 07 03 May Jun Oct Sep Oct Oct Oct Jun May Oct Sep Habitat Use Microhabitat characteristics were measured at 39 grouse-use sites and 39 random sites within 400 m of the grouse flush site. Detailed analysis has not been completed but there appears to be no selection for sandsage density or for height of shrubs, forbs, or grasses. Comparative analysis of species composition and relative frequency has not been completed. LITERATURE CITED Aldrich, J. W. 1963. Geographic orientation of American Tetraonidae. Wildl. Manage. 27:529-545. Amstrup, S. C. 214-217. 1980. A radio-collar for game birds. J. J. Wildl. Manage. 44: �175 Baker, M. F. 1953. Prairie chickens of Kansas. Misc. Publ. 5. 68 pp. Univ. Kansas Mus. Nat. Hist. Bendire, C. E. 1892. Life histories of North American birds with special reference to their breeding habits and eggs. U.S. Natl. Mus. Spec. Bull. 1. 446 pp , Bent, A. C. 1932. Life histories of North American gallinaceous birds. Natl. Mus. Bull. 162. 490 pp. U.S. Canfield, R. H. 1941. Application of the line intercept method in sampling range vegetation. J. For. 39:388-394. Crawford, J. A. 1980. Status, problems, and research needs of the lesser pra1r1e chicken. Pp. 1-7 in P. A. Vohs and F. L. Knopf, eds. Proc. Prairie Grouse Symp. Oklahoma State Univ., Stillwater. Giesen, K. M., T. J. Schoenberg, and C. E. Braun. 1982. Methods for trapping sage grouse in Colorado. Wildl. Soc. Bull. 10:224-231. Hoffman, D. M. 1963. The lesser prairie chicken in Colorado. Manage. 27:726-732. Johnsgard, P. A. 1983. Lincoln. 413 pp. The grouse of the world. J. Wildl. Univ. Nebraska Press, Judd, S. D. 1905. The grouse and wild turkeys of the United States and their economic values. U.S. Dep. Agric. BioI. Surv. Bull. 24. 55 pp. Loeffler, C. 1983. The status and management of the lesser prairie chicken in Colorado. Unpubl. Rep., Colorado Div. Wildl., Colorado Springs. 9 pp. Mohr, C. o. 1947. Table of equivalent populations of North American small mammals. Am. MidI. Nat. 37:223-249. Rash, M. T. 1985. Survey of the lesser prairie-chicken in Colorado, 3 April 25 May 1985. Unpubl. Rep., Colorado Div. Wildl. Colorado Springs. 22 pp. Robel, R. J., J. N. Briggs, J. J. Cebula, A. D. Dayton, and L. C. Hulbert. 1970. Relationships between visual obstruction measurements and weight of grassland vegetation. J. Range. Manage. 23:295-297. Sands, J. L. 1978. Game bird studies. Performance Rep., Proj. W-l04-R-19. New Mexico Dep. Game and Fish Proj. Albuquerque. 5 pp. Taylor, M. A., and F. S. Guthery. 1980. Fall-winter movements, ranges and habitat use of lesser prairie chickens. J. Wildl. Manage. 44:521-524. Prepared by _J..:.!dJ::=...r;::..:.=:.=....;_...;..!n___;_::. ~====. :.:...;._ Kenneth M. Giesen Wildlife Researcher _ ��177 Colorado Division of Wildlife -::-- JOB PROGRESS REPORT State of Colorado Project 01-03-045 (W-37-R) Work Plan Job 9 Job Title: 8 --- Food Selection and Nutritional Ecology of Blue Grouse During Winter Period Covered: Author: Avian Research 01 January through 31 December 1986 T. E. Remington Personnel: C. E. Braun, R. W. Hoffman, T. E. Remington, Colorado Division of Wildlife ABSTRACT Trials were conducted with captive blue grouse (Dendragapus obscurus) to measure metabolizable energy (ME) and nitrogen (MN) of needles of palatable and unpalatable conifers. The effect of monoterpenes and benzoic acid on palatability, ME, and nitrogen balance was measured. Total nitrogen, cell wall-bound nitrogen, and fiber were assayed as possible correlates of ME and MN. Fat stores of wintering blue grouse were measured. ME content of Douglas-fir (Pseudotsuga menziesii) and lodgepole pine (Pinus contorta) did not differ (p > 0.05) but exceeded (p < 0.05) ME of subalpine fir (Abies lasiocarpa) and Engelmann spruce (pIcea engelmanni) (1.75 vs. 1.53 and 1.15 Kcals/g, respectively). Palatability of preferred species (Douglas-fir and lodgepole pine) exceeded (P <0.05) that of the 2 non-preferred species (80 vs. 56 g/day). Birds were in positive nitrogen balance only on Douglas-fir. Monoterpenes added to Douglas-fir needles decreased their palatability (p< 0.05), increased weight loss (p < 0:05), and affected digestibility (p < 0:-05). Digestibility increased when 0:-565 g were added to 100 g of needles but declined sharply (P. < 0.05) at twice this level. Addition of benzoic acid to Douglas-fir needles (0.28% DM) had no effect (p> 0.05) on ME, palatability, or nitrogen balance. Palatability did not differ-(P> 0.05) between tree ages (old vs. young) or among needle ages (1-2, 3-4, 5-7) of Douglas-fir. Differences between tree ages and needle ages were complicated by interactions, but 1 and 2-year-old needles of old Douglas-fir were clearly superior in ME, digestibility, and nitrogen balance to other tree and needle ages. Cell wali-bound nitrogen comprised from 20 to 40% of total nitrogen; preferred needle groups generally had less nitrogen bound. Neutral detergent fiber (NDF) digestibility varied from 10 to 26% among species. NDF digestibility was a function of how much NDF entered the caeca (13 to 29%) and the efficiency of fermentation within the caeca (74 to 91%). Average fat content of adult and yearling male and female blue grouse was 2.01, 1.49, l.85, and L 77% of live weight, respectively. ��179 FOOD SELECTION AND NUTRITIONAL ECOLOGY OF BLUE GROUSE DURING WINTER Thomas E. Remington Trials were conducted with captive blue grouse to measure metabolizable energy content of (1) needles of palatable and unpalatable conifer species; (2) 1-2, 3-4, and 5-6 year-old needles of young and old Douglas-fir; and (3) needles of Douglas-fir treated with benzoic acid or monoterpenes. Work continues on nutritional analysis of the 375 samples of excreta (fecal and cecal) and conifer needles generated by these trials. This report summarizes progress to date; statistical analyses may be incomplete and/or preliminary. In addition, 4 blue grouse were captured and needles of Engelmann spruce collected for a trial to be conducted during January 1987 to investigate the digestibilityreducing effects of tannins. P. N. OBJECTIVES The objectives of this study are to investigate (1) winter food habits, and (2) winter food preferences of blue grouse, and (3) measure the nutritional quality (protein, fat, minerals) and anti-quality components (tannins, terpenoids, phenolic resins, fiber) of blue grouse winter foods and their relationship to diet preferences. Specific objectives are to: 1. Identify winter foods of blue grouse. 2. Investigate blue grouse/winter use of conifers for food by species, tree ages, and growth forms in relation to their availability. 3. Quantify tree species composition and physical characteristics of blue grouse winter feeding sites. 4. Measure protein and fiber content of conifer needles from trees fed-upon by blue grouse and from randomly-located trees. 5. Measure tannin, phenolic resin, and mono-, di- and sesquiterpene levels in conifer needles from trees fed-upon by blue grouse and from randomlylocated trees. 6. Rank blue grouse preference for Douglas-fir, subalpine fir, lodgepole pine, limber pine, and Engelmann spruce as winter foods. 7. Measure protein, fiber, and total digestibility by blue grouse of randomly-· selected needles of Douglas-fir, lodgepole pine, limber pine, subalpine fir, and Engelmann spruce. 8. Investigate the possible deterrent effects of specific tannins, phenolic resins, and mono-, di- and sesquiterpenes to browsing by blue grouse and to blue grouse digestibility of conifer needles. The objective of these trials was to test the hypothesis that (1) foods avoided by blue grouse contain less metabolizable energy (ME) or metabolizable nitrogen than preferred foods, and (2) benzoic acid or monoterpenes reduce ME �180 when added to palatable needles. Preferred vs. non-preferred comparisons were made between conifer species (trial 1), and between and among different tree and needles ages within Douglas-fir. Differences in preference were es~ablished previously with captive birds (Remington 1986). SEGMENT OBJECTIVES 1. Collect and process (clip 1-4 year-old segments, then freeze, crush, and filter to isolate needles) 9 kg of Douglas-fir needles. 2. Capture 4 blue grouse with telescoping noose poles or by darting. 3. Conduct trials to evaluate the effect of added tannins on the metabolizable energy (ME) and nitrogen content of conifer needles. 4. Prepare needles and droppings for nutritional analyses by freeze drying and grinding in a Wiley Mill. 5. Measure nitrogen, uric acid, urea, gross energy, and dry matter content of blue grouse droppings collected during ME trials. Measure nitrogen, gross energy, and dry matter content of food samples used in ME trials. 6. Compile and analyze data and prepare progress reports. METHODS The design, principle, and methodologies for all 4 trials were essentially the same. A randomized block design with 4 birds and 3 replicates per cell was used. Variables measured were dry matter intake, metabolizable energy, nitrogen balance, and weight change. The general principle was that metabolizable energy (and nitrogen) content of food items can be determined by subtracting energy excreted from energy ingested and dividing by the grams of dry matter ingested to correct for variations in intake. Endogenous energy losses in urine must be determined and subtracted from the energy excreted. Treatments consisted of needles from 4 different species of conifers (Douglas-fir, lodgepole pine, subalpine fir, and Engelmann spruce in trial 1 and 1-2, 3-4, and 5-6 year old needles from both old (>75 years) and young Douglas-fir in trial 4. Treatments in the monoterpenes - added trial (trial 2) consisted of 3 levels of monoterpenes added to 1-4 year-old Douglas-fir needles. Levell was a control, no monoterpenes added; level 2 was 2 times the control level (or double the average level in fed-upon Douglas-fir, 0.565 g added per 100 g wet weight of needles); and level 3 was 4 times the control level (1.690 g added per 100 g wet weight of needles). Monoterpenes were dripped into 100 g subsamples of needles which were then mixed to insure thorough distribution of monoterpenes over the needles. The monoterpene mixture consisted of camphene (41.0%), bornyl acetate (25.8%), a-pinene (19.7%), and limomene (13.6%). The proportions mimic those in fed-upon Douglas-fir (Remington 1986). These 4 terpenes together constitute 74, 78, and 71% of the terpenes found in fed-upon Douglas-fir, young Douglas-fir, and subalpine fir, respectively. �181 Treatments in th~ benzoi~ acid-added trial (trial 3) consisted of a control (no benzoic acid) and twice the level found in Engelmann spruce. Benzoic acid was dissolved in alcohol (3.673 g in 50 ml) and 2 ml of this solution was dripped over 100 g subsamples of needles. Two mls of alcohol were dripped over 100 g subsamples of control needles. Treatment replicates were randomly ordered and birds randomly assigned to treatment arrays. Recovery periods, where the birds were given Douglas-fir needles plus approximately 10 g of Purina game bird chow (12.5% protein, 2.5% fat, 10% fiber), were used whenever birds lost more than 5 g to prevent carryover effects encountered in 1985. Four birds, 2 males and 2 females, were used for all trials. A mixture of all 4 species tested in trial 1 was fed to the birds (supplemented with 10 g of game bird chow) for 2 weeks prior to the start of the trial. Needles were collected from mid-canopy of randomly located mature trees. One to 4-year-old segments were clipped from branches and frozen in a super-cold (-70 C) freezer. Needles were separated from the stems by agitating the frozen segments and filtering with different sizes of wire mesh screens. Birds were weighed on a portable analytical balance (±l g) each morning and droppings cleared prior to presentation of needles. Needles (initially 220 ± 1 g) were placed in 250 ml beakers and set in holders at the front of each cage. The beakers were removed about half an hour after sunset after birds had ceased feeding. Droppings were collected at this time and the following morning (24 hrs after needle presentation), placed in plastic bags and immediately frozen. Spillage and droppings were collected on trays under each cage. Consumption was calculated (±0.01 g) by subtraction. Food and excreta samples were freeze dried and ground to a fine powder in a Wiley mill. Gross energy was measured by burning 0.5000 g samples in a Parr adiabatic bomb calorimeter. Nitrogen was measured by micro-Kjeldahl analysis (Horwitz 1980) on 0.2500 g samples. Uric acid was measured spectrophotometrically using a method developed for poultry excreta (Marquardt 1983). Urea was measured colorimetrically on 1.0000 g samples (Horwitz 1980). Cell wall-bound nitrogen was estimated by measuring pepsin-insoluble nitrogen in neutral detergent fiber (NDF) residue in needle samples following Reed et ale (1982). Briefly, the NDF fraction was isolated using a detergent solution, then incubated in a pepsin solution for 8 hours and the nitrogen remaining measured by kjeldahl analysis. All analyses were run in duplicate and are expressed as percent dry matter. ME was calculated as: ME(Kcal/g) = GEi(Qi)-{[GEef(Qef)+GEec(Qec)]-[Kua(Qua)+Ku(Qu)]} Qi where GEi Qi GEef Qef GEec Qec Kua Qua Ku Qu = gross energy content in food (Kcal/g DM), quantity of food consumed (g DM), gross energy of fecal droppings (Kcal/g DM), quantity of fecal droppings excreted (g DM), gross energy of caecal droppings (Kcal/g DM), quantity of caecal droppings excreted (g DM), constant for caloric value of uric acid (2,686 Kcal/g), quantity of uric acid excreted (g DM), constant for caloric value of urea (2.53 Kcal/g), and quantity of urea excreted (g DM). �182 RESULTS AND DISCUSSION Palatable and Unpalatable Species Trial There were differences in metabolizable energy among needles of the 4 species tested (AOV, F = 33.3, P<O.OOl) and among birds (F = 4.5, P = 0.009). The 2 preferred species (Douglas-fir and lodgepole pine)-did not differ (p = 0.58) in ME content, but both contained more ME (p <0.05) than the 2 non-preferred species, subalpine fir and Engelmann sPruce_(Table 1). ME'levels were similar to the incomplete results of the 1985 trial. Palatability (dry matter consumption, DMC) also differed among species (Table 1), and was closely related to preferences identified previously (Remington 1986); i.e., Douglas-fir lodgepole pine subalpine fir = Engelmann spruce. Palatability is defined as grams of dry matter consumed when only 1 species was offered, and preference is defined as dry matter consumption when 2 or more species were offered. Averaged across birds, the palatability of preferred species greatly exceeded that of the 2 non-preferred species (80 vs. 56 g/day). The net effect of decreased consumption of less palatable foods containing a lower ME was greatly reduced retention of calories (CR, ME x DMC, Table 1) and concomitant weight loss. Weight changes did not seem to be linearly related to caloric retention. For instance, bird #397 lost an average of 3 g retaining 175 Kcals feeding on Douglas-fir yet lost only 8 g on an average caloric retention of 87 Kcals eating spruce. This bird also retained an average of 87 Kcals feeding on subalpine fir yet lost 27 g. There are several factors which could explain this discrepancy. Birds on poor foods (with caloric deficiency) were sedentary and conserved energy expenditures. Birds on good foods were flighty and excited. The source of endogenous energy reserve used to make up deficits in caloric intake could also effect weight losses. Relatively minor caloric deficiencies are likely to be met largely by liver and muscle stores of glycogen. About 3 g (2.7, Hill 1976:607) of water are excreted for each gram of glycogen catabolized which inflates apparent weight losses. Moderate caloric deficiencies are met largely by catabolism of fat reserves. Fat in birds has 2.2 times the caloric value of glycogen or protein (hence only 45% as much must be burned to yield the same amount of calories) (Dargolts 1973) and is only 5-7% water (Maynard et al. 1979). Thus, catabolism of fat should result in apparent weight losses only l/8th that of glycogen or protein (75-78% water). Large, chronic weight losses are met largely by protein reserves which should result in large apparent weight losses. Protein appeared to be catabolized to some extent during this trial since excretion of uric acid (a by-product of protein catabolism) was correlated to weight loss (p< 0.05). Excessive (>60 g) overnight weight losses in 1985 on unpalatable foods were apparently due to protein catabolism since caloric retention was similar ~nd uric acid levels much higher. Previous weight losses presumably used up glycogen and fat reserves. Nitrogen balance (net gain or loss of nitrogen) differed greatly among species (Table 1). Birds were generally in positive nitrogen balance (8 of 12 replicates) on Douglas-fir. The birds were in negative nitrogen balance on all replicates of the other 3 species. Uric acid content of excreta was not consistently related to high nitrogen losses on these species. Definitive conclusions on the cause of these high nitrogen losses cannot be made until urea (and possibly NH3) levels of excreta samples are completed. �183 Table 1. Calorie retention (CR, Kcals, ME x DMC) , metabolizable energy (ME, Kcals/g DM), nitrogen balance (NB, mg), dry matter consumption (DMC, g) and digestibility (DMD, %), and weight change (g) by blue grouse eating needles of 4 ~pecies of conifers; X(SD), ~ ~ 3. Species a PSME CR MEb NB DMC DMD l:;WT 422 (2+F) Bird number (age and sex) 424 (l-M) 425 (I-F) 397 (2+M) 141 1.81 (0.09) +30 77.7 (3.2) 36.6 (2) +13 (2) 152 1.60 (0.16) -22 94.7 (3.1) 31.8 (3) +5 (5) 145 1.79 (0.08) +12 81.0 (1.9) 35.2 (3) +4 (3) 175 1.88 (0.04) -14 93.0 (1.5) 37.3 (1) -3 (6) III 1.68 (0.11) -117 65.9 (5.2) 33.4 (2) -9 (3) 139 1.78 (0.23) -95 78.3 (4.6) 34.8 (3) -6 (6) 120 1.79 (0.10) -144 66.8 (5.2) 34.5 (2) -11 (6) 138 1.67 (0.31) -77 82.9 (9.5) 32.3 (6) -23 (15) 75 1.45 (0.05) -118 51.7 (1.7) 28.2 (0) -16 (3) 81 1.29 (0.25) -93 63.2 (7.2) 25.8 (5) -28 (15) 51 1.11 (0.03) -247 46.0 (8.0) 24.7 (0) -20 (1) 71 1.00 (0.16) -238 71.4 (5.1) 22.5 (2) -12 (4) PICO CR ME NB DMC DMD l:;WT ABLA CR ME NB DMC DMD l:;WT -363 37.8 (9.9) 35.4 (5) -15 (2) 87 1.62 (0.25) -285 53.8 (7.4) 30.0 (3) -27 (6) 85 1.30 (0.11) -236 65.1 (7.0) 27.6 (4) -6 (10) 87 1.19 (0.17) -216 73.0 (2.3) 27.0 (3) -8 (2) 67 1. 78 (0.23) PIEN CR ME NB DMC DMD l:;WT apSME = Pseudotsuga menziesii, PICO = Pinus contorta, ABLA = Abies lasiocarpa, PIEN - Picea engelmanni. DCorrected for uric acid excretion. �184 Preliminary results suggest urea levels are low, which suggests endogenous sources of nitrogen remain relatively constant across species and intake levels. Indigestible plant protein is the only other likely source of nitrogen in excreta and appears to be the major source of nitrogen losses on unpalatable foods. This strongly suggests tannin-like activity, or precipitation of plant protein, rendering it unavailable. Dry matter digestibility (DMD) varied across species (Table 1) and was highly correlated to ME, even across species (Fig. 1). The significance of this is that a gram of dry matter of "good" food appears equivalent to a gram of dry matter of "poor" food in energy content. The difference in ME within, and across, species is derived from the amount of dry matter digested. This supports the hypothesis that digestibility-inhibiting compounds within unpalatable food reduce DMD and ME. The high correlation between DMD and ME, even across species, means ME can be accurately predicted from DMD. DMD is a much simpler, less expensive parameter to measure. Future trials of this type may effectively use DMD to infer ME. Monoterpenes-added Trial Adding monoterpenes to Douglas-fir needles decreased their palatability (F = 18.6, P <0.001, Table 2), increased weight loss (F = 6.84, P = 0.004, Table 2) and afrected DMD (F = 10.9, P <0.001, Figs. 2, 3): DMD and-ME increased from level 1 to level 2-but declined sharply at level 3 (Table 2). The level 2 mono terpene treatment is approximately equivalent to the monoterpene level of subalpine fir and young «75 years) Douglas-fir, both of which are unpalatable to blue grouse (Remington 1986). The minor suppression of palatability at this level is much less than the 33-53% decrease in dry matter consumption of subalpine fir relative to Douglas-fir in trial 1 (Table 1). It is unlikely that monoterpenes are a major contributor to the low palatability of subalpine fir or young Douglas-fir. The level 2 treatment increased DMD (F = 2.80, P = 0.10) over level 1 (no monoterpenes added) for 3 of the 4 birds (Fig. 3). This is consistent with previous work indicating that monoterpene hydrocarbons and low levels of oxygenated terpenes stimulate bacterial fermentation in-vitro (Oh et al. 1967). Caecal bacteria may have benefited from the addition of relatively small amounts of monoterpenes (presumably by using them as a substrate for fermentation) and increased VFA production and DMD. At the higher level, or twice the average concentration found in subalpine fir or young Douglas-fir, DMD plummetted (F = 8.59, P = 0.006). Averaged across birds, DMD went from 31.6% with no monoterpenes-added to 33.6% at levelland to 28.2% at level 3. At higher concentrations, monoterpenes within Douglas-fir inhibit bacterial fermentation (Oh et al. 1967, 1968). Apparently a threshold of monoterpene exposure exists, above which significant depression of DMD occurs (Hobbs et al. 1986). This threshold appears to be above the level of monoterpenes in any of the species tested for at least 3 of the 4 birds. Only 1 of 10 subalpine fir trees and 4 of 10 young Douglas-fir trees previously sampled had a monoterpene content above the level 2 treatment. No needle samples approached the level 3 treatment (Fig. 2). Monoterpenes are probably not responsible for DMD depression in unpalatable foods. Consistent with decreased intake and DMD, weight losses were larger when birds were eating level 3 treated food (-5 vs. +3 g on level 1 + 2). �• +422 36 .. * 34 * ~38 +425 36 * 34 32 .. 32 30 r •• 28 * •• 26 .at .• , 24 24 22 I!! •• 22 11•29 I. 99 I I. 49 11.8e 11•69 * 36 38 * 32 .. .. 0 •• 24 i •• .. •• .,_t 20 18 r.ea I. 59 2.90 ~: ME (Keats/g) Fig. 1- Relationship between metabolizable conifer needles by 4 captive blue grouse. •• •• •• 26 .. .2.99 +397 t: 28 ~ 26 .1.89 t: ••.. •• ~r30 .1.69 6 •• 34 • 1.~9 1.29 #424 38 22 * .. 28 ~ ~26 0 * •• ;,30 • • • .. • * 11.29 11.~9 11.6e I 1.89 ME (Keats/g) energy (ME) and dry matter digestibility of t-' CP \J1 �~ .. 11 ENGELMANN SPRUCE c» 0\ ~ 0 LIMBER PINE HJ 9 8 33 LODGEPOLE D o z w ::> a w a: 7 6 f- DOUGLAS-FIR : 31 SUBALPINE III ->"#- PINE 32 ~ >- r-' 30 C/) w C!:) Cl 29 28 5 m f- FIR YOUNG OOUGLASFIR :J LL a: w ffe:::( ~ 27 4 3 .",IIt..", ... t 26 •• 2 1 o 0.2 0.4 0.6 0.8 1.0 1.2 1.4 1.6 1.8 2.02.2 2.4 2.6 2.83.0 3.2 3.4 3.6 5.0 MONOTERPENE CONTENT (% OF OM) Fig. 2. Dry matter digestibility by blue grouse of Douglas-fir needles treated with 3 levels of monoterpenes and distribution of mono terpene content among needle groups. >a: o �'"?fi. '-" 36 :J 34 >I- rn - I(/) 32 L .."",,- W o C 0: 30 I- W I I~ >- C ~ "- "" " " """- 28 ~ 0: / I o ~ ""- " I 26 , 'II 1.0 2.0 3.0 4.0 5.0 =11= 425 =11= 424 =11= 397 =11= 422 6.0 MONOTERPENE CONTENT (% OM) Fig. 3. Dry matter digestibility by 4 captive blue grouse of Douglas-fir needles treated with 3 levels of monoterpenes (control = 1.25% of DM, level 2 = 2.50% of DM, level 3 = 5.00% of DM). f-' 00 -....J �188 Table 2. Calorie retention (CR, Kca1s; ME x DMC), metabolizable energy (ME, Kcals/g DM), nitrogen balance (NB, mg), dry matter consumption (DMC, g) and digestibility (DMD, %), and weight change (g) by blue grouse eating needles of Douglas-fir and Douglas-fir treated with 2 levels of monoterpenes. Treatment Control CR MEa NB DMC DMDa WT 422 ~2+F~ Bird number (age and sex) 424 (l-M) 42S (l-F) 397 (2+H) 101 (21) 1.38 (0.24) +24 (36) 72.6 (2.7) 30.0 (2.8) +2 (10) lSO (3) 1.66 (0.11) +83 (43) 90.S (S.O) 32.8 (1.1) +14 (11) 11S (26) 1.46 (0.30) +26 (SO) 78.S (2.0) 29.3 (3.6) +4 (1) l3S (9) 1.48 (0.06) +44 (34) 90.8 (2.S) 30.7 (1.9) +2 (9) WI 117 (7) 1.60 (0.03) +32 (20) 73.4 (2.8) 32.2 (0.4) +4 (6) 141 (1) 1.60 (0.12) -1 (4S) 88.7 (6.8) 30.9 (2.3) -2 (7) l4S (S) 1.76 (0.02) +73 (41) 82.1 (2.3) 34. S (1.0) +1 (10) lS3 (19) 1.67 (0.19) +46 (31) 91.8 (1.3) 32.9 (2.8) +1 (12) Level 2 CR ME NB DMC DMD WT 78 (12) 1. 20 (0.21) -60 (62) 64.9 (4.2) 24.8 (1.6) -9 (2) 134 (7) 1.67 (O.lS) -33 (32) 81.0 (S.l) 28.S (2.2) -S (3) 112 (9) 1. Sl (0.11) -1 (9) 74.3 (1.3) 28.9 (1.4) -4 (S) 114 (38) 1.41 (0.37) -S3 (70) 79.9 (6.1) 26.4 (7.1) -7 (5) Level 1 CR ME NB DMC DMD aNot corrected for uric acid excretion. Benzoic Acid-added Trial Benzoic acid is an organic acid (C6HSC02H) occurring in leaves of some plants and in many berries. Benzoic acid has strong antibacterial and antifungal properties (Windholz 1976), and could act to decrease bacterial fermentation analogous to monoterpenes. Benzoic acid was found in Engelmann spruce needles (Remington 1986), and my be responsible for the low palatability and ME of this species. Addition of benzoic acid (at double the concentration of spruce) to Douglas-fir needles had no effect on DMD (F = 1.13, P = 0.30), ME (F = 1.40, = 0.2S) or nitrogen balance (F = 2.8, P = 0.11; Table 3). Addition-of this concentration of benzoic aCid-may be a-small nitrogen drain. Nitrogen gains averaged 92 ± 44 mg on control food and S9 ± S8 mg on benzoic acid-treated food. Benzoic acid is detoxified and excreted as ornithuric acid in birds (Baldwin et ale 1960) which would increase nitrogen excretion. R �189 Table 3. Calorie retention (Cr, Kcals, ME x DMC), metabolizable energy (ME, Kcals/g DM), nitrogen balance (NB, mg), dry matter consumption (DMC, g) and digestibility (DMD, %), and weight change (g) by blue grouse eating Douglas-fir needles treated with benzoic acid (0.28% DM) and untreated; ~(SD), N = 3. Bird number Untreated 422 424 425 397 Untreated CR ME NB DMC DMD DWT 124 1.82 (0.06) 44 68.2 (4.6) 34.8 (2) 162 1.90 (0.09) 94 84.4 (1.4) 35.7 (1) 156 2.04 (0.24) 124 76.3 (1.2) 39.7 (4) 173 2.09 (0.05) 108 83.0 (2) 40.6 (1) -5 (11) Treated CR ME NB DMC DMD DWT -4 (6) 122 1.86 (0.05) 58 65.9 (4.0) 36.0 (2) 1 (6) -4 (3) 170 1.99 (0.21) 65 85.3 (10.5) 37.7 (4) (-3) (13) -6 (1) 134 1. 91 (0.11) 96 70.2 (5.8) 36.8 (2) 3 (7) 132 1.84 (0.11) 16 71.6 (4.0) 35.2 (2) -14 (5) DMD and ME of both treated and untreated Douglas-fir needles were substantially higher than Douglas-fir needles used in the species or mono terpene added trials, suggesting significant tree-to-tree variation in ME. Tree and Needle Age Trial Calorie retention, ME, and DMD declined with needle age in both old and young Douglas-fir (Table 4). This could be due to differences in the absolute levels of nitrogen or energy across needle ages, or to variation in the amounts which are metabolizable. Palatability (DMC) did not differ beteween tree ages (F = 0.04, P = 0.84) or among needle ages (F = 0.23, P = 0.80) (Table 4). The-preferences for young needles and old trees observed in the wild and documented in captivity (Remington 1986) were not translated to differences in palatability as were preferences among species. This may indicate that factors affecting preferences within Douglas-fir are qualitatively or quantitatively different from factors affecting preferences among species of conifers. Dry matter consumption over the course of ths trial was only 69, 72, 77, and 75% of the amount of Douglas-fir consumed in trial 1 for birds 422, 424, 425, and 397, respectively. Bird weights were similar or slightly higher during the latter trial so decreased consumption was not due to decreased body size. It is probable that dry matter consumption declined concomitantly with energy expenditures as birds acclimated to captivity. �190 Table 4. Calorie retention (CR, Kca1s, ME x DMC) , metabolizable energy (ME, Kca1s/g DM), dry matter consumption (DMC, g) and digestibility (DMD, %), nitrogen balance (NB, mg), and weight change (g) by blue grouse eating 1 and 2, 3 and 4, and 5 and 6-year-01d needles of old and young Douglas-fir; K(SD) , N = 3. Bird number (a~e and sex) Tree and needle age 422 (2+F) OLD 1-2 CR MEa DMC DMDa NB t.WT 94 (10) 1.67 (0.07) 56.6 (5.4) 32.2 (1.0) -6 (8) +2 (10) 120 (15) 1.75 (0.06) 68.6 (8.2) 32.5 (0.9) +17 (58) -4 (22) 98 (2.7) 1.65 (0.12) 59.7 (5.4) 31.0 (1.7) -2 (19) o (7) 119 (21) 1.67 (0.14) 70.7 (6.2) 32.5 (3.4) -6 (53) -4 (7) OLD 3-4 CR ME DMC DMD NB t.WT 93 (3) 1.58 (0.02) 58.9 (2.0) 30.7 (0.6) -94 (19) -6 (5) 102 (11) 1.48 (0.13) 69.4 (4.4) 28.0 (0.9) -137 (24) -6 (4) 101 (10) 1.57 (0.08) 64.5 (3.8) 30.2 (2.3) -111 (27) o (3) 104 (21) 1.62 (0.11) 63.6 (9.0) 32.5 (2.1) -132 (53) -26 (15) OLD 5-6 CR ME DMC DMD NB t.WT 70 (13) 1.42 (0.24) 48.9 (3.0) 26.7 (4.0) -143 (55) -17 (5) 98 (9) 1.46 (0.09) 66.9 (3.8) 26.8 (1.6) -164 (18) -5 (12) 94 (14) 1.50 (0.23) 62.7 (1.8) 28.9 (1.8) -110 (24) -5 (6) 127 (12) 1.76 (0.09) 71.9 (4.3) 33.6 (1. 8) -108 (25) -7 (7) YOUNG 1-2 CR ME DMC DMD NB t.WT 82 (6) 1.49 (0.12) 54.8 (2.0) 28.9 (2.0) -82 (29) -6 (9) 115 (16) 1.69 (0.20) 68.3 (3.8) 30.2 (2.1) -'104 (34) +1 (12) 92 (10) 1.55 (0.14) 59.5 (1.3) 27.8 (2.9) -80 (41) -2 (9) 79 (22) 1.15 (0.24) 67.6 (4.8) 24.0 (4.3) -140 (25) +6 (7) YOUNG 3-4 CR ME DMC DMD NB t.WT 78 (4.8) 1.48 (0.08) 52.5 (3.3) 28.7 (1.9) -112 (3) -3 (8) 111 (9) 1.56 (0.06) 71.1 (5.3) 29.7 (0.7) -119 (11) -9 (3) 95 (3) 1.51 (0.20) 63.3 (6.2) 29.4 (3.7) -122 (30) -12 (7) 106 (17) 1.53 (0.29) 69.6 (3.3) 30.6 (4.9) -127 (49) -12 (6) YOUNG 5-6 CR ME DMC DMD NB t.WT 70 (1) 1.40 (0.04) 50.2 (2.5) 27.8 (1.4) -184 (33) -7 (8) 95 (2) 1.43 (0.03) 66.3 (2.9) 28.1 (0.7) -194 (36) -10 (5) 99 (1) 1.50 (0.03) 66.6 (1.9) 28.9 (0.4) -148 (12) -16 (1) 123 (17) 1.64 (0.19) 74.9 (3.5) 33.1 (3.7) -113 (41) -4 (11) aNot corrected 424 (l':"M) for uric acid excretion. 425 (I-F) 397 (2+M) �191 Differences in DMD between tree ages and among needle ages (main effects) were complicated by interactions (p <0.05) between birds and needle age and tree age and needle age (Table 4).- Needles from old trees were more digestible th~n needles from young trees (F = 6.78, P = 0.01); this was most pronounced for 1 and 2-year-old needles. Digestibility of needles from old trees declined with age but digestibility of needles from young trees remained relatively constant with age. Differences in nitrogen balance among treatments generally paralleled differences in DMD (Table 4). Effects of tree age and needle age were again complicated by interactions between birds and needle age and tree age and needle age. Birds were in positive nitrogen balance only on 1 and 2-year-old needles from old trees. Nitrogen losses increased with needle age for both old and young trees (p <0.05) and needles from young trees caused greater nitrogen losses for all 3 needle age groups than needles from old trees (Table 4). Increased nitrogen losses on older needles may be due in large part to differences in nitrogen content among needle ages and increased binding of plant protein to structural components (cell wall) in older needles. Tannins can decrease metabolizable energy or nitrogen by incorporating starch and protein into the cell wall (Reed et ale 1982, Zucker 1983) or by precipitating plant proteins, digestive enzymes, mucin, or salivary protein (Rayudu et ale 1970, McLeod 1974, Mitjavilla et ale 1977, Sell et ale 1985). The importance of cell wall-bound nitrogen in decreasing nitrogen availability was investigated by measuring pepsin-insoluble nitrogen in neutral detergent fiber (NDF) residue in needle samples. Pepsin-insoluble cell wall bound nitrogen comprised from 20 to 40% of total nitrogen (Table 5). Pepsin digestion has a minimal effect on reducing the amount of cell wall bound nitrogen. Douglas-fir needles contained more soluble nitrogen and a higher percentage of total nitrogen in soluble form than did needles of the other 3 species (Table 5). Within Douglas-fir, soluble nitrogen was generally highest in needles from old trees and deGlined with needle age. These differences are in general agreement with blue grouse food preferences (Remington 1986) and nutritional performance (i.e., nitrogen balance, ME) on these foods (Table 1). The poor palatability of subalpine fir, and the poor performance of blue grouse on fir would not be predicted from it's relatively high soluble nitrogen content. Similarly, the relatively small difference in soluble nitrogen content between needle age 1-2 of old and young Douglas-fir cannot explain the large difference in nitrogen balance when birds fed on them (Table 4). Cell wall-bound nitrogen appears to be one of several parameters affecting how much nitrogen and energy in conifer needles is metabolizable by blue grouse. An indirect measure of tannin protein precipitating activity in conifer needles can be derived from an analysis of nitrogen excreted in caecal droppings. The rationale for this is that when a protein is bound to a tannin further digestive degradation and absorption in the intestines is prevented; this complex should be small enough to pass into the caeca. While bacterial degradation of the tannin-protein complex is possible, absorption of amino acids is not (Mortensen and Tindall 1981). Thus, nitrogen in proteins bound to tannins in the gut should ultimately be excreted in caecal droppings. The percentage of total N excreted that was excreted in caecal droppings was highest for non-preferred foods, and generally correlated with negative �192 nitrogen balance (Table 6); i.e., much of the nitrogen imbalance could be explained by apparent tannin activity. Table 5. Total nitrogen, pepsin-insoluble neutral detergent fiber (NDF) nitrogen, and soluble nitrogen in conifer needles used in digestibility trials. Needle group Total NDF-bounda: Old 1-2 PSMEd Old 3-4 PSME Old 5-6 PSME Young 1-2 PSME Young 3-4 PSME Young 5-6 PSME Benzoic acid PSMEe Monoterpene PSt-lEe Species trial PSMEe Species trial PICOe Species trial ABLAe Species trial PlENe 0.97 0.81 0.80 0.84 0.82 0.76 0.86 0.82 0.84 0.78 0.93 0.73 0.258 0.188 0.190 0.171 0.207 0.143 0.273 0.270 0.170 0.317 0.295 0.254 Nitrogen _ Solublei:5 % SolubleC: 0.712 0.622 0.610 0.669 0.612 0.617 0.587 0.550 0.670 0.463 0.635 0.476 73.4 76.8 76.3 79.6 74.7 81. 2 68.2 67.1 79.8 59.3 68.3 65.2 aCalculated as: (% N in NDF after pepsin digestion x NDF wt)/sample wt. bCa1cu1ated as: total N (%) - (NDF-bound N). cPercent of total N that is soluble (extracted by NDF solution or pepsin). dDenotes tree (old or young) and needle (1-6) age of Douglas-fir (PSME). eDouglas-fir used in benzoic aCid, monoterpene, or species trial. PICO = lodgepole pine, ABLA = subalpine fir, PIEN = Engelmann spruce. Table 6. Nitrogen lost in caecal droppings on different needle groups (% of total N excreted). Group Douglas-fir (1-4 yrs) Lodgepole pine Subalpine fir Engelmann spruce Old Douglas-fir 1-2 yrs 3-4 yrs 5-7 yrs Young Douglas-fir 1-2 yrs 3-4 yrs 5-7 yrs Loss 27.8 25.6 38.8 36.9 28.4 38.3 42.5 31.2 37.4 45.0 Notes Highly palatable, positive N Palatable Unpalatable, very negative N Unpalatable, very negative N Highest preference, positive Negative N balance More negative N balance Negative N balance More negative N balance Most negative N balance balance balance balance N balance �193 Bacterial digestion of fiber has been thought to contribute significantly to the energy grouse derive from food (Leopold 1953, Moss and Hannson 1980). Moss (1977) found that red grouse (Lagopus lagopus scoticus) were more effective at digesting fiber in heather (Caluna vulgaris) than sheep or red deer (Cervus elaphus). This proficiency at fiber digestion is extraordinary for a non-ruminant, particularly since the morphology of grouse digestive tracts seems designed to exclude the bulk of dietary fiber from the caeca where it could be fermented (Fenna and Boag 1974). NDF analyses of conifer needles and excreta were available from trials conducted in 1985 to evaluate the extent and variability of fiber digestibility among species of conifers by blue grouse. NDF digestibility varied from 10 to 26% (Table 7) among species. These values are substantially below the 38% digestibility of cellulose and 44% digestibility of lignin in heather estimated by Moss (1977). The inadequacies of the proximate analysis methodology used by Moss to estimate fiber are likely responsible for his inflated digestibilities. The large variability in fiber digestibility among conifer species was unexpected. NDF in lodgepole pine and subalpine fir needles was 2 to 2.5 times more digestible than NDF in Douglas-fir and Engelmann spruce. Total NDF digestibility is a function of how much NDF is made available to cellulolytic fermentation (i.e., that enters the caeca), and the efficiency of digestion of NDF within the caeca. These parameters were evaluated to assess causes of variability in NDF digestibility. Substantially more NDF from lodgepole pine and subalpine fir was diverted into the caeca (29 and 25%, respectively) than from Douglas-fir and Engelmann spruce (13 and 14%, respectively). Digestibility of the NDF that entered the caeca was substantially higher for lodgepole pine and subalpine fir (Table 5). To make these calculations it is necessary to assume that all NDF digestion occurs in the caeca. That significant bacterial fermentation occurs only in the caeca has been demonstrated for willow ptarmigan (McBee and West 1969) and rock ptarmigan (Gasaway 1976). Apparently, both exclusion of NDF from the caeca and efficiency of fermentation effect NDF digestibility. Particle size of fibrous components following grinding of needles in the gizzard may be the determinant factor regulating both how much fiber enters the caeca and how efficiently it is fermented. Since only small particles can enter the caeca and small particles are more likely to be completely fermented, it may be that the distribution of particle sizes of lodgepole pine and subalpine fir needles following griding are skewed towards small particle sizes relative to those of Douglas-fir and Engelmann spruce. The diameter of plant cell walls has been found to be a significant correlate of NDF digestibility in vitro (Spalinger et ale 1986). Table 7. Digestibility of NDF in conifer needles by blue grouse. NDF Species % DM Di~estibility (%~ In caeca Total Douglas-fir Lodgepole pine Subalpine fir Engelmann spruce 0.38 0.50 0.42 0.47 10.3 26.2 21.2 10.4 77 91 87 74 aAssuming no fermentation outside of caeca. Available to caecaa % of total Amount (g) 4.1 11.4 7.7 3.4 13.4 29.0 24.6 14.0 �194 These data shed light on~the manner in which energy is extracted from different species. Fiber digestion is relatively low in Douglas-fir but this is more than compensated for by it's high cell content component which has a high digestibility (57%). Lodgepole pine needles are low in cell contents and high in fiber but this is mitigated by a high NDF digestibility. Uric acid levels in excreta samples were determined using a method developed for poultry excreta (Marquardt 1983). Uric acid absorbs light at 285 nanometers, consequently it's concentration can be determined spectrophotometrically assuming no interference from other compounds which absorb at this wavelength. Although this assumption was proven true with poultry excreta, it became suspect for grouse feces. Endogenous nitrogen losses calculated from uric acid content in excreta following spruce feeding exceeded total nitrogen excretion, clearly impossible. The extent of interference was estimated by measuring absorbance of excreta samples before and after incubation with uricase. Absorbance due to interference was 2-3 times absorbance due to uric acid content for droppings following feeding of fir, lodgepole pine, and subalpine fir, and about 10 times for droppings following spruce feeding. Absorbance due to uric acid content was not predictable from total absorbance (due to uric acid and interference) already measured. Thus, it will be necessary to repeat the analyses and incorporate a uricase incubation step as part of the procedure. Absorbance due to uric acid can then be calculated as initial absorbance - absorbance following uricase digestion. This has been completed for the species trial. Uric acid excretion was generally lowest when birds were feeding on Douglas-fir and lodgepole pine and highest on spruce and fir (Table 8). This variation is due to differential weight losses on these species. Uric acid is a by-product of protein catabolism, birds catabolizing protein to meet energy requirements (presumably after glycogen and fat depletion) should have increased uric acid excretions. Uric acid excretion increased as weight loss increased, particularly at weight losses greater than 10 g. Uric acid excretion was quite low, comprising from 0.14 to 1.8% of the DM of droppings and accounting for only 6.2 to 15.5% of total nitrogen excretion (Table 8). Since urea levels have been found to be negligible, endogenous nitrogen losses appear extremely low. In contrast to typical avian patterns, grouse excrete as much or more nitrogen in the form of ammonium salts as they do as urates (Moss and Hanssen 1980). This may be due to breakdown of urates by microorganisms in the caeca (Mortensen and Tindall 1981). Ammonia levels in excreta samples will be measured. Mean uric acid content of excreta samples from 4 blue grouse fed Table 8. Douglas-fir, lodgepole pine, subalpine fir, and Engelmann spruce. Bird number 424 422 SEecies PSMEb PICO ABLA PIEN (%)a (6.2) (7.8) (15.5) (11.1) 425 mg % DM mg % DM 191 210 404 424 0.44 0.52 1.31 1.43 148 94 119 139 0.27 0.20 0.31 0.27 397 m~ % DM mB % DM 64 148 354 138 0.14 0.37 1.82 0.31 133 229 519 166 0.26 0.45 1.62 0.41 apercent of total nitrogen excreted as uric acid nitrogen. bpSME = Douglas-fir, PICO = lodgepole pine, ABLA = subalpine fir, PIEN Englemann spruce. �195 Fat levels in blue grouse were determined by ether extraction of dried and ground carcass samples (Remington 1983). Fat content as a percentage of live weight was similar to mean values reported for ruffed grouse (Bonasa umbel1us) (2~3%, Thomas et ale 1975), rock ptarmigan (Lagopus mutus) (3.7-4.0%, Thomas and Popko 1981), willow ptarmigan (Lagopus 1agopus) (1.5-2.0%, Thomas 1982; 3.5-4.1%, Brittas and Marcstrom 1982), and black grouse (Lyrurus tetrix) (1.1-2.6%, Ijas et ale 1978) collected during winter (Table 9). These values are substantially lower than maximal values for Spitzbergen rock ptarmigan, which store as much as 32% of their body weight as fat prior to the onset of the continuous winter night (Mortensen et ale 1983). Table 9. Total body fat content of blue grouse collected in Middle Park, Colorado, November-April. Ad x SD Total bodl fat (Eercent live weight) Males Females Ad Yrlg Yrlg 1.29 1.39 1.52 1.65 1.75 2.04 3.07 3.38 0.73 1.12 1.29 1.45 1.52 1.82 2.49 1.33 1.34 1.40 1.65 1.98 2.29 2.97 0.92 1.40 1.59 1.67 3.26 2.01 0.79 1.49 0.56 1.85 0.61 1.77 0.88 Fat content across sex and age groups did not differ greatly, although adults had somewhat higher fat contents. Variability in fat content within sex and age groups was substantial, with maximal values exceeding minimal values by a factor of about 3.5 (Table 9). The ecological significance of this variability is not known. Even maximal fat levels would represent a modest reserve. LITERATURE CITED Baldwin, B. C., D. Robinson, and R. T. Williams. 1960. Studies in detoxification. The fate of benzoic acid in some domestic and other birds. Biochem. J. 76:595-600. Brittas, R., and V. Marcstrom. 1982. Studies in willow grouse Lagopus 1agopus of some possible measures of condition in birds. Ornis Fenn. 59:157-169. Dargo1ts, V. G. 1973. 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Chem., Washington, D.C. 1018 pp. Ijas, L., I. Nuvja, Palokangas, J. Soimajarvi, and P. Valkeajarvi. 1978. Body lipid composition in the winter fed and control populations of the wild male black grouse (Lyrurus tetrix L.) in autumn and spring. Compo Physiol. 60:313-317. Leopold, A. S. 1953. Intestinal morphology of gallinaceous birds in relation to food habits. J. Wildl. Manage. 17:197-203. Marquardt, R. R. 1983. A simple spectrophotometric method for the direct determination of uric acid in avian excreta. Poultry Sci. 62:2106-2108. Maynard, L. A., J. K. Loosli, H. F. Hintz, and R. G. Warner. 1979. Animal nutrition. Seventh edition. McGraw-Hill Book Co., New York, N.Y. 602 pp. McBee, R. H., and G. C. West. 1969. ptarmigan. Condor 71:54-58. Cecal fermentation in the willow McLeod, M. N. 1974. Plant tannins - Their role in forage quality. Abstr. and Rev. 44:803-815. Nutr. Mitjavilla, S., C. Lacombe, G. Carrera, and R. Derache. 1977. 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Effects of various essential oils isolated from Douglas-fir needles upon sheep and deer rumen microbial activity. Appl. Microbiol. 15:777-784. Rayudu, G. V. N., R. Kadirvel, P. Vohra, and F. H. Kratzer. 1970. Effect of various agents in alleviating the toxicity of tannic acid for chickens. Poultry Sci. 49:1323-1326. Reed, J. D., R. E. McDowell, P. J. VanSoest, and P. J. Horvath. 1982. Condensed tannins: a factor limiting the use of Cassava forage. J. Sci. Food Agr. 33:213-220. Remington, T. E. 1983. Food selection, nutrition, and energy reserves of sage grouse during winter, North Park, Colorado. M.S. thesis, Colorado State Univ., Fort Collins. 89 pp. 1986. Food selection and nutritional ecology of blue grouse during winter. Job Prog. Rep., Colorado Div. Wildl. Fed. Aid Proj. 01-00-045 (W-37-R). Apr. Pp. 355-377. Sell, D. R., W. M. Reed, C. L. Chrisman, and J. C. RogIer. 1985. Mucin excretion and morphology of the intestinal tract as influenced by sorghum tannins. Nutr. Rep. 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An �198 Prepared Approved �199 Colorado Division of Wildlife Wildlife Research Report April 1987 JOB PROGRESS REPORT State of Colorado Project 01-03-045 (W-37-R) 12 Work Plan Job Title: 15 --- Chronology of Breeding and Nesting Activities of Wild Turkeys in Relation to Timing of Spring Hunting Seasons Period Covered: Author: : Job Avian Reseqrch 01 July 1985 through 31 June 1986 Richard W. Hoffman Personnel: C. E. Braun, A. L. Cade, R. W. Hoffman, R. L. Holder, B. S. Linkhart, and T. J. Spezze, Colorado Division of Wildlife; R. S. Nuse, Colorado State University ABSTRACT Gobbling and nesting activities of 50 radio-marked (7 males, 43 females) Merriam's wild turkeys (Meleagris gallopavo merriami) were monitored in relation to timing of the spring hunting season in southeastern Colorado. Winter flocks dispersed in early (males) to mid- (females) April. Females moved 9.6 ±4.3 km from wintering to breeding areas and occupied a breeding home range of 653 ± 397 ha compared to movements of 3.7 ± 1.7 km and home ranges of 1,541 ±542 ha for males. Eleven of 13 mornings of above average ~27 gobbles/bird/hr) gobbling occurred between 16 April and 2 May in conjunction with the spring hunting season. A second peak of shorter duration occurred after the season from 13 to 20 May. Gobbling was sporatic and varied among males, but the tendency was for,males to gobble more in the morning than evening, more on than off the roost, and more when aione than when accompanied by females. Only.3 hens ini~iated incubation prior to the.end of the spring hunting season, 8 started between 'i9 and 31 May, while 2 started between 6 and 18 June. The peak incubation period coincided with the second peak in gobbling activity. Based on a 100% survey, 4,174 spring hunters harvested 563 turkeys (13% success) and 1,414 fall hunters harvested 313 turkeys (22% success). Las Animas County accounted for 23 and 31% of the spring and fall harvest, respectively. Public land supported 54 (spring) and 52% (fall) of the pressure, but produced only 45 (spring) and 36% (fall) of the harvest. Hunter success was better on private (spring = 20%, fall = 35%) than public (spring = 10%, fall = 15%) land. The wing collection program sampled 69% of the spring harvest and 78% of the fall harvest. Merriam's dominated (>91%) the harvest samples. Bearded hens comprised 2% (N = 11) of the total harvest. Seventy-six percent of the spring harvest of Merriam's were adults compared to 43% adults for Rio Grande's. The fall harvest (excluding Rio Grande's due to an inadequate sample) was comprised of 55% juveniles. Sex ratios favored females (1.5:1) for adults and subadults combined and males �200 (1.6:1) for juveniles. ~timated hatching dates based on available methods for classifying age of fall-harvested juveniles were not in agreement with known hatching dates for clutches of radio-marked hens. �201 CHRONOLOGY OF BREEDING AND NESTING ACTIVITIES OF WILD TURKEYS IN RELATION TO TIMING OF SPRING HUNTING SEASONS Richard \\1. Hoffman Wild turkeys have been legally hunted in Colorado since 1949 (Burget 1957). Fall-only hunting was permitted until 1964 when the first spring season was held; subsequent spring seasons were held in 1965, 1967, 1968, and 1970 (Colo. Div. Wildl. 1984). A continuous 23 to 30-day season, opening in mid-April and extending through early to mid-May, has been held every year since 1973. Fall seasons have ranged from 9 to 16 days, opening in late September and closing in early to mid-October. In 1973, an estimated 1,301 hunters (spring = 496, fall = 805) harvested 265 turkeys (spring = 64, fall = 201). By 1983, 4,894 hunters (spring = 3,030, fall = 1,864) were harvesting 1,147 turkeys (spring 645, fall = 502). The Colorado Division of Wildlife (CDOW) responded to this growing demand by focusing their management efforts on (1) restoring populations of Merriam's wild turkeys in western Colorado, and (2) establishing new populations of Rio Grande (~ •.a. intermedia) wild turkeys in eastern Colorado. The success of the transplant programs plus the increase in hunter participation generated a need for more refined data upon which to base season recommendations. Paramount is the need for reliable estimates of harvest by area, subspecies, age, and sex, and for better documentation of the chronology of breeding and nesting activities on a regional basis. Previous studies of the wild turkey in Colorado (Burget 1957; Hoffman· 1962, 1966, 1968, 1973; Myers 1973) failed to address these concerns. To date, no quantitative data exist upon which to justify the timing and length of spring seasons or to biologically justify these seasons. CDOW personnel in the SE Region feel that acquisition of such information is critical to convincing landowners to allow spring hunting and to meeting strategic plan objectives of increasing hunter opportunity and harvest through improved access. P. N. OBJECTIVES 1. Document the timing of winter flock dispersal, onset of gobbling, peaks of gobbling, nest initiation, onset of incubation, and peak of hatch in relation to timing of the spring season. 2. Describe the gonadal cycle of females and compare the reproductive condition of females in relation to timing of the spring season. 3. Measure the abandonment rate of incubating females to varying levels of human disturbance around the nest. 4. Monitor hunter activity and harvest of wild turkeys on a statewide basis. �202 SEGMENT OBJECTIVES 1. Review literature pertinent to the objectives of this study. 2. Trap and instrument 40 wild turkeys (6 males, 34 females) with radio transmitters. 3. Document timing of winter flock break-up. 4. Document onset of gobbling and peaks of gobbling activity. 5. Document onset of egg laying, incubation, and peak of hatch. 6. Measure effects of human disturbance on rate of nest abandonment. 7. Implement a hunter questionnaire-wing collection program using a permit system and mail wing survey. 8. Compile data, analyze results, and prepare progress report. DESCRIPTION OF STUDY AREA Trapping operations were confined to Longs Canyon and 2 tributary canyons, Sowbelly and Martinez, 17 km southwest of Trinidad, Colorado 'in Las Animas County. From here, birds dispersed into an area encompassing over 448 km2• This area is roughly bounded by 125 on the east, Lorencito Canyon on the west, Colorado Highway 12 on the north, and the Canadian River in New Mexico on the south. Topography is mountainous, varying in elevation from 1800 to 2600 m. Four large canyons in excess of 30 km in length occur within the area, each with numerous side canyons and adjacent smaller canyons. Major vegetation types include pinyon pine-juniper (Pinus edulis-Juniperus spp.), mountain shrub, and ponderosa pine (~. ponderosa). The mountain shrub type is dominated by Gamble's oak (Quercus gambelii), which extends into both the pinyon-juniper and ponderosa pine types. Douglas-fir (Pseudotsuga menziesii) and occasional white fir (Abies concolor) occur on north slopes within the ponderosa pine type. Most of the area is under private ownership except for 1100 ha of land around Trinidad Reservoir administered by the Colorado Division of Parks and Outdoor Recreation. Present use of private lands is limited to cattle grazing and some logging. METHODS Trapping, Marking and Radio-tracking Wild turkeys were baited with oat hay and livetrapped during February and March using drop nets (Glazener et al. 1964), cannon nets (Dill and Thornsberry 1950), and clover traps (Clover 1958). Unsuccessful efforts were made to capture birds with cracked corn laced with alpha-chloralose at a dosage of 2 gm per cup (Williams 1966, Williams et al. 1973, Bailey et al. 1980). Captured birds were classified to age (Hoffman 1962) and sex (Hoffman 1962)) and banded with serially-numbered aluminum leg bands and patagial wing tags (Knowlton et al. 1964). Ages were recorded as subadult (8-10 months) �203 and adult (>18 months). -~ody weight, and length of primaries, carpal, spur, and beard were measured on each bird. Fifty birds were equipped with lithium battery-powered transmitters (Wildlife Materials, Inc., Carbondale Ill.) attached with a poncho collar (Amstrup 1980) or tail-clip (Bray and Corner 1972). Tracking was conducted from the ground using a hand-held, 3-element, Yagi antenna and Telonics TR-2 receiver. An aerial search was made on 30 May 1986 to locate missing birds. All locations were verified by visual observations and recorded to the nearest 50 m as Universal Transverse Mercator (UTM) coordinates. Gobbling Indicies Flocks containing instrumented birds were monitored a m1n1mum of 3 times per week beginning on 26 Febru~ry to determine onset of gobbling and period of flock dispersal. Gobbling indicies were conducted from 26 March to 25 June and categorized as preseason (26 Mar-18 Apr), hunting season (19 Apr-18 May), and postseason (after 18 May). An attempt was made to conduct 3 valid indices per week per time period (i.e., AM and PM). A gobbling index was considered valid if (1) positive identification was made of the instrumented bird(s) that was gobbling, (2) the bird(s) was not disturbed prior to or during the gobbling index, (3) the time the bird left (AM gobbling index) or went (PM gobbling index) to roost was known, and (4) the index included time on and off the roost. While it was not considered necessary to know the exact number of other birds present during the index, it was necessary to know whether the male being monitored was alone or associated with other males, females, or both sexes. A gobbling index lasted 1 hour from one half hour before to one half hour after sunrise (AM index) or sunset (PM index). The 1 hour period was divided into 6 10-minute listening periods so gobbling intensity could be quantified within the I-hour interval. Gobbling was also recorded in relation to presence or absence of other birds and whether the bird was on or off the roost. When possible, the gobbling activity of the instrumented male was compared with that of uninstrumented males he was associated with to assess the influence of the radio package on gobbling. This could only be done for a portion of the index. Therefore valid indicies were limited to instrumented males. Instrumented males were monitored on a rotating basis, the initial order being randomly selected. This worked for the first 3 weeks, then 3 of 7 males made extensive movements and became increasingly more difficult to consistently locate. Consequently, the order could not be maintained, but was adhered to as best possible. For AM indicies, the gobbler was located on the roost either the evening before or 1 hour before sunrise on the monitoring morning. Gobblers selected for a PM index were located at least one hour before sunset. Nesting and Movements Radio-marked hens were located at least once every 3 days from time of capture until flock dispersal. Subsequent locations varied depending on how far the hen moved from wintering to breeding area. Birds moving longer distances were relocated less frequently because it took considerable searching time to find them initially. Priority was given to locating nests and documenting hatching dates. Timing of other nesting activities (i.e., nest initiation, onset of �204 incubation) was approximated by knowing the date of hatch. Clutch size, fertility, and nesting success were determined from egg shell characteristics after the eggs hatched or after the nest was abandoned or depredated. Hens were not approached closer than 5 m while on the nest nor were they deliberately flushed off the nest to ascertain clutch size. Breeding home ranges (area occupied from 15 Apr-18 Jun) were calculated for both males and females using the minimum area polygon (Mohr 1947). Activity centers (males only) were defined as the area within the home range containing >70% of all locations. Winter to breeding range movements of males were calculated as the minimum distance between the trap site and the center of their activity center. For females, the distance was measured from the winter trap site to the nest site or, in the case of hens not located on a nest, from the trap site to the center of their 15 Apr-15 Jun breeding range. Permits, Questionnaires, and Wing Collections Hunters were required to obtain 1 of 2 types of permits during the 1986 spring and fall seasons: 1. Special Unlimited Hunting Permit - Unlimited in number and free of charge, these permits were available to any holder of a valid turkey license. Whereas the license could be purchased at any license agent, the permits were only available from CDOW offices either on a walk-in basis or by mail application. The special unlimited permits were valid for one season (spring or fall). Unsuccessful spring hunters could hunt in the fall without purchasing a new license, but before doing so, they were required to obtain another permit valid for the fall season. Successful spring hunters who wanted to hunt the fall season needed to purchase another license in addition to obtaining a special unlimited fall permit. By requiring spring and fall permits, hunters could be categorized as spring only, fall only, or spring-fall hunters and surveyed accordingly. Special unlimited permits for the spring season were available from 1 March to 18 May 1986 (end of season); those for the fall, from 1 August to 5 October 1986. Special unlimited permits were valid for all areas not requiring a limited permit. 2. Limited Hunting Permits - Limited in number, free of charge, and available by public drawing. Only mail applications were accepted for limited permits. The drawing and issuance of permits was handled through the Denver office. Hunters could apply for a limited permit without first purchasing a license. However, if they succeeded in drawing a limited permit, they needed to purchase a license before going hunting. Drawings were held on 28 March and 29 August 1986 for the spring and fall seasons, respectively. Limited permits were only valid for the area, season, and time period indicated on the permit. Holders of a limited permit were required to obtain a special unlimited permit if they hunted outside the area for which their limited permit was valid. Consequently, some hunters obtained 2 permits and were subsequently mailed 2 harvest questionnaires. This problem was identified as the questionnaires were being processed and the duplicates were excluded. Questionnaires were mailed to all permit holders immediately after the spring and fall seasons. Non-respondents were mailed a followup questionnaire �205 approximately 3 weeks later. Mean values calculated from responses to the second mailing were used to project answers for those permit holders not responding to either questionnaire. Every hunter that obtained a limited or unlimited permit was also issued a wing envelope coded by permit number. The instructions on the envelope requested each successful hunter to (1) complete the questionnaire printed on the envelope including their name, address, time of harvest, and location of harvest (county, small game management unit, and nearest town), and (2) to remove the least damaged wing and 3 or 4 breast feathers as depicted by 2 schematic diagrams, place them in the envelope, and mail the postage-paid envelope. The envelopes were addressed to the Wildlife Research Center in Fort Collins. Upon delivery, the wings were frozen until they could be processed. Information provided by hunters was transcribed onto a standardized form. The envelope's contents were examined to determine if both a wing and breast feathers were enclosed. The samples were then identified to su~species and classified to age and sex. Whenever possible, the following measurements and feather characteristics were recorded for each wing: length, condition (growing, fully grown, empty, broken, worn, pointed) and status (adult or juvenal) of primaries I through X (numbered proximal to distal), quill diameter of P IX and P X at their insertion into the follicle, stage of primary molt, carpal length, and status (adult or juvenal) of the first secondary and tertials. Since the range of the Merriam's and Rio Grande wild turkey does not overlap in Colorado, except possibly along the Arkansas River west of Pueblo, the 2 subspecies were identified from wing samples based on the location of harvest. However, inspection of the wings suggested that subspecies could be distinguished by wing color. Rio Grande's tended to have blacker primaries and fewer white bars than Merriam's, thus, giving the wing a darker appearance. Sex was ascertained by the presence of buffy (female) or black (male) tipping on the breast feathers (Hoffman 1962). Wings were assigned to age classes based on the shape, color, wear, and barring pattern of primaries IX and X (Hoffman 1962). Juvenal primaries IX and X were pointed, grayish-brown, and lacked white barring near the tip. Comparatively, adult primaries IX and X were rounded and black or blackish-brown, with white barring extending almost to the tip. Subadults possessed the outer primaries characteristic of juveniles except they were faded and worn as a result of being retained longer. Wings obtained from the spring season were classified as adults (>22 months) or subadults (10-11 months). Those from the fall were separated into 3 age classes: adults (>26 months), subadults (14-16 months), and juveniles « 6 months). Two techniques, both involving measurements of the actively growing post-juvenal primaries, were used to calculate the approximate age (±7 days) of juveniles (Knoder 1959, McGuire 1964). Knoder (1959) developed his key using eastern wild turkeys (~.£.silvestris), while McGuire (1964) used Rio Grande's. The reliability of neither method had been previously tested on Merriam's; however,. McGuire (1964) compared the mean length of primary feathers of eastern and Rio Grande wild turkeys and found a significant difference (p <0.10). Samples from Colorado were classified to age using �206 both methods and the hatching dates calculated. The results were compared for agreement between methods and for agreement with known hatching dates as determined from observations of radio-marked hens. RESULTS AND DISCUSSION Capture and Marking Ninety-nine turkeys (10 males, 89 females) were trapped in late February-early March with drop nets (75), cannon nets (20), or clover traps (4). Four subadu1t hens, 39 adult hens, and 7 adult males were equipped with leg bands, wing tags, and radios, and released at the trap site (Table 1). Two birds died during trapping. The remaining 47 birds were leg banded and released (26) at the trap site or transplanted (26) to an area east of Colorado Springs as part of the "Ranching for Wildlife" program. Twenty-two (25%) of the 89 hens captured had beards. All bearded hens were adults. The beard length was 134 ± 52 mm ± SD). The shortest beard was 35 mm, but it was still clearly visible through the breast feathers. Beard length for adult males was 2l6± 17 mm. Only 1 immature male was captured and his beard was 73 mm long. Adult males weighed 7.6 ± 0.6 kg compared to 3.9 ± 0.2 kg for adult females and 3.0±0.5 kg for subadult females. The immature male weighed 4.7 kg. (:!: Two attempts were made to capture turkeys with drugged baits. The first attempt involved a flock of 14 hens that .included 2 radio-marked birds. Ten individual piles (1 cup cracked corn/pile) of drugged bait (2 gm a1phachloralose/cup) were placed at 0430 hours on 12 March 1987 on a site the birds had been visiting for 7 days previously. The bait was mixed the night before. Fourteen hens arrived at the bait at 0650 hours. They fed intermittently until 0710 hours. By 0715 hours the birds were off the bait, at which time only 1 hen was showing signs of being drugged. This hen went down at 0745 hours while the other birds moved off into the trees out of visual contact. We approached the bait site at 0815 hours and found TX 1244 lying 75 m from bait site. She was completely comatose. Examination of the bait piles indicated less than 50% was consumed. We removed the leftover bait and proceeded to track the main flock (including TX0538) for another 2 hours catching occasional glimpses of the birds. None appeared to be affected by the drug. We returned to the bait site the same evening (1700 hrs. on 12 Mar) to locate TX0538. At least 5 birds ran into the trees as we drove up. We found TX0538 in a comatose state approximately 1 km from the bait site within a 100 m of where we left the flock at about 1030 hours that morning. The minimum amount of time it took her to succumb to the drug was 3.5 hours. The effects of the drug lasted almost 34 hours. TX1244 and 0538 were completely comatose for 24 hours; TX1244 was still somewhat lethargic when released on 14 March. Both hens were located on 19 March with 11 other hens near the bait site. If these were the same hens they were initially associated with, then only 1 hen was lost from the flock possibly as a result of the drug. Our second trapping effort with drugged bait involved a flock of 8 adult males. The bait (8 piles) was placed out at 0553 hours on 16 March. The dosage was increased to 2.5 gms of drug per cup of cracked corn. The birds arrived at 0649 hours, fed for 6 minutes, then moved into the trees. There �207 _-.;,_ Table l. Colorado, Wild turkeys trapped and equipped with radios in Longs Canyon, February-March 1986. Band # Sex ABe 18 19 20 64 65 66 67 M M Ad Ad Ad Ad Ad Ad Ad 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 56 57 58 59 60 61 62 63 86 F F M M M M M F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad SubAd SubAd SubAd SubAd Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad Weight (kB) Carpal (mm) 6.6 7.1 7.7 8.2 7.9 8.5 7.5 522 523 525 519 518 531 533 4.1 4.1 3.8 4.1 4.0 4.2 3.9 3.3 3.6 3.6 3.2 2.4 3.5 3.0 4.1 4.2 3.6 3.4 4.2 4.0 3.9 4.0 3.8 4.0 3.6 3.7 4.1 3.3 4.0 3.7 4.0 3.8 3.6 4.2 3.9 4.0 4.2 3.8 3.9 3.8 3.9 3.6 4.0 451 434 432 422 423 450 454 440 432 427 420 395 417 399 444 434 440 444 438 446 435 438 435 435 432 452 445 440 453 438 434 434 443 441 450 440 440 430 437 451 448 439 Beard (mm) Spur (mm) Radio freguencr 240 200 225 230 215 210(160)b 190 16 12 16 20 18 17 21 150.502-T 150.441-T 150.555-T 150.619-T 150.837-T 150.679-T 150. 778-T . 35 65 73 82 172 185 191 181 155 165 105 180 130 160 aT = tail mount, P = poncho mount. bDoub1e beard. cRadio recovered from bird #28 (mortality) and put on bird #6l. dRadio recovered from bird #39 (mortality) and put on bird 1162. 15l.159-P 15l.100-P 15l.259-P 15l.425-P 15l.241-P 151. 144-P 15l.218-P 15l.282-P 15l.201-P 15l.178-P 15l.121-P 15l.319-P 150.180-P 15l.068-P 150.278-T 150.639-T 150.880-T 150.306-T 150.342-T 150.178-T 150.158-T 150. 577-T 150. 213-T 150. 399-T 150. 372-T 151.750-P 150. 513-T 150.139-T 150.261-T 150.661-T 150. 412-T 150.798-T 15l.440-P 150.358-T 150.860-T 150.536-T 150.502-T 150.760-T 150.815-T 15l.282-pc 150. 342-Td 150.700-T 150.457-T �208 was some preening dispersed between bouts of displaying, gobbling, and chasing amongst the males. By 0730, the birds had moved away. One male appeared to be partially drugged, but he left when the other birds did. Examination of the.bait revealed less than 1/3 cup of corn was consumed. The birds had scratched around the corn piles searching for oats remaining from the original bait used to attract them to the site. We revisited the bait site that evening and saw 6 males suggesting 2 had succumbed to the drug. This flock was not seen again at the bait site. Use of alpha-chloralose baits treated at a dosage of 2-2.5 gms/cup is not recommended because of its variable, delayed effects. Increased dosages may result in mortality and, thus, need further testing. Treated cracked corn was avoided or consumed at a lower rate than untreated cracked corn suggesting the birds could detect the drug. Any birds trapped with drugged baits must be held a minimum of 30 hours before being released. Onset of Gobbling and Flock Dispersal Gobbling was first heard on 11 March and continued through 25 June when gobbling indicies were terminated. Male and female flocks remained intact through March with males seldom joining the hens. Late March activities of males consisted of early morning gobbling bouts followed by periods of fighting and strutting. The males seemed more interested in confronting each other than pursuing hens, which were often nearby. The flock of radio-marked males dispersed during the first week of April; however, none associated with females until 8 April. Unmarked males were seen with females as early as 1 April. During the preseason (26 Mar-18 Apr) monitoring period, radio-marked males were accompanied by females in only 10 (32%) of 31 observations. In comparison, they were observed with females on 20 (69%) of 29 occasions during the hunting season (19 Apr-18 May) and 11 (65%) of 17 times after the season (19 May-25 Jun). Gobbling Indicies Ninety-four valid gobbling indicies were obtained on 7 adult, radio-marked males between 26 March and 25 June. Gobbling was sporatic throughout the monitoring period, but peaked between 16 April and 2 May in conjunction with the spring hunting season (Table 2). Eleven of 13 mornings of above average gobbling ~27 gobbles/bird/hr) occurred during this time. A second peak of shorter duration occurred between 13 and 20 May. Early peaks in gobbling resulted from intense competition among males seeking to establish dominance and attract hens. The second peak of gobbling coincides with the peak of incubation (Bevill 1975, Bailey and Rinell 1967). Hunting seasons should be timed to bracket the second peak of gobbling when males are still sexually responsive but hens are unavailable. Gobbling activity differed among males monitored, but the tendency was for them to gobble more in the morning than evening (Table 3), more on than off the roost (Tables 4 and 5), and more when alone than when accompanied by hens (Table 6). Whereas cloud cover (Davis 1971), precipitation (Bevill 1973), and wind velocity (Bevill 1973) have been found to inhibit gobbling, weather conditions experienced in this study appeared to have no influence on gobbling (Tables 7 and 8). However, gobbling indicies were only conducted on 6 days �209 Table 2. AM gobbling activity in relation to timing of the spring season, 26 March to 25 June, 1986. Date Descriptive statistic Preseason 26 Mar 30 Mar 31 Mar 07 Apr 08 Apr 10 Apr 11 Apr 15 Apr 16 Apr. 17 Apr 18 Apr gobblers monitored a Total gobbles b Gobb1es/ bird/hr X ± SD 5 2 3 1 3 1 2 3 1 1 2 2 2.2 ± 1.3 0 36 39 24 58 1 10 39 52 88 89 39 39.6 ± 30.8 0 18 13 24 19.3 1 5 13 52 88 44.5 18 25.2 ± 26.6 x Median ± SD 1 1 3 1 2 1 1 1 1 1 1 1 2 2 1 1 1 1 1 1 1.3 ± 0.6 84 3 181 40 6 68 28 34 41 113 23 1 23 37 8 7 9 76 37 34 43.1 ± 45.3 84 3 60.3 40 3 68 28 34 41 113 23 1 11.5 18.5 8 7 9 76 37 28. 35.0 ± 31.9 Median ± SD 43 182 8 3 28 58 0 1 2 2 5.5 32.7 ± 56.3 43 45.5 4 x 1 4 2 1 3 4 1 2 1 2 2 2.1 ± 1.2 9.3 14.5 0 0.5 2 1 3.5 12.3 ± 17.4 Median SD 1 1.7 ± 1.0. 31 39.6 ± 44.0 16.3 26.7 ± 28.5 Median Hunting Season 21 Apr 22 Apr 23 Apr 24 Apr 26 Apr 27 Apr 28 Apr 30 Apr 01 May 02 May 03 May 04 May 06 May 07 May 09 May 11 May 12 May 13 May 15 May Postseason 19 May 20 May 22 May 23 May 28 May 04 Jun 05 Jun 08 Jun 18 Jun 25 Jun All Seasons .!i :! ± )- aRadio-marked males only. bTota1 gobbles during I-hour interval from one half hour before to one hour after sunrise. �210 when precipitation was recorded at the weather station. Trace amounts «0.02 in.) fell on 3 of 6 days (8, 16, 26 Apr) and from 0.12 (18 Apr) to 0.14 in. (30 Apr, 1 May) were recorded on the other 3 days. Above average gobbling was recorded on 4 of the 6 days when precipitation occurred, including the 3 days of highest rainfall. Gobbling indicies were not conducted on days when rainfall exceeded 0.2 in. because of restricted access caused by muddy roads. Few days (N = 4) were encountered when the wind velocity was >10 mph during the listenIng interval as the early morning and late evening hours were usually the calmest periods of the day. Gobbling was above average on 3 of 4 days of >10 mph wind velocities. The available data are not sufficient to draw conclusions regarding the influence of precipitation >0.02 in. and wind >15 mph on gobbling. Table 3. 1986. Txll AM vs. PM gobbling activity of radio-marked wild turkeys, spring Observation time (min)a AM PM 0679 0504 0557 0837 0619 0441 0779 720 600 840 300 600 360 360 180 660 120 120 0 240 360 Al1c 2,764 1,380 AM 0.43 0.33 0.27 0.52 0.62 0.61 0.17 0.41 ± 0.66 0.03 ± 0.05 0.15b 0.29b 0.51 ± 0.49 0.12 ± 0.26 0.40 0.31 0.24 0.83 0.49 0.82 0.17 ± ± ± ± ± ± ± Gobbles/min (x ± SD) PM 0.07 ± 0.09 0.003 ± 0.008 aThe observation time was from one half hour before to one half hour after sunrise and sunset. bN = 2, no standard deviation calculated. cExcludes duplicate observations when 2 or more radio-marked males were together. �211 Table 4. On-off roost AM gobbling activity of radio-marked wild turkeys, spring 1986. TXfJ Observation time (min)a On Off 0679 0504 0557 0837 0619 0441 0779 287 175 354 114 204 129 106 433 425 486 186 396 231 254 Allb 1,024 1,740 Gobbles/min On 0.86 0.73 0.49 1.75 0.94 1.83 0.50 ± ± ± ± ± ± ± Ci ± SD) Off 0.83 0.67 0.53 1.14 1.08 1.52 0.47 1.07 ± 1.02 0.18 0.10 0.07 0.10 0.23 0.41 0.60 ± ± ± ± ± ± ± 0.30 0.11 0.17 0.13 0.41 0.50 0.11 0.20 ± 0.32 aThe observation interval was from one half hour before to one half hour after sunrise. bExcludes duplicate observations when 2 or more radio-marked males were together. Table 5. On-off roost PM gobbling activity of radio-marked wild turkeys, spring 1986. TXI! Observation time (min)a On Off Gobbles/min (x ± SD) Off On 0679 0504 0557b 0837b 0441 0779 69 235 25 42 86 74 III 425 95 78 154 286 0.48 ± 0.71 0.03 ± 0.05 0.52 0.83 0.32 ± 0.44 0 0.37 ± 0.63 0.03 ± 0.06 0 0 0 0.003 ± 0.008 Allc 468 912 0.25 ± 0.46 0.06 ± 0.23 aThe observation interval was from one half hour before to one half hour after sunset. bN = 2, no standard deviation calculated. cExcludes duplicate observations when 2 or more radio-marked males were together. �212 Table 8. Wind velocity, cloud cover and precipitation in percent occurrence, on 14 days of above average gobbling and 26 days of below average gobbling.a Weather condition Gobbling activity Above average Below average 57 22 14 65 31 Wind velocity (mph) .:s_5 >5 <10 > 10 ~15 >15 Cloud Cover <10% > 10 .:s_75 > 75 Precipitation None Trace «0.02 in.) Moderate (0.08-0.14 in.)b 4 7 o 64 22 14 54 19 27 79 73 15 12 7 14 aAverage gobbles based on data from Table 1 for each period (preseason, season, postseason). bGobbling indicies were not conducted on days when rainfall exceeded 0.2 in. because of access problems. Roosting Activity Males left the roost earlier during the preseason and hunting season monitoring periods than during the postseason period (Fig. 1). Males left the roost an average of 14 minutes before sunrise between 26 March and 18 April, 9 minutes before sunrise between 19 April and 18 May, and 15 minutes after sunrise between 19 May and 25 June. There were only 7 mornings prior to 20 May when males left the roost at or after sunrise. It was raining or had rained the night before on 5 of these mornings. Starting on 20 May, males only came off the roost after sunrise. This date coincides with the culmination of the second peak of gobbling and peak incubation. Males went to roost after sunset during the preseason (x = 8 min. after sunset) and first half of the hunting season (K = 9 min.). On 3 May there was a distinct shift in evening roost activity with males going to roost an average of 10 minutes before sunset (Fig. 2). Males went to roost before sunset throughout the postseason monitoring period. Movements and Home Range Seven adult males moved 3.7 ± 1.7 km from wintering to breeding areas (Table 9). Once on the breeding range, they were extremely mobile as evidenced by their large home ranges (1,541 ± 452 ha) and activity centers (415 ± 156 ha), and by daily movements between consecutive roosting sites of 2,036 ±75l m (Table 9). The home ranges of all 7 males overlapped. One radio-marked male �+30r I + = AFTER SUNRISE - W +20~ a: z ::J +10r- • = BEFORE SUNRISE I I I CI) CI) zg 0 - - - ....-.. ~ en 02 a: ::J u, .£; o 0 I I 1111 I1111··11 I IIII -10~ !I _1111111' I 11111-111 1 I I I I 4 ~ ~ > W - I I I -20 ~ .! !I I I I II ' • I I 4 • . 4 ~ 1 -30'2'613'11811'111611'812'212'412'713'012141711'111'311 912'212'81511'81 30 7 10 15 17 21 23 26 28 1 3 6 9 12 16 20 23 4 8 25 MAR Fig. 1. APR MAY Minutes before or after sunrise that radio-marked male wild turkeys left the roost. JUN N I-' W �r +3O t- + = AFTER SUNSET - = BEFORE SUNSET +20~ W z +1o~1 (J) 1 I I!, 0 ,£; zE, > .,. I I j a:"S 0 f u ~(j) 00> LL ..,.. j (J) ::J N I-' T , • I I I • ! -101- I 4 ~ W 0 • - I -201 • T -30 2'511 1911'512212'51215 30 8 10 21 23 30 3 MAR Fig. 2. Minutes APR 11'ol1 912'21 7 8 12 21 27 MAY before or after sunset that radio-marked JUN male wild turkeys went to roost. �215 shared a portion of its activity center with the other 6 radio-marked males, 4 had portions of their activity centers in common with 5 other males, and 2 had overlapping activity centers with 2 other males. Two (N = 23) or more (N = 8) radio-marked males were observed together during 31 (33%) of the 94 valid gobbling indicies. Radio-marked males were also observed with unmarked males on numerous occasions indicating additional overlap in ranges. Table 9. Dispersal, home range, activity center, and daily movements of adult male wild turkeys during the breeding season, 1986. TX# 0778 0502 0441 0555 0619 0837 0679 Dispersala (km) 4.2 3.0 6.0 2.3 1.9 5.8 2.7 Home rangeb (ha) 1,772 1,192 1,156 2,333 722 1,764 1,845 (22) (32) (15) (33) (27) (14) (34) Activity centerC (ha) Daily movementd (m) 569 290 471 468 231 256 620 2,010 (4) 1,360 (9) 1,888 (3) 2,914 (7) 1,136 (4) -__e 2,907 (3) aDistance from winter trap site to middle of the breeding activity center. bArea occupied from 26 March to 25 .June. Total number of observations given within the parentheses. cArea containing >70% of all observations between 26 March and 25 June. dMean distance between the morning and evening roost site on the same day. Sample sizes are in parentheses. e~ = 1, insufficient sample. Females moved longer distances (9.6 ±4.3 km) from wintering to breeding areas and occupied smaller home range (653 ±397 ha) than males (Table 10). Activity centers were not delineated for females because they were not relocated as often as males. Daily movements between roosting sites averaged 1498±1024 m (N = 10). Only 1 of 4 subadult hens was successfully tracked to her breeding range. She traveled 4.5 km and occupied a home range of 479 ha. Mortality Twenty-two radios were recovered between 1 March and 18 July; 21 from hens and 1 from a male. Contact was lost with TXl124 (subadu1t female) in mid-April and TX0837 (adult male) in early Mayas a result of their radios malfunctioning. Sixteen recoveries were classified as from birds (including the 1 male) that died of natural causes, 5 were from birds that possibly slipped their transmitters, and 1 was from a bird that was apparently suffering from capture myopathy. The critical mortality period for hens was late March-early April. Twelve of the 15 suspected natural hen mortalities occurred between 1 March and 10 April; 4 occurred between 1 and 15 March, 6 occurred between 16 and 31 March, and 4 occurred between 1 and 10 April. Two hens died in early May prior to onset of incubation and 1 hen was killed on �216 Table 10. Winter to breeding area movements and breeding home range of female wild turkeys, 1986. Txt;a Dispersal (km)b 0457 0639 0880 1100 1159 1144 1178 1241 1218 0158 1282 0261 0860 1425 0372 0342 0306 1319 1750 0399 0139 0577 0661 0798 1201 1259 6.9* 4.0* 14.8* 5.4* 6.1* 6.9* 5.0* 8.8* 3.9* 9.4* 14.9* 14.4* 16.1* 8.6* 9.3* 6.6** 3.4** 4.5** 8.5** 13.1*** 12.8*** 9.2*** 11. 7*** 18.1*** 15.5*** 12.1*** Home Range (ha)c 982(7) 1,465(6) 901(7) 584(8) 300(7) 304(10) 643(15) 439(7) 1,391(6) 533(6) 620(7) 291(5) 479(10) 205(8) aA11 adult hens except TX1319. bDistance from winter trap site to nest (*), center of home range (**), or mid-May location (***). Mid-May locations were used for hens known to be on the breeding range that failed to nest or were not relocated enough (~5) to delineate a home range. cArea occupied between 15 April and 15 June for hens relocated ~ times. Sample sizes are in parentheses. �217 the nest in early- June. <~'Total estimated mortality of hens from late winter (1 Mar) through the nesting period (30 Jun) was 35% (15/43). No males died during this period. The only documented mortality of a male occurred in mid-July. Four of the 5 transmitters thought to have been dropped (tai1-mounts, N = 4) or slipped (poncho mounts, N = 1) were recovered in late March during the critical mortality period for hens. All 4 tai1-mounts were still attached to the 2 central retricies when recovered. No other feathers.or body parts were found near any of the transmitters to indicate the birds were killed, nor had the transmitters sustained any damage (i.e., tooth marks, scratches, etc.). It is possible the tail feathers (calamus) were damaged when the radios were attached, causing the birds to molt them prematurely. Three instrumented hens were recovered <500 m from the trap site within 10 days of when they were trapped. Two had been killed (or possibly scavenged) by mammalian predators and 1 was captured alive by hand. The captured bird could not fly and fell over when attempting to run. The bird was scarificed and necropsied at the Colorado State University Diagnostic Lab. It weighed 1.1 kg less (4.2 vs. 3.1 kg) than when initially captured. Gross lesions, indicative of capture myopathy (Spraker et al. 1987), were found in the thigh muscles. No lesions were found in the wing muscles. Although the 2 hens killed near the trap site were classified as natural mortalities, they too may have suffered from capture myopathy, thus, predisposing them to predation. Nesting Five (20%) of 25 hens surv1v1ng into the nesting season nested successfully; whereas, 9 attempted to nest but failed, 2 were suspected of being in the process of laying when they were killed, 1 was depredated on the nest, and 8 either did not attempt to nest or abandoned their clutch prior to onset of incubation. One renesting attempt was documented. This hen abandoned her first clutch of 14 eggs on 22 May and was relocated on a second clutch of 12 eggs on 17 June. The second clutch hatched on 11 July. Clutch size of first nesting attempts averaged 11.7 eggs (range, 10-14). Fifty of 57 (88%) eggs in successful nests hatched. The earliest known date for onset of incubation for a first nest was 7 May and the latest was 6 June. Only 3 hens initiated incubation prior to the conclusion of the spring hunting season (18 May), 8 started between 19 and 31 May, and 2 started between 6 and 18 June (including the renest). Two of the hens starting before the season ended didn't do so until about 15 May. The peak period for onset of incubation was 15-31 May. This coincides with the second peak in gobbling activity (13-20 May). Three hens were disturbed during the early stages (1st week) of incubation. All 3 abandoned their nest and only 1 renested. Hunter Compliance - Permit System A total of 4,698 permits was issued for the spring season and 4,788 licenses were sold. Hunter compliance with the spring permit requirement was therefore 98%. Questionnaires were sent to all permit holders by 1 June 1986 of which 59% were returned. A fo110wup questionnaire was mailed to the non-respondents on 24 June 1986 and 578 additional responses were received. The combined return rate for the first and second mailing was 72% (Table 11). �218 There were more permits l~sued (1,819) for the fall season than there were licenses sold (809). The reason being that unsuccessful spring hunters who wanted to hunt the fall season were not required to purchase another license, but they were required to obtain a fall permit. Consequently, total license sales and total permits issued were not directly comparable in evaluating hunter compliance. Instead, hunter compliance was measured as the proportion of new license buyers who also obtained a permit. The estimate was 94% (759 of 809). Questionnaires were sent to the 1,819 fall permit holders on 17 October 1986. Sixty percent were returned by 17 November 1986. On 20 November 1986, a followup questionnaire was sent to the 676 non-respondents and 231 replied. Seventy-three percent of the fall hunters responded to the questionnaire (Table 11). Table 11. Response to the 1986 spring and fall turkey harvest survey. Surveys mailed Surveys returned Percent return Non-deliverable 1st Spring 2nd Total 4,698 2,787 59 98 1,902 578 30 21 4,698a 3,365 72 119 1st 1,819 1,097 60 24 Fall 2nd 676 231 34 21 Total 1,8l9a 1,328 73 45 aTotal hunters sampled. The permit system allowed quick access to the names and addresses of hunters so the survey could be conducted immediately following each season. The other option was having to wait for license agents to return license stubs containing hunter's names and addresses. Although agents have deadlines for returning licenses stubs, they do not always comply. In some cases, licenses stubs for the spring turkey season are not received by the CDOW until August. In 1984, an attempt was made to survey spring hunters from names on license stubs. Because of the time required to obtain all the license stubs from the agents, the first mailing was delayed until 23 July. By bypassing the license agents and issuing permits at DOW offices, the first mailing of the 1986 spring survey was out by 1 June, just 2 weeks after the season closed. Prior to 1984, the CDOW conducted 1 turkey harvest survey annually. The surveys were mailed in November and sampled about 50% of the hunters with no attempt to stratify by spring only, fall only, or spring and fall hunters. Incomplete addresses recorded on the license stubs plus the time lag between when the license was issued and when the survey was mailed resulted in a high percentage of non-deliverable surveys; i.e., 6-10% using license stubs vs. 2% using permits. Surprisingly, response rates did not differ between surveys conducted using names taken from license stubs or from permits. Since the permit system is only temporary, a 2-license system is recommended as an alternative means of enhancing the survey process. Hunters would be required to purchase a license for each season (spring and/or fall) they wanted to hunt regardless if they were unsuccessful in the spring season. The �Table 12. Spring turkey harvest and hunter activity, 1986. DescriEtive statistic 1st No. in sample No. hunters % hunters No. hunters observing turkebs % hunters observing turkeys No. successful hunters (harvest) % successful huntersb No. hunter days Days/hunterb Crippling loss % crippling loss Total kill 2,787 2,474 89 1,610 65 377 15 8,955 3.6 87 19 464 Mailings 2nd 578 514 89 302 59 56 11 1,772 3.6 19 25 75 Both 3,365 2,988 89 1,912 64 433 14 10,727 3.6 106 20 539 Projected for Totals 1,333a 1,186 89 700 59 130 11 4,269 3.6 32 25 162 4,698 4,174 89 2,612 63 563 13 14,996 3.6 138 20 701 -aNon-respondents. bBased only on those license holders who actually hunted. ',_"...N f--' 1.0 �N N Table 13. o Fall turkey harvest and hunter activity, 1986. -, -, DescriEtive statistic 1st No. in sample No. hunters % hunters No. hunters observing turkebs % hunters observing turkeys No. successful hunters (harvest) % successful huntersb No. hunter days Days/hunterb Crippling loss % crippling loss Total kill 1,097 888 81 489 55 209 24 2,240 2.5 28 i2 237 Mailings 2nd 231 168 73 89 53 33 20 432 2.6 7 17 40 aNon-respondents. bBased only on those license holders who actually hunted. Both 1,328 1,056 80 578 55 242 23 2,672 2.5 35 13 277 Projected for 491a 358 73 190 53 71 20 931 2.6 12 17 83 Totals 1,819 1,414 78 768 54 313 22 3,603 2.5 47 14 360 �221 problem would then be insuring license agents abide by the deadlines for returning license stubs. Perhaps fines could be levied against agents not complying with deadlines or chronic abusers could have their licensing privilege revoked. Hunter Activity and Harvest - Questionnaire Projected estimates for the spring season indicated that 4,174 hunters harvested 563 turkeys for a success rate of 13% (Table 12).- Total kill, including a reported crippling loss of 20%, was estimated at 701 birds. Spring hunters averaged 3.6 days afield over the course of the 30-day season from 19 April to 18 May. There were 66% fewer hunters that participated in the fall season (1,414 hunters) and 44% fewer birds (313) were harvested (Table 13). However, hunter success (22%) was higher and crippling loss (14%) was lower. Fall hunters spent 2.6 days afield or about one day less than spring hunters, but fall hunters had only 16 days (20 Sep-5 Oct) of hunting which included just 3 weekends. The spring season lasted 14 days longer, allowing for 2 additional weekends of hunting opportunity. Most hunting pressure and harvest occurred opening weekend during both seasons (Table 14). Pressure and harvest did not change substantially over the remainder of the fall season. There was an increase in pressure on weekends during the spring season, but this was not always associated with a corresponding increase in harvest. The Southeast Region accounted for over 80% of the spring and fall harvest (Tables 15 and 16). Las Animas County was the leading harvest area with 23 and 31% of the spring and fall harvest, respectively. Public land supported 54% of the spring hunting pressure and 52% of the fall pressure, but produced only 45 (spring) and 36% (fall) of the harvest (Table 17). Table 14. Chronological distribution of hunting pressure and harvest during the 1986 spring and fall wild turkey seasons.a Date 1st 1st 2nd 2nd 3rd 3rd 4th 4th 5th weekend week weekend week weekend week weekend week weekend Totals Sprin~ Hunter da~s N % Fall Harvest N % Hunter da~s % N Harvest N % 3,354 1,215 1,652 650 1,201 550 884 352 619 32 12 16 6 12 5 8 3 6 172 61 34 26 19 20 23 10 4 47 17 9 7 5 5 6 3 1 1,018 398 547 285 424 38 15 20 11 16 107 35 36 24 34 45 15 15 10 15 10,477 100 369 100 2,672 100 236 100 aBased on hunter days and harvest that could be assigned to specific time periods. �222 Table 15. Count~ Las Animas Custer Fremont Huerfano Pueblo Baca Larimer Mesa Morgan Yuma Kit Carson Teller Archuleta Jefferson Chaffee Douglas Logan Otero E1 Paso Bent Boulder Clear Creek Costilla Garfield Montrose Adams Gilpin LaP1ata Lincoln Park Weld Totals Unknown Wild turkey-harvest by county, SGMU, and Region, spring 1986 N % 93 50 47 38 31 13 12 11 11 11 8 7 7 7 6 6 23 13 12 10 8 3 3 3 3 3 2 2 2 2 2 2 2 2 1 1 1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 6 6 5 4 4 2 2 2 2 1 1 1 1 1 1 397 36 100 SGMU N 84 80 68 79 81 42 70 32 36 50 44 88 78 6 58 82 56 10 30 52 76 60 83 86 90 62 64 48 157 37 34 25 17 16 16 11 11 11 9 8 8 6 6 6 5 3 3 3 3 2 2 2 1 1 1 1 405 28 % Region 39 9 8 6 4 4 4 3 3 3 2 2 2 1 1 1 1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 SE NE Central SW NW 100 N % 321 30 21 13 12 81 8 5 3 3 397 36 100 �223 Table 16. Wild turkey harvest by county, SGMU, and Region, Fall 1986. N % Las Animas Fremont Pueblo Custer Huerfano Larimer Mesa Archuleta Kit Carson Teller Douglas E1 Paso Bent Jefferson Otero Baca Clear Creek Gilpin Boulder Yuma 71 29 27 25 21 10 10 6 5 5 4 3 3 2 2 2 2 1 1 1 31 13 12 11 9 4 4 3 2 2 2 1 1 1 1 1 1 <1 <1 <1 Totals Unknowns 230 9 100 Count;¥: SGMU N 84 80 79 70 37 68 58 42 88 81 50 52 76 60 10 83 82 56 30 III 39 11 9 8 8 7 6 6 4 4 4 4 2 2 2 1 1 1 230 9 % Region 48 17 5 4 4 4 3 3 3 2 2 2 2 1 1 <1 <1 <1 SE NE Central NW SW 100 N % 194 11 10 10 6 84 5 4 4 3 230 "9 100 Table 17. Distribution of hunting pressure and harvest by land status during the 1986 spring and fall wild turkey seasons. Fall Sprin~ Land status Public Private Both Totals Hunters % N Harvest N Harvest Hunters % N % N % 1,593 953 417 54 32 14 161 194 45 55 541 410 99 52 39 9 81 146 36 64 2,963 100 355 100 1,050 100 227 100 �224 Hunter success averaged 28% on the spring limited permit areas, exceeding the statewide success rate of 13% on all but 1 area (Table 18). The 4 eastern plains limited permit areas (units 6, 36, 42, and 44) supported the highest hunter success rate (38%). Of significance is that wild turkeys did not exist in any of these units until Rio Grande's were released there in 1981. It was not advantageous in terms of success to hunt on a limited permit area during the fall season as only 38 of 220 (17%) fall limited permit holders were successful (Table 18). Unit 42 was the only fall limited permit area where hunter success surpassed the statewide average. It was a definite advantage to hunt on private (spring = 20% success, fall = 35% success) vs. public land (spring = 10% success, fall = 15% success) in both seasons. Table 18. Hunter success on limited permit areas during the 1986 spring and fall wild turkey seasons. Fall SprinB Permit areas Permits issued Lake Dorothey Beaver-Skagway Unit 42 Unit 44 Unit 6 and 36 Spanish Peaks 75 20 40 20 50 __b Harvest Success (%) Permits issued Harvest Success (%) 13 2 16 9 17 17 10 40 45 36 75 30 25 __a __a 15 4 8 20 13 32 90 11 12 aClosed to fall hunting. bNo restrictions on numbers of hunters during the spring season. Seventy-four percent of the birds harvested during the spring season were taken before noon; 56% were harvested between 0500 and 0900 hours (Table 19). Only 20% were not associated with other birds at the time of harvest (Table 20). Most (57%) were taken from flocks comprised of 1-5 birds. The remaining 23% were taken from flocks with 6 or more birds. There was no indication that hunters encountered smaller flocks as the season progressed. Hens undoubtedly were present in the larger flocks, which suggests they were still receptive to males and not incubating. This contention is supported by the fact that in 37 observations of adult males from 19 April to 18 May, 26 (70%) of the males were accompanied by females. Wing Collection Program Limitations - The validity of any population indice calculated from harvest samples is dependent upon the assumption that the different age and sex classes are harvested in proportion to their occurrence in the population. Long-term population and harvest data are often necessary to test this assumption. Such data are not available for the Merriam's wild turkey in Colorado or elsewhere throughout its range. In addition, if wings are collected over a broad geographic area, they may not accurately reflect the characteristics of local populations. These limitations do not preclude the �225 Table 19. Distrihution-6f harvest by time period during the 1986 spring and fall wild turkey seasons. Time period 0400 0500 0600 0700 0800 0900 1000 1100 1200 1300 1400 1500 1600 1700 1800 1900 - 19-26 A:er (MST)a % H 0459 0559 0659 0759 0859 0959 1059 1159 1259 1359 1459 1559 1659 1759 1859 1959 Totals Fall 20 Sept-5 Oct (MDT)6 S:erinB 27 AEr-18 Max (MDT)6 N % N % 8 48 39 39 18 15 11 11 5 5 7 11 14 20 3 0 3 19 15 15 8 6 4 4 2 2 3 4 6 8 1 0 1 11 22 21 18 8 12 3 5 2 0 4 3 11 8 3 1 8 17 16 14 6 9 2 4 2 0 3 2 8 6 2 0 0 11 38 34 29 18 15 8 10 9 17 15 28 7 0 0 0 5 16 14 12 8 6 3 4 4 7 6 12 3 0 254 100 132 100 239 100 aMountain standard time. bMountain daylight time. Table 20. Size of flocks from which turkeys were harvested during the 1986 spring season. Flock size 0 1-5 6-10 >10 N % N % N % N % 19-24 Apr 25-30 Apr 1-6 May 7-12 May 13-18 May 47 9 8 13 4 19 17 26 30 21 137 34 17 22 12 56 63 57 51 63 43 8 3 2 3 17 15 10 5 16 19 3 2 6 0 8 5 7 14 0 Totals 81 20 222 57 59 15 30 8 Date of harvest �226 use of wing data as a use!ul tool in formulating management strategies. Biologists must be aware of them and use caution in interpreting the data. Compliance - Of the 433 successful spring hunters who responded to the questionnaire, 343 (79%) said they returned a wing. However, 390 wings were processed, meaning 47 successful hunters who did not respond to the questionnaire still returned a wing. The wing collection program sampled 69% (390/563) of the spring harvest. A total of 244 wings was collected from the fall wing survey representing 78% of the estimated fall harvest. Eight-five percent (206) of the successful hunters who responded to the fall questionnaire indicated they returned a wing and 38 successful hunters not responding also returned a wing. Subspecies Composition - The Merriam's wild turkey was the dominant subspecies indentified from inspection of wings. It comprised 91 and 97% of the spring (Table 21) and fall (Table 22) samples, respectively, and was harvested in 26 of Colorado's 63 counties. Rio Grande's were taken in 6 counties. Fremont was the only county where both subspecies were harvested. The 35 Rio Grande wings collected during the spring season originated from the following areas: 12 from Unit 42, 6 from Unit 44, 10 from Unit 6, 4 from Unit 36, and 3 from Fremont County near Florence. Three of these areas (Units 6, 36, and 44) were closed to hunting in the fall. The result was that only 8 Rio Grande wings were identified in the fall sample of 244 wings: 6 from Unit 42 and 2 from Fremont County near Florence. Table 21. Sex, age, and subspecies composition of the 1986 spring harvest based on wing analyses. Males SubsEecies Merriam's Rio Grande Totals Subadult 80 (23)a 20 (57) 100 (26) Adult Females Adult Subadult Totals 264 (76) 15 (43) 0 0 5 (1) 0 349b 35 279 (73) 0 5 (1) 384 apercent. bSix additional Merriam's wings were processed but could not be classified to age or sex bringing the total wings collected to 390. Sex Composition - Five (2%) wings examined from the spring sample were from females (Table 21). All were Merriam's and all were assumed to be from legal hens; i.e., bearded hens. Two percent of the total spring harvest (~= 11) is probably a reasonable estimate of the legal harvest of hens. At this level of harvest, spring hunting should have minimal impact on the female segment of the population. �227 Examination of wings (Merriam's only) originating from the fall season revealed the sex ratio of adults and subadults combined favored females (1.5 females: 1 male), while the sex ratio of juveniles favored males (1.6 males 1 female). The biological implications of these ratios are difficult to interpret without knowledge of the population structure. Hoffman (1962) reported winter counts of 1.6 females: 1 male along the Front Range in southeastern Colorado. The sex ratio of 171 turkeys harvested on the Uncompahgre Plateau during the fall seasons of 1961 and 1963-67 was 1.7 females: 1 male (Myers 1973). Both ratios were based on combined samples of adults, subadults, and juveniles. A comparable ratio using the 1986 fall harvest data would be 1.1 males: 1 female. Myer's data, broken down by age class, shows that the sex ratio of adults and subadults (2.5 females: 1 male) and that of juveniles (1.5 females 1 male) still favored females. Sex ratios should favor females in a promiscuous species such as the wild turkey where most hens breed but only a portion of the males do so. The observed sex ratio of adults and subadults in the fall sample supports this argument but the sex ratio of juveniles does not. Reasons for the preponderance of males in the juvenile segment of the harvest 'are unknown but may be related to (1) differential vulnerability of juvenile males to hunting, (2) better survival of males from hatching until the fall season, or (3) biases in the methods for distinguishing sexes of fall-harvested juveniles based on the coloration of breast feathers. Age Composition - Seventy-six percent of the spring harvest of Merriam's wild turkeys were adults compared to only 43% for Rio Grande's. Rio Grande populations in Colorado are the result of recent (since 1981) introductions. Available evidence suggest these populations are still increasing and expanding into unoccupied ranges, while populations of Merriam's wild turkeys have remained stable or declined. The age compositiion of the Rio Grande harvest reflects good production, survival, and recruitment of young birds, attributes indicative of an increasing population. Estimates of nesting success based on observations of radio-marked Rio Grande (58% success) (J. Schmutz, unpubl. data) and Merriam's (20% success) hens further suggest differences in reproductive output that should be reflected by harvest samples. Subadult males in the increasing Rio Grande populations may not be suppressed by mature birds from participating in breeding activities. If so, then subadults should be equally, if not more vulnerable due to their inexperience, to being lured in by the calling tactics of spring hunters. The following discussion regarding the age composition of the fall harvest pertains to the Merriam's wild turkey. Samples of Rio Grande wings from the fall season were inadequate for meaningful interpretation. Subadults comprised just 5% of the fall harvest including juveniles and 11% excluding juveniles (Table 22). Biologically, this could be interpreted as poor production in 1985 and subsequent low recruitment into the 1986 population. However, another factor contributing to the deficiency of subadults is associated with the onset of primary molt and whether the key feathers (primaries IX and X) for separating adults and subadults are still present at the time wings are collected. If these feathers have already molted, then subadults cannot be distinguished from adults. This was the case as 92% of the adult males and 46% of the adult females had completed their primary molt by the fall season. The problem of separating adults and �N N 00 Tables 22. Age, sex, and subspecies composition of the 1986 fall wild turkey harvest based on wing analyses.a Adults Females Males Subspecies Merriam's Rio Grande's Total Subadults Females Males Total Juveniles Females Males N % N % N % N % N % N % N % N % 38 1 42 25 52 3 58 75 90 4 40 57 2 0 18 0 9 0 82 0 11 0 5 0 75 1 61 33 47 2 39 67 Total N 122 3 % 55 43 Sample size 223 7 Poults/ hen 2.0 1.0 aA total of 244 wings was processed of which 230 could be classified to age and sex. One Rio Grande wing and 4 Merriam's wings could not be classified to age or sex. Six Merriam's wings were classified as juveniles but could not be identified to sex and 3 Merriam's wings classified to sex (1 male, 2 females), could not be assigned to an age class. blncludes adult and subadult hens. �229 subadults was compounded:~or males because they start and finish their primary molt sooner than females. A similar problem was confronted in interpreting wing data derived from harvest samples of blue grouse (Dendragapus obscurus) (Hoffman 1985). Misidentification of subadults is not a problem in the spring when birds are just beginning their primary molt. The percent juveniles in the harvest (55%) and the ratio (2.0:1) of juveniles to adult and subadult females were used as indicies to productivity. The 2 indices suggested only fair production of young in 1986. lhis was partially attributed to low nesting success. Nesting success, based on wing molt patterns (Hoffman 1985:6), was estimated at 35%. Adults (36%) were only slightly more successful than subadults (30%), although the small sample (N = 11) of subadults may not be representative of this age class. The overallinterpretation of fair production is somewhat better than indicated by field observations of Merriams wild turkeys in southeastern Colorado. Here, only 5 (20%) of 25 radio-marked hens surviving into the nesting season nested successfully. Of the 15 unsuccessful nesters, only 1 attempted to renest. Despite low nesting success, juveniles still comprised 55% of the harvest sample. This may be due to a greater vulnerability of poults to hunting or that low nesting success does not necessarily equate to poor production and a resulting low percentage of young in the harvest. Turkeys may compensate for low nesting success by having large clutches, high hatching success, and good survival of young birds. Hatching Dates - Wings from 93 Merriam's poults were classified to age following Knoder (1959) and McGuire (1964). Hatching dates were then calculated by backdating from the date of harvest. The peak of hatch ranged over a 3-week period from 19 May to 8 June 1986 with the 2 methods being in close agreement (Table 23). These data indicate that the spring season was correctly timed to coincide with the peak of incubation, which would have occurred 28 days prior to the peak of hatch or from 22 April-12 May. Table 23. Estimated hatching dates for Merriam's wild turkeys in Colorado based on wing analyses, 1986. Method Time Eeriod McGuire (1964) N hatchin8 % Knoder (1959) N hatching % 14-20 Apr 21-27 Apr 28 Apr-4 May 5-11 May 12-18 May 19-25 May 26 May-l Jun 2-8 Jun 9-15 Jun l6-22Jun 1 1 6 15 13 14 21 15 3 4 1 1 7 16 14 15 23 16 3 4 1 1 5 14 14 17 19 14 4 4 1 1 6 15 15 18 20 15 4 4 Totals 93 100 93 100 �230 It was surprising that the methods of Knoder (1959) and McGuire (1964) agreed so closely for age classification of Merriams wild turkeys since they were developed from measurements of Eastern (Knoder 1959) and Rio Grande (McGuire 1964) wild turkeys. Furthermore, McGuire (1964) found that his technique incorrectly classified Eastern wild turkeys because of significant differences in primary measurements between the 2 subspecies. Primary measurements of Merriam's wild turkeys may be intermediate between those of Eastern's and Rio Grande's, thus, either method would produce the same approximate, but not necessarily accurate, estimate of age. Onset of nesting by Merriam's wild turkeys near Trinidad did not agree with the chronology of nesting activities as revealed by wing analyses. Only 3 radio-marked Merriams hens initiated incubation prior to 18 May, 8 started between 18-31 May, and 2 between 6-18 June. This places the peak of hatch between 6-28 June almost 3 weeks later than indicated by wing analyses and suggests the spring season was incorrectly timed. Until more data become available on the development of the postjuvena1 primaries of known-age Merriam's poults, the reliability of Knoder's and McGuire's methods for age classification of Merriam's wild turkeys remains questionable. Although no Rio Grande poults were included in the wing sample, nesting data were collected on Rio Grandes in conjunction with a habitat study in northeastern Colorado (Schmutz 1987). The incubation period of all (12) known nesting hens along the South Platte River overlapped the spring season and therefore agreed with the findings of the wing survey. What these results indicate is that there are geographic differences in timing of nesting within the state. LITERATURE CITED Amstrup, S. C. 1980. 214-217. A radio-collar for game birds. J. Wi1d1. Manage. 44: Bailey, R. W., and K. T. Rine11. 1967. Events in the turkey year. Pages 93-111 in O. H. Hewitt, ed. The wild turkey and its management. The Wi1d1. Soc., Washington, D.C. ____~~--' D. Dennett Jr., H. Gore, J. Pack, R. Simpson, and G. Wright. 1980. Basic considerations and general recommendations for trapping the wild turkey. Proc. Nat1. Wild Turkey Symp. 4:10-23. Bevill, W. V., Jr. 1973. Some factors influencing gobbling activity among wild turkeys. Proc. Southeast. Assoc. Game and Fish Comm. 27:62-73. 1975. Setting spring gobbler seasons by timing peak gobbling. Proc. Nat1. Wild Turkey Symp. 3:198-204. Bray, O. E., and G. W. Corner. 1972. A tail clip for attaching transmitters to birds. J. Wi1d1. Manage. 36:640-642. Burget, M. L. 1957. The wild turkey in Colorado. Fed. Aid Proj. W-39-R. 68 pp. Colorado Game and Fish Dep. �231 Clover, M. R. 199-201. 1956. Sltigle gate deer trap. California Game and Fish J. 42: Colorado Division of Wildlife. 1984. Colorado Small Game, Furbearer and Varmint Harvest. Dep. Nat. Resour. Div. Wildl., Denver. 210 pp. Davis, J. R. 6-7. 1964. Spring weather and wild turkeys. Alabama Conserve 41(1): Dill, H. H., and W. H. Thornsberry. 1950. A cannon-projected net trap for capturing waterfowl. J. Wildl. Manage. 14:132-137. Glazener, W. C., A. L. Jackson, and M. L. Cox. 1964. turkey trap. J. Wildl. Manage. 28:280-287. The Texas drop-net Hoffman, D. M. 1962. The wild turkey in eastern Colorado. Game and Fish. Tech. Publ. 2. 49 pp. Colorado Dep. 1966. Merriam's turkey roost preferences on mountain ranges. Colorado Div. Wildl. Game Info. Leafl. 45. 6 pp. Colorado. 1968. Roosting sites and habits of Merriam's turkeys in J. Wildl. Manage. 32:859-866. 1973. Some effects of weather and timber management on Merriam's turkey in Colorado. Pages 263-271 in G. C. Sanderson and H. C. Schultz, eds. Wild turkey management: current problems and programs. Univ. Missouri Press, Columbia. Hoffman, R. W. 1985. Blue grouse wing analyses: methodology and population inferences. Colorado Div. Wildl. Spec. Rep. 60. 21 pp. Knoder, E. 1959. An aging technique for juvenal wild turkeys based on the rate of primary feather moult and growth. Proc. Natl. Wild Turkey Symp. 1:159-176. Knowlton, F. F., E. D. Michael, and W. C. Glazener. 1964. A marking technique for field recognition of individual turkeys and deer. J. Wildl. Manage. 28:167-170. McGuire, R. D. 1964. Aging Rio Grande wild turkey poults, Meleagris gallopavo intermedia (Sennett), by primary feather length and general body characteristics. M.S. Thesis, Oklahoma State Univ., Stillwater. 54 pp. Mohr, C. O. 1947. Table of equivalent populations of North American small mammals. Am. MidI. Nat. 37:223-249. Myers, G. T. 1973. The wild turkey on the Uncompahgre Plateau. Colorado Div. Wildl. Fed. Aid Proj. W-37-R. 153 pp. Final Rep. Schmutz, J. A. 1987. Habitat use by Rio Grande wild turkeys in Northeast Colorado. Colo. Div. Wildl. Fed. Aid Proj. 01-03-045 (W-37-R) Job Prog. Rep. April 1987. In press. �, t 232 Spraker, T. R., W: J. Adrian, and W. R. Lance. 1987. Capture myopathy in wild turkeys CMeleagris gallopavo) following trapping, handling and transportation in Colorado. J. Wildl. Dis. 23:447-453. Williams, L. E., Jr. 1966. Capturing wild turkeys with alpha chloralose. Wildl. Manage. 30:50-56. , D. H. Austin, T. ------~-turkeys with oral drugs. J. E. Peoples, and R. W. Phillips. 1973. Capturing Pages 219-227 in G. C. Sanderson and H. C. Schultz, eds. Wild turkey management: current problems and programs. Univ. Missouri Press, Columbia. �Colorado Division 233 of Wildlife JOB PROGRESS REPORT State of Colorado --~~~~------------------------- 12 Work Plan Job Title: : Job 16 Habitat Use by Rio Grande Wild Turkeys in Northeast Colorado Period Covered: Author: Avian Research 01-03-045 (W-37-R) Project 01 January through 31 December 1986 Joel Schmutz Personnel: C. Braun, J. Corey, T. Davis, M. Etl, R. Hoffman, S. Steinart, Colorado Division of Wildlife; W. Andelt, J. Ratti, J. Schmutz, Colorado State University ABSTRACT Rio Grande wild turkey CMeleagris gallopavo intermedia) habitat use was studied in the South Platte River floodplain in northeast Colorado during January-December 1986. Eighteen hens were trapped and radiomarked during February. Yearling hens dispersed an average of 39 km and adult hens 9 km from their capture site to nest sites. Twelve of 13 hens monitored through the breeding season nested. Eggs in 7 nests hatched. Primary vegetation at 11 nests was western snowberry (Symphoricarpos occidentalis) while pepperweed (Lepidium latifolium) dominated at 2 nests. Nest sites were characterized by more shrubs, forbs, and understory cover, taller understories, and less grass than non-use sites. Broods moved an average of 742 m from the nest site to the brood home range. Brood home ranges varied from 14 to 62 ha in size. Sixty-five percent of brood locations were in riverbottom habitat, 22% in agricultural habitat, and 13% in the edge between these 2 habitat types. Microhabitat analyses of riverbottom locations showed nonrandom habitat use, but the interpretation was not clear. ��235 HABITAT USE OF RIO GRANDE WILD TURKEYS IN NORTHEAST COLORADO Joel Schmutz P. N. OBJECTIVES The principal objectives of this study are to (1) describe nest sites and (2) brood sites of Rio Grande wild turkeys in the South Platte River floodplain. SEGMENT OBJECTIVES 1. Capture wild turkeys while concentrated in winter flocks. Mark with leg bands and obtain morphological measurements from all captured wild turkeys. Attach radio transmitters to hens. 2. Monitor hen dispersal from winter flocks to nesting areas. 3. Measure vegetative and geographic parameters at wild turkey nests. Measure identical parameters at non-use sites. 4. Obtain data on clutch size and nesting success of wild turkeys. 5. Measure vegetative and geographic parameters at wild turkey brood sites. Measure identical parameters at non-use sites. 6. Analyze data and prepare annual Job Progress report. DESCRIPTION OF STUDY AREA Rio Grande wild turkeys presently exist along portions of the South Platte River floodplain in Logan, Morgan, and Washington counties in northeast Colorado (Fig. 1). Highest water levels occur in late May-early June, inundating the floodplain in years of high snowfall in the mountains. The riparian ecosystem, or river bottom, is 0-1.0 km wide and dominated by plains cottonwood (Populus sargentii). A relatively open canopy exists with mature cottonwoods being common. Box elder (Acer negundo), green ash (Fraxinus pennsylvanica), and Russian olive (Elaeagnus angustifolia) are present and increasing in frequency. Willows (Salix spp.) occur in more mesic areas. Western snowberry (Symphoricarpos occidentalis) is the dominant shrub, although shrub willows (Salix spp.), poison ivy (Toxicodendron radicans), and Indigo bush (Dalea sp.) are fairly common. Shrubs usually occur. in discrete patches. The herbaceous understory becomes quite dense during the summer in undisturbed river bottom areas. Common grasses include salt grass (Distichlis stricta), sand dropseed (Sporobolus cryptandrus), prairie cordgrass (Spartina pectinata), wheatgrass (Agropyron spp.), and cheatgrass brome (Bromus tectorum). Common forbs include pepperweed (Lepidium latifolium), ragweed (Ambrosia spp.), sunflower (Helianthus spp.), thistle (Cirsium sp.), and poison hemlock (Conium maculatum). Grazing occurs on much of the privately-owned river bottom. Intensive agriculture occurs on lands adjacent to the river bottom with corn, alfalfa, and wheat or cattle grazing being the most common uses. �N W (j\ DOWNSTREAM AND UPSTREAM BOUNDARIES ••• STERLING 0 OF WILD TURKEY STUDY AREA ) ~ o DENVER FORT MORGAN o Fig. 1. Rio Grande wild turkey study area along the South Platte River in northeast Colorado. �237 METHODS Wild Turkey Capture and Marking Wild turkey flocks were baited with shelled corn and oat hay. Trapping was with a drop-net following procedures described by Lange (1983). Age, sex, body weight, and length of primaries, carpal, and beard were measured on each captured bird. Blood samples were also obtained to test for presence of Mycoplasma spp. Age was classified from characteristics of.primaries IX and X. Aluminum leg bands were attached to all birds. Radio transmitters within 150.101-151.701 were attached to hens. One transmitter was solar powered; the rest were powered by lithium batteries. Transmitters were either tail-clip (26-29 g) or poncho (29-32 g) mounted. Nest Habitat Use Radio-marked hens were relocated at least once every 4 days between release and nesting. Approximate locations were plotted on 1:24000 topographic maps. Radio locations were obtained with a Telonics TDP-2 receiver and a 3-element Yagi antenna. Dispersal is defined as the distance between the last location of the entire winter flock and the nest site. Dispersal distances parallel the river rather than straight-line distances as wild turkeys were not relocated more than 200 m from the riparian edge. After nest initiation, incubating, radio-marked hens were approached to within 20 m and sites flagged so that nests could be located after hatching. Clutch size and fertility and hatching rates were determined from eggshell characteristics. Nesting success is defined as hatching >1 egg in a nest. Habitat parameters were measured on a 0.04-ha plot centered over each nest site and on 4 0.04-ha plots established in random distances (within 75 m) and directions from each nest. These sets of 4 plots are hereafter referred to as adjacent non-use (AN) plots. Parameters measured included: tree diameter at breast height (DBH), basal area, canopy cover, understory height, understory cover, life form occurrence (shrubs, forbs, grasses, and bare ground), identification of the 3 most common herbaceous species, and distances to habitat edge, nearest tree >30 cm DBH, nearest road, and nearest active human dwelling. Basal area was computed from DBH data gathered using a Biltmore stick. Canopy cover (measured with a densiometer) and understory height (measured with a pole in 5-cm graduations) were measured at plot centers and plot peripheries in the 4 cardinal directions. A vegetation profile board (Nudds 1977) was placed at plot centers and read from plot peripheries in the 4 cardinal directions to ascertain understory cover. Life form occurrence was measured on 1 randomly oriented, diameter-length transect (Canfield 1941). Distances were determined by pacing or from maps. Additional 0.04-ha plots were established at 2.5-km intervals throughout the study area. These plots, referred to as riparian non-use (RN) plots, were at random, perpendicular distances from the river with the constraint that they were within the river bottom habitat. The same parameters were measured as at nest and AN plots. �238 Brood Habitat Uaeiand Movements Brood hens were radiolocated and 0.04-ha habitat plots established at flush sites. At least 3 days elapsed between flushes of individual brood hens to avoid a human-disturbance bias of their habitat selection. Parameters measured were identical to nest habitat parameters. Brood habitat anlayses were split into 2 periods: early (0-3 weeks) and late (4-6 weeks) brood periods. Two sets of non-use plots were therefore required. Data from the nest RN plots were used for early brood non-use plots. These plots were later remeasured to account for phenological change and these data used for late brood non-use plots. Individual broods were radiolocated 0-4 times between flushes using dual 5-element Yagi antennae. The frequency of these locations varied due to logistic constraints and environmental factors that limited signal reception. Standard deviation (SD) of this antenna system was 1.50• Two or 3 receiving points were used and the bearings were entered into a computer program to calculate the location estimate (White and Garrott 1984). Locations were also plotted on maps or aerial photos to ascertain habitat type. Both visual and telemetry locations were distributed among 3 diurnal periods of equal length. Home ranges were calculated from both visual and telemetry locations. Two methods of calculation were used: the minimum area polygon (Mohr 1947) and the Jennrich and Turner (1969) 95% confidence ellipse. The former was used as it is the most commonly applied method in other wild turkey studies. The latter is used as it is also commonly used and is less biased by sample sizes. Habitat use data from brood flushes were not gathered beyond a poult age of 6 weeks as by this age brood movements in response to my approach became apparent. Hens were still monitored at least once per week until transmitter loss or failure. RESULTS AND DISCUSSION Capture and Marking Four winter flocks of wild turkeys were located in the South Platte River floodplain (Fig. 1). Trapping of 2 flocks was attempted with success only for the flock near Merino. Twenty-seven wild turkeys were trapped on 10 February. Nine were males (1 adult, 3 yearlings) and 18 were females (6 adults, 12 yearlings). Birds termed "yearlings" were only 8-9 months old; this term seems more appropriate than "juvenile". Yearling females = 3.43 kg) weighed less than adult females (~ = 3.92 kg, ~ = 0.002) (Table 1). ex All captured wild turkeys tested positive for an unidentified Mycoplasma species. Mycoplasma galliseptica is pathogenic and has been demonstrated to decrease reproduction in wild turkeys. Unidentified Mycoplasma species are likely non-virulent (Amundson 1985, pers. commun.). �239 Table 1. Wild turkeys captured at Merino, Colorado 10 February 1986. Band II Sex Age Weight (kg) Carpal (mm) Beard (mm) 101 103 104 110 111 118 119 123 125 M M M M M M Yrlg Yrlg Yrlg Yrlg Yrlg Yrlg Ad Yrlg Yrlg 5.5 5.0 5.1 5.4 6.1 6.5 8.0 4.8 5.7 455 445 435 465 455 476 515 458 455 35 22 lOa 40 45 79 230 20 48 102 105 106 107 108 109 112 113 114 115 116 117 120 121 122 124 126 F F F F F F F F F F F F F F F F F Yrlg Yrlg Ad Yrlg Yrlg Ad Yrlg Ad Yrlg Yrlg Ad Yrlg Ad Yrlg Yrlg Yrlg Yrlg 3.4 3.5 3.7 3.6 3.3 3.9 2.9 3.7 3.5 3.6 4.1 3.5 4.3 3.6 3.8 3.3 3.2 328 396 412 393 394 406 385 432 396 411 415 390 432 406 395 392 386 20 11 M M aDouble beard. bp = poncho, T Radio Frequencyb l50.702-P 151. 622-P 151. 483-P l50.l0l-T 150. 724-T l5l.300-P l50.l2l-T l5l.435-P l50.483-T 151. 36l-P 151. 342-P l50.324-T 151. 46l-P l50.240-T l50.402-P l50.l96-T l51.028-PS tailclip, PS = solar-powered poncho. Dispersal to Nesting Areas Yearling hens (39 km) dispersed farther than adult hens (9 km, P < 0.05) (Fig. 2). Although wild turkey nests were up to 110 km apart, 9 of 12 nests were within 1 km of another radio-marked hen's nest (Fig. 2). Although age-related dispersal is documented for many avian species, the mean magnitude observed here is larger than previously reported fo~ wild turkeys. There may be several reasons for these unusually long dispersals. The linear, heterogenous nature of the habitat would result in longer movements as compared to other wild turkey habitats if a hen's phenotype allows for a given amount of 'searching' for a suitable nesting location. Alternatively, the relatively low population density may allow for all hens to nest in optimal habitat. A hen may achieve more reproductive success by moving a long distance to optimal habitat than by moving a short distance to sub-optimal habitat. �tv .p- o ILIFF o STERLING A - NEST OF ADULT HEN o "\~~ Y - NEST OF YEARLING HEN F - WINTER FLOCK LOCATION BEFORE DISPERSAL yA A ~'\~ ~ •.\i> AA ~V~ A MERINO o F I 'II ~~~ sa WELDONA o o 5 10 15 KILOMETERS o BRUSH FORT MORGAN Fig. 2. Dispersal to nesting areas by wild turkey hens in northeastern Colorado, 1986. 20 25 30 �241 Nesting Success Twelve of 13 hens monitored through the nesting season attempted to nest. The 1 hen that did not nest was a yearling. Three of 5 adult hens and 4 of 8 yearling hens nested successfully (Tables 2, 3). Eggs in successful nests hatched from approximately 23 May to 17 June. Using a 28-day incubation and 10-15 day laying period (Bailey and Rinnell 1967), nests were initiated between 10 April and 10 May. All successful hens began incubation before the conclusion of the turkey hunting season (19 Apr-18 May). Nest initiation dates for unsuccessful hens were difficult to estimate, although all but hen # 151.301 were believed to have initiated nesting during the same peirod as successful hens. Hen # 151.301 began incubating an II-egg clutch (her first known nesting attempt) between 4 and 15 June. On 13 July, an unmarked hen was observed with a young brood «1 week of age) indicating she also began incubation in early June. Table 2. Successful wild turkey nests in northeast Colorado, .1986. Hen Age 1.651 1.435 1.46la 0.196 1.028 1.701 1.621a Ad Ad Ad Yrlg Yrlg Yrlg Yrlg Date eggs hatched 23 28 05 26 27 29 17 N total eggs May May Jun May May May Jun Totals -+ x _ SD 14 14 10 11 8 10 11 12 14 7 11 8 10 10 78 72 (92%)b 11.1 ± 2.19 N hatched eggs N fertile eggs 10.3 ± 2.36 11 13 5 8 8 10 10 65 (83%)b 9.3 ± 2.56 aprobable 2nd nesting attempt. bpercent of total eggs. Table 3. Hen Unsuccessful wild turkey nests in northeast Colorado, 1986. Age 1.342 1.301 Ad Ad 1.621 0.121 1.402 0.240 Yrlg Yrlg. Yrlg Yrlg N eggs 7 11 1 5 9 7 Nest fate Probable skunk predation of eggs, hen found dead Probable observer-induced abandonment, followed by skunk predation Abandoned (successfully renested elsewhere) Hen survived attack on nest, abandoned Hen survived attack on nest, abandoned Entire clutch infertile, full term incubation �242 Three of 6 unsuccessfully nesting hens were attacked while on the nest (Table 3). Although predator species are unknown, vegetative cover at 2 of these nests seemed to preclude avian predators. The high rate of nesting attempts (5 of 5 adults, 7 of 8 yearlings), although possibly an artifact of small sample size, is greater than previously reported. The rate of yearling nesting attempts is particularly noteworthy as previous research has found little nesting by this age class (Reagan and Morgan 1980). Yearling nesting may occur more frequently in low population densities either through greater available nesting habitat or less behavioral inhibition from intraspecific interaction. Nesting Habitat Of all nests, 11 of 13 were in clumps of snowberry and 2 were in dense pepperweed stands. Pepperweed and sandbar willow (Salix interior) were minor vegetative components at 5 of the nests in snowberry clumps. All nests were in ungrazed riparian areas and 8 were on areas administered and/or managed by the Colorado Division of Wildlife. Nonrandom nest habitat use was indicated from variables measured at nest and non-use sites. Shrubs were more abundant and grasses less so at nest sites than at all non-use sites (Table 4). Understory cover «1 m) was greater at nest sites than non-use sites, although this significance was not unexpected as understory cover was correlated with the amount of shrubs (0-0.5 m, r = 0.35, P = 0.01; 0.5-1.0 m, r = 0.39, P = 0.006). Forbs were generally more abundant at nest sites than-non-use sites and this may have contributed (r = 0.3, ~ = 0.03) to the signficant1y taller understory at nest sites. There were no significant differences between nest and non-use sites for variables associated with the overstory. Within the understory, snowberry was the dominant woody shrub. In mid-April when most hens selected their nest sites, forbs and grasses were just beginning growth and, thus, provided little cover. The nest data analyzed here were measured after eggs had hatched and the understory grown. This unavoidable 6-7 week delay in data collection may have resulted in an overestimate of the nonrandom use of forbs (current year's growth). Also, the potential importance of residual forbs in providing nesting cover may be obscured. Comparison of AN and RN plots allowed for analysis of wild on a slightly larger scale. Vegetation in the vicinity of had more forbs, taller understories, less small trees, and dispersion of large trees than vegetation found throughout plots. turkey habitat use nests (AN plots) a more homogenous the study area (RN) Adults vs. Year1ings.--There was no significant difference between nests of adult and yearling hens for the vegetative parameters measured, although small sample sizes may have obscured any real differences (Table 5). There was a trend for nests of adult hens to have a taller understory, more shrubs, and less forbs than those of yearlings. �Table 4. Vegetative parameters at wild turkey nests and non-use plots in northeast Colorado, 1986. - x RN plots (N = 19) AN plots (N = 52) Nests (N = 13) SD x SD E_a x pb SD ~c Canopy cover, % 27 33 21 23 0.87 30 34 0.56 0.5,3 Understory cover, % 0-0.5 m 0.6-1.0 m 1.1-2.0 m 97 71 9 1 19 7 66 33 8 31 30 11 * * 0.17 34 15 9 30 18 9 * * 0.54 * 0.04 0.52 8 7 6 0 6 6 5 1 2 7 11 1 4 6 6 2 * 0.96 0.02 0.55 2 4 12 3 4 5 7 4 0.004 0.09 0.02 0.07 0.46 0.02 0.48 0.05 107 78 36 34 61 58 42 24 0.04 * 28 33 26 18 * * * * 8 14 12 0.68 23 31 0.38 0.07 0.34 0.54 1.0 0.12 0.92 0.45 0.65 0.74 Life form, m/22 m Shrubs Forbs Grasses Bare Ground Understory height, cm Plot center Total plot Distance to large (>30 em DBH) tree, m Basal area, m2/ha 0-25 em DBH 26-45 cm DBH >45 em DBH Totals 13 ' 0.6 1.9 3.4 1.4 4.1 7.9 0.5 2.6 3.5 1.2 4.9 6.1 0.70 0.28 0.48 2.3 2.2 4.1 6.0 9.4 6.7 8.8 0.76 8.6 5.4 . 3.1 9.5 11.8 I aprobabi1ity level for Nests vs. AN. bprobabi1ity level for Nests vs. RN. eprobabi1ity level for AN vs. RN. *p < 0.001. N -I:'- w �244 Table 5. Vegetative parameters at nests of adult and yearling wild turkeys in northeast Colorado, 1986. Adults (N = 5) x SD Yearlings (N = 8) x SD P Canopy cover, % 21 37 31 29 0.46 Understory cover, % 0-0.5 m 0.6-1.0 m 1.1-2.0 m 97 67 9 1 19 5 97 73 10 1 19 8 0.81 0.77 0.82 10 5 5 1 7 6 6 1 6 9 7 <1 4 5 4 1 0.34 0.21 0.38 0.37 114 98 50 41 102 65 21 16 0.94 0.16 15 9 12 7 0.71 Life form, m/22 m Shrubs Forbs Grasses Ground Understory height, cm Plot center Total plot Distance to large (>30 cm DBH) tree, m Basal area, m2/ha 0-25 cm DBH 26-45 cm DBH >45 cm DBH Totals 0.4 2.5 1.3 0.5 4.1 2.6 0.7 1.6 4.8 1.7 4.1 9.6 0.47 0.43 0.77 4.2 7.1 7.1 10.4 0.60 Successful vs. Unsuccessful Hens.--Nests of successful hens tended to have more shrubs and less forbs than those of unsuccessful hens (Table 6). Both nests within dense pepperweed stands were destroyed by predators. Preferential use of snowberry by successful hens was further indicated by greater amounts of understory cover than at unsuccessful nests. Canopy cover at successful nests was 40% whereas at unsuccessful nests canopy cover was 12%. Habitat Se1ection.--These data demonstrate nonrandom use of the riparian habitat by nesting wild turkeys. Dense understory, primarily in the form of snowberry, was used more than expected. The study design could not, however, elicit why this use pattern was observed. It was assumed that wild turkeys select nesting habitat based on vegetative structure with concealment from predators the primary selection criterion (e.g., Lazarus and Porter 1985). This assumption is not necessarily true as habitat selection in birds may have diverse causes. Habitat selection is probably genetically based in part, thus, observed use patterns may be dependent upon genealogy, not vegetation. Alternatively, environmental stimulus, such as parental teaching or dominance hierarchies, may affect habitat selection. Even if vegetative structure is �245 what a bird uses to assess habitat, food resources and not predator protection may be the most important criterion as is the case with yellow-headed blackbirds (Xanthocepha1us xanthocepha1us) (Orians 1980). For ground nesting, pre~ocia1 birds such as wild turkeys, food resources available to young are dependent upon the mobility of the young. Thus, hens may select nesting habitat primarily on its juxtaposition to good brood-rearing habitat. Table 6. Vegetative parameters at nests of successful and unsuccessful wild turkeys in northeast Colorado, 1986. Successful (N = 7) x SD Unsuccessful . (li = 6) x SD P Canopy cover, % 40 37 12 19 0.15 Understory cover, % 0-0.5 m 0.6-1.0 m 1.1-2.0 m 98 77 13 0 21 7 96 63 5 1 14 3 0.17 0.18 0.04 Life form, m/22 m Shrubs Forbs Grasses Ground 10 7 5 1 6 8 7 1 5 6 4 1 0.19 0.44 0.53 0.87 Understory height, cm Plot center Total plot Distance to large (>30 cm DBH) tree, m Basal area, m2/ha 0-25 cm DBH 26-45 cm DBH >45 cm DBH Totals 6 6· 5 1 111 76 46 39 102 78 15 22 0.74 0.50 13 9 13 7 0.96 0.9 3.3 2.3 1.8 5.2 3.7 0.2 0.3 4.8 0.3 0.8 10.7 0.34 0.39 0.91 6.5 7.9 5.4 10.8 0.66 From a management standpoint, a question more important than why does a wild turkey use snowberry is the question 'does a wild turkey require snowberry'. This question can be fully addressed only with a manipulative experiment. The study design used here is strictly correlative. Comparison of nesting habitat of successful and unsuccessful hens, although still only correlative, more closely approximates such an experiment as it relates a measure of fitness (the production of young) to habitat. Also, correlations are strengthened when coupled with additional, independent evidence such as knowledge of the high food availability and high predator densities along the South Platte River floodplain. �246 Brood Movements Eggs in 7 nests successfully hatched (Table 2). Only 4 brood flocks were monitored intensively due to transmitter failure, long travel distances, and flocking with other radio-marked brood hens. Three of these 4 brood flocks were composed of multiple (2 or 3) hens and 15 poults. Home ranges calculated via the minimum area method ranged from 14 to 62 ha in size (Table 7). Jennrich and Turner (1969) home range estimates were 106-209% larger. These home range estimates reflect brood ages of 1-9 weeks. Small sample sizes prohibited separate analyses of early and late brood home ranges. Mackey (1982) observed late brood ranges 3-4 times larger than early brood ranges, hypothesizing an areal increase due to increased poult mobility and a more diversified diet. Mean distance from nest to home range centers was 742 m (range 290-1070 m). Broods which made movements 700 m from the nest to the brood home range did so during the first 5-10 days after hatching. Three of these hens (2 brood flocks) nested on cnow areas and moved to adjacent, privately owned land for the brood-rearing period. Hen 151.621 also successfully nested on cnow property. Although no brood home range was calculated due to limited locations, she stayed on the wildlife area and within 1 km of the nest site during the early brood period. At the beginning of the late brood period (poults 3-4 weeks of age), she moved 5 km to an apparent late brood range. This hen nested on a small island. She apparently crossed the snowmelt-filled river with her flightless brood within 9 days after hatching. Hen 151.028 also crossed a section of river with poults <5 days after hatching. Table 7. Wild turkey brood home ranges (ha) in northeast Colorado, 1986. Brood hen Minimum area method Jennrich and Turner method N 150.196 151.701 l5l.65la 151.461 14 23 59 62 34 71 133 128 17 12 20 17 aBrood flock included hen 151.435. Brood Habitat Use Macrohabitat Use.--Ninety-seven locations were obtained for analysis of macrohabitat use. Wild turkeys were located within the river bottom 63 times (65%) (Table 8). Edge locations reflect either visual observations <40 minto the river bottom or triangulated location estimates where habitat type could not be confidently assigned. Agricultural locations included observations in barley, wheat stubble, plowed fields, and previously grazed pastures. Agricultural locations obtained via triangulation may also have included corn and sorghum. Forty-five of the 97 locations were triangulated location estimates. Significant location error may have occurred due to inherent error �247 of the telemetry system 6r from turkey movements during the location process (Schmutz, unpub1. data). However, I believe misc1assification was low as percent of locations per habitat type was similar for visual and triangulated locations (P >0.1). Distribution of locations among habitat types differed between hens. The 2 brood hens with the smallest home ranges (150.196 and 151.701) were always located within the river bottom. These hens may have been in higher quality brood habitat that did not require movement to edge or agricultural habitats (probably for food). Table 8. Habitat type of wild turkey brood locations obtained visually or via biotelemetry triangulation, northeastern Colorado, 1986. Brood hen River bottom (N) 150.196 151.701 151.651a 151.461 151.621 151.028 17 12 17 12 5 0 Totals 63 Edge (N) 0 0 5 6 2b 0 13 Agricultural (N) 0 0 11 7 0 3c 27 aBrood flock included hen 151.435. bEdge locations represent river bottom - road edge. cAll 3 locations in previously grazed pasture. Microhabitat Use.--Ear1y and late brood habitats were not significantly different for any of the measured variables. These 2 data sets were therefore combined and brood habitat compared to RN plots (Table 9). Greater understory cover in the first 0.5 m at brood sites was probably due to greater amounts of forbs. Shrubs were less common at brood sites than at non-use sites. Where shrubs occur the cover they provide is probably too dense for poults (Healy 1985). These analyses were performed with Mann-Whitney tests which examine differences in means. The river bottom habitat of the South Platte River floodplain is quite heterogenous and, thus, parameter values may have equal means but be from different segments of this heterogenous distribution. The Ko1mogorov-Smirnov 2-samp1e test was therefore performed on these same data (Table 9). Nonrandom brood habitat use was evident from the many significant differences. I am presently unsure of how to interpret these differences; further examination of these parameter distributions is required. Molting Two of 3 tail-clip mounted transmitters monitored through the nesting and brood~rearing period were recovered via molting. Tail feathers with transmitters were molted during the last week in August and the first 2 weeks in September. The third tail-clip transmitter apparently failed before molting. �248 Vegetafive parameters at wild turkey brood and non-use sites in Table 9. northeast Colorado, 1986. Brood 28) RN plots (N. = 19) (!! = x SD x SD pa pb Canopy cover, % 27 26 30 34. 0.72 0.99 Understory cover, % 0-0.5 m 0.6-1.0 m 1.1-2.0 m 55 19 9 26 20 10 34 15 9 30 18 9 0.02 0.31 0.88 0.02 0.01 <0.001 Life form, m/22 m Shrubs Forbs Grasses Bare ground <1 5 12 5 1 4 6 7 2 4 12 3 4 5 7 4 0.06 0.18 0.76 0.34 <0.001 0.17 0.99 0.004 Understory height, cm Plot center Total plot 37 42 25 19 28 33 26 18 0.12 0.40 0.10 0.15 Distance to large (>30 cm DBH) tree, m 30 47 23 31 0.14 0.04 Basal area, m2/ha 0-25 cm DBH 26-45 cm DBH >45 cm DBH Totals 0.6 2.8 5.6 1.0 3.4 8.4 2.3 2.2 4.1 5.4 3.1 9.5 0.48 0.52 0.26 <0.001 <0.001 <0.001 9.1 9.8 8.6 11.8 0.71 0.004 aprobability level for Mann-Whitney test. bprobability level for Kolmogorov-Smirnov 2-sample test. LITERATURE CITED Amundson, T. E. 1985. Health management in wild turkey restoration programs. Proc. Natl. Wild Turkey Symp. 5:285-294. Bailey, R. W., and K. T. Rinnell. 1967. Events in the turkey year. Pages 73-92 in O. H. Hewitt, ed. The wild turkey and its management. The Wildl. Soc., Washington, D.C. Canfield, R. H. 1941. Application of the line interception method in sampling range vegetation. J. For. 39:388-394. Healy, W. M. 1985. Turkey poult feeding activity, invertebrate abundance, and vegetation structure. J. Wildl. Manage. 49:466-472. �249 Jennrich, R. I., and F. B. Turner. 1969. Measurement of non-circular home range. J. Theoretical Bio1. 22:227-237. Lange, C. A. 1983. Fundamentals of successful wild turkey trap and transplant operations. Central Mtn. and Plains Sect., The Wi1d1. Soc. Annu. Mtg. Gunnison, Colo. 8 pp. Lazarus, J. E., and W. F. Porter. 1985. Nest habitat selection by wild turkeys in Minnesota. Proc. Nat1. Wild Turkey Symp. 5:67-82. Mackey, D. L. 1982. Ecology of Merriam's turkeys in southcentra1 Washington with special reference to habitat utilization. M.S. Thesis, Washington State Univ., Pullman. 87 pp. Mohr, C. o. 1947. Table of equivalent populations of North American small mammals. Am. Mild. Nat. 37:223-249. Nudds, T. D. 1977. Quantifying the vegetative structure of wildlife cover. Wi1d1. Soc. Bull. 5:113-117. Orians, G. H. 1980. Some adaptations of marsh-nesting blackbirds. Univ. Press. Princeton, N.J. Princeton Reagan, J. M., and K. D. Morgan. 1980. Reproductive potential of Rio Grande turkey hens in the Edwards Plateau of Texas. Proc. Nat1. Turkey Symp. 4:136-144. White, G. C., and R. A. Garrott. 1984. Portable computer system for processing biotelemetry triangulation data. Colorado Div. Wi1d1. Game Info. Leaflet 110. 4 pp. Approved by ~_,-"-7_~.....;;._ ~~=-..:..::~:;.::;._. C1ait E. Braun Wildlife Research Leader _ ��251 Colorado Division of Wildlife Wildlife Research Report April 1987 JOB FINAL REPORT State of Colorado Project 01-03-045 (W-37-R) 13 Work Plan Job Title: 8 -~- Population Characteristics and Habitat Use by Columbian Sharptailed Grouse in Northwest Colorado Period Covered: Author: : Job Avian Research 01 December 1980 through 30 June 1987 Kenneth M. Giesen Personnel: Michael D. Bauman, Clait E. Braun, Kevin Brennan, Kristi Coughlon, Kenneth M. Giesen, James Haskins, James L. Hicks, Richard W. Hoffman, Thomas Lines, Michael W. Middleton, Gary C. Miller, Daniel Schaad; Colorado Division of Wildlife ABSTRACT Lek attendance patterns, seasonal movement, home range, habitat perferences, nesting parameters, and population characteristics as measured by harvest of Columbian sharp-tailed grouse (Tympanuchus phasianellus columbianus) were studied from 1981 to 1985 in Routt and eastern Moffat counties, Colorado. Two study areas, Cedar Hill Gulch and Hayden, were selected for intensive study of movements and habitat preferences of radio-marked birds. Peak numbers of sharp-tailed grouse occurred on leks in late April and coincided with peak hen attendance and mating. Females (N = 20) dispersed farther from leks than males (~ = 18) for all months documented (Apr-Dec). Most (>95%) spring-autumn locations of radio-marked birds were within 3.0 km of the lek of capture. Home range size averaged 103.3 ha and was similar (p > 0.05) for males and females. Mountain shrub habitats dominated by big sagebrush (Artemisia tridentata), snowberry (Symphoricarpus sp.), and serviceberry (Amelanchier sp.) were used by sharp-tailed grouse in preference to all other available habitat types. Shrub densities were highest at nest sites and higher at sharptail use sites than at random sites. Observed clutch size (N = 13) averaged 10.8 eggs with eggs in 61.5% of nests hatching successfully. Most hunter harvest occurred on <5% of the occupied habitats (California Park, 20-mile Park). Examination of hunter-harvested birds indicated an even sex ratio and an estimated annual population turnover of 57.6%. �252 MANAGEMENT RECOMMENDATIONS 1. Ascertain status (active/inactive) of a sample (N = 25) of historic leks in Routt and Moffat counties annually between 1 April and 15 May. Leks should be visited within 2 hours of sunrise and all sharptails counted. Only 1 visit per active lek is needed annually if birds are observed. 2. Use intensive search, as time permits, and landowner contacts to locate additional sharp-tailed grouse leks. Priority should be given to areas of high hunter pressure (California Park, Slater Park, Twenty-mile Park) and to areas expected to be impacted by habitat changes due to mining, mineral exploration, reservoir development, subdivisions, etc. 3. Ascertain status (active/inactive) of all historic leks in Dolores, Mesa, Montrose, San Miquel, and Rio Blanco counties at least every 2 years. Leks should be visited within 2 hours of sunrise between 1 April and 15 May. 4. Hunter harvest of sharp-tailed grouse should be measured annually through the use of hunter check stations and/or wing barrels. Wings should be collected from each station each Monday morning and Friday afternoon and tagged with location and date. A sample of at least 100 wings from Routt and Moffat counties should be collected annually for estimates of annual production and population turnover. 5. Mountain shrub habitats within 3.0 km of leks should be maintained to provide suitable nesting, brood rearing, and loafing cover. A mixture of vegetation with total densities up to 30,000 shrubs/ha is recommended. Snowberry, serviceberry, and chokecherry (Prunus virginianus) are preferred by Columbian sharp-tailed grouse and should be maintained within occupied habitats. �253 POPULATION CHARACTERISTICS AND HABITAT USE BY COLUMBIAN SHARP-TAILED GROUSE IN NORTHWESTERN COLORADO Kenneth M. Giesen Concern over status of Columbian sharp-tailed grouse in Colorado intensified after a questionnaire survey indicated both population size and distribution of this subspecies had declined sustantially in western North America (Miller and Graul 1980). Reasons for this decline are not well documented although changes in land use resulting from agriculture, energy development, and human population growth have coincided with population declines (Hart et al. 1950, Kessler and Bosch 1981). Because of low recreational demand for this species in Colorado, there has been only sporatic attempts by wildlife managers to monitor population levels and hunter harvest. Previous studies of Columbian sharp-tailed grouse in Colorado delineated its distribution, provided general information on habitats used (primarily lek site descriptions), and documented low harvest levels (Rogers 1969, Giesen and Hoffman 1981). Because energy development and associated human population activities are expected to increase within the range of Columbian sharp-tailed grouse, this study was initiated to measure population parameters and quantify habitat use of this subspecies in northwest Colorado. P. N. OBJECTIVES The major objectives of this study were to: (1) measure sharp-tailed grouse population parameters including age and sex ratios, productivity, and population turnover, (2) document sharp-tailed grouse habitat preferences, (3) ascertain movement patterns of sharp-tailed grouse and measure home range, (4) measure harvest levels of sharp-tailed grouse in western Colorado, and (5) prepare standard operating procedures for monitoring sharp-tailed grouse populations in western Colorado. METHODS The initial year of investigation was primarily devoted to selecting study areas and developing and testing techniques for locating, capturing, and marking sharp-tailed grouse. Potential sites were limited to Routt and eastern Moffat counties where most historic leks were located and populations presumably highest (Giesen and Hoffman 1981). Local Division of Wildlife District Wildlife Managers in these areas were contacted and their knowledge of sharptail populations, habitat conditions, and.access was considered in selecting study sites. Active leks were located using a parabolic microphone listening device and searching with binoculars. Once located, leks on study sites were surveyed within 2 hours of sunrise 1 or more times per week from March through June. All sharptails were counted and classified to sex using behavioral criteria (Hjorth 1970). After several counts were obtained, grouse were flushed to obtain a count of all lek-attending birds including those not displaying or those hidden from view by vegetation. Other historic leks in Routt and eastern Moffat counties were surveyed when access and time allowed. All sharptails on active leks were classified to sex when possible. �254 Male and female sharp-tailed grouse were trapped on leks using walk-in funnel traps (Giesen et ale 1982). Mirror traps (Tanner and Bowers 1948), bait traps (Gullion 1961), and mist nets were tried at different times but no grouse were trapped using these methods. Each trapped grouse was marked using a numbered aluminum band and a unique combination of colored plastic bandettes. Sex of captured birds was ascertained from plumage characteristics of tail feathers (Ammann 1944), crown feathers (Henderson et ale 1967), and development of air sacs and eye combs. Age was ascertained by the shape and wear of the distal primaries, these primaries were more pointed and frayed in yearling «15 months of age) than in adults (>15 months of age) (Ammann 1944). All captured grouse were weighed to the nearest 5 gms on a pesola scale. Miniature lithium-battery or solar-powered transmitters (weight 20-30 gms) were placed on a sample of male and female sharp-tailed grouse each year using a poncho (Amstrup 1980). Transmitter packages weighed <4.0% of the birds . weight and did not appear to affect reproductive behavior, movements, or habitat preference. Attempts were made to visually locate all radio-marked birds weekly using a portable receiver and hand-held 3-element yagi antenna. When a signal was heard the radio-marked bird was approached by walking rapidly towards it until it flushed. Observations were recorded on a standardized form and locations plotted on a topographic map (scale 1:24,000). Home ranges were calculated using the convex polygon method (Mohr 1947). Nests were located by following radio-marked hens and with the use of a trained pointing dog. Clutch size was ascertained during early incubation and nest fate was monitored by inspecting nests weekly until hatch or depredation. Vegetative structure (density) and shrub species composition were measured at random sites and grouse use sites using cover board (Jones 1968) and point-centered quarter methods (Cottam and Curtis 1956). Random points were located using a table of random numbers to select UTM coordinates within 2.0 km of each lek on the study areas. Random points were plotted on a topographic map (scale 1:24,000) to the nearest 50 m. Topographic features were used to locate these points in the field and a random distance (0-25 m) was paced in a randomly selected cardinal compass direction to the measurement site. The flush site was used as the central point for measurement of vegetation at grouse use sites. The cover board consisted of 3 square panels, 15 cm on a side, each divided int·o 25 3-cm squares, alternately painted black and white. The cover board was placed on the sampling point with 1 side pointing north. Readings of each panel were taken from approximately 15 cm above the ground at a distance of 5 m, and at 450 above the cover board from the observers eye level. The number of colored squares at least 50% visible were recorded for each panel for a total of 6 scores. Additional cover board readings were taken at 5- and 10-m distances at each of the cardinal compass directions. When analyzed, scores for each point consisted of the sum of the panel scores for both ground level and 450 readings. Possible scores ranged from 0 (total obstruction) to 75 (total visibility). The point-centered quarter method (Cottam and Curtis 1956) was used to measure species composition and density of shrubs. Because of phytomorphological variance among shrub species in the study area, measurements were recorded and �255 analyzed separately for shrubs ~l.O m and >1.0 m in height. Species of shrub was recorded for the nearest shrub in each quarter and height and crown diameters were measured to the nearest centimeter. Crown diameter was calculated as the mean of the greatest vertical canopy intercept and the vertical canopy intercept at 900 to the former measurement. Relative frequency, relative density, importance, and absolute density were calculated following Dix'(196l). When the nearest shrub was >5 m for short (~1.0 m) shrubs or >10 m for tall (>1.0 m) shrubs, the species was recorded but shrub height and crown diameter were not measured. The distance from the center point to these shrubs was arbitrarily set at 6 and 12 m, respectively. Sharp-tailed grouse hunting pressure and harvest characteristics were measured in Routt and Moffat counties using check stations and wing barrels (Hoffman and Braun 1975). Check stations were normally operated only during the ' initial weekend of the hunting season and only in areas having the highest number of hunters. Wing barrels were placed in most accessible locations having sharptail populations where harvest was suspected. Wing barrels were checked twice weekly to document time of harvest and to separate weekend from weekday harvest. Whole body weights of harvested grouse were obtained when possible at check stations and sex was ascertained by gonadal examination. Age of harvested birds (Ammann 1944) was.recorded for all wings 'collected from wing barrels and at check stations. STUDY AREAS Two study areas of differing habitat types (Cedar Hill Gulch and Hayden) were selected in northwest Colorado (Fig. 1). Study area boundaries were delineated after the initial year of radio-tracking and included all l-km2 UTM grids within 2 km of active leks which received spring, summer, or autumn use by radio-marked sharp-tailed grouse. Cedar Hill Gulch was approximately 20 km northeast of Craig in eastern Moffat County. The study area of 2,200 ha included parts of sections 29-33, T9N, R89W, and parts of sections 4-9, T8N, R89W. Most of this area is in private ownership although there are small parcels of Federal (BLM) and State (Land Board) lands. Topography is moderately rolling with elevations ranging from 2,000 to 2,500 m. Drainage is primarily to the south and west through Cedar Hill Gulch and Little Bear drainages into Fortification Creek and eventually to the Yampa River. Vegetation consists primarily of mixed mountain shrub habitats (83.4%) with scattered irrigated hay meadows (14.2%) with some dryland wheat (2.0%) occurring on the southern preriphery of the area (Fig. 2). Cattle grazing is the primary use of the rangelands. Two sharp-tailed grouse leks were present on the area at different times although most research activity centered around the largest lek, Cedar Hill Gulch. The Hayden study area was approximately 6 km northwest of Hayden in Routt County. The specific study area of 1000 ha included parts of sections 20, 28-30, and 32, T7N, R88W. The land is entirely in private ownership or under private control (State Land Board). Topography is gently rolling with hills bisected with drainages to the south (Coal Bank Gulch), west (Rock Spring Gulch), and north (Corral Gulch, Buck Gulch). Maximum topgraphic relief is approximately 100 m and elevations ranged from 2,100 to 2,200 m elevation. The vegetation was a mixture of mountain shrub habitats (72.8%) interspersed �256 with Much to a over edge cultivated hay meadows (7.2%) and dry1and wheat fields (18.8%) (Fig. 3). of the area has been subdivided into small ranches and grazing is limited few horses and cattle. One rancher pastures his cattle in a hay meadow winter and sheep graze in mountain shrub pastures adjacent to the north of the study area. One active sharp-tailed grouse 1ek was on the area. RESULTS AND DISCUSSION Lek Surveys The number of leks inventoried in recent years (1964-85) varied from none to 26 annually and increased with the initiation of the research project (Table 1). Initially it was believed that 1ek counts (i.e., average number of males or total birds on leks) would provide a reasonable, if not accurate, estimate of population change over time, or at least indicate population trends for this species (Ammann 1957, Pepper 1972, Hillman and Jackson 1973, Kirsch et a1. 1973, Kobriger 1975, Yde 1977, Mattise 1978, Nielsen 1978, Ziegler 1979). Beck and Braun (1980) reported that 1ek counts have no proven value as a population index for sage grouse (Centrocercus urophasianus). ~obriger (1975) earlier concluded that sharp-tailed grouse 1ek counts were not correlated to any population estimator, thus suggesting spring 1ek counts have little value in measuring population size or documenting population trends. Table 1. Counts of sharp-tailed grouse on leks in Routt and Moffat counties, Colorado, 1964-85. Year 1964 1965 1966-76 1977 1978 1979 1980 1981 1982 1983 1984 1985 N active grounds counted Total birds counted birds/1ek 7 15 0 2 6 15 5 24 26 24 12 7 38 91 0 16 54 123 36 335 317 311 107 73 5.4 6.1 0 8.0 9.0 8.2 7.2 14.0 12.2 13.0 8.9 10.4 Evidence indicates that not all sharp-tailed grouse attend leks daily (Rippin and Boag 1974) and only a portion of the adult male population becomes territorial on leks in spring (Moyles and Boag 1981). As a result, numbers of males on leks may be expected to vary daily and seasonally. The number of males attending leks (lek size) generally follow a predicted pattern. Counts in March were variable, probably the result of daily weather and precipitation. Lek attendance became regular in April and high counts of �257 males and total birds occurred during the last 2 weeks of April (typically the peak of hen attendance and mating). Numbers of males attending leks declined throughout May and early June and by mid-June lek attendance became sporatic. Data from 4 leks (Cedar Hill Gulch, Pelly's, Smith's, Wiseman's) checked periodically from 21 March to 20 June 1983 show the typical trend in lek counts during the breeding season (Table 2). Because of vegetative obstruction at several of these leks I was not always able to classify all birds to sex, thus the high count in April, which coincided with the peak of hen attendance in 1983, partially accounted for the high counts during this period. However, counts of sharp-tailed grouse on leks in March or early April before hen attendance were typically 10-30% higher than counts after 11 May suggesting that fewer males attend leks later in the breeding season. Not only did fewer males attend leks after the peak of hen attendance but displays and vocalizations were also reduced. Thus, when comparing trends in lek . counts (Table 3) the data are usually biased by the timing of counts which is often related to observer schedules in spring and access to leks each year. Counts of males or total birds on leks may vary within a year even though the population is constant. Thus, if lek counts are to be used to measure population trends, an effort should be made to survey leks at the same time each year. In recent years use of lek surveys to replace lek counts to obtain indices of population changes in prairie grouse has been tested (Cannon and Knopf 1981, Martin and Knopf 1981). These studies indicated a strong positive correlation between numbers of leks and densities of breeding males. These studies suggest that when prairie grouse populations increase, males respond by "creating" more leks instead of increasing the average number of males on each lek. Cannon and Knopf (1981) indicated that small leks (i.e., few males) and temporary leks must be included if lek surveys are to be sensitive indices to population changes. Traditionally the large and more stable leks are generally surveyed and counted first, followed by smaller, peripheral, temporary, and inaccessible leks. There are several important values of lek counts and surveys in addition to their possible use as population indices. The location of the lek is of extreme value to a grouse for biological purposes (reproduction). Lek surveys allow observers to subjectively evaluate habitats and document major changes (Beck and Braun 1980). The knowledge of lek locations and activity also has public relation values and may stimulate a greater appreciation for the species and its environment. Most importantly, sharp-tailed grouse leks are easier to locate and quantify than nesting, brood rearing, or winter habitats, thus being important in land use planning and for potentially mitigating impacts to sharptail populations. Movements and Home Range Radio transmitters were placed on 18 male and 20 female sharp-tailed grouse from 3 study leks during 1982-85 to facilitate their location for measurement of seasonal movements and home range. Initially attempts were made to locate each radio-marked grouse at least weekly but this was not accomplished. The number of visual locations of radio-marked birds was directly related to their survival from spring to fall, movements, and transmitter life. The number of visual locations per radio-marked grouse ranged from 2 to 29 and represented �N VI Table 2. Seasonal variation in counts of sharp-tailed grouse observed on leks, Routt and Moffat counties, Colorado, 1983. Lek Mar 21-30 Apr 11-20 21-30 Cedar Hill Gulch Pe11y's Smith's Wiseman's 19 3 16 7 9 9 16 11 14 8 15 10 16 8 18 9 Totals 45 45 47 Average 11.2 11. 2 Percent of total high count 78.9 78.9 1-10 Jun Mal 1-10 00 11-21 21-31 11 7 16 7 10 6 14 6 9 6 13 6 10 5 15 6 5 2 14 7 51 41 36 34 36 28 11.8 12.8 10.2 9.0 8.5 9.0 7.0 82.5 89.5 71.9 66.7 59.6 66.7 51.9 1-10 11-20 �259 Table 3. Trends in sharp-tailed grouse 1ek activity, Routt and Moffat counties, 1981-85. High count of all birds 1982 1984 1983 Lek 1981 Annans 20-mi1e Barnes Bear Creek Ca1i~ornia Park Road #1 California Park Road 112 Cedar Hill Gulch Cottonwood Creek E1khead Road #1 E1khead Road 112 (Wisemans) E1khead Road 113 Elk Mountain tl2 (Masiare11i) Elk Mountain 113 Elk River Cemetary Energy Fuels Fish Creek Fly Creek Green Acres Hick's Hinkle Maneotis McKinney Ranch Morapas Gas Field Morgan Creek Noland Ranch Pe11y's Salt Creek til Salt Creek #2 Schneider's Scott Place Smith's Taylor's Villard's Wingate Yellow Jacket tl2 14 NC 3 9 1 24 11 0 15 10 9 3 3 10 24 NC 2 15 6 40 17 NC 46 NC 13 13 9 NC NC NC NC 18 NC 20 16 5 4 0 NC 14 11 NC 15 11 26 4 3 NC 11 0 0 0 5 30+ 17 NC 25 NC 14 13 9 NC 3 24 NC 25 NC 7 7 NC 0 0 0 19 9 NC 11 18 24 2 1 1 8 0 0 NC 3 10+ 14 36 31+ 11 9 12 4 0 0 18 0 4 0 2 14.0 13.3 11.5 Average/active 1ek with birds seen aNC = Lek not checked; 0 Lek surveyed, inactive. NC NC 0 2 0 17 0 NC 10 5 2 NC NC NC NC NC NC NC NC NC 11 14 19 15 3 2 0 NC NC 7 NC NC NC NC 8.9 1985 0 NC 1 1 1 10 0 NC 7 5 9 NC 0 NC NC 0 NC NC NC 0 21 0 15 6 0 NC NC NC NC 2 NC 2 NC NC 6.7 �260 10-252 days of observation. Few yearlings of either sex were radiomarked (2 males, 2 females). Their movements and home ranges were similar to those of adults and the data were pooled for analysis. Movements.--Mean monthly dispersion from lek of capture was calculated for each radio-marked grouse and summarized separately for males (N = 18) and females (N = 20) (Table 4). For each month (Apr-Dec) the median dispersion distance was less for males than females (Table 5). The overall mean dispersion distance was less for males than for females (605 vs. 1,475 m, t 3.6395, P <0.01). When mean dispersion distances were calculated separately for each-season, males remained closer to leks than females for spring (16 Mar-3l May, 229 vs. 1,394 m, t = 5.8566, P < 0.001) and summer (1 Jun-3l Aug, 858 vs. 1,603 m, t = 1.9455, P <0.05). This was not true for Autumn (1 Sep-3l Oct, 1,276 vs. 1,606 m, t = 0~5227, P <0.30). Table 4. Dispersion from lek of capture by radio-marked male (~ = 20) and female (N = 18) Columbian sharp-tailed grouse in Routt and Moffat counties, Colorado, 1982-85. Distance from lek (km) ::0.5 0.5 - 1.0 1.0- 1.5 1.5 - 2.0 2.0 - 2.5 2.5 - 3.0 > 3.0 Male 200 65 6 1 1 3 15 11 locations Female Totals 11 66 62 42 15 11 10 211 131 68 43 16 14 25 Male 68.7 91.1 93.1 93.5 93.8 94.8 100.0 Cumulative % Totals Female 5.1 35.5 64.1 83.4 90.3 95.4 100.0 41.5 67.3 80.7 89.2 92.3 95.1 100.0 Table 5. Mean monthly dispersion of male and female sharp-tailed grouse from lek of capture, Routt and Moffat counties, Colorado, 1982-85. Month N Apr May Jun Jul Aug Sep Oct Nov Dec 3 15 15 11 11 8 6 5 3 Males Median (m) Range (m) N 0 150 360 425 600 585 625 800 1,100 0 0-680 0-3,850 140-6,100 100-3,250 100-3,730 400-3,700 250-1,200 700-1,550 1 20 17 12 7 5 5 0 1 Females Median ~m) Ta Range ~m) 1 050 1 130 1,290 1,175 1,300 1,550 1,000 N/A 400-3,650 500-4,570 700-4,400 400-2,520 650-2,550 350-5,000 6,700 N/A N/A <0.001 <0.001 0.01 <0.005 > 0.10 > 0.10 aprobability of males dipsersing less than females, Mann-Whitney Test. N/A �261 Male attachment to lek sites was expected as males have been reported to display or visit leks every month of the year (Ammann 1957, Hillman and Jackson 1973, Twedt 1974). The fidelity and close attachment of males to leks may be a method for dominant males to maintain their position on leks and secure breeding position, and for subordinate males to secure a central territory on leks. Holding a central territory is thought to be a prerequisite for breeding. The advantage to females in dispersing from leks (or maintaining home ranges away from leks except for breeding) may be to decrease competition for resources, lessen predation, or more likely, to obtain more or better resources for feeding, nesting, and brood rearing. The similarity in mean dispersion distances from leks by males and females in autumn is the result of males moving farther from leks. These movements may indicate a decline in resources near the lek because of colder temperatures and desiccation/ exfoliation of vegetation or a movement to wintering habitat. Defining critical habitats (breeding, nesting, brood-rearing, foraging, wintering) is difficult without intensive studies of seasonal habitat use. However, one can assume that if a population maintains itself over time all components for survival must be available within the populations' (vs. individuals') home range. Analysis of >500 telemetry locations of sharp-tailed grouse from mid-March through December indicates >90% of their activities occurred within 2.5 km of the lek of capture (Table 4) and >95% within 3.0 km of the lek of capture. Although winter habitat requirements of Columbian sharp-tailed grouse are not well known, it is likely that maintaining all habitats within 3.0 km of leks will provide all seasonal habitat requirements for sharp-tailed grouse in northwest Colorado. Home Range.--Thirty-six of 38 radio-marked sharp-tailed grouse (18 males, 18 females) provided some information to estimate home range size. Home range was not strongly correlated to the number of locations (r = 0.07) or the time interval (r = 0.38). A realistic estimate of home range-size was calculated using data-from birds located a minimum of 10 times (ave. = 18.2, range 11-29) over a minimum of 60 days (ave. = 162.5, range 75-252). No differences occurred between areas (Cedar Hill Gulch, Hayden) and data for each sex were pooled. There was no difference in home range size of males (N = 13, K = 106.7 ha) and females (N = 6, x = 96.8 ha) (Mann-Whitney U-Test, T = 30.5, P> 0.1). Average spring-autumn home range was 103.3 ha for 20 sharp=tailed grouse observed an average of 18.2 times over an average of 162.5 days (Table 6). Comparable home range data for Columbian sharp-tailed grouse are few. Marks and Marks (1987) calculated spring-autumn home ranges of 15 radio-marked sharptails in Idaho to be 187 + 114 ha, somewhat larger than observed in this study. They suggested home range size was a reflection of habitat quality with larger home ranges being observed in pastures with heavy grazing pressure by cattle. Spring-fall home ranges of prairie sharp-tailed grouse (!. E. compestris) range from 13 to 105 ha (Artmann 1970, Ramharter 1976, Gratson 1982) and for plains sharp-tailed grouse (T. E. jamesi) from 32 to 200 ha for individual birds, (Christenson 1970). �262 Table 6. Home range size of male (N = 13) and female (N = 7) Columbian sharp-tailed grouse, Routt and Moffat counties, Colorado: 1982-85. Transmitter freg,uency Year Age 151.160 151.309 151.187 151.264 151.453 151.552 150.177 151.600 150.960 151.021 150.228 151.361 151.502 151.041 150.480 151.311 150.993 151.360 150.420 150.480 1982 1982 1982 1982 1983 1983 1983 1983 1983 1983 1983 1984 1984 1984 1984 1984 1984 1984 1984 1985 Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad Ad Yrlg Ad Ad Ad Ad Yrlg Ad Yrlg Ad Average N days Home range size (ha) Sex Location N locations M M M M M M F M M M F F M F F F M M M F Hayden CHG CHG CHG CHG Hayden Hayden Hayden CHG CHG Hayden Hayden Hayden CHG CHG CHG CHG CHG CHG CHG 11 15 11 14 27 24 20 29 11 18 13 14 21 16 22 26 19 24 18 11 187 252 215 252 91 231 169 205 89 152 169 121 123 163 157 219 170 127 75 83 43.1 406.2 48.6 309.7 15.3 100.7 111.8 72.9 25.0 76.4 98.6 127.8 28.5 132.6 93.1 84.0 170.1 61.1 29.9 29.9 18.2 162.5 103.3 These studies suggest that spring-fall home ranges of sharp-tailed grouse are typically less than 200 ha and that habitat quality may influence home range size. Few data are available for year-long home ranges, primarily because winter data are lacking. Robel et ale (1972) winter-trapped 5,680 plains sharp-tailed grouse in South Dakota and had subsequent fall band recoveries as far as 28 miles (44 km) from the trap site. They documented greater movements for juveniles than adults, and females than males. The long distances between trap sites and recovery location may have represented dispersal of juveniles and a migration between breeding and wintering habitats for this population. Habitat Use Habitat Preference.--Four habitat classes were delineated on each study area, plotted on large scale aerial photographs (approximately 1:8,000) and the percent of each type occurring on the study areas measured using a dot grid. The habitat type was recorded each time a radio-marked sharptail was located (Apr-Dec) to ascertain possible habitat preferences. Data were analyzed separately for each sex to elucidate potential differences in habitat preferences and separately for each area because of habitat differences between Cedar Hill Gulch and Hayden. The null hypotheses was that each sex would use each habitat type in proportion to its availability. �263 There were differences (p <0.05) in relative habitat preferences between males and females. Although both sexes were located most often in mountain shrub habitats, males tended to select for hay pasture habitat more often than expected from habitat availability (Tables 7,8). In contrast, females generally avoided all habitats other than mountain shrub. Although the data were pooled for all months, the preference of males for hay pasture was strongest in summer (Jun-Aug). In fall and early winter, both sexes used mountain shrub habitats almost exclusively. The use of wheat fields was limited to a few weeks after harvest when waste grain was readily available prior to fall snow cover. Table 7. Habitat use by radio-marked male and female sharp-tailed grouse at Cedar Hill Gulch, Moffat County, Colorado, 1982-85. Habitat type Mountain shrub Hay pasture Wheat/stubble Miscellaneous Totals ax2 b:X2 Percent of area Observed Males Expecteda Females Expected 15 Observed 83.4 14.2 2.0 0.4 142 42 0 0 153 26 4 1 144 7 0 0 126 21 3 1 100.0 184 184 151 151 15.637, O.Ol<P <0.025. 15.905, 0.01 <: P < 0.025. Table 8. Habitat use by radio-marked male and female sharp-tailed grouse near Hayden, Routt County, Colorado, 1982-85. Habitat type Mountain shrub Hay pasture Wheat stubble Miscellaneous Totals Percent of area Observed Males Expecteda Females Expected15 Observed 20 1 63 2 1 0 48 5 12 1 107 66 66 72.8 7.2 18.8 1.2 49 49 9 0 78 100.0 107 8 227.957, R< 0.005. 17.571, 0.005 < P < 0.01. The preference of males for hay pasture habitats in summer is not well understood. Structurally, hay fields were more similar to wheat fields than mountain shrub habitats, yet males (and females) generally avoided wheat fields. It is possible that food resources, especially insects, may have been relatively more abundant in hay pastures. Yet females, even hens with broods, �264 rarely used hay fields even though chicks are generally suspected to feed heavily on insects the first few weeks after hatch. The relative amount of mountain shrub rangeland necessary to maintain Columbian sharp-tailed grouse populations is unknown as both areas studied had high proportions of mountain shrub habitats (>70%) and stable sharptail populations. Marks and Marks (1987) reported differential selection for sagebrush habitats although their study areas had relatively little non-shrub habitats. Habitat Characteristics.--Visual obstruction readings (VOR) (Jones 1968) were recorded at 378 random sites at Cedar Hill Gulch and 225 random sites at the Hayden study area (Table 9). Ground level VOR was much less than at 450 in all habitat types. The high ground level VOR of hay pasture sites was caused by constant cattle grazing of these pastures resulting in little vegetative height. Table 9. Visual obstruction readings on random vegetation plots at Cedar Hill Gulch and Hayden study areas, northwest Colorado, 1982-85. Habitat tlpe N Cedar Hill Gulch 0 45 N Halden 0° 45° Mountain shrub Hay pasture Wheat Wheat stubble Fallow field 315 54 9 0 0 3.1 40.5 0.2 N/A N/A 44.7 66.0 66.9 N/A N/A 54 63 54 18 36 0.0 4.3 14.3 32.3 51.4 36.0 52.4 64.6 75.0 74.8 Visual obstruction at sharptail use sites (Tables 10, 11) was similar to random sites with 1 exception. Males at Cedar Hill Gulch selected (p < 0.05) dense cover (lower VOR) in hay pastures, typically by using patches of ungrazed vegetation within the pasture. The selection of dense cover may provide sharptails with protection from predators, shelter from the elements (i.e., shade in summer) and a place to meet their nutritional and energetic requirements. The similarity in 450 VOR among all habitat types from both study areas suggests that concealment from predators may be the least important factor in selecting habitats for feeding and loafing or that all habitats exceed minimum cover requirements. Table 10. Visual obstruction readings at male sharp-tailed grouse feeding-loafing sites, Cedar Hill Gulch and Hayden study areas, northwest Colorado, 1982-85. Habitat tlpe· Mountain shrub Hay pasture N 215 77 Cedar Hill Gulch 0 5.3 2.3 Hayden 45° N OC 45° 50.5 44.1 19 71 0.3 4.6 44.0 57.7 �265 Table 11. Visual obstruction readings at female sharp-tailed grouse feeding-loafing sites, Cedar Hill Gulch and Hayden study areas, northwest Colorado, 1982-85. Cedar Hill Gulch Hayden Habitat type N 0 45 N Mountain shrub Hay pasture 215 0.4 N/A 44.5 N/A 56 o o 0.1 'N/A 36.5 N/A Characteristics of Mountain Shrub Habitats.--Vegetative characteristics of mountain shrub habitats were measured at 123 random sites and 186 sharp-tailed grouse use sites and analyzed separately by study area and for random sites, male use sites, and female use sites (Tables 12-23). Two species of short (~1.0 m) shrubs (snowberry, big sagebrush) dominated the landscape on both study areas and were the most important shrubs at grouse use sites. In all situations, snowberry had a higher importance value than any other short shrub indicating that, although it was structurally similar to big sagebrush (shrub height, crown diameter), it was relatively more abundant at grouse use sites. Densities of all short shrubs varied widely with slope and aspect and were apparently related to soil type and structure as well as moisture and temperatures differences. Generally the highest densities of shrubs occurred on north and east slopes. The most important tall shrub was serviceberry and sharp-tailed grouse on both study areas were associated with this shrub species. The only exception was equal male association with chokecherry at the Hayden study area. In contrast to other areas (Marks and Marks 1987), serviceberry was widely distributed on both study areas and generally throughout the range of Columbian sharp-tailed grouse in Colorado. The importance of serviceberry may be highest in winter (Dec-Feb) when all observations of sharp-tailed grouse were associated with this species. Sharp-tailed grouse apparently selected serviceberry buds in preference to other food during winter (Marks and Marks 1987). Investigation of habitats occupied by sharp-tailed grouse in western Colorado suggests the limits of sharptail distribution coincide with the distribution of serviceberry. Nesting Clutch Size and Nest Success.--Nests of 13 hens were examined for clutch size and nest success. Ten complete clutches averaged 10.8 eggs (range 8-14) with 10 eggs being observed most often. Two incomplete clutches had 6 eggs. Nest success was 61.5% with eggs in 8 of 13 clutches hatching successfully. Two hens were known to have been killed on the nest with hens surviving the other depredated nests. Clutch size varied from 4 to 12 eggs from 12 other nests in northwest Colorado (J. Monarch, pers. commun.) and from 9 to 12 eggs elsewhere (Marks and Marks 1987). Nesting Cover.--Females selected relatively dense cover (32,563 plants/ha) for nesting with snowberry and big sagebrush the most abundant shrubs at nest sites (Table 24). Densities of short shrubs (~1.0 m) were 5 times higher at . �Table 12. Vegetative characteristics of mountain shrub habitats at 105 random plots, shrub height ~1.0 m, Cedar Hill Gulch, Moffat County, Colorado Species Heighta Ame1anchier sp. Artemisia tridentata A. cana Cercocarpus montanus Prunus virginianus Purshia tridentata Quercus gambe11ii Rosa woodsii SlmphoricarEus sp. 47.8 41.4 47.8 71.0 22.2 34.2 44.2 22.0 44.0 -- Mean Crown diam.a 49.0 44.6 47.0 86.0 14.2 61.1 33.3 20.0 45.2 Totals N points N _plants Relative fr~qu_ency Relative densdty Importance va1ue~ Absolute _densityC 18 70 3 1 3 12 7 1 68 26 182 11 1 5 17 9 1 168 9.8 38.2 1.6 0.6 1.6 6.6 3.8 0.6 37.2 6.2 43.3 2.6 0.2 1.2 4.0 2.1 0.2 40.0 16.0 80.5 4.2 0.8 2.8 10.6 5.9 0.8 77 .2 480 3,429 201 16 93 310 162 16 3,096 183 420 100.0 99.8 199.8 7,803 aCentimeters. bSum of relative frequency and relative density. cP1ants/ha. Table 13. Vegetative characteristics of mountain shrub habitats at 105 random plots, shrub height >1.0 m, Cedar Hill Gulch, Moffat County, Colorado. Species HeLghtf' Ame1anchier sp. Artemisia tridentata CercocarEus montanus Juniperus communis Prunus virginianus Quercus gambe11ii SlmphoricarEus sp. 194.1 116.0 123.9 124.0 180.0 179.1 107.0 Totals Mean Crown diam.a 184.6 111.9 132.4 81.2 93.2 100.2 138.2 N points N plants 68 51 4 4 2 15 5 211 115 12 7 2 65 8 45.6 34.2 2.7 2.7 1.3 10.1 3.4 149 420 100.0 aCentimeters. bSum of relative frequency and relative density. cPlants/ha. Relative frequency Relative density Importance va1ueb Absolute dens t ty? 50.2 27.4 2.9 1.7 0.5 15.5 1.9 95.8 61.6 4.6 4.4 1.8 25.6 5.3 80 44 5 3 1 25 3 100.1 200.1 161 N 0\ 0\ �Table 14. Vegetative characteristics of mountain shrub habitats at 75 male sharp-tailed grouse use sites, shrub height 21.0 m, Cedar Hill Gulch, Moffat County, Colorado. Species Heighta Ame1anchier sp. Artemisia tridentata A. cana -Crataegus·sp. -Prunus virginianus Purshia tridentata Symphoricarpus sp. Quercus gambe11ii 57.7 47.0 26.0 65.0 50.3 30.5 37.5 24.4· Mean Crown diam.a 45.1 44.3 23.0 78.7 23.1 58.7 28.5 13.2 Totals Relative freguency Relative density Importance va1ueb Absolute densityC N N poInts plants 13 35 1 2 5 6 57 6 19 78 1 5 16 6 167 8 10.4 28.0 0.8 1.6 4.0 4.8 45.6 4.8 6.3 26.0 0.3 1.7 5.3 2.0 55.7 2.7 16.7 54.0 1.1 3.3 9.3 6.8 101.3 7.5 825 3,404 39 223 694 262 7,292 353 125 300 100.2 100.0 200.2 13,092 aCentimeters. bSum of relative frequency and relative density. cP1ants/ha. ~ Table 15. Vegetative characteristics of mountain shrub habitats at 75 male sharp-tailed grouse use sites, shrub height >1.0 m, Cedar Hill Gulch, Moffat county, Colorado. Species Ame1anchier sp. Artemisia tridentata Crataegus sp. Prunus virginianus Quercus gambe11ii Rosa woodsii Symphoricarpus sp. Totals Mean Heighti'! Crown diam.a 205.7 115.6 384.0 142.0 277 .4 169.5 109.5 167.9 91.9 367.0. 60.2 110.8 84.2 106.0 N N poInts plants Relative freguenc~ 58 23 2 5 16 2 2 182 58 7 10 38 2 3 53.7 21.3 1.8 4.6 14.8 1.8 1.8 108 300 99.8 Relative density Importance va1ueb Absolute densityC 60.7 19.3 2.3 3.3 12.7 0.7 1.0 114.4 40.6 4.1 7.9 27.5 2.5 2.8 167 53 6 9 35 2 3 100.0 199.8 275 N 0\ -...J aCentimeters. bSum of relative frequency and relative density. cP1ants/ha. �Table 16. Vegetative charactertistics of mountain shrub habitats at 75 female sharp-tailed grouse use sites, shrub height ~1.0 m, Cedar Hill Gulch, Moffat County, Colorado. Species Ame1anchier sp. Artemisia tridentata A. cana - -CercocarEus montanus Prunus virginianus Purshia tridentata Mahonia repens Quercus gambe11ii Rosa woodsii Symphoricarpus sp. Mean N N Relative Relative Importance Heigh_ta_ CroWIl.di_Cl~.~ _ ___ll9'int_s __ _p_lAP._t!3 freClu~_n_cy de11sJty___ va1ueb 45.6 56.1 74.5 57.0 40.7 29.0 7.7 29.5 31.5 44.8 34.8 55.1 62.4 55.5 22.0 55.8 9.8 26.4 22.2 35.1 Totals Absolute denad ty? 11 42 3 1 4 6 1 6 4 62 17 81 8 1 4 10 3 10 4 162 7.9 30.0 2.1 0.7 2.9 4.3 0.7 4.3 2.9 44.3 5.7 27.0 2.7 0.3 1.3 3.3 1.0 3.3 1.3 54.0 13.6 57.0 4.8 1.0 4.2 7.6 1.7 7.6 4.2 98.3 1,360 6,443 644 72 310 788 239 788 310 12,887 140 300 100.1 99.9 200.0 23,841 aCentimeters. bSum of relative frequency and relative density. cP1ants/ha. Table 17. Vegetative characteristics of mountain shrub habitats at 75 female sharp-tailed grouse use sites, shrub height >1.0 m, Cedar Hill Gulch, Moffat County, Colorado. Species Heighta Ame1anchier sp. Artemisia tridentata A. cana CercocarEus montanus Prunus virginianus Quercus gambe11ii Symphoricarpus sp. 191.0 116.0 106.0 113.5 148.0 222.7 116.9 -- Mean Crown diam.a 184.0 115.7 71.0 94.2 105.7 125.1 121.6 N _p_o'ints N plants Relative f:r~quenc_y 70 17 1 1 7 11 8 226 20 1 2 11 29 11 60.9 14.8 0.9 0.9 6.1 9.6 7.0 115 300 100.2 Relative d_ensity Importance va1ueb Absolute dens Lty" 75.3 6.7 0.3 0.7 3.7 9.7 3.7 136.2 21.5 1.2 1.6 9.8 19.3 10.7 208 18 1 2 10 27 10 100.1 200.3 276 \ Totals aCentimeters. bSum of relative frequency and relative density. cPlants/ha. N 0\ 00 �Table 18. Vegetative characteristics of mountain shrub habitats at 18 random plots, shrub height ~1.0 m, Hayden study area, Routt County, Colorado. Species Heighta Mean Crown diam.a Amelanchier sp. Artemisia tridentata Prunus virginiana Purshic tridentata Rosa woodsii Symphoricorpus sp. 32.0 35.0 69.0 68.0 35.0 51.1 16.8 33.4 71.7 62.0 11.5 51.0 Totals N _~9Ints N pl~nts Relative frequency 2 10 3 1 1 15 2 30 3 1 1 35 6.2 31.2 9.4 3.1 3.1 46.9 32 72 99.9 Relative density Importance valueb Absolute densityC 2.8 41.7 4.2 1.4 1.4 48.6 9.0 72.9 13.6 4.5 4.5 95.5 93 1,390 140 47 47 1,620 100.1 200.0 3,337 aCentimeters. bSum of relative frequency and relative density. cPlants/ha. Table 19. Vegetative characteristics of mountain shrub habitats at 18 random plots, shrub height >1.0 m, Hayden study area, Routt County, Colorado. Species He Lght=' Amelanchier sp. Artemisia tridentata A. cana -Prunus virginianus S~mEhoricarEus sp. 240.8 N/A 125.0 338.8 125.2 Totals Mean Crown diam.a 251.3 N/A 127.0 168.4 105.9 N N po'in_t_s~pl~nts Relative frequency Relative density Importance valueb ~ Absolute densityC 12 6 1 2 3 39 24 1 4 4 50.0 25.0 4.2 8.3 12.5 54.2 33.3 1.4 5.6 5.6 104.2 58.3 5.6 13.9 18.1 68 42 2 7 7 24 72 100.0 100.1 200.1 126 N CJ\ I.D aCentimeters. bSum of relative frequency and relative density. cPlants/ha. �Table 20. Vegetative characteristics of mountain shrub habitats at 12 male sharp-tailed grouse use sites, shrub height ~1.0 m, Hayden study area, Routt County, Colorado. Species Heighta Amelanchier sp. Artemisia tridentata Prunus virginianus Purshia tridentata Symphoricarpus sp. 41.0 29.3 50.6 36.6 42.7 Mean Crown diam.a 21.8 17 .5 22.2 58.2 36.9 Totals N N Relative ..Po_i"nt:s .p_lA'nts frequency Relative density Importance valueb Absolute densityC 2 6 3 3 7 2 15 5 8 18 9.5 28.6 14.3 14.3 33.3 4.2 31.2 10.4 16.7 37.5 13.7 59.8 24.7 31.0 70.8 27 198 66 106 238 21 48 100.0 100.0 200.0 635 aCentimeters. bSum of relative frequency and relative density. cPlants/ha. Table 21. Vegetative characteristics of mountain shrub habitats at 12 male sharp-tailed grouse use sites, shrub height >1.0 m, Hayden study area, Routt County, Colorado. Species Heighta Amelanchier sp. Artemisia tridentata Prunus vir~inianus Symphoricarpus sp. 241.5 106.0 183.7 103.0 Totals Mean Crown diam.a 252.0 115.5 106.7 117.5 N _ poIIl_ts N Relative Relative Importance piant~ ____lj.eJll.lenc_y densi_D'_.. valu.e_~ 8 4 6 1 15 14 18 1 42.1 .21.1 '31.6 5.3 19 48 100.1 -aCentimeters. bS~m of relative frequency and relative density. cPlants/ha. Absolute densj_!yc 31.2 29.2 37.5 2.1 73.3 50.3 69.1 7.4 50 47 60 3 100.0 200.1 160 N '-J o �Table 22. Vegetative characteristics of mountain shrub habitats at 24 female sharp-tailed grouse use sites, shrub height ~1.0 m, Hayden study area, Routt County, Colorado. Species Ame1anchier sp. Artemisia tridentata Gutierrezia sarothrae Prunus virginianus Rosa woodsii Symphoricarpus sp. Mean HeLght;" Crown diam.a 56.3 60.9 37.3 48.7 49.5 49.0 33.8 58.8 47.7 25.3 24.2 47.0 Totals N points N p1ap.t_~ Relative fr_egll~n~_ Relative density· Importance va1ueb Absolute densityc 5 9 3 4 2 22 6 30 3 6 2 49 11.1 20.0 6.7 8.9 4.4 48.9 6.2 31.2 3.1 6.2 2.1 51.0 17.3 51.2 9.8 15.1 6.5 99.9 260 1,307 130 260 88 2,136 45 96 100.0 99.8 199.8 4,181 aCentimeters. bSum of relative frequency and relative density. cPlants [ti«, Table 23. Vegetative characteristics of mountain shrub habitats at 24 female sharp-tailed grouse use sites, shrub height >1.0 m, Hayden study area, Routt County, Colorado. Species Heighta Ame1anchier sp. Artemisia tridentata Prunus virginianus Rosa woodsii Symphoricarpus sp. 209.2 124.0 121.8 130.0 115.3 Totals Mean Crown diam.a 186.9 138.2 33.0 78.5 119.3 N points N plants Relative frequency Relative ~e~~it~_ Importance va1ueb Absolute densityC 23 5 2 1 3 79 8 5 1 3 67.6 14.7 ·5.9 2.9 :8.8 82.3 8.3 5.2 1.0 3.1 149.9 23.0 11.1 3.9 11.9 117 12 7 1 4 34 96 99.9 99.9 199.8 141 aCentimeters. bSum of relative frequency and relative density. cPlants/ha. N -....J I-' �N -....J Table 24. Vegetative characteristics northwest Colorado, 1982-85. Species Amelanchier sp. Artemisia tridentata Chrlsothamnus sp. Gutierrezia sarothrae ,Mahonia repens Prunus virginianus Symphoricarpus sp. Totals aplants/ha. Relative frequency N of mountain shrub habitats at 8 sharp-tailed grouse nest sites, Shrub height <1.0 m Relative Absolute densitya density Relative frequency Shrub height >1.0 m Relative Absolute density a density 7.3 36.6 2.4 7.3 7.3 7.3 39.0 3.1 40.6 1.0 4.2 7.3 3.1 43.8 973 12,746 314 1,319 2,292 973 13,750 52.0 24.0 N/A N/A N/A 12.0 12.0 60.3 20.6 N/A N/A N/A 14.7 4.4 118 40 0 0 0 29 9 99.8 100.1 32,367 100.0 100.0 196 �273 nest sites than at sites 10 m from nests or at random sites on either study area. Tall shrubs (>1.0 m) averaged 50% higher density. Apparently, females selected clumps of mountain shrub vegetation which were denser than surrounding vegetation. Only 1 nest was found in habitat other than mountain shrub, this nest was in a clump of alfalfa in a hay pasture. The preference for shrubs, especially snowberry has been reported elsewhere (J. Monarch, pers. commun., Oedoekoven 1985; Marks and Marks 1987). An earlier study in Utah (Hart et ale 1950) reported the majority of nests in alfalfa, wheat or stubble fields, possibly the result of extremely heavy grazing of native rangelands. Hunter Harvest Hunting seasons for sharp-tailed grouse in northwest Colorado opened the 2nd Saturday in September (1981-85) and varied from 16 days (1981-83) to 23 days (1985) to 30 days (1984) in length. During this period, 673 sharp-tailed grouse wings were collected at 4 check stations, 19 wing barrels, and field checks of hunters (Table 25). Most wings came from southern Routt (N = 338, 50.2%) and northern Routt County (N = 314, 46.7%) with relatively few wings received from eastern Moffat County or elsewhere. The distribution of wings received is believed to fairly represent the distribution of sharp-tailed grouse hunters and harvest and coincides with the distribution and relative abundance of sharp-tailed grouse in northwest Colorado. Within each Game Management Unit, harvest and presumed hunting pressure varied. For example, in northern Routt County over 70% of the wings received were from California Park although this small area represents <2% of the area and <5% of the sharp-tailed grouse habitat in Units 4, 5, 14, 214, and 441. The relatively high hunting pressure in this area is due to hunter tradition (several hunting parties traditionally camp there on the initial weekend of the hunting season), the availability of public lands in the area, and the availability of sage grouse and blue grouse (Dendragapus obscurus) which provide additional recreational opportunity. Although there are larger sharp-tailed grouse populations which are equally accessible to hunters, most are on privately-owned lands and are unknown to most hunters. This pattern of unequal hunting pressure and harvest is also true for Unit 13, 26, 131, and 231 (southern Routt County) with most of the harvest centered around 20-mile Park. Except for 1984 when the season length was 30 days, over 50% of the sharp-tailed grouse harvest occurred during the initial weekend of the season (Table 26). One possible reason is that grouse hunters in Colorado, especially sage grouse and sharp-tailed grouse hunters, have traditionally experienced extremely short season lengths. Prior to 1978, the sage and sharp-tailed grouse season in Routt and Moffat counties was 3 days forcing all hunters to participate on the opening weekend or miss the season entirely. The effect of longer hunting seasons has been to spread out hunter pressure and harvest over time, without any apparent increase in total harvest. This is similar to the effects of longer sage grouse hunting seasons on sage grouse harvest (Braun and Beck 1985). There is no evidence from Colorado or elsewhere that hunter harvest is selective for any age or sex class of sharp-tailed grouse. Age and sex �Table 25. Origin of sharp-tailed grouse wings, northwest Colorado, 1981-85. 1981 Location Units 4, 5, 14, 214, 441 California Park check station Gould check station Black Mountain wing barrel Ralph White Res. wing barrel Moffat Co. Rd. 29 wing barrel Hayden Cog Rd. wing barrel California Park wing barrel Elk Mountain wing barrel Hahn's Peak Rd. wing barrel Fields checks Subtotal Units 3, 301 Cedar Mountain check station Cedar Mountain wing barrel Moffat Co. Rd. 3 wing barrel Subtotal N % Totals N 1984 1983 % 35 0 0 5 24.6 0.0 0.0 3.5 34 0 0 0 19.1 0.0 0.0 0.0 N/A N/A N/A N/A N/A N/A 2 1.4 9 3 5.1 1.7 N/A N/A N/A N/A 0 24 0.0 16.9 1 17 66 46.5 1 0 2 3 N % N N 1985 % 0.6 9.6 45 2 1 0 11 3 16 16 17 10 20.1 0.9 0.4 0.0 4.9 1.3 7.1 7.1 7.6 4.5 15 0 0 5 0 9 1 0 0 7 25.0 0.0 0.0 8.3 0.0 15.0 1.7 0.0 0.0 11.7 64 36.0 121 54.0 37 0.7 0.0 1.4 0 0 0 0.0 0.0 0.0 4 5 0 1.8 2.2 0.0 2.1 0 0.0 9 26.8 4.2 3.5 0.0 6.3 8.5 2.1 0.0 24 7 36 2 28 13 0 0 13.5 3.9 20.0 1.1 15.7 7.3 0.0 0.0 N/A N/A 0.0 0.0 0.0 0 1 3 73 51.4 142 100.0 Units 12, 13, 15, 26, 35, 36, 131, 231 Twentymi1e Park check station 38 Twentymi1e Park wing barrel 6 Hayden Gulch wing barrel 5 Milner wing barrel 0 Steamboat Springs wing barrel 9 Oak Creek wing barrel 12 Wolcott wing barrel 3 Rock Creek wing barrel 0 N/A Sage Creek wing barrel Five Pines South wing barrel 0 Muddy Creek Wing Barrel 0 Field checks 0 Subtotal 1982 N -...J % 10 0 14.5 0.0 N/A N/A 0 0 0 10 1 2 3 0.0 0.0 0.0 14.5 1.4 2.9 4.3 61.7 26 37.7 2 0 0 3.3 0.0 0.0 6 0 0 8.7 0.0 0.0 4.0 2 3.3 6 8.7 3 15 23 3 24 9 3 3 1.3 6.7 10.3 1.3 10.7 4.0 1.3 1.3 N/A N/A N/A N/A 5 7 8.3 11.7 22 2 31.9 2.9 N/A N/A N/A N/A N/A N/A N/A 0.0 0.6 1.7 0 0 11 0.0 0.0 4.9 3 0 0 0 0 0 0 6 5.0 0.0 0.0 0.0 0.0 0.0 0.0 10.0 0 7 0 0 0 1 0 5 0.0 10.1 0.0 0.0 0.0 1.4 0.0 7.2 114 64.1 94 42.0 21 35.0 37 53.6 178 100.1 224 100.0 60 100.0 69 100.0 .p.. �Table 26. Time distribution of sharp-tailed grouse wings received, northwest Colorado, 1981-85. 1981 Period First weekend First week Second weekend Second week Third weekend Third week .Fourth weekend Fourth week Fifth weekend Totals N 1982 % N 73.2 104 90 26 18.3 22 4.9 19 7 2.1 18 3 2 1.4 29 ------------------------------------------------------------------------------------142 99.9 178 1983 % N 1984 % 50.6 103 46.0 12.4 12.9 29 10.7 37 16.5 10.1 2.2 5 16.3 22.3 50 closed ---------------------closed ---------------------closed ---------------------closed ---------------------100.0 224 99.9 N 1985 % 22 7 10 6 2 0 1 4 4 36.7 11. 7 16.7 10.0 3.3 0.0 1.7 13.3 6.7 60 100.1 N % 62.3 43 11 15.9 7.2 5 2 2.9 1 1.4 0 0.0 7 10.1 -- closed -- closed -69 99.8 N -...J V1 �276 composition of hunter-harvested birds should reflect the actual structure of the fall population. Harvest data from 1976 to 1985 indicate that young of the year typically comprise over 50% of the fall population and the sex ratio of juveniles and adults does not differ from 1:1 (0.25 < P < 0.5) (Table 27). This suggests that if there are equal sex ratios at hatch, there is an equal mortality rate for males and females. The proportion of young in the harvest indicates the total year-to-year population turnover is approximately 58%, assuming a stable population. Studies of banded sharptails suggests that annual population turnover may exceed 70% (Robel et al. 1972). Hickey (1955) suggested that hunter harvest rates equaling 50% of the annual population turnover rate could occur without reducing grouse populations. Thus a safe harvest level for sharp-tailed grouse in northwest Colorado would be 25% of the fall population. Table 27. Age and sex composition of harvested sharp-tailed grouse, northwest Colorado, 1976-85. Year Adultsa % N 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 10 47 13 32 25 83 60 74 29 19 Totals 71.4 65.3 46.4 40.5 39.7 58.4 33.9 33.0 48.3 27.5 393 10-year average (%) Young N 4 25 15 47 38 59 117 150 31 50 % 28.6 34.7 53.6 59.5 60.3 41.6 66.1 67.0 51.7 72.5 534 42.4 alncludes yearlings. bKnown sex only. 57.6 Sample size Adultsa,b Males Females 14 72 28 79 63 142 177 224 60 69 5 5 1 3 14 9 5 3 2 927 4 Youngb Males Females 6 4 18 7 7 8 2 7 3 13 24 18 3 3 10 3 15 19 23 5 7 42 51 77 86 45.2 54.8 47.2 52.8 �277 LITERATURE CITED Ammann, G. A. 1944. Determining the age of pinnated and sharp-tailed grouse. J. Wildl. Manage. 8:170-171. 1957. The prairie grouse of Michigan. Tech. Bull. 200 pp. Amstrup, S. C. 214-217. 1980. A radio-collar for game birds. Michigan Dep. Conserve J. Wildl. Manage. 44: Artmann, J. W. 1970. Spring and summer ecology of the sharp-tailed grouse. Ph.D. Thesis, Univ. Minnesota, St. Paul. 129 pp. Beck, T. D. I., and C. E. Braun. 1980. The strutting ground count: variation, traditionalism, management needs. Proc. West. Assoc. Fish and Wildl. Agencies. 60:558-566. Braun, C. E., and T. D. I. Beck. 1985. Effects of changes in hunting regulations on sage grouse harvest and populations. Pp. 335-343 in S. L. Beasom and S. F. Roberson, eds. Game Harvest Management. Caesar~eberg Wildl. Res. Inst., Kingsville, TX. 374 pp. Cannon, R. W., and F. L. Knopf. prairie grouse populations. 1981. Lek numbers as a trend index to J. Wildl. Manage. 45:776-778. Christensen, C. D. 1970. Nesting and brooding characteristics of sharp-tailed grouse (Pedioecetes phasianellus jamesi Lincoln) in southwestern North Dakota. M.S. Thesis, Univ. North Dakota, Broqkings. 53 pp. Cottom, G., and J. T. Curtis. 1956. The use of distance measures in phytosociological sampling. Ecology 37:451-460. Dix, R. L. 1961. An application of the point-centered quartet method to the sampling of grassland vegetation. J. Range. Manage. 14:63-69. Giesen, K. M., and D. M. Hoffman. 1981. Distribution and status of mountain sharp-tailed grouse. Final Rep. Colorado Div. Wildl. Fed. Aid. Proj. W-37-R. April 1981. Pp. 183-189. __________ , T. J. Schoenberg, and C. E. Braun. 1982. Methods for trapping sage grouse in Colorado. Wildl. Soc. Bull. 10:224-231. Gratson, N. W. 1982. Habitat, mobility, and social patterns of sharp-tailed grouse in Wisconsin. M.S. Thesis, Univ. Wisconsin, Stevens Point. 91 pp. Gullion, G. W. 1961. A technique for winter trapping of ruffed grouse. Wildl. Manage. 29:109-116. J. Hart, C. M., O. S. Lee, and J. B. Low. 1950. The sharp-tailed grouse in Utah. Its life history, status, and management. Utah Dep. Fish and Game, Fed. Aid Div. Publ. 3. 79 pp. �278 Henderson, F. R., F. W. Brooks, R. E. Wood, and R. B. Dahlgren. 1967. Sexing of prairie grouse by crown feather patterns. J. Wildl. Manage. 31:764-769. Hickey, J. J. 1955. Some American population research on gallinaceous birds. Pages 326-396 in A. Wolfson, ed. Recent studies in avian biology. Univ. of Illinois Press, Urbana. 479 pp. Hillman, C. N., and W. W. Jackson. 1973. The sharp-tailed grouse in South Dakota. South Dakota Dep. Game, Fish and Parks Tech. Bull. 3. 64 pp. Hjorth, I. 1970. Reproductive behavior in Tetraonidae with special reference to males. Viltrevy 7:185-596. Hoffman, R. W., and C. E. Braun. 1975. A volunteer wing collection station. Colorado Div. Wildl. Game Infor. Leafl. 101. 3 pp. Jones, R. E. 1968. A board to measure cover used by prairie grouse. Wi1dl. Manage. 32:28-31. J. Kessler, W. B., and R. P. Bosch. 1981. Sharp-tailed grouse an4 range management practices in western rangeland. Pp. 133-146 in J. M. Peek and P. D. Dalke, eds. Proc. Livestock Relationships Symp., Univ. Idaho For., Wildl. and Range Expt. Stn. 10. Kirsch, L. M., A. T. Kleff, and H~ W. Miller. 1973. grouse population relationships in North Dakota. 37:449-453. Kobriger, G. D. parameters. Land use and prairie J. Wildl. Manage. 1975. Correlation of sharp-tailed grouse population North Dakota Outdoors 28(5):10-13. Marks, J. S., and V. S. Marks. 1987. Habitat selection by Columbian sharptailed grouse in west-central Idaho. U.S. Dep. Inter., Bur. Land Manage., Boise District, Idaho. 115 pp. Martin, S. A., and F. L. Knopf. 1981. Aerial survey of greater prairie chicken leks. Wildl. Soc. Bull. 9:219-221. Mattise, S. N. 1978. Effects of grazing systems on sharp-tailed grouse habitat. M.S. Thesis, South Dakota State Univ., Brookings. 46 pp. Miller, G. C., and W. D. Graul. 1980. Status of sharp-tailed grouse in North America. Pages 18-28 in P. A. Vohs, Jr., and F. L. Knopf, eds. Proc. Prairie Grouse Symp. Oklahoma State Univ., Stillwater. Mohr, C. O. 1947. Table of equivalent populations of North American small mammals. Am. MidI. Nat. 37:223-249. Moyles, D. L. J., and D. A. Boag. 1981. Where, when, and how male sharptailed grouse establish territories on arenas. Can. J. Zool. 59:1576-1581. Nielsen, L. S. 1978. The effects of rest-rotation grazing on the distribution of sharp-tailed grouse. M.S. Thesis, Montana State Univ., Bozeman. 52 pp. �279 Oedekoven, O. o. 1985. Columbian sharp-tailed grouse population distribution and habitat use in south central Wyoming. M.S. Thesis, Univ. Wyoming, Laramie. 58 pp. Pepper, G. W. 1972. The ecology of sharp-tailed grouse during spring and summer in the aspen park1ands of Saskatchewan. Saskatchewan Dep. Nat. Resour. Wi1d1. Rep. 1. 55 pp. Ramharter, B. G. 1976. Habitat selection and movements of sharp-tailed grouse hens during the nesting and brood rearing periods in a fire maintained brush prairie. Ph.D. Thesis, Univ. Minnesota, St. Paul. 78 pp. Rippin, A. B., and D. A. Boag. 1974. Recruitment to populations of male sharp-tailed grouse. J. Wi1d1. Manage. 38:616-621. Robel, R. J., F. R. Henderson, and W. Jackson. 1972. Some sharp-tailed grouse population statistics from South Dakota. J. Wi1d1. Manage. 36:87-89. Rogers, G. E. 1969. The sharp-tailed grouse in Colorado. Game, Fish and Parks, Tech. Pub1. 23. 94 pp. Colorado Div. Tanner, W. D., and G. L. Bowers. 1948. A method for trapping male ruffed grouse. J. Wi1d1. Manage. 12:330-331. Twedt, C. M. 1974. Characteristics of sharp-tailed grouse display grounds in the Nebraska sandhi11s. M.S. Thesis, Univ. Nebraska. 72 pp. Yde, C. A. 1977. Effects of rest-rotation grazing on the abundance and distribution of sharp-tailed grouse. M.S. Thesis, Montana State Univ., Bozeman. 70 pp. Ziegler, D. L. 1979. Distribution and status of the Columbian sharp-tailed grouse in eastern Washington. Completion Rep., Washington Fed. Aid Wi1d1. Rest. Proj. W-70-R-18. 26 pp. Prepared by Kenneth M. Giesen Wildlife Researcher ��281 Colorado Division of Wildlife Wildlife Research Report April 1987 JOB PROGRESS REPORT State of Colorado Project 01-03-045 (W-37-R) 14 Work Plan Job Title: 3 ----- Seasonal Movement and Habitat Use By Greater Prairie-chickens Period Covered: Author: : Job Avian Research 01 January through 31 December 1986 Michael A. Schroeder Personnel: Mike A. Schroeder and Gary White, Colorado State University; Clait E. Braun, Jack Corey, Francis Pusateri, and Mary Rasm~ssen, Colorado Division of Wildlife· ABSTRACT Investigations were begun to examine seasonal movements and habitat use of greater prairie-chickens in northeastern Colorado. Preliminary results suggest that during the breeding season, female prairie-chickens move throughout home ranges that often encompass 2 or more leks. Females tended to have larger home ranges than males during most seasons. Females nested an average of 2.67 km from the lek at which they were captured. Although most males seemed to display site fidelity to a particular lek, a few males moved between 2 or more leks. Habitat use was variable within each season, and differences between observed and available habitat sites were unclear. Additional work is planned to increase sample sizes for all seasonal categories. ��283 SEASONAL MOVEMENT AND HABITAT USE BY GREATER PRAIRIE-CHICKENS Mike A. Schroeder P. N. OBJECTIVES 1. Quantify seasonal habitat use, movements, and lek attendance of greater prairie-chickens in northeastern Colorado. SEGMENT OBJECTIVES 1. Review literature on Tetraoninae, prairie grouse, greater prairiechickens, habitat selection, lek behavior, dispersal, and migration. 2. Trap and mark all greater prairie-chickens on up to 4 leks. 3. Radiomark up to 50 (30 females, 20 males) greater prairie-chickens. 4. Relocate all radio-marked greater prairie-chickens at least 20 times per season. 5. Prepare vegetative cover map of the entire study area. 6. Describe vegetation (macro and micro habitat analysis) at all pra1r1echicken relocation sites. An equal number of random sites will be described in the same interval as measurements at use sites. 7. Document reproductive parameters, movements, and survival for all radiomarked greater prairie-chickens. 8. Compile and analyze data and prepare progress reports. METHODS An area centered 20 km northwest of Wray, Colorado was chosen for research on greater prairie-chickens. A study area of approximately 200 km2 was monitored during March-May to determine lek density and male lek attendance (Table 1). Trapping efforts were concentrated on a smaller core area of 75 km2• Birds were captured using walk-in traps and cannon nets between March and June 1986 (Table 2). All captured birds were banded with a numbered aluminum band and a unique combination of 3 colored plastic bands. The age category was generally determined by examining patterns of feather wear (Ammann 1944). Additional measurements suggested that age categories were relatively distinct (Table 3). Bird ages were: yearlings, 5 to 17 months of age (1 Nov of first year to 31 Oct of second year) and adults, older than 17 months of age (after 31 Oct of second year). Five battery- and 36 solarpowered radio transmitters were attached to poncho-type markers (Amstrup 1980) and placed on 15 males (9 adults, 6 yearlings) and 26 females (13 adults, 13 yearlings). Radio weights ranged between 1.8 and 2.3% of each birds' body weight. �Table 1Location of active greater prairie-chicken leks on study area in northeastern Colorado and the median number of males attending the lek during the breeding season (a plus sign symbolizes an active 1ek without an accurate count of males). Universal Transverse Mercators Meters East Meters North 712080 714880 716370 716610 716670 717050 717100 717150 717250 718280 718580 719070 719340 719920 720160 720980 721900 722100 722840 722990 723300 723760 724110 724270 724420 724830 724960 725280 725870 726290 726520 726700 727280 727450 728280 728630 728730 730390 731000 732420 733080 4453140 4453420 4453120 4456480 4453840 4455600 4457710 4448870 4454290 4450000 4453690 4445600 4451710 4449630 4453060 4458240 4455230 4453700 4456700 4456570 4458750 4450330 4453510 4459920 4449180 4448130 4447950 4450980 4445950 4447050 4455730 4450140 4444810 4454630 4447390 4448610 4445330 4446390 4447120 4445100 4446330 Township (North) 3 3 3 3 3 3 3 2 3 2 3 2 3 2 3 3 3 3 3 3 3 2 3 3 2 2 2 3 2 2 3 2 2 3 2 2 2 2 2 2 2 Legal location Range (West) Section 46 46 46 46 46 45 45 45 45 45 45 45 45 45 45 45 45 45 45 45 45 45 45 45 45 45 45 45 45 45 44 44 44 44 44 44 44 44 44 44 44 27 25 25 13 24 18 7 7 19 6 29 20 32 5 28 9 22 27 15 15 2 2 26 2 2 11 12 36 13 13 18 6 19 19 17 8 20 16 16 22 14 guarter NW NW NE NE SE SW SW NW SW NE NW NW NE SE SW NE NW NW NE NE SW SW NE NE SE SE SW SW SE NE SW NW SE SE NW NW NW SW NE SE SW Attendance 3 + 3 5 6 3 6 7 14 8 3 + 4 14 + 7 15 11 4 2 4 + 2 5 10 11 7 7 3 2 11 8 7 + 3 7 9 7 8 6 4 N CXl ~ �285 Table 2. Distribution of 81 captured greater prairie-chickens by sex, age, and capture technique during 1986 in northeastern Colorado. Category Walk-in traps Radios~ Bands Cannon nets Radios Bands Totals Radiosa Bands Males Adults Yearlings 5 3 2 12 8 4 11 7 4 6 4 2 16 10 6- 18 12 6 Females Adults Yearlings 22 9 13 20 13 7 5 2 3 0 0 0 27 11 16 20 13 7 Totals 27 32 16 6 43 38 aBirds given radios were also banded. The seasonal collection periods were defined as early spring (15 Feb-31 Mar), late spring (1 Apr-15 May), early summer (16 May-30 Jun) , late summer (1 Ju115 Aug), fall (16 Aug-31 Oct), and winter (1 Nov-14 Feb); designation of seasons was based on aspects of breeding behavior and movement (Robel et a1. 1970). Radio-marked prairie-chickens were tracked using a portable receiver and a 3-e1ement yagi antenna. Each bird was visually observed once every 7-14 days. Numerous additional observations were obtained by triangulation; directional readings were obtained within 1.0 km of the target transmitters and at ang1es-of-incidence greater than 350 and less than 1450• Initial estimates of accuracy suggested that locations derived by triangulation had a 95% probability of being within 200 m of the actual location. All locations were recorded using Universal Transverse Mercator coordinates (nearest 10-m interval). Home range size was estimated as the area within a 75% probability contour generated with harmonic means for each radio-tracked bird for each season. Habitat was examined at both observed and 'available' sites. Available sites were chosen relative to observed sites (modification of a stratified sampling method), and were randomly selected within a 0.5-km circle (0.5 km was representative of a typical prairie-chicken flight distance) centered over the observed site. To eliminate problems associated with measuring habitat variables in a changing environment (e.g., snow cover, plant growth, grazing pressure), both sites were examined on the same day. Two 18-m perpendicular transects were established in the center of the site. The orientation of the initial transect was randomly determined. Ten point-intercept locations, 2 m apart, were located along each transect (total of 20 points). All plant species intercepted at each point were identified and recorded. A heightdensity-index (HDI) was recorded from a height of 1 m and at a distance of 4 m to one side of the transect for all 20 points. The HDI was recorded as the height of vegetation obstructed on a Robel Pole (to the nearest 5 cm). Heights for sand sagebrush (Artemisia fi1ifo1ia), grasses, and forbs were recorded to the nearest 5 cm. A single 25-m2 circle was centered on each site (2.82 m diameter) and all plant species within the circle were identified and recorded. Location, slope, and aspect were also recorded at each site. �N Table 3. Measurements of greater prairie-chickens Males H SD Primaries (cm) I II III IV V VI VII VIII IX X 18 18 18 18 18 18 18 19 18 19 11.86 12.19 12.82 14.15 16.67 17.43 17.52 17.40 16.40 13.02 0.42 0.27 0.25 0.47 0.46 0.47 0.44 0.29 0.38 0.34 Diameter (rom) Primary IX 7 3.53 Pinnae (cm) 16 7.99 Weight (g) 24 1039.8 Adults Yearlin~s x x SD 11 10 10 11 11 11 10 10 10 11 11.35 11.86 12.48 13.74 16.37 17.23 17.45 17.22 15.76 13.20 0.41 0.28 0.27 0.38 0.33 0.34 0.42 0.43 0.36 0.42 0.08 8 3.29 0.40 11 7.84 48.8 ()\ Females Adults Measurement oo captured in northeastern Colorado in 1986. N 13 999.2 Yearlings x SD 20 20 20 20 19 20 19 23 23 23 11.46 11.85 12.40 13.66 16.03 16.64 16.78 16.67 15.73 12.68 0.38 0.32 0.31 0.35 0.33 0.25 0.35 0.30 0.48 0.36 0.06 15 3.32 0.58 19 3.73 24.1 N 26 905.2 x SD 21 21 21 21 21 21 20 23 22 23 10.81 11.30 11.99 13.03 15.36 16.23 16.41 16.10 14.90 12.33 0.22 0.26 0.32 0.32 0.30 0.31 0.29 0.33 0.36 0.40 0.15 15 3.17 0.12 0.52 21 3.76 0.63 72.0 N 26 896.7 42.1 �287 RESULTS Density Thirty-nine active leks were documented on the study area during 1986 (Table 4). The densit of leks on the study area was approximately 0.20 1eks/km2 (0.05 leks/mile). Densit of displaying males was approximately 1.22 ma1es/km2 (3.12 males/mile). Male attendance at leks appeared to be fairly constant from early March to early June, whereas female attendance (as represented by trapping success) showed a dramatic peak during the first 2 weeks of April (Table 4). 2 2 Table 4. Trapping success (percent captures per trap per day) for capturing. greater prairie-chickens in northeastern Colorado, 1986. Trapping week 04-10 11-17 18-24 25-31 01-07 08-14 15-21 22-28 Mar Mar Mar Mar Apr Apr Apr Apr (N Trap days traEs x da~s) 62 59 105 105 106 96 112 91 Males Rate Females 3.2 1.7 1.0 1.0 0.9 9.4 0.9 4.4 0.0 0.0 1.9 7.6 14.2 18.8 2.7 1.1 Total 3.2 1.7 2.9 8.6 15.1 28.1 3.6 5.5 Survival Radio-marked greater prairie-chickens were monitored an average of 87 days (some still being monitored) past the date of their capture. Twenty-one birds died; examination of kill characteristics suggested that 15 were taken by coyotes (Canus 1atrans), 2 by raptors (at least 1 Swainson's hawk, Buteo swainsoni), and 4 by unknown predators (as estimated from kill characteristics). Ten radio-marked birds were 'lost', either because of broken radios, or the birds moved too far from the study area to be found. Reproductive Success Twenty nests had a mean (median equal to mean) date of hatch of 10 June (some hatch dates were predicted based on the known onset of incubation). First nests had a mean hatch date of 1 June (N = 10), while probable, or known renests (based on the presence of a brood patch on a female visiting a 1ek or the known failure of a female's first nest), had a mean hatch date of 19 June (N = 10). Six nests (4 probable renests, 2 first nests) successfully hatched an average of 4.8 chicks (30% hatching success). Two of the broods did not survive 1 week. The other 4 broods (3, 4, 4, and 8 chicks) survived at least in part, past September (3, 1, 4, and 6 chicks). �288 Movement Sixteen radio-marked males were monitored. All but 3 (2 yearlings and 1 adult), exclusively visited the 1ek at which they were captured. One of the yearling males visited at least 5 leks. Females frequently visited more than 1 1ek and most had more than 2 leks within their home range. Twenty-one females nested an average of 2.67 km (SD = 2.15) from the 1ek where they were first captured and 0.86 km (SD = 0.49) from the nearest 1ek. Only 5 females nested closer to their 1ek of capture than to another 1ek. Home range size was estimated for radio-marked greater prairie-chickens for all seasons (Table 5). Females appeared to have larger home ranges than males during most seasons. Yearlings of both sexes tended to move more than adults. Home range size of females seemed to be influenced by their brood status. Four hens with broods had home ranges that averaged 499.7 ha, substantially larger than most other birds. During the early and late summer, numerous birds could not be locat.ed on the study area. Several birds were sub$equent1y found 9-48 km from the study area. As most of the birds (excluding brood females) appeared to move to their winter ranges in June and July, the long distance movements were tentatively considered migratory movements (tentative based on the assumption that they will return to their breeding areas in spring 1987). Analysis of distances between probable winter and breeding season ranges showed that females tended to move further (x = 13.8 km, N = 12, median = 5.0 km) than males ~ = 4.4 km, ~ = 5, median = 1.2 km). Habitat Habitat variables were compared between observed and available sites for nesting females (Table 6), during the late summer (Table 7), fall (Table 8), and winter seasons (Table 9). Few of 17 variables examined differed (probability level <0.05) for the late summer, fall, and winter seasons. Sample sizes were minimal for seasonal comparisons, which made differences difficult to detect. Greater prairie-chickens used a wide variety of basic habitat types (from corn fields to dense sand sagebrush), which increased the statistical variances. Differences between observed and available sites were detected during the nesting season when females appeared to select areas with taller vegetation and thicker cover, possibly for nest concealment. DISCUSSION Few data have been gathered on mortality, reproduction, 1ek attendance, movement, and habitat use by greater prairie-chickens in Colorado. Sample sizes in this study are still too small to infer clear conclusions, however certain trends are apparent. Males apparently have site fidelity to a particular 1ek and can be found on the same 1ek every day. However, little is known about the proportion of males remaining faithful to a particular 1ek and those that move between leks. �Table 5. Home range size (area within 75% probability contours generated with harmonic means, ha) by sex, age, and season for greater prairie-chickens in northeastern Colorado during 1986. Category Adults Males Yearlings All Adults Females Yearlings All All birds Early spring N X SD 4 26.2 15.1 0 4 26.2 15.1 2 210.6 64.5 1 26.6 3 149.3 115.6 7 78.9 94.3 8 99.9 65.0 4 407.1 621.8 12 202.2 362.0 10 335.2 302.4 11 701.2 985.9 21 526.9 749.8 33 408.9 649.3 5 160.7 115.0 3 280.3 148.4 8 205.6 133.0 9 208.2 186.7 6 210.5 208.1 15 209.1 188.1 23 207.9 167.8 4 96.3 39.4 2 122.6 6.1 6 105.1 33.5 6 180.4 163.7 6 216.9 143.3 12 198.6 147.9 18 167.4 128.7 2 152.2 150.0 2 216.8 113.2 4 184.5 114.7 4 592.8 531.6 6 252.9 204.8 10 388.9 385.1 14 330.5 338.9 1 413.5 0 1 413.5 3 413.8 530.2 0 3 413.8 530.2 4 413.8 432.9 Late spring N X SD Early summer N X SD Late summer N - X SD Fall N - K SD Winter N X SD N co \0 �290 Table 6. Habitat at nest sites and available locations of greater prairiechickens in northeastern Colorado in 1986. Observed (24) SD ~ Habitat variable a P-va1ue t Fmax 4.47 2.14 2.63 1.79 >0.05 0.002 Height (cm) Sand sagebrushb Grasses Forbs 57.92 99.17 46.88 28.05 17.55 27.50 39.38 84.79 44.17 26.72 21.94 23.80 >0.05 >0.05 >0.05 0.023 0.016 0.717 Cover (%) Bare Sand sagebrush Grasses Cool season grasses Need1e-and-threadc Six weeks fescued Warm season grasses Blue gramae Sand dropseedf Prairie sandreedg Forbs 12.08 9.38 76.04 37.92 22.92 14.38 43.96 10.00 13.75 11.88 11.25 7.65 6.13 14.89 24.27 17.69 14.91 18.94 11.42 13.21 9.87 9.12 20.21 7.71 67.92 28.75 15.83 13.54 45.63 11.04 10.21 8.54 10.21 15.84 8.34 17.75 17.83 15.51 13.95 20.50 11.13 13.14 12.02 8.14 <0.05 >0.05 >0.05 >0.05 .:>0.05 >0.05 >0.05 >0.05 >0.05 >0.05 >0.05 0.434 0.093 0.143 0.147 0.842 0.771 0.750 0.357 0.299 0.678 Species richness (N) 18.13 5.06 17.92 4.79 >0.05 0.884 2.17 1.97 2.54 2.75 >0.05 0.590 Height-density-index Slope (degrees) (cm) Available (24) x SD alf the Fmax test was significant, the t test was not done. bSand sagebrush = Artemisia fi1ifo1ia. cNeed1e-and-thread = Stipa comata. dSix weeks fescue = Vu1pia octof1ora. eB1ue grama = Boute1oua gracilis. fSand dropseed = Sporobo1us cryptandrus. gPrairie sandreed - Ca1amovi1fa longifo1ia. �291 Habitat at observed and available locations of greater prairieTable 7. chickens in northeastern Colorado during late summer 1986. Observed (40) SD x Habitat variable Available (40) SD x a P-va1ue t Fmax 4.38 2.60 4.03 3.23 >0.05 0.590 Height (cm) Sand sagebrush Grasses Forbs 45.88 92.75 49.50 19.18 16.41 20.72 36.50 88.13 48.13 18.72 17.86 22.47 >0.05 >0.05 >0.05 0.030 0.231 0.777 Cover (%) Bare Sand sagebrush Grasses Cool season grasses Need1e-and-thread Six weeks fescue Warm season grasses Blue grama Sand drop seed Prairie sandreed Forbs 13.63 9.75 74.00 32.38 21.88 12.38 50.88 13.13 8.63 14.50 14.00 8.09 9.60 15.62 17.47 15.96 10.38 15.10 8.96 9.13 9.86 11.56 16.88 7.75 71.75 35.13 22.63 10.88 44.50 14.00 7.38 11.75 13.75 11.36 9.67 18.86 18.76 17.90 8.39 16.90 13.60 7.59 9.03 12.65 <0.05 >0.05 >0.05 >0.05 >0.05 >0.05 >0.05 <0.05 >0.05 >0.05 >0.05 Species richness (N) 19.83 4.97 18.08 4.69 >0.05 0.109 2.38 2.46 2.93 3.18 >0.05 0.389 Height-density-index Slope (degrees) (cm) alf the !max test was significant, the t test was not done. 0.356 0.563 0.499 0.844 0.479 0.079 0.508 0.197 0.927 �292 Table 8. Habitat at observed and available locations of greater prairiechickens in northeastern Colorado during fall 1986. Observed (39) x SD Habitat variable Height-density-index P-va1ue a t Fmax 5.48 9.37 10.05 18.09 <0.01 Height (cm) Sand sagebrush Grasses Forbs 40.90 91.92 48.33 29.38 34.46 27.82 30.77 105.26 61.15 27.25 46.20 7.10 >0.05 >0.05 <0.01 0.119 0.153 Cover (%) Bare Sand sagebrush Grasses Cool season grasses Need1e-and-thread Six weeks fescue Warm season grasses Blue grama Sand dropseed Prairie sandreed Forbs 16.54 8.33 73.72 24.49 12.56 6.15 59.62 12.82 15.38 10.90 13.08 13.09 10.72 17.20 19.19 15.43 7.65 16.36 15.93 15.06 13.57 16.13 18.59 4.49 74.49 16.79 13.33 3.33 64.74 16.03 14.62 9.87 11.28 12.24 5.71 16.17 17.38 16.52 5.42 17.88 19.54 15.41 15.37 11. 79 >0.05 <0.01 >0.05 >0.05 >0.05 < 0.05 >0.05 > 0.05 >0.05 > 0.05 > 0.05 0.477 Species richness (N) 16.90 6.73 16.10 7.10 > 0.05 0.613 2.64 3.54 2.77 3.24 > 0.05 0.868 Slope (degrees) (cm) Random (39) x SD alf the !max test was signficant, the t test was not done. 0.839 0.067 0.832 0.190 0.430 0.824 0.756 0.576 �293 Table 9. Habitat at observed and available locations of greater prairiechickens in northeastern Colorado during winter 1986-87. Observed (11) x SD Habitat variable Available (11) x SD P-va1ue F a t 6.06 3.03 6.16 4.37 >0.05 0.951 Height (cm) Sand sagebrush Grasses Forbs 33.18 83.18 40.45 35.59 34.66 37.58 37.73 105.00 48.64 28.58 37.62 32.26 >0.05 >0.05 >0.05 0.745 0.173 0.590 Cover (%) Bare Sand sagebrush Grasses Cool season grasses Need1e-and-thread Six weeks fescue Warm season grasses Blue grama Sand dropseed Prairie sandreed Forbs 36.36 5.91 54.55 5.91 5.91 0.00 50.45 11.82 7.27 14.55 10.91 33.77 7.69 29.62 10.68 10.68 0.00 27.97 19.27 10.09 17.95 12.21 32.27 2.73 61.36 12.73 10.91 0.45 51.82 8.64 8.18 7.73 7.73 17.37 3.44 17.04 18.76 19.34 1.51 20.53 18.04 8.74 15.23 9.58 0.05 0.05 > 0.05 > 0.05 . > 0.05 < 0.01 > 0.05 > 0.05 > 0.05 > 0.05 > 0.05 Species richness (N) 9.73 7.04 11.36 6.52 > 0.05 Slope (degrees) 2.64 2.46 4.64 7.97 < Height-density-index (cm) alf the !max test was significant, the t test was not done. < < 0.01 0.516 0.307 0.462 0.898 0.694 0.824 0.348 0.504 0.578 �294 The extent of female movement during spring is an important and unanswered question. Female behavior and habitat requirements, especially those relating to nesting and 1ek breeding, have been difficult to examine. Although numerous hypotheses have been proposed to explain the evolution of 1ek behavior from a dispersed mating system, most of these hypotheses have been rejected (Davies 1978, Wittenberger 1978, Bradbury 1981, Bradbury and Gibson 1983, Payne 1984). Of 2 that remain, female preference for clustered males (Bradbury 1981), and settlement of males on 'hotspots' of female traffic (Bradbury et a1. 1986), both predict a positive correlation between female home range size and the spacing of male clusters (leks). The first hypothesis ('female preference' hypothesis), which suggests that each 1ek should have its own population of females, apparently is not supported by the preliminary findings of this study which suggests that most females visit more than one 1ek. Thus far, little evidence has been gathered that either supports or rejects the 'hotspot' hypothesis. Movement of females in relation to leks is significant with respect to greater prairie-chicken management. The fact that.females nested an average of 2.67 km from the 1ek at which they were captured, suggests that leks may not be an important indication of local habitat quality. Examination of habitat has been limited to a stratified sampling method that limits examination of available habitat to areas relatively close to the 'observed' habitat. Although certain differences were detected between observed and available habitat, especially during the nesting season, low sample sizes preclude a detailed analysis of seasonal habitat use. Certain difficulties exist with interpretation of these data. Biases associated with trapping are not clearly understood. Trapping methods improved as spring progressed in 1986. Consequently, numerous females were caught that had already failed with their first nesting attempt. The bias toward late trapping success might partially explain why the estimated hatching success rate was only 30% (previous research suggests that second nests are less successful than first nests, Robel and Ballard 1974). LITERATURE CITED Ammann, G. A. 1944. Determining the age of pinnated and sharp-tailed grouses. J. Wi1d1. Manage. 8:170-171. Amstrup, S. C. 214-217. 1980. A radio-collar for game birds. J. Wildl. Manage. 44: Bradbury, J. W. 1981. The evolution of leks. Pages 138-169 in R. D. Alexander and D. W. Tinkle, eds. Natural selection and social behavior: recent research and new theory. Chiron Press, New York, N.Y. , and ----=-~-P. Bateson, , ----~~-leks. Anim. R. M. Gibson. 1983. Leks and mate choice. Pages 109-138 in ed. Mate choice. Cambridge Univ. Press, Cambridge, Mass. , and I. M. Tsai. Behav. 34:1694-1709. 1986. Hotspots and the dispersion of �295 Davies, N. B. 1978. Ecological questions about territorial behavior. Pages 317-350 in J. R. Krebs and N. B. Davies, eds. Behavioral ecology: an evolutionary approach. Sinauer Associates: Sunderland, Mass. Payne, R. B. 1984. Sexual selection, 1ek and arena behavior, and sexual size dimorphism in birds. Ornith. Monogr. 33. 52 pp. Robel, R. J., and W. B. Ballard, Jr. 1974. Lek social organization and reproductive success in the greater prairie chicken. Am. Zool. 14:121-128. ____ ~ __--' J. N. Briggs, J. J. Cebula, N. J. Si1vy, C. E. Viers, and P. G. Watt. 1970. Greater pralrle chicken ranges, movements, and habitat usage in Kansas. J. Wi1d1. Manage. 34:286-306. Wittenberger, J. F. 80:126-137. Prepared by 1978. The evolution of mating systems in grouse. ~1JL::--=:-=·:-.:-U-:==-::-i1~,......:;~,.=----=--==--..:~_ Michael A. Schroeder Graduate Research Assistant Approved by ~C4i...::;-=v:::::::;;.---,-:::-~Z--,-;...__,o.~-,",--,,C1ait E. Braun Wildlife Research Leader _ Condor ��297 Colorado Division of Wildlife Wildlife Research Report April 1987 JOB PROGRESS REPORT State of: Project: Colorado W-152-R (W-37-R) Work .Plan: 17 Job Title: Population Dynamics of White-tailed Ptarmigan Period Covered: Personnel: : Job Avian Research 7 01 January through 31 December 1986 Clait E. Braun and Kenneth M. Giesen, Colorado Division of Wildlife ABSTRACT Long-term studies of populations of white-tailed ptarmigan (Lagopus leucurus) were continued at hunted (Mt. Evans) and unhunted (Rocky Mountain National Park) areas in Colorado through 1986. Densities of breeding ptarmigan decreased slightly at Mt. Evans and continued to decrease at Rocky Mountain National Park. Densities at this site were the lowest documented in the 1966-86 interval, the result of decreased adult survival and juvenile recruitment. Nesting success was variable at the 2 sites~ being good in Rocky Mountain National Park and below average at Mt. Evans. Harvest increased at Mt. Evans in 1986 (22.9% of the population banded in 1986 was harvested), and was twice that documented in 1985 (11.6%). ��299 POPULATION DYNAMICS OF WHITE-TAILED PTARMIGAN Clait E. Braun and Kenneth M. Giesen Long-term studies of trends in population size and investigation of reasons for fluctuations in size of tetraonid populations are lacking. Studies on the population dynamics of unhunted and hunted populations of white-tailed ptarmigan were initiated in Colorado in 1966 and have continued essentially uninterrupted at 2 sites. Studies of the unhunted population (Rocky Mountain National Park) have identified possible short-term cycles of 7-8 years with an amplitude of 25-30% between high and low breeding densities. Conversely, studies of the manipulated population (hunted) at Mt. Evans through 1980 have not indicated any cyclic pattern and it would appear that controlled hunting may mask any long-term trend that may occur. This study is designed to examine the question whether white-tailed ptarmigan are truly cyclic and whether hunting affects the apparent oscillations. P. N. OBJECTIVES The goals of this investigation are to be able to predict the length and amplitude of cycles in white-tailed ptarmigan in Colorado, to examine the impact of hunting on cycles, and to clarify underlying causes of the apparent cycles. SEGMENT OBJECTIVES 1. Conduct breeding (May-Jun) and brood (Aug-Sep) censuses of white-tailed ptarmigan using tape-recorded calls of males (breeding) and chicks (broods). 2. Censuses will be conducted on previously established, defined study areas at Mt. Evans (hunted) and at Rocky Mountain National Park (unhunted). 3. Capture (noose poles) and band (aluminum and plastic color-coded bands) all unmarked white-tailed ptarmigan encountered on study areas at Mt. Evans and at Rocky Mountain National Park. 4. Individually identify all ptarmigan observed on study areas at Mt. Evans and Rocky Mountain National Park through use of binoculars. 5. Make hunting season and bag limit recommendations for Mt. Evans and collect hunting data through use of volunteer wing barrels and hunter field checks. 6. Compile data, analyze results, and prepare progress reports. �300 STUDY AREA AND METHODS Areas investigated were Mt. Goliath-Mt. Evans in Clear Creek County and at Tombstone Ridge-Sundance Mountain to Fall River Pass in Rocky Mountain National Park in Larimer County. The physiography, geology, location, and vegetation of these study areas have been previously described (Braun 1969, 1971; Braun and Rogers 1971; Giesen 1977). Ptarmigan were located through use of tape-recorded calls (Braun et al. 1973), captured through use of telescoping noose poles (Zwickel and Bendell 1967) as described by Braun and Rogers (1971), and classified to age and sex and banded following Braun and Rogers (1971). Age of chicks was estimated following Giesen and Braun (1979). Numbered plastic bandettes were not used as in earlier years (Braun and Rogers 1971) as a color-code system using up to 4 different colored plastic bandettes was instituted in 1977-78. A check station was operated on the Mt. Evans highway during the opening weekend of the ptarmigan season in that area. A volunteer wing collection station was available to hunters in the area when the check station was not in operation until the season closed. RESULTS AND DISCUSSION Breeding Densities Mt. Evans.--Timing of breeding events in the Mt. Evans area was "normal" in 1986. During the late May-early June interval, 12 pairs and 2 single males were identified. Thus, breeding densities decreased slightly from levels measured in 1984-85. During the breeding season, only 4 of 14 males identified were yearlings while 4 of 12 females identified were yearlings. Thus, recruitment decreased in 1986 similarly to the decrease in recruitment documented in 1985. Rocky Mountain National Park.--Surveys of ptarmigan present on breeding territories along Trail Ridge Road in RMNP during May and June indicated the minimum breeding population was 28 birds, which included at least 12 pairs. This represents the lowest breeding density (4.5 birds/km2) since initiation of the long-term study in 1966 and a continuation of the population decline initially documented in 1977. The declining breeding densities appear to result from continuing low overwinter survival of adult hens and secondarily from low survival and recruitment of juveniles. Survival of banded adults from 1985 was 58% (29 of 43 males, 7 of 19 females). Recruitment of chicks banded in 1985 was 21% (4 of 19) although yearlings comprised 34.3% (24 of 70) of all adult ptarmigan identified in 1986. �301 Table 1. 1966-86. White-tailed ptarmigan breeding densities (birds/km2), Colorado Study area Year Rocky Mountain National Park (5.5 km2) 1966 1967 1968 1969 1970 1971 1972 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 1986 11.3 9.8 11.5 12.0 9.6 9.1 8.7 7.8 8.0 11.1 13.5 12.9 10.7 8.7 8.4 8.2 7.8 6.7 5.8 6.0 4.5 Mt. Evans (4.0 km2) 3.0 2.7 2.7 2.2 2.0 4.2 7.5 6.2 6.2 6.2 6.7 >6.0 7.5 10.3 9.5 9.0 6.5 6.5 8.0 8.0 6.5 Nesting Success and Brood Size Mt. Evans.--Nineteen hens were located during mid July-early September 1986 on or immediately adjacent to the study area. Seven hens (36.8%) were with broods while 12 were apparently unsuccessful nesters (without chicks). Average brood size to 1 September was only 2.4 chicks/successful female. Data were available from 22 chicks captured and banded or taken during the hunting season. Of this total, all but 1 (20 Jul) hatched between 6 and 15 July. The one late-hatched chick was probably the result of a renest. Rocky Mountain National Park.--July and August surveys indicated average nesting success (60%) in 1986 (6 of 10 hens identified were with broods). Hatch dates were estimated from primary molt and growth for 29 chicks captured for banding. Hatch dates ranged from 9 to 17 July (median = 14 Jul), similar to the long-term average. Survival of chicks to 1 September was excellent with an average August brood size of 4.8 chicks, the highest recorded since 1980. �302 Harvest Mt. Evans.--The hunting season at Mt. Evans in 1986 opened on 20 September and closed on 5 October (16 days) with a bag and possession limit of 3 and 6. Thus, the season was delayed 2 weeks from the statewide opening as it has been each year since 1978 (except 1981 which was delayed 1 week). A check station was operated from dawn to dusk on the opening weekend (20-21 Sep) similar to operations in previous years. In addition, a volunteer wing collection station was available through 5 October. During the 2-day check, 15 hunters with 21 ptarmigan were checked. Twelve of the 21 ptarmigan (59.1%) were banded. Bands from 2 other harvested ptarmigan were received during the 1986 hunting season. Thus, at least 11 of 48 (22.9%) ptarmigan banded in 1986 on the study area were harvested. Nine of the 21 birds (42.9%) checked on the opening weekend were chicks of which 4 (44.4%) were banded. The harvest rate in 1986 was about twice that documented in 1985. LITERATURE CITED Braun, C. E. 1969. Population dynamics, habitat, and movements of white-tailed ptarmigan in Colorado. Ph.D. Thesis, Colorad·State Univ., Fort Collins. 189 pp. _____ 1971. Habitat requirements of Colorado white-tailed ptarmigan. West. Assoc. State Game and Fish Comm. 51:284-292. , and G. E. Rogers. 1971. ----Colorado Div. Game, Fish and Proc. The white-tailed ptarmigan in Colorado. Parks Tech. Publ. 27. 80 pp. , R. K. Schmidt, Jr., and G. E. Rogers. 1973. Census of Colorado white-tailed ptarmigan with tape recorded calls. J. Wildl. Manage. 37:90-93. Giesen, K. M. 1977. Mortality and dispersal of juvenile white-tailed ptarmigan. M.S. Thesis, Colorado State Univ., Fort Collins. 55 pp. , and C. E. Braun. 1979. A technique for age determination of juvenile -----white-tailed ptarmigan. J. Wildl. Manage. 43:508-511. Zwickel, F. C., and J. F. Bendell. J. Wildl. Manage. 31:202-204. Prepared by t/JY ~ C aitE.Braun Wildlife Research Leader Kenneth M. Giesen Wildlife Researcher B 1967. A snare for capturing blue grouse. � https://cpw.cvlcollections.org/files/original/e9968d47cf076a3cb0674e6b5eebdce2.pdf 926d2bcca76f175291637355450f4d29 PDF Text Text Colorado Division Wildlife Research April 1987 303 of Wildlife Report JOB PROGRESS State of Colorado ----------------------------------- Project 01-03-045 21 Work Plan Job Title: Warren Personnel: (W-37-R) Avian Research 2 : Job Dynamics Period Covered: Author: REPORT of Cottonwood 01 January Regeneration through 31 December 1986 D. Snyder J. F. Corey, J. P. Palic, A. F. Palic, and W. D. Snyder, Division of Wildlife Colorado ABSTRACT Statistical analyses, including testing for differences among strata from early to recent inventories of tree stands, were completed by D. C. Bowden for the 4 major Colorado Rivers. Comparisons included canopy cover and age class among strata, individual age class-canopy cover categories by stratum, and combined data. Most sample sizes within strata were too small to provide meaningful data. Extensive and intensive transects of natural regeneration of plains cottonwoods (Populus sargentii) established in 1983-84 showed that while natural attrition within and among stands continued, survival within better sites remained good. Within intensive transects, survival averaged 76% in 1986 compared to 65% during the previous year. Green ash (Fraxinus pennsylvanica) continued to increase in importance along the lower South Platte River. Extensive transects also documented transition toward increased perennial herbaceous vegetation and reduced cover of annuals as soils and vegetation became stabilized after the 1983 flood. River flow and adjacent groundwater levels at stem cutting· sites along the South Platte continued to fluctuate dramatically in 1986 and were believed to be the primary factor in only moderate stem cutting survival which averaged 45% among all 1986 plantings. Although survival was much improved over previous years of testing, it remained below levels acceptable for recommending it to management personnel. This is especially true when considering the labor intensive efforts needed to place the cuttings below the lowest groundwater levels. Monitoring of controlled burns conducted by management personnel within the. South Platte riverbottom floodplain continued. These included impacts of fire on woody species and herbaceous cover, and the composition and height of herbaceous vegetation within plowed fireguards. Stands of tall seed-producing annuals within fireguards appeared to be one of the most beneficial aspects of the burns to wildlife. Characteristics of the Arkansas River in southeastern Colorado including channel width, channel depth, relief between adjacent mature trees and river bed, and stream flow characteristics were compared between sites below and above John Martin Reservoir. ��305 DYNAMICS OF COTTONWOOD REGENERATION Warren D. Snyder P. N. OBJECTIVES Quantify changes in stand density and changes in area of riparian cover types over a recent time span approximating 30 years within the South Platte, Arkansas, Colorado, Rio Grande, and South Republican (below Bonny Reservoir) rivers in Colorado. Analyses and evaluations will be based on aerial interpretation by the Colorado State Forest Service. Document conditions conducive to natural regeneration and survival of plains cottonwoods and willows, including site characteristics and frequency of occurrence, in streamside riparian habitats of the lower South Platte River in northeastern Colorado following high water conditions in 1983 and 1984. Test methods for establishing woody vegetation within streamside riparian zones where natural propagation cannot be expected. These include: (a) using stem cuttings of dormant specimens and planting them so their bases extend into the groundwater as a method for propagation of selected tree, shrub, and vine species for use where natural propagation no longer occurs, (b) create exposed bare ground sites using tillage or scarification for natural establishment of plains cottonwoods, peachleaf willow (Salix amygdaloides), or other woody vegetation. METHODS Primary methods used in this study were reported earlier (Snyder 1985, 1986). Supplemental or modified procedures are identified here. Modified planting methods were used in spring 1986 in efforts to place stem cuttings deeper into the groundwater and to remove competing vegetation. Competing vegetation was removed by tillage from small tracts (approximately 20 m/side) within the Red Lion, ll-E, and Check Station Meadow sites on the Tamarack Wildlife Area along the South Platte River. Subsequent to planting, these cultivated sites were overlaid with black plastic and wood chip mulch layers to eliminate competitive weeds and the need for subsequent tillage. A portion of the plantings was also placed in natural competitive vegetation adjacent to the cultivated sites for survival comparison. Within the Duck Creek site cultivation was not used, however, the herbicide, glyphosate (Roundup), was applied in mid spring to reduce competition from inland saltgrass (Distichlis stricta). Plastic sleeves were temporarily placed over the cuttings to prevent herbicide contact. The plastic-organic mulch treatment was applied to an approximate 1-2 m area surrounding one-half of the cuttings. The plastic organic mulch treatment was used around all plantings within the South Republican site. Holes for planting stem cuttings were dug with a tractor-mounted auger of l-dm diameter to a depth approximating 12 dm. A l-dm diameter plastic tube was then inserted and repeatedly pushed down as sand was extracted with a hand-operated sand auger. This permitted stem cutting placement to greater depths into the water table. �306 Sampling of dominant vegetation occurrence within fireguards and disturbance tillage strips included random selection of starting points and listing of dominant vegetation within 1-2 m samples at 5 or 10 step intervals (depending upon the size fo the tract) along the strip. Sampling was conducted in late September. A transit-level and rod were used to obtain stream profile samples along the Arkansas River. Samples were obtained at accessible locations within the random mile locations previously selected for sampling inventory data. Twelve locations above and 12 locations below John Martin Reservoir were sampled. A line was stretched across the channel to permit sampling at 1 and/or 2 m intervals. Channel width and vertical distance from river level to cottonwood tree bases were also obtained. RESULTS Analyses of Cottonwood Inventory Data Preliminary analyses of the photo-interpretation inventory data revealing status and trends of cottonwood stands, shrubs, and other vegetation or land use types along lower portions of Colorado's 4 major rivers were previously reported (Snyder 1984, 1985). More detailed analyses of the cottonwood data have since been completed by D. C. Bowden, with strata being the primary basis for analyses. Findings are summarized in the following text and tables and reference is made to data in Snyder (1984, 1985) for a more complete perspective. Arkansas River.--Attempts to analyze early to recent changes in hectares of cottonwood stands within individual strata by age class-canopy cover (cc) classification did not provide meaningful data as mean changes were highly variable (both + and -) and probability values tended to be high (Table 1). Sample sizes within strata were not adequate, and the same findings occurred for all 4 rivers. When strata were combined, the most consistent declines of hectares and probabilities were within the 35-55 cc c1assificatin (Table 1). Ignoring age class-canopy cover, the downstream (3rd) stratum (below John Martin Reservoir) contained the greatest average decreases in hectares of cottonwoods with the greatest reliability (p = 0.03, Table 2). Combined data, ignoring strata, revealed a highly significant overall decline in stands of trees along the lower Arkansas River. Analyses were also conducted within and among strata considering cc but summing over age class (Table 3) and vice versa (Table 4). Due to the high variability in mean changes among strata and among cc (Table 3), few meaningful data could be derived. In general, it appeared that the most pronounced changes occurred within the 35-55% cc level and within the 3rd (downstream) stratum. Among age classes averaged over the 3 strata, marked changes (P<0.05) were noted in the older age classes but differences among strata were not evident (Table 4). �307 Table 1. Mean changes (ha) of cottonwood stands from early to recent intervals within 3 sampling strata and combined strata along the lower Arkansas River. x Stratum P x P C1Aa 1 2 3 Combined 4.29 -0.14 -0.44 0.63 C1B 0.15 0.78 0.23 0.25 -1.04 -0.53 -0.01 -0.43 8.21 -0.26 -3.84 0.002 0.08 -2.96 -5.17 -3.23 -4.50 -0.19 -1.84 -1.71 C -0.85 -0.64 0.10 -0.39 -0.39 0.31 -0.60 -0.12 -0.71 -0.46 C3B 0.27 0.93 0.49 0.22 -7.51 -1.66 1.13 -1.71 C4A 1 2 3 Combined 0.38 0.35 C2C 0.97 0.26 0.17 0.06 C3A 1 2 3 Combined -1.83 -0.06 C2B 0.03 0.86 0.04 0.998 0.01 0.10 0.53 0.10 -1.63 -0.93 -0.44 -0.88 0.41 0.82 0.12 0.11 C3C -3.70 0.07 0.005 0.94 -0.71 0.07 C4B 0.05 0.03 0.86 0.12 p C1C 0.10 0.10 0.36 0.01 C2A 1 2 3 Combined x C4C 0.15 0.02 0.36 0.005 0.10 0.14 0.81 0.28 0.08 0.39 aCanopy cover (%) and age class of cottonwoods: A = < 35%, B = 35-55%, 55% canopy cover; 1 = < 6, 2 = 6-15, 3 = 16-30, 4 = >30 in. dbh. Table 2. Changes (ha) from early to recent intervals in cottonwood stands among strata along the lower Arkansas River, Colorado. Stratum x P 1 -9.00 -7.30 -11.20 -9.21 0.27 0.15 0.03 0.002 2 3 Combined average �308 Table 3. Changes (ha) in cottonwoods among strata from early to recent intervals by canopy cover along the lower Arkansas River, Colorado. Canopy cover (%) <35 Stratum 2 3 7.10 0.22 -1.23 0.68 -6.03 0.02 -1.48 0.36 -10.10 0.07 -6.08 0.02 -4.50 0.25 -6.25 0.003 -6.03 0.08 0.03 0.98 -0.71 0.12 -1.48 0.05 1 x P 35-55 ~ P z. >55 P Combined x Changes (ha) in cottonwoods among strata from early to recent Table 4. intervals by age class along the Arkansas River. Age class (in. dbh) <6 :R P 6-15 :R P 16-30 x P >30 x P 1 Stratum 2 3 x 1.42 0.74 -0.73 0.29 -0.46 0.23 -0.19 0.81 7.68 0.06 -3.33 0.22 -9.73 0.04 -3.69 0.06 -15.70 0.06 -1.78 0.55 -0.71 0.83 -4.14 0.05 -2.38 0.06 -1.43 0.009 -0.34 0.29 -1.19 0.0004 South Platte River.--Because of inadequate sample sizes, few meaningful data were available (Table 5 and Table 1). A highly significant decline in cottonwood stands from early to recent intervals was noted in the upper stratum (1) below Greeley but, in contrast, an increase in trees was noted in the 2nd stratum (Table 6). The 3rd stratum showed a decline ",hereas little change was noted in the lowest stratum. Because of these inconsistent findings, differences among strata may be more a factor of sampling than of actual trend in differences among strata and should be interpreted with caution. �309 Table 5. Changes (ha) in cottonwood stands from early to recent intervals within 4 sampling strata and for combined strata along the South Platte River, Colorado. Stratum P x P x C1B 1 2 3 4 Mean -0.60 2.83 -3.36 0.59 -0.44 0.27 0.17 0.09 0.76 0.53 -0.08 -0.27 -2.86 3.40 -0.49 C2A 1 2 3 4 Mean -2.55 -1.02 -9.60 2.94 -3.56 2 3 4 Mean -0.82 -8.70 1.70 -3.10 -2.28 0.77 0.06 0.83 0.70 0.40 4 Mean C 0.97 1.40 -0.78 -0.19 0.37 C2C 0.28 0.29 0.93 0.51 0.94 -0.52 5.31 4.00 5.38 3.128 1.87 2.54 -1.13 -1.82 0.59 0.50 0.43 0.59 0.42 0.62 C3C 0.84 0.11 0.26 0.17 0.32 0.07 0.47 0.68 0.61 0.39 -9.00 2.10 2.20 -1.40 -1.76 C4B 0.03 0.23 0.45 0.33 0.33 0.37 0.28 0.17 0.33 0.14 -0.97 1.56 -2.89 2.21 -1.11 C3B C4A 1 2 3 0.88 0.59 0.03 0.32 0.36 -1.69 4.50 -0.19 -4.50 0.10 C3A 1 C1C C2B 0.30 0.75 0.02 0.55 0.02 P C4C 0.90 -0.58 0.01 0.07 0.20 0.78 0.14 0.37 -0.07 0.05 -0.32 -1.00 0.04 0.76 0.81 0.28 0.31 0.82 aCanopy cover (%) and age class of cottonwoods: A = <35%, B = 35-55%, 55% canopy cover; 1 = <6, 2 = 6-15, 3 = 16-30, and 4 = >30 in. dbh. Mean changes in cottonwood stands relating cc to strata were not evident (Table 7) except that declines were noted in density levels within stratum 1. The most pronounced decline apparently was within open stands (p = 0.02). Analysis of mean changes based on age class within and among strata yielded highly variable results with no pronounced trends for either age class or strata (Table 8). �310 Table 6. Changes (ha) in cottonwood stands from early to recent intervals among strata along the South Platte River. x Stratum 1 (upstream) 2 3 4 (downstream) P -12.60 9.74 -13.10 0.20 -5.48 x 0.009 0.019 0.09 0.99 0.046 Table 7. Changes (ha) in cottonwood stands from early to recent intervals among strata and canopy cover, South Platte River. Stratum Canopy cover (%) 1 2 3 4 -3.01 0.20 -5.46 0.15 -12.0 0.11 0.30 0.97 -5.91 0.02 P -1.40 0.69 8.94 0.03 1.00 0.83 4.30 0.54 2.67 0.16 }f P -8.18 0.01 6.25 0.14 -2.10 0.81 -4.40 0.45 -2.25 0.42 x <35 P 35-55 x >55 x Table 8. Changes (ha) in cottonwood stands from early to recent intervals by age class among strata along the South Platte River. Age class (in. dbh) Stratum 1 2 3 4 x <6 K P -1.66 0.16 4.12 0.16 -9.10 0.01 1.77 0.69 -2.04 0.087 6-15 x -2.40 0.52 6.00 0.50 -10.90 0.06 -3.30 0.76 -3.08 0.29 -10.40 0.08 -1.30 0.86 8.00 0.26 0.90 0.90 -0.91 0.76 1.80 0.06 0.88 0.49 -1.09 0.31 0.80 0.38 0.55 0.25 P 16-30 K P >30 x P �311 Colorado River.--Data available (Tables 9-12) provide evidence that cottonwood stands through all age class and cc densities declined within the upper (1st) stratum and that the overall decline was significant (p = 0.021). A lack of young trees «6 in. dbh) was evident among all strata (Tables 9-12). A similar decline in large trees (>30 in. dbh) was noted although sample sizes were inadequate. Most of the decline occurred in the intermediate to dense cc categories and was especially evident when strata were combined (Table 11). Rio Grande River.--Mean changes in hectares of cottonwoods from early to recent intervals were generally positive along the Rio Grande (Table 13-14). The upper stratum appeared to gain the most and a slight (nonsignificant) decrease was noted in the lower stratum (Table 14). Based on the available data (Table 15), trends were away from open stands toward the intermediate (35-55%) cc density which showed a pronounced increased (p = 0.015). Limited evidence that trees within the 6-15 in. dbh age class declined in the lower 2 strata are available (Table 16). Trees in the 16-30 in. dbh age class increased within all strata. Natural Establishment and Survival of Cottonwood Seedlings Intensive Transects.--F100ding along the South Platte River in spring and summer 1983 and 1984 stimulated considerable natural reproduction of cottonwoods in many sites. Intensive transects established randomly were monitored in 1984 and 1985 (Snyder 1986). Sampling continued in early fall 1986 (Table 17). One transect, within the Brush Wildlife Area was added to the 24 previously established sites. A controlled burn had been conducted in late winter 1986 and impacted the seedlings within the transect. However, rapid resprouting from basal areas occurred and the resprouts appeared moderately healthy going into fall 1986. Seedling mortality among all transects averaged 24.2% from fall 1985 through fall 1986 compared to a 35% mortality rate during the previous year. Livestock crossed the river in summer 1986 and noticeably browsed all seedlings within transect 4 on the Sedgwick Bar Property. However, attrition on most other transects was attributed to natural stand thinning or to desiccation. Growth rates have varied widely among. and within sites but most seedlings that germinated in 1983 have attained heights of 1 m or more. All seedlings within 6 of the 25 transects had died by fall 1986 (Table 17). Extensive Transects.--Thirty transects established along the South Platte River in late summer 1984 were inventoried in September 1986 (Table 18). Cottonwood seedlings established by natural reproduction in 1983 and 1984 occurred in 0.86% of 1,984 sample frames in early fall 1986. Frequency of occurrence was 1.61% in 1984 and 0.96% in 1985 revealing the attrition also documented within the intensive transects. In 1984, 17 of 30 transects contained 1983-84 seedlings compared with 10 of 30 transects in 1986. Because of the few occurrences of 1983-84 seedlings encountererd, a continuous meter-wide belt along all transects was also measured in fall 1986. This sampling approach increased the number of occurrences only slightly to 22 tallies within 12 of the 30 transects. �312 Table 9. Changes (ha) in cottonwood stands from early to recent intervals within 4 sampling strata and combined strata along the Colorado River. Stratum x P x C1Aa 1 2 3 4 x -0.08 -1.06 -0.36 -0.77 -0.62 0.39 0.56 0.66 0.22 0.16 x -0.15 5.93 0.86 -0.86 1.20 -1.93 -0.58 -0.60 -0.79 0.24 0.03 0.36 0.25 0.03 x -1.48 -2.00 2.15 1.37 0.25 x 0.02 0.009 -1.00 0.62 -1.95 -0.81 0.37 0.69 0.37 0.16 0.82 0.09 0.54 0.31 0.85 0.59 -0.12 -3.24 -0.46 0.07 -0.84 -2.05 0.55 0.07 -0.32 0.91 0.90 0.20 0.35 0.85 0.34 C3C 0.22 0.05 0.08 0.88 0.02 -0.34 0.45 -0.66 0.91 0.23 C4B -0.13 -1.84 -0.32 0.11 -0.47 0.49 0.25 0.10 0.09 C2C C3B C4A 1 2 3 4 0.43 0.27 0.09 0.15 -1.05 1.54 -2.05 -0.14 -0.34 P C1C C2B C3A 1 2 3 4 x C1B C2A 1 2 3 4 P 0.60 0.64 0.71 0.28 0.61 C4C 0.19 0.13 0.25 0.26 0.07 -1.83 -0.43 -0.06 -0.53 0.20 0.30 0.27 0.09 aCanopy cover (%) and age class of cottonwoods: A = <35%, B = 35-55%, C = > 55% canopy cover; 1 = <6, 2 = 6-15, 3 = 16-30, and 4 = >30 in. dbh. �313 Table 10. Changes (ha) in cottonwood stands from early to recent intervals among strata along the Colorado River. Stratum x P 1 2 3 4 -3.37 -7.00 -0.67 -1.87 -3.04 0.021 0.29 0.77 0.57 0.08 x Table 11. Changes (ha) in cottonwoods among strata from early to recent intervals by canopy cover along the lower Colorado River. Stratum Canopy cover (%) 1 2 3 4 p -1.71 0.34 2.90 0.66 2.65 0.18 -0.24 0.90 0.83 0.58 K -1.31 0.09 -5.48 0.08 -3.41 0.16 -0.56 0.56 -2.44 0.004 -0.34 0.60 -4.44 0.07 0.08 0.97 -1.02 0.43 -1.43 0.05 x <35 35-55 p x >55 p x Table 12. Changes eha) in cottonwoods among strata from early to recent intervals by age class along the Colorado River. Age class (in. dbh) <6 X p 6-15 X p 16-30 >30 Stratum 1 2 3 4 x -0.08 0.39 -4.00 0.14 -0.32 0.81 -3.32 0.03 -2.22 0.004 -0.63 0.81 -0.92 0.55 0.54 0.52 -1.21 0.048 5.43 0.075 p -1.94 0.17 -4.78 0.29 1.03 0.15 2.35 0.16 -0.36 0.71 x p -0.14 0.19 -3.67 0.06 -0.75 0.26 0.08 0.74 -0.997 0.019 X �314 Table 13. Changes (ha) in cottonwood stands from early to recent intervals within 3 sampling strata and combined strata along the Rio Grande River. Stratum x P x ClAa 1 2 3 x 0.47 1.41 -0.15 0.60 0.30 0.35 0.75 0.25 0.83 0 -0.16 0.14 x 0.08 -0.22 0.22 0.02 x -3.19 -2.07 -0.40 -1.73 0.056 0.16 0.08 0.005 x C -0.04 -0.29 0.18 -0.05 C2C 0.71 0.31 0.17 0.045 2.09 6.08 2.38 3.73 0.41 -0.30 -1.86 -0.703 0.77 0.74 0.25 0.27 C3C 0.52 0.01 0.20 0.002 4.19 1.66 0.67 1.92 C4B 0.81 0.20 0.36 0.58 0.15 0.11 0.18 0.09 0.74 -1.33 -0.47 -0.50 C3B C4A 1 2 3 0.29 1.00 0.36 0.57 -0.46 -1.14 -1.61 -1.15 C3A 1 2 3 C1C C2B 0.92 0.89 0.34 0.98 P X C1B C2A 1 2 3 P 0.23 0.60 0.36 0.13 C4C -0.31 0.34 0.19 0.46 0.05 0.72 1.13 -0.52 0.18 0.14 aCanopy cover (%) and age class of cottonwoods: A = <35%, B = 35-55%, >55% canopy cover; 1 = < 6, 2= 6-15, 3 = 16-30, and 4 = >30 in. dbh. 0.12 0.09 0.36 0.39 �315 Table 14. Changes (ha) in cottonwood stands from early to recent intervals among strata along the Rio Grande River. Stratum 1 2 3 x x p 5.92 3.60 -1.01 2.47 0.10 0.36 0.65 0.12 Table 15. Change (ha) in cottonwoods from early to recent intervals by canopy cover among strata along the Rio Grande River. Canopy cover 1 Stratum 2 3 x p -2.68 0.11 -1.17 0.68 -0.15 0.77 -1.17 0.25 35-55 ~ p 2.15 0.55 >55 x 6.46 0.06 (%) <35 x p 5.27 0.028 -0.49 0.87 0.62 0.52 -1.47 0.45 2.78 0.015 0.86 0.49 Table 16. Changes (ha) in cottonwoods from early to recent intervals among strata and by age class along the Rio Grande River. Age class (in. dbh) <6 1 Stratum 2 3 x 2.03 0.07 0.08 0.96 -0.77 0.10 0.25 0.69 0.02 0.99 -1.67 0.52 -3.25 0.19 -1.84 0.13 x 3.09 0.50 5.67 0.09 2.65 0.27 3.92 0.16 x 0.78 0.18 -0.47 0.45 0.36 0.36 0.15 0.56 x p 6-15 x p 16-30 p >30 p �Table 17. w Cottonwood seedling density along South Platte River transects from fall 1984 through 1986. Site Transect 1/ Grazing status Samp1es/ transect Total seedlings 1985 1986 1984 Grazed Ungrazed Ungrazed Ungrazed Grazed Grazed Grazed Grazed Grazed Ungrazed Ungrazed Ungrazed Ungrazed Ungrazed Ungrazed Ungrazed 25 9 25 25 25 20 7 17 25 25 16 25 25 25 25 25 69 82 28 3 54 27 35 172 199 13 92 51 268 67 48 29 24 4 1 26 3 24 129 119 8 81 44 0 54 27 226 54 33 184 Grazed Grazed Ungrazed Grazed Ungrazed Ungrazed Grazed Grazed Ungrazed 25 25 18 25 25 17 20 25 25 360 84 158 94 195 48 216 334 106 2 1 51 1 95 9 56 37 62 0 0 39 0 95 7 12 11 43 1984 I-' ~ Mean density/m 1985 1986 Seedlings established in 1983 Sedgwick Bar Haxtun R&G Tamarack Bravo Tamarack Brush 4 2 4 2W 4W1 4W2 4W3 6W1 6W2 21W 22W 24W 1 2 9Wa 1 26 14 1 0 13 0 12 99 85 4 75 37 2.76 9.11 1.12 0.12 2.16 1.35 5.00 10.12 7.96 0.52 5.75 2.04 10.72 2.68 21.33 1.16 2.67 0.16 0.04 1.04 0.15 3.43 7.59 4.76 0.32 5.06 1. 76 1.04 1.55 0.04 0 0.52 0 1.71 5.82 3.40 0.16 4.69 1.48 2.16 12.00 9.04 2.16 14.67 7.36 0.08 0.04 2.83 0.44 42.22 5.89 31.11 16.44 27.56 0 0 2.17 0 42.22 4.56 6.67 4.89 19.11 Seedlings established in 1984 Tamarack Bravo Sedgwick Bar Haxtun R&G Tamarack Jones 13W1 13W2 3 Sa 3a 2Ea 6W3a 6W4a 3a 14.40 3.36 8.78 41. 78 86.67 31.33 96.00 148.44 47.11 aSamp1es were obtained within a 0.09-m2 frame and then corrected to a m2 density for comparison. �Table 18. Frequency of occurrence (m2) of cottonwood and green ash seedlings and other cover typesa obtained within random transects along the South Platte River, northeastern Colorado, September 1986. Site Julesburg Sedgwick Bar Haxtun R&G Tamarack Bravo Transect E-1 E-2 W-1 W-2 W-3 Jones 1 1 1 2 3 4 1 2 3 4 E-6 E-5 E-3 E-1 W-1 W-2 W-ll W-17 W-18 W-19 1 2 3 Dune Ridge Cottonwood 83-84 85-86 1 2 4 1 2 1 1 1 1 1 1 2 1 1 2 2 2 3 21 1.06 5 7 4 4 3 1 1 7 6 5 Grass Annual Peren. 6 7 5 5 12 26 13 19 23 33 2 3 4 3 2 5 6 7 3 10 26 11 21 1 6 2 3 8 9 20 9 3 2 6 6 20 36 39 17 6 23 11 19 11 7 3 1 2 5 4 1 1 9 1 2 125 6.30 100 5.04 12 0.60 Peren. 3 2 6 7 11 3 5 17 0.86 2 3 2 3 5 8 11 2 1 1 Forbs Biennial 9 15 1 5 5 3 1 2 6 Annual 2 3 9 3 6 6 3 3 1 3 1 1 2 Totals Percent Ash 1 Common reed Low 7 6 8 7 4 12 8 Shrubs Tall Vines 1 2 5 4 13 11 1 10 5 2 5 4 4 3 5 5 1 5 1 5 3 2 Bare grnd. 4 1 3 2 5 15 18 8 7 1 38 56 40 39 12 6 9 15 3 34 8 7 1 17 6 4 2 6 2 35 19 13 6 3 5 7 7 1 12 1 1 6 13 1 13 1 1 10 10 1 33 36 14 7 5 8 1 7 265 13.36 1 7 2 85 4.28 701 35.33 2 17 0.86 40 23 4 6 14 5 6 4 25 11 10 8 6 20 23 8 4 16 4 6 221 11.14 216 10.89 5 2 5 5 1 3 Sandbar 2 2 1 3 6 Down timbo Litter 2 1 5 2 12 8 1 3 1 18 7 2 5 2 1 1 1 7 1 2 1 4 7 1 1 1 2 2 1 6 4 1 2 37 1.86 47 2.37 2 2 1 Total 63 48 41 78 106 38 62 27 45 69 83 89 61 105 130. 82 93 86 65 33 65 48 66 88 67 34 21 52 75 64 90 4.54 17 0.86 13 0.66 1,984 aCover types were classified by dominant cover present where more than one type was present within a 1-m2 sample. W I-' -....J �UJ r-' CIJ 0 - - ~ 2 I I 4 l- ._.-.-._ •••••••••••••••• ,t tI I I I d CHECK STATION MEAOOW T,l,MARACK 11-&: RED LION DUCK CREEK I \ \ I I I , I ,, , ~ --' \ .•... - . 6 ',.... ........ , (\ ,, ,, lQ ~ _.~ \ .. ...... ,, \ I \ \ , I 8 .......•. I ~ l:l _' \ \ I , .' '. '\' ~( 10 ', ,.~. ,,' ." I n \ • I , ~ 12 , I \ ,," .tI.~ • • I ,{ .···f .....\ ,.'~., ,,, ,,.'r:•..• " \ \ - --..J!,. \. \' \ ~ ,.... , I,: \ ," I I '.• I ' I \ ' , \, 14 16 ~ It >0 ::c < ~ § t') sO>fr~~~ t') ~ 1984 III ~ It ~!: 0 Z Q ::c < ::c ~ r-4 0> ~ ~'J: 0. ~ 8 > ~ M ~ Q t') ~ 0> 0. ~ ~ It ~>0 ~!: r-4 ~ ~ J: 8 ::c < 1985 Fig. 1. Groundwater fluctuations (dm) within South Platte River, Logan County, Colorado. ~ M Z c 1986 stem cutting planting sites from 1984 through 1986, �319 Cottonwood seedlings established in 1985-86 occurred in 1.06% of the sample frames. This latter group of seedlings, established during low river conditions, are highly vulnerable to inundation mortality in subsequent years and are not expected to significantly contribute to future tree stands along the river. Green ash were recorded within 12 frames (0.60% occurrence) in 1986 compared to 9 frames (0.45%) in 1985. This species was found within 5 of 30 transects in 1985 and within 7 of the transects in 1986. Green ash seedlings were tallied 25 times within 9 of the 30 transects when sampled on a continuous belt approach in 1986. Almost no old green ash trees exist along the South Platte River and only a modest number of young seed-producing trees are scattered along the river. However, extensive seedling reproduction is expanding from the parent stock, often in moderate to dense stands. Much of the reproduction is attributed to the 1983-84 high water conditions. The species is less subject to browsing by livestock, is more shade tolerant, and apparently can withstand moderate flooding innudation. As a consequence, it may gradually become the dominant tree in many locations along the South Platte. When seedlings were not tallied within sample frames, the dominant vegetation type within each frame was tallied as in previous years. Annual forbs were tallied as dominant in 6.3% of the frames, biennial forbs occured 5.0% of the time, and in combination the 2 represented 11.3% of the occurrences. In contrast, annual and biennial forbs represented 23.9% of the occurrences in 1984 and 14.9% in 1985. Perennial grasses and perennial forbs in combination were dominant in 35.6% of the frames in 1984, 40% in 1985, and 48% in 1986. These data provide evidence of trends toward stable perennial herbaceous stands following flooding inundation and disturbance that promoted annuals during 1983-84 floods. Decreased abundance of annual forbs is indicative of decreased seed abundance to wildlife, although it must be recognized that many annual and biennial forbs (e.g., Lactuca spp., Conyza spp.) do not yield highly nutritious seeds. However, on average the seeds of annuals are much more nutritious than those of perennials and are preferred as food by wildlife. River Volume and Groundwater Measurements Increased frequency of groundwater sampling at stem cutting planting sites along the South Platte River during the spring and summer of 1986 (Fig. 1) revealed frequent river and adjacent groundwater fluctuations. Similar patterns occurred in 1985 and 1986 with major river volume occurring in spring during the interval of mountain snowmelt followed by dramatic declines from July into September. The Red Lion, ll-E, and Check Station Meadow sites (Fig. 1) all are near the river channel and are influenced by stream flows. The Duck Creek site lies within a seepage area from the Jumbo Reservoir intake canal and is influenced by canal flows and vegetation at the site. Stem Cutting Propagation Trials As previously reported (Snyder 1985, 1986), stem cutting trials were disappointing during 1984 and 1985 because of poor survival. Three factors, competing vegetation, insufficient planting depth, and repeated fluctuations in groundwater levels were suspected as primary factors affecting survival. �320 Therefore, intensive approaches were used in an effort to determine which, if any, of these factors were responsible. The labor-intensive plastic sleeve-hand auger approach was effective in obtaining adequate planting depths. The lowest 1986 groundwater levels at the South Platte sites were in the 12-15 dm range (Fig. 1). Average stem cutting depths among all species were 17.1 dm at the Check Station Meadow site, 17.7 dm at site 11-E, 19.4 dm on the Red Lion site, and 19.5 dm at Duck Creek (average planting depths were about 10 dm in 1985). All 1986 planting depths were below the lowest water levels recorded in summer 1986 indicating the bases of most cuttings remained continuously in saturated soil. In addition, the plastic-organic mulch treatment eliminated competing vegetation and undoubtedly retained moist soil at higher levels than at adjacent plantings where vegetation usurped considerable soil moisture in mid to late summer. Survival results markedly improved in 1986 contrasted to those in previous years (Table 19). Survival to late summer among 49 unmu1ched plantings averaged 33% compared to 22% survival among the same combined species at the same sites in 1985. This provides limited evidence that increased planting depth in 1986 was at least a partial factor in survival since groundwater levels and fluctuations tended to be similar between the 2 years. Survival averaged higher within the mulch treatment on the Red Lion and 11-E sites which contained a majority of the cuttings and was nearly equal at the other 2 sites. The combined average among sites and species was 51% survival for mulched plantings but analysis provided no evidence that survival was significantly greater than that within untreated plantings (t = 0.79, df 31, P>0.05). The overall average of 45% survival was below acceptable limits when conSidering this as a viable planting technique. Part of the mortality may have resulted from plant injury especially to crooked stems that did not readily pass through the plastic sleeve. Injury was suspected among a few plantings that failed to leaf out but the overall impact of injury related mortality is uncertain. The lowest survival rate occurred on the Red Lion site (Table 19) where observations indicated deer or rabbits were browsing some of the new leaves. New plantings apparently cannot withstand defoliation because of lack of energy reserves. Available evidence, however, leads to the SUsplClon that fluctuating groundwater levels continue to be the most important variable contributing to low survival of stem cuttings. There were rapid and repeated fluctuations in groundwater levels with declines in March, increases in April, declines in June, and major declines in July 1986 (Fig. 1). New roots forming in moist soil above the water table might be repeatedly inundated and desiccated and therefore severely stressed. Swenson and Mullins (1985) noted reduced survival among cuttings placed in fluctuating water tables in their New Mexico trials. Holes for placement of stem cuttings cannot be easily dug into the groundwater table because the sands repeatedly collapse as soon as the water table is reached in most sites along the South Platte and South Republican rivers. Thus, a retaining sleeve is needed and the procedure becomes too labor intensive. If high survival cannot be achieved, then the approach becomes completely impractical and cannot be recommended. It would be more practical to plant rooted seedlings and apply a mulch protection. �Table 19. Survival of stem cutting plantings from early spring to fall 1986 within 4 South Platte River and 1 South Republican River planting sites. Site Treatment Populus sargentii Salix Salix ~. .l.Y.t.ea Amorpha b:.ui.t.i. Vitis yulpina _ 3 o 1/4 0/4 Salix .in.t..eJ:.. Salix "gold". Elaeag. anauat.. Combined planted survival South Platte River Check Station Meadow l1-East Red Lion Duck Creek Mulched Unmulch 5a 2 3 Mulched Unmulch 1 3 1 3 5 1 3 1 Mulched Unmulch 1 o 1 o Mulched Unmulch 2/2 2/2 3 3 5 3 3 1 0/1 0/3 0/2 1/2 1/1 1/2 24 10 12 5 29 29 17 6 30 10 11 1 6 3 9 4 10 7 South Republican River West Hale Ponds Mulched Combined Mulched Unmulch Total Percent survival 2 5 9/17 5/17 6/16 1/8 10/18 7/12 13/20 2/10 4/13 5/5 3/5 0/2 1/2 99 49 50 16 14/34 7/24 17/33 15/30 4/13 5/5 3/5 1/4 148 66 41 29 52 50 31 100 60 25 45 aFive stem cuttings per trial were used unless otherwise listed. W N I-' �322 Findings for the lower South Platte River may not be the same as for the lower Arkansas River where supplemental establishment of trees is critically needed. Based on a few samples, the water table averages about 20 dm below existing old cottonwoods downstream from John Martin Reservoir and in many places cottonwoods are no longer present. However, the stream may not fluctuate to a major extent most years. Fine sands are suspected to again be a problem there but testing would be needed. Special equipment would be needed for deep hole boring and almost all of the riparian habitat is privately owned. Dense stands of tamarisk (Tamarix gallica) might pose a problem in severely competing for available groundwater in many locations. As noted earlier (Snyder 1986), the species identified as coyote willow (S. exigua) did not fit published descriptions. Therefore, Dieter H. Wilken of the Colorado State University Botany Department was consulted for correct identification. The species, which has excellent growth form for wildlife, was identified as S. lutea or diamond willow. Soil Scarification-Natural Reproduction Efforts were made in spring 1986 to maintain the 18 soil scarification plots established in fall 1984. However, nearly all remained well above groundwater levels in 1986 with dry soil surface. Canada thistle (Cirsium arvense) invasions became an increasing problem to the extent that continuence of these plots as weed-free bare soil environments was impractical. Repetitious tillage and/or herbicide treatments are needed to control noxious weeds and would not provide the proper soil environment for seedling establishment. The 3 low sites where limited seedling establishment was noted in 1985 were destroyed in fall 1986 with construction of new bridges across the river. Therefore, the soil scarification effort has been discontinued. Monitoring of Controlled Burns in Riverbottom Division management personnel established fireguards and burned 2 small (1-2 ha) tracts within the Brush Wildlife Area in late winter 1986 along the South Platte River in Morgan County. The east burn contained only 10 young to overmature cottonwood trees, 6 peachleaf willows, and a few other woody species whereas the west burn contained only 1 each of peachleaf willow, Russian-olive (E1aeagnus angustifolia), American elm (Ulmus americana), and boxelder (Acer negundo). One old cottonwood and one peachleaf willow clump were excluded with fireguards from the fire. Fires were started at the bases of most large trees and kept at low intensity by using pumper spray so that little damage to the trees occurred. Several peachleaf willows were burned severely but new basal growth (to 2 m) was attained on most in 1986. Regrowth of Virginia creeper (Parthenocissus quinquefo1ia), indigo bush (Amorpha fruiticosa), Russian olive, American elm, and sandbar willow (Salix interior) was noted. Survival of cottonwood seedlings within the random transect established in fall 1985 was good (Table 17). The fire was not intensive at the transect site and most seedlings resprouted and made rapid regrowth. One of the primary objectives of the riparian burns within the Brush Wildlife Area was to renovate herbaceous vegetation that no longer remained standing into or through winter and which contained excessive residual. Opportunity to obtain pre-treatment height-density (HDI) samples within the 1985 burn was not available. However, late February 1986 height-density sampling along randomly �323 located transects was conducted within proposed 1986 burns and controls (Table 20). Post-treatment sampling of the 1985 burn was also conducted but showed no evidence that a major increase in HDI occurred based on comparisons with nearby pre-treatment 1986 samples (Table 20). Subsequent sampling should provide more precise comparisons of burning impacts on riparian herbaceous vegetation quality. Table 20. Height-density (dm) of grass-forb and low shrub vegetation in the 1985 (post-burn) and 1986 (pre-burn) sites and controls within the Brush Wildlife Area, South Platte River, late winter 1986. Shrub Grass-forb Transect status East East West West East East West West West East 1-86 2-86 1-86 2-86 1-86 2-86 1-86 2-86 1-85 1-85 Pre-burn Pre-burn Pre-burn Pre-burn Control Control Control Control Post-burn Post-burn N x N 44 56 67 56 80 58 57 74 64 64 0.56 0.72 0.65 0.33 0.47 0.31 0.33 0.30 0.63 0.56 8 8 17 8 2.62 3.56 2.56 4.19 2 3 18 3.50 0.58 3.11 4 0.81 x Disturbance Tillage Data concerning dominant vegetation occurrence within fireguards and other disturbance tillage transects varied (Tables 21-22). Within the Brush Wildlife Area, sampling was conducted within each of the fireguards established to contain the 2 1986 burns. In addition 2 transects were established along the perimeter of the 1985 burn where a 2nd growing season had elapsed. The available data (Table 21) also include samples from a 1986 hay meadow burn site within the Elliott Wildlife Area and 2 disturbance tillage strips initially plowed in fall 1985 within Area I-Won the Tamarack Wildlife Area. Findings reveal similarity within 3 of the 4 Brush Wildlife Area transects but high variance among sites in species occurrence and quality. Sunflower (Helianthus sp.), cocklebur (Xanthium strumarium), giant ragweed (Ambrosia trifida), Chenopodium spp., sweetclover (Meli1otus spp.), and western ragweed (A. psilostachya) were important dominants in most samples and most were also d~minant species within the Elliott sample (Table 21). However, major differences were noted within the 2 Tamarack sites. The exterior disturbance tillage strip had been plowed within dense stands of poison ivy (Rhus radicans) and snowberry (Symphoricarpos albus), no annuals were tallied as dominants within samples (Table 21). The interior strip, within a more open site, contained a more diverse composition with numerous annuals. �Table 21Occurrences of dominant vegetation within samples along fire guards and disturbance tillage strips along the South Platte River, Fall 1986. Species ANNUALS Helianthus sp. Ambrosia trifida Thlaspi arvense Euphorbia spp. Amaranthus spp. Lactuca spp. Chenopodium spp. Salsola kali Kochia scoparia Iva xanthifolia Xanthium strumarium Cenchrus sp. Echinochloa sp. Subtotal BIENNIALS Melilotus spp. Conium maculatum Subtotal PERENNIALS Ambrosia psilostachya Lepidium sp. Solidago sp. Asclepias syriaca Circium arvense Rumex crispus Glycyrrhiza lepidota Rhus radicans SymEhoricarpos albus Carex sp. Spartina pectinata Distichlis stricta Subtotal Rating a FC FC F F F FC C C C 1-86 33 3 Brush Wildlife Area 2-86 1-85° 22 3 F C C Elliott 3 5 1 28 3 12 2 1 1 3 3 7 1 3 6 1 1 6 1 55 32 14 25 14 3 4 1 38 16 1 1 21 26 15 13 20 F C 2-85° Tamarack Ext Int 14 25 15 13 2 2 3 1 9 1 11 1 20 17 4 4 4 4 8 9 3 5 9 14 2 12 30 2 3 1 1 C C 1 4 1 2 C 10 6 1 1 12 15 33 aF=food, C=cover, If not rated are of little value to most wildlife. bHad completed their second growing season after tillage. 1 w N -'" �325 Dramatic height differences were also noted within and among transects based on vegetation and soil variations. Three of the 4 Brush Wildlife Area transects contained excellent tall stands whereas the 4th, in a sandy site yielded much shorter vegetation (Table 22). Within the Elliott fireguard, dominant vegetation ranged from 0.6 to 2 m in height and averaged slightly less than 1 m. Vegetation within both Tamarack strips tended to be short and open providing marginal feeding cover for wintering wildlife. Table 22. Average height (m) of dominant vegetation within fireguard samples in the Brush Wildlife Area, South Platte River, fall 1986. Species ANNUALS Helianthus sp. Ambrosia trifida Lactuca sp. Chenopodium spp. Salsola kali Kochia scoparia Iva xanthifolia Xanthium strumarium Cenchrus sp. Echinochloa sp. Euphorbia spp. BIENNIALS Melilotus spp. PERENNIALS Ambrosia psilostachya Lepidium sp. Asclepias syriaca Circium arvense Rumex crispus Glycyrrhiza lepidota Symphoricarpos albus Spartina pectinata Distichlis stricta Carex sp. Transect x 1-86 Transect 1-85 2-86 1.85 2.23 1.54 1.83 1.74 0.91 1.73 2.86 1.52 2-85 x 0.81 1.83 1.63 1.22 1.62 1.52 1.66 1.76 1.01 0.30 0.31 0.30 0.30 0.54 0.83 0.91 1.58 0.61 0.99 0.91 0.91 0.91 0.91 0.56 0.91 0.36 1.22 0.40 1.22 0.45 1.04 1.69 0.91 0.61 0.91 0.80 0.30 0.91 0.30 1.60 1.18 1.36 0.25 0.25 0.44 Based on observations, the plowed fireguards were one of the major benefits of the controlled burns to riparian wildlife. Ring-necked pheasants and coveys of northern bobwhite were observed using the fireguards within the Brush Wildlife Area where tall stands of annual forbs created protective feeding, loafing, and movement cover as well as interspersion. Sunflowers and ragweeds were considered especially important as food and cover. Other species providing either food or cover to wildlife are also listed (Table 21). Preliminary findings indicate pre-treatment vegetation and soil type are �326 important considerations when establishing productive disturbance tillage strips. Annual, or at most, biennial tillage is needed to retain vigorous open stands of annuals. Characteristics of the Arkansas River Channel Several major irrigation canals divert water from the Arkansas River near Pueblo and continuing downstream toward John Martin Reservoir. As a consequence, the river's volume is usually progressively diminished from west to east. Water containment and diversion also occur at John Martin Reservoir and a few downstream canals divert water before the river passes into Kansas. Monthly and annual flow volumes at points along the river vary (Fig. 2, Table 23). They illustrate progressively diminished flows but also show fluctuations among seasons. It must be remembered that tributaries including the Pugatorie River, Butte Creek, and others contribute to the Arkansas as it passes through southeastern Colorado. As a consequence of these and other factors including controlled releases from Pueblo Reservoir, the character of the Arkansas River changes drastically as it progresses across southeastern Colorado. Channel width is one readily noticeable affect. Mean channel width (Table 24) at 12 sites upstream from John Martin Reservoir averaged 87 m contrasted to only 35 m at 13 downstream sites (! = 10.21, 23 df, R < 0.05). The width sampled was between the banks of the stream at normal river levels. The original, much wider channel is still discernable in some downstream (below reservoir) locations but tamarisk (Tamarix gallica) has invaded the previously occupied channel in most locations. Comparison of early (1930's and 1940's) aerial photos with those taken in recent years also reveal dramatic reductions in downstream channel width. Mean daily stream flows (CFS) averaged per year at 5 gaging Table 23. stations along the lower Arkansas River in southeastern Colorado, 1965-l984.a Year Avondale Las Animas Location J. Martin Res. Lamar Stateline 1965 1967 1969 1971 1973 1974 1975 1977 1979 1980 1981 1982 1983 1984 1,094 622 920 802 970 592 818 363 910 1,308 559 1,363 1,344 1,645 493 48 105 112 120 82 127 68 148 404 75 256 392 587 435 389 187 189 220 119 127 120 187 358 229 305 405 388 361 193 17 49 86 39 28 46 46 189 86 97 169 177 1,067 285 199 131 151 60 51 34 20 171 66 96 226 272 951 215 261 113 202 x aData from U.S. Geological Survey Water-Data Reports for Colorado. �327 2,500 2,000 (near Pueblo) o 1,500 z o ~ ~ t f:! 1,000 900 800 ..•..... ,. 700 "'.<, 600 500 "~Below ,• 400 300 line ~ • .~'.' Lamar / p.r- • ..... f_.~',.. ..~ ,L" 100 •••• I , / ••....... , • \ ....::......• .~--" ..... ...., I 200 50 J• ..•. /, " "", ., ~ -, ., ··0 • .., --.. -~ ..... Jan \ .•...• ..".................. ..._..... • • ".., " v- • •••• , ....., ... •.....• Feb. Mar Apr May JUri Jul Aug Sep Oct Nov Dec MONTH Fig. 2. Mean daily stream flows (CFS) averaged per month at 5 gauging stations along the lower Arkansas River, southeastern Colorado during 14 years (from 1965 to 1985). �328 Table 24. Width (m) of the Arkansas River Channel and vertical distance (dm) from water level to base of nearby cottonwood trees at random sites upstream and downstream from John Martin Reservoir, southeastern Colorado, 1986. Sample mile 04 08 12 15 30 45 47 52 55 58 81 82 Water to tree (dm) Sample mile Downstream Width (m) 105 65 93 88 94 70 100 75 109 81 70 93 13.0 12.2 12.7 15.2 15.2 11.2 15.5 22.3 16.0 10.8 9.9 9.1 089 095 106 107 119 130 136 143 144 147 149 155 162 32 37 50 50 42 19 21 24 37 50 30 32 27 10.4 87 13.6 35 20.4 Upstream Width (m) Water to tree (dm) a 26.3 17.8 21.3 14.2 18.3 19.8 19.3 aCottonwoods were not available for sampling in several locations below John Martin Reservoir. Another concern was whether the river channel had deepened appreciably below John Martin Reservoir in relation to upstream areas. However the dense tamarisk stands along the river banks made it extremely difficult to obtain stream profile samples comparing upstream and downstream sites based on a standard 100 m width because transit-level sightings could not be conducted through the dense growth. Stream profile samples that varied in length but averaged 98.2 m wide were conducted on 12 upstream samples. In comparison 9 downstream sites averaging 97.2 m wide were sampled. The average of the 8 deepest profile measurements from upstream samples was 17.8 dm whereas those from downstream profiles averaged 23.1 dm (t = 2.93, 19 df, P< 0.05). Thus, the data provide evidence that the downstream channel had deepened. Personnel of the Corp of Engineers at John Martin Reservoir provided historic degradation from 13 sample points lying between John Martin Reservoir and Lamar. The original survey conducted in 1943-44 had been repeated at some sites in 1962, 1966, 1972, and 1981. Graphic presentations of the data show that at several sample points within a few km of the Reservoir, a narrow channel had deepened about 0.61 to 1.2 m beyond the original channel depth. However, further downstream near Lamar deposition had occurred filling much of the original channel leaving a narrow channel at about the same elevation as that present in 1943-44. Progressing on toward the Kansas border, observations indicate the river continues to show less relief between streambed and adjacent riverbottom. A major flood occurred immediately below John Martin Reservoir and downstream in 1965 when Muddy Creek, Clay Creek, Butte �329 Creek, and other streams flooded and undoubtedly deposited considerable sediment in sites containing tamarisk and other dense vegetation. Channel deepening would also impact the ability of cottonwoods and willows to sustain themselves by natural reproduction. Even if occasional flooding occurred and tamarisk did not dominate, young seedlings established following high water would desiccate when the river returned to normal stream flow levels. Thus, greater relief means there is less chance for natural cottonwood reproduction to occur. The vertical distance from river water level to the base of nearby cottonwoods was measured within sampling sites at several locations upstream and below John Martin Reservoir (Table 24). Trees were not present for sampling at several downstream locations so sample size was reduced. The average vertical distance from water to tree base among upstream samples was 13.6 dm contrasted to an average of 20.4 dm among downstream samples. The results were statistically different (t = 2.61, 18 df, P <0.05), however additional samples should be obtained before-placing strong confidence in the data. These and previous data do not provide much encouragement for retention or propagation of cottonwoods and willows over extensive areas of the Arkansas floodplain. LITERATURE CITED Snyder, W. D. 1984. Lowland riparian cottonwood studies. Colorado Div. Wi1dl. Game Res. Job. Prog. Rep. Fed. Aid Proj. N-4-R-2. Jan:295-370. 1985. Dynamics of Cottonwood Regeneration. Colorado Div. Wildl. Avian Res. Job Prog. Rep. Fed. Aid Proj. 01-00-045 (W-37-R). Apr. (In Press). 1986. Dynamics of Cottonwood Regneration. Colorado Div. Wildl. Avian Res. Job Prog. Rep. Fed. Aid Proj. 01-03-045. Apr. (In Press). Swenson, E. A., and C. L. Mullins. 1985. Revegetating riparian trees in Southwestern floodplains. Pages 135-138 in R. R. Johnson et al. (Tech. Coord.) Riparian Ecosystems and their management: reconciling conflicting uses. First North American Riparian Conference. U.S. Dep. Agric. For. Servo Gen. Tech. Rep. RM-120. Prepared by ~~ ~ Warren D. Snyder Wildlife Researcher C ��~ULULauu ULVLbLUU UL jjl WLLOLLLe Wildlife Research Report April 1987 JOB PROGRESS REPORT Colorado State of 01-03-045 (W-37-R) Project 21 Work Plan Job Title: 3 Sandsage-Bluestem Prairie Renovation Period Covered: Author: : Job Avian Research 01 January through 31 December 1986 Warren D. Snyder Personnel: W. Brown, L. Budde, M. Creamer, L. Crooks, T. Davis, K. Dillinger, M. Etl, M. Gardner, T. Kroening, W. Miles, F. Pusateri, J. Saunders, K. Inouye, and W. Snyder, Colorado Division of Wildlife ABSTRACT A warm early spring and above average precipitation in 1986 promoted vegetation growth and seed head production on the Tamarack Prairie, especially by needle-and-thread (Stipa comata). Habitat modifications completed by management personnel included 4 controlled burns (108 ha) and tillageherbicide renovation of previously interseeded tracts (24 ha) during spring 1986. The height density index (HDI) of residual grass-forb and sandsage (Artemisia filifolia) vegetation in late winter 1986 increased from that of the previous year but did not attain qualities equal to pre-treatment levels within 1984 burns. Within 1985 burns, first year post-treatment HDI values for grass-forb and sandsage vegetation were below pre-treatment levels (p < 0.05). Analyses of crown cover revealed that among dominant native grasses, controlled burning tended to suppress needle-and-thread the first year but rapid recovery was noted the second year. Prairie sandreed (Calamovilfa longifolia) was enhanced the first year after fire and blue grama (Bouteloua gracilis) and sand dropseed (Sporobolus cryptandrus) showed uncertain but generally positive responses to fire. Evidence that sand bluestem (Andropogon hallii) was also enhanced by fire continued to increase. Sandsage was suppressed but not killed by spring burns and the same was true for cactus (Opuntia ~.). Little impact of fire on perennial and annual forbs was noted. Tillage-herbicide revegetation manipulations initiated in spring 1985 continued to yield dramatic results in establishment of tall, warm-season grasses with greatly enhanc~d HDI's. Tillage-herbicide renovation of previously interseeded tracts likewise yielded dramatic results in promoting dominance by tall, warm-season grasses. Comparisons among treatments and years for 2 deep-rooted warm season grasses were conducted revealing that the tillage-herbicide renovation approach �332 yielded the most pronounced results. Evaluations showed that 1985 efforts to control sandsage with an aerially applied herbicide were too effective, however, understory forbs made considerably recovery in 1986. �333 SANDSAGE-BLUESTEM PRAIRIE RENOVATION Warren D. Snyder P. N. OBJECTIVES Monitoring of environmental and vegetation conditions and changes continued as follows: 1. Precipitation was monitored throughout the year supplementing electronic rain gauge data with information from nearby weather stations through the winter months. 2. Soil moisture accumulations, plant phenology, and weather were monitored primarily in spring and especially at time of controlled burns. 3. Visual obstruction (height-density) measurements were obtained on treatments and controls where applicable in late winter and/or early spring prior to major new growth. 4. Crown cover, species composition, and frequency of occurrence measurements were obtained from mid-summer to early fall. 5. Photos of treatment and controls were taken in October. Data compilation and writing the annual job progress report was conducted during fall and winter 1986-87. METHODS Approaches used within this study were previously summarized (Snyder 1985, 1986) and are outlined in the Segment Objectives. In an effort to evaluate responses of prairie sandreed and sand bluestem, 2 deep-rooted, warm-season, tall grasses, the following procedures were used. Several transects within 1984, 1985, and 1986 burn sites, their controls, and within 1985 revegetation strips and 1986 renovation (interseeded) strips that contained nearly pure proximal stands of the 2 grass species were randomly selected. At each transect, percent seeded status was evaluated by ocular estimate. In addition, 12 height measurements and 12 height-density measurements were obtained within stands of each of the 2 grass species. RESULTS AND DISCUSSION Environmental Conditions and Controlled Burning - 1986 Precipitation.--Two automatic recording precipitation gauges were installed in mid-February 1986 and were supplemented by 2 additional automatic recorders in mid-May. All were removed in mid-November. Except for several instances where the recorders became plugged, all provided usable data. November through February data were obtained from nearby U.S. Weather Service stations. Spring through late summer readings revealed that considerable �334 variation occurred among locations within the Tamarack Prairie (Table 1). An exceptionally heavy rain (1 dm, 3.95 in.) was recorded in early April within the east portion of the Prairie and provided excellent subsoil moisture. Greater precipitation than in surrounding areas was received in the central portion of the Prairie in August and in the west portion in September (Table 1). Table 1. Monthly precipitation (in.) recorded at 4 sites within the Tamarack Prairie in 1986 and supplemented in winter by data from nearby U.S. Weather Stations. Location Month East South Middle West Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec 0.08a 0.93a 0.48 4.67 3.15 2.73 1.21 0.84 1.63 1.49 0.31 0.44 0.08 0.93 0.08 0.93 0.08 0.93 0.60 3.21 2.30 2.96 0.52 1.32 2.64 1.45 0.48 0.44 2.16 3.51 1.50 1.34 2.04 2.81 0.46 0.44 2.78 1.34 2.60 2.37 0.45 0.44 Annual x 0.08 0.93 0.54 3.94 2.54 3.00 1.14 1.53 2.10 2.03 0.43 0.44 18.70 aDecember through February data from nearby U.S. Weather Stations. Monthly readings among stations were averaged for comparison with data from previous years (Table 2). April through June amounts were greater than those for 1984 and 1985 and also above the long-term average. Precipitation in 1986 was also supplemented by November-December 1985 snows (Table 2) that prevented soil drying over winter and partially accumulated in soils during January 1986 thaws. Soil probe sampling in spring 1986 before top soils became too dry to penetrate revealed that subsoils were consistently moist to over 1 m in depth and frequently approached a 2-m depth (Table 3). The soil probe was only 1.83 m (6 ft.) long and could frequently be inserted the entire distance especially in the eastern part of the prairie and within revegetation strips (Table 3). Since sandy soils retain about 2.5-3.8 cm of moisture/0.3 m (1-1.5 inc./ft.) (D. Smika, pers. commun.), the Tamarack Prairie contained 10-20 cm of moisture (4-8 in.) through mid-spring before major vegetation growth occurred. Temperature and Phenology.--Average monthly temperatures were above normal (at Sterling) during most of the first half of 1986 and were exceptionally warm in March (Table 2). Only February and May temperatures averaged below long-term means. As a consequence, vegetation phenology was especially rapid in March �Table 2. Precipitation (in.) and monthly average temperature (F) departures recorded from 1984-86 in relation to long-term averages, Tamarack Prairie and nearby U.S. Weather Stations. Departure from x tem2erature Precipitation Long-terma x Month Jan Feb Mar Apr May Jun Ju1 Aug Sep Oct Nov Dec Annual 1984 1985 1986 0.32 0.35 0.78 2.02 2.55 2.52 2.09 2.04 1.37 0.99 0.44 0.49 0.47 1.12 0.66 3.42 2.01 2.80 0.49 0.66 0.44 1.39 0.01 0.80 0.47 0.13 0.38 2.24 2.49 0.82 3.78 0.83 1.42 0.37 1.06 1.20 0.08 0.93 0.54 3.94 2.54 3.00 1.14 1.53 2.10 2.03 0.43 0.44 15.96 14.27 15.19 18.70 1984 1985 1986 -4.8 0.7 0.4 -5.9 0.6 -1.8 0.3 3.5 -0.7 -3.0 2.9 1.2 0.1 -7.1 7.1 4.4 3.6 -1. 7 0.5 0.5 -1.8 -2.1 -7.1 -7.8 3.8 -0.5 9.5 2.7 -1.9 3.0 0.4 0.5 0.9 0.4 1.0 -1.1 aAverage of long-term data from Sterling and Sedgwick weather stations. w w VI �w w Table 3. Average soil moisture accumulations based on soil probe samples (m) within selected vegetation and dates during spring 1986, Tamarack Prairie. May 19 ~r Site Vegetation Burn 1-84 Burn 2-84 Burn 1-85 Burn 2-85 Burn 3-84 East Tam. Cent. Tam. East Tam. Native Native Native Native Native Native Reveg. Reveg. 1 1.18 1.40 1.14 1.17 7 (6)a (5) (5) (5) 1.36 (5) 1.46 (5) 1.75 (3) 1.63 1.79 1.83 1.79 (5) (3) (3) (3) 1.67 (5) 1.63 (5) 30 1.74 (3) 1.25 (3) 1.83 (3)b 1.83 (4) 1.65 (3) (j'. Jun 18 0.82 (3) 1.80 (2) 1.80 (2) a( ) sample size. b1.83 m (6 ft.) was the total prob1e length so total moisture base was not reached in all samples. �Table 4. Phenological SEecies Allium textile Artemisia filifolia A. ludoviciana Astragalus sp. Chenopodium album Cymopteris montanus Eri~eron bellidiastrum LathyruB EolymorEhus Leucocrinum montanum Mentzelia nuda Penstemon angustifolius Phlox andicola Psora lea lanceolata Sphaeralcea coccinea Tradescantia occidentalis Tragopogan sp. Agropyron smithii Bouteloua gracilis Calamovilfa longifolia Panicum virga tum PasEalum stramineum SEorobuluB crYEtandrus Stipa~ Cyper~ sp. conditions 3 of selected vegetation during spring 1986, Tamarack Prairie. E. bud May A r Mar 11 25 1 5-7a Bud 2.5 9-11 23-24 30 2.5 E. flower 2.5-5 2.5 2.5 Emerging Leafing 2.5 E. bloom M. leafing 5 5 E. bloom E. flower 7.6 13 5-7.6 5 E. bloom Bloom 10 Emerging 2 2 10-15 Bloom F. bloom 10-13 F. bloom E. bloom .7.6 5-7.6 10 Bloom Bloom 2.5-5 5 30 F. bloom E. leafing Bloom Emerging 5 7 5 30 Bloom 15-20 2.5-5 7-15 5-7.6 Emerging 5-7.6 15-25 25-30 15-20 5-7.6 15 15 F. bloom 10 36 Bloom F. bloom Bloom 10 30 8 25 Heading Headed -aHeight in cm. I..;.) I..;.) -..J �338 and April but slowed in May (Table 4). A hard freeze in mid-April may have also slowed growth. Emergence of warm-season grasses in early to mid-April was especially notable and most had made several cm of growth at the time of controlled burns on 23 and 30 April. Most were slightly ahead of the early phenology in 1985 (Snyder 1986). No green-up of any warm season species was noted at the time of the 4 May 1984 burns (Snyder 1985). Controlled Burns.--The early phenology in 1986 prompted efforts to conduct controlled burns ahead of those in previous years. Burn sites 2-86 and 3-86 were completed on 23 April, however, a wind change forced termination of burning on site 1-86 shortly after the fire was started. Weather conditions on 23 April permitted moderately hot and fast burns. Winds up to 15 mph or more prompted fast sweeping headfire tongues into the sites followed by slower flanking fires on remaining vegetation. The temperature was approximately 75 F and relative humidity was low to moderate. Sites composed mostly of needle-and-thread did not burn completely due to excessive green vegetation. The large burn on site 1-86 was completed under less windy conditions on 30 April. Residual vegetation was dry but considerable green vegetation promoted an excellent moderately cool burn. A portion (approximately 16 ha) of the herbicide treated site (Snyder 1986) was also burned on 30 April. Approximate areas within the 1985 and 1986 burns are summarized in Table 5. Table 5. Approximate sizes of controlled burns and number of transects (N) per burn in 1985 and 1986, Tamarack Prairie. Site Ac 1985 Burn Ha N Ac 1986 Burn Ha N Ac Combined total Ha N 1 2 3 4 44 66 36 17.8 26.7 14.5 5 8 4 114 47 67 40 46.0 19.0 27.1 16.1 7 4 8 158 113 103 63.8 45.7 41.6 12 12 12 Total 146 59.0 17 268 108.2 19 374 151.1 36 Vegetation Sampling and Evaluation on Burned Sites Height-Density Sampling.--Snyder (1986) reported that controlled burns on sites 1 and 3 in 1984 markedly reduced the height-density (HDI) quality of residual grass-forb and sandsage cover in spring 1985 (~<0.05). Subsequent sampling in late winter 1986 revealed that post-fire vegetation recovery continued on both sites but neither had returned to pre-treatment HDI quality (Table 6). Although mean HDI values in 1986 for grass-forb vegetation remained below those in pre-treatment status (1984) on both sites, analyses could not detect that differences existed (p >0.05). In contrast, visual obstruction by sandsage was still markedly reduced on both sites as it was when vegetation was combined (Table 6). Both grass-forb and sandsage showed significant recovery from 1985 to 1986 within burn 1-84 when analyzed separately but not when combined. Significant vegetation recovery within burn 3-84 from 1985 to 1986 was not detected, if it occurred. �339 Table 6. Height-density (dm) means of residual grass-forb and sandsage vegetation within 1-84 and 3-84 burn sites and their controls from late winter 1984 (pre-treatment) through 1985 and 1986 (post-treatment) intervals, Tamarack Prairie. Vegetation Treatment Pre1984 1985 Post1986 1984-85 F values 1984-86 1985-86 Burn 1-84 Grass-forb Fire Control 0.253 0.252 0.136 0.296 0.232 0.301 47.5 4.24 8.19 Sandsage Fire Control 0.871 0.762 0.310 0.671 0.358 0.643 40.1 14.98 8.06 Fire Control 0.373 0.337 0.162 0.355 0.256 0.368 66.1 10.51 0.02 Combined Burn 3-84 Grass-forb Fire Control 0.222 0.183 0.021 0.191 0.106 0.200 22.5 5.96 2.32 Sandsage Fire Control 0.827 0.935 0.121 0.797 0.356 1.044 114.1 35.31 0.91 Fire Control 0.492 0.531 0.047 0.503 0.201 0.629 187.0 53.18 0.14 Combined ap <0.05. A preliminary analysis was conducted comparing pre-treatment and post-treatment HDI by combining all 17 transects within the 3 sites as being within one burn. Sandsage was not present on all transects and was excluded from preliminary analysis. However, grass-forb and combined vegetation HDI's were both greatly reduced in post-treatment sampling (p < 0.05, Table 7). Covariance analyses of site means provided evidence of marked reduction in cover quality for grass-forb, sandsage, and total vegetation (p< 0.05). Differences between pre- and first year post-treatment site means (ignoring possible year effect) were analyzed using paired t tests by including data from burn 1-84, burn 3-84, and the 3 1985 burns. -Findings indicated that the HDI was reduced (p< 0.05) for grass-forb, sandsage, and total vegetation. �340 Table 7. Height-density (dm) means of residual grass-forb and sandsage vegetation within 3 sites burned in 1985 and their controls from pre-treatment 1985 to post-treatment 1986 intervals, Tamarack Prairie. Vegetation site Treatment 1985 1986 1985 Control 1986 0.507 0.381 0.180 0.379 0.138 0.119 0.053 0.110 0.374 0.346 0.158 0.323 0.325 0.280 0.180 0.277 1.008 0.679 0.642 0.744 0.417 0.342 0.438 0.388 0.872 0.615 0.537 0.627 0.828 0.745 0.767 0.771 0.142 0.130 0.088 0.124 0.444 0.369 0.323 0.380 0.391 0.335 0.465 0.382 F 1,31 F 1,3 Grass-forb 1 2 3 x 96.62a,b 49.82a No data 95.85a Sandsage 1 2 3 x Combined ve~etation 1 2 3 ~ 0.565 0.407 0.302 0.429 115.25a 33.31a ap < 0.05. bY1 31based on transect means combining sites. Data were inadequate for analysis within sandsage. F1,3 based on site means. Two years of pre-treatment HDI have been collected for the burns conducted in 1986. The impact of the fire on the residual vegetation quality will not be measured until late winter 1987. Site means for grass-forb, sandsage, and total vegetation in 1985 and 1986 varied (Table 8). �341 Table 8. Height-density (dm) means of grass-forb, sandsage, and combined vegetation during pre-treatment late winter 1985 and 1986 intervals within 3 sites burned in spring 1986, Tamarack Prairie. Control Treatment Vegetation Site Grass-forb 1 2 3 x Sandsage 1 2 3 x Combined 1 2 3 x 1985 1986 1985 1986 0.326 0.314 0.259 0.292 0.304 0.337 0.256 0.290 0.294 0.316 0.233 0.275 0.277 0.312 0.213 0.265 0.621 0.375 0.395 0.566 0.838 0.500 0.650 0.790 0.829 0.417 0.618 0.682 0.756 0.300 0.690 0.690 0.421 0.315 0.272 0.335 0.470 0.339 0.281 0.366 0.345 0.317 0.298 0.320 0.342 0.331 0.323 0.332 Removal of sandsage by fire not only reduced HDI in subsequent years but also reduced the percentage of times it was the primary visual obstruction. This is illustrated (Table 9) when pre- and post-treatment percentages are compared for burn sites and their controls. These data also provide an index of the relative abundance of sandsage among sites. Table 9. Percent total height-density samples obstructed by sandsage among the 1984, 1985, and 1986 controlled burn sites, Tamarack Prairie. Year burned 1984 1985 1986 Site 1 3 Treatment 1985 1984 19.4 44.7 /a / 13.3 26.2 1 2 3 x 11.6 8.7 26.3 13.8 1 2 3 x 32.1 0.9 8.4 15.5 a/ denotes burn treatment occurred. / / / / 1986 1984 12.7 38.1 16.7 46.3 Control 1985 1986 15.9 51.5 12.9 50.8 1.4 4.8 9.1 4.8 13.9 8.7 42.5 18.6 13.2 11.8 48.6 21.3 30.9 1.2 7.5 15.2 9.6 1.3 17.0 11.0 13.7 3.1 23.3 15.7 �342 Crown Cover, Composition, and Frequency of Occurrence.--Samp1ing was completed, using the point frame method, within the 3 sites burned in 1985, which included 17 transects (432 samples/transect) and their controls, and the 3 sites burned in 1986 (19 transects and their controls) during August 1986. For the 1985 burns this was the second post-treatment sample whereas it was the first post-treatment sample for the 1986 burns. Sampling was not conducted in 1986 within the 1984 burns or their controls. Crown cover was summarized by site and combined sites to show combined composition and frequency of occurrence data for the 1985 and 1986 burns and their respective controls (Tables 10-13). Crown cover data are most meaningful when compared with data from pre-treatment or previous year samples. Therefore, mean crown cover tallies per transect for selected species, groups of species, or covers along with analysis of covariance findings are summarized for the 1985 and 1986 burns (ignoring site analysis) (Tables 14, 15). In addition, pre- to post-treatment analyses, using paired t tests of site differences for selected or species groups were obtained and are included in the following discussions. Analyses which compare transect results and do not consider sites (analysis of covariance) for interpretive purposes, yield results pertinent only to the location of the burns and controls. In contrast, results based on site differences (paired t test) yield findings pertinent to the Tamarack Prairie as a whole (D. Bowden, pers. commun.). Blue Grama.--Based on analysis of covariance of transect means, fire did not enhance blue grama from pre- to first-year post-treatment intervals within the 1985 burns (Table 10). However, pre-treatment to second year (1986) post-treatment data were greater within the burn (p< 0.05) as were first to second year post-treatment results. Likewise, 1985 (pre-treatment) to 1986 (post-treatment) analysis revealed that the 1986 fires potentially enhanced blue grama crown cover (Table 15). Paired t tests of pre- to post-treatment site differences failed to discern differences for either the 1985 or 1986 burns (~>0.05) and the same was true when the 2-year data were combined ignoring possible differences among years. Thus, the evidence, while not conclusive, indicates fire may enhance blue grama. Point frame sampling within controls revealed pronounced decreases in blue grama crown cover within controls (Tables 14, 15). When managing sandsage-b1uestem rangeland in eastern Colorado for prairie grouse, blue grama, which offers extremely marginal cover, is not a preferred species. Thus, if fire caused it to decrease rather than increase, it would probably be better for prairie grouse management. Need1e-and-Thread.--Above average precipitation received in spring 1986 accompanied by excellent subsoil moisture prompted exceptional growth and seed production by this species. Cro,vu cover within controls increased dramatically from levels in preceding years (Tables 14, 15). Fire had apparently suppressed (p <0.05) need1e-and-thread in burn 1-84 and within the 1985 combined burns (p <0.05, Snyder 1986) based on first-year post-treatment analysis. The same finding was evident following the 1986 burn (p <0.05, Table 15). Need1e-and-thread had attained considerable growth at-the time of both 1985 and 1986 burns. Therefore, it is not surprising that fire suppressed first year post-treatment growth of this species. However, data (Table 14) and visual inspections revealed that need1e-and-thread recovered markedly during the second growing season following fire within 1985 burns �343 Table 10. Crown cover (point frame), species composition (%), and frequency of occurrence of vegetation within treatment transects on 3 sites burned in 1985, Tamarack Prairie, August 1986. Vegetation Bare ground Dead vegetation Boute1oua gracilis Stipa comata Sporobo1us cryptandrus Ca1amovi1fa longifo1ia Andropogon ha11ii Agropyron smithii Aristida sp. Bromus tectorum Panicum virga tum Cyperus & Carex spp. Artemisia fi1ifo1ia Opuntia sp. Ambrosia psi10stachya Artemisia 1udoviciana Tradescantia occidenta1is Evo1vu1us nutta1ianus Psora1ea tenuif10ra Physalis subg1abrata The1esperma megapotimicum Mentze1ia nuda Ipomoea 1eptophy11a Liatris sp. Oenothera nutta11ii Croton texensis Chenopodium album Cryptantha sp. Pepidium densif10rum He1ianthus petio1aris Cirsium sp. Lactuca sp. Crown cover 2 3 1 436 783 59 303 50 361 23 21 674 1,163 89 425 226 552 47 129 432 520 202 92 83 86 87 27 3 36 3 43 108 4 11 2 1 2 2 9 1 1 43 13 51 13 17 1 2 9 15 8 1 1 10 2 5 17 1 3 17 1 1 1 5 4 3 2 1 1 Total Compo Freq. occurr. 1,542 2,466 350 820 359 999 157 177 3 36 43 16 202 17 28 3 6 19 16 9 1 11 2 1 1 6 39 4 4 5 1 1 10.49 24.58 10.76 29.95 4.71 5.31 0.09 1.08 1.29 0.48 6.06 0.51 0.84 0.09 0.18 0.57 0.48 0.27 0.03 0.33 0.06 0.03 0.03 0.18 1.17 0.12 0.12 0.15 0.03 0.03 0.82 1.00 1.00 1.00 0.65 0.94 0.06 0.06 0.12 0.18 0.71 0.35 0.29 0.18 0.18 0.35 0.18 0.18 0.06 0.24 0.06 0.06 0.06 0.12 0.71 0.12 0.18 0.18 0.06 0.06 �344 Table 11. Crown cover (point frame), species composition (%), and frequency of occurrence of vegetation within 3 1985 control sites, Tamarack Prairie, August 1986. Vegetation Bare ground Dead vegetation Boute1oua gracilis Stipa comata Sporobo1us cryptandrus Ca1amovi1fa longifo1ia Andropogon ha11ii Paspa1um stramineum Agropyron smithii Aristida sp. Bromus tectorum Muh1enbergia pungens Cyperus & Carex spp. Artemisia fi1ifo1ia A. fi1ifo1ia (dead) Opuntia sp. Ambrosia psi10stachya Artemisia 1udoviciana Tradescantia occidenta1is Evo1vu1us nutta1ianus Psora1ea tenuif10ra Physalis subg1abrata The1esperma megapotimicum Hap10pappus spinu10sus Mentze1ia nuda Sphaera1cea coccinea Erigeron sp. Croton texensis Chenopodium album Cryptantha sp. Pepidium densif10rum Conzya canadensis Erioqonum annum Euphorbia sp. Sa1so1a ka1i Cirsium sp. He1ianthus petio1aris Lactuca sp. Argemone sp. Crown cover 2 3 1 176 1,059 69 405 43 159 34 1 30 1 443 1,570 50 643 205 224 10 1 155 532 565 60 95 128 66 34 2 6 1 3 131 2 1 6 7 9 1 1 65 5 1 195 7 16 4 2 5 7 6 2 2 2 1 8 5 5 6 10 11 1 2 18 2 14 2 3 2 1 1 4 1 Total Compo Freq. occurr. 1,151 3,194 179 1,143 376 449 78 2 187 1 6 1 4 391 14 18 6 2 16 16 1 6 3 4 1 6 10 13 2 1 3 26 5 7 14 2 3 2 1 5.97 38.11 12.54 14.97 2.60 0.07 6.24 0.03 0.20 0.03 0.13 13.04 0.47 0.60 0.20 0.07 0.53 0.53 0.03 0.20 0.10 0.13 0.03 0.20 0.33 0.43 0.07 0.03 0.10 0.87 0.17 0.23 0.47 0.07 0.10 0.07 0.03 1.00 1.00 1.00 0.88 0.41 0.12 0.82 0.06 0.06 0.06 0.18 0.76 0.29 0.35 0.12 0.06 0.59 0.29 0.06 0.12 0.12 0.06 0.06 0.18 0.12 0.24 0.06 0.06 0.18 0.47 0.12 0.18 0.12 0.12 0.06 0.06 0.06 �345 Table 12. Crown cover (point frame), species composition (%), and frequency of occurrence of vegetation within treatment transects on 3 sites burned in 1986, Tamarack Prairie, August 1986. Vegetation Bare ground Dead vegetation Boute1oua gracilis Stipa comata Sporobo1us cryptandrus Ca1amovi1fa longifo1ia Andropogon ha11ii Agropyron smithii Aristida sp. Bromus tectorum Panicum capi11are Cyperus & Carex sp. Artemisia fi1ifo1ia A. filifo1ia 0Euntia sp. Ambrosia psi10stachya Artemisia 1udoviciana Tradescantia occidenta1is Evo1vu1us nutta1ianus Psora1ea tenuif10ra The1esperma megapotimicum Mentze1ia nuda Ipomoea 1eptophy11a Sphaera1cea coccinea Oenothera nutta11ii Eriqeron sp. Croton texensis Cryptantha sp. Pepidium densif10rum Lactuca sp. Euphorbia sp. Cirsium sp. Amaranthus sp. He1ianthus sp. Conzya sp. Crown cover 2 3 1 1,304 524 238 246 146 218 118 11 2 2 703 197 82 183 154 304 6 39 1,458 366 404 302 166 331 5 130 1 10 5 6 13 3 35 40 43 18 8 19 6 128 6 3 24 5 15 Total 4 4 10 5 18 4 2 4 2 1 2 9 6 7 2 16 45 4 5 14 1 2 7 6 1 3,465 1,087 724 731 466 853 129 180 3 2 15 12 141 9 38 64 43 27 23 19 4 2 26 68 4 13 18 1 2 1 18 6 7 6 1 Compo Freq. occurr. 19.80 19.99 12.75 23.33 3.53 4.92 0.08 0.05 0.41 0.33 3.86 0.25 1.04 1.75 1.18 0.74 0.63 0.52 0.11 0.05 0.71 1.86 0.11 0.36 0.49 0.03 0.05 0.03 0.49 0.16 0.19 0.16 0.03 0.89 1.00 1.00 0.84 0.47 0.63 0.11 0.05 0.16 0.21 0.53 0.26 0.37 0.42 0.16 0.32 0.26 0.21 0.05 0.05 0.21 0.37 0.05 0.32 0.32 0.05 0.11 0.05 0.26 0.11 0.05 0.05 0.05 �346 Table 13. Crown cover (point frame), species composition (%), and frequency of occurrence of vegetation within three 1986 control sites, Tamarack Prairie, August 1986. Vegetation Bare ground Dead vegetation Boute1oua gracilis Stipa comata Sporobo1us cryptandrus Ca1amovi1fa longifo1ia Andropogon ha11ii Agropyron smithii Aristida sp. Muh1enbergia pungens Panicum capi11are Cyperus & Carex spp. Artemisia fi1ifo1ia A. fi1ifo1ia (dead) Opuntia sp. Ambrosia psi10stachya Artemisia 1udoviciana Tradescantia occidenta1is Phlox andico1a Evo1vu1us nutta1ianus Physalis subg1abrate The1esperma megapotimicum Ipomoea 1eptophy11a Cichorium intybus Sphaera1cea coccinea Erigeron sp. Croton texensis Chenopodium album Pepidium densif10rum Eriogonum annum Conzya sp. Cirsium sp. Crown cover 2 3 1 232 1,560 201 461 50 278 5 10 185 775 20 347 92 181 9 53 468 1,557 121 361 148 247 8 38 4 3 5 157 9 13 15 4 7 13 4 1 5 2 1 1 4 1 1 5 1 19 1 22 1 1 15 238 21 26 24 7 3 11 8 1 3 Total Compo Freq. occurr. 885 3,892 342 1,169 290 706 22 101 4 3 5 15 399 30 39 15 31 20 5 8 1 5 2 1 30 1 23 5 1 9 2 3 10.40 35.56 8.82 21.48 0.67 3.07 0.12 0.09 0.15 0.46 12.14 0.91 1.19 0.46 0.94 0.61 0.15 0.24 0.03 0.15 0.06 0.03 0.91 0.03 0.70 0.15 0.03 0.27 0.06 0.09 0.95 1.00 1.00 1.00 0.42 0.68 0.05 0.05 0.11 0.05 0.79 0.21 0.58 0.05 0.21 0.47 0.11 0.32 0.05 0.05 0.05 0.05 0.11 0.05 0.11 0.16 0.05 0.11 0.11 0.05 �Table 14. Crown cover (mean point frame tallies/transect) for selected species and species groups among pre- and post-treatment samples within combined 1985 burn sites, Tamarack Prairie. Species or group Pre 1984 Boute1oua gracilis Stipa comata Sporobo1us cryptandrus Ca1amovilfa longifolia Andropogon hallii Combined warm-season gr. Artemisia fi1ifo1ia Combined perennial forbs Combined annual forbs Bare ground Dead vegetation 28.5 31.4 4.8 36.8 2.6 39.4 22.5 4.4 4.1 30.9 260.1 Treatment Post 1985 1986 21.2 27.7 21.6 54.5 7.9 61.8 8.2 3.4 2.8 110.3 170.1 20.6 48.2 21.1 58.8 14.3 70.5 11.9 5.7 3.5 90.7 145.1 Pre 1984 30.8 31.3 8.1 33.1 4.5 35.9 24.7 2.9 3.6 42.2 249.3 Control Post 1985 1986 21.9 55.2 22.7 28.5 8.8 32.8 22.6 2.1 3.5 47.1 217.9 10.5 67.2 22.1 26.4 9.8 31.1 23.8 4.2 4.2 67.7 187.9 84-85 F values 84-86 0.11 15.99a 0.46 17.11a 0.68 19.02a 30.01a 0.18 0.10 92.20a 32.35a 5.91a 4.70 0.64 20.77a 5.65 37.20a 15.96a 0.19 0.34 30.10a 14.74a 85-86 8.00a 6.98a 0.07 1.69 4.93 6.96a 3.06 0.06 0.09 3.80 3.68 ap <0.05. VJ .p. -....J �w p. Table 15. Crown cover (mean point frame tallies/transect) for selected species and species groups among preand post-treatment samples within combined 1986 burn sites, Tamarack Prairie. Species or group Pre 1984 Boute1oua gracilis Stipa comata Sporobo1us cryptandrus Calamovilfa longifo1ia Andropogon ha11ii Combined warm-season gr. Artemisia fi1ifo1ia Combined perennial forbs Combined annual forbs Bare ground Dead vegetation 37.3 42.8 10.3 26.9 1.9 28.9 18.1 7.2 6.6 39.5 234.2 ap <0.05. Treatment Post 1985 1986 36.1 40.0 13.7 25.3 2.4 27.6 16.2 4.7 1.6 41.1 241.4 38.1 38.5 24.5 44.9 6.8 51. 7 7.9 15.4 3.2 182.4 57.2 00 Control Pre 1984 1985 Post 1986 F value 1985-86 D.F. 40.6 28.1 4.6 39.5 0.6 40.1 14.4 5.1 3.1 28.9 263.9 34.6 52.0 14.8 33.8 1.1 34.9 19.3 3.0 1.3 32.6 226.6 18.0 61.5 15.3 37.2 1.2 38.3 22.6 6.3 2.3 46.6 204.8 8.93a 6.14a 8.11a 17.00a 7.67a 20.83a 21.16a 4.26 0.15 180.89a 137.04a 1, 1, 1, 1, 1, 1, 1, 1, 1, 1, 1, 35 35 35 34 15 35 26 31 24 35 35 �349 (p <0.05, Table 14). Thus, any negative impacts of fire on this species are apparently short lived. Paired! test analyses of site differences provided no evidence of significant impacts of fire on this species. Needle-and-thread and blue grama were both listed as increasers under controlled burning management by Kirsch and Kruse (1973). Although needle-and-thread attains much greater height and cover value than blue grama, it does not stand well over-winter especially under snows and does not provide residual cover of high height-density quality for spring nesting in residual by prairie grouse. Sand Dropseed.--Whereas the 1985 burns showed no evidence of impacting sand dropseed, the 1986 fires apparent.Ly stimulated increased growth when compared with controls (Tables 14, 15)" The species had also. shown evidence of increased crown cover follo.wing fire in burn 1-84 (Snyder 1986). Both 1985 and 1986 appeared to.be better grotdng years for the species because it was less abundant in most 1984 samples" Paired t t.estanalyses of site differences could not detect significant changes if they occurred for this species, and the different findings lead to unc.ertainty that the species was enhanced markedly by fire. The species is less abundant: than needle-andthread but possesses cover values and stand:i.ngqualities that are similar. Prairie Sandreed.--Among the 4 dominant grasses within the Tamarack Prairie, this species seems to respond most favorably to fire. Within the 1985 burns, pre- (1984) to post-treatment 1985 and 19.86 data showed major responses of prairie sandreed to fire (p < 0.05) and the same was evident in 1986 burns (p < 0.05) (Tables 14, 15). Paired t tests of differences also were noted withi~ 1986 burns and when 1985 and 1986 data were coabfned for analysis (~< 0.05). Controlled burns were conducted after initial growth of prairie sandreed had begun in both 1985 and 1986 and the fires stimulated rapid new growth of the species within a few days following the fires. In 1984, when no major response to fire was noted, burning occurred well ahead of greenup of the species. At present, prairie sandreed provides the best co.ver, both green and residual among dominant native grasses within the Tamarack Prairie. However, in most locations its HDI value is still considered marginal in quality for use by nesting prairie grouse. Sand Bluestem.--This species, while widespread and common within the Tamarack Prairie, is one of the less abundant species7 and therefore is not frequently found within point frame sampling. Data from the 1985 burns (Table 14) provided limited, but inconclusive evidence that the species was enhanced by fire (P > 0.05). Evidence was more positive within the 1986 burns (p < 0.05, Table IS). Samples were not adequate to detect differences if they-occurred within sites using paired t tests. However, visual observations and data support the contention that sand bluestem is an increaser under fire management (also reported by Kirsch and Kruse 1985). Combined Warm-season Grasses.--This group is dominated by pra1r1e sandreed and sand bluestem, but also includes switchgrass (Panicum virgatum) and sand paspalum (Paspalum stramineum) (Tables 10-13). As would be expected, the group, like its dominant species, shows strong evidence of enhancement by fire from both the 1985 and 1986 burns (p< 0.05, Tables 14-15). Fire response apparently even carried into the second year following fire based on data CTable 14) for the 1985 fire. Paired t tests of pre- to post-treatment site �350 differences were also noted from the 1986 fires and for the 1985 and 1986 burns in combination (P < 0.05). Sandsage.--Fire removed the canopy cover of sandsage and burns conducted after considerable leafing had occurred tended to retard regrowth but caused little if any mortality. Within the 1985 burns, sandsage canopy cover was still greatly reduced after the second post-treatment growing season (p <0.05), (Table 14) and the 1985 to 1986 recovery was not significant (R >0.05). Annual and Perennial Forbs.--Data presented this year (Tables 14, 15) as in previous years, (Snyder 1985, 1986) provided little evidence that either annual or perennial forb crown cover was enhanced or hindered (P>0.05) by controlled spring burns. Some species such as spiderwort and sweet pea had made considerable growth prior to burns conducted in 1985 and 1986 (Table 4), but these perennials recovered quickly since soil moisture reserves were adequate. Bare Ground and Dead Vegetation.--Resu1ts, as in preceding years, show that fire drastically reduces dead vegetation and increases the amount of bare ground. The transition back to increased litter and dead vegetation is a slow process (Table 14). Influence of Time of Burning on Sandsage.--Spring controlled burns, while showing evidence of retarding sandsage, have not significantly impacted survival (Snyder 1985, 1986). Sampling in late August 1985 of sandsage within the 24 June 1985 fire set by lightning provided uncertain evidence that 11 of 50 plants had been killed but some late budding was still occurring. Therefore, the marking pins were left for subsequent observations in 1986, which revealed that 44 of the 46 plants where pins could be found were still alive. Thus, the preliminary tally had been inaccurate and sandsage continued to resprout into fall. A cooler second fire had burned following a late afternoon shower at the site. August 1985 inventory revealed 79 of 81 survivors and 20 May 1986 tallies showed 78 of 80 survivors. Two fires burned north from Interstate 76 on 5 July 1986 that were caused by a burning truck tire. Late August 1986 inventory of sandsage survival at the 2 locations was conducted. Only 14 of 50 plants showed evidence of life at that time on the upper site, however, as during the previous year, some plants were just starting to sprout so the marking pins were retained for subsequent inventory. At a lower hillside location, where the fire was not as intense, 42 of 50 sands age plants showed evidence of survival. Revegetation Treatments Ti11age-Reseeding.--Successfu1 revegetation of 19 strips (approximately 20 ha) to a tall, warm-seeded grass mixture dominated by switchgrass in 1985 was reported earlier (Snyder 1986). Use of a low rate of atrazine herbicide (Snyder 1985) prior to seeding was a major factor in stand establishment and rapid first-year growth. A low rate of the herbicide was applied again in early spring 1986 to continue suppression of annual forbs while the grasses gained further growth. Results were highly successful as indicated by HDI averaging 2.6 dm among samples obtained at transects within 4 of the strips in fall 1986. Height-density within nearby stands of need1e-and-thread and prairie sandreed dominated undisturbed vegetation averaged 1.0 dm. Subsequent resamp1ing of these transects in late winter 1987 will provide a comparison of overwinter standing between the 2 vegetation types. �351 First year vegetation crown cover, composition, and frequency of occurrence data were summarized earlier (Snyder 1986). Comparison of crown cover between 1985 and 1986 late-summer intervals varied (Table 16). Little change was noted in crown cover of perennial grasses that were dominant prior to tillage. Needle-and-thread had been almost totally eliminated by tillage, whereas the other species showed moderate survival. Blue grama and western wheatgrass (Agropyron smithii) both increased moderately from 1985 to 1986. Bluestems (primarily seeded) and switchgrass in combination increased markedly (p <0.05) within the 12 transects from 1985 to 1986 showing their increased dominance within the strips (Table 16). Witchgrass (Panicum capillare), an annual, declined along with Texas croton (Croton texensis) the only annual forb tallied within 1985 samples. Tillage Renovation of Interseeded Tracts.--Based on results of preliminary 1985 trials (Snyder 1986), management personnel shallow disked, harrowed, applied atrazine herbicide at a low rate, and seeded switchgrass, where needed, in approximately 23.6 ha (58 ac) of tracts that had previously been interseeded in 1981 and 1982. These renovation treatments were completed during spring 1986. The majority of the 30 small tracts contained fair to good stands of seeded tall, warm-season grasses but their growth was suppressed by other vegetation. Tillage and herbicide reduction of competition prompted excellent growth of the deep-rooted seeded species during the 1986 growing season. One tract, where the impact of controlled burning had been evaluated in 1985 (Snyder 1986) was used to monitor the impact of tillage renovation in 1986. One-half of the site, containing 15 random transects, was disced, harrowed, and treated with atrazine, whereas the rest containing an equal number of transects, was retained as a control. One transect within the treatment portion was lost during tillage. Evaluation of pre- and post-treatment vegetation crown cover samples revealed that seeded tall grasses (primarily bluestem and switchgrass) increased dramatically within the tillage renovation tract (~< 0.05), (Table 17). Prairie sandreed, a native deep-rooted species showed no evidence of change (p> 0.05). Blue grama, needle-and-thread, sand dropseed, and other minor grasses were reduced markedly by the tillage (~< 0.05), (Table 17). Some reduction of perennial forbs from treatment seemed apparent but sampled were too small to detect a significant reduction, if it occurred. Controlled Burning vs. Tillage Renovation.--Observation of deep-rooted tall, warm season grass responses to fire and tillage treatments within the Tamarack Prairie prompted an attempt to compare seed production and growth responses among treatments and years. Two species, prairie sandreed and sand bluestem, which are not destroyed by the tillage used in revegetation (1985) and renovation (1986), were selected. These grasses, especially prairie sandreed, are expected to provide the dominant cover for prairie grouse habitation in future years on the site. �352 Table 16. Crown cover between 1985 and 1986 sampling intervals from 12 random transects within the 1985 revegetation strips, Tamarack Prairie. Vegetation or cover Bare ground Dead vegetation Tally 1985 Subtotal 804 212 Tally 812 156 968 1,016 Bouteloua gracilis Stipa comata Calamovilfa longifolia Sporobolus cryptandrus 25 40 66 126 57 1 128 226 217 Agropyron smithii Panicum capillare Cyperus & Carex Paspalum stramineum 17 10 4 64 5 5 11 78 Andropogon sp. (native & seeded) Panicum virga tum 8 37 75 400 303 311 Artemisia filifolia Opuntia sp. 5 475 7 3 1 6 Psoralea tenuiflora Sphaeralcea coccinea Physalis subglabrata Ipomoea leptophylla Oenothera sp. Cichorium intybus Evolvulus nuttalianus Liatris sp. Croton texensis 1986 Subtotal 1 2 10 1 1 3 5 8 3 1 6 1 1 22 76 12 �353 Table 17. Average crown cover/transect of selected species and species groups from pre-treatment (1985) to post-treatment (1986) intervals within a tillage renovation site and its control, Tamarack Prairie, 1986. Species or species group 1985 Tillage 1986 1985 Control 1986 Andropogon-Panicum Calimovilfa longifolia Boutelous, Stipa, Sporobolus, Agropyron, etc. Perennial forbs 13.7 11.3 17.6 24.2 12.4 5.4 16.4 10.7 17.1 16.6 10.6 16.8 1.5 0.6 2.2 1.7 a(p < Fl,26 value 34.68a 1.04 62.lla 1.76 0.05). Results of late summer 1986 sampling varied (Table 18). Findings for both were similar but prairie sandreed consistently yielded greater HDI's than sand bluestem among all treatments and controls illustrating its greater cover value. Among the 3 years of burns, results provided evidence that HDI's peaked following the second growing season after fire and then began to regress in the third year. Mean heights and seed production also tended to regress by the third year (Table 18). Comparisons in subsequent years are needed to determine if these patterns will continue or are a result of other environmental influences. The tillage (plus herbicide) treatments in 1985 and 1986 were much more severe treatments than burning but stimulated more dramatic responses from these deep-rooted species since shallow-rooted competition was reduced. Second-year lillI'swere considerably greater than those following the first growing season, in part because of the accumulation of residual by the second year. Mean height and seed head production also were better following the second growing season but samples in tillage sites also tended to have greater heights and seed head abundance than samples in burned sites. Continued sampling is needed to document trends over several years. Findings provide evidence that tillage renovation may be the best approach for dramatically increasing the HDI value especially when accompanied by revegetation to alter composition toward increased tall, warm-season composition. However, costs are much greater than when controlled burning is used. Data provided in Snyder (1986) indicated that once the composition was altered toward tall, warm-season grasses, fire would be a more effective renovation technique for use on the prairie in the future. Strip Spraying of Sandsage Treatment of an 80.8-ha tract within the Tamarack Prairie with 2,4-D herbicide aerially applied in June 1985 was previously reported (Snyder 1986). An approximate l6-ha tract within the site was control burned in 1986. Twelve point-frame sample transects had been randomly positioned within the spray site and sampled in May 1985. First-year post-treatment sampling of vegetation crown cover was completed in May 1986. Pre- and post-treatment �354 Table 18. Rate of seeding (%), mean height (dm), and height-density (dm) of prairie sandreed and sand b1uestem among burn and tillage renovation treatments and controls within the Tamarack Prairie, late summer, 1986. % N Treatment or control Transects Samples X K HDI Seeded Height 84 84 72 72 72 72 0-20 0-25 25-75 25-50 50-75 40-70 4.9 4.7 7.8 6.3 10.4 8.0 0.59 0.43 1.00 0.63 2.16 1.07 84 84 72 72 72 72 0-20 '\,25 >50 >75 >75 50-80 2.9 4.0 8.6 8.7 10.2 7.9 0.19 0.32 0.88 0.36 1.93 0.40 Prairie sandreed Untreated controls 1984 burns 1985 burns 1986 burns 1985 revegetationa 1986 tillage renovationa 7 7 6 6 6 6 Sand b1uestem Untreated controls 1984 burns 1985 burns 1986 burns 1985 revegetationa 1986 tillage renovationa 7 7 6 6 6 6 aSamp1ing was conducted within old native stands of the 2 species that survived tillage because of their deep-rooted growth form. crown cover tallies indicated that sandsage mortality was nearly 100% with only 1 tally of a live plant in May 1986 (Table 19). A walking tally of sandsage survival was also conducted in June 1986. Only 17 of 689 plants (2.5%) were alive providing additional evidence of the degree of herbicide impact. Post-treatment (1986) sampling also provided evidence of a pronounced increase in perennial grasses (424 to 784), especially need1e-and-thread. However, increases in this species were evident throughout the rangeland in 1986 (Table 14, 15). Perennial forbs, which were virtually absent in summer 1985 following herbicide treatment, recovered markedly in 1986 (Table 19). However, the total number of tallies (96) was much reduced from pre-treatment sampling levels (223) (P < 0.05) and the number of species was also reduced. Data from 1985 and 1986-burn site controls provide no evidence that perennial forbs as a group should have been significantly lower in 1986. Therefore, the reduction is directly related to impact of the herbicide treatment. Numbers and species of annual forbs, while much less abundant, were also reduced in 1986. �355 Table 19. Crown cover (point frame) of vegetation and ground cover within 11 random transects obtained during pre- and post-treatment (May) intervals within the June 1985 herbicide treated tract, Tamarack Prairie. Vegetation or cover Bare ground Dead vegetation Perennial grass Bouteloua gracilis Stipa comata Sporobolus cryptandrus Calamovilfa longifolia Andropogon hallii Muhlenbergia pungens Koleria cristata Aristida sp. Annual grass Bromus tectorum Festuca sp. Pre-treatment (1985) Post-treatment (1986) 402 1,425 529 1,324 142 194 34 51 148 510 46 65 3 2 3 1 9 1 70 117 4 Artemisia filifolia (alive) A. filifolia (dead) 436 112 259 Opuntia & Echinocereus spp. 44 50 Perennial forbs Ambrosia & Artemisia spp. Tradescantia occidentalis Lathyrus polymorphus Psoralea tenuiflora Phlox andicola Evolvulus nuttalianus Abronia fragrans Cymopteris montanus Allium textile Mentze1ia nuda Leucocrinum montanum Penstemon angustifo1ius Thelesperma megapotimicum Annual forbs Croton texensis Chenopodium sp. Pepidium & Lesguerella sp. Cryptanthia sp. Unid. forbs 5 18 114 55 4 6 2 2 1 1 9 77 6 1 1 1 12 1 2 2 3 24 1 3 2 1 1 �356 LITERATURE CITED Kirsch, L. M., and A. D. Kruse. 1973. Prairie fires and wildlife. Timbers Fire Eco1. Conf. 12:289-303. Proc. Tall Snyder, W. D. 1985. Sandsage-b1uestem pralrle renovation. Job Prog. Rep., Colorado Div. Wi1d1. Fed. Aid Proj. 01-03-045. Apr. (In press). 1986. Sandsage-b1uestem prairie renovation. Job Prog. Rep., Colorado Div. Wi1d1. Game Research Rep. Fed. Aid Proj. 01-03-045. Apr. (In press). Prepared by ~~~~~~~~.Warren D. Snyder Wildlife Researcher ~ _ �357 Colorado Division of Wildlife Wildlife Research Report April 1987 JOB PROGRESS REPORT Colorado State of: Project: W-152-R (W-37-R) Work Plan: 21 Job Title: Response of Selected Wildlife Species to Aspen Silvicultural Practices Period Covered: Author: : Job Avian Research 4 01 July 1985 through 31 December 1986 Richard W. Hoffman Personnel: C. E. Braun, R. W. Hoffman, and T. E. Remington, Colorado Division of Wildlife ABSTRACT Literature review and communications with agencies interested in the aspen-type were continued as were efforts to identify research needs concerning aspen-wildlife relationships that could be addressed by the CDOW. ��359 RESPONSE OF SELECTED WILDLIFE SPECIES TO ASPEN SILVICULTURAL PRACTICES Richard W. Hoffman Quaking aspen (Populus tremuloides) dominated plant communities comprise nearly 2.3 million acres of commercial forest lands in Colorado (Green and Setzer 1974). Much additional acreage is occupied by non-commercial stands and mixtures ot aspen and conifer species. Together these types offer important water, forage, wildlife, and recreational values. Aspen typically occurs in clones produced asexually by adventitious sprouting from a single parent root system (Schier 1981). Seedling establishment is rare due to short-lived seed and demanding seedbed requirements (Maini 1968, McDonough 1979). Sprouting can be stimulated by destroying the existing stand (Crouch 1981, Schier 1981). For centuries, fire was the natural regenerative force responsible for the perpetration of aspen (Gruell and Loope 1974). Twentieth century forest management policies have emphasized fire suppression. The result is that most aspen stands in the Central Rockies have reached the mature to overmature categories and are in need of treatment if they are to be maintained. In the absence of fire, clearcutting is the most effective regenerative procedure (Jones 1975, Schier and Smith 1979, Crouch 1981). Until recently, there has been little commercial demand in Colorado for aspen for manufacturing wood products; consequently, mass treatments (clearcuts) within the aspen type have been virtually non-existent. This situation is changing as the Louisiana Pacific Corporation (LP) plans to process aspen chips into waferboard, a compound type panel that LP feels may eventually replace conventional plywood. LP has built 2 mills, 1 near Montrose and another in Kremmling, and plans to harvest 2,500 acres of aspen/year/mill to maintain an annual production of 25 million board-feet of waferboard. The realization that large scale aspen manipulation programs may soon become common practice in Colorado has generated a pressing need for information on the impacts these treatments will have upon other associated resources. Aspen-wildlife relationships are of particular interest because little quantitative information exists. Aspen has long been recognized for its cover and forage benefits to wildlife, but this understanding has not progressed beyond the descriptive stage. Aspen will be managed for its timber producing values and manipulation will occur with or without input from wildlife managers. Ironically, forest managers are interested in, and have requested input from wildlife specialists in developing aspen management strategies that are congruent with wildlife requirements. �360 P. N. OBJECTIVES The objectives of this planning study are to: 1. Examine and evaluate existing literature on aspen-wildlife relationships to identify information needs pertinent to the development of a detailed study plan. 2. Interview selected personnel of the CDOW, USFS, BLM, and Louisiana Pacific Corporation concerning research needs within the aspen type. 3. Prioritize research needs and identify funding sources. 4. Prepare a detailed study plan on an approved research topic dealing with aspen-wildlife relationships. SEGMENT OBJECTIVES 1. Review literature pertaining to aspen ecology, aspen silvicultural practices, aspen-wildlife relationships, effects of aspen maniplation on wildlife populations, and chemical and nutritional characteristics of aspen. 2. Interview selected personnel of the CDOW, USFS, BLM, Colorado State Forest Service, and Louisiana Pacific Corporation concerning aspen-wildlife research needs. 3. Interview selected personnel involved in aspen research and management in other states to identify potential problems and assist in the selection of an appropriate research topic. 4. Visit proposed apsen timber sales in Colorado that are potentially suitable for conducting research. 5. Prepare a detailed study plan on an approved research topic and select study areas to meet research needs. 6. Obtain funding to support the research project. RESULTS AND DISCUSSION Cooperation from the U.S. Forest Service and Louisiana Pacific (LP) and control over the treatments to be imposed are prerequisites to the further development of a study plan. No commitments were made due to the political and environmental issues surrounding the proposed treatment of aspen in Colorado. The future of LP in Colorado is uncertain. As a result, the decision was made not to proceed with an aspen study, but to continue open communications with other agencies interested in the aspen type and to pursue other avenues of aspen-wildlife research that could be addressed by the CDOW. A community study in the aspen type is outside the realm of the CDOW research section and it is questionable whether such a study would provide meaningful �361 information for making management decisions. A study designed to measure changes in abundance and distribution of wildlife species in response to aspen manipulation is premature and should not be implemented until the forementioned political and environmental issues are settled. Such a study cannot proceed without complete control over the treatments to be imposed. Studies of how and why selected wildlife species use the aspen type have important management implications, fall within the research capabilities of the CDOW, and can be conducted independent of other agencies. By understanding not only what species use the aspen type, but why and how they use it, wildlife managers will be better equipped to predict and mitigate the impacts of aspen manipulation on wildlife populations. This information is also prerequisite to designing studies to measure wildlife responses to aspen manipulation. LITERATURE CITED Crouch, G. L. 1981. Regeneration of aspen clearcuts in northwestern Colorado. U.S. Dep. Agric., For Servo Res. Note RM-407. 5pp. Green, A. W., and T. S. Setzer. 1974. The Rocky Mountain timber situation, 1970. U.S. Dep. Agric., For. Servo Resour. Bull. Int-lO. 78pp. Gruell, G. E., and L. L. Loope. 1974. Relationships among aspen, fire, and ungulate browsing in Jackson Hole, Wyoming. U.S. Dep. Agric., For. Serv., Int-unpubl. Rep. 33pp. Jones, J. R. 1975. Regeneration on an aspen clearcut in Arizona. Agric., For. Servo Res. Note RM-285. 8pp. U.S. Dep. Maini, J. S. 1968. Silvics and ecology of Populus in Canada. Pages 20-69 in J. S. Maini and J. H. Cayford, eds. Growth and utilization of poplars in Canada. Canada Dep. For. and Rural Dev. Publ. 1205. McDonough, W. T. 1979. Quaking aspen-seed germination and early seedling growth. U.S. Dep. Agric., For. Servo Res. Note Int-234. l3pp. Schier, G. A. 1981. Aspen regeneration. Pages 15-21 in N. V. DeByle, ed. Situation management of two intermountain species: -aspen and coyotes. Part 1 - Aspen. Utah State Univ., Logan. , and A. D. Smith. 1979. Sucker regeneration in a Utah aspen clone following clearcutting, partial cutting, scarification, and girdling. U.S. Dep. Agric., For. Servo Res. Note Int-253. 6pp. ��Colorado Division of Wildlife Wildlife Research Report April 1987 363 JOB PROGRESS REPORT State of Colorado Project 01-03-045 21 Work Plan Job Title: 5 Evaluation of Habitat Quality on Conservation Reserve lands in Eastern Colorado Period Covered: Author: : Job Avian Research 01 July through 31 December 1986 Warren D. Snyder Personnel: C. E. Braun, G. P. East, J. F. Lipscomb, D. L. Schrupp, and W. D. Snyder, Colorado Division of Wildlife ABSTRACT Study progress included obtaining background information as a basis for developing a sampling design. Coordination was conducted to include the Colorado Division of Wildlife within a nation-wide effort to assess the value of the Conservation Reserve Program for key wildlife on a region-wide basis. Limited monitoring of the Division's efforts to enhance vegetation quality for wildlife on retired croplands was conducted. Progress toward development of a revised program narrative for the study continued. ��365 EVALUATION OF HABITAT QUALITY ON CONSERVATION RESERVE LANDS IN EASTERN COLORADO Warren D. Snyder The Conservation Reserve Program (CRP), as part of the 1985 federal farm program administered by the U.S. Department of Agriculture, was implemented in early 1986. Its purpose was to place perennial vegetation on highly erodible croplands to reduce soil erosion and to remove these croplands from grain production for 10-year intervals. The program's acceptance by farmers in many dryland farming areas of Colorado was demonstrated by commitment of 14.25% of the eligible cropland in eastern Colorado to the program by the end of the fourth (Feb 1987) sign-up interval. The 25% maximum of eligible cropland per county had been reached in several southeastern counties and others are approaching this maximum. In better farmland areas of northeastern Colorado, the commitment to CRP remained less intensive. This cropland retirement program may have considerable benefits to wildlife, especially in better farmlands along the eastern edge of Colorado and in states to the east. However, since much of eastern Colorado is marginal for farming as well as for wildlife, uncertainty as to benefits to wildlife remains. Concern for the species of perennial covers being planted and their values for wildlife led to implementation of a cost share incentive program wherein the Division of Wildlife committed more than $270,000 to encourage the use of taller grasses that will benefit wildlife as well as to encourage planting of shrub thickets within CR tracts. This evaluation study was implemented to assess the potential impacts of the CR program on wildlife within the plains of eastern Colorado. The program could potentially impact several species of plains upland game as well as many non-game wildlife species. Evaluations will also attempt to assess the value of the Division's supplemental cost-share efforts to enhance cover quality. P. N. OBJECTIVES Identify (1) the geographic relationship between distribution of Conservation Reserve lands in eastern Colorado and the distribution and relative density of ring-necked pheasants, lesser prairie-chickens, and greater prairie-chickens, (2) the height-density quality of the herbaceous vegetation occurring on Conservation Reserve lands, and (3) the general vegetation composition of these lands. In addition, a comparison will be made between the height-density quality of tracts receiving supplemental CDOW cost share payments and those not receiving supplemental payments to plant selected grasses. SEGMENT OBJECTIVES 1. Coordinate with the Colorado State ASCS office of the USDA, Don Schrupp, Gordon East, and regional management personnel to develop a computer mapping system to relate Conservation Reserve lands to wildlife distribution and density in eastern Colorado. Examine available data on a sample basis to project costs. �366 2. Develop a stratified random sampling design for visual obstruction (height-density) and general composition measurements of randomly-selected CR tracts in eastern Colorado. 3. Develop a comprehensive Program Narrative covering Conservation Reserve Program in eastern Colorado. 4. Prepare an annual progress evaluation of the report. RESULTS Possible data transfer from USDA - Agriculture Stabilization and Conservation Service (ASCS) state office computer files directly to software for use by D. Schrupp to plot the distribution of the Conservation Reserve (CR) in Colorado using Colorado Division of Wildlife computer mapping facilities was explored. Gordon East coordinated a meeting among personnel of the 2 agencies. However, it was determined that State ASCS office files did not contain a legal description of approved CR tracts preventing direct data transfer for computer mapping. Discussions with personnel of district offices of the USDA Soil Conservation Service (SCS) revealed that at least some county or district offices were plotting CR tracts on county maps. Therefore, plans are being formulated to photocopy or manually transfer CR data from district offices. If the essential data are not available at SCS offices, personnel of county ASCS offices will be contacted to obtain the necessary information. Field inspections and contacts were made in fall 1986 to monitor the Division's supplemental cost share program designed to improve cover quality on selected CR tracts. Recommendations for program modification were submitted. Plans are in progress to include an evaluation of these efforts within the study design. Considerations for developing a stratified random sampling design have been discussed with the Division's statistician, D. C. Bowden. Background information has been collected concerning monitoring the impact of CRP on certain wildlife species. An interagency meeting concerning evaluation of the values of the Conservation Reserve Program (CRP) on wildlife was attended in March 1987. Personnel of the National Ecology Center (U.S. Dep. Interior, Fish and Wildlife Service), State agencies, ,and other agencies and organizations were in attendance. A proposal was developed for the National Ecology Center to coordinate nationwide effort by state agencies to monitor the impact of the CRP on selected wildlife species. Tentative plans are to use Natural Resource Inventory (NRI) quarter-sections that previously have been randomly selected by the SCS as a basis for monitoring CRP, assuming enough NRI tracts contain CR. Habitat suitability index models (HSI) will potentially be used to determine pre- and post-CRP qualities of these tracts for wildlife. Within Colorado, this effort will be one part of a more comprehensive monitoring program. a As of March 1987, 14.25% (1,412,659 acres) of 9,010,800 acres of cropland has been committed to the CRP within 23 eastern Colorado counties (Table 1). Most southeastern Colorado counties have reached or slightly exceeded their maximum allowable commitment of 25% of eligible cropland. Farmers in most east-central Colorado counties had committed 10-20% of eligible cropland. Less �367 than 10% of the cropland in northeastern Counties had been committed, and in better dry1and farming areas, the rat~ was 5% of less. However, during the recent, fourth (Feb 1987) sign-up interval, farmer commitment to the CRP was much greater than during previous intervals. Table 1. Acres and percent of eastern Colorado cropland accepted into the Conservation Reserve Program as of March 1987. Conservation reserve % Acres County Total croE1and Adams Arapahoe Bacaa Bent Cheyenne Crow1eya Elberta E1 Paso Huerfanoa Kiowaa Kit Carson Las Animas Lincoln Logan Morgan Otero Phillips Prowersa Pueb10a Sedgwick Washington Weld Yuma 621,800 201,400 1,032,000 116,000 642,000 84,700 208,000 96,500 6,800 685,000 885,000 90,000 512,000 654,700 242,300 84,600 373,000 530,000 114,700 211,000 838,000 1,037,000 644,300 21,499 31,947 278,757 26,808 119,724 23,672 52,031 12,963 1,950 192,108 103,465 19,359 88,359 22,636 21,926 2,597 8,800 144,847 32,239 4,828 76,580 93,295 32,448 3.46 15.86 27.01 23.11 18.65 27.95 25.01 13.43 28.68 28.04 11.69 21.51 17.22 3.46 9.05 3.07 2.36 27.33 28.11 2.29 9.14 9.00 5.04 Total/Mean 9,910,800 1,412,659 14.25 aDesignates counties where the maximum allowable percent of Conservation Reserve has been attained. �368 Prepared by 11)a»uMJ ~ Warren D. Snyder Wildlife Researcher C �369 Colorado Division of Wildlife Wildlife Research Report April 1987 JOB PROGRESS REPORT Colorado State of: Project: W-152-R (W-37-R, W-88-R) 1 (6-1) Work Plan: 22 Job Title: Avian Research Publications Period Covered: Author: : Job Avian Research 01 January through 31 December 1986 Clait E. Braun Personnel: C. E. Braun, P. O. Dunn, K. M. Giesen, J. K. Ringelman, W. D. Snyder, M. R. Szymczak ABSTRACT Publications accomplished under this job in 1986 are: Braun, C. E. 1986. Review-Bobwhites in the Rio Grande Plain of Texas by V. W. Lehman. Wilson Bull. 98:492-493. , and K. M. Giesen. 1986. Population characteristics of white-tailed ptarmigan in Colorado. Proc. Conf. Science in the National Parks-1986. Abstract and Poster Paper. .:~c.-· Dunn, P.O., and C. E. Braun. 1986. Late summer-spring movements of juvenile sage grouse. Wilson Bull. 98:83-92. -----', and • 1986. Summer habitat use by adult female and juvenile sage grouse. J. Wildl. Manage. 50:228-235. Snyder, W. D. 1986. A matter of respect. Colorado Outdoors 35(6):11-12. Szymczak, M. R., and J. K. Ringelman. 1986. Differential habitat use of patagial-tagged female mallards. J. Field Ornith. 57:230-232. Prepared by -:=-:;---:->-,tIa~V,,---::-_f __ ~~~=-c.;:.__ Clait E. Braun Wildlife Research Leader _ ��371 Colorado Division of Wildlife Wildlife Research Report April 1987 . JOB FINAL REPORT Colorado State of: Project: 01-03-045 (N-5-R) : Job Avian Research 2 Work Plan: 23 Job Title: Ciconiiform Reproductive Success and Public Viewing in the San Luis Valley, Colorado Period Covered: Author: 01 January 1984 through 30 June 1986 Janet L. Schreur Personnel: E. Decker, D. A. Hein, R. A. Ryder, R. G. Walter, Colorado State University; C. E. Braun, J. L. Schreur, B. Widhalm, Colorado Division of Wildlife ABSTRACT The reproductive success of colonially-nesting Ciconiiformes and effects of public viewing were investigated in the San Luis Valley, Colorado between April 1984 and August 1985. Black-crowned night-herons (Nycticorax nycticorax) snowy egrets (Egretta thula), cattle egrets (Bubulcus ibis), and white-faced ibises (Plegadis chihi) nested at Russell Lakes, Russell Lakes State Wildlife Area, Head Lake, Adam's Lake, and Monte Vista National Wildlife Refuge. Ninety-eight percent of the nests were in hardstem bulrush (Scirpus acutus) and 2% were in common cattail (Typha latifolia). Overall black-crowned night-heron nesting success was 54% and differed (X2, P<O.05) among heronries. Productivity ranged from 0.0 to 1.9 nestlings/nest-attempt. Overall snowy egret success was 49% in 1984 and 60% in 1985. Productivity ranged from 0.3 to 2.4 nestlings/nest attempt. Black-crowned night-heron, snowy egret, and white-faced ibis productivity may not b~ sufficient to maintain stable populations. One cattle egret nest was observed in 1984 'and it was successful. All 3 cattle egret nests failed in 1985. Overall white-faced ibis nest success was 47% in'1984 and 68% in 1985. Productivity ranged from 0.3 to 2.4 nestlings/nest attempt. Flooding, predation, and human disturbance caused serious nest losses. Mean flushing distances of black-crowned night-herons, snowy egrets, and white-faced ibises from a person on foot were 152.5, 93.3, and 94.5 m, respectively. Mean flushing distances from a motor vehicle were 46.5, 56.8, and 53.7 m respectively. ��373 ACKNOWLEDGMENTS Funding Division for this investigation of Wildlife. for initiating I extend this project and patience. field assistance Refuge. I am grateful My sincere Decker, special thanks and his unfailing The Alamosa/Monte provided was provided to Dr. to these my major advisor, support, agencies for their is also extended and Dr. and understanding. Professor R. G. Walter for his encouragement I thank friendship, cold marsh at first E. Davey, ship, light. S. MacVean, and G. Davey contributed Lakes. data analysis Finally, egrets, for their and Dr. D. A. Hei.n design. my field assistant, balking E. I also thank of the study and never and information Russell cattle criticism support. to Professor R. A. Ryder assistance, support, enthusiasm, Vista National Wildlife Refuge guidance, In addition, C. E. Braun for work on Monte Vista National Wildlife appreciation for his constructive by the Colorado E. Edmiston, for at wading into the murky, K. Crandall, valuable B. Widhalm, assistance, friend- at Monte Vista National Wildlife Refuge S. Geor ge+s patience and and help were invaluable during and writing. I thank the black-crowned and white-faced They were tolerant ibises of my presence night-herons, snowy egrets, which made this study and their beauty possible. and grace never �374 ceased to fascinate species me. I expect and people with a desire this study to benefit to view them. these �375 TABLE OF CONTENTS ABSTRACT OF THESIS ACKNOWLEDGMENTS v LIST OF TABLES ix UST • OF FIGURES xii Chapter I II INTRODUCTION METHODS •.•. Census and Reproductive Nest Site Characteristics. Flushing Distance 1 Success III STUDY AREA IV" RESULTS Nesting Phenology . Arrival and Nest Construction • • . . Egg Laying, Incubation, and Hatching . Nest Census . . . . • • . . . • .• Nesting Success . . • . • . • • Black-crowned Night-heron. Snowy Egret .••• Cattle Egret . White-faced Ibis Productivity . . . . Black-crowned Night-heron. Snowy Egret. . • • White-faced Ibis . Nest Site Description. Flushing Distance . • 4 4 6 7 8 16 16 16 17 20 22 22 24 26 26 29 29 33 34 36 47 �376 TABLE OF CONTENTS (Cont'd) Page Chapter V VI DISCUSSION. • . . Census. . . .. Nesting Success and Human Disturbance. Monte Vista National Nest Sites . . . • • .... .. . . . . . Productivity .•. • . . • • • • Wildlife Refuge 71 RECOMMENDATIONS. LITERATURE APPENDIX. CITED . • . . . . . . • • • . .• Marsh Bird Disturbance Trials American Coots • . • • • Redheads •.. Ruddy Ducks Mallards . • . Gadwalls •..• American Avocets Conclusions . • • • • . 50 50 53 64 65 67 .... 73 83 84 85 85 89 89 89 91 91 �377 LIST OF TABLES Table 1 2 3 4 5 6 7 8 9 10 11 12 Colonial-nesting Ciconiiformes nests in the San Luis Valley, Colorado, 1984- 85 • • . . 21 Nesting success of black-crowned night-herons in the San Luis Valley, Colorado, 1985 . • . • . 22 Nesting success of black-crowned night-herons in the San Luis Valley, Colorado, 1985 . . . . • . • . . 23 Nesting success of snowy egrets in the San Luis Valley, Colorado, 1984- 85. . . . • . • . . • • . 25 Nesting success of snowy egrets Valley, Colorado, 1984-85 27 in the San Luis . . • • . • Nesting success of white-faced ibises Luis Valley, Colorado, 1984-85 in the San . . • . • . 28 Nesting success of white-faced ibises Luis Valley, Colorado, 1984- 85 in the San . • . . . . 30 Distribution of clutch size of black-crowned nightherons in the San Luis Valley, Colorado, 1985 31 Distribution of brood size and mean number of nestlings per successful nest and nest attempt for black-crowned night-herons in the San Luis Valley, Colorado, 1985 • . . • . • . . • • • . . . . . • . 32 Distribution of clutch size of snowy egrets in the San Luis Valley, Colorado, 1984- 85 • . . . 33 Distribution of brood size and mean number of nestlings per successful nest and nest attempt for snowy egrets in the San Luis Valley, Colorado, 1984- 85. . • • • . . • • • • . . . • . . • . . . . 35 Distribution of clutch size of white-faced ibises in the San Luis Valley, Colorado, 1984-85 . . . . • • 36 �378 LIST OF TABLES (Cont'd) Table 13 14 15 16 17 18 19 20 21 Distribution of brood size and mean number of nestlings per successful nest and per nest attempt for white-faced ibises in the San Luis Valley, Colorado, 1984- 85 . . . . . . . • . . 37 Distances from nests water in 5 heronries Colorado, 1985 44 Distances from nest and random sites to the nearest Ciconiiformes nest in 5 heronries in the San Luis Valley, Colorado, 1985 • 45 Water depth at Ciconiiformes nest and random sites in the San Luis Valley, Colorado, August 1985 . . . . . . . . . . . . . . . . . . . . . . 46 Height of Ciconiiformes nests above the water's surface in 5 heronries in the San Luis Valley, Colorado, 1985. . . • . . . • • . • • . . . . . 48 Flushing distance of black-crowned night-herons, snowy egrets, and white-faced ibises approached in a vehicle or on foot, Russell Lakes SWA, Colorado, 1985. . . . . . . . . . . . . . . . . 49 Nesting success of black-crowned night-herons in the United States • . . • . . . . • • . • . 54 Productivity of black-crowned night-herons North America . . • . . . . • • • • • • • 56 Clutch size of black-crowned night-herons North America. • . . . • • • . 22 Productivity 23 Clutch 24 Quantification bance 25 and random sites to open in the San Luis Valley, • . . . . . • . of snowy egrets size of snowy egrets of marsh bird . . . . in in in the United States 59 . in the United States responses 61 62 to distur. . . . . 85 Marsh birds counted on a wetland before, immediately after, and at 15-minute intervals for 1 hour after disturbance by a person walking. • • . • . • 86 �379 LIST OF TABLES (Cont'd) Table 26 Page Marsh birds counted before, immediately after, and at 15-minute intervals for 1 hour after disturbance by a motor vehicle, 1985 . . • . . . . . • 87 �380 LIST OF FIGURES Figure 1 San Luis Valley in southcentral 2 Russell 3 Heronries Colorado 4 5 6 7 8 9 10 11 12 Lakes area, Colorado San Luis Valley, 9 Colorado .• on Monte Vista National Wildlife Refuge, . . . . . . . . . . Head Lake and San Luis Lake, Colorado 11 13 San Luis Valley, ................. 15 Incubation and hatching of colonial-nesting Ciconiiformes in the San Luis Valley, Colorado, 1984 . . . . . . . . . . . . . . . . . . . 18 Incubation and hatching of colonial-nesting Ciconiiformes in the San Luis Valley, Colorado, 1985 .... . . . . . . . . . . . . . . . . . 19 Ciconiiformes nest locations in the Head Lake heronry in 1985 . • • . • • . • . . • . 38 Ciconiiformes Lake heronry nest locations in the Harrence in 1985 . . • • • • • • . . • 39 Ciconiiformes SWA heronry nest locations in 1985 40 Ciconiiformes nest locations heronry in 1985 ..•.•...•... in the Russell Lakes . • • . • . . in the Parker Pond 41 Ciconiiformes nest locations in the Unit 18 heronry at Monte Vista National Wildlife Refuge in 1985 . . . . . . . . . . . . . . . . . 42 Frequency of American coot responses to (a) a person on foot and (b) a motor vehicle during disturbance trials in 1985 . . • • • . • • . . • 88 �381 UST OF FIGURES (Cont'd) Figure 13 14 15 Frequency of redhead responses to (a) a person on foot and (b) a motor vehicle during disturbance trials in 1985 . . • . • • • • 90 Frequency of gadwall responses to (a) a person on foot and (b) a motor vehicle during disturbance trials in 1985 • . • • . . . • . 92 Frequency of American avocet responses to (a) a person on foot and (b) a motor vehicle during disturbance trials in 1985 • . . . . . 93 ��383 CHAPTER I INTRODUCTION Ciconiiformes, attention market Plumage Bill was passed resulted Audubon hunters, ended in the country's Society of the century sportsmen, and poachers great were close to extirpation (Casmerodius when the Audubon in New York in May 1910. first to protect This law was one laws in the United States the plumage trade. Poaching wardens Florida's of and wildlife managers. Consequently, wildlife conservation successfully have a long history birdwatchers, 1928, Allen 1957). and snowy egrets of the first allies, value of plumes at the turn professional (Thompson albus) and their from conservationists, The high market attracted herons being of egrets hired remaining and also by the National heronries (Anderson 1978) • Although tions today, concentrations birds. illegal shooting has less poor reproductive success and loss of breeding Ciconiiformes night-herons, ing in the western on Ciconiiformes related habitat United States South America where DDT is still used contaminants. and white-faced winter this group of and insectivorous, to environmental snowy egrets, popula- to high pesticide still affect are lar gely piscivorous which makes them susceptible crowned effect ibises in Mexico, Central, (Ryder 1967, Ryder Blackbreedand 1978, �384 Ryder et al. 1979). Ciconiiformes Recently, poor reproductive has been correlated DDT) concentrations 1981, Findholt (Capen breed and Leiker and forage in wetlands. and actual loss of wetlands declines (Allen 1957). et al. documented water (1979), nest losses In Colorado Graul failures and Utah, (1980), caused 1980, Findholt 1984). Changes have caused in the population Alford (1978), and McEwen et al. by flooding of (1984) or abruptly lowered levels . . Nonconsumptive their 1979, Steele 1984, McEwen et al. stability Ryder of with high DDE (a metabolite 1984, Henny et al. Ciconiiformes success large bodies (Gray for bird wildlife users, and showy plumage, 1975). watchers and photographers, Investigation of the effects on sensitive wildlife species of the United States' is needed population Furthermore, are predicted U.S. (Yeosting activities to increased and indications Ciconiiformes, habit nest subjects of nesting to disturbance. wildlife recreation because participate by a large (Tyre proportion and James 1971, 1975, Fazio and Belli 1977, Shaw et al. Dep. Inter. numbers to increase and Burkhead In response their vulnerable 1979, More 1979, U.S. users are desirable of nonconsumptive Aney and Cowan 1975, Gray Samson 1983). to Ciconiiformes may also be causing While Ciconiiformes colonially makes them particularly 1978, Arthur attracted based 1982, Boyle and of nonconsumptive on demographic wildlife trends in the 1973, Belli 1977). demand for nonconsumptive of low reproductive the Colorado Division success wildlife of of Wildlife and Colorado State �385 University Area initiated this investigation. (RLSWA) and the surrounding for study cattle (Graul because egrets, black-crowned and white-faced 1977, Ryder was being considered The objectives des, to breeding night-herons, for development were to: methods of public and nesting success snowy egrets, were known to nest there 1979, McEwen et al. of this study Lakes State Wildlife San Luis Valley were selected ibises (2) document reproductive and (4) test Russell 1984). of public Also, RLSWA viewing opportunities. (1) locate and census success, (3) describe viewing that were least of Ciconiiformes. nest heronsites, detrimental �386 CHAPTER II METHODS Census and Reproductive Field work began ended on 16 August heronries Success on 18 April in both years. were searched estimated were located Care was taken the sun's to prevent were surveyed, numbered number of eggs and/or April 1985. was effect blocked nest early in the acute angle of Calm con- requirements. egret, were marked species, on nest of nestlings. were additional cattle each heronry. and white-faced but in 1985, all colonial-nesting tag and the date, nests were conducted heat exhaustion Active nests had on foot due to access wading through vegetation and no precipitation were included. during of snowy egret Nest searches In 1984, only snowy egret, nests activity to minimize investigator-related when emergent rays historically in 1985. by quietly and production. the morning that on 15 May 1984 and 3 June but the number from aerial photos Nests success aircraft 1985 and in the San Luis Valley were at Adam's Lake was not visited and time constraints, ditions All wetlands from a fixed-wing The heronry Wetlands on foot for nesting and May in both years. searched 1984 and 5 April ibis Ciconiiformes with a 2 x 2-cm number, young were recorded. location, A nest was and �387 considered Adults active if it contained were not marked Therefore, the number Each active nest incubated and could not be individually of renesting was revisited attempts every 10 days of age nestlings Nesting would run from which nest success traditional estimate, had at least the total number the nestling tivity at my approach using the nests. Nests of these nests and success period by hatching success or or produc- 1982). into incubation incubation usually (Maxwell and Kale 1977). (Erwin and Custer for snowy egrets 1 20-day period period for white-faced In this and nestling begins after Therefore, 1982, Custer (J enni a more accurate 1978, Johnson were combined into 1 25-day period The nestling which would have overesti- (Miller and Johnson was divided In Ciconiiformes, day period during to produce and Nichols 1981, Erwin and Custer night-herons (nests the and productivity. Hensler incubation nests in nesting of nest egg is layed method To calculate found estimate vals. making it diffi- ltraditional' of successful The Mayfield method is believed the nesting At to 10 days of age) was divided Inclusion 6-8 they originated. were not included calculations. mated success survive of active period 10 days of age. (Mayfield 1961, 1975). the number 1 nestling was not determined. reached was calculated and the Mayfield method distinguished. 9-11 days in 1984 and every days in 1985 until it failed or young cult to ascertain eggs or live young. 1979, study inter- the first egg laying and for black-crowned et al. 1983), 1 22- 1969, Maxwell and Kale 1977), ibises (Archibald was 10 days for all 3 species. et al. 1980). The probability �388 of a nest rate surviving these X 100 to yield nest 2 periods was multiplied by the hatching success. Nest Site Characteristics Heronries Refuge at Russell (MVNWR) were cover old) aerial photos and the U.S. obtained described cover National Wildlife mapped in 1985 from recent of Agriculture, was cover by Mosby (1980). «3 year- Soil Conservation mapped by intersection Nest locations Service. as were plotted on each selected in 1985 map. Variables were: measured (.1) distance Ciconiiformes depth nest, (±1 cm), at each randomly (±0.1 m) from nest (2) distance and (4) height An area was considered diameter and contained variables, A sign except test nest 'open height, nest to the nearest neighboring (±0.1 m) to open water, (±1 ern) of nest water' vegetation. 20 m in All of these were also measured (Steel and Torrie (3) water above the substrate. if it was at least no emergent of the medians at random sites. 1980) was used to for differences. All Ciconiiformes sites and Monte Vista from the Colorado Diviaion of Wildlife Department The Head Lake heronry test Lakes were measured Due to the large sample of nest random sites nests at the Russell number sites and a comparative of nests Lakes of random and Head Lake heronries. at MVNWR heronries, for each species were measured. number a random and an equal number of �389 Flushing Distance FIushing egrets, distances and white-faced Snowy egret flushing 1984 and April-June and black-crowned 1985. located of black-crowned Groups ibises distances 1985. were measured Flushing on clear calm mornings. at a slow, steady motor vehicle ance agent measured rate of speed until it flushed. during distances by either to the spot where the bird(s) with a Leitz rangefinder. birds was then a person The distance June-August during and feeding Each group Lakes. of white-faced were measured of resting snowy at Russell were measured night-herons (1-19 birds) night-herons, ibises April-June were approached on foot or in a (±2 m) from the disturb- had been standing was �390 CHAPTER III STUDY AREA The San Luis Valley, in the southcentral portion lake bed is bordered and the Continental west. The mountain ranges (Fig. to the south winters. (Fig. 1) had an average annual Dep. the Russell Lakes 5 C during 1982 through during 29 C have occurred area), into At Saguache through June 1984). summers (16 km north annual of temperature was as low as - 30 C have February, while highs (U.S. Dep . Commerce and July and in the San Luis Valley Temperatures December short of of 4.8 C from 1931 to the average 1983. during on the has a maximum width temperature Commerce 1978). study ancient on the the Valley extends drainage is to form the by cool, sunny, The Rio Grande 1960 (U.S. been recorded This flat, from 2,285 to 2,400 m (Svoboda The climate is characterized dry park, Mountains meet at Poncha Pass varying cold, 1). de Christo The Valley is 240 km long, 80 km , with elevation intermountain Divide in the San Juan Mountains limit of the Valley; New Mexico. largest of the state by the Sangre east northern Colorado's of 1985, 1986). The growing season date of the last frost is 18 September (U.S. is short is 4 June Dep. and dry. In Saguache, the mean and the mean date of the first Commerce 1978). Average annual frost �391 COLORADO • DENVER • GRAND JUNCTION • COLORADO SPRINGS • PUEBLO o, 10 , 20 , KILOMETERS ~ 00 RUSSELL LAKES Fig. 1. San Luis Valley in southcentral Colorado. �392 precipitation was 26.8 em from 1931 through months were July and August crn , respectively Although tion run-off marshes. A low divide north half is drained a high water shallow, of the Rio Grande basins (Ryder northern and Sangre closed basin. River separates 1951) (Fi g. 1). Streams de Christo draining mountains This water is used for irrigation of black of the valley floor is dominated greasewood (Chrysothamnus (Sarcobatus spp.), vermiculatus), and inland salt grass Wetlands are .dominated by hardstem balticus), emergent dominates plants and common cattail. Baltic rush in water .::.15em in depth. in water include (Lemna minor), water crowfoot It by an association rabbitbrush stricta). baltic rush (Juncus is the dominant Hardstem from 15 to 150 ern in depth. sago pondweed flow into (Distichlis bulrush, the or evaporates. San Luis Lake is the low point of the "Clos ed Basin. Vegetation The to the south by the Rio Grande River, half is a closed basin. this table and irriga- alkaline lakes and while the northern San Juan of 3.6 and 4.2 Commerce 1978). in numerous 2 drainage The wettesf with monthly averages the climate is arid, result the Valley into southern (U. S. Dep. 1960. bulrush or cattail Associated (Potomogeton pectinatus), (Ranunculus aquatic duckweed trichophyllus), and water milfoil (Myriophyllum spicatum). Ciconiiformes nested at Russell Head Lake in the San Luis Valley. l06°07'W) included artesian wells (Fig. 6 naturally 2). Lakes, RLSWA, MVNWR, and Russell Lakes occurring Ciconiiformes lakes nested (37°57'N, fed by springs in hardstem and bulrush �t TO SAGUACHE I I ! , I COUNTY ROAD R WELL ~'J; ~ j \ ., "', I ~",-"A "'"", '''"'\ """\ <, "'T'''-'''-/ ! I() CD N RUSSELL LAKES I.. ui ::i JOHNSON LAKE N 1 1 MILE 1,2 KILOMETERS W Fig. 2. Russell Lakes area, San Lui s Valley, Colorado. 1.0 W �394 in Trites 2 (2.1 km of surface Lake 1.5 m) during free 1984. of emergent hardstem submerged feeding plant activities were Harrence Trites the Russell island lake's 0.35- km2 surface. tained carp of Russell 0.5-km2 were aquatic creating bulrush (Ondatra Lake, 2). 20% of the it was murky, aquatic con- vegetation. with a maximum depth of 2.1 rn , 2). covered Hardstem Division bulrush pers. was clear. habitat commun.). However, due to stable trout of Wildlife and resting (D. Lan glois, probably of in 1985 (Fig. covered as breeding and the water water submerged levels soil conditions. 3). 106°05'W) contained Both wetlands for waterfowl production. with a maximum depth of 1. 5 m, was open water, heronry Colorado and waterfowl t managed Muskrats RLSWA was a privately-owned It is managed MVNWR 37°28'N, and Unit 18 (Fig. (Fig. by the were sparse anaerobic wetland Lakes surface. few carp plants of wave action and and had sparse until purchased for non game birds Lakes As with Trites and muskrats, (CDOW) in 1982. There by with little of 1.1 m) , 0.4 km east of hardstem RLSWA was a man-made hatchery murky carpio). (maximum depth A crescent-shaped 60% of its was 50% was dominated as a result (Cyprinus surface abundant. contained 1. 4 km north 50% of the lake's was constantly growth of carp Lake Lake, with a maximum dept h of while the western The water aquatic zibethicus) The eastern plants, bulrush. water 25% was hardstem 2 heronries, were man-made Parker Pond Approximately bulrush, and Parker Pond and were covered 0.48 km 60% of the surface 15%was common 2 �~~ '9-~ ~o '1"k N '1( 1 1 MILE 1.2 KILOMETERS EIGHTMILE ROAD W Fig. 3. Heronries on Monte Vista National Wildlife Refuge, Colorado. \.0 \Jl �396 cattail. The water down. It supported populations. was clear and the wetland dense Muskrats aquatic was open water, common cattail. there Ciconiiformes (Fig. 4). depth of 1. 3 rn , nested Hardstem Herons between and ibises flowing, tions bulrush covered was dense nested and there areas area of open water. plant growth. The Carp in a wide portion of the channel point and the average was dense The channel was 0.1 water varied August. aquatic plant depth The water was growth. No carp were observed. were observed night-herons, snowy egrets, in the hardstem 105°51'W) on the from the air indicated emergent with a maximum 97%of the surface aquatic May to 0.2 m during Black-crowned (37°24'N, and were not observed. from 1. 2 m during ibises and 15%was of Head Lake in 1984 Head Lake and San Luis Lake in 1985. or muskrats of its growth. 3%was small scattered km in width at its widest clear, percent no carp were seen, 1. 4 km northwest was clear and there and muskrats bulrush, 2 was in a O.25-km wetland The heronry and the remaining water plant and invertebrate Seventy 15%was hardstem aquatic drawn carp were not observed. 2 0.30 km. The water was clear, was moderate growth were common, but Unit 18 was smaller and covered surface plant was regularly bulrush in Adams Lake 1984 and 1985 census that hardstem with willows (Salix spp.) and white-faced bulrush flights. Observa- was the dominant along the shoreline. �397 SPR ..._ING ...,,- CRE~K: __ .._ __r-- .....•-. ..._. ".._ .., ~\~ ....,-. (" --.( ... ,,: ""1,., ) " dJ· .. y N 1 } 1 MILE 1.2 KILOMETERS ) SAN LUIS LAKE SIXMILE LANE Fig. 4. Head Lake and San Luis Lake, Colorado. San Luis Valley, / �398 CHAPTER IV RESULTS Nesting Arrival and Nest Construction Snowy egrets, herons arrived All 3 species white-faced crowned were observed night-herons at Parker and herons and nest Parker at Parker were observed struction incubating Pond probably this heronry until then. at RLSWA on 1 May. not observed in the San Luis Valley on 10 April Lakes. sticks Lakes. began into the Two herons date. several was not visited Snowy egrets Arrival courtship On 16 April on this began ibises 1985. night-herons eggs 1985, White-faced carrying Pond and Russell day of and black- within a few days of arrival. but since it was not observed April in both years. Lakes on 17 April and black-crowned at Parker 16 April, observed at Russell night- Lakes on the first Four days later at Russell Pond were already construction than arrived construction 1985, both species heronries and black-crowned In 1985, snowy egrets were first observed Snowy egrets at Russell 1984. Pond on 6 April. were first ibises, in the San Luis Valley during field work on 18 April egrets Phenology began and nest at Head Lake or Unit 18 on MVNWR. at Nest days earlier every nest day, con- construction were �399 White-faced 1985. ibises began On 5 May, ibises nests at RLSWA. Parker nest construction were defending They began nest during territories construction May in - and constructing at Unit 18 and Pond on 21 and 30 May, respectively. Egg Laying, Incubation, Egg laying, incubation, within heronries. over a several faced ibises and Hatching and hatching Nests were initiated, week period nesting (Figs. at Parker and eggs laid and incubated 5 and 6). was completed in a 4-day period, initiation was highly with year, synchronous heronry, and incubation tion usually (Fig. indicating (Figs. and incubation 5 and 6). were combined since, egg is laid. In both years, Ciconiiformes began laying Snowy egret Parker Pond and Russell (2 Jun 1984) at Head Lake. laying several However, Lakes. days before exact dates first Egg laying Black-crowned snowy egrets began egrets laid eggs incuba- at Russell Lakes 14 days later night-herons began and white-faced ibises. night-herons laid eggs at Parker respectively. eggs were layed on 16 and 20 April at Russell Pond, The egg laying eggs were laid on 19 MaY' 1984 at and Russell Lakes on 11 and 12 April, Parker varied are not known for 1984. In 1985, black-crowned egret nest in Ciconiiformes, after the first Pond. that 5). begins and Parker of white- 1984 were an exception. of egg laying and species periods The 7 pairs Pond during Hatching Dates for the beginning were not synchronous respectively. When comparing 1 month earlier at Parker Pond The first snowy Lakes and the 2 years, snowy Pond and Russell Lakes �~ o o ~~SE ::r:....J W~ WFII ~~~ I ....J ....JCI) ~W CI)~ SE ~,~ -'!....J a: a: w 0 SE ~Z ~~ WFIr------------------------------------------[============~~ 5 9 13 17 MAY 21 25 29 31' 2 6 10 14 18 JUN 22 14 18 JUL Fig. 5. Incubation and hatching of colonial-nesting Ciconiiformes in the San Luis Valley, Colorado, 1984. The open portion of the bar represents the time period when eggs were being incubated, but no nests contained nestlings. The shaded portion of the bar represents incubation and hatching. During this period some nests contained nestlings and some contained only eggs being incubated. ' �o~ ~:5 ~(/) ~~ BCNH """""""""""""""""""""""""""",,'t :>:5 CI: SE 3: oct SE ~ WFI CI: CI: o ~Z ~~ a.. c:o .,.. I- Z :> ,,"""""""""""""'1 r-...."""""""""""""""""""""""""""""""""""""""""" "'1 """"""""""""""""" BCNH .,_" '" ~"""""""""""""","""","",""""",","",",'1 -0,.",,, BCNH -0,."'1 l'\."""" SE .' ,,,,,,,,,"'1 WFI """""""x""""- BCNH SE WFI I 2 4 e 8 10 12 14 18 .8 2022242828 APR JO • 2 I 4 8 8 1012 14 18 18 2022242830 MAY 1 • 2 4 1 8 8 1012 14 18 18202224282830' JUN ,,,- ,,,- "- I 2 4 .1 8 _L 8 1012 14 18 18 20222428 2830 JUL Fig. 6. Incubation and hatching of colonial-nesting Ciconiiformes in the San Luis Valley, Colorado, 1985. The open portion of the bar represents the time period when eggs were being incubated, but no nests contained nestlings. The shaded portion of the bar represents incubation and hatching. During this period some nests contained nestlings and some contained only eggs being incubated. +:- o I-' �402 in 1985 than in 1984. crowned General night-herons Egg laying began ibises Snowy egrets night-herons in heronries began laying numbered ibis nests they blac k+ in 1985. At Parker was greater the 2 species began (Figs. 1). laying egret or 5 and 6). white-faced Pond, than snowy egrets nested 30 days before by 3-7 times (Table of ibis nests nests, layed eggs before where over Pond in both years. of egret that also laid eggs one month earlier at Parker the number indicate at RLSWA and Unit 18 on 5 and 6 May in 1985. Black-crowned white-faced observations ibises nests In heronries out- where or equal to the number within 2- 8 days of each other. Nest Census During white-faced Colorado 1984, there ibis nests (Table 1). were 181 snowy egret, in 3 heronries All 3 species Lake from a fixed-wing aircraft, cattle egret, and in the San Luis Valley, were also observed but a ground at Adam's census was not attempted. During cattle egret, (Table 1). estimate 1985, 498 black-crowned and white-faced The 1985 nest of the number Luis Valley because ibis nests snowy egret, were counted in 6 heronries count is a much more representative of colonial-nesting (1) black-crowned included, (2) the number estimated from aerial photos, thorough. night-heron, night-heron of snowy egret and Ciconiiformes (3) nest nests nests in the San were at Adam's Lake was searches were more �Table 1. Colonial-nesting Ciconiiformes nests in the San Luis Valley, Colorado, 1984-85. N nests Black-crowned night- heron Heronry 1984 Head Lake Russell Lakes RLSWA Parker Pond Unit 18 Adams Lake Totals aBlack-crowned bDiscovered Snowy egret 1985 1984 1985 a a 0 a b c 33 52 5 126 30 c 27 35 0 73 ·b c - 246 135 night-herons during 1985. nested White-faced ibis egret 1984 1985 1984 0 16 13 70 8 32 1 0 0 0 b c 0 0 0 3 0 c 139 1 3 during It is not known cCiconiiformes were observed on nests ground census was not accomplished. The photos in 1985. Ca ttle 1984, but were if Ciconiiformes not Totals 1985 1984 1985 25 14 2 7 b c 5 0 22 18 68 c 52 49 0 80 b -- 38 68 40 214 106 32 46 113 181 498 counted. nested from a fixed-winged aircraft number of snowy egret nests at this site in previous years. in 1984 and 1985, but a was estimated from aerial ~ o w �404 Par ker Pond was the largest 43% (217) of Ciconiiformes (Table 1). nests In 1985, 106 nests Pond in Unit 18 on MVNWR. Unit 18 were combined, heronry con tainin g 44% (80) and in 1984 and 1985, respectively were located 1. 6 km east When the nests at Parker of Parker Pond and 64% (320) of the known Ciconiiformes nests in the San Luis Valley were on MVNWR in 1985. Nesting Black-crowned Overall eggs that Night-heron nesting were 2). during heronry (percent 1 young Nesting The poorest abandoned successful success had at least was 54% (Table heronry. Success survive success success of nests containing to 10 days differed 2 (X , of age) !:< was at RLSWA where incubation. Parker with 73% of the nests 1 or more for 1985 0.05) with all 5 nests Pond was the most producing young to 10 days of age. Table 2. Nesting success of black-crowned Luis Valley, Colorado, 1985. night-herons N Heronry Head Lake Russell Lakes RLSWA Parker Pond Unit 18 Totals Nests Successful Unsuccessful in the Success (%) 33 47 5 91 24 16 15 0 66 11 17 32 5 25 13 48 32 0 73 46 200 108 92 54 San �405 Nesting Between flooded success at Head Lake 5 and 14 June, water depth 11 (33%) black- crowned great-horned (48%) was reduced increased ni ght- heron owls (Bubo virginianus) to 32%at Russell Lakes. Nestlings Although predators were not observed indicated great-horned in a plains portion occurred cottonwood at night. tree by up were killed and the heronry, evidence for the losses. were found within A pair (Populus success by of the heronry. within 2 owl pellets Predation nesting owls were responsible were dismembered, and predation nests. from 15 nests in the southeast lings from 55 to 120 cm and lowered some of them eaten by floodihg. of great-horned sargentii), Nest- the heronry, owls nested 0.7 km north of the heronry. Nesting (Table 3). success Success was also calculated calculated using the Mayfield method by this method was 2.1 to 37.3% lower in each heronry than success calculated with traditional percentages, nests by the traditional at Parker method. As Pond were most success- ful (49. 1%). Table 3. Nesting success (Mayfield method) of black-crowned herons in the San Luis Valley, Colorado, 1985. night- Nest success Heronry Head Lake Russell Lakes RLSWA Parker Pond Unit 18 Incubation (A) Hatching (B) Nestling (C) 0.168 0.588 0.000 0.627 0.966 0.741 0.656 0.823 0.753 0.827 0.500 0.947 0.909 Nest success (%) (AxBxCx100) 10.3 29.1 0.0 49.1 43.9 �406 Snowy Egret Overall nesting 0.10<P <0.25) ranging 1984. success of snowy egrets from 1984 (49%) to 1985 (60%) (Table from 10 to 82%, differed In 1985 success 2 (X , did not differ 4). Success, (X2, P < 0.005) among heronries did not differ in (X2, 0.50 < P < 0.75) among heronries. The lowest nesting this study was at Russell scheduled nest abandoned. Nestlings, containing Lakes in 1984 (Table appearing was probably the heronry but not on 24 June. obtained froin landowners The photographers which was in the center and triggered photographers hours reported on 2 consecutive the bulrush around the under nests. by 2 photographers I observed their their activity on their activity of the heronry. who on activities was with the photographers. around nest #40, A camera was placed on a from an island 80 m away. The they were in the heronry from 0700 to 1000 days. were destroyed, the central Although nests surviving nests were on the periphery from nest #40. Nesting Lakes in 1985 to 71%. During but the eggs were cold. and correspondence remotely during 90% (35) of the nests Information concentrated 4). directly caused on 24 June. observed to have been dead for 2-4 days, or in the water 23 June, tripod I found only eggs were intact, Abandonment visited for snowy egrets count on 27 June, were lying in the nests Nests success success no nests was trampled. of the heronry improved The only 3 and farthest (X2, P < 0.005) at Russell �Table 4. Nesting success of snowy egrets in the San Luis Valley, Colorado, 1984-85. N Successful Nests Heronry 1984 1985 Head Lake Russell Lakes RLSWA Parker Pond Unit 18 17 29 0 39 -- 0 14 13 49 8 85 84 Totals 1984 7 3 Unsuccessful 1985 1984 1985 1984 -- 10 26 -- 41 10 -- 42 50 32 successful 1985 10 7 29 4 -- Percent 7 -- 4 6 20 4 -- 71 54 59 50 43 34 49 60 -- -82 .I>- o -....J �4Ub Success calculated using lower in each instance method (Table Parker 5). than the Mayfield method was 7.5 to 25% success calculated As with traditional percentages, Pond in 1984 were most successful. at Russell Hatching Lakes in 1984 reflects success success the effects were lost during calculation the traditional nests at The low (0.3%) success could not be calculated 90%of the nests using of nest for Russell the hatching abandonment. Lakes because period. The nesting would have been even lower if hatching success could have been included. Cattle Egret One cattle egret nest was located Head Lake in 1984 (Table lings surviving observed before Three cattle 3rd nest White-faced Overall ibises A pair Lakes heronry egret nests incubation. was Pond in 1985. of 2 nests disappeared was abandoned. Ibis nesting success increased 6). in both years (Table 6). 2 (X , 0.025 < P< 0.050) from Although increase occurred because success increased in the 2 heronries Success 0% and from 67 to 45%at Head Lake and Parker The overall egrets at Parker The contents from 1984 to 1985, it decreased nested of cattle with 3 nest- in 1984, but a nest was not were located 47%in 1984 to 68%in 1985 (Table overall was successful the mass abandonment. All 3 failed during and the This nest to 10 days of age. in the Russell identified 1). in the San Luis Valley at where decreased Pond, of the discovery from 73 to respectively. of the �Table 5. 1984- 85. Nesting success (Mayfield method) of snowy Nest Heronry Year Head Lake Russell Lakes Parker Pond Russell Lakes RLSWA Parker Pond Unit 18 1984 1984 1984 1985 1985 1985 1985 Incubation 0.459 0.031 0.726 0.916 0.777 0.783 0.726 (A) egrets in the San Luis success Hatching 0.467 --- 0.818 0.644 0.771 0.657 0.586 (B) Nestling 1. 000 0.103 0.956 0.861 0.572 0.818 1. 000 (C) Valley, Colorado, Nest success (AxBxCx100) 21. 4 0.3 56.8 50.8 34.3 42.1 42.5 +-- o 1.0 �.j> I-' o Table 6. Nesting success of white-faced ibises in the San Luis Valley, Colorado, 1984-85. N -- Successful Nests Unsuccessful Heronry 1984 1985 1984 1985 Head Lake Russell Lakes RLSWA Parker Pond Unit 18 11 13 0 6 8 2 -- -- 5 0 14 11 64 4 -- 9 5 50 30 94 14 64 Totals -- 0 Percent successful 1984 1985 1984 1985 3 11 -- 2 67 -- 5 6 14 -- 64 45 78 16 30 47 68 -- 5 73 15 -- 0 �411 Unit 18 heronry in 1985. ber of ibis nests Unit 18 contained (64) than any other over heronry 4 times the n um- and 78% of the nests were successful. 2 In 1984, success Lakes was lower (X , 0.005 < P< 0.010) at Russell than at Head Lake or Parker graphers resulting abandonment in abandonment nests and farthest by photo- also caused The 2 surviving of the heronry ibis nests from snowy egret #40. In 1984, nesting than The disturbance of snowy egret of 85%of the ibis nests. were on the periphery nest Pond. success at RLSWA, Parker were incubated but no eggs downstream nests, 2 (X , P was lower Pond or Unit 18. Eggs in ibis nest from the heronry and found 0.005) at Head Lake At Head Lake, for more than the normal hatched. = 5 ibis nests 20-day incubation period, #90, located 100 m over 2 weeks later than the other also did not hatch. Success instances, calculated except the traditional using the Mayfield method was lower in all at RLSWA in 1985, than method culated for Russell number of nests (Table 7). Hatching Lakes or Parker surviving success success calculated using could not be cal- Pond in 1984 due to the small incubation. Productivity Black-crowned Average Only nests Night-heron clutch size for 179 nests found during incubation, in 1985 was 3.8 (Table but after egg laying was 8). �~ I-' N Table 7. Colorado, Nesting success 1984- 85. (Mayfield method) of white-faced Nest Incubation (A) ibises in the San Luis Valley, success Hatching Heronry Year Head Lake Russell Lakes Parker Pond 1984 1984 1984 0.908 0.256 0.560 0.815 Head Lake RLSWA Parker Pond Unit 18 1985 1985 1985 1985 0.811 0.862 0.576 0.789 0.000 0.878 0.632 0.836 (B) Nestling (C) Nest success (AxBxCxl00) 0.862 1.000 1. 000 63.8 25.6 56.0 0.851 1. 000 0.941 0.0 64.4 36.4 62.1 �413 completed were used to calculate size estimates were not biased clutch size. Therefore, clutch by nests that failed before egg laying was completed. Mean clutch Mean clutch Head Lake size varied size was highest (3.2). Although were most frequent 2 (X , 0.025 <P < 0.050). among heronries at Russell Lakes mean clutch (3.9) and lowest size varied. 4-egg at clutches in all heronries. Table 8. Distribution of clutch size of black-crowned in the San Luis Valley, Colorado, 1985. night-herons N eggs Heronry 2 3 Head Lake Russell Lakes Parker Pond Unit 18 2 1 3 4 6 7 28 4 10 45 Totals The mean number ranged For example, per N 8 32 54 11 2 4 12 0 1 0 0 0 19 44 97 19 3.2 3.9 3.8 3.4 4 4 4 4 105 18 1 179 3.8 4 of nestlings successful the highest of nestlings per The mean number on mean clutch nest mean clutch attempt (Tables 9). (3.9) (3.1) 2 and 9). /nest a high mean clutch size and mean number hatching The mean number sizes. and highest to 10 days and depended in 0 nestlings after mean were at Russell surviving at RLSWA resulted Mode to mean clutch size nest x 10 days (Table was related of nestlings size surviving nest successful from 0.0 to 1. 9/nesting than 6 from 2.6 to 3.1/successful of nestlings number 5 4 ranged more on nest Zero hatching attempt. Russell of nestlings Lakes. success success Lakes per had successful �.pI-' .p- of brood size and mean number of nestlings per successful Table 9. Distribution attempt for black-crowned night-herons in the San Luis Valley, Colorado, 1985. N nestlings Heronry Head Lake Russell Lakes RLSWA Parker Pond Unit 18 0 1 2 3 4 5 17 32 5 25 13 1 1 0 11 3 7 3 0 17 2 6 5 0 22 3 2 5 0 15 3 0 1. 0 1 0 ~ nestlings / successful nest 2.6 3.1 --2.7 2.6 nest and nest ~ nest lin gs / nest attempt 1.2 1.0 0.0 1.9 1.2 �415 nest (Tables 8 and 9), but ing in a low mean number contrast, Parker mean number Pond had low nest of young had the of nestlings per per highest nest success nest attempt nesting attempt (Table 2) result- (Table success 9). In (73%) and (1. 9) . Snowy Egret Average Mean clutch 1984 (3.9) clutch size for 138 nests size was higher (Table while 5-egg 10). clutches in 1984 and 2 (X , P < 0.005) In 1984, 4-egg Table 10. Distribution Luis Valley, Colorado, of clutch 1984- 85. in 1985 (4.6) clutches were most frequent 1985 was 4.4. than in were most frequent, in 1985. size of snowy egrets in the San N eggs Year 2 3 4 5 6 N x Head Lake Parker Pond Totals 1984 1984 1984 1 2 3 3 7 10 3 17 20 5 6 11 0 1 1 12 33 45 4.0 3.9 3.9 5 4 4 Russell Lakes RLSWA Parker Pond Unit 18 Totals 1985 1985 1985 1985 1985 0 0 2 0 2 0 1 1 1 3 3 4 29 0 36 9 5 24 5 43 1 1 6 1 9 13 11 62 7 93 4.9 4.6 4.5 4.9 4.6 5 5 4 5 5 Heronry Mean clutch (3.9-4.0), (Table again 10). but sizes varied sample sizes Mean clutch sample sizes slightly among heronries were too small to test sizes varied in 1984 for significance more in 1985 (4.5-4.9), were too small to test Mode for significance. but �416 Mean number of nestlings surviving 10 days after ranged from 2.3 at Head Lake in 1984 to 3.4/successful Russell Lakes nestlings and RLSWA in 1985 (Table per nest attempt number of nestlings per reflected low nest success White-faced ranged nest 11). nest at Mean number of from 0.3 to 2.4. attempt caused at Russell by nest hatching The low mean Lakes in 1984 abandonment. Ibis Average Mean clutch clutch size for 129 nests size was not different when compared to 1985 (Table in 1984 and 1985 was 3.7. (X2, 0.95 < P < 0.975) in 1984 12). Four-egg clutches were most freq uen t in both years. Mean clutch heronries in 1984 (Table from 3.4 to 4.1. 2 (X , 0.25 < P < 0.50) size did not vary 12). In 1985, mean clutch A chi-square . and Unit 18 indicated test differences comparing (P < 0.005) because clutches at Head Lake, (Table were most frequent 12). Three-egg clutches size ranged RLSWA, Parker Pond, among the heronries. - Head Lake was not included among of small sample sizes. Parker were most frequent Pond, Four-egg and Unit 18 at Russell Lakes in 1984 and at RLSWA in 1985. The ranged mean number from 2.0 to 3.1/successful of nestlings attempt of nestlings surviving (Table Lake resulted 13). nest to 10 days (Table ranged Zero hatching in no successful The abandonment surviving nests of 85% of the nests 10 days 13). after hatching The mean number from 0 to 2.4/nesting success for the 5 nests and 0 nestlings at Russell Lakes at Head per nest in 1984 attempt. �of brood size and mean number of nestlings per Table 11. Distribution attempt for snowy egrets in the San Luis Valley, Colorado, 1984- 85. N of nestlings Heronry Year 0 1 2 3 4 5 Head Lake Russell Lakes Parker Pond Russell Lakes RLSWA Parker Pond Unit 18 1984 1984 1984 1985 1985 1985 1985 10 26 7 4 6 20 4 1 1 3 1 0 2 1 3 0 6 1 1 8 0 3 0 16 2 2 11 2 0 2 6 5 4 8 0 0 0 1 1 0 0 0 successful ~ nestlings / successful nest 2.3 3.0 2.9 3.4 3.4 2.9 3.0 nest and nest ~ nes tlin gs / nest attempt O. 9 0.3 2.4 2.4 1.9 1.7 1.5 .Pi-' -...J �418 resulted in 0.3 nestlings Inest nest success (78%) and the nest attempt (2.4) . Unit 18 had the greatest attempt. greatest mean number of nestlings Table 12. Distribution of clutch size of white-faced San Luis Valley, Colorado, 1984-85. li per ibises in the eggs Heronry Year 2 3 4 5 N x Mode Head Lake Russell Lakes Parker Pond Totals 1984 1984 1984 1984 1 0 0 1 8 6 ·2 16 9 4 5 18 2 0 0 2 20 10 7 37 3.6 3.4 3.7 3.6 4 3 4 4 Head Lake RLSWA Parker Pond Unit 18 Totals 1985 1985 1985 1985 1985 0 0 1 3 4 1 11 1 10 23 2 6 7 41 56 1 0 5 3 9 4 17 14 57 92 4.0 3.4 4.1 3.8 3.8 4 3 4 4 4 Nest Site Description Heronries Head Lake, were vegetatively Harrence only one emergent Ciconiiformes stands. Although stem bulrush Pond nests and Lake heronries bulrush. Thus, at Unit 18 were in hardstem were in common cattail stands. all were in hardstem all Ciconiiformes night-heron 7-11). and RLSWA contained Pond and Unit 18 contained and common cattail, 85% of those in 1985 (Figs. Lakes), hardstem at these Unit 18, 2 black-crowned nests (Russell species, Parker mapped and bulrush both nests hard- at Parker bulrush. 14 white-faced ibis At �Table 13. Distribution attempt for white-faced of brood size and mean number of nestlings per successful ibises in the San Luis Valley, Colorado, 1984-85. N nestlings x nestlings! successful nest nest and per x nestlin gs I attempt Heronry Year 0 1 2 3 4 Head Lake Russell Lakes Parker Pond 1984 1984 1984 3 11 2 0 0 0 2 2 2 6 0 2 0 0 0 2.8 2.0 2.5 2.0 0.3 1.7 Head Lake RLSWA Parker Pond Unit 18 1985 1985 1985 1985 5 5 6 14 0 0 0 0 2 2 10 0 6 2 25 0 1 1 15 --- 0.0 1.9 1.3 2.4 O· 2.9 2.8 3.1 nest nest ~ I-' 1.0 �420 HEAD LAKE - N I, L 50 m ~ SCIRPUS • Fig. 7. in 1985. Ciconiiformes nest locations NEST SITES in the Head Lake heronry �421 HARRENCE LAKE N 1 100 m ~ SCIRPUS • NEST SITES Fig. 8. Ciconiiformes nest locations in the Harrence Lakes) heronry in 1985. Lake (Russell �422 RUSSELL LAKES SWA N 1 100 m ~ SCIRPUS D UPLAND • Fig. 9. Ciconiiformes nest locations heronry in 1985. Upland vegetation inland salt grass. NEST SITES in the Russell Lakes SWA included rabbitbrush and �423 PARKER POND MVNWR N e ~ e~fJ .~ €) 1 100 m ~ ~ SCIRPUS ~ TYPHA • Fig. 10. heronry, NEST SITES Ciconiiformes 1985. nest locations in the Parker Pond �424 UNIT 18 MVNWR e (J~~@ N t; s~ 1 C)~. @~~;;;T' @~ 100 m ~ SCIRPUS ~ TYPHA • NEST SITES Fig. ll. Ciconiiformes nest locations in the Unit 18 heronry at Monte Vista National Wildlife Refuge in 1985. Nests were in Scirpus (85%) and Typha (15%). �425 Ciconiiformes nested in small, island-like, emergent stands emergent (Figs. Mean distances corresponding mean distances from nests (Table sites (Sign Test, 14). 2 from random were greater than to the nearest distance nests nest than These 15). to open water in the mean distance the ibis from were not significant from Ciconiiformes nests = 12.00, (Sign Test, outlier X2 ibis nests to the from random points The one exception One of the 5 nests which was less than was the mean to the nearest nest (61. 2 m) was 296 m from the nearest was excluded, the adjusted mean was 2.6 m, the mean distance from random points to the nest. Water depth sites than the and white-faced differences were less (Table from white-faced When this nearest sites the mean and median distances at Head Lake. nest. were less = 1. 33, P = 0.25). neighboring p < 0.005) the edge of large night-heron The mean and median distances nearest were eit her Head Lake was the exception; to open water. X or near to open water from black-crowned to open water random stands Nests 7-11). mean distances 4 of 5 heronries nests close to open water. (Table were slightly corresponding decreased was measured 16). at all Ciconiiformes and random Ten of the means and 7 of the medians greater (Sign Test, means and medians markedly nests X 2 = 1. 33, ~ at random = sites. 0.25) depth was measured than the Water depth at Head Lake and Unit 18 between in May and when water at nests in August. nest initiation �.j:;'- N (J\ (rn) from nests Table 14. Distances Valley, Colorado, 1985. to open water in 5 heronries in the San Luis Head Lake Russell Lakes RLSWA Parker Pond Unit 18 night-heron x SD Median Snowy N sites Heronry Species Category Black-crowned N and random 16 5.2 3.8 4.7 13 4. 9 2.9 3.9 12 10.4 3. 9 9.4 17 6.2 3. 1 5.4 6 8.0 7.0 8.1 0 14 21. 7 13.4 21. 8 11 4.0 3.4 2.8 15 4.6 3.1 3.4 32 6.7 2. 3 54 5.6 2.1 26 16.5 11. 5 44 5.3 3.5 34 5.3 3.1 25 5.0 2.4 5.0 40 6.9 4.3 6.0 15 23.3 24.4 14.0 21 15.1 11. 6 13.7 14 11. 6 9.7 7.7 27 6.8 2.4 6.6 41 5.8 2.3 5.1 0 13 5.4 1.5 4.7 5 6.0 1.6 6.5 0 egret x' SD Median White-faced N x SD Median All species N x SD Random N :x SD Median ibis �(m) from nests and random Table 15. Distances in the San Luis Valley, Colorado, 1985. SD Median Snowy egret N x- Head Lake SD Median All species N x SD Random N x SD Median Ciconiiformes nest in 5 heronries Russell Lakes RLSWA Parker Pond Unit 18 27 4.7 4.2 3.7 41 4.8 5.3 3.5 0 16 6.3 5.0 5.6 14 7.1 8.2 4.7 0 13 1.7 1.0 1.5 12 2.0 0.7 1.9 17 2.6 2.0 1.8 6 1.5 0.7 1.4 5 61. 2 131.2 2.7 0 14 10.4 8.0 8.5 10 5.1 6.3 2.3 15 9.2 10.6 4.4 SD Median x nearest night-heron x White-faced N to the Heronry Species Category Black-crowned N sites ibis 32 13.55 51.7 54 4. 1 4.8 26 6.5 7.2 43 4.5 4.7 35 6.2 7.6 25 16.3 18.2 8.7 40 21. 8 17.0 18.8 15 127.3 99.0 139.0 21 124.4 122.0 109.3 14 29.8 34.5 15.3 +'N ......• �~ N 00 Table 16. Water Colorado, August depth 1985. and random sites Head Lake 27 21. 6 2.6 20.3 SD Median Russell Lakes RLSWA 41 21. 0 6.9 20.3 0 0 SD Median SD Median All species N :x SD Random N x SD Median Valley, Parker Pond Unit 18 14 10.2 6.6 12.7 12 39.8 6.0 41. 9 17 38.8 8.2 38.1 6 11. 9 3.1 10.2 , x :x San Luis 16 40.3 10.6 43.2 egret White-faced N in the night-heron x Snowy N nest Heronry Species Category Black-crowned N (cm) at Ciconiiformes 13 25.4 6. 1 25.4 ibis 5 24.4 5.8 20.3 0 14 43.5 6. 8 43.2 11 32.1 8.4 33.0 15 13.7 7.6 15.2 32 22.0 3.3 54 22.1 6.9 26 41. 8 6.6 44 37.7 3.8 35 11. 9 6.6 25 19.8 3.3 20.3 40 20.7 9.3 17.8 14 40.6 16.8 44.5 21 32.3 10.2 30.5 8 9.9 8.9 11. 4 �429 The height varied (Table of Ciconiiformes 17). Water levels and Unit 18 and comparisons Black-crowned than nests decreased markedly nested and white-faced closer surface at Head Lake can only be made within night-herons snowy egrets above the water's heronries. to the water surface ibises. Fl ushin g Distance The mean flushing distance of snowy egrets approached either a vehicle or a person on foot was 96 m (N Range = 27- 200 m) in 1984. Flushing when the opportunity year. No attempt vehicles arose while conducting was made to separate or persons areas separated outside into difference vehicle flushing !. test, (l-tailed (Table 18). when approached Disturbances groups 46 m, field work that types Into were approached Disturbance types on foot. on foot than not statistically on were There was no distances when approached different, on foot was approximately in a vehicle. The small sample size measured outside was due to the difficulty of the heronries to a heronry passed during were observed on foot passed on a dike separated in a the mean from a person of 2 to 16 persons Disturbances other P > O.5) in the mean flushing Although black- crowned ni ght- herons birds = were measured disturbance or 1 person when approached distance approaching species of heronries. 1 motor vehicle for all 3 species 19, SD on foot. In 1985, all 3 Ciconiiformes feeding distances = by twice that for of locatin g an d daylight. when vehicles and by the RLSWA heronry. from the nearest nests by �.j:'- w o Height (cm) of Ciconiiformes Table 17. Luis Valley, Colorado, 1985. above Head Lake Russell SD 36 27.2 9.9 Snowy egret N 0 12 32.5 9.1 4 60.5 13.7 0 x x SD x SD water's surface in 5 heronries in the San Lakes RLSWA Parker Pond Uni t 18 night-heron 2 40.6 3.6 White-faced N the Heronry Species Category Black-crowned N nests 16 23.3 5.9 13 38.6 13.7 10 34.0 11. 4 14 27.4 15.2 6 62.7 22.6 8 41. 9 21. 6 11 65.4 21. 1 9 48.3 17.7 0 ibis �431 85 m of open water generally stood faced ibises that was 0.75 - 1. 50 m deep. and watched the disturbance and black-crowned night-herons the hardstem bulrush. person wading into the water began On 2 occasions, flushed when the person water. Disturbance Russell Lakes, Parker the disturbance agent had traveled on the shoreline Snowy egrets pass, remained most birds toward less while whitehidden flushed the heronry. than and Unit 18. entered the water when a The birds 0.5 m into the was also tolerated Pond, in Birds separating by birds at would flush if the shoreline from the heronry. Table 18. Flushing distance (m) of black-crowned night-herons, snowy egrets, and white-faced ibises approached in a vehicle or on foot, Russell Lakes SWA, Colorado. 1985. Species Disturbance N x flushing -distance SD Range Black- crowned ni ght- heron On foot In vehicle 4 2 152.5 46.5 66.1 26.2 75-210 28-65 Snowy egret On foot In vehicle 16 28 93.3 56.8 42.7 26.6 57-150 18-110 White-faced ibis On foot In vehicle 42 14 94.5 53.7 48.2 26.8 38- 205 18-105 �432 CHAPTER V DISCUSSION Census Black-crowned ibises have nested 1979) . night-herons, were children Although and landowners, the 3 species nesting now in their at Russell (E. Davey and K. Schrnittel, the presence of nesting Valley has been known, trends and white-faced in the San Luis Valley since 1875 (Ryder Local ranchers 60's,. can recall snowy egrets, nest 50's and Lakes since they pe rs , commun.). Ciconiiformes censuses et aL. in the San Luis and breeding population have not been well documented. The black-crowned night-heron nest census of 33 nests at Head Lake in 1985 can be compared to Gr aul ts (1980) estimate 20 black-crowned at the same site in 1978.· In contrast, Graul while I counted The overall night-heron (1980) counted indicate number of snowy egret at Russell Lakes in 1978, Nest searches nests counted from 135 in 1984 to 139 in 1985. the number to 1985, the number 1984. 81 nests only 52 in 1985. Luis Valley increased figures nests of of nests of nests increased slightly in the San While these from 1984 may have been underestimated and censuses were more thorough in in all �433 heronries in 1985 and snowy egret Unit 18 were counted. in both years, nests the 3 heronries When comparing the overall number at Adam' s Lake and of nests decreased censused from 135 to 99. The total number of snowy egret Pond, and Russell Lakes decreased Lake, the number of nests to 27 in 1984 and then number of nests but When examining it is more accurate heronries heronries nested egrets subcolonies Lakes, Pond, At Russell Lakes instead nests years, and RLSWA. of discrete In 1984, Ciconiiformes night-herons Lakes and white-faced night-herons between but not at RLSWA. with black-crowned of snowy egret populations at Russell close proximity. at Russell and 5 black-crowned number in nesting are probably divided At Parker from 35 in 1984 to 16 in 1985, to combine nests in one of the Russell nesting to 0 in 1985. 1980), from 0 to 13. changes due to their the heronry from 10 in 1978 (Graul declined increased Parker At Head from 73 in 1984 to 70 in 1985. of nests at RLSWA nests These decreased at Head Lake, from 1984 to 1985. increased decreased Lakes the number nests nesting and snowy ibises, snowy egrets, at RLSWA. in the Russell In 1985, Lakes The overall area decreased from 35 in 1984 to 29 in 1985. When making inferences extent of fidelity considered. different of breeding There heronries about nesting adults to specific is evidence that in different years. found dead from unknown causes population breeding trends, heronries adults the must be may nest in Two adult snowy egrets in the MVNWRheronry during were the �434 beginning banded of the 1985 nesting as nestlings (pers. commun.) Ciconiiformes breeding at Russell believes between population heronries Both birds Lakes over the interchange heronries trends egret nest ling egret banding nested was banded of young egret. plumage are indirect cattle was observed cattle egrets observed with cattle of cattle nesting egrets San Luis Valley increased before 1985. during 1 Sightings of adult egrets of nesting. in breeding cattle One adult Lakes during and cattle egret nests this indicate in 1985 and ibises record study. con- 1977. ibis nests ibises colony plumage were regularly at Russell whether the annual in counted from 46 in 1984 to 113 in 1985. It is not certain No written at least was recovered evidence of white-faced were more thorough at Unit 18. that Pond The bird in the San Luis Valley since The overall number searches evidence by W. D. Graul in a snowy egret in 1978 and adult Observations at Parker as a snowy egret and herons. as a cattle foraging schedule. at Russell Lakes in 1977. In 1977, a nest- in breeding egret estimatin g of all were counted (1979) reported and identified egrets 1978 and identified tinued Therefore, was found in the San Luis Valley Miller and Ryder pair of cattle egrets Ryder adult on a total census 1984 at Head Lake and 3 nests in 1985. earlier. of breeding is common. depend had been 4 years in the San Luis Valley on a regular One cattle during season. Nest were found nesting nested of Ciconiiformes in the at Unit 18 nesting in Unit 18 could be found in the MVNWRor Colorado Division of Wildlife records. Personnel at MVNWRthought it was possible herons, egrets, �435 and ibises had nested seeing nests during the breeding nests or adults counted no change in Unit 18 previously, but no one could recall flying in and out of the emergent season. When comparing in both years, excluding the number Unit 18, there were large changes in the number heronries decreased from 25 in 1984 to 5 in 1985 at Head Lake. ibis nests increased ov~r the 2 years. Lakes, respectively. Russell Lakes area has fluctuated to 9 in 1978 (Graul ber of ibis nests The number in number stayed In contrast, of ibis nests Pond and in the from 11 in 1976 (Graul has also fluctuated Black-crowned every night-heron United States (73%) was comparable ing success at Head Lake, able to that observed white-faced 19). success may in the San Luis in most other Nesting success success reported Unit 18, and Russell in Idaho ibises year. nesting to nesting while the overall and Productivity observed (Table heronries breeding Success Valley was lower than that The num- at Head Lake from 7 in 1976 within the same suggests Nesting 1977), 1980), 25 in 1984, and 5 in 1985. of nests not nest in the same heronry in of nests 1980), to 16 in 1984, and 22 in 1985. 1977), 0 in 1978 (Graul The changes The number from 7 to 18 and 16 to 22 at Parker Russell western was almost of ibis nests individual number of ibis (46 in 1984 and 45 in 1985). There (Graul stands (Findholt 1981). heronries in the at Parker elsewhere. Pond Nest- Lakes was comparLow nest success, �~ w (J\ Table 19. Nesting success State Year Georgia Idaho Idaho Idaho Washington Washington Washington Washington Oregon Oregon Oregon Oregon Nevada Nevada 1955 1979 1979 1979 1979 1979 1980 1979 1980 1979 1979 1980 1979 1980 of black-crowned N nests 8 103 63 118 ----- --- --- --- --------- --- night-herons Percent successful 87 87 41 47 79 83 80 78 0 90 67 68 0.4 67 in the United States. Age nest followed to (days) 15-21 15-21 15-21 14 14 14 14 14 14 14 14 14 14 Reference Teal 1965 Findholt 1981 II " Henny " "al. et 1984 " " " " " " " " " " " " " " " " " " " " " " " " " " " �437 such as observed and Nevada at RLSWA, has also been observed (Henny et al. Nesting success 1984). calculated with the Mayfield method was lower in the San Luis Valley than at 7 heronries (Custer et al. 1983). Although by Custer I calculated the Mayfield method so comparisons is questionable was violated because in several human disturbance, nests on the Atlantic .coas t In the San Luis Valley, from 0 to 49.1% while estimates from 53 to 86.7%. et al. in survival blue-winged cable to single estimate nest disasterous success rates such as flooding, of many Klett and Johnson for age-related (Anas platyrhynchos) but their events. using survival the failure of hours. for mallards teal (A. discors), success of constant caused in a matter rates (1983) ranged Single events, and predation ranged could be made, its application (1982) modified the Mayfield method to account variation success nesting the assumption heronries. within a heronry in Oregon and modification is not appli- The Mayfield method may not in colonial birds as accurately as traditional methods. The mean number of productivity and indication was the most productive averaging of nestlings 1. 9 nestlings per nest of nesting heronry surviving others The mean number Pond, Russell Lakes, is a measure Parker in 11 of 31 heronries of nestlings Pond night-herons to 10 days per nest was observed Head Lake, success. for black-crowned Higher productivity (Table 20). attempt attempt. studied per nest by at and Unit 18 was less than at Parker but was within the range observed in other heronries. Zero �po. w co Table 20. Productivity Location Russell Lakes Arapaho NWR Head Lake Lake John Annex Arapaho NWR Adams Lake Hutton Lake NWR Russell Lakes Latham Reservoir Minidoka NWR Blackfoot Reservoir Mud Lake WMA Moses Lake Three-mile Island Three-mile Island Foundation Island Summer Lake Ladd Marsh Malheur Lake Malheur Lake Ruby Lake Ruby Lake of black-crowned State Colorado Colorado Colorado Colorado Colorado Colorado Wyoming Colorado Colorado Idaho Idaho Idaho Washington Washington Washington Washington Oregon Oregon Oregon Oregon Nevada Nevada Alberta Alberta Maine Maine North Carolina North Carolina North Carolina Rhode Island Rhode Island night-herons Year 1978 1984 1979 1979 1979 1979 1979 1979 1979 1979 1979 1979 1979 1979 1980 1979 1980 1979 1979 1980 1979 1980 1964 1965 1975 1979 1975 1979 1979 1979 1979 N nests 13 --- 6 13 20 20 7 20 20 103 63 118 ----- ------------- ----- 67 55 71 127 29 93 15 165 31 in North N young fledged per attempt 0.23 1.5 0.0 1.9 2.0 1.7 1.4 1.0 0.7 2.5 1.1 1.1 2.2 2.5 2.2 2.0 0.0 2.6 1.7 1.8 O. 3 1.2 1.1 0.1 2.3 2.7 1.2 1.5 1.9 2. 4 2.1 America. Age (days) young survived 15 Reference Graul 1980 Eisemann 1985 McEwen et al. 1984 II 14 14 14 14 14 14 14 14 14 14 14 14 14 40 40 15 15 15 15 15 15 15 II II II II II II II II II II II II II Findholt 1981 II Henny II II et al , 1984 II II II II II II II II II II II II II II II Wolford II Custer and Boag II et al. II 1983 II II II II II II II II II II 1971 II �439 nestlings per nest attempt was observed well as at Head Lake by McEwen et al. Oregon nest by Henny et al. attempt 1979 (1.0) at Russell (1984). Black-crowned night-heron equal to other areas nestling productivity mortality in actual nest less than estimated daily mortality rate paring success nesting in separate nestlings studies Findholt maintain a stable in this nesting than observation nestlings that Only 8 heronries attempt per nest period (Table 20). per nest attempt was 2.7. The accuracy predicts per The unmeasured when com- per nest 40 days, an average attempt while other of 2.3 young night-heron female to studied by other attempt of fledglings this num- of nestlings 2.17 in all heronries Of these, must (1972) calculated The number with which the number the number at 30-40 of fledglings of nestlings while Henny 2.17 nestlings 1980). were 10-21 days of age. ing to 10 days per nest was less than greater and Wolford and Boag (1971) observed ber to be 2.17 to 2.25 young/pair. San Luis Valley. study. black-crowned population, (Graul must also be considered until they (1981) calculated fledge per successful and number at approximately nestlings study) United States. or mean number fledged per in the San Luis Valley is success For example, until they in 1985 (this from day 10 to fled gln g for nestlings studies. observed nesting as of nestlings 1984) than in 1978 (0.23) days results attempt The mean number of the western study (1984) and at Summer Lake, Lakes was greater (McEwen et al. Additional at RLSWA in this survive authors surviv- in the had the 14-15 day the greatest number of fledglings per nest per successful nesting of �440 black-crowned nestlings night-heron per nest fledglings female is unknown. attempt probably per female because if their first ever, nestling mortality additional would result underestimate success of ospreys ni ght- herons (Goering night-herons fails (Findholt between in unstudied disturbance are 1981).· (Parsons success heronries may consistently due to investigator had no effect and Burger 1971). had negative effects ni ght- herons (Trembley on reproductive 1982), and in a Texas In contrast, on reproductive investigator success heronry disturbance of black-crowned and Ellison 1979), gulls (Larus spp.) (Robert and Ralp h 1975, Fetterolf 1978, Ollas on and Dunnet Burger 1981), and double-crested cormorants (Ellison and Cleary Investigator success visits, disturbance Adults but they returned heat exhaustion nestlings Occasionally, regurgitated Considering mortality, black-crowned black-crowned eggs during No mortality nest from and young was night-heron and snowy egret which may have affected effects, night-heron on reproductive from the nest on exposed observer and Reed 1981). effect 30 minutes. food boluses survival. in the San Luis Valley. flushed within or predation had little 1980, (Phalacrocorax 1978, Des Granges probably in this study. observed. How- (Pandion haliaetus ( (Poole 1981), black-crowned and Cherry auritus) of day 10 and fledging of Ciconiiformes reproductive Investigator success attempt the number in overestimation. Estimates effects. underestimates black-crowned known to renest The number of renests, populations nestling and unmeasured may be declining �441 Clutch measured size is another measure for black-crowned within the range (Table 21). night-herons of clutch Clutch of productivity. than in the United the overall size in the San Luis Valley were observed studies (Table was 4, equal (Table Table 21). The modal clutch to or greater than that aiz'e in the San Luis Valley is sizes measured sizes larger Clutch States average clutch in only 3 of 16 other size for the San Luis Valley measured by other authors 21). 21. Clutch size of black-crowned America. State N nests Idaho Idaho Idaho Utah Alberta Alberta Maine North Carolina Maine Rhode Island Rhode Island North Carolina North Carolina Maine Maine North Carolina nesting 4 4 3 4 " " " " " " " 4 3 4 4 4 3 3 Custer " II II et al. 1983 " " " " " " " " " II II " " " II " " " " Custer " and Osborn " II than (37%), but less than that (1977) (84%). nesting success observed success observed " by Teal by Maxwell and Kale calculated using 1975 " " 11 in the San Luis Valley in 1984 (1965) Snowy egret 1981 Findholt Wolford and Boag 1971 success 1985 (60%) was greater in North Reference Mode 3.7 3.7 3.4 4. 1 4.0 3.2 3.8 3.1 3.7 3.7 4.0 3.4 3.5 3.7 3.7 3.1 103 63 118 41 67 49 68 26 124 154 26 81 14 25 25 25 Snowy egret (49%) and x night-herons the �442 Mayfield method ranged at Parker Pond in 1984. 4 heronries in Florida in this Idaho, study Florida, number Black et al. to be greater, The mean number attempt from 0.3% at Russell of nestlings of nestlings per nest Pond in 1984 and Russell nestlings other per nest authors. increased per nest stable Therefore, mortality, number at Head Lake in the San Luis Valley and in other mean nestlin gs 3.2 fledglings populations per to maintain observer areas Snowy egrets Lakes, female are necessary snowy egret by 1980), 0.3 in 1984, to that considering of studied attempt At Russell mean was 2.4 at Parker 1980) to 0.9 in 1984. (1984) calculated snowy egret study A higher from 0.8 in 1978 (Graul population. and unmeasured The greatest per nest at Head Lake in 1985. Findholt in Colorado, in 3 of 12 heronries The mean nestlings nesting reported in this Lakes in 1985. was observed fluctuated 2.4 in 1985. successful attempt for to 10 days per nest to that 22). success from 54 to 100%. surviving (Table from 0 in 1978 (Graul did not nest (1984) calculated ranging was comparable and New York Lakes in 1984 to 56.8% effects, a r enest s , may be declining of the western United States. The average Valley during measured clutch size of snowy egrets 1984 was greater in other snowy egret clutch areas clutch size measured than 15 of 19 average of the United States size averaged in other (Table 4.6 which is greater heronries (Table size was 4 in 1984 and 5 in 1985 compared 1969) and 3 in Georgia nesting (Teal 1965). 23). in the San Luis clutch 23). than sizes In 1985, any The modal clutch to 4 in Florida (Jenni �Table 22. Productivity of snowy egrets in the United X Location State' Adams Lake Head Lake Russell Lakes Minidoka NWR Mud Lake WMA Black Foot Res. Colorado Colorado Colorado Idaho Idaho Idaho Florida Florida Florida Florida California New York Salton Sea Long Island N nests 10 10 18 25 14 16 21 States. nestlings I nest attempt 0.0 0.0 0.8 2.0 2.0 2.5 3.1 1.6 2.3 1.8 2.9 1.5 Age (days) young survived 7-10 7-10 7-10 14 14 14 14 to Reference Graul 1980 II II II II Findholt 1984 II II II II 1984 Black et al. II II II II II II II II II Platter 1976 St. Claire Ray and Burger 1979 ~ ~ w �444 Table size of snowy egre~s in the United Clutch 23. N nests x Idaho Idaho Idaho Florida Florida Florida Florida Florida Georgia Florida California Maine Maine Virginia North Carolina North Carolina South Carolina South Carolina South Carolina Florida 9 30 85 24 11 10 19 102 29 77 3.7 4.3 3.7 4.1 4.0 3.4 3.4 3.9 3.2 2.9 4.0 4.3 3.6 3.2 2.9 3.2 3.6 3.3 3.0 2.8 25 25 25 12 25 25 25 16 60 White-faced and ibis nesting to be quite The mean number with heronry 4 3 (Graul 1977), At Russell 1978 (Graul Ryder then 1980), varied Ibis nest 0.3in 1.9 young/pair 1980), 11 11 II II 11 11 II II 11 II II II 11 II II 11 11 11 II II II II II II 11 11 II II II If 11 If II 11 11 and Taylor with heronry success Utah, 1979 in 1984 was also ranging there from 3.2 attempt were also varied 0.43 in 1976 2.0 in 1984, and 0 in 1985. 1.55 in 1976 (Graul 1985, and (1967) calculated II per nest At Head Lake, were 1984 11 1980). of ibis nestlings there II Jenni 1969 Teal 1965 Maxwell and Kale 1977' Platter 1976 Custer and Osborn 1975 in northern 0 in 1978 (Graul Lakes, II II Girard (Steele and year. 1984 II Black et al. success variable to 94.1% in 22 heronries year, Findholt 1985 in the San Luis Valley. reported Reference Mode State States. 1977),0.44 1.9 at RLSWAin that if all ibises would be sufficient breed in 1985. in their to maintain first a stable �445 population. Productivity at Head Lake in 1984 and at RLSWA and than or equal to 1. 9 young Inest Unit 18 in 1985 was greater attempt. The average Valley clutch size of white-faced ibises (3. 6 in 1984, 3. 8 in 1985 was greater 1970, Kaneko 1972, Steele found an average Steele (1980) found average 22 heronries clutch 1980). in Utah. were also greater Kotter in the San Luis than in Utah (1970) and Kaneko size of 3.2 in a Utah heronry, clutch size ranged The average than Graul (Kotter clutch (1972) while from 2.7 to 3.5 in sizes in 1984 and 1985 (1980) ( 3. 3) meas ured at Russell Lakes in 1976. Ciconiiformes, lands exhibit number which nest fluctuations of nests colonially in nesting within individual are a common cause of nest Ryder et al. 33%of black-crowned study) and all nests 1979, McEwen et al. of black-crowned and Hutton Lake, water levels failure unstable productivity, Fluctuating night-heron nests water noted nests Wyoming. Graul 1978, At Head Lake, were flooded in 1985 (this were flooded in 1979 (McEwen et al. (1984) also wet- and in the West (Alford 1980, McEwen et ale 1984). night-heron caused success, heronries. levels 1979, Graul in shallow, that flooding caused at Lake John Annex, (1980) observed complete abandonment 1984). losses Colorado severely of heronries In lowered at Adams Lake and Head Lake in 1978. Predators (Teal have caused nest losses 1965, Dusi and Dusi 1968, Jenni Boag 1971, Callahan and Carey in colonial nesting 1969, Kotter 1979, Findholt species 1970, Wolford and 1981, Henny et al. �446 1984, Jehl and Chase 1987). nest lin gs in 15 black-crowned nest at Russell of predation (Callahan and Carey State Univ.) night-heron in bulrush Russell (Corvus production 1984). ravens Lakes, Eisemann, nearly (Procyon and raccoons no evidence Oregon destroyed California Rep , , Colo. 1987). black-crowned in 1980 (Henny many egret (Platter were observed of predation heronries (Jehl and Chase eliminated lotor) at Salton Sea, in other 1985 Unpubl. gull colonies corax) and 1 snowy egret owls have been Ciconiiformes at Summer Lake, Raccoons marshes Although 1979; J. nests Great-horned on nesting and in nesting Common ravens et al. night-heron Lakes in 1985. suspected owls killed all of t he Great-horned nests 1976). at MVNWRand near was observed. Human Disturbance Recreation-related reproductive shorebirds gulls success correlated adeliae) colonies turbed Kushlan Findholt (Thomson 1977). (Ream 1976), Ciconiiformes flying over 1982), and have been in wading shorebirds (Butorides and Adelie penguin to aircraft 1979, Black et al. (Pygoscelis may not be dis(Grubb 1978, 1984, Vos 1984), but recreation-related can have serious 1981, Vos 1984). 1984), herons with lowered (Greer declines activities green-backed and Fritzell by and habituate disturbance population in recreational 1980, DeRoos 1981), (Kaiser (Gavia immer) and Flood 1980), mallards In addition, to increases striatus) has been correlated in common loons (Robertson (Hand 1980). (Erwin disturbance effects (Anderson The activities 1978, Dusi 1978, of fishermen and �447 bird-watchers 1978, Findholt can cause complete abandonment the snowy egret of 2 photographers nests, number Lakes in 1984. caused the failure of 90%of 85%of the white-faced ibis nests, .an d an of black-crowned Why Ciconiiformes 120 minutes within heronries not the photographers ment through (DUsi 1981). The activities uncounted of heronries night-heron tolerated the colony is better at Russell my presence while counting is not known. nests eggs and young, It is possible tolerated for over that but slow move- than intense, prolonged disturbance. Ciconiiformes in heronries on foot or in vehicles heronry Russell disturbed (Schultz approach herons, by at least were less disturbed when the observers 85 m of water. Lakes support by observers snowy egrets, not statistically measured observers at on foot A motor vehicle could on foot to black-crowned and white-faced significant than 1984). twice as close as a person distances from the that wild animals are less in motor vehicles and Bailey 1978, Profera were separated Flushing the observation by observers ibises. night- The difference due to large individual variation was and ~.. small sample sizes. Monte Vista National Wildlife Refuge Heronries at MVNWRcontained any of the other heronries had 44%of all snowy egret Luis Valley during 1984. greater numbers in the San Luis Valley. and white-faced ibis nests of nests Parker than Pond in the San In 1985, 64%of all colonial-nesting �448 Ciconiiformes Unit 18. in the San. Luis Valley nested Parker as any other Pond contained heronry Ciconiiformes outside nests ful and productive mented in this study nesting success 3 times as many nests nesting success were at Parker success heronries. The highest The highest and productivity and productivity more. success- and productivity Pond. Pond in 1984 while the highest nesting Pond and MVNWR. than in most other night-heron Parker more than on MVNWRwere consistently black-crowned snowy egret at Parker in this overall docu- overall study white-faced were at ibis were at the Unit 18 heronry in 1985. Heronries to flooding, managed at MVNWRdid not experience human disturbance, the portion heronries Few nests prey past nest surveys during at MVNWRthan nesting and refuge great-horned at the other 2 mainte- few days. owls, black-billed A possible waterfowl the spring at these on a dike every although were present. were more ground species) access (Mephitis mephitis) ~ raccoons, and ravens were Pond and Unit 18 were within the heronry were lost to predators, (Pica pica), there during due Pond and Unit 18 from The only human disturbance driving skunks nest losses Water levels Parker closed to public was my presence nance workers that Parker of the Refuge and summer months. striped or predation. at MVNWRwhich prevented flooding or becoming dry. large magpies explanation and rodents heronries is (buffer in the San Luis Valley. Clear water contributed and periodic to increased draw down of wetlands nesting success may have also and productivity through �449 increased prey contained dense water populations aquatic plankton, and availability. plant and clearer RLSWA, and Head Lake. probably ians. supported Periodic nique plant plants prey water plant of fresh- and plankton populations of fish and amphib- soil and water disturbance. Lakes, a common management populations. of prey populations populations but were controlled the availability at MVNWR when comp ared to Russell anaerobic and plankton Lakes, increase prey greater large draw down of wetlands, by causing Russell water The dense at MVNWR, prevents inhibit growth, Wetlands Carp Carp conditions also inhibit that aquatic were common at at MVNWR. to herons tech- Clear water and egrets may which locate by sight. Nest Sites Ciconiiformes species of emergents 1977, Alford bulrush though in hardstem (Giles and Marshall 1978, Bray common cattail 1984). bulrush over other 1954, Burger and Miller In the San Luis Valley, Ciconiiformes bulrush predators. or shrub-covered islands hardstem night-heron nested may offer nests in shrubs, Nests in bulrush land by open water (Mustela spp.), ibis were in common cattail. trees, more protection even were available. located in 1984 and 1985, only 14 white-faced 2 black-crowned weasels to nest was the most common n es t+s uppor-ttn g substrate 679 nests dry prefer or on islands. Of and No Hardstem from both mammalian and avian were over water which would discourage and coyotes and separated raccoons, (Canis latrans). from skunks, The dense �450 cover of hardstem bulrush may also prevent nest detection by avian predators. Although Alford common cattail (1978) thought support. offers hardstem bulrush He found white-faced significantly higher saturated, selected provided ibis nests above the water's to flooding were significantly found ibis pairs similar protection surface began nest bulrush and nest losses than cattail. sites in bulrush at which time ibises superior in hardstem less in bulrush nest from predators, stands selecting nest to be due He also until they were sites in cattail stands. Maps of heronries adjacent noted to open water were preferred. that or near in the San Luis Valley suggest Ciconiiformes interior nested openings distance different from Ciconiiformes than on the periphery nests the mean distance the mean distance the 5 heronries. in a small bulrush of random points from nests from random points may explain stand Therefore, mize the distance water. (1954) stands Wildlife the median to open water. to open water was less to open water in 4 of but the difference. in the outlet Ciconiiformes sites to open water was not signficantly was only 70 m wide and the bulrush wide. at the Stillwater Head Lake was the exception, size of this heronry of bulrush In the San Luis Valley, from the median distance In contrast, Giles and Marshall within stands Management Area in Nevada. that nest The heronry of Head Lake. stand and was The channel was approximately may have selected from dry land more than the location nest the proximity sites 5 m to maxi- of open �451 Although mean distances random sites, a parametric for significance because Colonial-nesting birds other birds selected of other of predators to nest near and catch adults open water. test the assumption within the colony. Visibility access statistical select nest independently advantage to open water to the heronry based Therefore, approaching open water. advantage flying selves on the nest. difficult Ciconiiformes nests. a bird Valley (61.2, Argentina egret 10.4, Nevada meters of When approaching a nest of bulrush and then interior In contrast, stands must hover slowly lower them- nests appeared to be nest ibis inter-nest 5.1, sites close to other distance than within the San Luis 9.2 m) was greater than observed (1977). in the San Luis Valley Girard and Taylor Ciconiiformes Likewise, 0.7, 2.0, (1954) reported in snowy 2.6, in (0.91 m). Water depth period several more energy. distance 1. 5 m) was greater may be one on the nest. (1.2 m) by Miller and Burger inter-nest are not could glide in. and do an abrupt over the nest selected White-faced of could not surprise had to cross birds. Approaching and required sites Predators to a nest in the interior from 1 to 2 m directly nest the heronry stall in the last meter and land gently adults on the proximity may be that open water improves for nesting on the edge of open water, was violated. sites. on the nest if they Another could not be used to test of independence sites nest were less for nest -t han at nest and between nest sites varied sites. considerably At some nest during the nest sites it was only a few �452 centimeters presence deep and at others of some water flood the nest, nested the nest, was more important Nest height bulrush, under species, was also variable and water ibis nests 1 m deep. depth. depending on the height Black-crowned and white-faced why 33%of black-crowned were flooded at Head Lake. The but not so much as to than depth. lower than snowy egrets may explain it was over night-heron of night-herons ibises, nests, which but no �453 CHAPTER VI RECOMMENDATIONS 1. a year Heronries to monitor breeding in each heronry of Mayor be counted populations it causes after 10 days, 2. should over measuring and unknown this time. this time than 3. 15 August constructing during Public access should success should the nestling to heronry be by guided be censused A breeding success estimates mortality In are uncer- of nestlings of observers. containing 15 May. nests, will desert of nests among heronries. unmeasured effects once the last week reproductive uses of wetlands Ciconiiformes during and is less expensive. be allowed from 1 April through nest sites, The number method of monitoring renests, No recreational selecting these adults of reproductive tain due to unidentified be censused All heronries less disturbance the reliability trends. on 1 trip week of June. is the preferred Ciconiiformes addition, population for movement of breeding census because should the first to account nest in the San Luis Valley should Ciconiiformes and laying nests heronries eggs during (Vos 1984). wetlands from 16 May through tour or in motor vehicles. methods of public viewing restrict during much more readily stages are visitor disturbance Both of and �454 access. closer There is evidence distances 4. that motor vehicles than persons Water levels 15 August to prevent at a constant forage ically to 'recharge' Draw downs decrease conditions 5. aquatic prey and increase plant production. to visually densities 6. 'island' orientated will result Hardstem stands should be drawn down period- the productivity Carp should be controlled bulrush to increase bulrush is preferred support characteristics. water clarity enhances predators. Enhanced should be encouraged openings to other soil and the visibility aquatic of plant available. (from 20 to 100 m in diameter) 0.5 ha should include anaerobic of wetlands. Clear water in more prey depth flooding or abandonment. Wetlands where Ciconiiformes the soil. at- on foot. should be maintained from 1 April through are tolerated at least emergents to grow in small and stands 20 m in diameter. because larger than Hardstem of its superior �455 LITERATURE CITED Alford, E. H. 1978. to habitat: an adaptive Young Univ., Allen, R. P. birds. S. Winckler, Aney, Utah. M.S. Thesis, Brigham 42 pp. appeal Protection for information and management on the wacling eds. of wading IV, J. C. Ogden, 99-103 in A. Sprunt, Wacling birds. and C. D. Cowan. important Archibald, Natl. recreational 1975. resource. G. W., S. D. H. Lantis, 1980. Endangered tivity. IntI. L. M. ibises: Daniel, resource Rep. Audubon and Soc. Res. 1979. values. Oregon Wildl. L. R. Lantis, future value and sportsmen. Dep. 30(2): 8-9. and I. Muhet chi ka , in the wild and in cap- of wildlife: Pages and B. L. Driver, U.S. shows wildlife 20:6-17. The esthetic E. H. Zube, Survey Agric., perceptions 32- 34 in T. eds. Accessing Fo:r:-.Servo Gen. 1977. M.S. Survey Thesis, of nonconsurnptive Univ , Idaho, wildlife users Moscow. llO pp , of C. amenity Tech. RM-68. L. A. Idaho. their Zoo Yearbook the American public Belli, advantage. ibis in relation 7. W. W., Arthur, by white-faced Field Notes ll: 458-460. 1978. Pages nesting An urgent Audubon J. M. Rep. Provo, 1957. birds. Anderson, Early in �456 Black, B. B., P. M. W. Collopy, G. Bohall. flights Fish 1984. on wading H. F. Percival, Effects bird and Wildl. Res. A. A. Tiller, of low level military Unit, Univ. training Florida colonies in Florida. Tech. Florida. and Coop. Rep. 7. 190 pp , Boyle, S. A., and F. recreation: an annotated actions. Rep. Bray, U.S. 252. M. P. habitat Burger, J. 1984. 1981. Fish An evaluation effects Effects outdoor of human wildlife inter- and Wildl. Servo Spec. gulls. and other and egret of burning. Colonial Waterbirds 1977. and glossy R~, and W. Carey. D. E., of herons Murrelet of human disturbance and L. M. Miller. of preying Capen, bibliography Inter., and possible in white-faced Callahan, Dep. Nonconsumptive Sci. 113 pp , particularly ---- , B. Samson. 4: 28-36. site selection Auk 94: 664-676. 1979. upon snowy egret. 65: 57-59. on colonial species, Colony and nest ibises. marsh Great-horned Florida owl suspected Field-Nat. 7: 29. and T. J. Leiker. tissues of white-faced ibis. Environ. G. Osborn. 1975. Wading birds 1979. DDE residues in blood Pollut. 19: 163-171. Custer, T. W., and T. logical indicators: and Wildl. Servo ---- , G. L. Spec. Hensler, reproductive Atlantic 1975 colony survey. coast success, Sci. Rep. Wildl. and T. E. Kaiser. and organochlorine black-crowned night-herons. U.S. Dep. 206. as bioInter., Fish 28 pp. 1983. Clutch contaminants size, in Auk 100: 699-710. �457 DeRoos, G. T. wader 1981. species Nature Conserv. J. L., of selected and A. Reed. Agric. 1981. Disturbance colonies of double-crested Colonial Waterbirds D'usi , J. L. sites. Univ., Wagen- 1978. Proc. to nesting Stability L. N., of heron Colonial Waterbird failure in Quebec. 1968. colonies in swamp and upland Group Con£. pp. Ecological factors in a heronry. and L. Cleary. ances on breeding cormorants and control 4: 12-19. , and R. T. Dusi. Ellison, Dep., The Netherlands. DesGranges, ---- upon some breeding on the isle of Vlieland in the Netherland's Wadden Sea. ingen, The impact of tourism Wilson Bull. 1978. Effects of double-crested 38-40. contributing 80: 458- 466. of human disturb- cormorants. Auk 95: 510-517. Erwin, R. M. 1980. waterbirds regimes Breeding in two mid-Atlantic in colonial waterbirds: use by colonially nesting U. S. regions of human disturbance. , and T. W. Custer. ---- habitat under BioI. Conserv. 1982. Estimating an evaluation. different 18: 39-51. reproductive success Colonial Waterbirds 5:49-56. Fazio, J. R., and L. A. Belli. sumptive wildlife users and Nat. Resour. Fetterolf, P. M. response Con£. pp. 1978. in Idaho. Characteristics Trans. of noncon- North Am. WildI. Con£. 42: 117-128. The human artifactor: to the scientist. 48. 1977. Proc. gull behavior Colonial Waterbird in Group �458 Findholt, S. L. 1981. and reproductive Thesis, Idaho 1984. Organochlorine success State in black-crowned Univ , , Pocatello. Organochlorine reproductive pollutants, success eggshell quality night-herons. M.S. 40 pp. residues, eggshell of snowy egrets nesting thickness, in Idaho. and Condor 86: 163-169. Giles, L. W., and D. B. Marshall. colony on the Stillwater 1954. Alar Wildlife Management ge heron Area, and egret Nevada. Auk 71: 322-325. Girard, G. T., for 9 avian Florida. Goering, species Florida D. K., Texas Graul, and W. K. Taylor. W. at Moore Creek, Sci. 1977. Wilson Bull. Nesting Colorado-1976. Reproductive Merritt Island parameters N. W.R. , 42: 94-102. and R. Cherry. heronry. 1979. success Unpubl. 1971. Nestling mortality in a 83: 303-305. of the white-faced Rep • , Colorado ibis in Div , Wildl., Denver. 5 pp , 1980. species Jan. Gray, in Colorado. 1980. G. C. municipal Thesis, Greer, Population D. M. Part 1975. surveys Colorado Div. Wildl., bird and mammal Wildl. Res. Rep. 1:98-123. Nonconsumptive conservation Urban tion of management demand commissioners Univ , Massachusetts, 1982. of selected in Massachusetts. Amherst. waterfowl and planning. for wildlife by M. S. 94 pp. population: Environ. ecological Manage. evalua- 6: 217-229. �459 Grubb, 1978. M. M. herons Effects and egrets. of increased Proc. noise levels on nesting Colonial Waterbird Group Con£. pp. 49-54. Hand, J. 1980. L. occidentalis specific Henny, livens) predation. selected Blus, G. L., results. Jr., 1. west. nesting Krynitsky, Bur. Wilson Bull. J. R., and prey 1981. a model, Sport Fish. of and 1984. night-herons in the 48: 1-13. The Mayfield method of estimators and simulation 93:42-53. and C. Chase, selection dynamics and C. M. Bunck. J. Wildl. Manage. success: by intra- 99 pp. and J. D. Nichols. estimating Jehl, A. J. amplification of the population Dep. Inter., Rep. gull (Larus 18:59-63. impact of DDE on black-crowned intermountain Hensler, U.S. Wildl. Res. Current colonies and possible An analysis species. , L. J. ---- in western Biol , Conserve 1972. C. J. Wildl., Human disturbance 1987. III. by avian predators: two colonies of California gulls. Foraging patterns a comparative Studies study in Avian BioI. 10: 91- 101. Jenni, D. A. herons County, Johnson, 1969. during A study Florida. D. H. of the ecology of four species the breeding 1979. Ecol. Monogr. Estimating M. S., and E. K. Fritzel1. tionists on green-backed 48: 561-567. of at Lake Alice, Alachua 39: 245-269. nest success: the Mayfield Auk 96: 651-661. method and an alternative. Kaiser, season 1984. Effects heron behavior. in of river recrea- J. Wildl. Manage. �460 Kaneko, K. D. chihi) Nesting of Utah Lake. Provo, Klett, 1972. Utah. A. T., of the white-faced M.S. Thesis, Brigham (Plegadis Young Univ., 76 pp. and D. H. Johnson. survival ibis rates 1982. and implications Variability to nesting in nest studies. Auk 99: 77-87. Kotter, B. S. ibis (Plegadis Utah, Kushlan, A. chihi) J. G. R., of five species H. F. Utah. M.S. of helicopter Wildl. Manage. Thesis, Univ. of herons in coastal Nesting 73: 255-261. 1975. Suggestions censuses on wadin g 43: 756-760. and H. W. Kale, II. 1961. of the white-faced 126 pp. Wilson Bull. Bull. life history in northern 1979. Effects colonies. Mazwell, II, Mayfield, An ecological Salt Lake City. J. bird 1970. 1977. Breeding Florida. success Auk 94: 689-700. calculated for calculating biology nest from exposure. success. Wilson 87: 456-466. McEwen, L. C., C. J. Organochlorine from Colorado Stafford, residues and G. L. Hensler. in eggs and Wyoming. 1984. of black-crowned Environ. Toxic. night-herons and Chern. 3: 367-376. Miller, G. C., Western Miller, and R. A. Ryder. Birds Cattle egret in Colorado. 10: 37- 41. H. W., and D. H. Johnson. of nesting 1979. studies. 1978. J. Wildl. Manage. Interpreting 42: 471-476. the results �461 Miller, L. M., success More, and J. Burger. of the glossy T. A. users: Gen. H. S. Pages ibis. Factors 1980. Rep. U.S. NE-52. mapping 277- 290 in S. D. S chemnitz, 4th ed. Dep. wildlife Agric., For. 16 pp. Reconnaissance manual. nesting Auk 95: 353-361. of the literature. Tech. techniques affecting The demand for non consumptive a review Servo Mosby, 1979. 1978. ed. and map use. Wildlife management The Wildl. Soc , , Washington, D.C. J. C., Ollason, fulmar: Parsons, and G. M. Dunnet. the effect nestling behavior Nest failures in the J. Field Ornith. of observers. and J. Burger. K. C., 1980. 1982. 51: 39-54. Human disturbance in black-crowned night-herons. Breeding of cattle and Condor 84: 184-187. Platter, M. F. egrets 1976. at the Salton San Diego State Poole, A. 1981. J. tours, Res. Ream, success. 1984. North Rep. C. H. pesticide Univ., The effects reproductive Profera, Sea, ecology southern San Diego, California. Calif. Colonial Waterbirds of sage Colorado. grouse Colorado and snowy M. S. Thesis, 122 pp. of human disturbance Development Park, egrets on osprey 4: 20-27. public viewing Div. Wildl., Wildl. 2: 373- 386. 1976. residues 88: 427-432. Loon productivity, in northern human disturbance Minnesota. Wilson Bull. and �462 Robert, H. C., turbance and C. J. Ralph. on the breeding 1975. success Effects of gulls. of human disCondor 77: 495- 498. Robertson, R. J., and N. J. use of shorelines Flood. on breeding 1980. bird Effects of recreational populations. Can Field-Nat. 94: 131-138. Ryder, R. A. 1951. Colorado. M.S. Waterfowl production Thesis, in the San Luis Valley, Colorado State Univ., Fort Collins. 130 pp. 1967. faced ibis Distribution, (Plegadis migration chihi) in North and mortality America. of the white- Bird-Banding 38: 257- 277. 1978. Breeding snowy egrets IV, ---- in North J •. C. Ogden, Audubon Soc. Schultz, R. D., park Shaw, andJ. 1978. Proc. of Ciconiiformes Group West. Assoc. of 197-205 in A. Sprunt, eds. and G. C. Miller. Con£. A. Bailey. Witter, Nonhunting Pages and mortality Wading birds. Natl. 7. elk to human activity. W. W., D. J. movements, Winckler, Rep. and movements Colonial Waterbird America. and J. Res. , W. D. Graul, tion, distribution, Status, in Colorado. distribuProc. 3: 49-58. 1978. J. 1979. Responses Wildl. Manage. D. A. King, wildlife enthusiasts and M. T. of national 42: 91-100. Richards. and wildlife management. Fish and Wildl. Agencies 58:255-263. �463 St. Clair Raye, S., and J. of nestling success MidI. Nat. 102: 76-85. Steele, B. B. effects 1980. Steele, Reproductive Univ., R. G. D., and J. of statis tics: Co., New York, P. L. thula). Am. of the white-faced ibis: M. S. Thesis, 1980. Principles approach. and proced- McGraw- Hill Book 633pp. Natural free-tailed M.S. determinants 53 pp. a biometrical 1984. Colorado. (Leucophoyx success H. Torrie. N.Y. of the Brazilian Behavioral and colony characteristics. Logan. ures Svoboda, 1979. of snowy egrets of pesticides Utah State Burger. Thesis, history and geographic relationships bat in the San Luis Valley of Fort Hays State Univ , , Hays, Kans. 67 pp. Teal, J. M. 1965. Georgia. Thomson, Wilson Bull. R. B. penguin Co., Trembley, J. 1977. rookery Antarctic Thompson, Nesting success of egrets and herons in 77:257-263. Effects of human disturbance and measures of control. on an Adelle Proc. SCAR Symp. BioI. 3: 1177-1180. M. 1928. Pinehurst, J., The witchery N.C. The Archers 259 pp. and L. N. Ellison. ance on breeding of archery. 1979. of black-crowned Effects of human disturb- night-herons. Auk 96: 364- 369. Tyre, G. L., and G. A. James. participation U.S. Dep. in various Agric., For. 1971. activities Length and rate on recreation Servo Res. Note. sites SE-161. of individual and areas. 4 pp. �464 U.S. Department Gale Res. of Commerce. Co., 1985. 1978. Detroi t , Mich. Climates of the United States. 606 pp. Climatological data annual summary: Climatological data monthly Colorado 1984. 89: 1-29. 1986. summary: Colorado 1985. 90: 1-29. U.S. Department hunting Fish of Interior. and wildlife-associated and Wildl. Serv., Vos, D. K. 1984. Colorado State recreation. in northcentral J. W., and D. A. Boag. night-herons of fishing, Inter., 156 pp , great Colorado. Collins. survey U. S. Dep. D.C. of breeding Univ , , Fort of black-crowned 1980 National Washington, Responses human disturbance Wolford, 1982. blue herons M.S. to Thesis, 65 pp , 1971. Distribution in Alberta. and biology Can Field-Nat. 85: 13-19. Yeosting, D. R., childhood exploratory and D. L. Burkhead. recreation analysis. experience J. Leisure 1973. Significance on adult leisure Res. 5: 25- 36. of behavior: an �465 APPENDIX MARSH BIRD DISTURBANCE TRIALS �466 APPENDIX Marsh Bird Disturbance Trials A method of investigating ance on marsh reactions vehicle birds was tested. of marsh birds traveling April-July trials were conducted were counted immediately after During a disturbance, following a disturbance. person on foot for all trials 2 disturbance vehicle. Disturbance a slow, steady intervals agents were entered in 1984, a disturbance, of disturbance for was 1 a person count interval. on foot or a motor along a dike and returned were counted a disturbance trials 1985 was modified and a shortened traveled of marsh birds disturbance trials during to disturbance in 1985 (Table trial blinds which area. agents 24). at at 5-minute from elevated out of view of the disturbance Responses fied during trials Marsh birds for 1 hour after 1984 and in 1984. were either speed. July disturbance The agent agents of disturbance during and at IS-minute intervals The method for disturbance The agents the on foot or a motor immediately before 1 hour to include a person disturb- an area used by marsh birds. 1985 at RLSWA. marsh birds of recreation-related The method compared to either through Disturbance efforts were quanti- These values �467 were tested square for independence agents with a chi- test. Table 24. Quantification Response of marsh bird values 2 3 4 American Coots.--The counted (Table 25). (where they remained hidden open water of American coots a disturbance to pre-disturbance levels Coots swam out of sight could not be counted) during soon after Disturbance number immediately after to almost 0, but returned minutes to disturbance. No obvious response SUbjects watched disturbance Subjects swam away from disturbance Subjects "ran" on water or dove Subjects flew from wetland 1 americana) responses Response o stands from disturbance the disturbance the disturbance did not affect wetland in 1985 (Tables in 1984 decreased within 15 into hardstem bulrush when disturbed. and began They returning to disappeared. the number 25 and 26). (Fulica of coots counted The difference between on a 1984 and 1985 count data can be explained by coot behavior. coots over 20 m from the disturbance swam away and hid as they did in 1984. before Coots hidden the disturbance, intensity ance (X = 3.3, number alon g the trail into the open when disturbed. was independent 0.25 < ~ < 0.50) Redheads.--The counted flushed of coot responses 2 in hard stem bulrush In 1985, (Fig. of disturb- 12). of redheads on a wetland immediately after of the agent The (Aythya disturbances americana) decreased from �468 Table 25. Marsh birds counted on a wetland immediately before. immediately and at IS-min ute intervals for 1 hour after disturbance by a person walking. after. N present N counts Species Year American Before After disturbance (min. ) Post-disturbance 15 30 45 60 Coot 1984 6 Range Median x SO 1985 3-29 16 15.3 10.5 0-2 0 0.5 0.8 4-23 17.5 14.8 7.6 2-40 22.5 21.8 12.9 6-31 26.5 23.3 9.3 3-36 26.5 23.0 10.9 5-22 10.5 11.6 5.0 5-20 10.0 11.4 3.4 2-18 8.5 9.4 4.5 3-26 9.0 10.3 6.0 4-21 10.5 11.7 5.3 4-25 9.5 11.6 5.9 1-5 3.0 2.B 1.5 0-3 0.0 1.0 1.4 0-1 1.0 0.8 0.5 0-4 2.0 loB 1.5 0-6 1.0 2.4 2.9 0-5 1.0 2.0 2.0 2-17 7.0 7.5 4.4 0-14 6.0 6.6 4.4 o-n 6.0 5.5 4.0 0-13 3.0 4.B 4.7 0-13 6.0 6.4 4.7 0-16 3.0 4.8 4.9 1-12 2.0 4.4 4.0 0-2 1.0 1.0 1.0 2-12 3.0 4.7 3.7 2-10 3.0 4.4 3.0 0-7 4.0 3.7 2.5 0-10 2.0 2.7 3.4 1-4 1.0 1.8 1.5 0-1 0.0 0.3 0.5 0 0.0 0.0 0 0.0 0.0 0-3 0.5 1.0 1.4 0-6 0.0 1.5 3.0 1-11 9.0 7.5 4.7 0-2 0.5 0.8 1.0 0-3 1.0 1.3 1.3 0-3 0.5 1.0 1.4 0-3 1.0 1.3 1.5 0-4 1.0 1.S 1.7 1-5 2.0 2.3 1.1 0-2 0.0 0.6 0.9 0-2 0.0 0.3 0.7 0-1 0.0 0.4 0.5 0-2 0.0 O.B 1.0 0-2 0.0 0.6 0.7 1-10 5.0 5.3 3.0 0-6 2.0 2.7 2.5 O-B 2.0 2.7 3.0 O-B 1.0 2.2 3.1 0-12 3.0 4.2 3.8 0-9 2.0 3.3 3.6 1-4 2.0 2.3 1.1 0-4 2.0 2.1 1.1 0-5 2.0 1.9 1.4 0-6 2.0 2.2 1.7 0-4 2.0 1.9 1.2 0-8 2.5 2.9 2.5 16 Range Median x SO Redhead 19B4 5 Range Median x SO 19B5 21 Range Median x SO Ruddy Duck 1984 7 Range Median x SO 1985 4 Range Median x SO Mallard 19B4 4 Range Median x SO 1985 9 Range Median x SO Gadwall 10 1985 Range Median x· SO American Avocet 1985 Range Median x SO 12 �469 Table 26. Marsh birds counted on a wetland immediately before, immediately after, and at 15-minute intervals for 1 hour after disturbance by a motor vehicle, 1985. ~ present Species counts American Coot Range ~edian x SD 14 Redhead Range Median 13 x SD Ruddy Duck Range Median x SD 2 Mallard Range Median 2 x SD Gadwall Range Median Before 15 30 45 60 4-18 9.0 9.1 3.5 3-17 9.0 9.5 3.9 4-15 8.0 8.6 3.7 3-16 10.0 9.1 3.9 3-19 8.0 8.5 4.4 3-18 8.0 8.6 4.7 2-12 7.0 7.2 3..6 0-14 7.0 7.0 5.5 0-11 7.0 6.1 4.0 0-12 4.0 4.9 4.4 0-12 3.0 4.9 4.5 0-12 4.0 4.5 4.2 1-3 2.0 2.0 1.4 0 0.0 0.0 0 0.0 0.0 0 0.0 0.0 0 0.0 0.0 0 0.0 0.0 1-13 7.0 7.0 8.5 1-6 3.5 3.5 3.5 0-6 3.0 3.0 4.2 2-5 3.5 3.5 2.1 2-4 3.0 3.0 1.4 3-4 3.5 3.5 0.7 1-8 3.5 3.4 2.0 0-9 2.0 2.1 2.8 0-3 1.0 1.1 1.2 0-5 1.0 1.5 1.8 0-6 3.0 2.9 2.4 0-7 2.0 2.4 2.6 1-5 3.0 3.1 1.5 1-4 2.0 2.4 1.3 0-5 2.0 2.4 1.7 0-6 2.0 2.1 2.2 0-5 3.0 2.6 1.7 0-14 3.0 4.3 4.7 10 x SD American Avocet Range Median x SD (min. ) Post -disturbance After dist urbance N 7 �470 A. 68% 13 12 11 10 9 lilill 8 :::::::: ........ ........ ........ 0 7 ~~~~~~~~ ........ CJ) W 6 CJ) W CJ) 2 c, a: 5 21 4 ........ :::=:=:: 26% :.:.:.:. {{ ........ 3 :::::::: :::::::: :::::::: ........ 2 1:1111 ........ .;.:.:-: ........ :-:.:.:. :::::::: :::::::: ........ 5% rr I~j~j 1 :::::::: 0 1 0 2 3 4 8. 13 12 11 10 CJ) 9 CJ) 8 w 2 0 7 CJ) W 6 a: 5 21 4 o, 53% 40% 1 ::.:.:.:. ~: ~: ~: ~ -:.:-:.: ~:~:~:~: 'i'i';'i ~:~:!:~: ........ ::;::::: .:.:.:.: iI~i~ 3 2 ........ .:.:.:-: ........ :.:.:-:. ........ :::::::: 7% ::::': 0 1 0 :=:::::: ~~~~~~~~ .:.:.:.: :-:.;.:. :::::::: :::::::: ........ .:-:.:.: :.:.:.: ......... 2 3 4 RESPONSE VALUE Fig. 12. Frequency of American coot responses foot and (B) a motor vehicle during disturbance (X2 3.3, 0.25 < P < O. 50) • = to (A) a person trials in 1985 on �471 predisturbance heads counts 25). During flew from the area or hid in hardstem trials, redheads they did not return could be counted) the number on a wetland redheads had no affect on the number were independent <!: < 0.50) (Fig. counted pre-disturbance ducks stem bulrush of the agent number they during 1985, the number of ruddy 1 case turbance counted values (X of 2 = 2.15, (Oxyura a disturbance the hour after ducks decreased When disturbed, resurfaced ruddy in the hard- They began the disturbance. decreased after to pre-disturbance from return- During the disturbance levels, except in in 25 and 26). , Mallards. --Mallards the wetland after ducks and probably and did not return (Tables of redheads could not be counted. ing to open water all trials by disturbance. of disturbance in 1984 (Table 25). the water where In one trial, Response of ruddy on wetlands counts dove under 25 and 26). (where 13). Ruddy Ducks.--The jamaicensis) was unaffected red- In these activity the following hour. counted in 1985 (Tables 4 trials, bulrush. to predisturbance during of redheads Disturbance 0.25 in 1984 (Table responded and not returning in most trials n umber returned (Tables to disturbance during from the hour following the dis- 25 and 26). to pre-disturbance by flying The post-disturbance levels within 1 hour in 3 trials in 1985. Gadwalls. --The after a disturbance trials, disturbance mean number of gadwalls in most trials (Tables decreased 25 and 26). had no affect on the number immediately In 7 of 20 counted. In most �472 A. 11 109- CI) W CI) 8- 2 7- 0 o, 6- CI) W 5- a: 4- 21 3- 26% IIIIII 2- !!/II!J! 10 I o I I T J 1 2 3 4 8. 11 109CI) W CI) 2 87- 0 6- CI) W 54- ZI 3- n, a: 25% {f~ 2iiiil/lii 1- 0 I o I I 1 234 I r RESPONSE VALUE Fig. 13. Frequency of redhead and (B) a motor vehicle during 0.25< P< 0.50). responses disturbance to (A) a person on foot 2.15, trials in 1985 (X2 - = �473 trials, the number of gadwalls returned within 1 hour. the 2 agents There was no difference (X2 of disturbance American Avocets.--Neither on American avocets Response values ance (X2 = to pre-disturbance = 3.22, in gadwall responses to 0.50 < P < 0.75) 14). agent of disturbance (Recurvirostra americana) were low and did not depend 0.50 < P < 0.75) 1.18, (Fig. levels (Fig. had.an (Tables effect 25 and 26). on agents of disturb- 15). ConcI usions that Although sample sizes were small, disturbance different marsh bird turbances. several These hours apart to pre-disturbance consider groups responses disturbance species data indicate to visitor-induced 1 conducting desirable behavior present should be conducted the test) to dis- patterns and their to return tours should potential Further testing using elevated with blinds <20 m Use of 2 persons in evaluation indicated should be spaced for visitor disturbances. wetland. differentially disturbances Also, planning of avian species from the edge of the study blind, that to allow mars h bird levels. trials responded trials of disturbance (1 in the trials is �474 A. 10 9 8· CI) W CI) 50% 7 6 2 36% 5 0 0.. CI) W 4 a: 21 3 2 1 0 0 1 2 3 4 8. 10 9 CI) W CI) 2 0 0.. CI) W a: 21 8 50% 7 6 5 29% 4 21% 3 2 1 0 0 1 2 3 4 RESPONSE VALUES Fig. 14. Frequency of gadwall responses to (A) a person on foot and (B) a motor vehicle during disturbance trials in 1985 (X2 3.22, 0.50<P <0.75). - = �475 A. 10- 987- CI) W CI) 2 6- a.. S- 0 CI) W 4- 21 32- .. a: 10 1 o I I I I 1 2 3 4· B. 10 CI) W 9 8 2 7 6 a.. 5 W 4 CI) 0 CI) a: ZI 3 2 1 0 70% :::;:;:: ........ :.:.:.:. :::::::: ....... ::;:;:: ::::::: ':.:':'. ::::::: .:.:.:. ....... :-:.:.:. .:.:.:.: ........ :-:.:.:. .:.;.;.: ........ 20% 10% :.:.:.:. ........ :~:~:~:~ 0 1 2 3 4 RESPONSE VALUES Fig. 15. Frequency of American avocet responses to (A) a person on foot and (B) a motor vehicle during disturbance trials in 1985 (X2 1.18, 0.50 < P < 0.75) . = �476 Prepared by .~ L S'~ Janet L. Schreur GB Graduate Research Assistant Approved by ~£ ~ Clait E. Braun Wildlife Research Leader �Colorado Division of Wildlife Wildlife Research Report April 1987 477 JOB PROGRESS REPORT State of Colorado --~~~----------------------------- Project 01-03-045 24 Work Plan Job Title: 1 --- Habitat Selection and Behavior of Whooping Cranes Period Covered: Author: : Job Avian Research 01 January through 31 December 1986 Tanya Shenk Personnel: C. Braun, J. Ringelman, Colorado Division of Wildlife; D. Hei.n, T. Shenk, Colorado State University ABSTRACT During fall 1985 2 whooping cranes (Grus americana) used wetlands near Severence and Hudson, Colorado for over a month. Despite frequent forays away from their roost wetlands, these birds consistently relied on 2 sites during this period. The habitat requirements and preferences of whooping cranes during the migratory period are not known. Although mechanisms of habitat selection 'in water birds are poorly understood, physical characteristics of the wetland environment are believed to be important. Because whooping cranes are beginning to use non-traditional stopover sites in eastern Colorado, it is important to quantify the characteristics of wetlands sought by cranes as an aid to understanding the habitat characteristics of wetlands used, how habitat vital to their needs can be managed or preserved, and if it is possible to predict which wetlands are likely to be used as future stopover sites. Selected biological and physical characteristics of the 2 ,non-traditional use sites and 20 similar surrounding non-used wetlands were measured. Statistical comparisons between use and non-used sites will be made. Similar wetland characteristics were measured for IS'traditionally used wetland sites in the San Luis Valley for comparison with the 2 non-traditional use sites. Behavioral data were collected on the whooping crane using the non-traditional site during the fall 1986 stopover period for comparison with behavior data collected in the San Luis Valley. ��479 HABITAT SELECTION AND BEHAVIOR OF WHOOPING CRANES Tanya Shenk P. N. OBJECTIVES The primary objectives of this study are to: (1) compare habitat at non-traditional use and non-used whooping crane stopover sites in eastern Colorado, (2) compare habitat at non-traditional use sites and traditional use sites in the San Luis Valley, (3) compare whooping crane behavior at traditional use sites to behavior at non-traditional use sites, and (4) develop standardized methodology for habitat evaluation of future whooping crane stopover sites. SEGMENT OBJECTIVES 1. Review available literature on whooping crane and sandhill crane habitat use and behavior. 2. Develop, submit for review, and finalize a study plan for examining habitat selection by whooping cranes on fall stopover sites in Colorado. 3. Quantify wetland characteristics (physical features, vegetation, water chemistry, water level regimes), invertebrates, and disturbance factors for used and non-used, non-traditional stopover sites. 4. Measure selected biological and physical characteristics of used traditional stopover sites for comparison with non-traditional sites. 5. Document behavior of whooping cranes at non-traditional stopover sites for comparison with behavior at traditional stopover sites. 6. Conduct statistical comparisons of habitat selection at used vs. unused sites. 7. Compare behavior of whooping cranes at traditional vs. non-traditional stopover sites. 8. Develop standardized methodology for habitat evaluation of future stopover sites and provide recommendations for management of whooping cranes during fall. 9. Prepare quarterly progress reports and submit a report by 30 June 1987. DESCRIPTION OF STUDY AREA The Severence and Hudson study sites are within Weld County in northeastern Colorado. Southern Weld County lies entirely within the Colorado Piedmont section of the Great Plains physiographic province. Topography is broadly rolling. Elevation ranges from about 1450 to 1700 m. The land is irrigated by a system of reservoirs and ditches. The irrigated farmland of the area �480 supports a wide variety of crops including corn, alfalfa, sugar beets, pinto beans, potatoes, and onions. The San Luis Valley in south-central Colorado is the major spring and fall migration stop for the Rocky Mountain population of greater sandhill cranes and cross-fostered whooping cranes (Drewien and Bizeau 1974, 1978, Kauffeld 1981). This population currently numbers 17,000-20,000 sandhill cranes and 30-35 whooping cranes (Brown et ale 1985). Cranes use the valley for 3-4 months annually, primarily from October through mid-November and mid-February through mid-April. Barley, wheat, and potato farming are the primary land uses in the valley. METHODS Literature Review A literature review was done and knowledgeable persons were consulted on: a. whooping crane habitat selection at stopover sites, including the location and description of other known stopover sites, b. whooping crane natural history and the foster parent program, c. migration routes of whooping cranes, d. water quality descriptions of wetlands used by whooping cranes, e. vegetation identification and measurement techniques, f. invertebrate identification and sampling techniques, g. fish identification and sampling techniques, and h. time budget models. The literature review was completed to the extent of completing the study plan for the 1986 fall field season. I have continued the literature review throughout the entire project and will continue to do so until the thesis is finished. Study Plan The study plan was completed following the above literature search. The initial study plan was reviewed by Drs. J. K. Ringelman, C. E. Braun, R. C. Drewien, and D. A. Hein, and W. M. Brown. The suggestions and revisions of the above mentioned persons were incorporated into the study plan and a final draft of the study plan was submitted and accepted by Dr. C. E. Braun. Quantification of Habitat Characteristics of Eastern Colorado Sites The criteria for habitat evaluation of all study sites were based on current literature and knowledge concerning habitat requirements of migrating whooping cranes. �481 The 2 non-traditional use sites in eastern Colorado were located with help of CDOW personnel (J. Dennis, T. Lynch, and D. Crawford) who were involved in monitoring whooping cranes during fall 1985. Selection of non-use sites in eastern Colorado was based on similarity of wetland hydrology. Non-use sites were selected from within a 3-km radius circle centered at the use site with the same wetland classification (Cowardin et al. 1979) as the 2 use sites. This classification is (system) palustrine, (class) emergent wetland, and (modifier) saturated/semipermanent/seasonal (PEMY). The following criteria were measured on all eastern Colorado study sites: a. Distances to power lines and fences were measured with an optical rangefinder or topographic maps. Measurments were made from the nearest power line and fence to the wetland edge. b. Distance to nearest feeding site was measured with an optical rangefinder or topographic map. A potential feeding site was defined as the nearest grain field. Land use will be compared with that in 1985 documented by the Agricultural Stabilization and Conservation Service. c. Distances to potential disturbance areas such as roads, houses, farm buildings, and railroad tracks were measured with an optical rangefinder or topographic map. Distances were measured from the potential disturbance to the nearest wetland edge. d. Horizontal visibility was measured with a vegetation profile board (Nudds 1977). Viewing points were set at 10-m intervals along the wetland edge. From the viewing point, a 90-m transect was conducted perpendicular to the shoreline and percent visibility was measured at 15-m intervals. The profile board was 3 m high and divided into 0.5-m sections with the first 0.5-m section divided into 0.25-m increments for visibility measurements. Volunteers from the Department of Fishery and Wildlife Biology assisted in collecting the horizontal visibility measurements. e. Vegetation composition and distribution were mapped for each wetland from ground inspection. Aerial photographs were taken of all study sites to facilitate in mapping the wetland more accurately and ascertaining the percent and distribution of vegetation. f. Specific conductivity was measured at each wetland with a conductivity meter. g. Measurements of pH were taken at each wetland with a portable pH meter. h. Visual indicators of water depth, mudflat area, and turbidity were recorded. Turbidity was recorded on a scale of 1 (100% visibility to substrate), 2 (partial visibility to substrate), or 3 (0% visibility to substrate). Mudflat area, if present, was measured along transects at 10-m intervals across the mudflat. �482 Measurement of Wetland Characteristics of Traditional Stopover Sites Observation of whooping cranes in the San Luis Valley began 28 October and continued through 8 November. Roost sites were located by observing foraging whooping cranes just before sunset and watching them fly to roost and by observing whooping cranes on roost sites just before sunrise. Personnel at Monte Vista NWR and W. M. Brown were contacted for suggestions on possible roost site locations. All of the above variables were measured for the traditional roosting sites of whooping cranes in the San Luis Valley except water depth profiles, percent submergent vegetation, and invertebrate sweeps because the wetlands were frozen at the time of investigation. Documentation of Whooping Crane Behavior Whooping cranes and sandhill cranes were observed at Grays Lake NWR from 31 July to 5 August to facilitate familiarity with time budget methodology. Time budget data were collected on whooping crane #8422 during stopover period.s in eastern Colorado and in the San Luis Valley. Coded behaviors were recorded at 15-second intervals during the observation periods. Observation periods occurred between dawn and dusk. Night-time observations were not made at the suggestion of Dr. R. C. Drewien as whooping cranes sleep through the night. This methodology corresponds to that used by W. M. Brown and Dr. R. C. Drewien for whooping cranes of the Grays Lake flock. Time budget data collected on whooping crane #8422 will be statistically compared to data collected on whooping cranes using traditional stopover areas in the San Luis Valley. Statistical comparisons will also be made between the data collected at the non-traditional site and data collected at the traditional stopover site on #8422. Statistical Comparisons Data on habitat evaluation and time budget are being entered into LOTUS for statistical analyses. Development of Standardized Methodology The development of standardized methodology for habitat evaluation of future stopover sites and recommendations for management of whooping cranes during fall will be completed following the evaluation of data collected during fall 1986 and the proposed data collection for fall 1987. RESULTS AND DISCUSSION Quantification of Habitat Characteristics of Eastern Colorado Sites Distance measurements to power lines, fences, potential disturbances, and feeding site were completed. Because of change in land use from 1985 to 1986, the Agricultural Stabilization and Conservation Service CASCS) for Weld County was contacted for land use information during 1985. This information will be used to identify any changes in land use from 1985 to 1986. Corrections will be made in distance to nearest forage site as would have been present during 1985. �483 Horizontal visibility, vegetation composition and distribution, and visual indicators of water depth measurements were completed. It is recognized the water depth measurements taken in fall 1986 will be different from water depths that would have occurred in fall 1985. Also, water depths at a number of sites fluctuated within a season due to control of water levels by irrigation companies. Specific conductivity and pH measurements fluctuated throughout the.day. A range of these measurements for each wetland was recorded. The pH measurements ranged from 7.1 to 9.3. Specific conductance ranged from 1000 to 7500 micromhos/cm. Sweep net sampling of sites in eastern Colorado for invertebrate and fish composition and density yielded few organisms per sample. A net with l-cm mesh was used to retain only those macroinvertebrates of a size large enough to be potential whooping crane food items. The sweeps that produced invertebrates and/or fish were taken from wetlands with submergent vegetation. Measurement of Wetland Characteristics at Traditional Stopover Site One juvenile and 21 adult whooping cranes were observed in the San Luis Valley and 15 roost sites were located. The whooping cranes all roosted with sandhill cranes with a number of roost sites being used by more than one whooping crane. Three roost sites were on Monte Vista NWR, 1 was on Alamosa NWR, 7 were in the Rio Grande bottomlands, 1 was at a starch factory sewage pond, and 3 were approximately 1.2 km from the river. Habitat evaluations of the used roost sites could not be completed while the sandhill cranes and whooping cranes remained in the San Luis Valley. Because loafing areas occurred adjacent to the roost sites, there were birds present at or near the roost sites at all times. To avoid disturbing the cranes, I planned to return to the valley after the cranes had migrated to New Mexico. I returned to the San Luis Valley on 13 November with a volunteer to complete the habitat evaluations of the 15 used roost sites. All measurements were completed except water depth profiles, percent submergent vegetation, and invertebrate sweeps because the wetlands were frozen. Documentation of Whooping Crane Behavior Whooping crane #8422 was first observed in eastern Colorado by a landowner on 17 October. I returned from the San Luis Valley on 18 October and began time budgeting that day and continued through 26 October when I observed the bird spiraling and leaving the area. Over ~2 hours of time budget data were collected during the fall 1986 stopover period. During the 10-day stopover, whooping crane #8422 was observed using the same roosting site of the 2 previous periods (fall 1985 and spring 1986). On 2 nights the bird was observed using a different roost site 0.5 km from the used site. The bird was observed foraging in a volunteer barley field adjacent to the corn field used during the 1985 fall stopover. This cornfield had severe hail damage and no ears had been formed providing no grain for the whooping crane to forage on. The whooping crane was also observed to forage in wet areas adjacent to the barley field. �484 Whooping crane #8422 was first observed in the San Luis Valley by Alamosa NWR personnel on 2 November. The whooping crane was observed foraging in a barley field on Alamosa NWR with a group of sandhill cranes and 1 adult whooping crane. I began collecting time budget data on 3 November. I was able to collect time budget data during the foraging periods in the barley field. I did locate the roost site and adjacent loafing area used by this flock but could not approach the birds to collect time budget data without flushing the flock. Approximately 19 hours of time budget data were collected during the foraging periods for comparison with time budget data collected during the Hudson stopover period. Whooping crane #8422 was first observed by Bosque del Apache NWR personnel on 12 November. The bird remained in the vicinity of the refuge for 1-2 days and was then observed at Casa Colorada State Refuge, New Mexico. Casa Colorada SR is a traditionally used winter site by whooping cranes of the Grays Lake flock. LITERATURE CITED Brown, W. M., R. C. Drewien, and E. G. Bizeau. 1985. Mortality of cranes and waterfowl from powerline collisions in the San Luis Valley, Colorado. Pages 128-136 in J. C. Lewis, ed., Proc. 1985 Crane Workshop, Natl. Audubon Soc., Tavernier, FL. Cowardin, L. M., V. Carter, F. C. Golet, and E. T. Laroe. 1979. Classification of wetlands and deepwater habitats of the United States. U.S. Dep. Inter., Fish and Wildl. Serv., BioI. Servo Prog., FWS/OBS-79/3l. 103 pp. Drewien, R. C., and E. G. Bizeau. 1974. Status and distribution of greater sandhill cranes in the Rocky Mountains. J. Wildl. Manage. 38:720-742. , and 1978. Cross-fostering whooping cranes to sandhill foster parents. Pages 201-222 in S. A. Temple, ed., Endangered birds: management techniques for preserving threatened species. Univ. Wisconsin, Madison. ----crane Kauffeld, J. S. 1981. Management of migratory crane habitat on Alamosa and Monte Vista National Wildlife Refuges. Pages 117-121 in J. C. Lewis, ed. Proc. 1981 Crane Workshop. National Audubon Soc., Tavernier, FL. Nudds, T. D. 1977. Quantifying the vegetative structure of wildlife cover. Wildl. Soc. Bull. 5:113-117. Prepared by -=-_____,...-_i_,:I..M.J.j_+-o--__ W\ __ . _~ Tanya M! Shenk Graduate Research Assistant Aprpoved bY~Rti£~ Wildlife Researcher _ � https://cpw.cvlcollections.org/files/original/9d3d2ddfb1fd93be1de76035e5265a19.pdf c0f03df391a48291b4b585a183389e27 PDF Text Text Colorado Division of Wildlife Wildlife Research Report July 1987 1 JOB PROGRESS REPORT State of Colorado Project No. 01-03-047 (FW 26 p) ------------~------~-- Mammals 1 Research Work Plan No. 1 Multispecies Jnvestigations Job No. 7 Terrestrial Research Publication, Editing and Library Service Period Covered: July 1, 1986 - June 30, 1987 Authors: Personnel: M. W. Hershcopf, L. H. Carpenter R. B. Gill, N. McEwen, L. Lovett, S. Gaylord, L. Ruminsky, and all Mammals Researchers ABSTRACT During the 1986-87 Segment, 15 books were purchased for permanent reference by DOW personnel. Sixty-five additional publications were located, ordered, and obtained free of charge for use. Eighteen theses were purchased, obtained on interlibrary loan, or given to the library. An additional 840 individual references requested by Mammals Researchers were located by library staff and made available for use. About 20 of these requests were not available locally and were obtained through interlibrary loans. Fifteen manuscripts were published and 12 manuscripts were in the review process. ��3 MAMMALS PUBLICATION EDITING AND LIBRARY SERVICES Marian W. Hershcopf and Len H. Carpenter P. N. OBJECTIVE To provide a centralized support program for manuscript editing and library services to facilitate publishing results of research conducted in projects 01-03-047 - 11700 and 01-03-048 - 11700, 11400, 16700. SEGMENT OBJECTIVES To provide coordinated and efficient editing and library services and publish findings for all Colorado Mammals Research programs. SUMMARY OF SERVICES Publications purchased with Mammals Research funds and placed in the Research Center Library Chanin, P. l79pp. 1985. The natural history of otters. Facts on File, New York, NY. Davis, D. E. (ed.). 1982. CRC handbook of census methpds for terrestrial vertebrates. CRC Press, Boca Raton, FL. 397pp. Fennessy, P. F., and K. R. Drew (eds.). 1985. Biology of deer production. Proc~ of an Interntl. Conf., Dunedin, New Zealand. February 13-18, 19~3. The Royald Soc!. of New Zealand, Wellington. Bulletin 22. 482pp. . What Indian water Folk-Williams, J. A. 1982. Water in the West. VoL L means to the west. Western Network, Santa Fe, NM. l53pp. _____ , and J. S. Cannon. 1983. Water in the West. Vol. 2. Water for the energy market--a source-book. Western Network, Santa Fe, NM. l62pp. , S. C. Fry, -----Western water and L. Hilgendorf. 1985., Wat~r i~ the West._ VoL 3. flows to the cities. Western Network, .Santa Fe, NM, and Island Press, Covelo, CA. 2l7pp. Gentry, R. L., and G. L. Kooyman (eds.). 1986. Fur seals: material strategies on land and at sea. Princeton Univ. Press, Princeton, NJ. 29lpp. Grey, G. W., and F. J. Deneke. York. 299pp. Griffin, D. R. 237pp. 1986~ 1984.· Animal thinking. Urban forestry. 2nd ed. Wiley, New Harvard Univ. Press, Cambridge, MA. �4 Halfpenny, J. 1986. A field guide to mammal tracking in western America. Johnson Books, Boulder, CO. l6lpp. Lee, J. A., S. McNeill, and 1. H. Rorison (eds.). 1983. Nitrogen as an ecological factor. 22nd Symp. of British Ecological Soc. 1981, Blackwell Sci. Publi., Boston. 470pp. National Research Council (U.S.). Subcommittee on Mineral Toxicity in Animals. 1980. Mineral tolerance of domestic animals. Nat'l. Acad. of Sci., Washington, DC. 577pp. Northern Wild Sheep and Goat Council. Symp. April 14-17. Missoula, MT. 1986. Proceedings of the 5th Biennial 435pp. Pearce, S. C. 1983. The agricultural field experiment: a statistical examination of theory and practice. Wiley, New York, NY. 335 pp. Peek, J. M. 1986. A review of wildlife management. Cliffs, NJ. 486pp. Prentice-Hall, Englewood Roberson, J., and F. Lindzey (eds.). 1984. Proc. of the 2nd Mountain Lion Workshop, Zion Nat'l. Park, UT, November 27-29, 1984. UT Div. of Wildl. Resources, Salt Lake City, UTe 27lpp. White, R. J. 1987. Big game ranching in the United States. Goat Internatl., Mesilla, NM. 355pp. Wild Sheep and Publications Obtained Free or at Low Cost In addition to books purchased with Federal Aid Funds, about 65 free reports and short publications from state or federal agencies or from private sour~es, were located, ordered, and -obtained for use by Mammals Research personnel. Theses Purchased, Obtained on Interlibrary Loan or as Gifts for Use by Researchers Anderson, C. F., Jr. 1959. Nocturnal activities of the Columbian blacktailed deer, Odocoileus hemionus columbianus.Richardson,- affecting spotlight census results in the Oregon Coast Range. - M.S. Thesis, Oregon State Univ., Corvallis. 86pp. Blank, D. L. 1984. Forage quality comparison of burned and nonburned aspen communities. M.S. Thesis, Utah State Uriiv., Logan. 74pp. Bromley, P. T. 1967. Pregnancy, birth, behavioral development of the fawn, and territoriality in the pronghorn (Antilocapra americana Ord) on the National Bison Range, Moiese, Montana. M.A. The!3is, Univ. of Montana, Missoula. l32pp. �5 Campbell, G. R. 1967. white-tailed deer. The dental cementum technique for age determination of M.A. Thesis, S. Ill. Univ, Carbondale. 27pp. Fox, J. L. 1983. Constraints on winter habitat selection by the mountain goat (Oreamnos americanus) in Alaska •. Ph.D. Thesis, Univ. of Washington, Seattle. l47pp. Garris, R. S. 1983. Habitat utilization and movement 'ecology of black bears in Cherokee National Forest. M.S. Thesis, Univ. of Tennessee, Knoxville. 98pp. Harpman, D. A. 1984. An economic analysis of the nongame check-off. Thesis. Colorado State Univ., Fort Collins. 70pp. Jackson, D. H. 1986. Ecology of bobcats in east-central Colorado. Thesis, Colorado State Univ., Fort Collins. l16pp. M.S. Ph.D. Johnson, R. E. 1977. Road and aerial surveys for deer in the North Dakota Coteau region. M.S. Thesis, N. Dakota State Univ., Fargo. 47pp. Kennedy, P. K. 1985. Genetic variability in white-tailed deer (Odocoileus virginianus) and its relationship to-environmental parameters and herd origin. Ph.D. Dissertation, Memphis State Univ. King, M. M. 1985. Behavioral responses of desert bighorn sheep to human harassment, a comparison of .disturbed and undisturbed populations. Ph.D. Dissertation, Utah State Univ., Logan. Kralka, R. A. 1983. Development and transmission of Protostrongylus boughtoni (Nematoda: -Metastrongyloidea), a lungworm of the snowshoe hare (Lepus americanus). M.S. Thesis, Univ. of Alberta, Edmonton, Alberta, Canada. 2l0pp. Ludwig, J. L. 1967. Comparison of age determination techniques for the whitetailed deer of southern Illinois. M. A. Thesis, S. Ill. Univ., Carondale. 30pp. Ravey, R. R. 1984. Reintroduction of desert bighorn sheep into western Colorado. M.S. Thesis, Colorado State Univ., Fort Collins. 93pp. Robson, M. S. 1982. Monitoring river otter populations: scent stations vs. sign indices. M.S. Thesis, Univ. of Florida, Gaine~ville. -4lpp'. Sayre, R. 1987. Effect of shrub stems on microhistological estimates of ruminant diets. M.S. Thesis, Colorado State Univ., Fort Collins. 52pp. Trowbridge, B. J. 1983. Olfactory communication in the European otter, Lutra 1. lutra. Ph.D. Thesis, University of Aberdeen. Wigley, T. B. 1981. A model of beaver damage and control. Ph.D. Dissertation, Miss. State Univ., Mississippi State. l5lpp. �6 Reference Document Location and Delivery The Research Center Library staff also located and delivered about 840 individual articles on request for Mammals Researchers during this segment; about 20 were not available locally and were obtained through interlibrary loan procedures. Manuscripts Published Job Progress Reports; Federal Aid. All studies. Anderson, E. M. 1987. A critical review and annotated bibliography of literature on the bobcat. Colo. Div. Wildl. Spec. Rep. No. 62:61. Baker, D. L., and N. T. Hobbs. 1987. Strategies of digestion: digestive efficiency and retention time of forage diets in montane ungulates. Can J. Zool. (In Press). Bartmann, R. M., G. C. White, L. H. Carpenter, and R. A. Garrott. 1987. Areial mark-recapture estimates of confined mule deer in pinyon-juniper woodland. J. W ildl. Manage. 51:41-46. Carpenter, L. H., and R. B. Gill. 1987. Antler point regulations: The good, the bad, and the ugly. West. Assoc. Fish and Wildl. Agencies. Proc , (In Press). Freddy, D. J. 1987. The White River Elk herd: Colo. Div. Wildl. Tech. Publ. No. 37. a perspective, 1960-1985. Garrott, R. A., G. C. White, R. M. Bartmann, and L. H. Carpenter. Movements of female mule deer. J. Wildl. Manage. 51:634-643. Hobbs, N. T. 1987. Fecal indices to dietary quality: Wildl. Manage. 51:317-320. __ 1987. a critique. / J. :-'and L. H. Carpenter. 1986. Viewpoint: animal-unit-equivalents be weighted by dietary differences. J. Range Manage. 39:470. should Kufeld, R. C., D. C. Bowden, and J. M. Siperek, Jr. 1987. Evaluation of a telemetry system for measuring habitat usage in mountainous terrain. Northwest Sci. 61: (In Press). ---of, , and D. L. Schrupp. 1988. Influence of hunting on movements female mule deer. J. Range Manage. 41: (In Press). , , and ---mark-recapture 1987. Estimating mule deer density by combining and telemetry data. J. Mammal. 68: (In Press). Miller, M. W., N. T. Hobbs, W. H. Rutherford, and L. L. Miller. 1987.· Efficacy of injectable ivermectin for treating lungworm infections in mountain sheep. Wildl. Soc. Bull. 15:260-263. Pojar, T. M. 1987. Taxonomic history of the pronghorn. Pronghorn Workshop. Proc: (In Press). �7 Tiedeman, J. A., R. E. Francis, C. Terwilliger, Jr., and L. H. Carpenter. 1987. Shrub-steppe habitat types of Middle Park, Colorado. USDA Forest Servo Res. Paper RM-273. 20pp. White, G. C., R. A. Garrott, R. M. Bartmann, L. H. Carpenter, and A. W. Alldredge. 1987. Survival of mule deer in Northwest Colorado. J. Wildl. Manage. 51:852-859. Manuscripts in Review Anderson, A. E. , and ---Colorado Movements of a young translocated puma. Southwest Nat. Seasonal variation in mule deer densities on a northcentral winter range. Southwest Nat. , , and D. E. Medin. ---~deer:--southwest Nat. Kidney fat index and kidney fat weight in mule Armeder, H., D. A. Leckenby, D. J. Freddy, and L. Hicks. Integrated management of timber and deer--interior forests. Handbook for Habitat Futures Workshop. Ministry of Forests, British Columbia. Bear, G. D. Seasonal distribution and population characteristics of the Estes Valley, Colorado, elk herd. Colo. Div. Wildl. Spec. Rep. , G. C. White, L. H. Carpenter, R. B. Gill, and D. Essex. Observability ---bias in mark-resighting estimates of elk populations. J. Wildl. Manage. Kufeld, R. C., D. C. Bowden, and D. 1. Schrupp. Habitat selection and activity of female mule deer in the Front Range, Colorado. J. Range Manage. Minimum convex polygons, home range utilization, and --- , and nonparametric tolerance regions. J. Mammal. Home range and movements of female mule deer in the ---- , and Rocky Mountain foothills. J. Mammal. Reed, D. F. Activity patterns of sympatric mountain goats and mountain sheep. J. Wildl. Manage. Alpine habitat selection in sympatric mountain goats and mountain sheep. Great Basin Nat. Torbit, S. C., L. H. Carpenter, R. M. Bartmann, and A. W. Alldredge. 1988. Calibration of carcass fat -indices in wintering mule deer. J. Wildl. Manage. Prepared by M c.V\'c.v--. IJ. f-/Q./"skc.or/Marian W. Herschcopf Librarian Len H. Carpenter Wildlife Research Leader ��Wildlife Research Report July 1987 9 JOB PROGRESS REPORT , State of Colorado Project No. 01-03-047 (FW 26 P) --------------~------~- Mammals 1 Research Work Plan No. 2 Deer Investigations Job No. 6 Winter Habitat Selection and Activity Patterns of Mule Deer in Front Range Range Shrubland and Forest Habitats Period Covered: July 1, 1986 - June 30, 1987 Author: R. C. Kufeld Personnel: D. Bowden ABSTRACT Six manuscripts were prepared describing results of the study. One was accepted for publication in the Journal of Mammalogy. Another has been accepted by Northwest Science. Others will soon be submitted to appropriate scientific journals, thus completing the study. :;.:.... ��11 WINTER HABITAT SELECTION AND AC1IVITY PATTERNS OF MULE DEER IN FRONT RANGE SHRUBLAND AND FOREST HABITATS P. N. OBJECTIVE 1. To test Telonics telemetry equipment to determine its accuracy at various distances in locating a transmitter on the Horsetooth Mountain study area and the ability of an observer using that equipment to correctly detect deer activity patterns by habitat type. 2. To determine habitat selection and activity patterns of mule deer within habitat types in the Horsetooth Mountain area during winter. SEGMENT OBJECTIVE Prepare manuscripts summarizing results of the recently completed study. ACKNOWLEDGMENTS D. Bowden played an important part in manuscript preparation. METHODS Six manuscripts: (1) A method for estimating mule deer density by combining mark-recapture and telemetry data; (2) evaluation of a telemetry system for measuring habitat usage in mountainous terrain; (3) Minimum convex polygons, home range utilization, and nonparametric tolerance regions; (4) Home range and movements of female mule deer in the Rocky Mountain foothills; (5) Habitat selection and activity of female mule deer in the Front Range, Colorado; (6) Influence of hunting on movements of female mule deer;_have been prepared describing results of this study. RESULTS Manuscript Mammalogy Science. journals, number 1 has been accepted for publication in the Journal of and number 2 has been accepted for publication in Northwest Other manuscripts will soon be submitted to appropriate scientific thus completing the ~tudy. Prepared by ~c~ Roland C. Kufeld Wildlife Researcher ��~OLOraQO VlvlSion or Wiidlife Wildlife Research Report July 1987 13 JOB PROGRESS REPORT State of Colorado Project No. 01-03-047 (FW 26 p) --------------~------~- Mammals 1 Research Work Plan No. 2 Deer Investigations Job No. 7 Development of Census Methods for Deer in Plains Riverbottom Habitats Period Covered: July 1, 1986 - June 30, 1987 Author: R. C. Kufeld Personnel: D. Bowden, D. Whittaker ABSTRACT Sixty-eight mule and white-tailed deer were trapped in Clover traps in the South Platte riverbottom between Kuner and Sterling, Colorado, during 19 January - 17 February 1987. Twenty-five adults (13 mule deer and 12 whitetails) were radiocollared and eartagged. Remaining deer, mostly fawns, were eartagged. The most effective trap bait was eared corn. Less effective baits in decreasing order were apple pulp, deer-elk wafers, oats soaked in molasses, and corn silage. Instrumented deer were located by aerial telemetry at 2-week intervals to determine movements and home range size. More locations are needed before conclusions can be made. Tests to measure observer accuracy in locating transmitters by aerial telemetry showed a mean distance error between estimated and actual transmitter locations of 197 m with a range of 25 m to 789 m. Altitudes and airspeeds at which transmitter locations were estimated during the experiment did not affect distance error. ��15 DEVELOPMENT OF CENSUS METHODS FOR DEER IN PLAINS RIVERBOTTOM HABITATS Roland C. Kufeld P. N. OBJECTIVES 1. To determine seasonal movements and home range size of mule and whitetailed deer in plains riverbottom habitats. 2. To develop and test methods for estimating size of deer populations in plains riverbottom habitats. SEGMENT OBJECTIVE To determine seasonal movements and home range size of mule and white-tailed deer in plains riverbottom habitats. ACKNOWLEDGMENTS Don Whittaker and numerous personnel from Colorado Division of Wildlife Northeast Region were instrumental in capturing deer. Don whittaker was also active in evaluating deer preference for baits and accuracy of determining deer locations by aerial telemetry. STUDY AREA The study area is the South Platte River basin extending from Greeley, Colorado, to Nebraska. Riparian vegetation in the floodplain is dominated primarily by cottonwood (Populus sargentii), and willow (Salix spp~). The riverbottom is bordered along much of its length by agricultural lands, mainly cornfields. Other stretches are bordered by rangelands dominated by mixed prairie or sand sagebrush (Artemisia filifolia) (Costello 1954). Land status includes both private lands and state-owned wildlife areas. Results from the South Platte should apply to most other plains riverbottom habitats such as those found along the Arkansas River. METHODS AND MATERIALS Mule and white-tailed deer were trapped and marked in the South Platte riverbottom between Kuner and Sterling, Colorado, during the period 19 January 17 February 1987 (Table 1). Each adult doe and most adult bucks received a radiocollar and 2 orange, numbered eartags. Remaining adult bucks and all fawns received only eartags. Deer were captured in Clover traps (Clover 1956). Each trap was baited with alfalfa hay, apple pulp, and eared corn. An experiment to determine deer preference for trap baits was conducted concurrently with the trapping operation by seasonal employee, Don Whittaker, �16 in fulfillment of requirements for a special studies class at Colorado State University. Five baits were compared; apple pulp, eared corn, corn silage, oats soaked in molasses, and deer-elk wafers. Baits were weighed and placed into buckets and offered to wild deer in a randomized block design involving 2 replications of each bait. The experiment involved 3 trials each lasting at least 3 nights. Buckets were weighed daily to measure kg of bait removed. Deer always had adequate amounts of bait from which to select. Test statistics used to determine if there was a significant difference in amounts of bait removed were Wilks' Lambda, Pillai's Trace, Lawley-Hotelling Trace, and Roy's Greatest Root (Morrison 1976). Selection or avoidance of baits was determined using Duncan's multiple range test of means (SAS Institute, Inc. 1985). Radio-collared deer were located by aerial telemetry at approximately 2-week intervals beginning 23 February 1983. The aircraft was a Cessna 185 with a 2-element yagi antenna mounted on each strut. A switchbox permitted the telemetry operator, a passenger in the aircraft, to operate antennas jointly or separately. Deer locations were plotted on 1:50,000 scale USGS maps and identified by UTM coordinates to within 10 m. A test was conducted to measure observer accuracy in locating transmitters by aerial telemetry. Transmitters were placed on a post about 1 m above ground level at 29 known locations within the area where deer had been instrumented. Transmitters were placed by a second party, and actual locations were unknown to the aerial telemetry operator. Estimated locations were subsequently determined by the aerial operator, assigned to periodically locate instrumented deer, using the same aircraft and telemetry equipment. Altitude and airspeed were recorded at the moment each location was determined. However, the pilot attempted to fly as low and slow as possible for the given situation when each location was estimated. Actual and estimated transmitter locations were plotted on 1:50,000 scale USGS maps and described by UTM coordinates to within 10 m. Location error was measured as the straight-line distance between actual and estimated locations. Data were subjected to regression analysis with distance error as the dependent variable and airspeed and true altitude (altitude above ground level) as independent variables, a~d to analysis of variance for unbalanced designs. RESULTS AND DISCUSSION Sixty-eight deer were captured and marked, of which 25 adults (13 mule deer and 12 whitetails) were radiocollared (Tables 1 and 2). Eared corn was by far the most successful bait for attracting deer into Clover traps. Deer in the study area are accustomed to feeding in cornfields and readily recognized corn as a preferred food. In traps, apple pulp was not as preferred as corn, and alfalfa hay was rarely eaten by deer. Based on our experience, however, we suggest that the smell of apple pulp and green color of hay may help attract deer to trapsites. Thus, attractiveness of a baited trap may be increased if all 3 baits are used in the same trap. Mean amounts of bait consumed per night in bait preference experiments were 2.77 kg, 2.76 kg, 1.53 kg, 0.86 kg, and 0.45 kg for eared corn, apple pulp, deer-elk wafers, oats, and corn silage, respectively, over all trials. Analysis of variance indicated that choices were definitely being made by deer (p < 0.05 in tests by all 4 methods). Deer in trial 1 showed no preference �17 for any bait, but eared corn and apple pulp were preferred baits in trials 2 and 3. In all 3 trials, deer avoided corn silage. Aerial tracking of radio-collared deer is in its initial stages, and the number of flights made to date is insufficient for basing conclusions concerning deer movements or home range size. Plans call for aerial tracking of deer for at least 1.5 years. Error distance between actual and estimated transmitter locations varied from 25 m to 789 m (x = 197 m, s = 166 m). Airspeed varied between 65 and 80 knots = 71.6 knots, s = 4.4 knots), and altitude varied between 24 and 171 m (x = 74.9 m, s = 36.5 m). Regression analysis of distance error vs. altitude and distance error vs. airspeed, produced r2 values of 0.05 and 0.0002, respectively. Altitude and airspeeds at which transmitter locations were estimated during the experiment did not affect distance error (p = 0.21 and 0.20, respectively). The magnitude of observer error in locating transmitter by aerial telemetry appears to be sufficiently small to enable use of the technique to achieve study objectives. ex LITERATURE CITED Clover, M. R. 1956. Single-gate deer trap. Calif. Fish and Game 42:199-201. Costello, D. F. 1954. Vegetation zones in Colorado. Pages iii-x in H. D. Harrington, ed. Manual of the plants of Colorado. Sage Books, Denver, Colo. Morrison, D. F. York. 1976. Multivariate statistical methods. McGraw-Hill, New SAS Institute, Inc. 1985. SAS/STAT Guide for personal computers, version 6 edition. Carey, No. Car. Prepared by 'ld~~1-C ~ d I} d Roland C. Kufeld Wildlife Researcher ~ �13 Table 1. Mule and white-tailed deer trapped and marked in the South Platte riverbottom between Kuner and Sterling, Colorado, during 19 January 17 February 1987. Radiocollared Species mule deer Sex buck doe whitetail buck doe TOTAL Age adult fawn adult f awn adult fawn adult fawn and eartagged 3 Eartagged only 3 9 10 2 3 1 16 9 25 Total 6 9 10 2 4 16 9 12 12 - - 43 68 �19 Table 2. 2-17-87. Deer tagged along South Platte River between Hardin and Sterling, Colorado, 1-19-87 through Eartag number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 SEecies Sex whitetail whitetail whitetail whitetail mule deer mule deer mule deer whitetail whitetail whitetail whitetail mule deer mule deer mule deer whitetail whitetail whitetail whitetail whitetail whitetail mule deer mule deer mule deer whitetail mule deer mule deer whitetail whitetail mule deer whitetail whitetail whitetail mule deer whitetail whitetail whitetail whitetail whitetail whitetail whitetail whitetail whitetail whitetail whitetail whitetail whitetail whitetail mule deer mule deer mule deer mule deer mule deer mule deer mule deer mule deer whitetail mule deer whitetail mule deer whitetail mule deer whitetail whitetail whitetail mule deer mule deer mule deer whitetail buck buck buck buck buck buck buck doe buck buck buck doe doe buck doe doe buck buck doe buck buck doe doe buck buck buck doe buck doe doe doe doe buck buck doe doe doe doe buck buck doe buck doe doe buck doe doe doe buck doe buck buck doe buck doe doe doe buck buck doe doe doe buck doe buck buck doe buck Age when caEtured fawn fawn fawn fawn fawn fawn fawn adult fawn fawn fawn adult adult fawn fawn adult fawn fawn fawn fawn fawn adult adult fawn adult adult fawn adult adult fawn fawn fawn adult fawn adult adult fawn fawn adult fawn fawn adult adult adult fawn adult fawn adult adult adult fawn adult fawn fawn fawn fawn adult adult fawn fawn adult adult fawn adult fawn adult adult fawn Date caEtured 1-19-87 1-19-87 1-19-87 1-19-87 1-19-87 1-19-87 1-19-87 1-19-87 1-19-87 1-19-87 1-19-87 1-19-87 1-19-87 1-19-87 1-20-87 1-20-87 1-20-87 1-20-87 1-20-87 1-20-87 1-20-87 1-20-87 1-20-87 1-20-87 1-21-87 1-21-87 1-22-87 1-22-87 1-22-87 2- 2-87 2- 2-87 2- 2-87 2- 2-87 2- 2-87 2- 3-87 2- 3-87 2- 3-87 2- 3-87 2- 3-87 2- 3-87 2- 3-87 2- 3-87 2- 3-87 2- 4-87 2- 4-87 2- 4-87 2- 9-8(_ 2- 9-87 2- 9-87 2-12-87 2-10-87 2-10-87 2-10-87 2-10-87 2-10-87 2-11-87 2-11-87 2-11-87 2-11-87 2-11-87 2-12-87 2-13-87 2-13-87 2-17-87 2-13-87 2-17-87 2-17-87 2-17-87 Capture location 4 mi E of Atwood Bridge 4 mi E of Atwood Bridge 1 mi E of Hillrose Bridge 1 mi E of Hillrose Bridge 1 mi E of Hillrose Bridge 1 mi E of Weldona Bridge 1 mi E of Weldona Bridge 1 mi E of Weldona Bridge 3.5 mi E of Hardin Bridge 3.5 mi E of Hardin Bridge 3.5 mi E of Hardin Bridge Hardin Bridge Hardin Bridge Hardin Bridge 4 mi E of Atwood Bridge 4 mi E of Atwood Bridge 4 mi E of Atwood Bridge 4 mi E of Atwood Bridge 3.5 mi E of Hardin Bridge 3.5 mi E of Hardin Bridge Hardin Bridge Hardin Bridge Hardin Bridge 1 mi W of Hardin Bridge Hardin Bridge Hardin Bridge 1 mi E of Weldona Bridge 1 mi W of Hardin Bridge Hardin Bridge 2.5 mi E of Merino Bridge 1 mi E of Merino Bridge 2 mi E of Fort Morgan Bridge 1 mi E of Weldona Bridge 3.5 mi E of Hardin Bridge 2.5 mi E of Merino Bridge 1 mi E of Merino Bridge 1 mi E of Merino Bridge 1 mi E of Merino Bridge 1 mi E of Merino Bridge 2 mi E of Fort Morgan Bridge 2 mi E of Fort Morgan Bridge 3.5 mi E of Hardin Bridge 3.5 mi E of Hardin Bridge 1 mi E of Merino Bridge 3.5 mi E of Hardin Bridge 3.5 mi E of Hardin Bridge 2.5 mi E of Merino Bridge 2 mi E of Orchard Bridge Hardin Bridge 1 mi W of Hardin Bridge 1 mi E of Weldona Bridge 1 mi E of Weldona Bridge Hardin Bridge Hardin Bridge Hardin Bridge 4 mi E of Atwood Bridge 2 mi E of Orchard Bridge 1 mi E of Hardin Bridge Hardin Bridge Hardin Bridge 1 mi W of Hardin Bridge 2 mi E of Fort Morgan Bridge 2 mi E of Fort Morgan Bridge 2.5 mi E of Masters Bridge 2 mi E of Orchard Bridge 3.5 mi E of Hardin Bridge Hardin Bridge 1 mi E of Hardin Bridge Frequency of radiocol1ar attached 149.511 149.521 149.531 149.538 149.551 149.631 149.641 149.651 149.660 149.671 149.701 149.731 149.741 149.770 149.780 149.791 149.800 149.809 149.859 149.839 149.869 149.889 149.921 149.951 149.970 ��'VU..LU.LdUU lJ..LV..L;:,.LUH UJ. W.l.J.u.J..l.le 21 Wildlife Research Report July 1987 JOB PROGRESS REPORT State of Colorado Project No. 01-03-047 Work Plan No. Job No. 10 Period Covered: Author: 2 (FW 26 p) Mammals 1 Research Deer Investigations Energy Development and Mule Deer in Piceance Basin - Cooperative Study July 1, 1986 - June 30, 1987 R. M. Bartmann Personnel: T. Abbott, R. Barks, R. Brazie, L. H. Carpenter, G. DeBella, J. Depperschmidt, R. Fithian, A. Gigliotti, J. Jackson, D. Saltz, D. L. Weybright, G. C. White, and numerous others from the Division of Wildlife and Colorado State University. ABSTRACT Three pastures from 66 to 169 ha in size were stocked with mule'deer (Odocoileus hemionus) at densities of 44, 89, and 133 deer/km2 to simulate removal rates of 67, 33, and 0%, respectively. Each pasture contained about 50 radio-collared fawns with adults added to achieve desired removal rates. Starvation rates were negatively related to removal rate (density) and survival rates positively related. Both starvation and survival rates differed (R < 0.01) among removal rates. ��23 ENERGY DEVELOPMENT AND MULE DEER IN PICEANCE BASIN - COOPERATIVE STUDY Richard M. Bartmann P. N. OBJECTIVES To estimate the degree of compensatory winter mortality forces operating within the mule deer fawn age class in Piceance Basin. SEGMENT OBJECTIVES 1. To trap and radiocollar 120 fawns in Piceance Basin to estimate overwinter survival rates. 2. Live-trap and remove 340 does and fawns on the treatment area to effect a 20% population reduction. 3. Stock 3 pastures with deer at 3 densities to simulate 3 removal rates. 4. Measure over-winter fawn mortality in the pastures. 5. Measure over-winter changes in body condition of all deer in the pastures. The study is divided into 2 parts: a field experiment and a ment. The field experiment is the primary responsibility of Colorado State University, and the pasture experiment is the responsibility of the Division of Wildlife. Therefore, only experiment is reported here. pasture expericooperators at primary the pasture METHODS AND MATERIALS Deer that were live-trapped and removed for the field experiment were used to study the compensatory mortality process under more controlled conditions in fenced pastures. Three pastures at the Little Hills Wildlife Area were stocked during late November and early December with mule deer at 3 densities: 44, 89, and 133 deer/km2• These stocking rates can be viewed as simulating removal rates of 67, 33, and 0%, respectively. Fifty fawns were put in each pasture for estimation of survival rates. Each fawn was weighed and radiocollared before being placed in the pastures. Enough adults were added to each pasture to attain the target densities, but they were neither weighed nor marked in any way. Extra adults not needed for ,this experiment were placed in a separate pasture. Fawn radiocollars contain-a motion sensor set with a 4-hour delay to enable detecting mortalities. Radio signals were monitored 5-7 days per week during winter and all mortalities checked for cause of death as soon as possible. In the spring when deer started to migrate, gates in all the pastures were opened and the deer allowed to leave voluntarily. It was decided not to retrap and reweigh deer as was done the previous spring as it was considered more important to monitor survival through the migration period. Monitoring �, 24 continued at 2-week intervals into the summer to enable retrieving collars as they dropped off. RESULTS Pasture stocking occurred from 10 November-22 December 1986. Poor success capturing fawns on the field study treatment area prompted use of dropnets in other areas to catch 39 fawns to help meet stocking needs. Even then, the 50fawn goal was not met in the low and medium density pastures (Table 1). Predation on fawns by bobcats (Lynx rufus) was, again, a problem in all pastures (Table 2). However, starvation rates still displayed a negative relationship to removal rate (density) and survival rates a positive relationship (Fig. 1). Fawn survival differed among pastures (p < 0.01) as did mortality due to starvation (p < 0.01). But predation was independent of pasture density (P = 0.40). Survival rates in all pastures were 24 to 31 percentage pOints-lower than in 1985-86, suggesting deteriorating forage conditions at all stocking densities. Prepared by &vp~~Richard M. Bartmann Wildlife Researcher �25 Table 1. Stocking summary for 3 pastures at the Little Hills Wildlife Area during winter 1986-87. Number of deer Stocking Fawns densitya Low Medium High Adults Fawn:adult M F M F ratio 20 29 23 26 19 28 8 7 6 19 35 170:100 114:100 128:100 19 96 Storage 34 Pasture size Hectares Deer/area Acres km mi 43 169 101 66 419 249 162 89 138 112 231 360 146 361 79 204 aStocking densities simulate removal rates as follows: Medium = 33%, and High = 0%. Low = 67%, Table 2. Fate of mule deer fawns stocked at 3 removal rates in pastures at the Little Hills Wildlife Area during winter 1986-87. Percentages are in parentheses. Mortality Removal rate (%) N 46 49 51 67 33 o Starvation Predation Other Total 18 (39) 19 (44) 34 (68) 6 (13) 8 (19) 1 (2) 25 (54) 4 (9) 11 (22) 1 (2) 31 (72) 46 (92) Survived 21 (46) (28) i2 4 (8) aAdjusted totals due to radio failures and escapes are 43 fdr the 33% removal rate and 50 for the 67% removal rate. All calculations are based on adjusted totals. -~ �26 c .-+--0 ~ 0 o, 0 ~ n, 0.9 0.8 0.7 0.6 0.5 0.40.3 0.2 0.1 Survival 1985-86 1986-87 •••• •••• ••••• ... , •••••• ••••• •••• ••••••..••••• 0 0.9 0.8 0.7 0.6 0.5 0.40.3 0.2 0.1 0 Starvation ••• ••• ••• ••• ••••••••••• ••• 1986-87 ._ . 1985-86 o 33 Removal Fig. 1. •••• •••• •••• 67 (%) Survival and starvation rates of radiocollared mule deer fawns stocked in pastures at 3 simulated removal rates, 1985-86 and 1986-87. �Colorado Uivision of Wildlife Wildlife Research Report July 1987 27 JOB PROGRESS REPORT State of Colorado Project No. 01-03-047 Work Plan No. Job No. 2 11 Period covered: Author: (FW 26 p) Mammals 1 Research Deer Investigations Testing of Mule Deer Census Methodology July 1, 1986 - June 30, 1987 R. M. Bartmann Personnel: L. H. Carpenter, D. L. Weybright, and G. C. White ABSTRACT Aerial line transect surveys were made to estimate mule deer (Odocoileus hemionus) density in pastures at the Little Hills Wildlife Area in Piceance Basin from 15-18 December 1986. Although density estimates were generated, they are preliminary and work is continuing to determine an appropriate estimator to use for deer in pinyon-juniper habitat. Computer simulations using the group size data collected during the surveys will be done after an estimator is selected • ~ . ��29 TESTING OF MULE DEER CENSUS METHODOLOGY Richard M. Bartmann P. N. OBJECTIVES 1. To evaluate the effects of different criteria for determining group size on deer density estimates and associated confidence interval widths in aerial line transect surveys. 2. To gather data on spatial distribution of deer within groups to enable computer simulations to test effects of different deer groupings on density estimates and associated confidence intervals. 3. To evaluate the appropriateness of size-bias techniques for estimating deer density with aerial line transect surveys. SEGMENT OBJECTIVES Same as P. N. Objectives. METHODS AND NATERIALS Procedures were generally the same as described by Bartmann (1986) with the following exceptions: Pastures 8 and 9, which were used to hold extra deer not needed in the compensatory mortality study, were included in the 1986 surveys. This allowed 7 more transects to be established. Also, the intensity of line transect surveys was reduced by only flying mornings and afternoons for 3 days. Additional information collected during line transect surveys was notations on which groups of deer might be combined as single groups because of their close proximity to each other. Information on sizes of deer groups that occur in pinyon-juniper habitat can then be used in computer simulations of the effects of this variable on density estimates from line transect surveys. RESULTS Line transect surveys were flown in the pastures on 15-18 December 1986. Deer density in the pastures was 0.664 deer/ha, or a total of 320 deer. Preliminary estimates of deer density are shown in Table 1. Evaluations to determine an appropriate line transect density estimator to use with deer data are still proceeding. Computer simulations to determine the effects of different deer groupings on density estimates and their associated confidence intervals will not be run until this process is completed. �30 LITERATURE CITED Bartmann, R. M. 1986. Testing of mule deer census methodology. \Uldl. Game Res. Rep. July: 41-46. Prepared by Colo. Div. tLL~~--Richard H. Bartmann Wildlife Researcher Table 1. Estimates of deer density (deer/km2) from aerial line transect surveys in the Little Hills pastures in Piceance Basin, 15-18 December 1986. True density was 66.4 deer/km2• Estimator Fourier series Fourier series with 1 extra term Exponential polynomial Exponential power series Negative exponential Half-normal Density 58.3 65.3 72.8 73.2 97.7 62.2 Coefficient of variation 8.3 9.7 13.7 14.9 9.7 8.8 �Wildlife Research Report July 1987 31 JOB PROGRESS REPORT State of Colorado Project No. 01-03-047 Work Plan No. Job No. 14 Period Covered: Author: 2 (FW 26 p) Mammals I Research Deer Investigations Coyote Control and Compensatory Mortality in Mule Deer Populations July 1, 1986 - June 30, 1987 R. M. Bartmann Personnel: L. H. Carpenter, J. D. Depperschmdit, A. Foster, G. Papez, R. Raley, G. Rowley, D. Saltz, D. L. Weybright, G. C. White, and numerous others from the Division of Wildlife and Colorado State University. ABSTRACT Sixty mule deer (Odocoileus hemionus) fawns were radio collared on CB Tract from 11-19 November 1986. ADC personnel removed 78 coyotes from the study area. The predation rate on fawns was 39%, and fawn survival was 32%. ��33 COYOTE CONTROL AND COMPENSATORY MORTALITY IN MULE DEER POPULATIONS Richard M. Bartmann P. N. OBJECTIVES 1. To estimate over-winter predation rates on mule deer fawns associated with lowered coyote densities. 2. To estimate the degree of compensatory winter mortality of mule deer fawns in response to expected changes in coyote predation rates. SEGMENT OBJECTIVES Same as P. N. Objectives. METHODS AND MATERIALS Mule Deer Mule deer fawns were trapped with dropnets and radiocollared on the CB Tract study area in November, 1986. Radiocollars were of break away design and contained a mortality sensor set with a 4-hr delay. Signals from all fawns were monitored 5-7 days per week beginning immediately after trapping and continuing until they migrated to summer range in April. Thereafter, monitoring was done twice monthly to enable retrieval of collars that dropped off. Each fawn mortality was located as soon as possible to determine cause of death. Coyote Coyote control was done primarily by Animal Damage Control (ADC} ,personnel from the U.S. Department of Agriculture. Control methods included trapping, shooting from the ground, and shooting from a helicopter and a fixed-wing aircraft. RESULTS Mule Deer Sixty mule deer fawns, 29 males and 31 females, were captured and radiocollared on the CB study area from 11-19 November 1986. The predation rate (39%) was the lowest recorded thus far, and was largely due to a marked reduction in coyote-kills during spring (Table 1). Fawn survival (32%) was close to the 34% recorded in 1982-83 (Fig. 1). Coyotes Seventy-eight coyotes were removed from the CB study area from 2 September 1986 through 28 February 1987 CTab1e 2). As in 1985-86, the age composition �34 of coyotes removed switched from predominantly juveniles during fall trapping (70%) to mostly adults during aerial gunning in winter (63%). 4, / /)/ ~. --~.c---- - Prepared by .f~:L::_/11.:~:J4.o..a:.•.",-,,:::-dh~. £_~/.:_0.Y2:::."'~/--="/~:::::::::...:'u::!.:::·;:;:;t;;::::~!::::::~::::::::-_'·_--_---_'~--"._.. Richard M. Bartmann Wildlife Researcher Table 1. cause of mortality for radiocollared mule deer fawns on CB Tract study are from 1 December-15 June 1981-82 through 186-87. Percentages (in parentheses) are calculated from the number of nonfailing radio collars. Winter N Mortality cause Starvation Road kill Predation Radio failure Other ------------------------------------------------------------------------------1981-82 1982-83 1983-84 1984-85 1985-86 1986-87 66 61 60 60 57 59 25 28 30 44 29 23 1 (2) 4 (7) 23 (40) 5 (9) 11 (19) 8 (14) (53) (46) (52) (76) (51) (39) 1 (2) 2 (3) 5 (11) 5 (8) 2 (3) 1 (2) 3 (5) 9 (15) 19 2 2 1 Table 2. Coyotes removed from the CB Tract study area from 2 September 1986 through 28 February 1987. Coyotes removed Removal method Male Female Unk Adult Juvenile Unk Male Female Hale Female Unk ------------------------------------------------------------------------------Trapping Ground shooting Helicopter gunning Fixed-wing gunning 6 2 5 3 1 4 4 1 1 7 5 1 5 1 2 1 2 14 13 �35 1.0 Predation 0.9 0.8 1:: 0.7 0 .~ 0.6 !- 0 0.5 0 0.4 c, !- u, 0.3 Survival ------ Starvation ••••••••••••• - - --~.... ... .. .. _.... . •••••• .•. ~ 0.2 0.1 ; --. ----- -- ••••••.... .....t.~··*'······ .••• .... •••~ .......... -- --~ ; ~ ,~- ~ 0 81-82 82-83 83-84 84-85 85-86 86-87 Winter Fig. 1. Survival and mortality rates of radio collared mule deer fawns on the CB Tract study area, 1981-82 through 1986-87. ��Colorado Division of Wildlife Wildlife Research Report July 1987 37 JOB PROGRESS REPORT State of Colorado Project No. 01-03-047 (YW 26 p) Mammals 1 Research Work Plan No. 3 Elk Investigations Job No. 2 Trapping, Transportating, and Maintenance of Elk at Livestock-Elk Grazing Study Period covered: July 1, 1986 - June 30, 1987 Author: G. D. Bear Personnel: D. Baker, L. Carpenter, T. Hobbs, T. Lines, C. Reichert, M. Miller, C. Woodward, R. Berry, T. Veeder, J. Papas, J. Hicks, M. Bauman, K. Navo, B. Sheridan ABSTRACT Twelve female elk calves were captured during June to start a captive elk herd. These calves will be used for grazing and nutritional experiments •. . . A total of 163 elk (71 cows, 75 calves, 6 yearling cows, 8 yearling males, and 3 mature bulls) were c-aptured with a portable group trap near Maybell, Colorado. Fifty-four cows were used to stock experimental grazing pastures 19 mi north of Maybell from December to mid-April. Ten additional elk (6 cows, 4 bulls) were trapped on the Godiva Rim·winter range which is 15 miles north of Maybell and were fitted with te~emetry collars. Seasonal movements of these elk were· measured. .. '. ~.: .: ,': :'-:-_~-'"I' . "-,. ' ��39 TRAPPING, TRANSPORTING, AND MAINTENANCE OF ELK AT LIVESTOCK-ELK GRAZING STUDY George D. Bear -- P. N. OBJECTIVE To provide assistance to the Livestock-Elk Grazing Study near Maybell, colorado, by capturing and maintaining experimental elk herd. SEGMENT OBJECTIVES 1. Capture 12 female elk calves for tame-elk herd. 2. Capture and maintain 54 adult cow elk at the experimental facilities near Maybell, Colorado. 3. Determine seasonal movements of radiocollared elk in the vicinity of Maybell, Colorado. METHODS AND RESULTS Capturing Elk Calves Rocky Mountain National Park, near--Bst es Park, Colorado, was selected as the capture site due to the high density of elk and previous experience at capturing elk calves in the park. Individuals searched £alving areas on foot in early morning and late evening in early June, 1986. In late June, a helicopter was used to locate and capture elk calves on alpine ranges. Twelve female calves were desired; 18 calves (12 females, 6 males) were captured before this quota was achf.eved, Captured calves were blindfolded and placed. in burlap bags until they could be transported to a vehicle. They wer.ethen transported to the animal-rearing facilities at Ft. Collins to become part of a captive herd used in experimental grazing trials. Capturing Elk for Pastures Trapsites were established at Godiva Rim (approximately 15 mi north of Maybell) and Axial Basin (25 mi southeast of Maybell) using alfalfa hay and livestock salt to attract elk. A portable-group trap was used. After a group of elk was captured, indiviudal elk were isolated into a work area. All elk were marked with numbered eartags. Only adult cows. (2 yrs and older) were selected for the experimental pastures. Blood samples were taken for brucellosis and leptospirosis tests, and collars were placed on each cow before they were loaded into a livestock trailer for transportation to the study site. All other elk were released at the trapsite. . At the study site located at the Little Snake River Management Grazing. Facility, 19 mi north of Maybell, elk were placed in a pole corral. They were kept in the corral for at least 24 hrs to permit them to adjust to the �40 enclosure. Next, they were permitted to move out into the "hub" area (approx. 3-acre work area). for 24-48 hrs to adjust to electric fences. Elk were then placed in the pole corral once again and worked through the chutes, weighed, and released into designated pastures. Objectives of this job were to maintain the. elk at desired stocking rates, maintain a reserve' group of cows, and maintain the fences and other facilities until __the elk were released in April. A total of 163 elk were captured to meet the stocking quota at the experimental pastures (Appendix A). Classification of these elk are as follows: adult cows 71; calves, males 39, females 34, unknown 2; yearling cows 6; yearling males 8; and adult males 3. Several replacement cows were needed after the initial stocking of 54 cow elk. On January 18, rabbit hunters caused some of the elk in 1 pasture to force their way through the fence; 4 replacements were needed. In February, another elk escaped from the pasture and had to be replaced. Two cows died of apparent malnutrition during March and April and had to be replaced from the reserve group being held in the pole corrals. Overall, the electric fences were very effective in containing the elk. All elk used at the experimental pastures were taken from the Axial Basin site. Elk in that area occupy very large wheat fields and open sagebrush rangelands and, thus, are easily seen and pursued by hunters during the hunting sesons, which results in animals being in poorer body condition t~an would be normally anticipated. It is recommended that an effort be made to obtain cows in better condition from a different locality in the future_ Elk Movements Ten elk (6 cows and 4 bulls) on Godiva rim were fitted with telemetry collars in February. An additional 4 cows were fitted with telemetry collars before releasing them from the experimental pastures. The collared elk on Godiva Rim remained on that winter range through the first part of April. By April 15, 2 bulls had moved 35 mi east to the Black Mountain area north of Craig. By May 1, only 2 collared cows remained on the Godiva winter range. On May 15, only 1 cow was still on the winter range; however, she migrated eastward within the next 10-day period. All collared elk migrated from the Gidova Rim area to the Black Mountain area and by mid-June were scattered on summer ranges to the vicinity of Steamboat Lake. This is approximately 55 mi north of the Godiva winter range. Three of the 4 collared cows released from the experimental pastures returned to the Axial Basin area where they were trapped •..This is a distance of approximately 30 mi. They had to cross U. S.:" Highway 40 (U.s. 40) and the Yampa River to complete the journey. By May 11, these cows were southeast of Hamilton.in the Wilson Mesa area. On June 12 they were on the sUInmer range around Pagoda Peak and Sand Peak. This is approximately 25 mi east of the Axial Basin winter range. The fourth collared cow remained on a cedar ridge east of Maybell, apparently refusing to cross U.S. 40. ~repared by ~,u.[1~. GeOrgei)Bear Wildlife Researcher �41 APPENDIX A ELK TRAPPED AT AXIAL BASIN CAX) AND GODIVA RIM CGR) DURING THE PERIOD FROM DECEMBER, 1986, THROUGH FEBRUARY, 1987 Eartag number Date Trap location Age Sex 7 8 9 9 10 11 12 13 14 15 16 17 18 19 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 49 50 51 52 53 54 55 57 58 59 60 61 62 63 65 66 67 69 70 71 72 74 76 77 78 79 80 81 83 84 85 86 87 88 89 90 91 92 93 12- 9 12-12 12-12 12-12 12-15 12-12 12-12 12-12 12-12 12-12 12-12 12-12 12-12 12-12 12-12 12-12 12-12 12-12 12-19 12-19 12-19 19-19 12-19 1- 2 12-19 12-19 12-19 12-19 12-19 1- 2 12-22 12-22 12-22 12-22 12-23 12-23 12-30 12-29 12-29 12-30 12-29 12-29 12-29 12-29 1- 9 1- 8 1- 8 1- 9 1- 9 1- 9 1- 8 1- 9 1- 9 1- 9 1- 9 1- 9 1- 9 1- 9 1- 9 1- 9 1- 9 1- 9 1- 9 1- 9 1- 9 1- 9 1- 9 1-12 1-12 1-12 1-12 1-15 1-15 1-15 1-15 1-15 2-11 .2-11 2-11 2-17 AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX Ax AX AX AX AX AX AX AX AX GR GR AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX GR GR GR GR GR GR GR GR GR Calf Calf Yearling Calf Calf Calf Yearling Yearling Calf Calf Yearling Calf Yearling Calf Calf Calf Calf Calf Calf Adult Calf Calf Calf Calf Calf Calf C-;'if Calf Yearling Calf Calf Calf Calf Calf Yearling Calf Calf Calf Adult Calf Calf Calf Calf Calf Calf Calf Calf Yearling Calf Calf Yearling Calf Calf Calf Calf Calf Calf 'Calf Calf Calf Calf Calf Calf Calf Calf -Calf Calf Calf Calf Calf Calf . Adult Calf Yearling Yearling Adult Adult Calf Adult Adult M Unk M F F M F M M F F M M M M F F Remarks F Unk M M F M F M M M M M M M F F F M F F M M Died Release - Maybell RC 148.25 F F F M M F F M F F M M M F F M F M M F F F F M' M F M F M M F F F M M M F M M F .F RC 148.17 =-- RC RC RC RC 148.11 148.10 148.03 148.02 RC 148.08 RC 148.01 .. ..... �APPENDIX A - continued 42 Eartag number Date Trap location 94 95 96 101 102 103 104 127 128 129 130 130 131 131 132 132 133 133 134 134 135 135 136 137 142 143 144 145 146 147 148 149 150 151 152 153 154 155 156 157 158 159 160 161 162 163 164 165 166 167 168 169 170 171 172 173 174 175 176 177 178 179 180 181 182 183 184 185 186 187 188 189 190 191 192 193 194 195 196 197 198 199 200 2-11 2-11 2-11 1-27 1-27 1-27 1-27 1- 5 1- 5 1- 5 1- 5 1-27 1- 5 1-27 1- 5 1-27 1- :s 1-27 1-27 1- 5 1- 5 1-27 1- 5 1- 5 1-12 1-12 1- 9 1- 9 1- 9 1- 9 1- 9 1- 9 12- 9 12- 9 12- 9 12- 9 12-10 12-12 12-12 12-12 12-12 12-12 12-12 12-12 12-12 12-12 12-12 12-12 12-12 12-15 12-15 12-15 12-15 12-15 12-15 12-19 12-19 12-19 12-19 12-19 12-19 12-19 12-19 12-19 12-19 12-19 12-22 12-29 12-29 12-29 12-29 12-30 12-30 1- 5 1- 5 1- 5 1- 5 1- 5 1- 7 1- 7 1- 9 1- 9 1- 9 GR GR GR AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX AX Ase Adult Adult Calf Calf Calf Calf Yearling Calf Calf Calf Calf Adult Calf Adult Adult Adult Calf Adult Adult Adult Calf Adult Calf Calf Adult Yearling Adult Adult Yearling Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult AdultAdult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Sex F F M M M M F F F M F F M F F F F F F F M F M M F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F Remarks RC 148.07 RC 148.00 Elk pen Elk pen RC 148.18-Elk pen Elk pen Elk pen Elk pen RC 148.31-Elk pen Elk ·pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk ~n Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen llk pen Blue 33-Elk pen Elk pen Elk pen Blue 32-Elk' pen Elk pen Elk pen Elk pen Elk pen Elk pen .Elk pen Elk pen Elk pen E.lk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen Elk pen -; Elk pen �Colorado Division of Wildlife Wildlife Research Report July 1987 43 JOB PROGRESS REPORT State of Colorado Project No. 01-03-047 (FW 26 p) ~~~~~~~~~~- Mammals 1 Research Work Plan No. 3 Elk Investigations Job No. 4 Evaluation of Elk Harvest Methodology Period Covered: July 1, 1986 - June 30, 1987 Author: D. J. Freddy Personnel: B. J. T. P. Adrian, R. Bartmann, G. Byrne, L. Carpenter, J. Corey, G. Craig, Ellenberger, B. Gill, V. Graham, J. Gray, B. Hernbrode, Hobbs, T. Lines, J. Madison, T. Pojar, D. Reed, C. Reichert, Schnurr, L. Stevens, J. Toolen ABSTRACT During the 3 rifle hunting seasons in 1986, 118 elk harvested in the White River DAU were checked at the White River check station. Of these elk, 79 were legal bulls (>4-points on 1 antler beam) and none were yearling-bulls. Statewide harvest summaries indicated 10,259 bull elk hunters were in, the White River DAU during all 3 seasons, combined, and harvested 1,498 bulls. , Number of bull hunters increased 40% compared to 1985; but numbers were still depressed and similar to numbers of hunters common to the late 1960s.. The total harvest of bulls was 4 times the 1985 harvest but was similar to harvests during the early 1960s. The 4-point restriction has markedly reduced numbers of hunters and bulls harvested during the first 2 years of implementation:' The reduction in'mortality on bulls has allowed postseason bull:cow ratios to increase from 4":'7 bulls:lOO cows prior to the antler restriction ,to 17, bUlls:lOO cows in 1986. Opinion surveys were completed by. 272 hunters. In general; all hunters were'concerned about by numbers, of. >'4-point bulls in the elk-population, derived more satisfaction from the oppo~tutiity to hunt than from harvesting an elk and supported unlimited numbers'of hunters more than limited permits. However, resident and nonresident hunters differed in some of their viewpoints with nonresidents being more concerned about harvesting. a mature bull. Estimates of wounding loss suggested 0 •.51 spike. bulls were abandoned in the field for every legal bull harvested, an increase from 0.37. spike bulls in 1985. ;." . '._ ~ .': ��45 EVALUATION OF ELK HARVEST METHODOLOGY David J. Freddy P. N. OBJECTIVE To evaluate hunting systems that would promote older-aged structures among bull elk. SEGMENT OBJECTIVES 1. Establish a check station in the White River to monitor impacts of 4-point antler restrictions on harvest of elk, and conduct surveys to monitor hunter opinion. 2. Publish a Division of Wildlife report reviewing and analyzing existing data on the White River elk population. METHODS -AND MATERIALS Check Station A check station was established in the White River at the junction of the White River and Little Beaver roads during the first 4 or 5 days of the 3 combined deer-elk seasons. The check station was open from 10 AM until dark from 11-14 October, 18-22 October, and 1-4 November 1986, Elk were classified as to sex, and aged according to replacement and wear of teeth (Quimby and Gaab 1957). Additionally, an incisor was pulled for dental cementum estimates of age. Greatest number of antler-points/beam, basal circumference at the burr of the largest antler beam, and outside curvature of the longest antler-beam were recorded for each bull elk. Additionally, length of each brow tine (eye-guard) was measured. Hunter opinion was sampled using a survey (Appendix A). Hunters were given the survey and instructed to complete the survey after they had finished elk hunting and return the survey in a self-addressed, stamped envelope. Every other successful hunter and one hunter in every other vehicle having unsuccessful hunters were given surveys. Attempts were made to -sample only 1 hunter from each party of hunters. RESULTS AND DISCUSSION Check Station During all 3 rifle seasons, 118 elk harvested from the White River DAU (GMUs 12, 13, 131, 23, 24, 25, 26, 33, 34) were checked at the White River c~eck station (Table 1). Of these, 79 were legal bulls (> 4-points on 1 antlerbeam) and 47 (59%) were harvested during the first season. In 1985, 21 legal �46 bulls were checked at the same plus one additional check station (Freddy 1986). No legal yearling bulls were checked, which also occurred in 1985. About 10% of the legal bulls checked in 1972, when a 4-point antler restriction was also in effect, were yearlings (Boyd and Lipscomb 1976). Most of the elk checked in 1986 were harvested in GMUs 23 and 24 (97%) and harvested by resident hunters (58%). Nonresident hunters harvested 56% of all adult bulls checked and harvested 64% of the bulls during the first season (Tables 2, 3). One additional cow elk harvested in GMU 11 was also checked but not included in the data summaries for the White River herd. Numbers of hunters passing through the check station were generally low during all seasons with the most and least hunters during the second and third seasons, respectively. Elk were checked primarily on days 3 and 4 of each season (Table 4). Numbers of bull elk hunters in the \Vhite River DAU in 1986 increased to 10,259 compared to 7,316 in 1985 (statewide surveys), but this increase was comparable to numbers of hunters in the late 1960s (Fig. 1). Numbers of bull hunters in any 1 of the 3 rifle seasons in 1986 were <4,700, which was below numbers of hunters in any rifle season since 1960 (Fig. 1). Cow (antlerless) elk hunters numbered 2,016, which was a 12% decline compared to 1985 and similar to numbers of cow elk hunters in the late 1960s (Freddy 1987). , Total harvest of bulls from the White River DAU during the 3 rifle seasons was 1,498 (statewide surveys). This leyel of harvest was similar to harvests of the early 1960s (Fig. 1) but was ,g'reaterthan the 395 bulls harvested during rifle seasons in 1985 (Freddy 1986). Total harvest of antlerless elk during the rifle seasons was 1,078 (statewide surveys) which was similar to harvests during the late 1960s (Freddy 1987). Success of bull hunters was 20%, 13%, and 12% during the first, second, a~d third seasons, respectively, and success of cow hunters was 51% and 60% during the second and third seasons (statewide surveys). ~ Antler-point restrictions have depressed numbers of hunters and bulls harvested for 2 consecutive years. Declines in hunters and harvest also 'occurred in 1971 and 1972 when antler-point restrictions were implemented to protect yearling bulls (Fig. 1). The 3-season structure in 1986 markedly reduced hunters during anyone season. About 70% of the bulls and 46% of the cows coming'through the check station could be aged by tooth replacement and wear and dental cementum (Table 5). Two-year-old bulls comprised 86%,of the bulls that could be aged. Cows were generally <5 years old. As in 1985, numbers of cows that could be aged was low. Hunt~rs having cow permits were mailed tooth envelopes to collect lower incisors from elk they harvested. Unfortunately, tooth envelopes were mailed late from Denver, and hunters received envelopes after the seasons began. A more coordinated state-wide program encouraging hunters to collect incisor teeth from cow elk needs to be implemented. Most bulls checked had a maximum of 5 points on 1 antler-beam as did bulls in 1985 (point> 2.5 em length). Average basal circumference of the thickest antler-beam was 179 ± 3 (SE) rom in 1986 and 173 + 6 (SE) rom in 1985. 'Average outside curvature length of the longest antler-beam was 739 ± 10 (SE) rom in �47 1986 and 712 + 19 (SE) mm in 1985. There were no differences in antlers of all bulls measured in 1985 and 1986 (t-tests P > 0.20; Fig. 2). Antler measurements for bulls of unknown age did not differ from antlers of 2-3-year-old bulls in 1986 (p > 0.14; Fig. 2) suggesting most of the unknown aged bulls were 2 or 3 years old. The data suggest that antlers will not appreciably increase in size until bulls are 4 years old (Fig. 2). The intent of the 4-point antler restriction was to protect yearling bulls. However, concerns have been raised about wounding/illegal loss resulting from hunters hurriedly counting antler-points and, conversely, hunters failing to get a shot at a bull because they were counting antler-points (G. Byrne, Regional Biologist, pers. comm. 1986). Alternatively, the vast majority of yearling bulls would remain protected if legal bulls were defined as bulls having brow tines (eye-guards). Hunters could probably determine the legality of bulls more quickly and reduce the potential of wounding/illegal loss and missing opportunities to harvest legal bulls. If necessary, a minimum size (length) restriction could be used to define a legal brow tine. Both brow tines were measured on 75 bulls at the White River check station in 1986. All bulls had at least one brow tine >100 mm (4 in) in length (Fig. 3). Five bulls (7%) were missing or had broken I-tine. The largest of the 2 brow tines was <150 mm (6 in) for only 1 bull~{l%)(Fig. 3). Thus, if a legal bull was defined as a bull having at least 1 brow tine >100 mm (4 in) in length, all bulls checked would have been legal. Defining-a brow tine as 150 mm (6 in) in length would have made 99% of the bulls legal. Postseason Bull:Cow Ratios Yearly helicopter surveys of elk in January have been used to estimate postseason bull:cow:calf ratios (Freddy 1987). Postseason bull:cow ratios in 1983 and 1984 were 7 and 4 bulls:lOO cows for the White River DAU. Since the implementation of antler-point restrictions in the fall of 1985, postseason ratios have increased to 15 and 17 bulls:lOO cows in 1985 and 1986 •. During all 4 of these years, yearling bulls comprised at least 75% of the bulls observed during postseason surveys (data from Northwest Region files). Hunter Surveys During the 3 rifle seasons; 428 hunters were given a survey, and 272 usable responses were received (64%), including follow-up surveys (Table 6; Appendix B). Initial surveys were returned at a rate of 57%. Follow-up surveys were sent to 65 persons on 12 December 1986, and 33 (51%) were returned •..Completed surveys were accepted until 1 March, 1987, but 84% were received by 8 December 1986. Obtaining the hunter's mailing address at the time he/she was given the survey did not bias the rate at which surveys with and without addresses were returned (X2 2. 0.45, P > 0.40).Of the hunters that returned surveys, i33 (86%) were hunting bulls, 39 (14%) were hunting antlerless elk, 171 (63%) were residents, and 101 (37%) were nonresidents (Table 7). Nonresidents comprised the majority of hunters during the first season while residents dominated the second and third seasons (Table 7). During 1985, most of the hunters during the 2 rifle seasons were residents (Freddy 1986). �43 Success rates for bull hunters passing through the check station were 21% and 34% for residents and nonresidents, and success genera.LLy declined from the first to the third season (Table 7). Success rates for antlerless hunters were 77% and 63% for residents and nonresidents, and success also declined from the second to third seasons (Table 7)~ Antlerless elk were not legal during the first season. Overall, success of hunters coming through the check station was markedly higher than the projected success of all hunters in the White River DAU (statewide surveys), especially during ~he first season. Ex~erience of hunters in hunting elk in the White River differed among seasons (X = 13.4, P < 0.01). Persons hunting for their first time in the \lliite River comprised 41% of all hunters and 48% of the nonresident hunters during the first season (Table 8). During the second and third seasons, 42-47% of the nonresidents were hunting in the White River for their first time while first-year residents comprised only 13-17% of the resident hunters (Table 8). For all seasons, there were more first-time, resident hunters (X2 = 5.2, P < 0.06) and nonresident hunters (X2 = 6.0, P < 0.05) in 1986 than in 1985 (Freddy 1986). These data indicate that some hunters were attracted to the White River in 1986. Hunter Concerns The survey ascertained what concerned elk hunters (Appendix A questions 5-9) by asking hunters to select between pairs of choices that matched 5 items in various combinations: not enough bull elk, including spikes; not enough 4-point or better bull elk; not enough elk, cows and bulls; not enough days in the hunting seasons; and, too many hunters afield. Numbers of >4-point bulls \hereafter referred to as 4-point bulls) were of greatest concern to all hunters (X2 > 10, P < 0.001; Fig. 4, Q5-9). This likely reflects that most of the hunters surveyed were hunting bulls (Table 7). Hunters were more concerned about numbers of hunters afield than numbers of days to hunt (X2 = 32, P < 0.001; Fig. 4, Q9). . Opinions of residents and nonresidents differed little except that residents had equal concerns about numbers of all bulls and numbers of 4-point bulls and were less decisive in selecting between numbers of elk and numbers of 4-point bulls (Fig. 4, Q5 and 7). Opinions of residents and nonresidents were generally consistent among seasons (X2 > 4.2, P > 0.12). However, in choosing between numbers of all bulls and numbers of 4=point bulls (Fig~ 4, Q5), all hunters were less intense in their concerns about 4-point bulls during the second and third seasons, particularly nonresidents (X2 = 8.4, P ~ 0.02). Based on consistency and relative strength of responses, concerns of all hunters were ranked from most to least important as: numbers of 4-point bulls, numbers of all bulls, numbers of elk, numbers of hunters afield, and numbers of days to hunt. These concerns were similarly ranked by hunters surveyed in 1985 (Freddy 1986). . Hunter Satisfaction What constituted a satisfactory elk hunt was determined by asking hunters to choose between pairs of choices that matched 4 items in various combinations: �49 harvest only a 4-point bull; harvest any bull; harvest any elk, cow or bull; and opportunity to hunt (Appendix A questions 10-13). As a group, hunters derived more satisfaction from the opportunity to hunt than from any other choice (X2 > 5.2, P < 0.02, Fig. 4, QlO-12) and found equal satisfaction in harvesting any elk or a 4-point elk (X2 = 0.04, P > 0.50; Fig. 4, Q 13). However, differences between choices of residents and nonresidents were common. In choosing between harvesting a 4-point bull or the opportunity to hunt, residents decisively selected opportunity (X2 = 51, P < 0.001) while nonresidents were undecided (X2 = 1, P > 0.30; Fig. 4 QlO). Residents chose opportunity to hunt over harvesting any bull (Xi = 15, P < 0.001) es~ecially during the second season while nonresidents were again undecided (X = 2, P = 0.15, Fig. 4, Qll). Residents and nonresidents were both less decisive in choosing between harvesting any elk and the opportunity to hunt. Nonresidents chose opportunity to hunt over harvesting any elk (X2 = 3.8, P < 0.05), especially those hunting the first season (Fig. 4, Q12). Residents were undecided and placed nearly equal emphasis on harvesting any elk and opportunity to hunt (X2 = 1.9, P = 0.16; Fig. 4, Q12). Residents and nonresidents had opposing views in choosing between harvesting any elk or a 4-point bull. Residents selected harvesting any elk (X2 = 4.7, P < 0.03), especially those hunting the third season (Fig. 4, Q13). Nonresidents selected harvesting a 4-point bull (X2 = 9.7, P < 0.02), especially those hunting the first season (Fig. 4, Q13). Responses to question 13 seem to highlight fundamental differences in the needs of resident and nonresid~nt hunters and in the needs of persons hunting different seasons. Based on consistency and relative strength of responses, items that provided for a satisfactory hunt for residents and nonresidents were ranked from most to least important: residents--opportunity to hunt, harvest any elk, harvest any bull, harvest 4-point bull; nonresidents--opportunity to hunt, harvest 4point bull, harvest any bull, harvest any elk. Ranking of resident opinion was similar to results in 1985, but nonresidents placed more emphasis on harvesting a bull in 1986 than in 1985 (Freddy 1986). Harvest Systems Hunters chose between pairs of choices that matched 4 harvest systems in various combinations: any bull legal with unlimited numbers of hunters; only 4-point bulls legal with unlimited numbers of hunters; any bull legal with limited numbers of hunters; and only 4-point bulls legal with limited numbers of hunters (Appendix A questions 14-16). As a group, hunters decisively chose 4-~oint bulls with unlimited hunters over anr bull with unlimite~ hunters (X = 116, P < 0.001). This selection was consistent for residents and nonresidents among seasons (X2 .::2.9, P > 0.20; Fig. 4, Q14). Collectively, hunters decisively chose 4-point bulls with unlimited hunters over any bull with limited hunters (X2 = 63, P < 0.001). This selection was consistent among seasons but was less decisive during the third season (X2 = 5.8, P = 0.05; Fig. 4, Q15). Hunters collectively chose any bull legal with limited hunters over only 4-point bulls legal with limited hunters (X2'= 3.2, P < 0.001; Fig. 4, Q16). This selection primarily reflected opinions of residents as nonresidents were undecided between these choices that limited numbers of hunters (X2 = 0.9, P > 0.50). �50 Hunters generally favored harvest systems that maximized their opportunity to hunt (unlimited hunters) over systems that could possibly maximize numbers of older-aged bulls in the herd (antler restrictions plus limited hunters). Hunters made similar selections in 1985 (Freddy 1986). Choices of harvest systems were consistent with the viewpoint that a satisfactory elk hunt is determined primarily by the opportunity to hunt. Opportunity is seemingly more important than hunters' concerns about numbers of older bulls or the need to harvest older bulls. Hunters are willing to "protect" bulls only to the minimal extent of not harvesting yearlings and not by significantly reducing hunting pressure. Future Hunting Viewpoints As a group, hunters will continue to hunt in the White River if antler-point restrictions remain in effect next year (X2 = 120, P < 0.001; Fig 4, Q17) (Appendix A question 17). However, 12% of the residents and 25% of the nonresidents indicated they would not return to the White River. These percentages were not different from 1985 when 16% of the residents and 27% of the nonresidents indicated they would not return if antler-point restrictions continued (X2 ~ 1.34, P > 0.30)(Freddy 1986). The relatively high percentage of nonresidents not returning is surprising because nonresidents placed more emphasis on harvesting older bulls. These frequencies of failure to return may be the common rate of hunter turnover, or they could indicate a true rejection of antler-paint restrictions by some hunters. Collectively, hunters were not supportive of paying increased license fees to hunt mature, 6-point trophy bulls,(X2 = 12.6, P < 0.001; Fig. 4, Q18) (A~pendix A question 18). Nonresidents in particular were against higher fees (X = 11.6, P = 0.001) and although residents were more undecided, they were not supportive of higher fees (X2 = 3.4, P = 0.06; Fig. 4, Q18). License fes in 1986 were $25.00 for residents and $210.00 for nonresidents. The unwillingness by residents to pay higher fees is consistent with them placing higher value on hunting opportunity than on harvesting bulls. Although nonresidents placed more emphasIs on harvesting bulls, they were not willing to pay an even higher fee for hunting trophy bulls. Willingness to pay increased fees did vary somewhat among seasons. Hunters during the third season were the most unwilling supporters of higher fees, while hunters in the first and .second seasons were more undecided and less hostile to increased fees (Fig. 4, Q18). About 40% of the hunters were willing to pay increased license fees to hunt trophy bulls with 75% of these hunters willing to pay no more than twice the cost of a regular bull or cow license (Table ~)(Appendix'A question 19). Wounding Loss Bulls comprised ~65% of the elk reported abandoned in the field each season by hunters completing the survey. Spike bulls were the most frequentiy aban. doned animal. Spike bulls observed/hunter reporting were highest during the second season, and for all seasons, 0.51 spike bulls were abandoned for each reported legal bull harvested in 1986 (Table 10)(Appendix A 'questions 20, 20A). Bulls comprised 50% of the abandoned carcasses in 1985 and on average, 0.37 spike bulls abandoned for each reported legal bull harvested in 1985 (Table 10). The increase for 1986 in numbers of bulls abandoned, percent of abandoned elk that were bulls, and abandoned spike bulls/reporting hunter follows an increase in numbers of hunters in 1986. Illegal harvest of sp'ike bulls increased as numbers of hunters increased when Oregon instituted �51 antler-point restrictions on elk (Harper et al. 1985). Surprisingly, rates of abandoned calves-cows more than doubled in 1986 as ~ompared to 1985 (Table 10). Illegal loss of antlerless elk may increase as more hunters are forced to harvest only the branch-antlered bulls, which are ofte~ among the cows. An additional problem with abandonment of ant.Lerl.eas elk occurred during the first season when wounded antlerless elk could not be legally retrieved because there were no antlerless permits during the first season (Table 10). Bulls comprised <13% of the elk reported abandoned in the field by hunters questioned at check stations in the White River from 1966-68 (Table 11). During these years, any bull could be legally harvested. In 1971, spike bulls (lxl) were not legal and in 1972, legal bulls were restricted to bulls having 4-points on 1 antler-beam. During these 2 years, there were 32 yearling and 2 mature bulls, and 16 yearling and 6 mature bulls, respectively, reported abandoned by hunters queried at check stations in the White River (Boyd and Lipscomb 1976). Assuming no duplicate observations of carcasses, the 40 bulls reported abandoned in 1986 is the highest recorded--3l spike bulls and 9 mature bulls (Table 10). Our understanding of wounding loss associated with antler-point restrictions is, unfortunately, meager. Relying on information gathered at check stations is, at best, tenuous. However, the info~ation thus far obtained suggests a strong need to compare wounding loss in areas where any bull is legal with areas having antler-point restrictions to address the effects not only on bulls, but also on antlerless elk (Harper et ale 1985). Intensive studies involving radiotelemetry are needed to thoroughly address rates of wounding loss with different harvest regulations. Voluntary Comments by Hunters A variety of voluntary comments were received from 155 of the 272 hunters completing the survey (Table 12; AppendiX A question 21). For all hunters, the 3 most frequent comments were: support for the 4-point regulation, limit numbers of hunters, and need for fewer seasons. White River Elk Population Publication The manuscript, "The White River Elk Population: A Perspective, 1960-85" was completed and peer reviewed and is being published as Colorado Division of Wildlife Technical.Report No. 37 to be published in September, 1987. CONCLUSIONS· Antler-point restrictions reduced numbers of hunters and bulls harvested in the White River during 1985 and 1986 to levels common to t:heWhite·River in the 1960s. This reduction in hunting mortality on bulls increased postseason bull:cow ratios but at the expense. of decreased hunter participation. Limited data suggested that wounding loss increased from 0.37 to 0.51 spike bulls/legal bull harvested from 1985 to 1986. The majority of elk abandoned in the field were spike bulls. A commitment to a thorough evaluation of wounding loss is needed to properly determine the degree of wounding loss associated with antler-point regulations. ~ �52 Opinion surveys indicated hunters were: 1) concerned primarily with numbers of bulls in the population, 2) derived more satisfaction from the opportunity to hunt than from harvesting a 4-point bull, and 3) found the 4-point regulation acceptable and more desirable than limited-permit hunting. Nonresident hunters were, however, more concerned with harvesting bulls than were resident hunters. Hunters were not supportive of paying higher license fees to hunt mature, 6-point, trophy bull elk. LITERATURE CITED Boyd, R. J., and J. F. Lipscomb. 1976. An evaluation of yearling bull elkhunting restrictions in Colorado. Wildl. Soc. Bull. 4:3-10. Freddy, D. J. 1986. Evaluation of elk harvest methodology. Game Res. Rep. July(1):73-93. 1987. The White River elk population: Div. Wildl. Tech. Rep. 37 (in press). Colo. Div. Wildl. a perspective, 1960-85. Colo. Harper, J. A., and His Colleagues. 1985. Ecology and management of Roosevelt elk in Oregon, Revised Edition. O're, Dep , Fish and Wildl. Portland. 70pp. Prenzlow, E. J. 1967. Population components White River elk. Fish, and Parks Dep., Game Res. Rep. July:25l-276. Colo. Game 1968. White River elk pop~lation components. Parks Dep., Game Res. Rep. JuiY:383-42l. Colo. Game, Fish, and 1969. White River elk population components. Parks Dep., Game Res. Rep. July:179-234. Colo. Game, Fish, and Quimby, D. C., and J. E. Gaab. 1957. Mandibular dentition as an age indicator in Rocky Mountain elk. J. Wildl. Manage. 21:435-451. �53 Table 1. Age and sex of elk checked at the White River check station during 3 rifle seasons in 1986. All elk were harvested within Game Management Units 12, 23, and 24. Sex Season first (11-15 Oct) second (18-29 Oct) third (1-9 Nov) TOTALS Elk age Bull o Cow Unknown o o o 0 0 0 47 0 0 0 50 calf adult unknown 47 calf adult unknown 27 2 23 o o calf adult unknown o o 1 5 o 1 12 o o 0 0 1 2 35 1 0 calf adult unknown 79 TOTAL 80 (68%) o o 0 37 (31%) 1 (1%) Totals o o 2 o 2 17 o 4 114 o (3%) (97%) (0%) 118 (100%) Table 2. Summary by Game Management Unit of elk checked at the White River check station for all rifle hunting seasons, 1986. Game Management Unit Animal sex 12 23 24 Totals bull cow unknown 4 0 0 26 23 1 50 14 0 80 37 1 50 (43%) 64 (54%) TOTAL 4 (3%) 118 (100%) �54 Table 3. Residency of hunters bringing elk through the White River check station for 3 rifle seasons, 1986. All elk were harvested from Game Management Units 12, 23, and 24. Residency Resident Nonresident Totals Animal sex Season bull first second third ALL 17 16 3 36 30 11 3 44 47 27 6 80 cow first second third ALL 0 22 9 31 0 3 3 6" 0 25 12 37 unknown third 1 0 1 TOTALS ALL 50 (42%) 68 (58%) 118 (100%) Table 4. Numbers of elk checked on each day the White River check station was oEen during 3 rifle seasons, 1986. Day of season and check Season 1 2 3 4 5 first second third 2 4 1 12 7 1 18 15 6 15 11 11 15 TOTAL 7 (6%) 20 (17%) 39 (33%) 38 (32%) 15 (12%) - Totals 47 52 19 118 (100%) �55 Table 5. Ages of_elk checked during all 3 rifle hunting seasons at the \fuite River check station, 1986. Animal Jaw ageb Dental cementum age sex bull cowc unknown Season 0 1 2 3 4 5 6 7-20 unka ~1 or 2:2 Totals first second third ALL 0 0 1 1 0 0 0 28 16 4 48 4 0 1 1 1 0 0 0 0 0 0 0 0 0 0 14 10 0 24 14 10 0 24 47 27 6 80 second third ALL 2 0 3 2" "3 1 0 1 "3 third 1 0 0 0 0" O_ "5 2" 0" 0" 0" 0 3 2 1 1 2 0 0 "3 "3 0" 1 1 2" 15 5 20 15 5 20 25 i2 37 0 0 0 0 0 0 1 aNo tooth available for dental cementum. bElk of unknown age based upon results from dental cementum, but judged in the field as ~2-years old if bulls based on antlers and ~l-year old if cows based on body size. CAntlerless elk legal only during the second and third seasons. Table 6. Numbers of surveys given to and returned by elk hunters during 3 rifle seasons, White River, Colorado, 1986. Initia1 surve~s Fo1low-uE surve~s Sent Sent Total surve~s Returned 15 Returned 9 (60)b Usable first (11-15 Oct) 126 Returned 73 (61)a -second (18-29 Oct) 216 119 (55) 34 16 (47) 135 (63) 132 (61) 92 53 (58) 16 8 (50) 61 (66) 58 (63) 428 245 (57) 65 33 (51) 278 (65) 272 (64) Season third (1-9 Nov) ALL apercent of initial surveys sent. b _ Percent of follow-up surveys sent. 82 (68)a 82(68)a �(.]l 0'1 Table 7. Numbers, residency, and success of elk hunters surveyed.at a check station during 3 rifle seasons in the White River, Colorado! 1986. Numbers of hunters Successful Season Residencl Bull first (11-15 Oct) res nres TOTAL 13 22 'J5 second (18-29 Oct) res nres TOTAL 13 6 19 res nres TOTAL 3 4 res nres TOTAL third (1-9 Nov) ALL a Percent Totals Unsuccessful Bull 23 24 Cow 0 0 Bull 36 46 ""1)" 47 0 82 15 64 26 4 1 77 32 109 19 4 23 3 2 Cow 0 '0 _J: 18 90 "5 "7 9 2 11 3s 5 27 15 42 29 32 61 24 5 29 111 61 172 7 3 10 140 93 233 2411 Cow 0 0 Percent success All 36 (44)a 46 (56) 82 (100) Bull 36 48 43 Cow 0 0 0 96 (73) 36 (27) 132 (100) 17 19 17 79 75 78 12 4 16 39 (67) 19 (33) 58 (100) 11 27 17 75 50 69 31 8 39 171 (63) 101 (37) 272 (100) 21 34 26 77 63 74 0 �57 Table 8. Previous hunting experience in the White River area of elk hunters surveyed at a che~k station during 3 rifle seasons, White River, Colorado, 1986. Years hunted in White River Season ResidencI first (11-15 Oct) res nres TOTAL second (18-29 Oct) Occasiona111 First 19 (53)a 16 (35) 35 (43) 5 (i4) a 8 (17) 13 (16) 12 (33)a 22 (48) 34 (41) res nres TOTAL 69 (72) 15 (42) 84 (64) 11 (11) 4 (11) 15 (11) 16 (17) 17 (47) 33 (25) 96 (100) 36 (100) 132 (100) third (1-9 Nov) res nres TOTAL 29 (76) 11 (58) 40 (70) 4 (11) a (0) 4 (7) 5 (13) 8 (42) 13 (23) 38 (100) 19 (100) 57 (100) ALL res nres 117 (69) 42 (42) 20 (12) 12 (12) 33 (19) 47 (46) 170 (100) 101 (100) TOTAL 159 (59) 32 (12) 80 (29) 271 (100) a Most Totals 36 (100)a 46 (100) 82 (100) Percent Table 9. License fee increases acceptable to those hunters who were willing to pay more to hunt mature trophy 6-point bull elk in the White River based upon returned survels2 1986. Choices Bb CC Dd 1 0 1 2 24 1 1 2 a 2 29" res nres 32 7 5 1 5 0 6 1 48 9 TOTAL 39 6 5 7 57 res nres TOTAL 8 4 12 1 0" 1 0 0 1 0 0 1 10 4 100 res 49 26 7 2 6 0 Season ResidencI first (11-15 Oct) res -nres TOTAL -second (18-29 Oct) third (1-9 Nov) ALL n re s : TOTAL Aa 9 15 7s "9 "6 (75%) (9%) (6%) 9 1 10 (10%) aTwice the price of a regular bull or cow license b Three times the price of a regular bull or cow license c Four times the price of a regular bull or cow license ~ore than four times the price of a regular bull or cow license Totals 1-:3 16 71 29 100 (100%) �U1 0.; Table 10. Numbers and composition of elk reported dead and abandoned in the field during 3 rifle seasons, White River, Colorado, 1985-86. Numbers of abandoned elk and rates of abandonment based upon returned hunter surveys. Number of elk abandoned Season Calves Spike Mature cows bulls bulls Totals Number Spike Spike bulls Ca1f:cow/ hunters bu11s/ /lega1 legal calf: reEortins reEort bulla b cow, 1986 o (O)C 4 (33) 8 (66) second (18-29 Oct) o (0) 8 (27) third (1-9 Nov) 1 (7) ALL 1 (2) first (11-15 Oct) (0) 12 (100) 82 0.10 0.23 > 1.00 18 (60) 4 (13) 30 (100) 131 0.14 0.95 0.44 3 (21) 5 (36) 5 (36) 14 (100) 58 0.09 0.71 0.36 15 (27) 31 (55) 9 (16) 56 (100) 271 0.11 0.51 0.55 0 ------------------------------------------------------------------------------------------------------------1985 " 6 (55) 3 (27) 1 (9) 11 (100) 206 0.02 0.19 0.17 (0) 2 (29) 4 (57) 1 (14) 7 (100) 66 0.06 1. 33 0.18 1 (6) . 8 (44) 7 (39) 2 (11) 18 (100) 272 0.03 0.37 0.17 first (11-22 Oct) 1 (9) second (2-12 Nov) o ALL ~umber of legal bulls harvested by persons ~omp1eting the survey question. ,bNumher of cow or calves harvested by persons completing the survey question. legal during the first season, 1986. cPercent. Ant1er1ess elk not �59 Table 11. Numbers and composition of elk reported dead and abandoned by hunters questioned at check stations in the White River, Colorado, 1966-68. Data from Game Management Units 23 and 24. Hunters Year 1966 1967 1968 Number of elk abandoned questioned 924a 860b 1,153b Bulls 9 (9)c 13 (8) 13 (13) Cows Calves 81 (82) 129 (83) 70 (70) 9 (9) 14 (9) 17 (17) 99 (100) 156 (100) 100 (100) 2,937 35 (10) 280 (79) 40 (11) 355 (100) TOTALS Total aSuccessful and unsuccessful hunters; from Prenzlow 1967 bSuccessful hunters only; from Prenzlow, 1968, 1969 cPercent Table 12. Voluntary comments received from resident and nonresident hunters comJ2letin~ the surve~:, White River, Colorado, 1986. Rank Freguenc;l of resJ20nse Voluntar;l comments Res Nres Support 4-point regulation Limit numbers of hunters Need fewer seasons Need more elk Limit number nonresidents Dislike 4-point regulation Need better access to public land Hard to identify legal bull Concerned about wounding loss Limit vehicle access Need longer seasons Make spike bulls legal Did not like survey Do not limit residents Dislikes 3-season format Need more field officers Likes 3-season format No archery hunting Hunt deer and elk separately Do not hunt cow elk 22 36 7 6 2 10 7 4 0 10 3 6 6 1 2 4 0 5 3 1 1 3 2 1 2 1 2 0 2 0 1 '. 1 0 1 0 1 0 1 0 1 TOTAL RESPONSES 97 58 Total Res 58 13 1211 10 9 7 1 3 2 3 2 6 5 4 4 3 3 2 2 2 1 1 1 1 155 Nres 1 2 3 2 2 3 All 1 2 3 4 5 6 7 8 9 10 10 11 11, 12 12 12 13 13 13 13 ��APPENDIX A 61 OPINIONS OF ELK HUNTERS DURING RIFLE SEASONS WHITE RIVER AREA, COLORADO 1986 Thank you for taking a moment to complete this questio~naire. The Colorado Division of Wildlife values your opinions about elk hunting in Colorado and we hope you have an enjoyable hunt. Please complete ~eth e~q~! ef the questionnaire' after you finish elk hunting this year and then place it in the addressed, stamped envelope and mail promptly. 1. Are you a (circle one): A. Colorado one): license (bull) resident B. non-resident 2. Do you have an (circle A. antlered-only 3. Did you harvest B. antlerless~license an elk (circle one): A. yes B. (cow) no 4. Do you hunt elk in the White River area (circle one): A. most years B. occasionally C. this is the first year In the next 12 questions you will be asked to select each question. Please answer ~~£h question. from a pair of choices presented in 5. In the following pair, what £Q!l£~~!!! you the most as an elk hunter A. not enough bull elk, or B. not enough 4-point'or including spike bulls better bull elk (circle A or you the most as an elk hunter 6. In the +o l l owi nq pai r , what £~E!£~~!!! (circle A or B) : A. not enough days in the hunting season or B. B) : not enough 4:-point or better bull elk 7. In the following pair, what £~!!£~~!l! you the most as an elk hunter or B. not enough 4-point or A. not enough elk, cows better bull elk and bulls, in the herd (circle A or B) : B. In the +o l l ou i nq pair, what concerns -------- you the most as an elk hunter not enough 4-point or better bull elk (circle A or B) : you the most as an elk hunter what £Q!!£~~Q~ B. too many hunters in or the field (circle A or B) : A. too many hunters the field or in 9. In the foIl owi ng pair, B. A. not enough days in the hunti ng season . . ------------------------------------------------------------------------------------------ pair, what is mQ~i imQQ[i~!!iin determining §~ii~f~~iQ[Y elk hunt (circle A or B): 10. In the following A. you harvest only a 4-point or better bull elk or that you have a B. you simply have the opportunity to hunt in determining that you have a 11. In the following pair, what is mQ~t imeQ[t~!lt §~ii§f~fiQrY elk hunt (circle A or B): A. you harvest any bull elk or B. you simply have the (assume spike bulls legal) £Pportunity to hunt '-,:/ 12. In the following pair, what is ~Q~t i~eQrt~ntin determining that you have a §~ti~f~~!Qr~ elk hunt (circle A or Bl: A. you harvest any elk, cow or bull or B. you simply have the (assume spike bulls legal) opportunity to hunt MORE QUESTIONS ~~ ~~~~ �62 13. In the following pair, what is mQ~t tmRQtt~~t in determining that you have a ~~1i~f~f!Qr~ elk hunt (circle A or B): A. you harvest any elk, cow or bull or B. you harvest only a 4(assume spike bulls legal) point or better bull elk 14. In the following pair, which hunting system would you ~~!]! !Q i!!fr~~§~ numbers 4-point or better bull elk (circle A or B) : A. you hunt every year for any bull or B. hunt every year for only (assume spike bulls legal) with 4-point or better bulls unlimited numbers of hunters wi th .un l i e i ted numbers (!]QQ~r!!!H§) of hunters (!]Q 2~rmi.t§) of 15. In the following pair, which hunting system would you ~~!]! 1Q i!]fr~~§~ numbers 4-point or better bull elk (circle A or B) : A. you could hunt every other year or B. hunt every year for only for any bull (assume spike bulls 4-point or better bulls legal) with limited numbers of with unlilllited numbers of hunters (li!!!i.t~Q hunters (!]QQgrmH§) Q~r!!!H§) of In the following pair, which hunting system would you ~~!!! 1Q i!!fr~!§g numbers 4-point or better bulls (circle A or B) : A. you hunt every other year for or B. hunt every 3 to 4 years for any bull (assume spike bull s only 4-point or better legal) with limited numbers of bulls with limited numbers of hunters (!imilgg 2gr!!!i!§) hunters (!imi!gg 2~r!!!i!§) of lb. 17. If antler-point restrictions on bull elk remain in effect for the White River area, will you hunt in the White River area next year (circle A or B): A. yes B. no 18. If large, trophy b-point bull elk were available for harvest in substantial numbers, that is, elk 6 years of age or older, would you be willing to pay an increased license fee to hunt trophy bull elk (circle A or B): A. yes B. no 19. If you answered yes to question 18, how much would you be willing trophy bulls (circle one): A. twice the price of a regular bull or cow license B. three times the price of a regular bull or cow license C four times the price of a reguLar bull or cow license D. more than four times the price of a regular license 20. How many elk did you see that were shot dead this season (circle one number): Q ! £ ~ ~ ~ 20A. How many of these dead, abandoned -calves cows spike bulls 4-2oint or better 21. Your additional comments elk were and abandoned (circle the appropri~te in the fi eld nurabers): NUMBERS ~ bull s to pay to hunt large 1 1 1 1 2 2 2 2 3_ 3 3 3 _ �63 Appendix B. Frequencies of responses by hunters to survey guestions 5-18, 1986. Season Second First Hunter choice Question Hunter choice Third All Hunter choice Hunter choice Residency A B A B- A B A B 5 res nres TOTAL 15 11 26 21 35 56 42 10 52 54 24 78 19 11 30 20 7 27 76 32 108 95 66 161 6 res nres TOTAL 6 7 13 30 39 69 6 4 10 88 30 118 2 3 5 36 15 51 14 14 28 154 84 238 res nres TOTAL 13 10 23 23 36 59 39 10 49 55 24 18 7 25 20 11 31 70 27 97 98 71 169 res nres TOTAL 9 6 15 27 39 66 22 9 31 29 12 41 39 20 94 8 5 13 59 125 76 201 9 res nres TOTAL 13 22 35 23 22 45 17 9 26 68 20 88 11 5 16 22 10 32 41 36 77 113 52 165 10 res nres TOTAL 7 21 28 29 23 52 24 14 38 72 22 94 8 9 17 31 9 40 39 44 83 132 54 186 11 res nres TOTAL 13 20 33 23 24 47 33 14 47 63 22 85 14 8 22 24 10 34 60 42 102 110 56 166 12 res nres TOTAL 14 14 28 22 28 50 43 16 59 53 20 73 19 8 27 19 9 28 76 38 114 94 57 '151 13 res nres TOTAL 21 11 32 14 35 49 47 14 61 47 21 68 30 9 39 9 9 18 98 34 132 70 65 135 14 res nres TOTAL 4 10 14 29 35 64 10 7 17 85 25 110 8 4 12 29 14 43 22 21 43 143 74 217 15 res nres TOTAL 6 6 12 27 37 64 22 12 34 -'71 20 91 13 6 19 26 12 38 41 24 65 124 69 193 16 res nres TOTAL 24 21 45 6 20 26 60 16 76 25 14 39 31 11 42 6 5 11 115 48 163 37 39 76 17 res nres TOTAL 31 37 68 4 8 12 86 24 110 10 12 33 14 47 6 5 11 150 75 225 20 25 45 res nres TOTAL 13 17 30 21 28 49 49 10 59 10 6 16 28 13 41 72 '33 105 96 67 163 7 8 18 79 69 25 22 47 26 73 �64 0 0 0 ->< 20 en a= 16 IIU t- Z 12 ::::» ::c ••• ••• ::::» 8 4 ell 0 0 0 0 -)( 60 65 70 75 80 85 60 65 70 75 80 85 5 4 t- en IIU > 3 a= 2 ~ ::c 1 ••• ••• ::::» ell 0 YEAR Fig. 1. Numbers of bull el k hunters (TOP) and numbers of bull elk harvested (BOTTOM) in the White River DAU, 1960-86. Solid arrows denote periods of antler-point restrictions and the open arrow denotes an increase in license fees. �65 1986 ~ (79) (48) (5) (2) (5 3) (2 4) ct &1.1 c:a I- 4 • (19) • 6 <, en 1985 + t + + . + A. >< 2 ct ~ 0 E E a:: 260 u • 220 •••• ct en ct 180 >< 140 ~ 100 c:a ct + + t - + -- + t + t" E E ~1000 • ...• a:: 1&1 900 •••• I- Z ct >< 800 700 + + t 2 3 ct ~ 600 ALL AGE 4 2-3 OF UNK BULL ALL (YEARS) Fig. 2. Maximum antler length, maximum basal circumference, and maximum points/antler beam of bulls checked at the l-lhiteRiver check stations in 1985 and 1986. Averages bracketed by 957. confidence intervals and sample sizes are in parentheses. �66 N SMALLEST BROWTINE 40 o o 1 2 ~ BROWTINE = 75 LARGEST BROWTINE 4 0 1 2 3 4 LENGTH (mm xlOO) Fig. 3. Frequencies in sizes of smallest and largest browtines (eye-guards) on bull elk checked at the White River check station, 1986. �67 Fig. 4. Responses by hunters to choices A or B in survey questions 5-18 during each of 3 seasons and all seasons combined, White River, Colorado, 1986. See Appendix A for listing of survey questions. Significant differences tM!tween chodces denoted by ••• p ~n.001, •• p ~ 0.02, • P~0.05 for chi-square tests. �68 Q5 ••• bulls 150 • ALL HUNTERS f:] RESIDENTS " B= 4-point bulls o NON-RESI 0' DENTS 100 •• 50 0 2501 Q6 A=days , ••• to hunt 1.11 en B=4-point bulls Z 0 150 Q. en ••• 1.11 ICII:: 100 II. 0 ••• >- 50 u Z 1.11 :::» 0 0 1.11 ICII:: II. 150 and bulls bu lis 100 ' .. 50 o AB AB SEASON AB 1 AB AB AB SEASON 2 HUNTER A B A B SEASON CHOICE A B 3 ABABAB ALL �69 I Q8 1 200 A:hunters_ afield B = 4-point bu lis • ALL [ill RESIDENTS o 150 HUNTERS , ••• NON-RESIDENTS ••• 100 so ••• 0 1SO in season w .n Z 0 a. 100 .n afield ••• •••• D: ••• 0 SO •• >- U Z w 0 :) A=4-point w D: ••• ••• Q10 0 15 bu lis B= opportunity • •• •• AB AB AB SEASON 1 AB AB AB SEASON 2 HUNTER AB AB AB SEASON 3 CHOICE A B· AB A B ALL �70 Q 11 150 A=a ny bu I} • ALL B= opportunity [ill RESIDENTS 100 ••• HUNTERS o NON-RESIDENTS ••• 50 0 ••• '"Z •• 0 150 A. B= '"•••D: 0 pportunity 100 III. 0 >- 50 • u Z ••• ::;) 0 ••• D: III. 0 150 100 50 •• ·0 AB AB SEASON AB 1 AB AB A B SEASON HUNTER 2 AB A B A B SEASON CHOICE 3 AB AB ALL A B �71 200 l Q 14 A=any ••• bull unlimited B= 4-point 1SON bull unlimited • r.7r, ALL l!:!J D RESIDENTS HUNTERS 0 N-R ESID EN T S ••• 100 50 ••• o • 200 • •• permits B=4-point 1&1 bull unlimited '" 150 Z o a. '"0:: 1&1 100 ••• u. o >- 50 •• U Z 1&1 :) o o 1&1 ~ 150 A::.any bull permits B.::.4- poi n t bull per mit s 100 ••• 50 ••• o ABABAB SEASON AB AB 1 SEASON HUNTER AB' 2 AB AB AB SEASON 3 CHOICE AB AB AB ALL �72 250 200 1&1 en 125 Z ••• • ALL til. RESIDENTS 0 NON-REsrDENTS HUNTERS ••• ~100 en 1&1 CII: y" ••• 50 0 >u Z 0 1&1 ::;) o ••• 1&1 CII: y" -- 150 100 50 ••• o AB AB SEASON AB 1· ABABAB SEASON HUNTER 2 ABABAB S'EASON CHOIC E ABA8·AB 3 ALL �73 Wildlife Research Report July 1987 JOB PROGRESS REPORT Colorado State of Project No. 01-03-047 (FW 26 p) Mammals 1 Research Work Plan No. 3 Elk Investigations Job No. 5 Impact of Elk Winter Grazing on Livestock Production Project No. 01-03-048 (FW 26 p) Mammals 2 Research Work Plan No. 9A Job No. 1 Period Covered: July 1, 1986 - June 30, 1987 Author: D. L. Baker, N. T. Hobbs Personnel: G. Bear, M. Miller, L. Carpenter, B. Gill, B. Sheridan, K. Navo, C. Woodward, B. Seely, H. Seely, L. Lovett ABSTRACT We observed the effects of elk grazing during winter on forage production and cattle performance in spring in a sagebrush-grassland community. Twelve experimental pastures were stocked with 4 levels of elk density (0, 8, 15, 30 elk/km2) and 3 replications per level. Elk were allowed to graze pastures from January to April then replaced with 7 cow/calf pairs and 1 first-year heifer per pasture. Cattle grazed pastures for six weeks and were then moved to summer ranges. Grazing by elk resulted in a substantiaL decline in standing crop of live and dead perennial grass at low density (p = 0.26), however, moderate and high densities. caused significant reductions (p = 0.02). -Utilization of live and dead annual grass and forbs did ~ot differ from 0 in any treatment level (p > 0.12). All cattle gained weight during the c~urse of the spring grazing season. We observed no effects of elk grazing on-final weight and average daily gain of cows and calves (p > 0.60). In contrast to cow/calf pairs, weight performance of dry heifers tended to decline with increasing elk density. Heifers in control pastures gained significantly more weight and at a greater rate (p = 0.04) than t;.hosewhere elk density was, highest. , ��- 75 IMPACT OF ELK WINTER GRAZING ON LIVESTOCK PRODUCTION D. L. Baker and N. T. Hobbs -.. P. N. OBJECTIVES l. Assess impact of winter and spring grazing by elk on forage production and cattle performance in spring. 2. Determine optimum stocking densities of elk that are compatible with productive cattle operations in the sagebrush-grassland plant community. 3. Develop and test electric fence design for holding elk. METHODS AND MATERIALS Study Area We wish to make an inference to sagebrush-grassland communities of the mountain valleys and high plains of the western slope of the central Rocky Mountains. Such communities can be found near Montrose, Gunnison, Kremmling, Walden, Meeker, Rangely, and Craig, Colorado. To make that inference,·we conducted experiments on the Little Snake Wildlife Management Area in northwestern Colorado (township 9 north, range 95 west, sections 9, 10). The area is about 35 km (19 mi) north of Maybell, Colorado on.County Road 19. Although this area does not typically contain high concentrations of elk during winter, it is representative of areas that do have those high densities. Topography of the area includes level r!dgetops, rolling hills, and deep gullies, ranging in elevation from 1,800 to 2,000 m (5,900 to 6,600 ft). Aspects are southern and so~thwesterly with an average slope of 15 degrees~ Soils are generally sandy and sandy loam (Table 1). Climate of the area is dry and cold. The growing season averages only 81 days. __Annual mean temperature is 6.06 C (42.9 F). annual precipitation averages 27.5 cm (12.5 in). Table 1. Predominant soil types on the Little Snake Wildlife Management Area. Soil type Glendive loam Rocky River sandy/ loam Zeona loamy sand Unnamed sandy loam Re1sob-Unnamed sandy sandy loam _Zeona-Ryan Park loamy sands Slopes Rooting Depth (cm) . Permeability 0- 3% 3-12% 120 120+ moderate moderate 1- 8% 10-20% 3-15% 100 120+ 120+ high moderate-high moderate-high 2-15% 120+ high �76 Table 2. Pretreatment (May 20-25, 1986) herbage biomass (g/m2) summarized by pasture. Herbs incl ude standi ng 1ive + dead. Shrubs i nclude current growth only.l -----------~------------------------------------------------------------__Mean Pasture N 1 15 Herbage annual grass 4.07 22.32 7.56 33.95 3.39 annual grass perennial grass forbs total herbs shrubs total herbage 8.66 25.25 10.71 44.63 43.91 92.33 4.48 11.02 11.14 20.53 16.07 23.02 annual grass perennial grass' forbs total herbs shrubs total herbage 4.85 35.63 10.70 51.18 41.82 84.55 4.70 19.88 5.52 25.06 12.95 14.95 annual grass perennial grass forbs total herbs shrubs total forage 5.82 21.26 8.36 35.44 43.48 86.47 2.50 . 5.71 3.23 4.82 12.34 14.23 annual grass perennial grass forbs total herbs shrubs total herbage 5.82 20.16 11.22 37.21 33.20 74.61 2.97 4.83 9.15 10.98 12.17 14.03 annual grass perennial grass forbs total herbs shrubs total herbage 3.14 24.96 12.81 40.92 41.40 84.97 2.75 9.93 . 7.61 9.67 22.61 26.07 perennial grass o 2 15 10 3 15 10 4 15 10 5 15 10 6 15 10 Std Dev forbs total herbs shrubs total herbage 7.05 4.60 6.32 �77 Table 2. (cont.) Pretreatment herbage biomass summarized by pasture. PASTURE 7 N Herbage 15 annual grass perennial grass forbs total herbs shrubs total herbage 4.66 . 15.53 8.78 28.97 40.53 83.06 4.21 4.35 6.32 8.99 15.24 17.58 annual grass perennial grass forbs total herbs shrubs total herbage 5.42 16.37 6.55 28.34 28.21 70.51 6.19 5.61 5.15 7.89 11.29 16.14 annual grass perennial grass forbs total herbs shrubs total herbage 5.61 14.63 6.50 26.73 24.43 65.11 3.93 9.04 5.34 12.89 7.87 11.25 annual grass perennial grass forbs total herbs shrubs total herbage 7.30 20.59 7.98 35.88 27.40 69.35 10 8 15 10 9 15 9 10 15 10 11 10 12 15 10 Mean Std Dev 5.66 6.45 5.69 8.49 6.69 9.57 annual grass perennial grass forbs total herbs shrubs total herbage 7.18 .15.74 6.97 29.89 36.34 82.13 4.04 5.56 4.06 6.88 12.65 18.09 . annual grass perennial grass forbs total herbs shrubs total herbage ·6.26 20.78 9.88 36.91 36.35 82.15 3.49 "7.58 6.54 9.77 8.72 12.72 --------------------------------------------------------------------------- 1 Estipates of herbaceous biomass were based on harvested samples from 9plots systematically arranged in 50x50 m grids. Fifteen grids were randomly placed in each pasture. Estiinates of total biomass·and biomass of shrubs were based on double sampling using a herbage meter in 10 grids per pasture with 2 clipped plots and 9 readings per grid: Estimates of total herbage do not represent the sum of herbs and shrubs because of these differences in sampling. 0.25 m 1./ �,.1 : i~: . ~~ ~~: I i i i i i if ! i I, '" I" o .•r- -- . \.0 co CJ) s, " -r VI n::s VI +> ~ ~. <lJ ~~ :- VI 0.. n::s r- <lJ n::s +> s::::: E s; 'r- <lJ 0.. X Q). o 4-. n::s 0.. s, <lJ :::E .~ u, 0"1 "r- t.· �79 Vegetation is dominated by big sagebrush (Artemisia tridentata) with an understory predominated by needle and thread (Stipa Gomata), western wheatgrass (Agropyron smithii), Indian ricegrass (Oryzopis h~enoides), Junegrass (Koleria cristata), and cheatgrass (Bromus tectoru~. Important forbs include wallflower (Erysimum asperum), peppergrass (Lepidium ferfoliatum), silver lupine (Lupinus argenteus), and scarlet globe mallow Sphaeralcea coccinea). Herbage biomass at the approximate midpoint of the spring grazing season (May 20-25) is summarized in Tabl~ 2. The area has not been grazed by livestock during the last 5 years. Condition of the range is good. Experimental Design We observed the effects of elk grazing on cattle performance in a randomized complete block design with 4 levels of elk density (0, 8, 15, and 30 elk/km2) (0, 20, 40, 80 elk/mi2) and 3 replications per level. These levels were chosen to fall above and below the highest density of elk seen on any winter range in Colorado (19 elk/km2) (50 elk/mi2) in Rocky Mountain National Park). There are 3 blocks, each consisting of 4 pastures. Each pasture within a block was stocked with 1 level of elk density such that each block contained all levels. The 12 available pastures were blocked by pretreatment biomass of perennial grasses with the 4 lowest grass biomass pastures forming 1 block and the 4 highest grass biomass pastures forming a_second block, and the remaining 4 pastures serving as the third block. The 4 levels of elk density were randomly assigned to pastures within each block. After the initial randomization, the study design will be continued for 5 years without further randomization (repeated measures design repeating over 5 years). Treatments were imposed as-follows: Twelve 32.4-ha (80-acre) pastures were constructed within a 1.6 km x 2.4 km (1 mi x 1.5 mi) boundary fence (Fig. 1). Pastures were stocked with mature cow elk to achieve densities of 0, 8, 15, and 30 elk/km2 (Table 3). Elk were baited with alfalfa hay and trapped in portable corral traps during December-January. Adult females were f!tted with numbered collars and transported from the trap site (Axial Basin, Godiva Rim) to holding corrals at the Little Snake study site. Here, the elk were conditioned to electric fences for approximately 2-3 days, weighed, tested for serologic evidence of brucellosis and leptospirosis and released into pastures. All other animals were eartagged and freed at the trap site. Table 3. Stocking variables for experimental pastures. --------------------------------------------2-1-------------------------------2 # Elk Per Pasture Density (elk/km ) o o 3 8 15 30 5 10 Stocking Rate (ha/ADM) 9.55 5.74 3.24 ----T--------------------------------------~---------------~------------------20' 20, 40, and 80 elk/mi2• Assuming elk occupy pastures for 3.5 mos and that the ADM equivalent for elk is 3.1 elk per ADM. Rates given = 23.6, 14.2, and 8.0 acres/ADM. - �dO Fifty-four adult females were introduced to pastures between December 16 and January 9 (average pasture occupation date was December 30). Because elk were trapped in small groups during a 3-4 week interval, we were unable to stock all pastures simultaneously. Instead, we had. to accumulate animals in a central holding area and introduce 4 or 5 separate groups of elk into pastures. The number of animals stocked in a group was proportional to the target density of each pasture. In order to equalize grazing pressure in each pasture, elk were released from pastures on 3 different dates; April 14, 21, and 23. Under these conditions the average release date was April 19, 2 days earlier than our target date of April 21. This provided an average grazing period of 110 days. Actual grazing intensity (elk days use) varied according to stocking density (Table 4). Table 4. Grazing intensity and average body weight change (+ SE) of adult female elk released in Little Snake experimental pastures. Empty cells indicate pastures in which elk were released without being reweighed. Treatment (elk/km2) Pasture No.1 Total Elk Days__ Use Initial BW (kg) Mean BW Change (%) ------------------------------------------------------------------------------8 3 4 11 375 311 366 215 (5) 216 (8) 225 (12) 15 6 8 12 527 547 555 222 249 194 (5) (6) (9) 30 2 5 9 1,189 1,184 1,124 203 214 206 (6) (5) (7) - 5.4 * * * -11.2 (1.3) -9.5 (1.0) -11.0 (1.6) ..-7.9 (1.0) -12.9 (1.3) ------------------------------------------------------------------------------1 Pastures 1, 7, and 10 are controls containing no elk. Vegetation Sampling Utilization We estimated utilization of major forage classes by elk during winter and early spring using paired cage procedures (Klingman et ale 1943)(Table 5). Although this method is not without problems (Owensby 1969, Parsons et ale 1984), we know of no reliable q1,lantitativesubstitute. During late' September of year 1, we randomly located 30 0.7-m2, circular plots in each pasture. (For details on procedures for locating'random points in the field, see Appendix B).. Pretreatment sampling revealed that 30 clipped plots should estimate the true mean biomass of perennial grass within at least 10% of the mean, 90% of the time. In the immediate vicinity of each of the 30 randomly chosen plots, we subjectively chose an additional plot to form a duplicate set at each random location. The additional plots wire chosen such that_each �81 Table 5. Sampling regime for estimating standing crop, utilization, production. and --------------------------------------------------------------------------- Plot Time Caged Uncaged place place elk introduced elk removed clip 1 move clip 2 move clip 31 & forage qual ity move cattle introduced mid-season -. clip 4 move clip 5 & forage qual ity move cattle removed clip 6 & forage qual ity Calculations2 production before elk removal = clip 1 (green) forage removed by elk = clip 1 - clip 2 . utilization, dead herbs = (clip 1-- clip 2)/clip 1 (dead) utilization, live herbs = (c}tp 1 - clip 2)/clip-1 (green) spring standing crop = clip 3 . production, early season = clip 4 - clip 31 production, late season = clip 6 - clip 5 --------------------------------------------------------------~------------ 1 Cl ip 3 will be done only if cattle are introduced more than two weeks later than elk are removed. Otherwise, substitute clip 2 for ~lip 3 in all equations. 2 Pasture values for utili zat ion wi II be cal cul ated as the overall .rati0 of forage disappearance/forage available rather than the mean of 30 ratios from individual plots .. �82 resembles the original, random plot in biomass of perennial grasses and forbs. This provided 3G sets of matched pairs of plots within each pasture. Based on random assignment, one member of each pair was caged with a cone of reinforcing wire (mesh size = 13.2 x 13.2 cm, circumference at ground = 811 cm) staked to the ground, and the other marked with a brightly colored wooden stake placed in the ground to 5-cm height. The beating and distance to each stake was recorded. Immediately after the elk were removed in the spring, the caged and uncaged plots were clipped and harvested (for detail on clip and harvest methods, see Appendix C). Contents of these plots were separated by forbs and perennial grasses and by live and dead fractions. We estimated the amount of each category removed by elk as the difference in standing crop biomassl between caged and uncaged plots in the spring (Table 5). We estimated the standing crop of forage available to elk based on contents of caged plots. Utilization was calculated as the amount removed divided by the standing crop. We will repeat this procedure each year. After year 1, we will not rerandomize locations of paired plots but will move them in the immediate vicinity of the original set. Effects of treatment and year on paired plot measurements of utilization were examined with analysis of variance procedures outlined under experimental design. Forage Production We used the methods outlined above t'oestimate the standing crop of live and dead herbs at the beginning of the spring grazing season 2 and to estimate the subsequent production3 of live herbs (Table 5). Caged and uncaged'plots placed in the fall were clipped and harvested immediately after removing the elk from pastures. An additional pair of plots were chosen to mimic the original pair. Based on random assignment, one was caged, the other staked. These were clipped immediately before the cattle were introduced. We repeated this procedure at the midpoint of the spring grazing season (June 1) and at its end (June 28). At the beginning of the spring grazing season we estimated the standing crop of live and dead perennial grasses, annual grasses, and forbs in each pasture from 30 uncaged piots clipped before introducing cattle. Early season production was estimated as the increment in biomass of these categories as reflected by differences in green biomass between caged plots clipped at midseason and uncaged plots clipped before cattle were introduced. Late season production was estimated similarly as the difference between caged plots clipped at the end of the season and uncaged plots clipped at midseason. We examined effects of treatment on forage standing crop, production and utilization using the analysis of variance specified under experimental design. lWe define standing crop as the mass of herbage per unit area at a single poi~t in time. Units are kg/ha. 2we define the spring grazing season as the time interval during which cattle are in experimental pastures. This interval is intended to correspond to the time that cattle normally occupy spring ranges. 3we define production as the rate of addition of new plant biomass to the system. Units are kg/ha/d. We view production as synonymous with yield. '/ �83 Cattle Performance We observed reproductive performance and live weight changes of 7 Hereford cow-calf pairs and 1 nonlactating yearling heifer introduced into each pasture at the beginning of the spring growing season. This provided a stocking density of 2.71 ha/AUM (6.7 acres/AUM). Records on cowage and previous reproductive performance were used to stratify assignments of cows to pastures such that each pastures contained a mix of cows of similar age (4.8 years [+ SE = 0.36]) and reproductive potential. Cows were randomly chosen from each stratum and assigned to treatment groups. After this initial randomization, treatment groups will be maintained for 5 years, excepting replacements. One, 2-year-old cow with her calf will be added to each pasture during year 2 and 4 to replace old-age cows. Replacement cows will be recruited from a group of cattle outside the experiment. Any cow remaining unbred in the fall will be removed from the respective treatment groups and will be replaced with a cow-calf pair during the year she is without calf. However, replacements will be used to maintain stocking rates within the pastures and will not be included in observations of responses to treatments. Instead, we will assign a value of zero to the secondary production of those replaced, open cattle. Open cattle removed from the experiment will be returned the next year if they are pregnant. Measurements on calves were made as follows: Newborn calves and their dams were located each morning and evening. Calves were weighed using an electron balance (± 0.45 kg) and eartagged with numbers corresponding to the cow. In addition to body weight, we recorded birth date, sex, general health, and any problems related to parturition. If necessary, cows were assisted with delivery of calves. Calves showing symptoms of diarrhea or respiratory p~oblems were treated. On May 9, 84 cow/calf pairs and 12 first-year heifers were transported from the Seely Ranch near Pagoda, Colorado, to holding corrals at the Little Snake study site (=80 mi). Here, cows were provided ad libitum quantities of grass hay and water. The following day, all cattle were weighed (+ 0.45 kg) without shrinking and released into pastures. Cattle were allowed to graze pastures until June 18 (41 days) then reweighed without shrink and moved to summer pastures for breeding. In November we will reweigh cattle, wean calves, and pregnancy check each cow. Based on these weights, we will calculate net gain, average daily gain, and economic performance of cattle during the spring grazing season and during the interval from the end of that season to weaning. It is critical that our replication be adequate to allow a reasonable certainty of detecting treatment effects, even small ones. Calf weight gains are one of the most important and the most variable responses we plan to observe. We anticipate a coefficient of variation of roughly 20% for:daily weight gain of calves from 0-8 months of age (Cook 1986). The mean of a random sample of 8 of these calves would have a coefficient of variation of approximately 7%. Assuming 8 cow/calf pairs per replicate, we calculated the number of blocks (replications) in a randomized complete block design needed to achieve significance for treatment effects of different magnitudes (Snedecor and Cochran 1967:ll3-ll4)(Table 6). The needed number of blocks was calculated such that a treatment of a given size would be found significant by a test comparing �84 control and treatment grazing levels 1-8 proportion of the time. (The power of the test with significance level alpha under these conditions is I-B). These sample sizes are approximate for any given year of the study. Thus, it can be seen in Table 6 that for any year, we should be able to detect differences in average daily gain of calves of 20% at an alpha level of 0.10, 90% of the time. Treatment effects averaged over the 5 study years should reduce the number of blocks needed by a factor of 5 (Design effects and repeated measure correlations over time should effectively average out. ,See Appendix A.) It follows that for effects averaged over 5 years, a test at the 5% level with 3 replciations (=14/5) will detect a 10% treatment difference at least 95% of the time. If the response to treatment is linear, then individual levels would have to differ by as little as 3.3% per level to produce a detectable effect. Table 6. Number of blocksl needed to detect specified differences between the control and treatment levels at different values of alpha and l-B in a randomized complete block design. Size of Difference (%) 5 10 20 25 30 alpha=O.lO alpha=0.05 l-B = 0.80 32 9 3 2 2 0.90 0.95 0.80 0.90 0.95 42 11 25 7 35 10 44 12 4 52 14 5 3 3 4 3 4 3 3 2 3 3 2 2 2 lCalculated based on Snedecor and Cochran (1967:113-114). We examined treatment effects and their interactions with analysis of covariance in a randomized complete block design. Pretreatment body weight of cows and calves was used as a concomitant observation in our analysis. The data were analyzed using least squares procedures, and mean separations were performed suing orthogonal contrast (1 d.f.). The ~ priori comparisons were: control vs. all other treatments (inclusive); control vs. low, medium and high (individually) control and low vs. medium and high (inclusive), others·vs. high density. RESULTS AND DISCUSSION Elk Response Stocking experimental pastures with elk at the Little Snake Study Area proceeded on schedule during December and January, 1986-87. Despite exceptionally poor conditions for trapping caused by lack of snow cover, 122 animals were captured and processed at 2 trap sites, Axial Basin and Godiva Rim (Table 7). All adult females tested negative for seriologic evidence of exposure to brucellosis and 5 serovars of leptospirosls, �35 Table 7. Age and sex of animals trapped at Axial Basin and Godiva Rim sites during December 8-January 9. Age Class Adult females Adult males Yearling females Yearling males Calf females Calf males No. Animals 63 o 2 7 23 27 122 lIncludes 6 recaptures and 6 mortalities. After an initial adjustment period to confinement in the pastures (7-10 days), elk acclimated to their surroundings such that normal grazing patterns were established. Human activities in the central hub appeared to cause little disturbance in grazing behavior. During the day, elk were most often observed grazing or bedded in locations farthe~t from the hub, but tracks and overall plant use suggested that grazing activities included most of the pastures •.. Attempts to census elk in the pastures were marginally successful. We tried a variety of techniques. Counts from fixed-wing aircraft were generally unreliable due to variable snow conditions. Elk would often remain bedded, making sightings difficult even with ideal-background conditions. Low elevation flights using a helicopter were successful in moving elk from cover but, in some cases, frightened them into or through electric fences. Ground counts conducted by a single observer were the most successful and least disturbing to the elk. These counts were made eLther by driving a vehicle to several· prominent points on the perimeter where several pastures could be observed at once or by walking slowly into the pastures and counting the elk. Using this technique, we were able to make a complete census of all elk at least once each week. Losses of elk in the pastures during the yinter grazing period were attributed to a number of factors. Eight elk escaped or died in the experiment and had to be replaced during the course of the winter. Two elk died shortly ·after their release into the pastures from injuries incurred during the trapping and weighing process; 1 elk died as a result of entanglement in the electric fence, 3 of apparent malnutrition, and 2 elk apparently escaped since they were never relocated after their introduction into pastures. We experienced limited success in reweighing elk from the pastures at the end of the grazing period. In some instances, 2 people on foot were successful in walking elk out of the pasture and back into the holding corrals for reweighing and release. At other times, groups of 2 to 10 people were unsucessful and efforts resulted in elk being forced into adjacent pastures or injuring themselves in the fence , As a last resort to r-emove.elk, we were forced to cut the perimeter fence and allow elk to leave passively. �86 We were able to reweigh 31 of the 54 elk in the pastures. All elk that were reweighed lost weight during the winter grazing season (Table 4). Weight loss varied from 5 to 13%. Elk in the low density pastures lost the least amount of weight, and those in the medium and high density the greatest; however, the small sample size precludes definitive conclusions regarding the effect of elk density on elk body condition. In general, the solar-powered, electric fence system performed well and was effective in holding elk in captivity and precluding entry of large animals from the outside. Its effectiveness in holding elk in individual pastures should increase as elk are conditioned to electricity for longer periods of time prior to their introduction into the pastures. Last winter some groups of elk were exposed to the electric fence for as little as 24 hrs. This does not appear to be sufficiently long to prevent some animals from attempting to move between pastures or escape from the facility entirely. After initial calibration, the electric fence performed flawlessly most of the winter. Voltages in the perimeter fence and radial pasture fences were checked several times each week. The perimeter fence maintained a constant output of 5.4 kv throughout the winter and spring. Radial fences were less consistent and varied betwen 3.5 and 8.5 kv. To reduce this variability and to provide an emergency backup, we insta~~ed an additional solar panel and energizer for the radial pasture fences. Vegetation Response Effects on Vegetation Production and Utilization We observed substantial declines in the standing crop of green and d~ad perennial grasses in pastures grazed by elk (Table 8). The magnitude of differences between levels tended to decrease at higher densities. Combined effects of treatment, block, and the ungrazed standing crop accounted for 89% of the variation in the standing crop of grazed, dead perennial grass. Effects of treatment and ungrazed standing crop influenced the response of dead perennial grass (Table 8). Effects of block were not significant in the presence of the covariate (p = 0.72), but approached significance in its absence (p = 0.09)(Table 10). We saw no effect of elk grazing on the standing crop of dead perennial grass at low (8 elk/km2, P = 0.26) density; moderate and high densities caused significant reductions (Table 10, P = 0.02).Although the combined model of treatment, block and covariate accounted for 86% of the variation in standing crop of green perennial grass, no single effect was significant (P > 0.10, Table 9). However, individual contrasts showed patterns similar to those for standing dead; control and low density levels offered more green perennial grass than higher levels did (p = 0.04, Tables 8, 10). Patterns in standing crop of perennial grasses were reflected in patterns of their utilization by elk (Table 8). Estimated utilization of green and dead perennial grass in the control pastures did not differ from 0 (p :> 0.26); in all others, utilization was significant (p < 0.02, Table 8). Block, treatment, and the covariate accounted for 82% of the variation in utilization of dead perennial grass; however, only the treatment ef~ct was significant (p = 0.02, Table 9). Individual contrasts revealed that treatment effects began to be detectable at the 15 elk/km2 level (Table 10). �87 Effects of treatment and ungrazed standing crop influence estimates of utilization of green perennial grass (P < 0.03, Table 9). There was no block effect when the covariate was included in the model (p = 0.82), although it approached significance (p = 0.14) when the effect of the ~ovariate was removed. Utilization of green perennial grass in control and low levels differed from utilization in moderate and high_levels (p = 0.03, Tables 8, 10). Utilization of live and dead annual grass and forbs did not differ from 0 in any treatment level (Table 9). Utilization and standing crop of live and dead annual grass and forbs were not influenced by treatment or block; however, the effect of ungrazed standing crop was significant, and models for these responses accounted for a substantial proportion of their variation (76-90%). Consequently, we conclude that treatment effects were absent because use of these forage groups by elk was negligible. �38 Table 8. Least squares meansl for uncaged standing crop biomass of herbaceous vegetation at the beginning of the spring grazing season and overwinter utilization of vegetation by elk. P value corresponds totest of hypothesis that the mean differs from 0.0. Response Standing crop (g/m ) Perennial grass, dead Perennial grass, green Annual grass, dead Annual grass, green Forbs, dead Forbs, green Utilization (%) Perennial grass, dead Perennial grass, green Annual grass, dead Annual grass, green Forbs, dead F-orbs, green Level (elk/km2) Mean 0 8 15 30 0 8 15 30 0 8 15 30 0 8 15 30 0 8 15 30 0 8 15 30 16.7 13.5 11.4 9.5 6.7 6.1 5.7 4.8 0.74 0.34 0.40 0.56 1.15 1.33 1.29 1.15 0.10 0.04 0.13 0.04 3.60 2.35 3.37 2.87 0 8 15 30 0 8 15 30 0 8 15 30 0 8 15 30 0 8 15 30 0 8 15 30 . -0.4 15.0 29.9 45.5 5.5 15.3 . 16.6 31.0 -70.6 26.1 6.1 12:3 -28.5 -9.3 -12.0 12.0 67.7 56.6 65.7 56.3 -20.9 26.9 0.5 13.9 S.E. P > t: HO:LS Mean = 0 1.42 1.66 1.27 1.40 0.45 0.45 0.51 0!4~ 0.12 0.12 0.12 0.12 0.30 0.29 0.30 0.29 0.03 0.03 0.04 0.04 0.36 .0.37 0.39 0.38 0.0001 0.0004 0.0003 0.0011 0.0001 0.0001 0.0001 0lQQQ2 0.0021 0.0409 0.0255 0.0064 0.0137 0.0061 0.0081 0!0111 0.0407 0.3351 0.0186 0.3041 0.0002 0.0014 0.0003 0.0007 8.0 9.3 7.2 7.9 4.3 . 4.3 4.9 4.6 56.9 37.0 39.9 37.7 24.1 22.9 23.6 22.8 12.2 12.6 17.4 ..0.9 16.9 16.9 17.8 17.7·. 0.9611 0.1692 O.0088 ~ 0.0022 0.2584 0.0165 0.0198 0.0011 0.2827 0.5190 0.8858 0.7610 0.2900 0.7012 0.6330 0.6212 0.1135 0.1396 0.1653 0.1048 0 •. 2733 0.1729 0.9798 0.4672 ------------------------------------------------------------------------------lAdjusted for effects of the covariate, ungrazed (caged) standing crop. ~ �89 Table 9. Results of analsyis of covariance1for effects of elk density (level = 0, 8, 15, 30 elk/km2) on the standing crop of uncaged vegetation at the beginning of the spring grazing season and on overwinter utilization by elk. Ungrazed (caged) standing crop was used as a covariate to increase precision in the analysis. Response Standing Crop (g/m ) Perennial grass, dead Effect Mean sq. .error Pr > F Level Block Covar 29.121 19.192 30.762 0.0193 0.0905 0.0520 Perennial grass, green Level Block Covar 2.0756 0.7573 1.9867 0.0730 0.3664 0.1317 Annual grass, dead Level Block Covar 0.1404 0.1534 0.2511 0.1305 0.1202 0.0667 Annual grass, green Level Block Covar -- 0.0573 0.4045 1.4926 0.8773 0.2962 0.0612 Forbs, dead Level Block Covar 0.0069 0.0347 0.0799 0.3032 0.0276 0.0078 Forbs, green Level Block Covar 0.0914 0.0410 7~5477 0.1988 0.4265 0.0075 Level Block Covar 1157.5 .24.5 4.0 0.0248. 0.8544 0.8769 Level Block .Covar 319.9 165.5 557.5 0.0470 0.1408 0.0249 Annual grass, dead Level Block Covar 3033.1 209.1 8807.4 0.5706 0.9494 0.2107 Annual grass, green Level Block Covar 797.1 215.6 5403.9 0.6931 0.8745 0.1223 Forbs, dead Level Block Covar 57.•. 2 4411.5 827.5 0.8671 0.1629 0.3147 Forbs, green Level Block Covar 1243.8 338.5 134.9 Utilization (%) Perennial grass, dead Perennial grass, green ·.0.3362 0.6951 0.7091 ----l------------------------------------------------~---------------~--------Degrees of freedom: Level = 3, Block = 2, Covar = 1 �90 Table 10. Orthogonal linear contrasts (1 d.f.) for effects of level (elk/km2) on the standing crop of vegetation at the beginning of the spring grazing season and on overwinter utilization by elk. o o Response vs. Others 8 Contrast (p > F) vs. 15 30 o & 8 vs. 15 & 30 o & 30 vs. 8 x 15 ------------------------------------------------------~-----------------------Standing Crop Perennial grass, dead Perennial grass, green Annual grass, dead Annual grass, green Forbs, dead Forbs, green 0.0273 0.0295 0.0425 0.0096 0.0161 0.7361 0.0654 0.3718 0.1785 0.0284 0.0437 0.8197 0.0944 0.0720 0.1312 0.3643 .0.6736 0.0758 0.7767 0.5401 0.1407 0.6823 0.2942 0.0602 0.7701 0.5792 0.6772 0.9987 0.3728 0.2240 0.9425 0.6262 0.7123 0.6194 0.7769 0.3701 0.0245 0.3229 0.0419 0.0060 0.0096 0.9946 0.0275 0.1703 0.1607 0.0092 0.0278 0.6536 0.2251 0.2076 0.3476 0.3188 0.5279 0.3451 0.4101 0.6748 0.1372 0.5903 0.6631 0.1027 0.6586 0.9637 0.4278 0.2786 0.5847 0.2133 0.4636 0.9366 0.8164 0.9233 0.9623 0.3792 Utilization Perennial grass, dead Perennial grass, green Annual grass, dead Annual grass, green Forbs, dead Forbs, green Cattle Response Calves A total of 101 calves were born at the Seely Ranch between March 20 and April 28, 1987. Mean parturition date was April 9•. Calving conditions during this period varied from cold and wet in late March and early April to warm and dry for the remainder of the calving season. The majority of calves bqrn were in excellent condition. Later in _the calving season several calves were exposed to a virulent infection of Escherichia coli and had to be treated for diarrhea and associated dehydration and hypoglycemia. Infected calves were lethargic and often refused to nurse from the cow. Sick calves were removed from the cow, placed under a heat lamp in a dry barn, and bottle fed an electrolyteglucose solution until they could be returned to the dam. Nineteen calves showed various degrees of infection and 12 eventually succumbed to the disease. All surviving calves were healthy and symptom free prior to t~eir �91 release into the experimental pastures. This loss of calves, however, prevented us from stocking pastures with 8 cow/calf pairs. Instead, we placed 7 cow/calf pairs in each pasture plus 1 first-year heifer. Performance of heifers was separated from that of lactating cows in our analysis. Birth weight and growth rates of calves were related to sex. heavier at birth and grew faster from birth to 28 days of age calves (P < 0.04). Males weighed an average of 35.5 kg,(SE = and grew at a rate of 0.54 kg (SE = 0.2)/day; birth weight of 33.7 kg (SE = 0.8) and averaged 0.47 kg (SE = 0.03)/day. Male calves were than female 0.9} at birth females was All calves gained weight while in experimental .. pastures, however, we abserved no effects of elk grazing on final weight or ADG of calves (p > 0.25, Tables 11, 12). There was no block effect (p > 0.43). The covariate (pretreatment weight) was significant (P < 0.06) in the model and accounted for 87% of variation in calf final weight and 78% of the variation in average daily gain. Coefficients of variation for both measurements were less than 5%, suggesting that failure to detect differences among treatments was not due to large variances in calf response. Calves gained an average of 29.4 kg (SE = 1.4) during the spring grazing season. �92 Table 11. Least squares means1 ·for final weight and average daily gain of calves, cows and heifers in Little Snake grazing experiment. P value corresponds to test of hypothesis that the mean differs from 0.0. . .. ------------------------------------------------------------------------------- Response Level (e1k/km2) Mean S.E. Ho: P > t LS mean = 0 Calves Final wt. (kg) 0 8 15 30 79.7 77 .8 79.6 75.9 4.20, 4.40 4.40 4.20 0.0001 0.0001 0.0001 0.0001 ADG (kg/day) 0 8 15 30 0.75 0.72 0.80 0.68 0.09 0.09 0.09 0.09 0.0001 0.0001 0.0001 0.0001 Final wt. (kg) 0 8 15 30 468 454 475 46_~ 24.00 23.00 23.00 24.00 0.0001 0.0001 0.0001 0.0001 ADG (kg/day) 0 8 15 30 1.44 1.19 1.59 1.32 0.26 0.26 0.26 0.27 0.0028 0.0059 0.0016 0.0044 Final wt. (kg) 0 8 15 30 244 236 236 231 2.90 3.10 4.00 3.10 0.0001 0.0001 0.0001 0.0001 ADG (kg/day) 0 8· 15 30 1.54 1.35 1.34 1.22 0.07 0.08 0.10 0.08 0.0001 0.0001 0.0002 0.0001 Cow Heifers 1 Adjusted for the effect of the covariate, pretreatment body weight. �93 Table 12. Results of analysis of covariance for effects of elk density (level = 0, 8, 15, 30 elk/km2) on the final weight and average daily gain of calves, cows, and"heifers in the Little Snake grazing experiment. Pretreatment body weight was used as a covariate to increase precision of analysis. Effect Mean Sq. Error Pr > F Final wt. (kg) Level Block Covar 46.80 27.20 637.02 0.460 0.589 0.013 ADG (kg/day) Level Block Covar Response Calves 0.03 0.20 0.1.1 0.256 0.427 0.063 Cow Final wt. (kg) Level Block Covar 1092.30 839.80 341. 60 0.606 0.626 0.666 0.43 0.35 2.51 0.726 0.711 0.167 Cow ADG (kg/day) Level Block Covar --.- Heifer Final wt. (kg) Level Block Covar ·370.20 50.80 11293.90 ADG (kg/day) Level Block Covar 0.24 0.03 0.08 0~155 0.685 .0.0007 0.155 0.685 0.363 �94 Cows All cows gained weight during the spring grazing season (Table 11), however, weight gain was not related to differences in elk density ('P> 0.60, Table 12). We observed no treatment or block effect (p > 0.60) and the covariate (pretreatment body weight) accounted for only 48% ~f the variation in cow final weight and ADG. Although coefficients of variation were acceptable for cow final weights «5%), we observed high variation in cow ADG (C.V. = 32%). Cows gained an average of 53 kg (S.E. = 5) and averaged 1.39 kg (S.E. = 0.08)/day while in the pastures. Heifers In contrast to lactating cows, we observed substantial decreases in weight performance of first-year heifers in pastures grazed by elk (Table 11). Similar to cows and calves, all heifers gained weight; however, weight gain tended to decrease at higher elk densities. Combined effects of treatment, block and covariate accounted for 97% of the variation in heifer final weight and 72% of the variation in ADG. Although average treatment effect was not significant for either measure (p > 0.15, Table 12), individual contrast revealed that treatment effects were detectable at the 30 elk/km level (p = 0.04) (Table 13). Heifers in th~ control pastures were on average 5% heavier and gained weight at a rate that was 21% greater than heifers in the high density elk treatments. Summary of Measurements A number of measurements have been--collected this year but not analyzed and included in this report. Among these are: net primary production and botanical composition and utilization of individual species for each pasture. Additionally, we will complete cattle performance measurements this fall. These data will be collected each year of the study. Supplemental measurements on elk and cattle are planned for future years (Table 14). �95 Table 13. Orthogonal contrasts (1 d.f.) for effect of level (elk/km2) on final weight and average daily gain of calves, cows, .. and heifers in Little Snake grazing experiment. o vs. Others Response Contrast (p > F) o vs. 15 30 8 o o & 8 vs. 15 & 30 & 30 vs. 8 x 15 ------------------------------------------------------------------------------Calf Final wt. (kg) ADG (kg/day) 0.408 0.742 0.466 0.580 0.951 0.444 0.234 0.300 0.685 0.377 0.657 0.284 0.808 0.813 0.394 0.514 0.670 0.701 0.881 0.765 0.447 0.621 0.759 0.986 0.057 0.057 0.173 0.174 0.190 0.190 0.041 0.041 0.146 0.146 0.085 0.085 Cows Final wt. (kg) ADG (kg/day) Heifers Final wt. (kg) ADG (kg/day) Table 14. Summary of progress and future studies planned for elk-cattle grazing experiment, 1987-1991. Year Response Forage utilization by elk Forage production Cattle performance Cattle diet composition and quality Cattle intake Cattle grazing behavior Elk diet composition and quality Elk intake 1987 * * * 1988 * * * 1989 1990- * *'Ie * *'Ie 'Ie )\: )\: 'Ie 'Ie )\: 'Ie 'Ie 1991 * 'Ie ---------------------------------------------~----------------------~---------LITERATURE CITED A.O.A.C. 1980. Official methods of analysis. Analytical Chemists. Washington, D.C. Association of Official Cook, C. W. 1986. Can grazing be studied with statistical validity? .Pages 9-17 in Statistical Analysis and Modeling of Grazing Systems. Proc. 39th Ann. Meeting of Soc. Range Manage. Kissimmee, F~ *'Ie * �96 Klingman, D. L., S. R. Miles, and G. O. Matt. 1943. The cage method for determining consumption and yield of pasture herbage. J. Amer. Soc. Agron. 35:739-746. Owensby, C. E. 1969. Effect of cages on herbage yield in true prairie vegetation. J. Range Manage. 22:131-132. Parsons, A. J., B. Collett, and J. Lewis. 1984. Changes in the structure and physiology of perennial ryegrass sward when released from a continuous stocking management: implications for the use of exclusion cages in continuously stocked swards. Grass and Forage Sci. 39:1-9. Pearson, H. A. 1970. Digestibility trials: in vitro techniques. Pages 120-127 in Range and Wildlife Habitat evaluation. U.S. Dept. Agr. Forest Servo Misc. Pub1. No. 1147. Snedecor, G. W., and W. G. Cochran. 1967. State Univ. Press, Ames. 593pp. Statistical methods. The Iowa Tilley, J. M. A., and R. A. Terry. 1963. A two-stage technique for in vitro digestion of forage crops. J. Brit. Grassl. Soc. 18:401-411. Prepared by9MI.~· Dan L. Baker Wildlife Researcher C N. Thompsod:HObbs Wildlife Researcher C �97 Appendix A Sample Size Considerations David Bowden Statistics Department Colorado State University Assignment of cows to pastures through stratified random sampl ing (stratified by age) should produce a smaller error mean square than that obta ined by strict random samp1ing (age not cons idered in ass ignment) . Measurement of the same cows for the same pasture for five years would be expected to produce a mean with the ~ame standard error as a random sample of 40 cows because of correlations over time through the use of many of the same cows to produce the actual calves measured. time correlation effects should be "counter-balancing. The strat ificat ion and Thus, with regard to Table 3, the effective sample size increase of a factor-of 5 is recommended for testing for effects averaged over five years. .. �98 Appendix Locating Locations are strictly within independent. That on the fa 11 ows. A scale were surveyed will from a random following in the number each the is, specified with of the pastures 0.25 inch and we will choose Once the by navigating points points with and distances. will are pasture each compasses and wi 11 produce Each sampling chosen, as when they pasture. the maps. have no be achieved This within the they will prepared gri d. within on the such that of one point This distributed bearings be chosen point. evenly table. Field the placement wi 11 be plotted field in the will of any other be marked be numbered, to pastures map (1 in = 300 ft) 160 points point located placement will approximately di stances Random Points of points influence B frame bear inqs Poi nts paced and will intervals be �99 Appendix C Procedures We will sampling will for Harvest estimate date place Sample using frames disturbance clipped to stubble the plot each will to forced other samples air ovens in the dead herbs, grasses, annual placed from A.O.A.C following bility frame to forbs, Fort Each plot cause will be over the plane plot. in labelled Collins, they The contents of The contents #6 paper laboratory ·of and sacks, be dried All categories will be weighed assigned be and to the will Five ~omposites randomly in then (live 5g of each category analysis. on five, will of each sack will grasses) approximately based to bags. and perennial pasture 40 cm). = as we perenn i a 1 grasses; and dead categories. sample for so each point, (radius location to be in the in' burlap weighing, each sampling at be will sets of plots. Composites ing After in a composite be prepared eight g. live At each extending be placed at SOC for 48 hrs. into 0.10 will Samples pastures plot. Plants fi e 1d into returned removed ~nd sorted nearest 1 cm). the and Diet within a each within be considei~d samples are methods. placed (about be sorted herbage by a circular vegetation Each category with After defined will of Herbage of and weigh height grasses. and stored biomass be carefully boundaries plot annual clip will minimal the m2 plot a 0.5 and Analysis will (1980), Tilley calculations be analyzed for dry matter, and analyzed and Terry will (1963) for in vitro as modified be corrected for ash, and nitrogen digestible by Pearson ash in sample follow- organic matter (1970). Digesti- res idues (Alexander �100 and McGowan 1966). rumen fistulated We will course laboratory develop the study. runs a regression mean of digestibility obtain inocula for in vitro digestions cow maintained on grass hay. many separate in vitro digestion The frequency to eliminate standards running correct Holstein be conducting of the standardize We will of these variation i n all in vitro between the standard coefficients runs. standard values. of all runs during experiments between requi res that them. We will Following values We will samples each in the use from a this the we include 6 run we will current run, regression in the current run. and to �Colorado Division of Wildlife Wildlife Research Report july 1987 101 JOB PROGRESS REPORT State of Colorado Project No. 01-03-047 (FW 26 p) ------------~----~~-- MammalS 1 Research Work Plan No. 3 Elk Investigations Job No. 6 Effect of Elk Harvest Systems on Elk Breeding Biology Period Covered: July 1, 1986 - June 30, 1987 Author: D. J. Freddy Personnel: D. Hopper, M. Cousins, C. Wetherill, M. Miller, DVM, S. Torbit, Colo. Div. Wildl.; Dr. L. Johnson, DVM, Colo. State Univ.; L. Wyman and Wyman Ranch personnel; E. Ryland and Forbes-Trinchera Ranch personnel; D. Keller and M. Snyder, students, Colo. St. Univ. ABSTRACT Rectal palpation and real-time ultrasound were comparable in detecting pregnancy in elk. Ultrasound has the p~tential advantage, however, to allow quantitative staging of pregnancies: Two blood progesterone analyses, radioimmunoassay and "Heifercheck", accurately predicted pregnancy in elk and mule deer of known reproductive status. The simple-to-use and-less expensive "Heifercheck" commercial pregnancy testing kit appears suitable for field use. Reproductive tracts from elk and mule deer were collected on the ForbesTrinchera Ranch. Pregnancy races for elk were 33% for yearling cows and 93%'" for mature cows while fetal rates were 0.33/yearling cow and 0.96/mature cow. Fetal sex ratios were l7M:6F and 2 sets of twins were observed. Estimated conception dates spanned 34 days from 17 September-23 October and peaked 24-28 September. Pregnancy rates for deer were·lOO% for all ages of adult does, and fetal rates were 1.83 fawns/doe. Fetal sex ratios were 20M:13F, and fawns occurred as 4·singletons, 13 twins, and 1 set of triplets. Estimated·conception dates spanned 15 days from 26 November~lO December and peaked between 2-7 December. .. Most does were in fair body condition having kidney fat indexes of 26-28% and Kistner condition indexes of 53. ' . ��103 EFFECT OF ELK HARVEST SYSTEMS ON ELK _.BREEDING BIOLOGY .David J. Freddy P. N. OBJECTIVE To evaluate effects of harvest systems on breeding biolo~y of elk. SEGMENT OBJECTIVES 1. Select a study area for evaluating effects of harvest systems on breeding biology of elk. 2. Complete a peer reviewed study plan. METHODS AND MATERIALS Harvest Study The study envisioned would have compared breeding behavior and reproductive success in 2 elk populations: one population would primarily have had yearling bulls as breeding bulls while the second population would have had yearling and mature bulls for breeding. To achieve these types of male breeding systems, one population would have been subjected to intensive any-bull hunting, while in the second population, bul~ hunting would have been restricted by using either limited permits and/or antler-point restrictions. Breeding problems resulting from low bull:cow ratios have been strongly considered as the cause of declining calf:cow ratios in several elk herds in Colorado. In spite of this concern and the identification of possible study areas, there was not the needed support from within the CDOW to aggressively pursue this experiment. Thus, a study plan was not developed. Pregnancy Testing Evaluating methods for detecting pregnancy rates and fetal development _in elk and possibly mule deer was seen as a necessary precursor to any similar future harvest studies. Rectal palpation (Greer and Hawkins 1967, Follis 1972), real-time rectal ultrasound (White et al. 1985), and serum progesterone assays (Weber et al. 1982, Wood et al. 1986) were thus evaluated for their potential to detect pregnancy in elk and to a lesser extent, mule deer. Fetal collections of elk and deer were done on the Forbes-Trinchera Ranch to provide known pregnancy status to evaluate. blood assays and to provide fetal data.from herds with high bull:cow arid buck:doe-ratios to compare with fetal data from herds having low male:female ratios. Postseason ratios in 1986 on the ForbesTrinchera Ranch were 40 bulls:lOO cows:50 calves and 52 bucks:lOO does:58 fawns (Forbes-Trinchera Ranch files). Captive semi-tame elk on the Wyman Elk Ranch were used to evaluate rectal palpation, rectal ultrasound, and blood assays. .. �104 RESULTS AND DISCUSSION Pregnancy Testing Wyman Elk On 23 January 1987, 27 female elk owned by the Wyman elk ranch near Craig, Colorado, were pregnancy tested. Elk were held in a cattle squeeze-chute and pregnancy status was determined by rectal palpation and rectal ultrasound. Blood was collected from the jugular vein of each elk, cooled immediately, centrifuged within 24 hrs, and serum frozen. Radioimmunoassays (RIAs) for blood progesterone concentrations and "Heifercheck" (American Diagnostic Sales, Inc.; Westport, CT) pregnancy blood tests were conducted in April, 1987. Serum was thawed in a water bath immediately prior to conducting blood assays. Pregnancy rates were 70% based on rectal palpation and 78% based on rectal ultrasound (Table 1). Differences involved 2, 2-to-4-yr old cows. In these cases, palpation was considered too dificult, but ultrasound indicated the cows were pregnant. The 6 cows determined to be nonpregnant by palpation were also nonpregnant based on ultrasound. Pregnancy rates for broad age classes were 25% for yearlings, 89% for 2-4 yr ol~s, and 80% for cows ~ 5 yrs. The primary criteria indicating pregnancy for both palpation and ultrasound was the presence of placentomes. Fetal development was advanced and fetuses had dropped into the abdominal cavity where they could not be felt by palpation or "seen" by the ultrasound probe. This advanced development negated attempts to stage pregnancies based on fetal images produced by the ultrasound video recorder·system. Once in the squeeze-chute, elk were blindfolded where they stood and did not resist pregnancy testing. There were essentially no problems with either palpation or ultrasound. Copius amounts of carboxymethylcellulose lubricant were used to aid entry into the rectum. About 10 mins/elk were needed to palpate, ultrasound, and collect blood. One cow (3 yrs old) was injured prior to entering the squeeze-chute, and later had to be destroyed. The fetus (male) of this elk weighed 1.33 kg~ had a crown-rump length of 300 mm, hind leg length of 153 mm, hind foot length of 120 mm, estimated age of 122 .days (Morrison et a1. 1959), and an estimated date of conception of 24 September 1986. . The commercial blood test "Heifercheck" accurately determined pregnancy status of 27 (100%) elk (Table 1). Blood from 11 elk was also tested 3 times, and results were consistent across replicates. This evaluation was not a true blind test. However, color densities of control samples (known pregnant and nonpregnant) were quite different, and determining pregnancy status of remaining samples was easy. This easy-to-apply pregnancy test is based on concentrations of progesterone in the blood. RIAs for progesterone concentrations in blood were completed for all elk (Table 1). RIAs for nonpregnant elk averaged 0.30 ~ 0.06 (SE) ng/ml (n = 12, all reps) and were <0.6 ng/ml for all samples with ~range of 0.05 to 0.59. RIAs for pregnant elk averaged 3.28 + 0.22 (SE) ng/ml (n = 30, all reps)-and were >1 ng/ml for all samples with range of 1.15 to 6.14. Replicates a �105 within samples had CVs of 3-42% for nonpregnant and 0.5-25% for pregnant elk. Weber et al. (1982) reported threshold RIA values for elk sampled in March and April as ::.. 3.70 ng/ml for pregnant and '::"1.30ng/ml for nonpregnant elk. In this same study, RIAs for pregnant elk averaged 6.43 + 0.52 (SE) ng/ml (n = 14) with a range of 3.70 to 9.92. These higher values may indicate progesterone levels increase as pregnancy advances. Forbes Elk Reproductive tracts and blood of harvested female elk were collected by hunters on the Forbes-Trinchera Ranch near Alamosa, Colorado, from 29 November5 December 1986. Blood was kept cool for 2-3 days in a refrigerator, and then the serum was collected and frozen. Serum samples varied in condition from little or no hemolysis to substantial hemolysis. "Heifercheck" pregnancy blood tests were conducted on 23 samples on 14 April 1987. Again, this was not a true blind test as investigators were aware of the pregnancy status of most samples. The test correctly determined pregnancy in 21 (91%) samples (Table 2). One "false positive" and one "false negative" were obtained. In the false positive, the cow had an active corpora luteum and was thus probably producing progesterone but also had an infected uterus, which prevented implantation (elk 1116, Table 2). In the false negative, the cow had an active corpora luteum and a fetus weighing 2.9g, which was estimated to be 47 days old (elk #18, Table 2). RIAs for these 2 elk w~re 1.92 and 0.91 ng/ml, respectively (Table 2). Both values corresponded to the lower threshold for RIAs for pregnant elk from the Wyman Ranch (Table 1). Although "Heifercheck" will detect. pregnancies at 20 days in domestic cattle, there may be problems in detecting·early pregnancies in elk. Color densities of the test solutions did not change when read at 2- and 5-min intervals, except in the case of the "false negative", which tended to become a correct "positive" after 5 mins. Forbes Deer Reproductive tracts and blood were collected from mule deer selectively shot by the principal investigator from 16-19 March 1987 on the Forbes-Trinchera Ranch. Blood was collected immediately upon death and cooled. Blood was then centrifuged and serum frozen within 24 hrs of collection. Serum samples varied in condition from little to no hemolysis to substanial hemolysis. "Heifercheck" tests were conducted on 20 samples, 'including 2 male deer, on 14 April 1987. Again, this was not a blind test'. The assay correctly determined pregnancy status of all samples (Table 3). Three replciate tests on blood from 7 deer were all accurate. "Heifercheck." tests of 2 pregnant yearling does (deer #10, 15, Table 3) differed from tests of other pregnant does. A slight blue colo~ was observed. in these 2 test solutions. Normally, test solutions of nonpregnant animals had a strong blue color, while those of.pregnant animals were clear in color. RIAs for these 2 yearlings were 1.86 and 2.26 ng/ml compared to an RIA of 2.55 ng/ml for another pregnant yearling (deer D19, Table 3), which tested correctly in color. RIAs for pregnant does> 2-yrs old averaged 5.16 + 0.55 (SE) . ng/ml (n = 14) with a range of 2.06....;8.93 (Table 3). The low progesterone �106 concentrations in.the 2 yearlings may account for the marginal test results and suggest that detecting pregnancy in yearling does may be subject to error. RIAs from all pregnant does were > 1.86 ng/m1 and were <0.15 ng/m1 for the male deer (Table 3). Wood et a1.-(1986) concluded that RIAs for progesterone > 2.0 ng/m1 indicated pregnancy in mule deer collected from January-April. Conclusions Rectal palpation and rectal ultrasound comparatively detected pregnancy in elk. An advantage of ultrasound should be the ability to quantitatively stage pregnancies by measuring fetal size on the video-output, but this aspect needs further testing. Neither palpation nor ultrasound appeared to significantly stress elk that were restrained within a squeeze-chute. The commercial pregnancy test "Heifercheck" correctly determined pregnancy in 48 of 50 elk and 20 of 20 deer, and was consistent when replicated within elk or deer. The test was not affected by hemolytic blood or by prolonged freezing of blood samples. Cost/animal was about $3.00, compared to $7.00/ animal for an RIA. Unlike RIAs, however,"Heifercheck" does not provide a quantitative measure of progesterone concentrations. "Heifercheck" appears to provide an accurate and inexpensive means to pregnancy test elk and deer with minimal trauma to the animals. Fet~~ Collections Forbes Elk Reproductive tracts from 35 female elk were collected by hunters on the Forbes-Trinchera Ranch from 29 November-5 December 1986.· Hunters were sent instructions via mail prior to the hunting season explaining how to collect the intact uterus and ovaries ~long with a blood sample and a median incisor~ from the elk they harvested. About 43 of 50 permittees actually hunted, and nearly all successful hunters complied with the request for reproductive tracts. Reproductive tracts from 1 calf, 6 yearling, and 28 cows ':_2-yrsold were obtained at check stations by study personnel, usually within a few hours after the elk were harvested. Average age of collected female elk was 5.3 + 0.8 (SE) yrs with a range of 0.5-15 yrs (Appendix A) • .Pregnancy was determined from the presence of fetuses or an active uterus, and rates were 0% for calves, 33% for yearlings, and 93% for cows > 2 yrs old. (Table 4). Uteri from 3 cows were judged to be infected and not capable of supporting an embryo (Dr. L. Johnson, DVM, pers. comm.). Ages of c9wS having an infected.uterus were 1,13, and 15 yrs. The rate of infection was 9% for all cows > 1 yr old, and this deserves further investigation in future years. Because 2-of the infected cows were old, the high incidence rate may be related primarily to age-associated factors. A total of 29 old (contents and 1F and 2F 0.33/year1ing fetuses was observed in reproductive tracts from 33 cows > 1 yr of one uterus were lost). Two sets of twins consisting of 1M were found in cows 9 and 12 yrs old. Fetal rates were cow and 0.96/cow .::. 2 yrs old. Fetal sex was determined for 23 �107 fetuses, of which 17 (74%) were male and 6 (26%) were female. The 5 fetuses of unknown sex were too small in size for development of genitalia, but even if all were female, the sex ratio would still have favored males. Of the 6 female fetuses, 3 occurred in cows < 3 yrs old, and 3 occurred in cows> 9 years old. The sex ratio favoring males may be because of the small sample, but behavioral dominance or nutrition might explain the sex ratio if, indeed, the sample represents the population. C1utton-Brock et a1. (1986) proposed that mature, behaviorally dominant, and physically larger female red deer (often older-aged) produced more male offspring. Doe white-tailed deer nutritionally stressed or bred late in estrus produced more males in-utero (Verme 1965, 1969; Verme and Ozoga 1981), and more male fetuses were associated with nutritionally stressed reindeer cows (Skog1and 1986). Conception dates were estimated from fetal crown-rump measurements (Morrison et a1. 1959). Using 1 December as the median date for collections, conceptions peaked (30.8%) between 24 and 28 September (Fig. 1). About 73% of the conceptions occurred between 19 September and 3 October with 100% of the conceptions occurring in the 34-day interval from 17 September to 23 October. A second peak in conceptions, possibly representing a second estrus cycle, occurred from 14-18 October. In 1965 and 1966, the average conception date for elk in the San.Juan Basin of Colorado based on fetal measurements was 2 October (Boyd and Ryland 1971). Date of conception may be related, in part, to age of cow.· Of the 6 cows bred between 14 and 23 October (latest breeders), 1 was a yearling and 3 were 2 yrs old. Of the 5 cows bred between 19 and 23 September (earliest breeders), 2 were 2 yrs old and 1 was 4 yrs old. All cows > 9 yrs old were bred between 24 September and 3 October (Fig. 1). There were no differences (p > 0.50) in body weight, crown-rump length, and hind foot length between male and female fetuses, although females were smaller for all measurements (Table 5). Fetuses of unknown sex all weighed ~ 4 g and were ~ 35 mm in length. In general, all fetal body dimensions increased as body weight or length increased. The best relationships were crown-rump length vs. body weight and hind foot length vs. crown-rump length (Figs. 2, 3). Conclusions from these initial reproductive collections were: 1) high .overa11 pregnancy rates, pregnancy in yearlings, and a 5% twinning rate suggested that elk received adequate nutrition from seasonal ranges, 2) uterine infections in older and unproductive cows may indicate that more older cows need to be culled, 3) in-utero sex ratios strongly favoring males may indicate "proper" social dominance among older cows or might indicate that nutrition is not optimum, and 4) conception occurred during a 34-day period with about 73% of the cows bred during the first estrus cycle, and younger cows comprised most of the "late" breeders. Forbes Deer Eighteen adult, female mule deer were selectively shot on the Forbes-Trinchera Ranch 16-19 March 1987 to obtain reproductive tracts and indices of body condition. Reproductive tracts were obtained from 3 ~ear1ing and 15 adult does. Average age of does was 4.4 + 0.7 (SE) yrs ranging from 1-14 yrs (Appendix B). �103 All does were pregnant. The 33 fetuses produced fetal rates of 1.83 fawns/doe, 1.33 fawns/yearling doe, and 1.93 fawns/doe> 2 yrs old. Fawns occurred as 4 singletons, 13 twins, and 1 set of triplets. One yearling doe had twins, and the triplets were found in a doe 6 yrs old. Fetal sex ratios were 20M:13F (61% male) overall and 3M:lF for singletons, l6M:lOF for twins, and IM:2F for triplets. Female fetuses predominated in does ~ 6 yrs old. Conception dates were estimated from fetal crown-rump measurements (Hudson and Browman 1959). Using 18 March as the median date of collections, conceptions peaked (66%) between 2 and 7 December (Fig. 4). All conceptions were estimated to have occurred in the IS-day interval from 26 November-lO December. Time of conception did not appear to be related to age of doe. In Middle Park, Colorado, breeding in mule deer peaked from 26-30 November with an interval from 11 November-IS December from 1969-72 (Gill 1972). During these years, postseason buck:doe ratios in Middle Park were 50-55 bucks:lOO does. There were no differences (p > 0.50) in body weight, crown-rump length, and hind foot length between male and female fetuses, although females were lighter in weight (Table 6). Linear relationships occurred between crown-rump length and body weight and between hind foot length and crown-rump length (Figs. 5, 6). That both relationships were linear, unlike the elk, may reflect the low variance in fetal size. Body condition of does was variable ranging form poor to excellent. Kistner body condition indexes averaged 53 + 4 (SE) (Table 7), which placed most does in the fair condition category (Kistner et ale 1980). However, 28% of the does were rated in poor condition, which was somewhat surprising considering snow depths on the Forbes winter ranges are considerably less than in areas such as Middle Park, Colorado. Percent kidney fat was 26-28 (Table 7). Percent kidney fat in adult does from the Piceance Basin of Colorado was 35 and 16 in February and April, 1983 (Torbit et al., In press) and 11-19 for adult does in Middle Park during March 1969-71 (Roper 1972). In summary, high pregnancy and fetal rates indicated deer on the ForbesTrinchera Ranch receive adequate nutrition during fall and winter. However, some does were in poor condition, which may be related to the quality of specific portions of the Forbes winter range. LITERATURE CITED Armstrong, R. A. 1950. Fetal development of northern white-tailed deer. Amer. Mid. Nat. 43:650-666. Boyd, R. J., and E. E. Ryland. 1971. estimated by fetal growth curves. Info. Leaflet 88. 2pp. Breeding dates of Colorado elk as Colo. Div. Game, Fish, and Parks Game Clutton-Brock, T. H., S. D. Albon, and F. F. Guiness. 1986. Great expectations: dominance, breeding success and offspring sex ratios in red deer. Anim. Behav. 34:460-471. �109 Follis, T. B. 1972. Reproduction and hematology of the Cache elk herd. Div. Wildl. Resources Publ. 72-8. l33pp. . Gill, R. B. 1972. Middle Park deer study-productivity and mortality. Div. Wildl. Game Res. Rep. July(2):18l-l98. Utah Colo. Greer, K. R., and W. W. Hawkins. 1967. Determining pregnancy in elk by rectal palpation. J. Wildl. Manage. 31:145-149. Hudson, P., and L. G. Browman. 1959. Embryonic and fetal development of the mule deer. J. Wildl. Manage. 23:295-304. Kistner, T. P., C. E. Trainer, and N. A. Hartmann. 1980. A field technique for evaluating physical condition of deer. Wildl. Soc. Bull. 8:11-16. Morrison, J. A., C. E. Trainer, and P. L. Wright. 1959.· Breeding seasons in elk as determined from known-age embryos. J. Wildl. Manage. 23:27-34. Roper, L. A. 1972. Middle Park deer study-physical characteristics and food habits. Colo. Div. Wildl. Game Res. Rep. July (2):199-209. Skogland, T. 1986. Sex ratio variation ~n relation to maternal condition and parental investment in wild reindeer Rangifer !. tarandus. Oikos 46:417-419. Torbit, S. C., L. H. Carpenter, R. M. Bartmann, and A. W. Alldredge. 1987. Calibration of carcass fat indices in wintering mule deer. J. Wildl. Manage. (In press). Verme, L. J. 1965. Reproduction studies on penned white-tailed deer. Wildl. Manage. 29:74-79. J. 1969. Reproductive patterns of white-tailed deer related to. nutritional plane. J. Wildl. Manage. 33:881-887. , and ---cycle. J. J. Ozoga. 1981. Sex ratio of white-tailed deer and the estrus J. Wildl. Manage. 45:710-715. Weber, B. J., M. L. Wolfe, and G. C. White. 1982. Use of serum progesterone levels to detect pregnancy in elk. J. Wildl. Manage. 46:835-837. White,!. R., A. J. F. Webster,!. A. Wright, and T. K. Whyte. 1985. Realtime ultrasonic scanning in the diagnosis of pregnancy and estimation of gestational age in cattle. The Vet. Record 117:5-8. Wood, A. K., R. E. Short, A. Darling, G. L. Dusek, R. G. Sasser, and C. A. Ruder. 1986. Serum assays for detecting pregnancy in mule and white-tailed deer. J. Wildl. Manage. 50:684-687. �Table 1. Pregnancy status of cow elk on the Wyman elk ranch as determined by rectal palpation, rectal ultrasound 2 and blood assa~s2 23 Januar~ 1987. 7 GR 14 YE 20 GR 25 OR 33 OR 34 OR 2 GR 6GR 11 GR 16 GR 23 OR 24 GR 24 OR 25 GR 26 OR 27 OR 29 OR 29 YE 30 OR 31 OR 32 OR 35 OR 37 OR 43 OR 46 YE 50 BL 52 YE Agea Pa1Eation Ultrasound 2- 3 8-10 2- 3 1 1/2 1 1/2 1 1/2 2- 3 2- 3 3- 4 2- 3 2- 3 3-;-4 3- 4 2 1/2 3- 4 3- 4 3- 4 6- 8 4- 5 2- 3 3- 4 2- 3 1 1/2 4- 5 10-12 6 4- 6 no no no no no .no yes yes yes yes unknown yes yes yes yes yes unknown yes yes yes yes yes yes yes yes yes yes no no no no no no yes yes yes yes yes yes yes yes yes yes yes yes yes yes yes yes yes yes yes yes yes 0 RIA serum Erogesterone ng/m1c Pregnanc~ status Elk no. •......• •......• Blood b no no no no no no yes yes yes yes yes yes yes yes yes yes yes yes yes yes yes yes yes yes yes yes yes ~ - CV(%) ReE 1 ReE 2 x 0.373 0.089 0.594 0.188 0.587 0.052 3.70 4.31 2.83 6.14 4.80 2.79 3.73 5.98 2.60 1.15 2.39 2.47 1.80 1.86 2.79 5.39 2.61 3.47 3.32 3.42 2.80 0.395 0.144 0.572 0.109 0.371 0.096 3.74 3.31 0.384 0.117 0.583 0.149 0.479 0.74 3.72 3.81 4.1 33.2 2.7 37.5 31.9 42.0 0.8 18.6 3.06 3.40 13.9 2.52 2.56 2.2 1.73 2.10 24.9 3.16 5.11 2.63 2.98 5.25 2.62 8.8 3.8 0.5 2.72 3.02 14.0 ------------------------------------------------------------------------------------------------------------aAge estimated from replacement and wear of incisor teeth (years). b"Heifercheck'\ commercial blood test '•. c Radioimmunoassays. �Table 2. Pregnancy status of elk on the Forbes-Trinchera Ranch as determined b:z:fetal collections and blood assals, 29 November-5 December 1986. Pregnancy status b Elk no. a Age 1 2 4 6 7 8 11 12 13 14 15 16 17 18 19 21 26 30 31 32 60 63 65 1 5 1 3 7 3 6 13 1 2 14 15 3 4 2 8 4 1 2 9 2 1 12 @ collections yes yes no yes yes yes yes yes no yes yes no yes yes yes yes yes no yes yes yes no yes single single none single single single single single none single single none single single single single single none single twins single none twins Coq~ora 1uteum Blood Fetal d e 2 mins -_ @ 5 mins Ovar:z:1 Ovar:z:2 yes yes no yes yes yes yes yes no yes yes yes yes no yes yes yes no yes yes yes no yes yes yes no unknown yes yes no yes yesC yes yes yes yes yes yes yes yes no no yes (2?) no no yes (2?) no no no unknown yes no yes no no yes no no yes no no no yes no yes no yes no no yes yes no yes yes yes yes yes no yes yes yes yes maybe yes yes yes no yes yes yes no yes ------------------------------------------------------------------------------aAges > 2 based on dental cementum; ages wear. b"H e~Oferch eck" commerc~-a ° 1 blood test. c . ~ . Corpora 1uteum appeared cystic. dRIA 1.92 ng/m1. eRIA = 0.91 ng/m1. = 1 based on replacement and �Table ,3. Pregnancy status of mule deer on the Forbes-Trinchera Ranch as determined by fetal collections and blood assays. Deer were collected 16-19 March 1987. RIA se rumf Pregnancl status Deer no. 1 2 3 4 5 6 7 Agea. ,8 3 6 6 6 5 3 2 3 1 4 4 14 3 1 1 5 2 1 5 Fetal collections . yes yes yes yes yes yes yes yes yes yes yes yes yes no yes no ', y es ,. yes yes yes twins twins triplets twins twins, twins twins twins single single .. twins twins single none single none twins twins twins t\>tins Blood b Progesterone ng/ml yes yes yes yes yes yes yes yes yes yes yes yes, yes" no yes no yes yes yes yes 3.04 3.38 5.12 2.08 8.48 3.32 3.75 5.77 6.39 1.86 8.93 6.86 2.78 0.10 2.26 0.14 5.32 6.13 2.55 3.64 Corx~ora luteum Ovarl 1 Ovarl 2 yes yes yes (2) yes yes (2) yes yes yes (2) yes no yes no yes (2) yes yes (2) no yes no yes no no yes yes yes (2) yes (2) no 8 9 10 11 12 13 14 (d") yes no 15 16 (<<1') yes yes 17 yes yes 18 ! yes yes 19 yes (2) 20 no ------------------------------------------------------------------------------------------------------------a . . Ages ~ 2 based on dental cementum; ages = 1 based on replacement and wear. b"Heifercheck" commercial blood test. c ,Radioimmunoassays. " . I-' I-' N �113 Table 4. Pregnancy rates for ages of female elk, Forbes-Trinchera Ranch, 1986. A e Calf Samples (N) Pregnant % pregnant Adult Yearling 1 28 26 93 .6 o o 2 33 Table 5. Summary of measurements for elk fetuses of known sex, ForbesTrinchera Ranch, 1986. Hind foot length (mm) Crown-rump length (mm) Bod~ wei~ht (g) Statistics Male Female Male Female Male Female x SD min max n 27.6 13.8 12.2 58.5 17 20.9 7.0 11.9 29.0 6 90.2 16.7 63.5 115.5 17 86.0 9.5 72.0 94.0 5 20.5 4.8 13.5 28.5 17 19.8 2.7 16.5 23.0 5 ------------------------------------------------------------------------------Table 6. Summary of measurements for deer fetuses of known sex, ForbesTrinchera Ranch, 1987. Crown-rump. length (mm) Bod~ weight (~) Hind foot len~th (mm) Statistics Male Female Male Female Male Female x SD min max n 243.9 54.8 164.0 382.0 20 234.2 68.7 156.0 388.0 13 188.1 11.2 175.0 213.0 20 188.4 13.9 171.0 215.0 13 66.8 5.6 60.0 81.0 20 67.1 7.8 56.0 82.0 13 Table 7. Summary of body condition measurements for adult female deer collected on the Forbes-Trinchera Ranch, 1987. x SD min max n Whole body wei~ht (k~) Eviscerated a body -wei~ht (k~) 57.4 7.0 45.0 71.0 18 40.0 5.0 30.0 50.0 18 % kidney fat ---------Ri~ht 26 18 7 62 18 Left 28 22 7 83 18 Kistner fat index 53 18 25 85 18 ------------------------------------------------------------------------,-----aLess only internal organs and fluids. �114 OCT OCT 18 28 OCT 8 SEP SEP SEP 28 18 8 en ~30 ••• A. &II ~ u Z20 o u ••• Z10 &II u ~ &II A. o I I I 35 I I I 45 FE TAL Fig. 1. -- f I 55 AGE 65 75 I 85 (DAYS) Estimated ages and dates of conception for elk fetuses collected 29 November-5 December 1986, Forbes-Trinchera Ranch. �115 . E E CL. :E: ::t a 83 Ct:: I Z 3 0 Ct:: L' 50 Y = 22.1 + 3.7X- 0.04X2 18 " : 1 Fig. 2. 20 ::::: ::::: 28 58 39 BODY WT (91 Relationship between crown-rump length and body weight in elk fetuses collected 29 November-5 December 1986, Forbes-Trinchera Ranch. = HFL •..•.. R :0.98 28.5 r:: 0.98 .282 CRL -4.76 a:I "=22 v :c t~ z 23.5 w _j t- a· O 0 IJ.. 18.5 CI Z t-4 z 13.5 ~ Fig. 3. 63:5 98.2 CROWN-RUMP LENGTH (MM) 80.8 . 115.5 Relationship between hind foot length and body length in elk fetuses collected 29 November-5 December 1986, Forbes-Trinchera Ranch. �116 DEC DEC 16 10 DEC 4 NOV NOV 28 22 40 U) ~30 to- a. 11&1 ~20 0 u to- Z10 11&1 U a.= 11&1 a. 0 Fig. 4. I I 98 104 110 FETAL AGE (DAYS) 116 Estimated ages and dates of conception for deer fetuses collected 16-19 March 1987, Forbes-Trinchera Ranch. �117 = FECRL ~ 21,5.0 z: + .178 FEWT 145.39 r= 0.88 a .••..' J: toL!) m200.3 a a a ...J a a a, z: a ~ Cf 185.7 z ::::: G:) Ct: a o a 171.0 a a a ~~--------~--------~--------~~ 156.0 233.3 310.7 388.0 BODY WEIGHT (G) Fig. 5. Relationship between crown-rump length and body weight in deer fetuses collected 16-19 March 1987, Forbes-Trinchera Ranch • FEHFL = . ro.. 82.0 r = 0.88 ::::: ::::: '.l .466 FECRL -20.74 n==33 a a J: I- z '-' 73.3 UJ ...J I- 0 0 IJ.. 64.7 0, Z 1-4 a J: a 56.0 ~. 171.0 Fig. 6. 185.7 200.3 CROWN-RUMP LENGTH (MM) 215.0 Relationship between hind foot length and crown-rump length in deer fetuses collected 16-19 March 1987, Forbes-Trinchera Ran~h. �119 Appendix A. Reproduct-ive measurements Trinchera Ranch. from female elk harvested 29 November-5 December 1986 on the Forbes- Fetal measurementsb Elk ID nO. 1 2 3 4 5 6 7 8 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 30 31d 32 32Bd 60 61 62 63 64d 65 65Bd 66e Cowa ase Body weisht (s) 1 5 0 1 13 3 7 3 8 6 13 1 2 14 15 3 4 2 2 8 13 7 6 2 4 1 2 9 9 2 11 3 1 1 12 12 2 Crownruml1 (mm) 1.8 27.5 29.0 90.0 12.5 11.9 20.0 12.2 26.5 55.2 16.8 69.5 Hind leg (mID) Ovary wts. (g) Hind foot (mID) AgeC (da::is) Larse 46 66 unk M 3.7 3.7 2.7 1.7 61 M F M M 1.3 3.5 1.2 2.0 3.8 3.5 2.9 3.6 2.9 2.3 2.7 4.4 3.8 2.6 3.7 3.4 3.0 3.2 3.1 4.2 3.4 3.2 2.9 1.3 4.4 3.7 3.7 4.3 3.6 3.2 2.0 4.1 3.5 3.5 2.5 0.5 1.8 2.7 1.5 0.9 2.4 2.0 2.4 1.1 1.9 1.7 1.3 1.8 1.8 1.8 1.5 1.9 1.6 1.1 2.0 1.1 1;1 2.0 2.4 2.0 1.4 2.1 1.5 1.5 1.8 32.0 20.5 93.0 63.5 91.5 115.5 72.0 22.5 32.0 32.0 19.0 28.0 36.5 24.5 13.5 19.5 21.5 13.7 21.0 28.5 16.5 67 57 67 74 61 15.9 23.6 72.0 85.0 24.5 27.0 16.5 19.0 61 65 F M 35.4 2.9 0.9 2.3 4.0 109.5 31.0 18.0 28.0 35.0 33.5 25.0 72· 47 40 46 49 M unk unk unk unk 18.1 38.5 24.0 41.0 78.5 111.5 85.5 109.0 30.5 36.0 26.0 42 •.0 17.0 25.0 20.5 26.5 64 73 65 72 M 2.2 27.9 29.0 17.1 14.5 58.5 33.0 92.5 94.0 80.5 72.0 115.0 32.0 33.0 24.5 23.0 41.0 23.0 23.0 17.5 15.0 28.5 48 67 67 64 61 74 unk F F F M 23.3 21.8 23.4 87.5 84.5 91.0 ---27.0 27.5 28.5 19.0 18.0 19.5 66 65 65 M M F unk c aAge in years for cows ~ 2 yrs old based on dental cementum and for cows ~ lyr cementum or replacement and wear. bSee Armstrong 1950 for description CAge based on Morrison et a1. 1959. dDenotes twins. eFetus lost, cow was pregnant. of measurements. -------Small Sex M M M M M M M old based on dental �120 Appendix B. Ranch. Reproductive measurements from female deer collected 16-19 March 1987 on the Forbes-Trinchera Fetal measurementsb Deer 1D no. Deera aae Body weight (g) CrownrumE (mm) Hind leg 187 8 309 195 187 8 200 193 287 3 209 176 287 3 245 190 387 6 195 173 387 6 156 171 387 6 225 182 487 6 289 210 487 6 259 196 587 6 193 180 587 6 177 175 687 5 195 181 687 5 191 190 787 3 211 181 787 3 207 180 887 2 235 200 887 2 164 175 200 987 3 272 231 1087 183 1 229 1187 4 184 1187 4 235 175 4 250 193 1287 4 283 192 1287 264 1387 14 193 200 1587 1 180 321 201 1787 5 1787 201 5 333 2 215 1887 388 1887 2 382 213 1987 1 275 193 273 192 1987 1 2087 5 170 172 2087 157 175 5 --------aAge in years for does ~ 2 yrs based on dental and wear. (mm) 103 94 86 89 81 79 83 102 101 91 80 81 84 86 87 90 77 102 70 86 84 90 93 97 85 95 95 102 103 91 95 80 73 Ovary wts. (g) Hind foot (mm) 72 71 66 66 60 61 61 71 70 60 59 60 62 64 64 66 60 70 66 66 65 68 70 73 64 74 75 82 81 72 74 58 56 AgeC (dazs) 105 105. 102 102 99 99 99 107 107 100 100 103 103 101 101 103 103 106 102 101 101 105 105 105 101 107 107 110 110 105 105 99 99 ------------------------ bSee Armstrong 1950 for description cementum and for does ~ 1 yr o~ of measurements. CAge based on Hudson and Browman 1959. Sex M F M M F F M M F M F M M M M F M M M M M F M F M F M F M M F F F Left Right 1.10 1.10 1.30 1.30 1.45 1.45 1.45 0.65 0.65 0.80 0.80 0.70 0.70 1.50 1.50 0.60 0.60 0.50 0.70 2.60 2.60 2.00 2.00 1.20 0.50 1.20 1.20 1.30 1.30 0.80 0.80 1.00 1..00 1.25 1.25 0.45 0.45 0.28 0.28 0.28 0.80 0.80 2.00 2.00 0.70 0.70 0.40 0.40 1.40 1.40 0.60 0.60 1.50 1.50 1.15 1.15 1.80 1.00 1.30 1.30 1.50 1.50 0.80 0.80 1.70 1.70 based on replacement � https://cpw.cvlcollections.org/files/original/07e1abe9d35a4bed0c8a5d9cc14bf821.pdf da07bd69e9606bf039ad1760af6cb589 PDF Text Text ~o~oraao Vlvlslon ot Wl~a~lte Wildlife Research Report July 1987 lLl JOB PROGRESS REPORT State of Colorado Project No. 01-03-048 (FW 26 p) --------------~------~- Mammals Research Work Plan No. 1 Multispecies Investigations Job No. 1 Animal and Pen Support Facilities for Big Game Research Period Covered: July 1, 1986 - June 30, 1987 Author: P. H. Neil ABSTRACT Thus far during this fiscal year, 11 elk calves, 11 pronghorn antelope fawns, and 2 bighorn sheep lambs have been successfully hand reared and trained • All were raised on undiluted, canned, evaporated milk. Research on various stress levels in bighorn sheep continued. Seasonal intake investigations with pronghor~antelope also continued and are still in progress. The waterfowl holOing aQd research facility was completed to include an incubation and storage room. Seven elk suffered rattlesnake bites in late June-early July, but have recovered. All animals at the facility are presently healthy. ��123 ANIMAL AND PEN SUPPORT FACILITIES FOR BIG GAME RESEARCH Paul H. Neil P. N. OBJECTIVES To provide and maintain populations of captive animals and pen facilities to support Mammal and Avian Research programs. SEGML~T OBJECTIVES 1. Continue to develop and maintain facilities at Foothills Wildlife Research Station. 2. Coordinate rearing, training, and research activities with captive wild and tame big game animals for all big game cervid and noncervid research projects. 3. Integrate big game animals and physical plant support facilities, personnel, and fiscal resources into a single budget. METHODS AND MATERl:J...S Routine neonate rearing procedures were used to hand-rea:" 11 elk calves during the fiscal year. Eleven pronghorn antelope fawns and 2 bighorn sheep lambs are presently being hand-reared and trained for future nutritional studies. All animals are being raised on undiluted, canned, evaporated milk. Research on various stress levels in bighorn sheep continued throughout the fiscal year with studies being conducted in isolation pens and digestion cages during the fall, winter, and spring months. Seasonal intake investigations also continued with 6 adult female and 5 adult, castrate, male pronghorn. Daily intake and weekly animal weight records have been maintained throughout the fiscal year. Several major rebuilding projects began during the year. Fifteen isolation pens were rebuilt to include new feed boxes. The main alleyway connecting the pens on the east side of the facility is presently being torn down and replaced due to age. Routine maintenance cont~~~ed, and a considerable amount of repairs were done as a result of animal aD~ ~ind damage. RESULTS AND DISCUSSION Twelve female elk calves were captured from Rocky Mountain National Park and transported to the Foothills :acility to be hand-reared. One of the calves died from unknown causes. On~ bighorn ewe had to be euthanized as a result of a broken leg during the winter, and 2 bighorn rams died from pneumonia. We �124 originally had 13 pronghorn antelope fawns. One was an orphan, 3 were captured from northeastern Colorado, and 9 were born at the facility from our captive does. Two of the fawns died at early ages. Research being conducted on the effects of various levels of stress in bighorn sheep has included several approaches and designs and is explained under work Plan 2, Job 4 (Mike Miller and Tom Hobbs, Principal Investigators). Seasonal intake measurements were taken using 6 adult female and 5 adult, castrate, male pronghorn antelope. Weekly animal weights were recorded throughout the fiscal year. Progress and results are reported under Work Plan 1, Job 4 (Bruce Gill, Principal Investigator). R~building many of the structures on the east half of the facility was necessary due to aging and weather damage. Some of the construction is still in progress. - - All animals 'at the facility are presently healthy. Seven elk suffered rattlesnake bites in late June-early July but have recovered. The big game research herd presently consists of 11 elk, 2 Rocky Mountain goats, 16 bighorn sheep, 22 pronghorn, and 1 domestic cow. Other activities at the Foothills Wildlife Research Station during the year consisted of educational tours for wildlife personnel from various state and federal agencies, Colorado State University personnel, local Boy/Girl Scout Troops, and foreign visitors. Prepared by ~~~-(L=>'C Paul H. Neil Wildlife Technician III �~o~oraao Ulv~~~un u~ w~~u~~~~ Wildlife Research Report July 1987 125 JOB PROGRESS REPORT State of Colorado Project No. 01-03-048 (FW 26 p) Mammals --~~~~~--~--~--~- 2 Research Work Plan No. 1 Multispecies Investigations Job No. 3 Mammals 2 Research - Research Administration Period Covered: July 1, 1986 - June 30, 1987 Author: R. B. Gill Personnel: .L. E. Lovett c ABSTRACT - As a prelude to developing a Research Project Selection process, considerable time was spent in assisting in the development of a comprehensive planned management system for the Colorado Division of Wildlife. A draft Strategic Plan was ~eveloped along with a process for assigning planning and evaluation responsibilities for the program planning process. ��127 MAMMALS 2 RESEARCH RESEARCH ADMINISTRATION R. Bruce Gill P. N. OBJECTIVE To administer research within the Mammals 2 Research Unit for the highest productivity at the lowest cost. SEGMENT OBJECTIVE Administer and evaluate accomplishments of Mammals 2 Research. RESULTS AND DISCUSSION In the past the Research Sections have been criticized internally because allegedly they have not addressed issues or questions of priority. To obviate that criticism, Research Leaders were assigned the task of developing a research problem-selection process which assured that the best research projects were selected which dealt with organizational issues or problems of the highest priority. It was concluded that the only appropriate vehicle to identify researchable issues or problems of highest priority was the planning process by which the Division of Wildlife decides priority issues and projects for every other Division organizational unit. Such a system, if it was comprehensive, would not only identify priority projects for each organizational unit, but would also clearly establish the linkage among projects across organizational units. In short, a com~rehensive planning system would not only establish priority projects, it ",·c)t,ld also clearly indicate why they were priority and how each organizational UL<t's work was linked to that of other units to accomplish the totality of the Division's business. The problem for the Research Leaders in developing research plans that were integrally linked to the Division's comprehensive planning process and comprehensive management plan was that the Division had neither a comprehensive process nor a comprehensive plan. As a result, most of my administrative efforts during FY 1986-87 were devoted ~owards developing a comprehensive management plan. A draft copy of that plan is attached as Appendix A. The planning process calls for the development of a strategic plan which addresses issues, goals, and objectives at the statewide level. From the strategic pJan, program plans will be developed which outline issues, goals, and objectives at the programmatic level. From the program plans, operation~ ~lans will be developed which describe the goals and objectives each o'rg e..i , zational unit will address in order to contribute towards the objectives c! the program plans. Lastly, work plans will be developed for cost centers and individuals which describe and assign the projects necessary to the accomplishment of the operations plans objectives. Following the approval and implementation of the strategic plan, program plans will be developed, followed by operations plans. It is during the operations planning phase that Research Leaders will develop research operations plans �126 which will identify priority researchable problems or issues, list the research projects which will be required to resolve those problems/issues, and estimate costs of those projects. Candidate issues are: 1. The Division has the perception that wildlife habitats, including big game habitats, are diminishing in both quality and quantity, yet the Division has no reliable habitat evaluation system which quantifies habitat abundance and rates habitat quality in ways that predict animal population peformance. 2. It is believed that plant successional changes and/or abusive use by wildlife and domestic livestock has caused a decline in wildlife habitat quality. 3. Ther,e seems to be a paradox among wildlife recreationists. Interest in wildlife seems to be increasing while support for the Division of Wildlife as the steward of the wildlife resource seems to be decreasing. 4. Responses of single species populations to harvest systems are not well known, and reponses of several species within trophic levels or species among trophic levels are largely unknown. 5. Wildlife production and agricultural production are in conflict in many areas of the state because of a perception that wildlife and domestic livestock compete for common forages within common ranges. 6. A vocal, politically influential minority of the general public believes that hunting and trapping are unethical and immoral because they cause unnecessary animal suffering. 7. As Colorado's population becomes larger and more urbanized, an increasing segment of that urban population will have little experience with and knowledge of wildlife and, hence, little interest in wildlife issues. 8. Some species of Colorado wildlife are either extinct, becoming less numerous, or their status is not known. 9. Demand for various kinds of wildlife recreation and willingness to pay for it are too poorly quantified to make reliable predictions of trends in that demand or to confidently structure programs to meet that demand. ( Prepared by~ ~ , ~ J) R. ~ruce Gill Wildlife Research Leader �129 APPENDIX A * * * PRELIMINARY DRAFT * * * TODAY'S STRATEGY ••• TOMORROW'S WILDLIFE A COMPREHENSIVE MANAGEMENT PLAN FOR COLORADO'S WILDLIFE DRAFT COLORADO DIVISION OF WILDLIFE JULY 8, 1987 t--r ,~'-r * * * \_I A- L.-l I) - FOR DISCUSSION PURPOSES ONLY * * * ��.. 131 DRAFT I" COMPREHENSIVE MANAGEMENT PLAN TABLE OF CONTENTS 1. Introduction Purpose of this document How the Division of Wildlife Organization of the Division 2. Th~.Division's Mission plans for the future of Wildlife and Major Program Areas 3. Major Issues Facing the Division of Wildlife Demographic Changes Population growth/ encroachment on habitat Changes in kinds of recr~ation in demand Urbanization of population Dow Financial Dependence on Deer & Elk Hunting Rising Costs of Providing Wildlife Recreation Opportunities Conflict Between Resident and Nonresident Sportsmen Wildlife-Related Conflicts Growth of the Division of Wildlife Field Presence Required to Manage Statewide Resource Management Technology Limitations 4. Major Objectives and Strategies General Hunting Recreation Fishing Recreation Watchable Wildlife Species Conservation Statistical Summary Appendices List of species Current Program Status for Achieving Them ��133 I' DRAFT INTRODUCTION Wildlife is one of Colorado's most important assets. The State is home to over 500 species of terrestrial wildlife and 300 species of aquatic wildlife. Sportsmen spend over one billion dollars each year in Colorado in connection wi th hunting and fishing, and the State's wildlife resource contributes heavily to tourism, the State'~ second largest industry. Because the wildlife resources of the state are so important, it is vital that they be soundly managed. Sound management in a complex and changing world is the result of sound planning. To manage properly we must plan ahead. This means understanding where we are headed, where we want to go, and how to get there. Planning is the process that leads to such an understanding. The Calora~o Division of Wildlife began formal long range planning almost two decades ago. Its first five-year Comprehensive Management Plan was published in 1974, and subsequent editions were published in 1977 and 1983. This document is the fourth in the Comprehensi ve Management Plan series. TABLES , . COLORADO'S WILDLIFE Class or Phylum Number of Species 000 000 000 WILDLIFE RECREATION Activity Hunting Fishing Wildlife IN COLORADO Recreation Watching 000,000 000,000 000,000 -1- Days Sportsmen Expenditures $000,000 000,000 000,000 ,- �134 DRAFT PURPOSE OF THIS DOCUMENT Planning in the Colorado Division of Wildife, as in many organizations, is a complex process. Many different kinds of plans - some long range, some short range, some addressing a single issue, some comprehensive in scope - are utilized. Some are updated annually through the Division's formal planning and budgeting process, while others are prepared and/or updated on an ~ hoc basis as circumstances require. Because of this, the public often finds it difficult to understand exactly how the Division intends to manage Colorado's wildlife resources. Our various constituents may not be aware of how, when, and to whom to express thei r vi ews on Di vision plans. The Comprehensi ve Management Plan is intended to address these problems. It provides a summary snapshot of the key elements of all the various Division plans at a specific po i n t in time. It presents our major long term goals and objectives and describes in overall terms how the we intend to manage the wildlife resources of the State. By presenting such a summary view in a single document, public discussion and input can be facilitated. The Division encourages extensive public discussion and comment on the proposals set forth in this Comprehensive Management Plan. -2- �135 ,. DRAFT HOW THE DIVISION OF WILDLIFE PLANS FOR THE FUTURE The Comprehensive Management Plan presents a synopsis of the key points contained in the many different plans in existence throughout the Division. To understand how the Comprehensive Plan was prepared, it helps to understand something of the various planning processes employed within the Division. Commission role Basic Framework Strategic Program . Operations 'Work Satellite plans Comprehensive Management Plan -3- �136 DRAFT ORGANIZATION OF THE DIVISION OF WILDLIFE -4-- �'I DIVISION OF WILDLIFE DIRECTOR'S OFFICE AQUATIC WILDLIFE SECTION TERRESTRIAL WILDLIFE SECTION HABITAT RESOURCES SECTION PUBLIC SERVICES SECTION ADMINISTRA'- :VE & TECHNICAL SERVICES SECTION NORTHWEST REGION NORTHEAST I I I REGION SOUTHWEST SOUTHEAST REGION REGION CENTRAL ~I--' LV REGION ) .- ~ ;4, -....,J �-6- �1::;3 STRUCTURE According is to: to state DRAFT MISSION AND OF MAJOR PROGRAMS statute, the Colorado Di vision of Wildlife Protect, preserve, enhance, and manage the wildlife Colorado and thei r environment for the use, benef it, enjoyment of the people of this state and its visitors. of and and to: '.' Offer the greatest possible variety of wildlife-related recreational opportuni ty to the people of this state and its visitors. In other words, the Division is in the business of both wildlife recreation and wildlife conservation. The mission of the Division is to perpetuate the fish and wildlife resources of Colorado and to provide the opportunity for people to enjoy them. The Division intends to accomplish its mission efforts on four major program areas: Hunting recreation Fishing recreation Nonconsumptive recreation Species (Wildlife by focusing its Watching) conservation Through its recreation programs the Division seeks to provide a wide diversity of recreational opportunities throughout the State. The goal of these programs is to enhance wildlife recreation in Colorado. Consequently, the primary objectives of these programs are stated in terms that measure the value the Di vision adds to wildlife recreation in the State. They will specify the level of recreational value which the Di vision is attempting· to .produce in three separate areas hunting, fishing, and wildlife watching. Through its conservation program the Division seeks to preserve a rich wildlife heritage for future generations. The goal of this program is to enhance the health and diversity of wildlife populations in the State. Consequently, the primary objectives of the conservation program are stated in terms that measure the health, abundance, and diversity of wildlife populations in the State. -7- �140 DRAFT By establishing these as major program areas, diverse activities undertaken for common purposes can be grouped together to aid in planning and decision-making. For example, stocking fish, acquiring access to streams, and maintaining public facilities on streams are different activities all undertaken for the common purpose of serving anglers and enhancing fishing recreation. At some point in the planning process these activities must be considered together, and trade-ofis made between them. The level of stocking required is going to be influenced by the quantity of waters open to fishing, and the quantity of waters open to fishing is going to influence the investment required to develop, maintain, and. operate public facilities. In order to intelligently decide how to make these trade-offs, decision-makers must have a clear idea of the overriding purpose behind each of the activities, and how they are related to one another in the pursuit of the Division's overall goals. Creating these four program areas and establishing objectives for each provides such a mechanism. The programs provide a logical basis· for evaluating different options and judging which will best serve the Division. The citeria for evaluating different alternatives is how well and how efficiently they contribute to the overall program objectives. Of course, it is no simple matter to categorize the many things the Di vision dce s according to which program objecti ves they serve. Some activities serve more than one purpose (i.e., they contribute to the objectives in more than one program area). For example, in order to meet the demand for elk hunting, the •.Di vision must insure that there are large, heal thy, stable populations of elk in the state. The numerous activities in which the Division engages for the purpose of maintaining elk populations can thus be regarded as part of the hunting recreation program. But in managing elk populations to meet hunting demand the goals of the conservation program are also served. In reali ty most things the Division does affect more than one of the four ma jor program areas. There are no sharp lines dividing the programs from each other, nor need there be. For planning and decision-making purposes, the programs are viewed as forming a hierarchy, with fishing and hunting recreation at the top, wa tchable wi ldl ife in the middle, and species conservation at the bottom. All acti vi ties which are undertaken for the purpose of meeting the objectives of the hunting and fishing recreation programs are considered to be part of those programs. The watchable wildlife program deals only with those activities, in addition to the activities making up the hunting and fishing programs, necessary to accompl ish the objectives of the watchable wi Idl ife prog ram. -8- �ILl" .1 DRAFT Similarly, the species conservation program deals only with those activities which are necessary, over and above the things done to serve the recreation programs, in order to meet the objectives of the species conservation program. -9- �142 MAJOR ISSUES FACING THE DIVISION OF WILDLIFE DRAFT This section presents an overview of the key issues facing the Division of Wildlife. While many others could be identified, the ones discussed here are the ones of most significance to Division decision-makers. Many of them overlap and most are highly interrelated. Each issue is first described briefly. Then its impact on the Division is discussed, and finally a number of policy questions relating to the issue are posed. How the Division plans to deal wi th these issues is addressed in the • next section, which describes the Division's rnajor objectives for the next several years and the strategies that will be utilized in pursuing them. 1. DEMOGRAPHIC CHANGES .a. GYowth in Population/ Encroachment on Habitat Colorado's population will grow from the current 3.2 million to 4.1 million by the year 2000. This represents an increase of 900,000 people, level of or 28%. The rate of growth will slow from 2.4% per year (average over last fifteen years) to 1.7% per year over the next .fift~en years. In the past, population growth and the development that go along with it have seriously impacted wildlife habitat. Over the past five years an average of 117,000 acres of habitat were lost each year to building construction, highways, grassland plowout, and ski area development. Of the 2,380 flowing streams in the state, 25% have been including affected to some degree by man's activity, diversion, channelization, mining pollution, and domestic pollution. In the 21-year period from 1964 to 1985, the number of reservoirs in the state increased from 1,900 to 2,902, or by 53 percent. Surface acres of water increased from 205,000 to 368,000, or by 80 percent. As of 1985 inundated. a total of 1,039 -10- mi1des of streams had been �143 DRAFT Human encroachment on wildlife habitat in the state is expected to continue but at a slower pace than over the past fifteen years. A d~pressed agricultural industry and increased farm productivity should substantially diminish grassland plowout, which represents 80% of the estimated habi tat loss over the past two decades. Changes in ski area technology and economics, with more restrictions on development, significantly slow. the rate of habitat loss mountains. coupled should in the All human encroachment on wildlife habitat is cumulative in its impact. While the pace of encroachment may slow, the wildlife resource will continue to be pushed closer to its limits. This encroachment will impair the Division's ability to maintain the kinds of wildlife, the size of wildlife populations, and the locations of wildlife populations that the pulbic demands. It will also increase the cost of maintaining wildlife populations in the state. The Di vision is only one of many players in the decision-making process, and cannot directly encroachment on wildlife habitat. land use control In the past, the Division has re1ied on mitigation (primarily on projects funded with ftderal dollars) and acqu i sit ion of cr itical habi ta ts to influence human encroachment on wildlife. Policy questions/alternatives include: Should the Division focus on locations, etc. of development, influence the process generally? certain types, or should it sizes, try to Should the Division at tempt to work more closely wi th municipalities and other government agenCies at all levels to participate more fully in the entire land use decision-making process? Should the Division implement an agressi ve mi tigation policy, and attempt to negotiate for mitigation on all projects with significant wildlife impacts? -11- �144 DRAFT Should the Division develop educational programs targeted at developers, municipalities, government agencies, and other parties involved in development and land use decisions, to increase their awareness of the impacts of development on wildlife and how to minimize those impacts? Should the Division' seek legal constraints on development, such as restrictive zoning ordinances, water quality standards, minimum stream flow r~quirements, etc.? Should the Division attempt to preempt development through the purchase of critical habitat, water rights, development rights, etc.? b. Changes in Kinds of Recreation While the state's population is changing in other ways as well. in Demand changing is size, it is Single-person households, and those headed by single parents and females, are increasing as a percentage of total households. These groups have not traditionally participated heavily in wildlife recreation. People born during the "baby-boom" years are and will be entering middle age. The middle age segment of the population is probably the most active in wildlife related recreation, and plays a key role in passing on a wildlife "ethic" to subsequent generations. Consumer tastes are diversifying (this includes sportsmen and other wildife recreationists as consumers). Whereas at one time the recreational preferences of sportsmen were more or less uniform, there has been a marked rise in preferences for different kinds of recreation. For example, some hUnters like to hunt with a bow, some with a rifle. Some are more interested in hunting large, "trophy-size" animals or less common types of animals, while ~thers are primarily interested in harvesting meat. Some fishermen fish primarily to catch and keep fish, while others fish for sport and readily return fish to the water after they are caught~ -12- �145 Numerous market surveys have found that different groups of people particpate for different reasons, and place different values on different aspects of the experience. These surveys have also shown that for most people a great many factors are involved in producing a satisfactory and enjoyable recreational experience. These surveys have shown strong and growing interest in "nonconsumptive" recreational activities. At the same time, demand for these types of activities is often not well defined, which makes it difficult to respond to. Special interest groups representing these different groups of people have grown more numerous and more vocal. Each has its own particular agenda. i There is some evidence that certain traditional forms of recreation, such as hunting, may be plateauing or even beginning to decline. Nationally, the percentage of the population that participates in hunting has been steadily declining over the past two decades. Attempts to meet the changing and growing variety of demand vastly complicates the task of wildlife management. Often the interests of two different groups are not compatible, and must be somehow balanced. Management costs increase with increased diversity of opportunity, as different areas must be managed differently, perhaps under different regulations or different biological management practices. The Division has in recent years attempted to meet the demand for greater diversity of wildlife recreation. Examples of this include establishing separate seasons for bow and rifle hunting, designation of Gold Medal waters, and increased emphasi s on nonconsumpti ve uses. Two yea rs ago antler restrictions were imposed on deer and elk hunting in response to the perceived demand by hunters for more "trophy-size" animals. The Division cannot be all things to all people, but recognizes the need to serve a broad range of people wi th the greatest possible diversity of recreational opportunities. The key challenge will be finding the balance between serving a narrow range of interests well, and serving a broad range of interests poorly. -13- �146 DRAFT Policy questions/ alternatives include: Should the Division specialize more in providing those kinds of recreational opportunities to which the state is well suited, or should it try to offer a very broad range, even if it means developing opportuni ties which the state cannot very efficiently provide? Should the state be managed 100% for "quality" hunting (i.e., large animals, low hun t er density, the "Vail" of hunting), 100% for "harvest" (i.e., maximum sustained yield), or some combination? Should the Division actively attempt to influence demand and engage in efforts to promote the use of the resource, or should the Division simply focus on providing wildlife and regulations governing its use so people will have the opportunity to enjoy it? Should the Division wildlife recreation? promote, market, and advertise Should the Division pursue a strategy of "vertical integration" in wildlife recreation? To what extent should the Division engage in providing services and products ancillary to wildlife recreation? Should the Division develop its properties with facilities for serving campers, sportsmen, etc.? If so, how should they be developed and to what extent? Should the Division offer tours, provide offer where to/ how to classes, etc.? guides, Should the Division attempt to serve all segments of the wildlife recreation market or a selected few? What kinds of tradeoffs is the Division willing to accept between license prices and participation? License prices and the need to limit participation? participation and catch or harvest rates? Size of fish and game harvested and numbers harvested? c. Urbanization of Population Most of the growth in population years will be concentrated along corridor. over the next fifteen the Front Range urban As populations become more urbanized, interest in wildlife and participation in wildlife recreation tend to decline. -14- �141 DRAFT In 1984 Region, and its to reach Policy the Division created essentially comprising immediate surroundings the urban population. questions/ alternatives a new region - the Central the Denver metropolitan area - to gi ve focus to efforts include: Should the Division focus on developing recreational opportunities within metropolitan areas, or on encouraging urban residents to go outside metropolitan areas to where these opportunities already exist? Should the division's efforts focus on developing expanded recreational opportunities (e.g., more wildlife areas) or on educating and informing people about the opportunities that already exist (e.g., publishing bird-watching guides, etc.>. Should the Division direct educational efforts at specific groups (certain age groups, certain income groups, certain household characteristics, certain geographical locations, etc.) or at the general populace? If the Division is going to develop recreational opportunities in urban areas, what should these be? Should they be interpretive areas? Wildlife refuges? Exclusive use areas? Many small areas or a few large areas with greater numbers and diversity of species? 2. DOW FINANCIAL DEPENDENCE ON DEER AND ELK HUNTING Of the division's annual expenditures of about $40 million, over 75% is funded by sales of licenses to sportsmen. Over 50% is funded by sales of deer and elk licenses alone. For most of the past two decades, revenues deer and elk licenses grew rapidly. In recent years fallen sharply. revenue growth from these from the sale of two sources has Over the next several years, Division operations will be significantly constrained as a result. If this situation is not rectified, future operations may have to be sharply curtailed. Many potential strategies for addressing the numerous other issues outlined in this section may be impossible to implement due to lack of funds. -15- �148 " ~At-.•. .r I Dr~ In the past, the Division could rely on license fee increases and its robust fund balance to cope wi th any temporary declines in, revenue growth. It now appears that hunting fee increases might not significantly increase revenues, and the division's fund balance, after several years of decline, no longer provides the kind of cushion it once did. Many observers believe that hunting as a sport is ei ther plateauing or declining. If that is the case, then the Division could be faced with continued weak growth in revenues. Other. observers believe that the "decline" in hunting ha-s been overstated, and that rlslng demand for hunting in Colorado can and should be maintained. If this is true, it may well be that dependence on deer and elk license revenues is not a weakness but a strength. Policy questions/ alternatives include: Should the Di vision focus on reducing the proportion of its revenues obtai ned from dee rand elk license sales by developing alternative funding sources, or should it focus on strengthening these programs so revenues from them will continue to increase? Should 1icense fees be rai sed? If so, by how much? What tradeoff is the Division willing to accept between higher license fees and reduced participation? What kind of differential should exist between resident and nonresident license fees? Should the nonresident Di vision at tempt to increase the ratio to resident sportsmen to boost revenues? of Should the Division "market" wildlife recreation, encouraging more people to participate in hunting and fishing, and thereby increase revenues? If so, where should these efforts be targeted? Or is this not a proper role for the division? Should the Division refrain from attempting to influence demand and invol ve itself instead solely in providing the opportunity for people to use the resource? If· alternative funding sources should what should they be and at what groups targeted? -16- be developed, should they be �149 DRAFT 3. RISING COSTS OF PROVIDING WILDLIFE RECREATION OPPORTUNITIES Over the past decade the Division has embarked on many new programs and has broadened the scope of its acti vi ties in many areas. This trend years. has, if 'anything, accelerated over the past few Since 1977 the number of permanent employees at the Division has increased by over 25% and annual expenditures have increased by 100%. During the 1970s and early 1980s this expansion was easily accomodated because revenues were growing rapidly. In recent years the growth in revenues has slowed markedly. Yet the Di vision has continued to add new programs, and many programs initiated in earlier years are now entering period~ of peak funding requirements. The Division will continue to be under pressure to broacen both the scope and intensity of its activities and to undertake new programs. To accomodate past decisions and future demand in the face of slowly growing revenues, the Division must either curtail certain activities, stop providing certain services, or find significantly more efficient ways of doing the things it is already doing. Policy questions/ alternatives include: Should the Division invest in long term steps to improve efficiency (e.g., acquire a new accounting system, up~rade managerial capabilities through training, lmprove wildlife management techniques through research, etc.), recognizing that in the short term this will only increase the financial squeeze? Should the Division curtail certain activities or reduce or cease providing certain services it now provides? If so, what should these be? For example, should the Division expand hatchery production to maintain current catch rates (which under current fee structures would result in an increased drain on Di vi sion resources) or forego hatchery expansion wi th the realization that catch rates will decline? Should the Division cease acqu ar i nc land (and perhaps even liquidate some of its holdings) and accept some -17- �150 DRAFT reduction in access to the resource? Should the Division reduce the number of publications, number of brochures, phone answering service, etc., and accept a lower level of public awareness and a reduction in public "goodwill" towards the division? Should the Di vision cease maintaining publ ic facili ties on its properties? Should the Division reduce field law enforcement efforts and accept higher violation rates and reduced safety? Should the Division enter into more cooperative agreements with other bodies (e.g., Parks and Recreation, municipaliti£s,· etc.) for sharing costs (and benefits) of wildlife and recreation management? Should the Division liquidate or upgrade its property holdings in cases where wildlife and/or recreational benefits are not commensurate with the investment in the property? 4. CONFLICT BETWEEN RESIDENT AND NONRESIDENT SPORTSMEN While resident sportsmen account for 66% of the total number of sportsmen in the state, they account for only 37% of the license revenues. Conversely, while nonresident sportsmen account for only 34% of the total number of sportsmen in the state, they account for 63% of the license revenues. The cost of serving resident and nonresident hunters is essentially the same. Therefore, from the divisionis standpoint, the greater the proportion of nonresident sportsmen, the more the Division can do to maintain the resource and enhance recreational opportuni ties. In fact, without nonresident sportsmen, the Division would be unable to serve resident sportsmen as well as it does now. From the state's standpoint, the posi ti ve economic impact of nonresident recreationists is much greater than that of resident recreationists. Since the state is currently very concerned about economic impacts and economic development, this might suggest, that efforts should be made to increase the number of nonresident recreationists. However, the wildlife resource is finite (although renewable) and more nonresident sportsmen means reduced opportunities for resident sportsmen. The Division has always followed a policy of preferential treatment of' resident sportsmen. -18- �151 Policy questions/ alternatives include: Should the Division attempt to increase the number nonresidents who come to the state to hunt, fish, otherwise enjoy wildlife? of or Should the Division attempt to limit the number of nonresidents (either through pr ices, limi ting licenses, or some other means) who corne to the state to hunt, fish, or otherwise enjoy wildlife? What kind of differential resid~nt and nonresident fees? should exist between t· S;", : the Division attempt to provide the kinds rec!€ation opportunities most desired by residents, those most desired by nonresidents? 5. WILDLIFE RELATED CONFLICTS Many species of wildlife forage and feed. Many agricultural crops. By statute, pay owners wildJife. of or compete species with domestic animals for of wildlife also damage the Division must under some circumstances for damage caused to crops and feed by Over the past ten years, the Division has paid farmers and ranchers over $2 million for damage caused by wildlife, and has spent over $3 million on measures designed to reduce damage caused by wildlife. Over that same period of time Division personnel have probably devoted $2 million in manpower to game damage related activities. Wildlife itself is a publicly owned resource, yet the lands on which it is found are often pri vately owned, or are subject to certain use limi tations which preclude hunting or fishing. 50% of land in the state is privately Sportsment are not allowed on State Land Board Lands. Trespass on private lands factcr to poor relations sportsmen. Land ownership public lands. patterns -19- owned. is a major contributing and between land owners often restrict access to �152 Policy questions/ alternatives include: Should the Division seek statutory changes damage laws to reduce its liability? in the game Should the Division continue to invest in, or expand its investment in, measures designed to reduce game damage (e.g., paying for fencing, etc., to prevent game damage, Ranching for Wildlife, etc.)? Should the Division reduce the size of wildlife populations in areas where game damage is a significant problem? Should the Division pursue statutory/ for opening State Land Board Lands? Should the Division wildlife? acquire legal lands to provide avenues access to Should the Division continue, expand, or reduce programs aimed at convincing private landowners to allow hunting and fishing on their lands? Should the Division develop and promote financial incentives (e.g., payments to landowners, "trespass fees", etc.) to encourage pri vate landowners to open their lands to the public? 6. GROWTH OF THE DIVISION OF WILDLIFE Over the past ten years the number of permanent employees at the Division has increased by more than 25%, and annual expenditures have increased by 100%. Over the past ten years the Division has ini tiated new programs, and expanded both the scope and intensity of its activities. This has led to a growth in the number of specialists employed by the Division and in the diversity of fields they represent. Recreation management has become more complex, wi th more issues and interest groups involved. At the same time, as the resource is managed more intensively, wildlife management has grown more complex. The result of this has been to greatly compound the task of coordinating and integrating all of the various operations and functions within the division. -20- �153 DRAFT The Division has not yet successfully made the transi tion from a small, narrowly focused organization to a large, multi-facted organization. policy questions/ alternatives include: Are the human resources (skills, training, experience, expertise, etc.) of the Division appropriate for the job(s) it is trying to do? If not, how should any gap be closed? Should the Division acquire expertise in key areas, or invest in training existing personnel? Should the Division alter its recruitment, selection, and training processes? Should the Division invest in management systems to improve coordination and integration of Division operations? Should the Division replace its accounting system? Should the Division develop a management information system? Should the Division devise and implement more formal management systems (e.g., di recti ves, Planned Management System, moni tor ing and evaluation, etc.)? Should the Division continue to expand its ADP capabilities? Is the division's organizational structure appropriate? How can the functioning of the Commission Commission be improved, and how can division/ communications be improved? 7. FIELD PRESENCE REQUIRED TO MANAGE STATEWIDE RESOURCE The nature of the wildlife resource practically requires continual, permanent field presence of Division officers. This means Division manpower must be distributed throughout the state. As new programs are implemented, it is difficult to staff up to meet the additional manpower requirements by drawing down manpower from the field. As a result, there has been a tendency to delegate additional program responsibilities to field personnel already in place. Because few internal controls exist and records on manpower use are poor or nonexistent, this practice has gone unchecked for years. As a result, Division field personnel have been saddled with tremendous responsibilities to perform a wide variety of functions requiring a broad range of skills and expertise. The demands the Division has placed on field personnel are excessive relative to the number of field personnel, and have not been accompanied by sufficient training. -21- �154 uttAFT When new functions are identified and given high priority, the Division has three ways of carrying them out. One is to acquire the necessary expertise from outside, increasing total staffing levels in the process. The second is to simply increase the work load for existing staff by adding the new fUnction to existing responsibilities. The third is to acquire the necessary expertise from outside, while at the same time reducing existing staffing levels by a like amount. The fourth ,is to train existing staff to perform the new function, and accomodate its performance by cutting back or eliminating existing duties. In the past, the Division has relied almost exclusively on -the fir st two al terna tives. Due to fundamental changes in the dT vi si on's financi al condi ti on, together wi th the fact that current work load is at or beyond saturation levels, these are no longer viable alternatives. Policy i questions/ alternatives include: Should more of the multi-purpose (all-purpose?) DWMs' current responsibilities be taken over by specialists (with a consequent decrease in DWM staff levels and an increase in specialist staff levels)? Should more of the work currently being done by specialists be taken over by DWMs (with a consequent reduction in specialist staff levels, an increase in DWM staff levels, and increased efforts devoted to DWM training)? Is too much currently expected of field personnel? If so, what current activities should be curtailed or eliminated? When new programs are implemented, what existing activities should be curtailed or eliminated? 8. MANAGEMENT TECHNOLOGY In the future, the wildlife resource will be managed closer and closer to biological capacity, and will be managed for a growing diversity of uses. Both of these will require that the resource be managed much more intensively. Frequently, however, a lack of understanding of management techniques and a lack of data which those techniques require prevents the Division from managing .the resource as intensively as it COUld. In many cases, the Division does not know how to manage to br ing about a desired response. Stated differently, in many cases we do not know what the effects of a given management approach will be. -22- �155 DRAFT In other cases, we might understand how the resource would respond in theory, but we cannot quantify what that response will be because we lack the necessary data for making such a prediction. These are not new problems. They are solved primarily through research and data collection, activities to which the Division has devoted significant efforts for years. However, if we are going to demand more from the resource, increased knowledge about how wildlife systems behave, how they respond to management, and what their current status is will be necessary • .. , policy questions/ alternatives include: Should the Division expand, reduce, or maintain current levels of research? What is an appropriate proportion of the division's budget that goes to research? Research and data collection serve to reduce the risk of management decisions, by allowing better predictions of the consequences of those decisions. What tradeoffs is the Division willing to make be t we e a reducing the risk of management decisions and the cost of reducing that risk? Are there particular investment in research especially critical? -23- areas of and data concern where collection are �156 MAJOR OBJECTIVES AND STRATEGIES FOR ACHIEVING THEM DRAFT The previous section described a number of issues which the division must confront over the next fifteen years. As the discussion indicates, these issues present the di vision wi th both problems and opportunities. In this section, a number of objectives and strategies are described that show how the division intends to respond to"these various issues. These are targets which the division will MANAGE to achieve: they are not simply projections or forecasts. More specific objectives in the areas discussed below will be developed each year in the annual update of Program and Operations Plans. THE OBJECTIVES AND STRATEGIES SET FORTH IN THIS PRELIMINARY DRAFT ARE FOR DISCUSSION PURPOSES ONLY, AND DO NOT NECESSARILY REFLECT DIVISION RECOMMENDATIONS OR DECISIONS. Furthermore, they are based on a number of assumptions, including the actions of .t he Colorado Legislature and the level of future revenues. If these assumptions change, the Division's objectives and strategies may need to be changed as well. I. GENERAL A. In the three years following FY 1987-88 (FY 1988-89 through FY 1990-91), total planned spending will not exceed forcast revenues. B. In subsequent years (i.e., during the period FY 1991-92 through FY 2002-03) total planned spending will not exceed 95% of forecast revenues. (NOTE: Thi s does not mean that total spending will necessarily be held to 95% of revenues.) C. The division's total fund balance (at fiscal year-end) will be maintained at a level of $10 million or 25% of revenues, whichever is greater. D. The division's total unobligated fund balance (at fiscal year-end) will not fall below $5 million or 10% of total revenues, whichever is less. -24- �157 DRAFT E. Over the next fifteen year s, the proportion of total revenues obtained from the sale of deer and elk hunting licenses will be reduced from its current level of 60-65% to 40%. F. Over the next five years, the proportion of total operating expenditures made up of personal services costs of permanent employees will be reduced from its current level of 56% to 50%. It will not rise above 50% over the following ten years (FY 1993-94 through FY .2002-03). (NOTE: The reduction to take place over the next five years will be at whatever pace can be accomplished through attrition.) 6. Over the next three y~ars, 7.5% of the division's total ·planned expenditures will be devoted to capital investment (down from the current level of 8.5%). Capi tal investment here means capital constuction fund expenditures excluding land acquisition. Of the division's total planned expenditures, 92.5% will be sevoted to operations (operating funds). H. Over the next five years, the proportion of the di vis ion's total planned oper at ing expend itures devoted to research will be held at the current level of 8.5%. I. Over the next fifteen years, the proportion division's total planned operating expenditures to administrative activities will be reduced from its current level of 14% to 11%. of the devoted by 20%, J. Over the next three years, no new programs will be initiated, and no existing programs expanded, without specifically identifying how the increased activities will be funded. K. Over the next five years, projects will come primarily 1. 2. 3. 4. funding from: for new programs/ Increased revenues following fee increases. Curtailments of existing programs/ projects. Reduction in administrative costs. A sl ight shift from capi tal construction spending to spending on operations. 5. Liquidation of assets where long term benefi ts are clear. 6. Funds obtained through mitigation. -25- �158 DRAFT L. By no later than FY 1990-91, the Division will begin implementing sweeping changes in the way it funds its operations. These changes may include restructuring license fees, development of alternative funding mechanisms, and giving the Commission author ity to set fees. M. Over the acquisitions necessarily place. ) next three years, land no additional will be planned. (NOTE: This does not mean that no land acquisitions will take N. The objectives and strategies presented here ignore the potential for recelvlng funds through mitigation efforts. To the extent that the Division is successful in negotiating mitigation payments, the objectives with respect to capital construction spending, acquisition of land, and construction of hatcheries, among others, may be modified. II. HUNTING RECREATION A. Big Game 1. Over the next fifteen years, the number of deer and elk hunters and hunting recreation days will grow slowly. The number of elk hunters, currently at the depressed level of 132,000, will increase .t o 175,000 by FY 1989-90, and will remain at that level through FY 2002-03. The number of deer hunters will increase gradually from 190,000 to 200,000 over the next fifteen years. Target numbers of hunters in FY 2002-03 will range from 90% of previous historical highs for elk to 95% of previous historical highs for deer. 2. To avoid will be population 3. Deer populations will be increased by about 6% from current levels (from the current post-hunt population of 564,000 to 600,000) over .the next ten years. They will then be held at that level through FY 2002-03. 4. Elk success rates will be maintined at levels (.15) over the next fifteen years. .. exacerbating conflicts, elk populations held at current levels (post-hunt of 160,000) over the next fifteen years. -26- current �159 DRAFT 5. Deer success current level years. rates will be improved from the of .31 to .40 over the next fifteen 6. The division will manage for a diversity of hunting opportuni ty. Over the next fifteen years, 45% of the elk harvested will come from areas managed for high yield, 45% will come from areas managed 100% under branch-antlered restrictions, and 10% will come from areas managed for trophy-size animals and low hunter densities. 7. The proportion of all big game licenses sold to nonresidents will increase from the current 30% to 35-37% over the next fifteen years. 8. The proportion of big game hunting recreation days from species other that deer and elk will remain small, but will increase by 33%, from the current level of 6% to 8% over the next fifteen years. 9. By no later than FY 1991-92, big game license fees for res idents wi 11 be increased 20-40% above current levels and maintained at that level (in real terms) through FY2002-03. By no later than FY199l-92, big game license fees for nonresidents will be increased 10-30% above current levels and maintained at that level (in real terms) through FY2002-03. These increases will restore both resident and nonresident license prices to their average level over the last twenty-five years (in real terms). 10. No significant next five years game herds. 11. Expendi tures to prevent game damage and compensate land owners for game damage will be held to current levels over the next five years. 12. game hunting exceeds the If demand for big division's ability to accomodate, licenses will be limi ted and/or season structures will be adjusted. License price will not be used to modulate demand. investment will be made over the to secure more public access to big -27- �1GO DRAft B. Small Game 1. Over the next fifteen years, the number of small game hun t e rs and recreation days will show strong growth. Number of hunters will grow by 27% and recreation days will grow by 53%. 2. Average s~all game harvest per day will increase by 33%, from 1.2 per day to 1.6 per day, over the next fifteen years. 3. Emphasis will be placed on improving access to small game. Small game hunting properties managed by the division will increase from the current 175,000 acres to 197,000 acres over the next fifteen years. No acquisitions will be planne~ until after FY1989-90, however. 4. By no later that FYl99l-92, small game license fees for both residents and nonresidents will be increased by roughly 100% above current levels and maintained at that level (in real terms) through FY2002-03. III. FISHING A. By the year FY 20 02-0 3, the number of licensed angl er s and recreation days will be 40% above current levels. This means the number of anglers will grow faster than the state's population as a whole, reflecting the division's belief that per capita participation in angling will increase. B. The gap expenditures between fishing program revenues will be closed by the year FY200l-02. and C. catch per day will be allowed to decline from the current 2.8 per day to 2.3 per day by FY2002-03. Emphasis will be placed on Lncr eas i riq the average size of fish caught. D. By no later that FYl99l-92, resident license prices will be increased by 50-60%, and by another 50-60% by FY2002-03. By no later that FYl99l-92, nonresident license prices will be increased by 40-50%, and by another 40-50% by FY2002-03. These increases will put both resident and nonresident license fees significantly above their historical averages (in real terms) over the last twenty-five years. -28- '. �161 DRAFT E. Over the next three years, no planned to increase angler access. acquisitions will be F. Wi th the exception of the Buena Vista project, investment in hatchery production capacity expansion will not be made until demand is evidenced at higher fee levels (i.e.,' sometime after FY 1989-90). Current plans assume hatchery production capacity will need to be increased by roughly 25% by the end of FY 2001-02. G. To increase the average size of fish caught, and to fish production costs, areas managed under catch and release regulations will be expanded. z.educe H. Fishing in urban and warmwater areas than fishing in the state as a whole. IV. WATCHABLE will grow faster W!DL!FE A. Over the next fifteen years, the number of "nonconsumptive" recreation days will grow at an average rate of 5% per year. This is a very aggressive target, and reflects the division's belief that there is tremendous untapped demand for this form of recreation in Colorado. B. Over the next three years, division efforts will concentrate on educating the public about nonconsumptive recreation opportunities, maintaining existing nonconsumptive recreation properties and facilities, planning pilot development projects, and developing addi tional funding sources for the Watchable Wildlife program. 1. Educational efforts include Project Wild and informational brochures and publicat·ions and will be focused on the Central Region. Resources devoted to this effort will remain at current levels. C. By no later that FYl990-91, the division will have in place alternative funding mechanisms capable of generating $1 million per year for the Watchable Wildlife Program within three years of implementation D. Beginning in FY1989-90, the division will develop a number of pilot Watchable Wildlife facilities and/or projects, and closely monitor public response to them. E. During the twelve year period FYl991-92 through FY2001-02, the division will develop Watchable Wildlife facilities and/or projects, based on public response to pilot projects, in accordance with public demand. -29- �162 DRAFT F. Over the period FYl99l-92 through FY200l-02, Watchable Wildlife direct program expenditures will not exceed program revenues. v. SPECIES CONSERVATION A. By the end of FY2002-03, a total of five out of the 21 species now classified as threatened or endangered will be recovered. B. Over the next two years (FYl988-89 through 1989-90), the focus will be on maintaining current efforts on recovery p-rograms now in place, and on identifying all species of special concern. following _ fi vee years, through the (FYI990-9l FYl994-95), the focus will be on maintaining current efforts on recovery programs now in place, and on developing threatened and recovery p l a.ns for additional endangered species and species of special concern. C. Over D. Over the following eight years (FYl995-96 through FY2002-03) the focus will be on implementing all recovery programs and downlisting/recovering species. By FYl995-96, twenty-seven recovery programs will be in place. -30- �163 APPENDICES -31- �164 CURRENT PROGRAM CURRENT PROGRAM -33- STATUS STATUS �Wildlife Research Report July 1987 lO~ JOB PROGRESS REPORT State of Colorado Project No. 01-03-048 (FW 26 P) Mammals ~~--~------~------~- 2 Research Work Plan No. 1 Multispecies Investigations Job No. 4 Big Game Forage Selection Dynamics Period Covered: July 1, 1986 - June 30, 1987 Author: R. B. Gill Personnel: M. A. Wild, M. W. Miller ABSTRACT Eleven of 13 pronghorn were hand-reared to 2.5 mos. of age. All fawns were conditioned to wear mock headgear which will subsequently be used to experimentally manipulate light wavelengths which they are permitted to see. Plans call for conducting foraging bouts with pronghorn females. Each female will be subjected to 6 experimental treatments which will consist of denying them access to red, blue, green, yellow, near infrared wavelengths compared to a control where all wavelengths are accessible. Treatment effects will be measured by comparing the quality of diets selected under each treatment regime. ��167 BIG GMIE FORAGE SELECTION DYNAMICS R. Bruce Gill P. N. OBJECTIVE To evaluate the role of color v~s~on foraging ruminant--the pronghorn. in diet selection of a small, selectively SEG!-lL"l'T OBJECTIVES 1. fu __~sh the results of intake experiment. 2. Design 3. 5ubmi t study plan for--Ro1e to pet:::review. 4. RaisE pronghorn 5. Publish second generation pronghorn intake experiment. of Vision in Forage fawns for "Role of Vision damage Selection by Pr ongho rns+-; study. study results. METHODS ~~D MATERIALS Intake Experiment Six, 3 year-old pronghorn females were fed a pelleted version of the winter feeding ration ad libitum. Individual intake was not measured; instead, total intake for all six females was measured daily and averaged among all 6 fe~ales to estimate individual daily intake. A pronghorn male was ~~aced with the females in October, 1986, so intake of pregnant females cou~~ be compared with intake from nonpregnant females from the previous year. R~sults have not yet been analyzed froID 3 years of intake data because the 3-year period will not be completed until October 10, 1987. The second generation intake study will test for the effects of ration size (pellets vs. wafers) on intake and body weight performance of pregnant females. Role of Color Vision in Pronghorn Diet Selection Currently, plans call for beginning this experiment in January, 1988. Handreared pronghorn will be allowed to graze without restriction in a short-grass prairie community north of the Foothills Research Facility. Each animal (up to 5) will be subjected to each of 5 vision treatments. Vision treatments will be imposed by camera lens filters which selectively fi:!.terout light in red, green, blue, yellow, and near infrared wavelengths. Diet selection will be quantified by counting bites of forage species. Selected plant parts of forage species which comprise more than 2% of the species intake by weight will be collected for nutritive analysis. Intake of plant parts by weight will be estimated by simulating pronghorn bites with hand plucking. Quality of diets selected by pronghorn subjected to various filter treatments will be compared to diets selected by animals subjected to clear glass filters. �Ibd Pronghorn currently are being trained to wear headgear similar to aviator goggles. Experiments will be replicated among each animal by grazing with each filter 2 times a week and grazing biweekly among months. Indexes to diet quality will be the cell soluble fraction of selected diets and the crude protein content of diets. Pronghorn Fawn Rearing and Training Six pronghorn females were placed with a mature buck in October, 1986, and remained with the buck for at least 30 days. Plans were to remove fawns from dams within 24 hrs of birth. Fawns would be reared in isolation for the first 3 weeks of life to insure imprinting on humans. Rearing diets consisted of undiluted, canned, evaporated milk offered ad libitum approximately 5 times a day from 4.5 weeks to 7 weeks when it was reduced to 3 times daily. Thrice-daily feeding continued from approximately 7 weeks until 11 weeks whereon frequency was reduced to twice-daily feedings until-14 weeks. Once daily feedings continued from 14 weeks until weaning at approximately-110 days of age. From 4 weeks of age fawns were trained to accept wearing a cloth headpiece that resembled a mask. Eyeholes were cut in cloth and the apparatus was held in place with Velcro fasteners. Pronghorn-Winter Wheat Damage Study Plans called for analyzing all data collected during the pronghorn-winter whea~ damase study. All data would be presented and discussed in a final report to the Agricultural Experiment Station of Colorado State University. Two manuscripts are to be prepared and submitted to professional journals for publication consideration. One will discuss results of the controlled grazing experiments with hand-reared pronghorn grazing wheat pastures for specified treatment lengths. The other will discuss the role of plant nutritional phenology in determining the period of winter wheat use by pronghorn. RESULTS Pronghorn Intake Study On October 10, 1987, collection of 3 year's data on pronghorn daily intake and weekly weight performance will be completed. Analysis will focus on describing seasonal trends and comparing those trends with other ungulate species. Preliminary examination of the data reveals strong seasonal trends both in body weight and in ad libitum food intake. Similar to trends of deer, elk, and mountain sheep, pronghorn intake and body weight peak in fall and then decline through winter. Intake inclines in spring and summer to peak again in fall. Role of Vision Study Plan Completion of a final draft of a study plan awaits completion of fawn rearing and training. it is not yet known how many proghorn fawns will tolerate the experimental apparatus well enough to forage normally during experimental grazing bouts. �169 Fawn Rearing and Training Nine fawns were torn to captive pronghorn does at the Foothills Research Facility. An additional 4 fawns were taken from wild dams and brought to Fort Collins to rear. Two fawns died of diseases early in the fawn rearing process. The remaining 11 have thus far been reared successfully. Eight of the 11 are being trained regularly to wear the experimental headgear during foraging bouts. ~ilk intake has been recorded daily for each fawn, and body weights have been recorded at weekly intervals. These data will be summarized into a publication on pronghorn fawn growth rates after weaning. In addition, forage selection dynamics have been recorded.separately for each grazing bout. Diet selection dynamics as a function of age will be analyzed and a manuscript prepared for publication consideration. Pronghorn Damage Study A draft final report to Colorado State U~iversity's Agricultural Experiment S~ation was prepared, reviewed, an~ edited by the 4 authors. Deadline for submitting the final report to the Agricultural Experiment Station is September 17, 1987. Following that sub~ission, 2 manuscripts will be prepared and submitted to professional journals for publication consideration. \ Prepared by ~i .. ,,~~ ... ._.-. - •.. -' --'-I...- R. Bruce Gill Wildlife Research Leader ��171 Wildlife Research Report July 1987 JOB PROGRESS REPORT State of Colorado Project No. 01-03-048 (FW 26 p) --~~~~~--~~~~~- Mammals 2 Research Work Plan No. 1 Multispecies Investigations Job No. 5 Home Range Model Evaluation Period Covered: July 1, 1986 - June 30, 1987 Author: G. C. White Personnel t D, '.' R. Anderson ABSTRACT Progress towards the objectives of this job include: 1. An evaluation of home range procedures was performed and copies were distributed to interested cnow personnel. 2. A 2-day workshop was conducted on line transect estimation. 3. A I-day workshop was conducted on home range estimation. 4. A menu-driven line transect microcomputer program was developed for wildlife inventory. 5. A I-day workshop was conducted on microcomputer statistical packages with emphasis on Statistical Analysis System (SAS). 6. A variety of consultations were held with cnow personnel on the application of microcomputer technology to wildlife analysis problems. ��173 HOPE RANGE MODEL EVALVATION Gary C. White P. N. OBJECTIVE Evaluate the home range concept as it applies to large and small carnivores and develop improved home range analysis models. SEGMENT OBJECTIVES 1. Evaluate home range models for large and small carnivores. 2. Conduct training workshops for DOW researchers and biologists in home range concepts and line transect methodology. RESULTS AND DISCUSSION Conceptual and quantitative models of home range were reviewed and critiqued. This review and critique resulted in a revision of the "Home Range Estimation" chapter in the book "wnite, G. C., and R. A. Garrott. 1987. Analysis of biotelemetry data - a primer. Colorado State University. Ft. Collins, CO." (chapter appended as Appendix A of this report). A I-day workshop was held on home range estimation. CDOW biologists and researchers participated. A menu-driven software program was developed fo" line transect surveys, and a 2-day workshop was conducted on line transect s~rvey estimates of deer, elk, and pronghorn populations. CDOW biologists and researchers attended. A l-day,"hands on" workshop was presented to CDOW biologists and researchers conce,ning microcomputer statistical analysis software programs. SPSS, SAS, Minitab, and others were reviewed. Utility of SAS program was explored in some detail. All participants had access to a microcomputer for "hands on" experience. Consultations were held with a variety of CDOW personnel on the application of microcomputer technology to specific wildlife analysis problems. Among those consulted were: L. H. Carpenter, R. M. Bartmann, R. B. Gill, G. D. Bear, D. J. Freddy, T. M. Pojar, D. F. Reed, and C. E. Braun. c' Prepared by l/t -.: ( .. .,i '7 V J(; /k Cr Gary C. White Associate Professor - CSU ��175 APPE\D IX _.1. / ..-- Chapter VII Home Range Estimation Home range is defined as "that area traversed by the individual in its normal activities of food gathering, mating and caring for young" (Burt 1943:351). Thus, the units of home range are area. In practice, the term "home range" is used to describe two aspects of animal movement. First, the basic map of locations produced from tracking an animal is often referred to as the animal's home range. Second is the numerical estimate of the area used by the animal. Most of this chapter will explore the mechanics of the numerical estimation process, but the importance of the map of locations should be emphasized. All too often, the assumptions of a numerical estimate are so seriously violated that the number is worthless, but the map of locations is not distorted. The practical information gained from the biotelemetry data is accommodated with the map, but is lost with the numerical estimate. The key word in the above definition is "normal." Home range is not all the area that an animal traverses during its life time, but rather the area where it normally moves. Excursions to the area outside its normal area should not be considered part of the home range (Burt 1943:351). Therefore, objective criteria with a biological basis are needed to select movements that are "normal." In the literature, two criteria have been used. The first is the subjective evaluation of the researcher, and thus does not provide a repeatable, objective approach for other researchers. The second is to use a probabiIity level; for example, the estimated home range includes the animal's locations 95% of the time. This leads to a more precise probabilistic definition of home range: the probability of finding an animal at a particular location on a plane. This density function has been called the "utilization distribution" (Jennrich and Turner 1969, VanWinkle 1975, Ford and Krumme 1979, Anderson 1982). The home range estimate is calculated by drawing equal height contours around the utilization distribution (Anderson 1982). The home range is specified by the contour such that 95% (or some other percen- �176 Analysis of Biotelemetry Page 126 tage) of the animal's locations are 'Within the contour. Data This criteria is certainly objective, and thus repeatable, but it lacks a sound biological basis. Why should 5% or any other percent of an animal's movements not be considered normal among studies, an objective repeatable A second inadequacy (cf. Anderson 1982)? However, for purposes of comparison method is needed, and a 95% area meets this criterion. of the above definition of home range is that a time frame is not specified. Some time period over which the home range is estimated must be defined to implement estimator. a home range This time frame could be as long as the life time of the animal, but generally shorter periods are of practical value. The primary consideration and do the methods in estimating home range must be what is the purpose of the effort, used to collect the data satisfy this purpose. tion of animals for the month of January, then an adequate number of observations sample of animals must be radioed and an on each animal must be taken during January to satisfactorily mate each home range. A 2-stage sampling effort is required. must be captured and radioed, "Withthe requisite assumption that this sample of animals is representa- Next, each of the radioed animals is sampled to determine during the month of January. Samples must be taken throughout order to be representative its locations the month at all hours of the day in of an animal's movements for the month. Generally a representative sample, with each animal equally likely to be radioed, likely to be chosen as the time to determine esti- First, a sample of the animal population tive of the entire population. means a random must be If the purpose of the study is to estimate the average home range for the popula- taken into account. adequate Sampling considerations sample and each time of day equally the location of the radio. Independence of Observations Biotelemetry data are three dimensional closer in time two locations as has been pointed out throughout are taken, the less likely they are to be statistically this text. Thus, the independent. Stated differently, given the animal's location at time t, the expected change in location would be small for a small increase in time, t + l:J. As the difference in the two times becomes greater, the probability that the second Chapter location is known, given the first, is less and less likely. VII - Home Range Estimation A general rule of thumb to Draft June 9, 1987 �177 Analysis of Biotelemetry determine Page I1i Data if two locations taken at time t 1 and t 2 can be considered statistically independent is whether sufficient time has elapsed for the animal to move from one end of its home range to the other. ever, more sophisticated approaches have been developed by Dunn How- (1978) and Dunn and Brisbin (1982). All but one of the estimators be statistically independent. discussed in this chapter require that the input data (i.e., locations) Each location contributes two locations are not independent, as much information the sum of the information contributed as every other location. by the two data points is not 2' units, but. less than 2 units, because one of the locations can be used to make a reasonable the other. guess of_ To further illustrate this point, 100 locations taken 5 seconds apart on a radio-collared would not contribute nearly as much information locations each taken 3 days apart. If elk on the elk's home range for a 300-day period as 100 With locations taken 5 seconds apart, the estimate of home range would be very small and might even be zero if the animal had been resting during the 500 second interval. Only one of the locations really contributes any information other 99 locations inflate the sample size, but contribute on the animal's following sections where 2 or more locations are not independent. dimension is much smaller than it appears. independence erroneous discussed in the The sample size used to determine Only one of the estimators of the data, and thus does not generate The almost no information. The same process of sample size inflation occurs for the home range estimators the estimate home range. takes into account the time answers when there is a lack of statistical in the input data. The above example is not a practical case, because the 100 locations time are not a representative sample of the 300-day interval. taken 5 seconds apart in If the objective of the study was to esti- mate the home range over the 500 second interval, the 100 locations taken at 5-second intervals would be a representative sample, and the argument The 100 observations independent are not statistically can be made that the home range is properly estimated. independent on the time scale of 300 days, but they are on the time scale of 500 seconds. Draft June 9, 1987 Chapter VII - Home Range Estimation �173 Analysis of Biotelemetry Page 118 The bottom line on the question of statistical independence of consecutive observations the time interval been properly sampled for which the home range estimate is to apply?" tative (translated independent to mean random) sample has been taken, then the observations Data is "Has If a r epr ese n- can be assumed to be relative to the time frame of the sample. Minimum Convex Polygon The oldest and most common method of estimating (Mohr 1947). The minimum area polygon is constructed convex polygon (Fig. VII-I), and then calculating efficient FORTRA'\' algorithm for determining home range is the minimum by connecting the outer locations to form a the area of this polygon. the coordinates area polygon Eddy (1977) provides an of the polygon. Given these coordi- nates clockwise around the polygon, the area (A) is calculated as 16 15 J 14 13 il) • 12 • +) ~ •.... •.... .•...• 0 0 o :>-. • • • •• ••• • • ••• • I •• • • • • • • • • • • • • • • • • 11 "0 ~ • 10 9 8 7 6 1 3 5 7 9 11 13 15 17 19 x coordinate Fig. VII-I. Construction of a minimum area polygon by connecting the outer locations to form a convex polygon. The home range estimate is 0.007668 ha. Data are given in Table VII-l so that the reader may verify this estimate. Chapter VII .;..Home Range Estimation Draft June 9, 1987 �179 Analysis of Biotelemetry Page 129 Data n-l A [Xl (y" - Y2) + L: x,-(Vi-l - _vi.d .•. X"(>',,.l - YI)] i=2 2 The advantages of the convex polygon are (1) simplicity, (2) flexibility of shape, and (3) ease of calculation. However, the disadvantages are major. The size of the home range estimate increases indefinitely as the number of locations increases (Jennrich and Turner 1969) because the minimum area polygon is estimating total area utilized, not the area utilized in normal movements. Thus, the home range estimate is a function of the number of locations utilized to generate the estimate. Two estimates are not comparable if one is based on 100.data points, while the second is based on 500 data P9ints, because the second estimate is expected to be somewhat larger. The additional 400 locations Table '\11-1. Simulated data from a bivariate normal distribution with and covtr, y) = O. Observation Number 1 2 3 4 5 6 7 8· 9 10 11· 12 13· 14 15 16 17 18 19 20 21 22 23 24· 25 • x 240cr86: 14:06:28 250cr86:21:32:18 260cr86: 11:13:00 270cr86:02:33:30 OlNOV86:08:02:56 02NOV86:18:09:32 OSNOV86:04:12:04 07NOV86:08:13:17 10NOV86:09:07:12 20NOV86:02:02:32 2ONOV86:03:19:29 23NOV86:15:56:39 24NOV86:09:51:48 25NOV86:14:52:56 26NOV86:05:40:39 02DEC86:04:50:57 03DEC86:00:35:40 03DEC86:12:19:43 OIDEC86:00:20:25 OIDEC86:10:05:10 OIDEC86:17:00:30 08DEC86:17:18:20 13DEC86:01:06:41 13DEC86:05:38:09 15DEC86:18:49:15 (m} 10.6284 11.5821 15.9756 10.0038 11.3874 11.2546 16.2976 18.3951 12.3938 8.6500 12.0992 5.7292 5.4973 7.8972 12.4883 10.0896 8.4350 13.2552 13.8514 10.8396 7.8637 6.8118 11.6917 3.5964 10.7846 J.I.: = iJ.v = 10, ": = 3, "y = 1.5, y (m} 8.7061 10.2494 10.0359 10.8169 10.1993 12.7176 9.1149 9.3318 8.8212 8.4404 6.1831 10.9079 15.1300 10.4456 11.8111 11.4690 10.4925 8.7246 9.9629 10.6994 9.4293 12.4956 11.5600 9.0637 10.5355 Locations on boundary of the convex polygon. Draft June 9, 1987 Chapter VII - Home Range Estimation .•. �130 Analysis of Biotelemetry Page 130 Data provide further outlying locations. which lead to a larger polygon. One approach is to eliminate to correct the problem of increasing home range size with increasing sample size the "outliers" polygons based on an ordering criterion for the locations. ordered (ranked) We have calculated 95CC before the home range polygon is calculated. in descending Consider a list of numbers. order, with the largest number last. Nonparametric This list can be confidence vals can be placed on the 95th quantile based on the ranking of values in the list (Conover 115). However, ordering two-dimensional list (i.e., x and y coordinates x or y coordinate a one-dimensional is not reasonable. area each location contributes list of numbers of location) is straightforward, whereas inter- 1971:110ordering a is not obvious, because ordering by only the We assumed that the list could be ordered based on the amount of to the minimum area polygon. Thus to select the first location (the one Table VII-I. Continued. Observation Number 26· 27 28 29· 30 31 32 33 34 35 36 37 38 39 40 41 42· 43 44 45 46 47 48 49 50 Chapter \ VII - Home Range Estimation 16DEC86:16:42:14 17DEC86:07:05: 19 17DEC86:21:29:58 18DEC86:15:01:57 20DEC86:13:03:37 21DEC86:20:49:01 21DEC86:22:26:11 22D EC86: 18:06:00 23DEC86:08:32:42 27D EC86:09:51:28 29DEC86:12:20:03 3ODEC86:01:21:24 31DEC86:22:43:14 04JAN87:01:18:52 04JAN87:11:20:56 07JAN87:22:10:49 OBJAN87:12:02:33 11JAN87:12:58:54 13JAN87:05:58:34 14JAN87:02:22:31 14JAN87:19:36:24 15JAN87:05:59:05 2OJAN87:23:37:29 27JAN87:04:17:46 3OJAN87: 13:40: 10 x y em} em} 11.0807 10.9715 11.0077 7.6522 12.0681 13.2519 12.6987 10.2604 10.5340 10.1214 10.8152 11.3384 8.6962 10.1955 8.4525 9.9342 6.7882 9.0810 11.4518 10.2106 9.8179 11.5134 9.6083 10.5646 11.8786 16.9375 9.8753 13.2040 6.1340 7.1120 8.8229 4.7925 15.0032 11.9726 9.8157 6.7730 11.0163 9.2915 4.4533 14.1811 8.5240 9.3765 10.8769 12.4894 8.6165 7.1520 5.5695 12.8300 4.4900 10.0929 Draft June 9, 1987 �181 Analysis of Biotelemetry contributing Page 131 Data the most area to the polygon), each of the points on the polygon boundary are deleted one at a time, with the area attributed to that point being the difference polygon and a new polygon constructed nal polygon that contributes cedure is then repeated in area between with the one data point eliminated. the original The location on the origi- the most area is selected as the highest ranking location. The entire pro- for the polygon formed when the highest ranking point is eliminated, tinues in this fash.ion until enough points have been ordered and con- to form the 95% polygon - 5% of the points are ranked. The inadequacy of the above procedure is demonstrated but are far from the majority of locations (Fig. VII-2). when two locations are closely spaced. The area contributed to the polygon by each of these two outlying locations is small when the other point is still included in the polygon boundary. That is, exclusion of one of these two points, but including the other, eliminates area from the polygon. only a small sliver of To effectively eliminate the two points from the analysis, they have to be simul- 16 .........•• 15 --------- -------- -- 14 \ ...) ("t) C • • I. C) \ • • \ \ \ • • • • •• • • • •• ~ • •• • •• • • • ~'~ ~ • • • • • • • • • • • • • 12 11 '"0 0 0 \ • 13 Q.) - ~\.... \ \ 10 9 ~ \ , \ 8 7 6 1 3 5 7 X 9 11 coordinate 13 15 17 19 Fig. VII-2. The problems encountered when only one location at a time is deleted from the minimum area polygon. Both of the outliers must be deleted to remove the area inside the dashed line triangle. Draft June 9, 1987 Chapter VII - Home Range Estimation �132 Analysis of Biotelemetry Page 131 taneously deleted. Therefore, to construct points in the data set must be considered. a 95% polygon, all possible combinations For example, to construct points, all possible sets of 5 points must be eliminated, lated. Data of 5% of the a 95% polygon for 100 data one group at a time, and a polygon area calcu- Note that if none of the 5 points are on the polygon boundary, then the polygon area is the same as the original, and so the area doesn't actually have to be calculated. rithm that finds the smallest convex polygon containing a proportion this approach, Hartigan (1987) gives an algo- (0:) of the sample locations. the sample size bias of the minimum polygon estimator could be eliminated. The minimum convex polygon area also does not allow for the time correlation the locations With are assumed to be statistically home range may result if an inadequate independent. of the data, i.e., As discussed above, a biased estimate of sample is taken. Even though the minimum convex polygon allows a .••• ide range of shapes, the method often leads to spurious results when a concave shape is needed. Fig. VII-3. estimate. Minimum Consider as an example an animal with a home area polygon that includes part of the lake (shaded Chapter \,11 - Home Range Estimation area) in the home range Draft June 9, 1987 �183 Analysis of Biotelemetry Data Page 133 range around the edge of a lake (Fig. \-11-3). A minimum convex polygon estimate includes part of the lake, even though a terrestrial animal would not inhabit this area. The obvious method of correcting the polygon concave. on the boundary. 4). To understand the defect of the minimum area polygon method is to make However, some objective rule has to be made regarding which points are to be Every point in a set of data can be on the boundary of a concave polygon (Fig. VIlthis, consider the mean of the x's and v's of a set of locations as the center. Then start with any point and begin working in a clockwise direction, connecting the current point to the next '. point encountered through 36cr(i.e., as the band of the clock pivots around the mean location. After the hand has swept entirely around the clock face), the original data point closest to the mean is encoun- tered again, and the polygon is closed. 16 '15 14 13 ~ .._) (t !: .....• ~ :... 0 0 (..l ~ 12 11 10 9 8 7 6 1 3 5 7 9 11 13 15 17 19 x coordinate Fig. VII-4. Concave polygon that connects all the locations taken on the animal. Draft June 9, 1987 Chapter VII - Home Range Estimation �134 Analysis of Biotelemetry Page 13..t Data In the example shown in Fig. VII-3, the researcher could quite objectively decide that the lake area in the minimum convex polygon should be excluded. Thus, a concave polygon could be derived on an objective (and thus repeatable) biological merits for determining basis. However in general, an objective procedure that is based on a concave polygon does not exist. In other words, we may only allow a concave polygon that has acute concave angles greater than 30°. Thus, many of the interior points in Fig. VII-4 would no longer be on the polygon boundary. However, the choice of 30° is no better than some other angle and is not based on biological grounds. Of course, a convex polygon is based on a choice of 0° angle (i.e., a convex polygon), and thus may not be any more appropriate angle. A very interesting O'Callaghan approach concerned (1974). The input parameters with human are specified to the computer the degree of concavity allowed in the home range polygon. could be prescribed, these procedures perception than some other of shapes is given by algorithm, which determines If biologically motivated decision rules would offer potential for home range estimation. Although the minimum area polygon estimate of home range size does not require the data to be uniformly distributed within the polygon, the characteristics how the data are distributed equally frequently, data against grammed distribution using randomly generated (1985) has been made in that the random the nearest telemetry locations are uniformly distributed, location should follow an exponential these distances is compared to the expected distribution Chapter uniform distribution would not be rejected, VII - Home Range Estimation This test has been pro- to the procedure locations are only generated proposed by within the then the distances from random points to distribution. The observed distribution with the Cramer-von For the data in Table VII-I, the WZ statistic is 0.157 (P bivariate (1985) describe a test of the locations. in the SAS code in Listing 1 of Appendix 7. A modification If the telemetry to be discussed can- means that each section of the home range is used i.e., there is no center of activity. Samuel and Garton Samuel and Garton polygon. whereas some of the other estimators under a uniform distribution a uniform on within the polygon. The minimum area polygon can estimate home range size for data that are uniformly distributed, not. Data distributed of the estimate will depend somewhat = of Mises statistc (WZ). 0.132). Thus the null hypothesis of a even though the data are known to be from a Draft June 9, 1987 �135 Analysis of Biotelemetry bivariate normal Data distribution. Page 135 This result emphasizes the low power of this goodness small sample sizes. To perform goodness of fit tests with reasonable of fit test for power, much larger sample sizes are required. Bivariate Normal Models Jennrich and Turner Estimator Hayne (1949) first suggested the use of a circle to estimate home range of animals captured on trapping grids. The approach was greatly improved by Jennrich and Turner (1969) when the circle was generalized to an ellipse. The underlying spatial model assumed in the Jennrich is a bivariate normal probability cally and independently to move randomly distribution. distributed Thai is, location vectors (x;, y;) are assumed to be identi- according to a bivariate normal model. about their home range, with the most probable center - the mean x and mean y location. and Turner approach Thus, animals are assumed location (mode) Then, a 95% (or any other percentage) being the very ellipse is calculated around the mean location, and the area of this ellipse is an estimate of the animal's home range (Fig. VII-5). To calculate the home range ellipse from the n pairs of (z, )';) coordinates, and covariances must first be calculated. the means, variances, These quantities are " D; ;=1 X= --, n " D; ;=1 y= --, n n ~(x, _X)2 52 - ;=1 :% (n - 1) " ~(}'; -Y)2 52 1/ = ;=1 (n - 1) Draft June 9, 1987 , and Chapter \11 - Home Range Estimation �136 " Page 1.36 Analysis of Biotelemetry 16 D:l!a I i • I 15 ....J 14 I 12 Co) ..;; ("j 11 -=~ •..~,... 10 • 0 o >. • • 13 9 • • • • • • • • •• • • •• • • • • •• I • • • • •• • •• • • • •• •• • • • • ,. 8 7 • 6 1 Fig. \11-5. VII - 1. 5 3 Jermrich-Turner 7 13 11 x coordinate 9 bivariate normal ellipse estimator 15 17 applied to the simulated 19 data of Table 2:" (Xi - X) (Vi - Y) S%Y i=1 = (n - 1) Iennrich and Turner (1969) suggest the use of n - 2 in the denominator of s;, s;, and s%Y to provide unbiased estimates, as opposed to n - 1. The covariance matrix. r;, is defined as f: = [S; S~j S S2' %Y Y with the correlation of x and y defined as The 1- Q confidence f:, ( l:t 11/2) home range area is then calculated as the square root of the determinant times", times the appropriate Xrl-a)(2) of statistic for the 1-0: level with 2 degrees of freedom. Thus A For Q = = Chapter 1f' 1 f: 11/2xrl-a)(2) . 0.05, a 95% confidence ellipse is obtained, and VII - Home Range Estimation X[1-a)(2) = 5.99. Draft June 9, 1987 �137 Analysis of Biotelemetry Data For the data presented Page 137 in Table VII-I, S:tJj = -1.2167, with n - 1 used in the denominator x = 10.138, Y = 10.347, s; of s;, s~, and s:tJj. Thus = 11.778, s; = 2.573i, and r; is • _ [11.778 -1.216~ -1.2167 2.S737J' r; = with r = 0.010106 -0.2210 andA ha. The value of A only represents the area of the home range estimate. on a map showing the (.x;,Yi) locations (Batschelet S~(X requires 1981). For the ellipse centered at -x? - 2s:tJj(x -X)(y -J) + s;(y _J)2 Before the general case can be considered, To actually plot the ellipse the use of the quadratic equation of the ellipse (x,Y), the quadratic equation is = X(1-a)(2) . 2 special cases must be checked. If s; = s;, the ellipse is a circle with radius (X(l - ;)(2) )1". If s; < s~ and s:tJj = 0, the major axis of the ellipse is parallel to the Y Sll axis. Excluding these special cases, the follo-wing quantities are calculated: R = res; - s;? a = [(S; b [ + s; + 4(s •••?r" \R a)(2l" , )X[l - (s; + s; -: 8 = arctan , )X[l - a)(2)]," , and 2 -2s: sy-s:-R Here a and b are the semi-axes (a > b), and 8 is the angle by which the major axis is inclined versus the For computer x = x + a cos,p Y =Y + plotting (Batschelet 1981:264) uses the parametric equations of the ellipse cos 6 - b sin,p sin 6 , a cos,p sin 6 + b sin,p cos 6 . Here, 1J; is a variable angle increasing from 0° to 360° in small steps. Batschelet a convenient step length for computer Draft June 9, 1987 (1981:265) suggests that plotting is 4°. Using this choice, the ellipse is created from 90 Chapter VII - Home Range Estimation �la8 Analysis of Biotelemetry Page 138 points. For tbe reader desiring a better understanding this subject in Batschelet That is, the 95% ellipse estimated study. Koeppl et al. (1975) suggest a)(2, ,,- 2) where F (1- (2, n - '2). The F-statistic but is estimated able sample sizes that the X(l- value should a)(2) is an F-statistic be replaced at level (1 - ex) and degrees (Xf1-0.Cl5)(:2) = of freedom 5.99). n F (1 _ 0.(0)(2, 5 10 20 30 9.55 4.46 3.55 3.34 42 3.23 62 122 3.15 3.07 3.00 ,,- 2) is almost 2 x larger when n 2(n - 1) F (n - 2) 2 X(l - O.Cl5)(2) 25.47 10.04 7.49 6.92 6.62 6.40 6.19 6.00 5.99 5.99 5.99 5.99 5.99 5.99 5.99 5.99 = 10 for ex = 0.05. The disadvantage of using the of Koeppl et al. is that the expected home range size is now a function of the sample size and Marcus 1978), just as the minimum area polygon was shown to be (Jennrich home range size. The sample size is obviously important in determining for estimating the 95% confidence VII - Home Range Estimation interval the expected tbe quality of the home range but it should not enter into the expected size of the home range. size should be used to calculate and Turner if the purpose of estimating A was to have 95% probability of including the next location taken on the animal, but it is not appropriate Chapter with the value takes into account the fact that the ellipse center (X;Y) is not known exactly. 1969). The F-statistic would be appropriate estimate, from study to from the data. As the following table shows, very little difference occurs for reason- The F-statistic estimator (Madden over of home range size is not a function of sample size. Thus, home range size is much more comparable a){2, ,,_ 2) 00 F-statistic (1969) provides a great improvement from 100 data points is expected to be the same size as the 95% from 500 data points. l(n-l) n _ 2 F (1- and Turner area polygon because the estimate ellipse estimated of the geometry of an ellipse, see the chapter on (1981), and also his chapter on bivariate methods. The bivariate normal model of Jennrich the minimum Data As shown below, sample of the home range estimate, which Draft June 9, 1987 �139 Analysis of Biotelemetry Data Page 139 provides an appraisal of the quality of the estimate. For the data in Table VII-I, the Koeppel et aI. estimate is 0.01079 ha. The programs DC80 and McPaal both provide this estimate instead of the Jennrich and Turner estimate provided by HO;\fER. Dunn and Brisbin (1982:46) have developed an equation to estimate the confidence interval for A: P[(JI :5 (2Jt - 4)Alxrl)(2n-4J] - 4)Alxfl-.)(2n-4):5A = where I + u Q in terms of usual :! percentage = 1- points. a, Nonsymmetric provide a shorter interval, but generally a symmetric interval (u = I = confidence intervals (1.1 f I) (/2) will be desirable. Thus a 95% confidence interval for A is ( a = 0.05 ) (21 - 4)A 2 Xtl-a/2)(2n-4) (2Jt - 4)1 A <A< 2 . X(a/2)(2n-4) For the data in Table VII-I, the confidence From these expressions interval is 0.007760-0.01371 ha. Dunn and Brisbin (1982:46-47) have derived an estimate size (n) needed to estimate A such that A bas probability of the sample at least 1 - a of incurring a relative error less than r: n2: where Zrl - a/2) r 2 Z[l _a/2) +2, is the 1 - a/2 percentage point of the standard normal distribution. Take as an example (from Dunn and Brisbin 1982:47) a = 0.05 and r ~ 0.10, i.e., A is desired to be within = 10% of the true A with probability 95%. Then fl > (1.96)2 + 2 ::: 386 - . (0.10)2 For r = 0.20, n 2: 98. Thus, these results stress the large sample size needed to adequately determine the home range of an animal. an estimate with a confidence Draft June 9, 1987 As a general rule of thumb, at least 100 locations are needed to achieve interval of =20%. Chapter VII - Home Range Estimation �190 Page 140 Analysis of Biotelemetry Data Weighted Bivariate Normal Estimator Samuel and Garton (1985) have proposed a modification of the Jennrich-Turner ellipse estimate where each data point is weighted by the distance it is from the mean of the locations. matrix notation used by Samuel and Garton in describing this estimator. We will follow the The weighted distance is cal- culared in terms of the probability of this particular value occuring this far or farther from the mean location: d·, = [(X; - x*),(S.zt1ex; - X*)]l12 where d·i is the weighted Malalanobis distance for location S"'z is the variance-covariance matrix of the locations. were missing in the equation presented variance-covariance x* = X;, x* is the vector of weighted means, and Note that several • denoting weighted estimates by Samuel and Garton (1985:519). The weighted mean and matrix are calculated as _i=_l__ and " ~ L:w; (X; - x·) (X; - x*)' ,=1 S*: = --------- n L: w, ;=1 where w, = I {2/d; 1 for d, > 2 for d, S 2 . Thus points close to the mean are weighted greater than those far away, and particularly outliers are weighted very Jow, to the extent that they will no longer affect the estimate. estimator To make the more stable, points within 2 standard deviations are all given the same weight, with any point beyond 2 standard deviations given less weight depending on its distance from the mean. The weighting of the individual data points makes the estimator mator is referred Chapter important, to as a robust ellipse estimator. VII - Home Range Estimation robust to outliers, hence the esti- The SAS code to generate this estimate is provided Draft June 9, 1987 �191 Analysis of Biotelemetry Data Page I-U in Listing 3 of Appendix 7. x= For the data in Table VII-I, 10.171, Y = s; = 10.604, s; = 1.8918, and 10.368, giving an area of 0.008240 ha (95% CI 0.006328-0.11750). Sry = -0.9476, Thus the weighted ellipse estimate is smaller than the unweighted estimate. Multiple Ellipses Don and Rennolls bution that incorporates (1983) have proposed biological assumes that the coordinates tree. by the variance) at each of the attraction for attraction of the attraction points, a l-dimensional points. in their paper). Their model den provides the relative use ("strength") and Because a circular normal distribution is fit variance is estimated. asymmetry around each of the attraction metric home range estimator. "nuclei" points are known, e.g., the location of a squirrel's points, their estimator points, and hence general normal distribution points (termed of the attraction Given one or more attraction range (measured attraction a horne range model based on a circular normal distri- Although their estimator of the home range, the assumption points seems inconsistent allows of a circular with the goal of a nonsym- For example, imagine the home range of a cougar whose den is at the base of an unclimbable cliff. Don and Rennol1s (1983) applied their estimator which seems reasonable unless a squirrel's den tree was next to an unuseable to grey squirrel data, area like a four-lane high- way! Dunn Estimator The problem of lack of independence of observations was solved for evenly spaced (temporal) locations by Dunn and Gipson (1977), who developed an estimate of home range from the multivariate Orhnstein-Uhlenbeck stochastic process (MOU). The MOU diffusion process describes the motion of a particle on a surface exhibiting a central tendency. + .6.t is recognized estimator The probability distribution to be a function of the location at time t (i.e., Markovian), of a particle at time t and thus this home range allows for the time series nature of the data. Locations closely spaced in time are recognized to not provide as much information Draft June 9, 1987 about home range size as points further Chapter apart in time. Relaxing VII - Home Range Estimation �192 Analysis of Biotelemetry Page 1~2 the assumption of independence Data by modeling the time series nature of the data is a valuable contribu- tion to home range estimation. Dunn's estimator but the time sequence requires the assumption of the observations are considered, fixes along the animal's path is recognized. information lute time or location. i.e., the correlation Two observations about the home range as two observations matrix between two observations close in time do not contribute as much The covariance taken a fixed amount of time apart is constant, regardless of the abso- stable over time and space, i.e., the tern- are homogeneous. Input data for the Dunn estimator to be independent, between the successive taken further apart in time. Hence, the home range is considered poral and spatial dimensions assumed that the animal is using its space as bivariate normal, consist of groups of observations, or "bursts". Bursts are i.e., enough time has elasped from the end of the previous burst to the start of the current burst that the two observations are statistically independent. . tor is currently developed, the time interval between observations in time. That is, the time interval between observations As the Dunn estima- within a burst must be equally spaced is the same. This limitation of the estimator can be removed, but will require significantly more computer time. An example is shown in Fig. VII-6 comparing In this case, the two estimates ha). The randomization gesting that significant the Dunn ellipse with the J ennrich- Turner ellipse. are nearly identical (Jennrich-Tumer test conducted by HOMER time series correlation ellipse 55.2 ba, Dunn ellipse 51.1 (Appendix 5) was not significant (P was not present in the data. = 0.853), sug- Thus we would not expect large differences in the estimates from these two home range estimators. Testing Bivariate Normality Conceptually, Animals bivariate do not move randomly pose to obtain resources normal models do not properly model the movements about a central point on a homogeneous available within their home range. The movement of animals. surface, but move with purassumed with the bivariate normal model is random except for the tendency to stay to the middle - there is no purpose or predictable direction to the movement. Chapter Also, there is no biological reason why an animal is expected to spend VII - Home Range Estimation Draft June 9, 1987 �193 Analysis of Biotelemetry 4391800 Data Page 143 j r <, I , ( I 4391600 .- ~ 4391400 . ~•... 4391200 c c \ \ • ,, , I \ \ I I I Dunn , \ I, \ \ Jennrich- I~ , t •• " I • Turner .1 • , I I , I~Jn., , 1'1 \ , \ ~I , I, ' \ \ 4390800 \ 1.. , ,. I I I I \ \ o ~ 4391000 \ . · ,t:\ _J -, \ , I I .1J I / / " -, -, / 4390600 0746700 0746900 0747.' 00 X 0747300 0747500 coordinate Fig. VII-6. Comparison of Dunn and Jennrich-Turner ellipses for a female mule deer summering on the Roan Plateau, Piceance Basin, Colorado. the most time at the exact center of its home range. The most likely location may be a den site, which is unlikely to be located at the mean x and y locations, as assumed by bivariate normal models. Further, the elliptical shape assumption will often be violated. The example data given in Fig. VII-3 would produce erroneous estimates if used with any of the ellipse estimators. Smith (1983) provides a chi-square goodness of fit test to evaluate whether the home range data are consistent with the assumption of bivariate normality, although the calculations shown in the paper are incorrect. The test looks at the frequency of locations within a series of concentric ellipses (contours) representing equal probability of occurrence (Fig. VII-1). That is, the test is analogous to a chi-square goodness of fit test for continuous univariate distributions, but with cells replaced by contours. The SAS test for this test is provided in Listing 1of Appendix 7. Draft June 9, 1987 Chapter VII - Home Range Estimation �194 Page 14-1 Analysis of Biotelemetry 16 15 .., 14 .,I .\. . ........ , ...', 313 212 '-11 v ' ;... cl0 c CJ 9 >, 8 I ./ ,~-------------_.------ .. I • J \ -, '-, • '" ...... -- ,- ~:"-"- .--.-~.- II. Ir', 11-- " .• ', -, .- ... ..-------.....- - • "", , .. --__. '_- , • ,' • • I '\ '\. • \. 1 / ..' - - - - - - - -- -- ~_J ( 80% 60% 40% 20% • I Data • I '" ....• • 6 5 4 1 7 5 3 9 11 13 x coordinate 17 15 19 Fig, VII· 7. Concentric ellipses for probability levels of 20, 40, 60, and 80% are shown plotted over the simulated data from Table VII-!. The goodness of fit test suggested by Smith (1983) depends on the number of observed locations within each of the areas defined by the concentric ellipses. For the simulated data in Table VII-I, the observed frequencies count these hypothesis locations from of a bivariate Fig. normal VII-7. The distribution expected are 12,8,11,9, frequencies would be 50/5 = = for each cell under 10, i.e., the 50 locations divided equally into the 0-20, 20-40, 40-60, 60-80 and 80-100 percentage null hypothesis is not rejected (P and 10. You can classes. the null should be For these data, the 0.911), suggesting that the data fit a bivariate normal distribution as is the case. Samuel and Garton (1985) provide another test of bivariate normality that is more powerful than the test suggested by Smith. If the data follow a bivariate normal distribution, then the squared proba- biliry distances from the mean of the data (i.e., the squared values of d, described above) should follow a x? distribution with 2 degrees of freedom. can be constructed to accompany Further, the robust estimate Chapter VlI - Home Range Estimation a robust version of the test using the d*; values of home range size proposed by Samuel and Draft June 9, 1987 �195 Page 1~5 Analysis of Biotelemetry Data Garton (1985). SAS code for this test is provided in Listing 3 of Appendix 7. For the data in Table VII-I, the unweighted goodness of fit test gives a 0.15). The weighted 'W'l value is 0.214 (P = 'W'l value of 0.116 (P > 0.062). Thus the weighted ellipse does not appear to fit the observed data as well as the unweighted ellipse. To illustrate that animal home ranges often do not follow a bivariate normal or bivariate uniform distribution, the following results from 38 mule deer home ranges are reported. The entries in the table are the number of home ranges that met the (P < 0.10) criterion: Goodness of fit Result P> 0.10 P < 0.10 Bivariate Uniform 2 36 Distribution Bivariate Weighted Bivariate Normal Normal 1 3 37 35 Thus we conclude that for these mule deer, neither the bivariate uniform nor the bivariate normal dis-tributioas is a good fit of the data. One example from this set of data is shown in Fig. VII-8. The impact of a few outlying locations causes the polygon estimate to differ substantially from the ellipse. Further, the effect of these outliers causes the ellipse to extend in the opposite direction from the outliers to compensate for their impact on the shape of the normal distribution. The bivariate normal distribution requires that the utilization distribution be symmetrical, and thus forces the ellipse out into vacant area opposite of the outlying locations. The bivariate normal goodness of fit test suggested by Samuel and Garton (1985) rejected the null hypothesis (P < 0.01), as did the concentric ellipse test suggested by Smith (1983) (P < 0.01). The goodness of fit test for the bivariate uniform distribution suggested by Samuel and Garton (1985) also rejected this hypothesis (P < 0.01). One of the pitfalls of testing a set of observations for fit to a distribution is that a small sample size precludes the test having any power. When the null hypothesis is accepted, the conclusion is not that the data fit the distribution, but rather that too few observations were taken to reject the null hypothesis. Draft June 9, 1987 Chapter VII - Home Range Estimation �196 Analysis of Biotelemetry Page 146 Data _-----_ 4391800 ". ,- ". I / I 4391600 / / \ I Q) ._; C'j .-- ..... ..,..... \ \ \ 4391400 ._, ;.... 0 0 \ 4391200 C,.) \ I I \ I \ \ • •• \ ~ \ 4391000 \ I I I J I I I •• \ \ 4390800 I \ ,,• • "'I J I I •• •.... " .•.. I I -- 4390600+---~--.---.---~--.---.---,---,---~ 0746600 0746800 0747000 0747200 0747400 X co or-d in a te Fig. VII-8. Impact of an asymmetrical utilization distribution on tbe Jennrich-Turner ellipse estimator. Although animal locations seldom fit a bivariate normal distribution, the use of the bivariate normal model for home range estimation is still worthwhile. The concept of a probability model to describe animal movements is practical. The ease of calculation, particularly the Jennrich-Turner estimare, and the availability of programs (Dunn 1978) make their application easy compared to other approaches. Also the many enhancements made by Dunn (1978) and Dunn and Brisbin (1982) are guaranteed to see the continued use of this model. Nonparametric Approaches Grid Cell Counts Methods utilizing the counts of location within grid cells or a map of an animal's home range (Fig. VII-9) are truly nonparametric approaches to home range estimation. No assumptions are made relative to the shape of the area utilized. The area over which an animal has moved is dissected by a Chapter VII - Home Range Estimation Draft June 9, 1987 �197 Analysis of Biotelemetry Page I-t7 Data grid into cells, or blocks. The number of animal locations are tabulated for each of these cells, and tbe sum of the areas of cells containing locations is taken as the estimate of home range area. Several serious problems arise in the practical application tend to be added together disappear to form the home range estimate. from one location and reincarnate tions happened to be taken. One approach ing two consecutive of this method. Obviously, the animal did not suddenly at the next, but it moved across the areas where no loca- Thus, the disjointedness to correcting Disjoint areas (areas not connected) is due to sampling precision. the problem is to assume that cells crossed by the straight line join- locations should also be counted as part of the home range area. However, use of a straight line is artificial, and selection of the maximum time interval for which locations will be connected is subjective. Grid cell methods have been' utilized for the analysis of movement Cedar Creek Natural History Area (SinifI and Tester 1965). These estimates avoid the problems tioned above because the locations are taken on animals every 45 seconds (Cochran the straight line assumption does not appear unreasonable, second length does not seem too subjective. and filling in cells for time intervals of 45 However, many times the data were sampled 1969, Rongstad (snowshoe bares) are used. Why use these values as opposed to 30 minutes or 5 minutes? Rongstad and Tester (Rorigstad 1971) and thus time intervals of 1 hour (deer) or 15 minutes of SUbjectivity. Also home range estimates sampling intensity of the experiment men- et al. 1965). Thus, and Tester method raises a question and Tester data from the do not seem to be a reasonable (1969) also describe procedures filled cells are also counted in the home range estimate. Again the that are direct functions of the approach. where cells within 2 units of distance of Again a subjective estimate of the connecting distance of 2 cells must be made. Complete counts of grid cell areas do not allow for removal of the outliers, i.e., a 95% confidence area is not defined as described above. However, this criticism. is easily overcome cumulative area versus the cumulative number of fixes can be constructed. any other percent) Draft June 9, 1987 because a plot of The area at which 95% (or of the fixes are included is thus a 95% area estimate. Chapter VII - Home Range Estimation �198 Page 148 II Analysis of Biotelemetry 2 15 I 0 14 I 0 4 3 I 5 6 7 8 9 10 11 12 13 14 15 16 Data 17 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 131 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 12 0 0 0 0 1 2 0 0 1 0 2 0 0 0 0 0 11 0 0 1 0 1 1 0 0 3 3 2 1 0 0 0 1 10 0 0 1 0 0 1 2 2 1 1 1 1 1 1 1 0 9 0 0 1 0 0 0 1 1 0 2 1 1 0 0 1 0 8 0 0 0 0 1 0 0 1 0 0 0 0 1 0 0 0 7 6 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 o· 0 0 0 0 1 0 0 0 0 0 5 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Fig. VII-9. Illustration of how a grid cell home range estimate can be calculated by summing the areas of the cells that contained one or more animal locations. For these data, the cell size is 1 m2, giving an estimate of 0.0038 ha. Data are listed in Table VII-I. The greatest problem with the grid cell approach ing the number of grid squares to draw on the map. Choosing too coarse a grid produces coarse esti- mates and tends to overestimate home range size. Choosing too fine a grid mesh produces mates of home range area and aggravates the problems causes many more disjoint areas to be connected an arbitrary is selecting the grid square size, i.e., determin- mentioned above. than a less fine grid mesh. decision for which no biologically based, objective procedures underesti- That is, too fine a grid Choice of grid cell size is are known. Possibilities include setting grid cell size based on the average (or median) distance between consecutive locations. However, the level of arbitrariness is just adjusted down to the assumptions of the rule, rather tban at the level of the data analysis. Fourier Series Smoothing Anderson (1982) has presented bivariate Fourier series. to be smoothed Chapter approach to estimating home range utilizing the Fourier series have often been used in statistics to smooth data. The function is decomposed For home range estimation, (x,y) coordinate a nonparametric into a series of sine and cosine components of different frequencies. the bivariate functions consist of the x-y plot with a "spike" plotted at each where the animal was located. VII - Home Range Estimation The two-dimensional Fourier transform smooths these Draft June 9, 1987 �199 Analysis of Biotelemetry Data Page 149 spikes to form a surface representing lest area encompassed the animal's use of area. by 95% (or any other percentage) Home range is calculated as the smal- of the volume of the Fourier transform sur- face, just as with the bivariate normal model. Anderson' found that the main problem with the use of the Fourier transform the edge of the home range are not estimated the home range are where few locations highly variable for large percentages the home range has to be incorporated Anderson was that areas at very well because of lack of data. That is, the edges of are taken. Therefore, the estimate of home range size is (such as 95%) because the poorly estimated surface at the edge of into the calculation of volume. (1982) suggests that a more reasonable approach is to utilize a 50% volume as an esti- mate of home range. The impact of the edges is much less because 50% of the volume lies near the center. as small as 50% seems to be a rather narrow description A percentage (1943:351) definition of home range. However, there is no biological justification this figure has probably been adopted by biologists because of the use of a: Thus, the selection experiment of any percentage can probably be justified data simulated of the Fourier estimator for the use of 95%; 0.05 in statistical tests. if the value fulfills the needs of the and the Jennrich-Turner from a bivariate normal distribution, from a bivariate normal distribution, in this situation. (Anderson the variance, or range of estimates, normal ellipse for almost as well as 1982). Because the data are has a distinct advantage does so well is encouraging. were very close to the true home range size determined Further, performed the bivariate normal estimator Hence the result that the Fourier estimator of both estimators bivariate the Fourier estimator the bivariate normal method for a 50% home range estimate distribution. = in Burt's being conducted. In comparisons simulated of "normal" The means analytically from the normal was roughly the same for the Jennrich- Turner and Fourier methods. However, we would expect that the Fourier estimate would not perform as well as the Jennrich-Turner if the percent of the space used is increased because the fringe of the distribution bution assumptions Draft June 9, 1987 from 50% to 95%. This is is much more difficult to estimate, as discussed above. The distri- made for the Jennrich-Turner estimator would greatly increase Chapter its effectiveness, VII - Home Range Estimation �200 Page 150 Analysis of Biotelemetry Data whereas the Fourier estimator would have only the available data to estimate the distribution. For the data in Table VII-I, the following table lists the home range estimate (calculated with the McPaal program) as a function of the percent of future fixes to be included. Percent locations Home range (hal 25 0.002636 0.005423 0.008717 0.01125 0.01232 50 75 90 95 For this particular data set, the 95% estimate is greater than the estimates area polygon (0.007668 ha) and tbe Jennrich-Turner tions of this experiment Harmonic produced (0.010104 ha) estimators. must be simulated to understand the relationships However, many replica- between these 3 estimators. Mean Dixon and Chapman (1980) have proposed monic mean of the areal distribution. the use of a bome range estimator The center(s) dual within its home range. Thus, the estimator has some worthwhile of occurrence features of greatest of an indivi- in that home ranges with 2 centers of activity can be properly defined, and no shape criteria are imposed of area are developed based on the har- of activity is located in the area(s) activiry, The activity area isopleth is then related directly to the frequency Contours by the minimum OD the estimator. from a grid of points, where the observed area is the harmonic mean of the distance from the grid node to the observed biotelemetry locations. The farther a location is from a grid node, the less influence the location has on the mean for the node (Fig. VII-IO). specific equation to calculate the harmonic mean dg1,g2 for a grid node located atxgl,Yg2 The is 1 dg1,a2 = 1 " -; ,E Grid nodes located 1 [ext -Xgl)2 at centers + {)Ii - Yg2?]l/2 of activity have the shortest distances and hence the lowest values of dg1•g2, those located far from centers of activity have the longest distances and hence highest values of Chapter VII - Home Range Estimation Draft June 9, 1987 �201 Analysis of Biotelemetry * (x, ' J2 * Page 151 Data *) * • *Y3 ) l~~' * * d1 (Xl ' J1 ) * x * >< d2 * * * :< * B Fig. VII-IO. Illustration of the mechanics of the Dixon-Chapman home range estimator. For grid node A, the reciprical of the mean of lid!> I/dz, and Ild3 is much smaller than for grid node B. Thus when the grid nodes are used to construct a contour map, the area around grid node A will be included in a lower probability contour than the area around grid node B. dgl.gZ' The grid of values are then contoured dgl.gZ Study of the above equation immediately the telemetry undefined, location (Xt, to estimate the isopleths of activity. exposes one of the problems Yi) is identical to the grid node location and a contour map cannot be constructed. grid some small increment location is positive. from a particular (XgI, of the estimator. Ygz)? The value of dgl,gz is A logical response to this problem is to shift the (&, .6.y), so that the distance from each grid node to each biotelemetry Unfortunately, this action identifies a second problem. The contours constructed grid are unique to that grid, i.e., if the grid origin is shifted, different estimates result. In practice, distances less than 1 unit length are given a value of 1, which also aggrevates of the units of dimension used to compute the estimate. tance between grid nodes and number Draft June 9,1987 What if of grid nodes. Further, the estimator In particular, Chapter the problem is sensitive to the dis- if the units of distance are can- VII - Home Range Estimation �202 Analysis of Biotelemetry Page 152 verted and a new grid used, different achieved. Thus, practical application However, the procedure estimates of home range size (and isopleths of this estimator Data of activity) are is sensitive to both the grid origin and cell size. still can be useful in defining a map of activity, if the investigator recognizes that the map may vary somewhat in shape and is only an approximation. For the data in Table VII-I, the DCSD program calculates the following estimates. Percent locations 25 50 75 90 95 Home range (ha) 0.0006234 0.002340 0.005590 0.008704 0.01168 Ten grid divisions were used. Had only 5 divisions been used, the following estimates result. Percent locations 25 50 75 90 95 Similar answers are produced Home range (ha) 0.001528 0.002736 0.006626 0.009983 0.01236 by the McPaal program, although we have been unable to dupli- cate estimates with these 2 programs. Problems Common to all of the Nonparametric Methods All of the non parametric data. methods so far considered have. ignored the time series nature of the As pointed out earlier, animals do not move about their environment thus the lack of statistical home range estimates. terms of assumptions independence Chapter locations about shape) that handles the time series nature of biotelemetry methods area polygon: none have an associated VII - Home Range Estimation decribed here share a problem confidence interval. fashion, and may lead to nonrepresentative The greatest need in estimation of home range is a nonparametric Second, all of the nonparametric minimum of the consecutive in a random approach (in data. in common with the Thus, an estimate is produced, Draft June 9,1987 �203 Analysis of Biotelemetry Page 1::3 Data but no estimate of its standard quality and usefulness error is provided. of the estimate, Not only is the researcher but no method of determining left to wonder about the sample size is available. Thus sample size criteria have not been provided in the literature. In contrast, sample the ellipse-based estimators size can be determined. assumptions However, provide estimates as discussed of precision above, and, hence, necessary ellipse estimators require strong about the shape and structure of the use of the home range. Computer Programs for Home Range Calculation In the previous sections, we have mentioned range calculations discussed. various computer Here, we will summarize codes that will perform the home the codes that we know about at this time. --~- 1985! 4391600 4391400 4391200 I 4391000 4390800 I ¢ I 0 I o e I I I I I I I I .-~ e 00 -- ;' -..- ;' ;' ;' 4390600+---~--~--~--~--~--~--~--~---0746600 0746800 0747000 0747200 0747400 x coordinate Fig. VII-H. Summer home ranges of a female mule deer from the Piceance Basin of western Colorado that was radio tracked for two consecutive summers. The home range was estimated with the minimum area polygon. Draft June 9, 1987 Chapter VII - Home Range Estimation �204 Page 1::'" Analysis Table \'11-2. Computer codes for the 1B;\1 PC and compatibles Pro~am HOMER of Biotelemetry that provide home range estimates. ~H~o~m~e~R~a~nag~e~E~s~ti~m~a~to~r~s~ (Appendix 5) Data _ Minimum convex polygon J ennrich- Turner ellipse Dunn ellipse Siniff- Tester grid cell counts SAS Listing 1, Appendix 7 Minimum convex polygon Test of bivariate uniform distribution SAS Listing 2, Appendix 7 Jennrich- Turner ellipse Test of bivariate normal distribution (Smith 1983) Samuel-Garton weighted ellipse Test of bivariate normal distribution (Samuel and Garton 1985) SAS Listing 3, Appendix 7 ~lcP AA:L Minimum convex polygon J ennrich- Turner ellipse Fourier series Dixon-Chapman TELE\1/PC Minimum convex polygon Jennrich-Turner ellipse Dixon-Chapman DC80 Minimum convex polygon J ennrich- Turner ellipse Dixon-Chapman HO:>1ERA""IGE Minimum convex polygon Jennrich-Turner ellipse Weighted bivariate normal ellipse Dixon-Chapman For straight forward home range estimates, the McPAAL program is probably the easiest for the novice to use. Graphics mates. However, abIes is desired, are provided on the screen, and printer output can be obtained if statistical summaries the SAS procedures of the estimates offer considerable SAS is available to perform data summaries, for the esti- across animals, or other classification power. visual presentations The full statistical vari- analysis power of of the data in plots or histograms, etc. Evaluating Home Range Estimators Part of the confusion about home range estimation lack of rigor in evaluating new estimators. A new estimator methodology is proposed, has been produced but the only evidence that it works is an example set of data from an animal where the true home range is not known. Chapter VII - Home Range Estimation from the To demon- Draft June 9,1987 �205 Analysis of Biotelemetry Page 155 Data strate that a home range estimator home range is known. "works", Monte Carlo simulations The bias and the precision (1982) does a realatively good job of developing Simulations are presented Jennrich- Turner normal ellise. The estimator is the Dixon-Chapman lished in the literature the statistical comparing the Fourier estimator bivariate statistical properties of the estimator that demonstrate where the true can then be assessed. properties Anderson of the Fourier estimator. to both the minimum area polygon and the most notably lacking simulations harmonic mean estimator. this estimator are required No simulations will provide unbiased to verify its have been pub- estimates for a known home range . . Part of the problem in demonstrating distribution of animal movements. extremes of the reasonable If an estimator and study the performance performs well at both ends of the spectrum, distribution defined, going out to 00. performs Thus we recommend formly distributed, points. that estimators i.e., no attraction to produce bivariate normal distributions irregular home range size can be calculated for a heavily with the boundaries are not of the of locations within the home range are uni- To achieve various shapes of home ranges, both convex shapes. Anderson distributions and mixtures of bivariate normal (1982) simulated can be developed. analytically, so that the bias and precision assessed. We feel that an extensive set of computer of their statistical properties be simulated through a mixture of three in his Fig. 6a. Thus a range of shapes and utilization used home range estimators. the two at the extremes. then it should perform reasonably and concave polygons can be simulated with the uniform distribution, Draft June 9, 1987 of the estimator At the other extreme is the uniform distribution, and the distribution combined is to simulate such as the bivariate normal, where the edges of the distribution home range are explicitly delineated distributions well is selecting the underlying We suggest the solution to this problem distributions, the middle range of distributions. center-weighted that an estimator Continued application simulations In each case, the true of an estimator can be are needed to evaluate the commonly of the estimators without a better understanding will only further cloud the issues relative to home range sizes. Chapter VII - Home Range Estimation �206 Analysis of Biotelemetry Page 156 Data Similarity of Home Ranges An approach common to measuring two or more home ranges. fidelity and animal association In the case of fidelity, home range estimates animal between two or more time periods. or more animals are compared. For measuring Continued for measuring VI) is to compare are compared animal association, for a single the home ranges of two data collection over several time periods or animals pro- vides a series of home range maps that can be compared on the assumptions (Chapter of the home range estimation for similarity. procedure. fidelity with data from mule deer summering The approach Figure VII-ll depends heavily illustrates this approach in the Piceance Basin, Colorado. A similar approach was used by Tierson et a!. (1985). Although summering not quantitative, illustrates the fidelity of this deer to a specific site. However, the influence of a few locations in 1985 greatly affects the estimate of over- lap, hence fidelity. the reader's the figure adequately As shown in Fig. VII-12, use of a Jennrich-Turner perception of the overlap of the home ranges. range overlap using the bivariate normal estimators Another ellipse estimator would change problem with estimating is that the utilization distribution home should be taken into account, making the estimate of overlap difficult to calculate for the bivariate normal ellipse. That is, the concentric probability ellipses should be used to estimate overlap, i.e., the integral of the utiliza- tion function, not just the amount of overlapping area of the two ellipses. The second disadvantage a selected number of presenting fidelity data in this manner is that usually only data from of animals can be presented, rest of the instrumented population. Quantifying sive periods allows the fidelity of all instrumented Many applications leaving the reader wondering in the literature the amount of home range overlap between succesanimals to be presented such as required Chapter can be performed For example, the MRPP test of Mielke and Berry et al. 1985) can be used to test for shifts in an animal's area of utilization, for testing for fidelity or association. (where both lack measures in a table. which have used home range estimates without actually calculating a home range estimate. (1982) (also see Zimmerman about the fidelity of the of precision), VII - Home Range Estimation Instead of comparing the method is testing whether 2 home range estimates the 2 sets of locations came Draft June 9, 1987 �207 Analysis of Biotelemetry Data Page 1::7 • 4391800 198~~, --.,--1985 .......... // / , / -, / / I 4391600 I o I / I c.> ...,_) cj $..0 I 4391400 0' 0 4391200 0 0 U .. • I c:: . "'0 0 , • 0 ••• • \ ° \ \ \ \ 0 \ \ \ I I I I I I I 0 I~ I e • 00 ~ '.»•• • e o III CI . • 1 1 °'. 01. 00 °li I "-~o °° 0 \ 0 o~" I I \ ~ \ ! · 1:*-1, \ 4391000 \ II 0 I. ·0 -, \ 0 \ 0 0 \0 ° -, \ <¢ OJ; II I I I I I \ \ \ 4390800 r ~r:10 -, ~~ 0 0 I I 00 0 I • I 0 00 I 0 () ,. .,- ". ------4390600 0746600 0746800 0747000 X 0747200 0747400 coordinate Fig. VII-12. Summer home ranges of a female mule deer from the Piceance Basin of western Colorado that was radio tracked for two consecutive summers. The home range is estimated with the JennrichTurner bivariate normal ellipse. from a common distribution. of the underlying distribution. The approach is nonparametric, making no assumptions The null hypothesis of the test is that the two (or more) utilization tributions are the same. The MRPP statistic is based on the within-group measures ignored. between locations about the shape compared The method is programmed to the average distances between dis- average of pairwise distance locations when groups are in Berry (1982). For the data shown in Figs. VII-ll and VII-l2, the MRPP statistic is not significac: (P = O.???), suggesting no large shift in home range as a function of year. Usefulness of the Home Range Concept There is no single preferred if a method estimator of home range. All estimators have their faults. However, must be used, there are some criteria by which the most appropriate Draft June 9, 1987 Chapter method may be VII - Home Range Estimation �208 Analysis of Biotelemetry Page 158 Data selected: 1) Time series randomization test: if each point is independent, secutive points should be the same regardless the average distance between con- of the order in which they are entered 1981). Swihart and Slade (1985) have extended this test by making additional 2) the underlying statistical distribution of the locations. Shape criterion: goodness apply a chi-square of fit test to determine (Schoener assumptions whether about the home range shape is close to the expected shape (Samuel and Garton 1985, Smith 1983). 3) Sample size: the large sample size required for nonpararnetric in a particular study. Estimators Home is a concept range estimators requiring additional assumptions that is frequently abused. may preclude their use may be preferred. Some of the problems go back to its definition as the area used by the animal in its normal activities (Burt 1943:351). What is use? What is normal? Normal is frequently defined as 95% of the time, but with no biological justification. Another approach to eliminate the need for a home range estimate is to use the raw locations to test the hypothesis of interest. As an example, home range size may be desired because it is a measure of an animal's energetic costs. That is, the larger the home range, the more energy expended in searching for food. However, the distance between consecutive locations (corrected for time differences) would also measure energetic costs, and does not suffer from the many assumptions a home range estimate. rather than comparing Comparison of distance home range estimates. example of where the problems required to obtain moved for 2 groups of animals seems preferable The use of the MRPP test described of home range estimates above is another can be avoided by applying statistical tech- niques to the raw data rather than comparing home range estimates. Finally, before computing Often home range estimates designed and tested. information should consider are the only product from a biotelemetry Home range estimates are a poor substitute data from "binoculars VII - Home Range Estimation what will be learned. study because no hypothesis was for good experimental about why an animal moves is provided in a home range estimate. more than just observation Chapter home range, the investigator that see in the dark". protocol. No Wildlife biologist need Wildlife science has moved Draft June 9, 1987 �209 Analysis of Biotelemetry Page lS!) Data beyond this point in its quest for knowledge. Summary Chapter VII 1) The concept of a home range is useful to biologists. into a rigorous statistical procedure lost when actual estimates is not easily done. understand 2) used estimators the estimate. are unknown, of the concept Most of the benefits of the concept are because of the many subjective Estimates and thus provide little biological insight. of the commonly implementation of home range size are constructed decisions that must be made to construct construction, However, Further, and Monte lack objective criteria in their the statistical properties Carlo simulations of most are needed to them where the true home range size is known. Because home range is a function of time as well as space, sampling time in a representative fashion. procedures Home range estimates should be reported must sample in the context of the time frame of the sample. 3) A serious limitation of most home range estimators a confidence interval estimator. is the lack of a precision estimate and hence Statisitical estimates that lack precision estimates provide a false sense of security to the user. 4) Often home range estimates research being conducted. are developed because the researcher The hypotheses to be tested were not formulated tion, so home range estimates research has not carefully designed the provide a quantitative summary of the data. Careful design of because a statistical analysis can usually avoid the pitfalls of home range estimation, based on the raw data can be developed to test important not used in the analysis, and thus the assumptions hypotheses. they require prior to data collec- Home range estimates are not needed, are and the biases inherent in the technique are avoided. Draft June 9, 1987 Chapter \11 - Home Range Estimation �210 Analysis of Biotelemetry Literature Data Cited Chapter VII Anderson, D. J. 1982. The home range: a new non parametric 112. estimation technique. Ecology 63:103- Batschelet, E. 1981. Circular statistics in biology. Academic Press, New York, N.Y. 371 pp. Burt, W. H. 1943. Territoriality 24:346-352. and home range concepts as applied to mammals. Cochran, W. W., D. W. Warner, J. R. Tester, and V. B. Kuechle. 1965. Automatic system for monitoring animal movements. BioScience 15:98-100. Conover, W. J. 1971. Practical Nonparametric Dixon, K. R. and J. A. Chapman. 61: 1040-1044. Statistics. 1980. Harmonic J. Mammal. radio-tracking Wiley, New York, N.Y. 462 pp. mean measure of animal activity areas. Ecology Don, B. A. C. and K. Rennolls. 1983. A home range model incorporating J. Animal Ecology 52:69-81. biological attraction points. Dunn, J. E. 1978. Computer programs for the analysis of radio telemetry data in the study of home range. Statistics Laboratory Tech. Rep. No.7, Univ. of Arkansas, Fayetteville, AR. Dunn, J. E. 1978. Optimal sampling in radio telemetry studies of home range. Pages 53-70 in H. H. Shugart, Jr. ed. Time Series and Ecological Processes. SlAM, Philadelphia, PA. Dunn, J. E. and I. L. Brisbin, Jr. 1982. Characterizations of the multivariate Ornstein-Uhlenbeck diffusion process in the context of horne range analysis. Statistical Laboratory Technical Report No. 16, Univ. of Arkansas, Fayetteville, AR. 72 pp. Dunn, J. E. and P. S. Gipson. Biometrics 33:85-101. 1977. Analysis of radio telemetry data in studies of home range. Eddy, W. F. 1977. A new convex hull algorithm for planar sets. ACM Trans. Math. Sofrvvare 3:398403. Ford, R. G. and D. W. Krumme. 1979. The analysis of space use patterns. Hartigan, J. A. 1987. Estimation 82:267-270. of a convex density contour in two dimensions. Chapter VII - Home Range Estimation J. Theor. BioI. 76:125-155. J. Am. Stat. Assoc. Draft June 9, 1987 �211 Analysis of Biotelemetry Page 161 Data Hayne, D. W. 1949. Calculation of size of home range. J. Mammal. 30:1-18. Jennrich, R. 1. and F. B. Turner. 22:227-237. 1969. Measurement of non-circular home range. J. Theoretical BioI. Koeppl, J. W., N. A. Slade, and R. S. Hoffmann. 1975. A bivariate home range model with possible application to ethological data analysis. J. Mammal. 56:81-90. Madden. R. and L. F. Marcus. 1978. Use of the F distribution ranges. J. Mammal. 59:870-871. in calculating bivariate normal home Mielke, P. W., Jr. and K. J. Berry. 1982. An extended class of permutation pairs. Commun. Statistics 11:1197-1207. techniques Mohr, C. O. 1947. Table of equivalent populations Nat. 37:223-249. of North American O'Callaghan, J. F. 1974. Computing the perceptual and Image Processing 3:141-162. boundaries Rongstad, O. J. and J. R. Tester. 1969. Movements J. \Vildl. Manage. 33:366-379. and habitat use of white-tailed Rongstad, O. J. and J. R. Tester. Wildl. Manage. 35:338-346. for matched small mammals. of dot patterns. Am. Midl. Computer Graphics deer in Minnesota. 1971. Behavior ':C1dmaternal relations of young snowshoe hares. J. Samuel, M. D. and E. O. Garton. 1985. Home range: a weighted normal estimate and tests of underlying assumptions. J. Wildl. Manage. 49:513-519. Schoener, T. W. 1981. An empirically based estimate of home range. J. Theoretical Siniff, D. B. and J. R. Tester. 1965. Computer telemetry. BioScience 15:104-108. analysis of animal movement Smith, W. P. 1983. A bivariate normal test for elliptical home-range and recommendations. J. Wildl. Manage. 47:613-619. Swihart, R. K and N. A. Slade. 1985. Testing for independence Ecology 66:1176-1184. BioI. 20:281-325. data obtained by models: biological implications of observations in animal movements. Tierson, W. c., G. F. Mattfeld, R. W. Sage, and D. F. Behrend. 1985. Seasonal movements ranges of white-tailed deer in the Adirondacks. J. Wildl. Manage. 49:760-769. Draft June 9,1987 and home Chapter VII - Home Range Estimation �Analysis of Biotelemetry Page 16: VanWinkle, W. 1975. Comparison 39:118-123. of several probabilistic home-range Data models. J. \Vildl. Manage. Zimmerman, G. M., H. Goetz, and P. W. Mielke, Jr. 1985. Use of an improved statistical method for group comparisons to study effects of prairie fire. Ecology 66:606-611. Chapter VII - Home Range Estimation Draft June 9, 1987 �Wildlife Research Report July 1987 213 JOB PROGRESS REPORT State of Colorado Project No. 01-03-048 (FW 26 p) ------------~------~-- Mammals 2 Research Work Plan No. 2 Bighorn Sheep Investigations Job No. 4 Plane of Nutrition and Bighorn Sheep Population Performance Period Covered: July 1, 1986 - June 30, 1987 Authors: Personnel: M. W. Miller, N. T. Hobbs J. A. Armstrong, K. A. Trust ABSTRACT Administering adrenocorticotrophic hormone (ACTH) gel to bighorns elevated urine cortisol concentrations (VCCs) (p < 0.01) and urine cortisol:creatinine ratios (VCCRs) (p < 0.01), and reduced urine creatinine concentrations (VCr) (p < 0.01), for 24 hrs after subcutaneous injection. From 24-48 hrs post-ACTH injection, VCCs, VCrs, and VCCRs were lower (p < 0.01) in treated bighorns than in controls. Cortisol in urine is stable for up to 48 hrs at 4°C (p > 0.05) and 36 hrs at 220C (p < 0.05); marked reduction in cortisol levels occurred in urine stored at 220C beyond 48 hrs. Estimated mean VCCR obtained from urine-stained snow samples did not differ (p > 0.05) from mean VCCR measured in free urine samples from bighorns; ACTH-treated and control bighorns could be differentiated using VCCRs measured in urine-stained snow samples. Our data suggest VCCR is a sensitive tool for detecting adrenal responses in bighorns. ��PLANE OF NUTRITION A..~DBIGHORN SHEEP POPULATION PERFORMANCE M. W. t-liller N. T. Hobbs P. N. OBJECTIVE To treat bighorn sheep to control disease where necessary. SEGMENT OBJECTIVES The specific objectives of our work include: 1. Determine if adrenal responses to acute stress can be used as a reliable indicator of chronic stress. 2. Correlate the magnitude of adrenal responses to acute stress with measures of immune competence in bighorn sheep. 3. Experimentally compare the utility of blood, urine, and fecal cortisol concentrations as indicators of exposure to chronic stress in bighorn sheep. PREFACE We continued developing techniques for detecting adrenal responses in bighorns by sampling and measuring cortisol in urine and feces. These techniques were used to measure adrenal responses of captive bighorns to alternate-day adrenocorticotrophic hormone (ACTH) injections. In separate experiments, we determined effects of time and temperature on cortisol stability in urine and examined techniques for estimating urine cortisol levels from urine-stained snow. CHANGES IN URINARY CORTISOL EXCRETION IN BIGHORNS IN RESPONSE TO ALTERNATE-DAY ACTH TREATMENT INTRODUCTION We continued a series of experiments in measuring and managing chronic stress in bighorns by exam~n~ng changes in excreted cortisol in response to alternateday ACTH injections. Only urine cortisol data have been analyzed to date; fecal cortisol assays and analyses are pending improvement of laboratory techniques. Pertinent literature reviews and a complete study plan for the stress experiments in bighorns can be found in Part 2 of the Colorado Division of Wildlife Research Report, Work Plan 2, Job 4, July, 1985. �216 METHODS Using alternate-day ACTH administration to simulate chronic stress, we evaluated the utility of cortisol levels in excreta as indicators of adrenal response in bighorns. Twelve tame bighorns entered into this blocked, 2-way crossover experiment. We paired sheep using sex, age, and body size as criteria. One animal from each pair was randomly assigned to ACTH treatment; treatment and control assignments were reversed for the second half of the study. Each half of the experiment lasted 41 days, and a 60-day recovery period separated halves. Animals were housed in bare-ground isolation pens (about 50 m2), and we provided alfalfa hay, a pelleted concentrate, and water ad libitum throughout the experiment. During the first pretreatment period (days 1-12), clinical signs of pneumonia developed in 4 bighorns representing 2 experimental pairs; those animals were removed from the study. Two of these recovered and subsequently entered the experiment after the recovery period. However, only data for 4 pairs-of bighorns that completed the crossover are reported here. Both halves of this experiment were conducted using the same schedule: We collected urine and feces on days 1, 6, 11, 16, 22, 28, 34, and 40. Each collection ~eriod lasted 48 hrs, and sheep were housed in metabolic cages (about 10 m ) for the entire period. Urine and fecal collections were made at l2-hr intervals, and we stored all samples at -200 C until processing. On day 13, we began administering ACTH gel (0.5 U/kg) subcutaneously to 1 randomly selected sheep from each pair. Alternate-day ACTH administration continued for the remainder of the experiment. The other bighorns served as controls, receiving equivalent volumes of 0.9% saline solution on the same schedule as treatment sheep (Fig. 1). Urine and feces from each collection were weighed, and we used 10% aliquots for composites representing 24-hr subperiods of each 48-hr collection period. The 24-hr subperiods preceeding ACTH treatment were termed "chronic", and 24-hr subperiods following ACTH administration were termed "acute." Urine cortisol concentrations (UCC) were measured by radioimmunoassay (RIA). Urine creatinine (UCr) concentrations were measured in samples diluted 1:20 by a colorimeteric method. We expressed UCC inpg/dl, UCr in mg/dl, and calculated urine cortisol:creatinine ratios (UCCRs) expressed as pg/mg. Fecal samples were lyophilized and are awaiting analysis. We analyzed data using analysis of variance for repreated measures with mean pretreatment values as covariates. Because mean pretreatment UCCRs did not differ between halves of the crossover, we used each bighorn as its own control in our analyses. Data from acute (0-24 hr post-ACTH) and chronic (24-48 hr post-ACTH) subperiods were analyzed separately. RESULTS AND DISCUSSION Alternate-day ACTH administration produced marked changes in urinary cortisol excretion in treated bighorns. Acute responses were characterized by increased mean UCCs (P < 0.01) and reduced UCrs (p < 0.01) in treated bighorns; both mean UCCs and UCrs were lower (p < 0.01 and P < 0.01) during �217 chronic response phases than those for controls (Tables 1 and 2). Combined changes in UCC and VCr account for more than a 4-fold rise (P < 0.01) in acute UCCRs of treated bighorns and for reduction (P < 0.01) of UCeRs on days when ACTH was not administrered (Table 3). Elevated UeeRs in ACTH-treated bighorns can be attributed to increasing cortisol production and secretion, combined with reduction in urine creatinine levels by day 22. Rates of creatinine metabolism and excretion are relatively constant and are correlated with muscle mass of an animal. We assume that neither ACTH nor elevated cortisol levels alter these rates. Therefore, reduction of UCr relates to urine dilution by diuresis, a mineralocorticoid effect of cortisol. It follows that UeCR more accurately estimates urinary cortisol excretion than simple measures of urine cortisol levels. Because the ratio corrects for dilution and measures changes in cortisol relative to a constant excretion of creatinine, UCCR should be a relatively sensitive indicator of adrenal responses in bighorns subjected to chronic stress. Moreover, using UCCR may allow remote sampling of urine in the field (see below). Reduced UCCs and UCCrs in treated bighorns 24-48 hrs after ACTH administration suggests that negative feedback mechanisms regulating the hypothalamopituitoadrenal axis remained intact throughout the experiment. Exogenous ACTH appeared to increase synthesis and secretion of cortisol for less than 24 hrs; elevated serum cortisol then reduced endogenous ACTH production resulting in lower cortisol levels between treatments. Despite reduced cortisol excretion on "chronic" days, some of our data suggest that physiological effects of elevated serum cortisol levels may have carried from treatment to treatment. By day 34, chronic UCrs in treated bighorns were lower (P < 0.05) than for controls (Fig. 3b); the trend for reduction of chronic UCr appeared by day 22. These carryover effects contributed to negative feedback that reduced chronic uce in treated bighorns to levels below those observed during the pretreatment period (fig. 3a). Observed changes in lymphocyte responses (unpublished data) are also thought to be related to chronic, cumulative effects of ACTH administered. Thus, our experimental model appears to have simulated at least some of the physiological phenomena believed to occur in wild bighorns subjected to chronic stress; adrenal responses associated with these are detectable using techniques we described. Mechanisms that allow manifestation of deleterious effects of chronic stress in bighorns remain largely undescribed, but availability of tools to monitor physiological responses associated with chronic stress should enhance future investigations. STABILITY OF CORTISOL IN URINE METHODS Urine collected from a single bighorn over 3 hrs was divided into 80, 2-ml aliquots, and these aliquots were randomly assigned to 1 of 16 temperature x time treatments Cn = 5 aliquots per treatment): �213 Temperature (OC) x Time (hr) o 4 12 24 36 48 72 120 168 22 After imposing treatments, all aliquots were stored at -200 C until assayed for cortisol concentrations; time = 0 samples for both temperatures were frozen immediately and served as controls. Urine cortisol concentrations were measured as previously described. We used Duncan's multiple range analysis to group mean urine cortisol concentrations under the 2 temperature treatments .. RESULTS AND DISCUSSIOl~ Cortisol concentrations were reduced by higher storage temperatures (P < 0.001) and by longer storage times (p < 0.001), and these appeared interactive (time x temperature P < 0.001). At refrigerator temperatures (40 C), cortisol concentrations in aliquots stored for up to 48 hrs did not differ (p < 0.05) from those of controls (Table 4A). Room temperature (220 C) aliquots stored for up to 36 hrs did not differ (P < 0.05) in cortisol content from controls (Table 4B); storage beyond 72 hrs at 220 caused marked reduction in urine cortisol concentration. These data suggest that cortisol in urine is relatively stable, particularly at cooler temperatures, although the half-life of cortisol in urine is somewhat shorter than for cortisol in serum stored under similar conditions. ESTIMATION OF CORTISOL:CREATININE RATIOS BY SAMPLING URINE-STAINED SNOW METHODS Urine samples from 4 tame bighorns (2 control animals and 2 subjected to alternate-day ACTH treatment) were used to investigate utility of urinestained snow for estimating urine cortisol:creatinine ratios. Each urine sample was mixed thoroughly and divided into 2, SO-ml subsamples. Aliquots (3 x 4 ml) from 1 subsample, poured directly into storage tubes, served as controls. We briefly warmed (-300 C) the second SO-ml subsample in a water bath, then poured the urine into a fresh snow drift. A core sample of urinestained snow was collected, melted, and mixed thoroughly; 3, 4-m1 aliquots from this melted snow served as treatment samples, and cortisol and creatinine concentrations were measured for each aliquot group. We compared UCCR estimates from snow samples and controls using a paired t-test. �219 RESULTS AND DISCUSSION Individual estimates of UCCR from snow samples differed substantially from actual UeCRs (Table 5). Despite these differences, the overall estimated mean UCeR for snow samples did not differ (p > 0.05) from the mean UCCR measured directly from urine. Low UeCRs appear to be estimated more poorly from snow than higher UeCRs (Table 5), probably a result of high variation associated with estimates of extremely low cortisol concentrations by RIA. However, we were able to use UCCR estimates from snow to correctly discern between ACTH treated and control bighorns. This ability, combined with the relatively accurate estimation of a simulated pop~lation mean, suggests promise for urine-stained snow as a remote sampling approach to estimating urinary cortisol excretion and adrenal responses in bighorns and other free-ranging species. ./ Prepared by . ./::;~. ../> l1ichael v. "11iller;DVH N. Thompson Hobbs Wildlife Researcher �220 Table 1. Urine cortisol concentrations (pg/dl) from treatment and control bighorns before and after alternate-day ACTH treatment. A) Acute response (0-24 hr post-ACTH). B) Chronic response (24-48 hr post-ACTH). All values are reported as mean (standard error). Urine cortisol concentration (pg/dl) A) Treatment* 2.35 (0.14) 2.75 (0.04) 3.22 (1.15) 7.94 (1.57) 1.92 (0.22) 2.07 (0.29) 2.61 Acute Response Control Treatment B) Pretreatment Chronic Response Control Treatment (9.77) 0.92 (0.26) Table 2. Urine creatinine concentrations (mg/dl) from treatment and control bighorns before and after alternate-day ACTH treatment. A) Acute response (0-24 hr post-ACTH). B) Chronic response (24-48 hr post-ACTH). All values are reported as mean (standard error). Urine creatinine concentration (mg/dl) A) Treatment* 210.0 09.0) 237.0 (29.2) 242.6 (39.1) 168.8 (21.4) 190.6 07.7) 195.7 01. 9) 237.2 (52.3) 178.0 (28.6) Acute Response Control Treatment B) Pretreatment Chronic Response Control Treatment Table 3. Urine cortisol:creatinine ratios (ug/mg) from treatment and cortisol bighorns before and after alternate-day ACTH treatment. A) Acute response (0-24 hr post-ACTH). B) Chronic response (24-48 hr post-ACTH). All values are reported as mean (standard error). Urine cortisol:creatinine ratio (ug/ml) Pretreatment A) Acute Response Control Treatment B) Treatment* o.cn (0.001) 0.011 (0.002) 0.014 (0.004) 0.052 (0.007) 0.011 (0.001) 0.010 (0.002) O.Oll (0.001) 0.005 (0.001) Chronic Response Control Treatment �221 Table 4. Duncan's multiple range analysis of mean urine cortisol concentrations (ug/dl) from samples stored at 40 (A) and 220 C (B) over time (0-168 hrs). Means with different letters differ (p < 0.05). A) B) TU1E Mean 36 3.524 0 3.476 12 3.316 24 3.304 48 2.998 72 2.968 168 2.926 120 2.868 TIME Duncan Grouping B B B B B B A A A A A A A B Mean C C C C C C C C C C C Duncan Grouping 24 3.578 0 3.382 B 12 3.246 B 36 3.074 B B 48 2.682 D D 72 2.340 D 120 0.650 E 168 0.154 F B A A A A A C C C Table 5. Estimates of urine cortisol: creatinine ratios from urine and urinestained snow samples. Values are represented as mean (standard error). Urine cortisol:creatinine ratio (ug:mg) Animal Treatment CTRLa ACTHb ACTH CTRL A B C D aCTRL bACTH saline control = ACTH-treated Control 0.009 0.039 0.053 0.024 (0.001) (0.003) (0.004) (0.003) Snow 0.003 0.050 0.047 0.007 (0.001) (0.005) (0.002) (0.002) �222 I DAY; 1 2 34 COLLECTION; CCX S 6 7 8 9 10 11 12113 14 IS 16 17 18 19 20 21 22 23 24 2S 26 27 28 29 30 31 32 3334 X X CCX I I I X X C CI X X X CCX X A X X X X CCX A X A X X X X CCX A X A X 3S 36 37 38 39 40 41 X X X CCX A X A X X X X C C A X A X A I TREAr.-<.L';"!'; X X X X X X X X X X X I I X, A A X A X A X A X r------------~------------------------------------~ Pretreatment Period Treatment Period Fdgu r e 1. Collection and treatment schedule for alternate-day ACTH experiment. PLOT OF CORTISOL 0.08 CREATININE (fg:mg) a c 0.07 + Control ;\ 1 , 0.05 ACTH a I· 1\ " 0.06 VS PERIOD + , ' \ I + :\' a + ! : i .J 0.04 !/a + a I I 0.03 I I + c ,, . 0.02 + 0.01 + \1. / 1/ 1 r----i »>1'/ 'T----b/ I ,, ~I /~ I: I cl---I i I , ,, 0.00 + --+---------+---------+---------+---------+---------+---------+---------+2 I I Ac:..'''' " Collection 5 6 7 8 Period Figure 2. Average urine cortisol:creatinine ratios (ug/mg) for ACTH-treated and control bighorns before and after treatment. Vertical lines = 2 standard errors. �Urine Cortisol:Creatinine Ratio Acute ~U~~~I~~~:~mo~) Fig..-e 3a. Acute (0- 24h) effects of alternate- day ACTH administrationon urine parameters of bighorn sheep. ~ 0.07 0.06 o.os 0.04 0.03 0.02 0.01 O~-------L--------~-------L--------~------~ o so 50 10 40 20 Day - ACTH -+-CTRl Urine Cortisol Concentration Urine Creatinine Ooncentration Acute Acute U_~OC~I~~/~dl~I ~ UO- (mg/dI) 350 10~ soc 8 ,t e 2liO 200 1150 4 100 2 50 O~-------L--------~------~--------~------~ so o 10 40 20 150 0 0 20 10 - ACTH -+- CTRl 30 40 liO Day Day - ACTH -+- CTRl N N W �. Urine Cortisol:Creatinine Ratio Chronic Fig"e 3b. Chronic (24-48h) effects of alternate- day ACTH administration on urine parameters of bighorn sheep. u v~CCA~~(U~g~:m~O~I 0.01. 1 ~ r- 0.012 00111<::: 1--7 0.009 o.ooe 0.004 0.002 o~------~------~-o 10 L- 30 20 -L -J 50 40 Day - ,acTH -- CTRl Urine Cortisol Concentration Urine Creatinine Concentration Chronic Chronic 5rU_CC~(U~g~/~~) ~ U_~=-(~m~O~/d~I)~ ~ ~Or •• GOO BegIn ICTH &eoln ACTH 260 3 200 HIO 100 60 o~------~------~------~------~------_J •• 0 o 10 20 30 50 O~------~------~~------~--------~------~ o 20 50 10 30 Day - ACTH -- CTAl Day - ACTH -- CTRl 40 N N -f:> �~u~urdUO UlVlSlOn OL WllGllle Wildlife Research Report July 1987 JOB PROGRESS REPORT State of Colorado Project No. 01-03-048 (FW 26 p) --~----------~------~- Mammals 2 Research Work Plan No. 3 Pronghorn Investigations Job No. 1 Pronghorn Population Dynamics Study Period Covered: July 1, 1986 - June 30, 1987 Author: T. M. Po jar ABSTRACT The experimental random strip and random quadrat pronghorn census was completed during 18-22 August 1986 in the Great Divide area. During the strip transect census, perpendicular distance from the flight-line was estimated for all groups of pronghorns detected. This data set was used to estimate the distance sightability curve and to obtain a line transect estimate of density. The density and herd structure estimates (+ 90% confidence limits) for the Great Divide Data Analysis Unit are: Strip transect, 7,719 + 15%; quadrat, 16,798 + 23%; and line transect, 25,960 + 14%. The population size estimate from the random transect census was 77.6% higher than the same census in 1985, and the random quadrat estimate was 79.6% higher in 1986. These are both far above increases that could be realized by natural reproduction and suggest emigration as the most likely explanation. The buck:lOO does ratio for the strip census is 43:100 (+ 19%) and the fawn:lOO does ratio is 82:100 (+ 10%). The random transects in Block 3 (the eastern one-third of the total area, 452 mi2) were reflown following strict line transect methodology. Three separate estimates of density were then available for Block 3: Random quadrat, 5,324 + 26%, random strip, 3,028 + 34%, and line ransect, 8,301 + 22%. Comparison of random'strip and line transect censuses was conducted in the Hugo Data Analysis Unit, which is typical shortgrass prairie. Using an observability decay curve generated from perpendicular distance estimates of groups from the flight-line, it is estimated that about'36% of the animals are missed during a conventional strip transect survey. ' ��227 PRONGHORN POPULATION DYNAMICS STUDY Thomas M. Pojar P. N. OBJECTIVES 1. Test the efficiency and precision of a quadrat and strip census design on rolling sagebrush steppe and shortgrass prairie. 2. Test the efficiency and precision of herd structure estimates when done concurrently with a quadrat and strip transect census on both rolling and sagebrush steppe and shortgrass prairie. 3. Test the reliability of doing strip transect and quadrat censusing during late summer when it is possible to do herd structure counts. 4. Measure and model the effects of reducing a pronghorn population density by 50% or greater. SEGMENT OBJECTIVES 1. Test the efficiency and preC1Slon of quadrat system sampling to estimate density and herd structure of pronghorns. 2. Simulate population changes to evaluate pronghorn population estimates derived from the censuses. 3. Evaluate the application of the line transect method for censusing pronghorn by estimating the parameters influencing the probability density function f(x). ACKNOWLEDGMENTS The following Colorado Division of Wildlife personnel contributed to the current phase of this study: M. Bauman and V. Graham served as navigator and observer on the strip transect and quadrat surveys, respectively. D. Bowden of the Statistics Department (Colorado State University) provided statistical consultation, and D. Anderson, Wildlife Coop Unit (Colorado State Unviversity) offered helpful discussions on the line transect methodology. T. Drummer (Michigan Technological University, Dept. of Mathematical Science) provided the analysis of factors that may influence detectability. METHODS AND MATERIALS The study area and methods used for the strip transect and quadrat censuses are described in Pojar (1983). Modifications in the strip survey to permit line transect analysis can be found in Pojar (1986). The only modification in this procedure for this segment was to estimate perpendicular distances of observed groups in 50-m increments rather than 100-m increments. As in the past, the program, TRANS?CT (Laake et al. 1979) was used to analyze the line transect data. I~ ~ll ~3ses the fourier series estimator was used. �2~d In the Hugo/Limon area (Hugo DAU, Units A36, A37, A38, A38l) the 16 random transects from previous experimental strip transect censuses were used. Half of these were randomly assigned as line transects and the other half as strip transects. The difference between the line and strip transect censusing is as follows: On strip transects, an attempt was made to count and classify all pronghorns observed. To do this, deviations were made from the flightline to get an accurate count and to classify each group detected. Groups that were detected after departure from the flight line were noted as"post-line" groups. Distance of the groups from the flightline was estimated as a possible correction factor for the strip data. During the line transect census, a strict flightline was maintained, and group sizes and perpendicular distance were estimated without deviation from the flightline. Two experienced observers (Elkins and Pojar) estimated perpendicular distances and classified independently. Short practice sessions were conducted each morning before the censusing effort began in estimating distance by hovering 400 m from a fence marked at 50-m increments with plastic flagging. Chi-square tests were used to test the independence of distance from the flightline and group size, activity, and proportion of bucks in the group. The probability of 0.05 was used as the significance level. RESULTS Great Divide Results of the experimental random strip and random quadrat pronghorn census in the Great Divide area are presented in Table 1. With the exception of the first year of this study (1982), the difference between the quadrat and strip censuses has remained very consistent (Fig. 1). The data from 1983-86 were collected using the same primary observer, navigator/observer, and the same type of helicopter (Bell-Soloy). During these 4 years, we used the same helicopter contractor and had only 2 different pilots, both of which have an excellent understanding of the census techniques and what is expected of them as pilots. Therefore, the correlation in Figure 1 includes only the data from 1983-86 in the calculated coefficient (R2 = 0.99). The estimate of 7,719 pronghorns based on the random transect method for 1986 was 77.6% higher than the 1985 estimate of 4,344, and the quadrat estimate was 79.6% higher than in 1985 (16,798 vs. 9,351). Both of these estimates represent increases that are far above what could be realized by natural reproduction in 1 year. The most likely explanation is emigration of unknown origin and, on a purely speculative basis, it may be from a backwash of movements that took place during the severe winter of 1983-84 when the population estimate declined by 50%. The distance of groups from the flight line was estimated during the strip transect sampling. This permits analysis of this data as line transect data. Program TRANSECT (Laake et al. 1979), using the fourier series estimator, was used. This approach gives a population estimate of 25,960 + 14.3%. However, tests of independence revealed that group size influences detectability (Chisquare = 26.34, P < 0.01, 6 d.f.) with larger groups being more detectable at greater distances from the flightline (Table 2). Quin (1979) proposes poststratification by group size as a means to alleviate the group size influence and obtain an unbiased estimate of the mean group size. Correction for the group size bias yields a population size estimate of 19,693. The proportion �of bucks in a group did not significantly (P > r.05) influence detectability, but group activity was a factor in detectabilit: ~able 2). The buck-to-doe ratio from quadrat data is alwa) higher than from the transect data, although not statistically significant because of the large variance. The quadrat estimate of buck-to-doe ratio is 52.8, which is 23.4% higher than the estimate of 42.8 from the transect data CTable 1). Over the 5 years' data, quadrat estimates have averaged 29% higher than transect data, which indicates that single bucks and small buck groups may be missed during transect sampling. Fawn-to-doe ratios have been fairly consistent between the 2 types of sampling. They are characterized by relatively tight confidence intervals because of sufficient sample size and low variance in both cases (Table 1). Block 3 of Great Divide A limited test of the line transect method was conducted in Block 3 of the Great Divide DAU. Block 3 is roughly the eastern one-third of the DAD and 452 mi2 in size. The random transects used in the strip transect census were reflown following strict line trans,ct methodology; that is, there were no deviations from the flightline. A comparison of 3 separate estimates of density (+ 90% confidence interval) were then available for Block 3. These estimates-are: Quadrat, 5,324 (+ 26.3%), strip, 3,028 (+ 34.5%), and line, 8,301 (~ 22.2%). Hugo/Limon Conventional strip transect sampling (mile-wide strips) yields the density and herd structure estimates presented in Table 3. The line transect data analysis in Table 4 points to the extreme variation in population size estimates than can be obtained using different cut-points (distance groupings) based on the distribution of group sightings in Figure 2. Also, in Table 4, the strip data was subjected to line transect analysis to correct for detectability bias as discussed in Anderson and Pospahala (1970). Using the 50-em cut-point analysis as the "corrected" density estimate and comparing it to the conventional strip estimate in Table 3, the correction factor for Elkins' strip count is 1.54 and for Pojar it is 1.56. Or put in different terms, Elkins "missed" 35.3% of the animals during the conventional census, and Pojar "missed" 35.8%. LITERATURE CITED Anderson, D. R., and R. S. Posaphala. 1970. Correction of bias in belt transect studies of immobile objects. J. Wildl. Manage. 34(1):141-146. Laake, J. L., K. P. Burnham, and D. R. Anderson. 1979. User's manual for program TRANSECT. Utah State Univ. Press, Logan. 26pp. POjar, T. M. 1983. Pronghorn investigations--pronghorn population dynamics study. Colo. Div. Wildl. Res. Rep. July, Part 4:379-385. �no Pojar, T. M. 1986. Pronghorn investigations--pronghorn population dynamics study. Colo. Div. Wild1. Res. Rep. July, Part 2:195-203. Quin, T. J. II. 1979. The effects of school structure on line transect estimators of abundance. Pages 473-491 in Contemporary Quantitative Ecology and Related Ecometrics, G. P. Pat11 and M. L. Rozenweig (eds~). Internat1. Co-op Pub1. House, Fairland, MD. 473-49lpp. -: -, Prepared by --I _-- - j,~ '(j e>: . //.) CofhomasH. Pojar :. wildlife Researcher a.7 �Table 1. Results of the random strip transect and random quadrat census for the Great Divide Pronghorn Data Analysis Unit (A3 and A301). 1983 1982 ~J.£ Quad 16,650 +17.3'£ 1985 StrlJ 4,689 +13.6% 4,052 Quad 9,345 +18.2% 7,644 7.?2 23,993 16.39 5,327 3.8? 11,046 7.60 4,347 +31.3'£ 2,985 13,770 7,373 Upper Mean Dens 1ty (Animals/sQ. mi.) 5,709 3.54 19,530 10,363 Bucks:100 Does 90'£C.l. Lower Upper Fawns:l00 Does 90'£C.l. Lower Upper No. Clas1fied Bucks Does 1984 Qu~rt 18,438 --+30.1% 12,883 Strit_ 8,868 +16.9'£ Pop. Est. 90'£C.l. Lower 13.55 _ 45.5 +18.6'£ 52.2 +19.3'£ 43.3 +14.4% 54.3 +17.7% 34.8 +lA.O% 37.1 54.0 66.8 +9.4'£ 42.2 6?3 83.2 +9.4'£ 46.6 66.0 70.6 60.5 73.1 75.4 91.1 36.7 49.1 65.6 +6.7% 61.7 70.0 78.5 41.0 47.7 +11.51· 47.2 53.2 +11.n, 61.4 76.8 336 738 324 620 609 1,404 325 598 Fawns 493 1,567 516 1 ,460 921 TOTAL 2,934 422 1,345 ~ ~. J _' +24.11 42.4 69.4 54.4 +20.4% 43.2 65.4 1986 St:.!::_!r Quad 4,344 9,351 +21.0% -+19.9% 3,431 7,490 Strip 7,719 +15.4% 6,527 Quad 16,79A +22.6% 13,004 5,258 3.53 11,711 7.61 8,912 6.28 20,597 13.67 40.2 +19.2% 51.8 +70.3% 42.8 +19.3% 47.9 32.5 A8.1 +7.8% 67.3 41. 3 98.5 +10.5% 108.8 88.2 95.0 81. 2 34.5 51.0 81.5 +9.7'f. 73.6 A9.4 323 929 443 218 390 277 689 173 334 526 1,230 212 1,695 820 607 1,573 329 836 ]_!_Q02 2,758 52.R +77.]'(. 38.2 67.5 7r..7 +13.4% 68.2 89.3 318 602 474 1,394 N W I-' ~ �232 Table 2. factors. Tests of independence between distance from flightline and other GrouE size Distance 1-3 4-5 6-10 11+ Total Distance and GrouE Size 100m 100-200m 200m+ 134 37 50 58 20 38 28 22 29 10 12 26 238 91 143 Total 221 116 79 56 472 Chi-Square 26.34, P 0.00, a.r , 6 Distance and Activity Activit Distance 100m 100-20Om 2000+ Total Bedded Missin~ 63 13 57 3 3 133 Chi-Square Standin~ Runnin~ Total 3 86 38 78 86 37 46 238 91 184 9 202 169 513 15.27, P 0.02, d.£. = 6 Distance and Proportion Bucks ProEortion bucks Distance < = 10% Bucks > 10% Bucks Total 100m 100-200m 200m+ 138 50 87 100 41 97 238 91 184 Total 275 238 513 Chi-Square = 4.86, P = 0.08, d.f. = 2 �Table 3. Density and herd structure estimates based on a sample of 8 random 1,600 m- (1 mi-) wide strip transects. Hugo pronghorn Data Analysis Unit (A36, A37, A38, A38l; 1,688 mi2), 1986. Observers: Mark Elkins and Tom Pojar. Herd Structure based on a sample of 436, Elkins; 449, Poj~;. Observer Pop. size estimate Elkins Pojar 2,384 2,644 90% C.1. +35.2% +31. 0% 90% C.1. B:10OD ratio F:100D ratio 90% 34.9 32.1 +30.2% +27 •6;~ +39.5% +37.3% 27.1 28.2 c.r . Table 4. Line transect pronghorn density estimates for the Hugo Data Analysis Unit (A36, A37, A38, A381). Estimates based on 8 randomly selected lines; observers: Mark Elkins and Tom Pojar. 50m 95% C.1. 100m 95% C.1. 1,3,5, 800 H 95% C.1. 3,173 5,407 +34.5% +35.8% 3,186 3,307 +34.4% +36.6% 3,321 2,596 +34.3% +34.0% 3,683 4,120 +28. 4~~ +26.5% 3,717 4,147 +28. 3~~ +26.4% 5,202 5,496 +31. 2% +30. 5~~ Line transects] Elkins Pojar Strip transects= Elkins Pojar lStrict flight-lines were followed with no deviations to classify observed pronghorns. =The perpendicular distance estimates made during the strip transect census were used as line transect data. Deviations were made from the flightline to classify all pronghorns observed. Table 5. Paired t-tests (d.f. = 15) for number of groups and total animals seen by each observer on the right vs. left side of the flight line and comparison of observers by number of groups and total animals seen. The seating in the Bell-Soloy Helicopter was from left to right: p:lot, Elkins, Po'ar. Difference ComEarison Mean SD T-value Probability -0.467 -1.800 -0.200 -4.933 4.373 26.889 3.802 20.631 0.413 0.259 0.204 0.926 0.686 0.799 0.842 0.370 -0.533 -0.267 1.667 -1.467 0.640 0.594 6.032 4.565 3.228 1.740 1.070 1.244 0.006 0.104 0.303 0.234 Right vs. Left Elkins, no. groups Elkins, total animals Pojar, no. groups Pojar, total animals Elkins vs. Pojar No. groups on right No. groups on left Total animals, right Total animals, left �234 20 - 18 '82 0 0 0 'r' - • 16 x UJ l- 14 ~ I- 12 -c (J) UJ l- 10 e:( c: 0 e:( '84 8 y- :J a -269.99 R2= 6 + 2.15x .991 O~ 0 5 6 7 8 TRANSECT ESTIMATE (x 1000) Figure 1. Relation of strip vs. quadrat population size estimates the Great Divide pronghorn DAU. The 1982 point is not included in the regression, see text. for �'t:J Q) :> c, Q) I/) .a a I/) ~ 0 11 ~ 10 c, -.•.. .,._. ----_._- 20 19 18 17 16 15 14 13 12 - . __ .- . _ .. -'_... -- _-_ .... ---- .- .. _. ,_ .. _------._--- ,,_._----- 9 ~rj ;;o- .,_ 8 " j ~ 0 7 c: 6 q:j 0 I- .•.. Q) 0 , l Q) Q. R 100 150 200 250 v, 300 350 400 ~I ::,j v,1 450 VI v r 500 v ;i C Vlu V1 v v 550 ,l v' v v v ~ I fl v ~ Vj ~, v v / ~ H ' '~' , 2. . , I 1/ Figlln:~. j ,', rl ~/l ,) 50 v', "~ ~ r1 >i t' 0 V ,i , I 1 VIr , J ~ :.1 ~ ;'j 2 - , j I I... v ,I ~/j i~j 3 ~. ::j n ~ 4 :.' ..i ~ /j 5 c. r-' 600 ~~ 650 I I 700 750 I BOO DIstance (meters) from rlf9~t Une ElkIns I r ,'I rder l", '-1 ! Pe r pend i cu La r d Ls t.anc o o s t Lm.tt c s b v o b s o r vc r s Elkins 1 i nu t r an S c (' tel' n s u s , I 9 H fJ • .ind I'Pj;lr for the Hugo pronghorn Di\U N W U'l ��Colorado Division ot Wildlite Wildlife Research Report July 1987 237 JOB FINAL REPORT State of Colorado Project No. 01-03-048 (FW 26 p) --------------~------~ Mammals 2 Research Work Plan No. 4 Rocky Mountain Job No. 1 Seasonal Habitat Selection and Activity of Sympatric Mountain Goat and Bighorn Sheep Populations Period Covered: July 1, 1986 - June 30, 1987 Author: Goat Investigations D. F. Reed ABSTRACT Two manuscripts titled "Alpine habitat selection in sympatric mountain goats and mountain sheep" and "Activity patterns of sympatric mountain goats and mountain sheep" were submitted to refereed journals. The first manuscript was not accepted, primarily because of confounding factors. The second manuscript will be revised and resubmitted to the Journal of Wildlife Management during the next segment. Findings relating to these manuscripts were: 1) mountain goats used alpine habitats disproportionately to their availability, 2) mountain sheep used alpine habitats proportionately to their availability, 3) mountain goats and mountain sheep did not use the same alpine habitats in toto, and 4) there was no difference in the activity patterns of the 2 species, however, there was variation between individuals and seasons. Data on marked animals and reproduction are summarized. Data on mortality are provided. ��239 SEASONAL HABITAT S::::_ECTION AND ACTIVITY OF MOUNTAIN GOAT AND MO~NTAIN SHEEP POPULATIONS Dale F. Reed P. N. OBJECTIVE Determine the degree of interspecific competition between mountain goats and bighorn sheep through research. SEGMENT OBJECTIVES 1. Test the hypothesis that mountain goats use alpine habitats disproportionately to their availability. 2. Test the hypothesis that mountain sheep use alpine habitats disproportionately to their availability. 3. Test the hypothesis that mountain goats and sheep use alpine habitats that are spatially and/or temporally discrete - their distributions are independent. 4. Test the hypothesis that mountain goats and sheep exhibit the same activity patterns in alpine habitats. ACK.~O\o.'LEDG?-l ENTS See Reed (1985). DESCRIPTION OF AREA The study area has been described by Reed (1982). METHODS AND MATERIALS Methods and materials have bee~ 5escribed by Reed (1981, 1982). RE·.;:LTSAND DISCUSSION Capture and Marking Seventy-one mountain goats were marked with radio telemetry collars, neck bands, or eartags (Table 1). Of this 71, 10 were harvested and 6 died during the study. Ten mountain sheep were marked with radio telemetry collars or neck bands. Three other sheep were individually identifiable from a previous marking program in 1977 (collars Yellow 19, Plain Yellow; eartag 4), and 1 was a readily distinguished, partial albino. Some radio collars were still active at the end of the segment (Tat~e 2). �240 Habitat Use The manuscript titled "Alpine habitat selection in sympatric mountain goats and mountain sheep" covers the results and discussion of habitat use. Activity Telemetry tip-switch activity collars were maintained on 9 mountain goats and 9 mountain sheep for varying periods during the study (Table 2). Generally, mountain goats were accessible for obtaining activity data. However, mountain sheep were more transitory and often dispersed into areas located near the boundary of the study area. Hence, more activity data were collected on mountain goats than on mountain sheep during the study (Table 3). Some of the results have been described previously (Reed 1985). Additional results and discussion will be available in the manuscript titled "Activity patterns of sympatric mountain goats and mountain sheep." Reproduction Of adult female mountain goats collared, reproductive status was estimated on 5, 11, 17, 19, 16, 17, 14, and 9 in 1980, 1981, 1982, 1983, 1984, 1985, 1986, and 1987, respectively (Table 4). The mean age of mountain goats (n = 12) having their first kid or kids was 3.2 + 0.9 (SD) years. The mean age of mountain goats (n = 8) having twins was-7.5 + 2.5 (SD) years. Mountain goat 27 (Red tele II)-had 2 sets of twins, 1 at age 9 and the other at age 11. She alternated twins and singletons during the 5-year period (Table 4). Eleven percent of the adult (> 4 years old) females had twins, 65% had 1 kid, and 24% had no kid or kids (Table 5). This percent of twinning is relatively high compared to some reports (Brandborg 1955:94) and low for others (Hayden 1984). The rate of twinning, as the rate of having singletons, is likely a minimum since some kids may die soon after birth. An example of this was reported previously (Reed 1983). No reproductive data obtained so far on mountain goats (Table 4) support Heimer's hypothesis of alternate-year production (Heimer and Watson 1982). Furthermore, applicability of this hypothesis to mountain goats in the Mt. Evans study area may be in doubt because dominant male behavior in mountain goats differs largely from that in sheep. Also, it would have to be assumed that success in harvesting dominant male mountain goats is comparable to success in harvesting dominant male mountain sheep. This mayor may not be the case. Of the marked or identifiable female mountain sheep, the reproductive status was estimated on 11 (Table 6). Data from mountain sheep 1 (Table 6) may be in support of the alternate-year production hypothesis. Mortality A number of marked animals were recovered during the study (Table 7). A mlnlmum of 11 deaths of collared mountain goats occurred. Based on this small sample, the mean age at death from natural causes was 7.6 yrs (n = 9). �241 Species Interaction See Reed (1985). Conc€Dtual Models of Competition See Reed (1985). LITE~~TURE CITED Hayden, J. A. 1984. Introduced mountain goats in the Snake River range, Idaho: characteristics of vigorous population growth. Pages 94-119 in M. Hoefs, ed. Proe. Bienn. Symp. North. Wild Sheep and Goat Counc. 4, Whitehorse, Yukon, Can. 5l3pp. Heimer, h. E., and S. Watson. 1982. Differing reproductive patterns in Dall sheep: population strategy or management artifact? Pages 330-336 in J. A. Bailey and G. G. Schoonveld, eds. Proc. Bienn. Syrnp.North Wild Sheep and Goat Counc. 3, Fort Collins, CO. 405pp. Reed, D. F. 1981. Rocky Mountain goat ecology study. Game Res. Rep. July, Part 2:209-222. _____ 1982. Rocky Mountain goat-bighorn shee Div. of wildl. Game Res. Rep. July:79-l28. Colo. Div. of Wildl. ompetition study. Colo. 1983. Seas~: habitat selection and ae. ~vity of sympatric mountain goat and bigho~. ;.eeppopulations. Colo. Div. of Wildl. Game. Res. Rep. July Part 2:403-4~2. 1985. Seasonal habitat selection and activity of sympatric mountain goat and bighorn sheep populations. Colo. Div. of wildl. Game. Res. Rep. July Part 2:193-224. C Prepared by ----- \-------~ '" ~ai~~_. Dale F. Reed wildlife Researcher \ �N +:> N Table 1. Date. age. sex. collar or tag. and selected measurements of mountain goats trapped in the Mt. Evans area. liorn lenglh (cm) No. 2 3 4 5 6 7 a 9 10 11 12 13 14 15 16 17 10 19 20 21 22 23 24 25 26 27 20 Date Age Sex 22 Aug 00 22 Aug 00 Y F K F 22 10 22 29 29 30 Aug Sep 5ep Sep 5ep Sep 00 00 00 00 00 00 A F 30 30 7 7 7 9 5ep 5ep Oct Oct Oct Jun 00 00 00 00 00 01 10 Jun 01 10 Jun 01 16' Jun 01 17 Jun 01 17JunOl 29 Jun 01 29 Jun 01 20 Aug 01 30 Aug 01 4 5ep 01 , 7 5ep 01 155epOl 6 Oct 01 14 Oct 01 A F A F K F A(6) 1 F 2 yr F K F A{O) F K M 2 yr F A{l) F 2 yr F Y A M M A(3) F A(3) M Y F A(6) F K M 3 F 4 F 5 2 M 6 Collar/tag Black col1ar Whi te eartag White tele Blue tele Yellow tele Orange eartag 000 Green 2 tele (Ch 2) Yellow dlag (Ch 1) Yellow eartag 79 Black collar 2 Yellow eartag (no no.) Black collar 3 Green 3 tele (eh 3) Red-White-Blue tele (Ch 2) Yellow w/Green "-" eartag Yellow w/Green "t" earlag Blue diag (Ch 4) Yellow w/Blue "t" eartag Black collar 4 Black tele r ((I-. I,) Trap l cce t ion [em ) l enqt h ( CI;I) 112 Left Right 16.5 16.5 Wt. (kg) Front leg fm scapula { (Ill) Hind foot (em) III III 112 5 Rl 53 53 52 TC 53 53 TC 52 III R2 112 51 51 S2 Yellow w/Blue "-" eartag Black collar 5 Black collar 6 Black collar 7 52 TC 112 F Green earlag Black collar 0 e F Red tele II (Ch 7) K 11 Black ee r taq F Girth I~rn to muzzle (cm) 112 RI 112 S2 52 114 116 159 165 77 116 102 152 9'1 69 116 72 110 99 142 113 95 121 100 128 147 14] 2.0 3.3 22.9 23.1 4.4 4.3 13.2 12.7 21.3 20.3 21.021.5 89 119 125 183 14.0 25.0 102 114 90 111 149 147 126 157 51 96 101 111 92 105 133 97 'i 21.6 21.3 27 74 2].9 79.4 41 21 25 41 77 22.2 80.0 14.I 24.0 10.5 26.0 60.0 57.0 80.0 30.0 27.0 34.0 19.0 25.5 19.0 73.0 07.0 51.5 73.0 30.5 31.5 27.0 01 146 145 159 21.7 22.4 2].8 23.0 15.0 15.5 24.0 23.0 0.4 0.4 19.] 19.4 23.0 2].4 21.4 22. I l]ll 17.8 17. I 147 21.9 21.7 140 114 21.8 5.3 21.2 5.3 74 45 25.0 12.0 20.5 22.0 23.5 19.5 58 79 ]3 23.0 2].8 15.2 9.9 42.0 76.0 82.0 81.0 73.0 76.0 31.0 21.0 30.3 29.5 30.5 30.0 78.0 30.0 61.0 26.ll �29 Page 2 (conttnued) Table 1. 51 118 153 23.4 22.7 51 87 107 4.2 4.1 F Green dlag (Ch 5) Blue eartag Grey eartag Y F F Black 1 dangle Black 2 dangle OAC OAC y M OAC 24 Jun 82 25 Jun B2 2 F Y M Black 3 dangle Blue dtag wide (Ch 4) "ld te eartag 1 25 25 25 25 26 26 26 y Unk Y M Y 6 y F F F 5 F •.;.I!,I_· Y F M Whl te 7 eartag White 6 eartag (right ear) 115 22.2 12.0 no 6 F K M 20 Jun B2 y 34 20 Jun 82 20 Jun B2 21 Jun 82 y 35 36 30 31 32 33 37 38 39 40 41 42 43 44 20 Oct 81 20 Oct 81 Jun Jun Jun Jun Jun Jun Jun 82 82 82 82 82 82 B2 84.0 16.0 64.0 22.3 12.1 79 - - 32.0 - 25.0 30.0 ~2.0 26.5 IlPP OAC eartag 2 Wid te eartag 3 Whtte eartag 5 (right ear) OAC OAC OAC Red tele 1 (Ch 7) 4 dangle III <1[?riag (left ear] 24.0 OAC .i" i le nl>r~ 75 30 OAC OAC OAC OI\C 45 46 26 Jun 82 27 Jun 82 27 Jun 82 47 29 5ep 82 5 4 4 Unk 48 29 Sep 82 Y F Black collar 8 11 OAC 90 150 119 49 30 5ep 82 K M Orange eartag I (left ear) 3 Y Black collar E Orange eartag 2 (left ear) 78 120 142 3.B 20.6 3.B 20.3 32 F F - - y F Orange eartag 3 (left edr) Black collar F OAC OAC OAC OAC OAC OAC OAC 118 182 23. I 23.6 - - - 32 .0 - - - 32.0 50 51 5.2 53 54 55 56 57 30 5ep 82 16 Jun 83 16 Jun 83 F F F Wid te 9 eartag White 8 eartag (left ear) Blad collar A R2 III OAC 2 F 3 M F 29 Jun 83 5 3 M Orange eartag 5 (right ee r] OAC (Ha rves t ed 10 Scp B3) SE Rogers 30 Jun B3 4 F Black collar II OIlC 119 16 Jun 83 16 Jun 83 29 Jun B3 Orange eartag 4 (right ear) Green dlag 112 (Ch 5) 171 23.7 23.7 N ~ W �<, N +> +> Table 1. Page 3 {continued} F F 58 59 60 6 Jul 83 7 Jul 83 7 Jul 83 2 9 1 61 62 63 7 Jul 83 7 Jul 83 11 Jul 83 6 3 5 64 65 66 67 83 84 84 84 84 84 3 1 3 2 F F 69 10 27 28 13 14 20 3 2 M 70 71 8 Aug 86 14 Aug 86 2 1 68 Aug Jun Jun Jul Sep Sep M F M F F F F M F Orange eartag 6 Black collar K Orange eartag 7 OIlC OIlC OIlC 120 162 24.1 24.4 - - - 31.0 Black collar M Orange eartag 8 Black collar N OAC OIlC 111 107 175 158 24.7 21.3 22.4 21.7 - - - 31.5 31.0 110 166 24.7 24.7 - 0 1 P V OAC OIlC OIlC OAC OIlC 112 88 114 110 157 113 1!>.1 137 OAC OAC 111 141 - - 16.2 11.9 23.1 lB.2 20.1 20.2 30.5 29.5 26.5 33.0 31.0 Black collar V Orange eartag 2 (left ear) 17.3 12.0 23.2 10.5 21.2 20.2 Black tele II (Ch 5) Black collar 1 II OAC OAC Black Blue Black Black collar eartag collar collar - - - - TT arc. Tumbling Creek at first switchback, Sl and S2 = Saddle below curve and mile post 6 (1 and 2. east and west trap, respectively). Rl and R2 • Rogers east above Lincoln Lake drainage and road between mile post 6 and 7 (1 and 2, north and south trap, respectively). DAC • Data acquisition center site at mile post 7. BPP • Below photo point on road shoulder between Rl - R2 and mile post 7. ~ - - - - 30.0 �245 Table 2. Number assigned to animal, telemetry collar, channel, status, frequency, pulse and activation date for radios outfitted on mount~:: goats and sheep in the Mt. Evans area. No. Collar Channel a Status Frequency (mhz) _:"c tivation Pulse per . min. (pprr. date Mountain goat 3 4 5 7 8 13 14 17 20 27 29 35 40 55 70 te Blue Fluorescent green Green 2 Yellow diag Green 3 Red-white-blue Blue diagd Black I Red II Green diag I Blue diag wide Red I Green diag II Black II \o."l.i 0 1 2 1 3 2 4 6 7 5 4 7 5 5 D A 148.130 172.487 ? ? 172.237 172.262 172.237 172.287 172.262 172.312 172.387 172.412 172.362 172.312 172.412 172.362 172.362 82 65-90c 65-90 65-90 75-120 65-90 60 90-65 65-90 65-90 172.362 172.237 172.287 172.237 172.462 172.437 172.262 172.287 172.387 65-90 65-90 65-90 90-65 90-65 90-65 65-90 75-120 65-90 D ? D D D D D ? D D AA 78 90-65b 22 Aug 80 26 Jul 83 11 11 Jul 83 48 29 30 7 9 30 11 14 20 24 25 29 8 Sep Sep Oct Jun Aug Aug Sep Oct Jun Jun Jun Aug 80 80 80 81 85 83 84 81 82 82 83 86 28 8 17 24 4 10 5 5 2S Dec Feb Jun Nov Dec Dec Sep Sep Oct 83 85 82 82 82 82 84 84 84 11 Mountain sheep 1 5 7 8 9 10 11 12 13 Black I Black & yellow Red I Yellow Black & white Blue White Red II Black II 5 1 3 1 9 8 2 3 6 D AA A A ? ? AA D AA -------------------------------------------------------------------------------aA = alive, AA = alive and active (transmitter), D = dead; as of 30 Jun 87. bActivity (tip switch) collars; 90 ppm head up, 65 ppm head down. CActivity (tip switch) collars; 65 ppm = head up, 90 ppm = head down. dDiag denotes diagonal. �246 Table 3. Mean duration of feeding, resting, and moving activity periods of 8 mountain goats and 6 sheep during day (sunrise-sunset) and night (sunsetsunrise) . Mean activity periods (min) Day (sunrise-sunset) No. Collar Feeding (n) Resting (n) Moving (n) Night (sunset-sunrise) Feeding (n) Resting (n) Moving (n) Mountain goat 4 14 17 20 27 29 40 70 Blue Red-white-b1ue Blue diagonal Black I Red II Green diagonal Red I Black II 83.3 (32) 72.7 (28) 137.1 48.1 8.0 ( 5) 24.6 (29) 175.3 (30) 24.1 (34) 67.1 (38) 11.7 33.9 (56) 177.5 (73) 16.5 13.7 ( 8) ( 9) 74.6 (72) 41.1 (72) 19.3 67.4 (42) 51. 3 (38) 8.0 ( 3) 37.5 (37) 207.4 (41) 80.5 (31) 53.7 (33) 13.8 (4) 28.8 (16) 364.0 (22) 62.0 101.0 5.0 12.4 ( 3) ( 5) (1) ( 5) 86.5 (4) 49.0 48.2 ( 3) ( 4) 89.8 (8) 37.9 68.8 (35) 58.4 (36) 10.0 69.4 36.5 ( 5) ( 8) 59.3 (19) 56.3 (19) 87.3 (6) 56.9 (29) 38.5 (28) 12.0 37.6 215.4 (1) (7) (11) 39.5 (46) 36.7 (43) 7.5 30.0 (25) 78.0 (29) ( 6) 48.0 (4) 117.5 (4) 116.8 (4) 317.7 29.1 (29) 168.6 (36) 7.0 (1) 70.8 49.6 (6) (5) 30.2 (4) 34.4 (7) (3) (6) (3 ) (3) Mountain sheep 1 5 8 9 11 13 Black I Black and yellow Blue Black and white White Black II ( 3) (7) 222.6 (10) 68.3 27.8 321.7 ( 6) ( 6) ( 2) 9.3 (3) ( 9) 2.0 (1) �247 Table 4. Estimated re roductive St~tus of collared female mountain Est'd No. 3 4 5 7 8 10 12 13 14 17 19 20 22 23 24 26 27 29 31 35 40 47 48 50 53 55 57 42 59 61 63 64 66 67 68 70 71 Date first collared Collar White tele (813) Blue tele (Ch 0) Fluorescent green (CH 1) Green 2 (Ch 2) Yellow diag (Ch 1) Black colla~ 2 Blacl< colla T 3 Green 3 (Ch 3) Red-white-blue (Ch 2) Blue diag (Ch 4) Black collar 4 Black tele (Ch 6) Blacl< collar 5 Black collar 6 Black collar 7 Black collar 8 I Red tele II (Ch 7) Green diag I (Ch 5) Black collar Zi Blue diag wide (Ch 4) Red tele I (Ch 7) Black collar A Black collar 8 II Black collar E Black c o Ll ; - F Gr e e z d i a .. :Ch 5) Bl_j_,.;, co: BlacK col, ~j B':c..:k col] Ii:- K BLci< co.Ll ar ~ Black Black Black Black Black Elack Black when first collared 22 Aug 80 18 Sep 80 22 Sep 80 29 Sep 80 30 Sep 80 30 Sep 80 7 Oct 80 7 Oct 80 9 Jun 81 16 Jun 81 17 Jun 81 29 Jun 81 28 Aug 81 30 Aug 81 4 Sep 81 15 Sep 81 6 Oct 81 20 Oct 81 18 Sep 84 24 JUD 82 25 Jun 82 29 Sep 82 29 Sep 82 30 Sep 82 16 Jun 83 29 Jun 83 30 Jun 83 30 Jun 83 7 Jul 83 7 Jul 83 11 Ju1 83 Ie Aug 83 28 Jun 84 13 Jul 84 14 Sep 84 8 Aug 86 14 Aug 86 collar N collar 0 collar P collar V collar Y tele II (Ch 5) collar 1 II oats 1n the Evans ~t. stud a!· _ age No. kids 4 3 4 6 2 8 2 7 3 3 1 6 3 4 5 6 8 6 3 2 6 U 1 3 2 5 4 5 9 6 5 3 3 2 3 2 1 1980 la 1981 1b 1 1 1 1 2 2 1 1 U _d o o U o 1982 o o 1 1 U o 1 1 1 1 1 1 1 1 1 U If Ih 2 1 1 o Ie 1 1984 U o 1 0 1 1 1 C 1 u U U u U 1 U U 0 U U 1 U U 2 U 1 2 1 1 1 0 U o 1 o 1 U U Ie o o 2 1983 o 1 1 U U U U U 1 o 1 U U 1 1 1 2 U 1 U o o o o o o 1 U U U 1 1 2 0 1 o 1 U o li 1 o o 1 1 o :?87 1 1 o o 1986 2 1 1 o o o u u u U t: r 1 U 1 1 1 1 1 1 1 o U 1 o o 1 1 u o o BWhite eartag 1bConfirmed with 1 kid 2 Jun; without kid 29 Jun and thereafter. ~Parturition 1 Jun yielded 2 kids; with only 1 kid 4 Jun and thereafter. Mortality fall 82 - summer 83. ~Blue "-"; after eartag pulled out, knoW!) as "i::ljuredear." Black eartag. gAt least 1 is evidenced by milk in udder at time of death. bBlue eartag. ipreviously Grey eartag. jPreviously White eartag 4. Table 5. Number of adult (~4 years old) marked female mountain goats associated with 0, 1, or 2 neonates or kids across years in the Mt. Evans study. No. of neonates 0 1 2 Years 1980 1 4 0 Total 1982 1983 1984 0 4 9 2 11 2 7 10 2 1981 1985 1986 4 4 4 9 3 13 0 7 3 1987 2 7 0 No. 26 70 12 Percent 24 65 11 --------------------------------------------------------------------Total 5 11 17 19 16 17 14 9 108 100 �'). " L'+0 :a:':o? 6. r e c r od c c t i ve :'s!!:oar€:c sta:t.:s of :::a!'Ke~ 0" Date Co!la~ So. Black te2.e I :~ ~ 2 Ye Ll ov 3 Plain yellow £artag 5 6 9 10 11 12 13 (Ch first collared 5> Black and yellow tele (Ch 1) Partial albino Black and white tele (Ch 9) Blue tele (Ch 8) 'White tele (Ch 2) Red tele (Ch 3) Black tele II (Ch 6) ::'de;,:!:!2~:e fe::21e shee:l mountain Est. age 1 Fet 77 1 ret 77 77 25 Jan 77 1 reb 77 Jun 84 198G 12 12 U 7 7 1 :S2: 1982 1 0 0 0 I,; c 0 0 0 0 I,; 4 Dec 82 10 Dec 82 t ~t. Evans ne :983 1984 1985 0 C C c U u u u 1 1 1 C U 12 0 1 1 U 1 V 0 U U 6 5 3 5 Sep 84 5 Sep 84 25 Oct 84 ~ :"l s,::u:-:v area . :986 :;87 . ki d s ~~o t: 1 1 1 1 1 U 3 U t; V u 1 u U V U apreviously Yellow 17; Black tele collar added 28 Dec 83. b~ost numbers have come off. Only stitching and faded areas make identification possible. ~~o stitching or faded areas apparent; collar frayed OD bottom-poste~ior. Previously eartag 18; Black and yellow telemetry collar added 8 Feb 85. Table 7. Mortali ty of collared female mountain goats and mountain sheep in the 11t. Evans study. Date No. Collar A~e narked Recovered/Death Estimated cause ~1ountain goats 1 8 26 40 20 3 55 29 14 17 Black 1 I Yellow diag Black 8 Red tele I Black tele I Black Xa Green diag II Green diag I R-W-B te1e Blue diag 27 Red te1e II 1 4 6 8 7 9 7 8 5 7 22 30 15 25 29 22 29 20 9 16 Aug Sep Sep Jun Jun Aug Jun Oct Jun Jun 80 80 81 82 81 80 83 81 81 81 13 6 Oct 81 unk 13 5 Sep 84 1 Feb 77 8 <29 18 21 8 16 17 22 1 >24 <5 23 Sep 80 Jun 83 Sep 82 Jul 84 Sep 84 Jun 85 Sep 85 Jun 83 Nov 83 HarJun 86 Aug 86 Harvested winter Harvested Disease 1"inter Ioiinter Winter Injury/winter Disease Disease/winter Injury/dispatched Mountain sheep 12 1 Red tele I Black te1e Ib 1 Jun 85 1 Sep 85 winter Age/winter apreviously White te1e. bpreviously Yellow 17; fitted with telemetry collar 28 Dec 83. �Colorado Division of Wildlife Wildlife Research Report July 1987 249 JOB FINAL REPORT State of Colorado Project No. 01-03-048 (FW 26 p) ------------~------~-- Mammals 2 Research Work Plan No. 5 Black Bear Investigations Job No. 1 Black Bear Population Ecology Period Covered: July 1, 1986 - June 30, 1987 Author: T. D. I. Beck ABSTRACT Data files and computer programs stored on a tape in a nonstandard FORTRAN format have been deciphered and converted to standard FORTIL~ and made .microcomputer usable. One black bear mortality and 5 observations of tagged bears were added to data from 1987. ��251 BLACK BEAR INVESTIGATIONS Thomas D. I. Beck P. N. OBJECTIVE Determine black bear life history and population characteristics through research. SEGMENT OBJECTIVES Analyze data and prepare manuscripts for publication. ACKNOWLEDGMENTS Assistance with data compilation and debugging computer programs was provided by Michael Smith. RESULTS AND DISCUSSION The major accomplishment in this segment was finally extracting the data and appropriate programs from a master computer tape. Most of the programs were written in a nonstandard FORTRAN language, and many hours were spent deciphering the programs. All data and standard FORTRAN language programs are now usable on microcomputer. In the clear vision of hindsight, it would have been faster and easier to start over on data entry rather than debug the master tape from the University of Montana. Analyses of denning and migration manuscripts were completed. All data for home range and movements analyses are now stored. All handling data are also in computer files. No tagged bears were reported killed during the Fall, 1986 season while one (M-16) was reported killed during the Spring, 1987 season. Additionally, 5 tagged bears were treed during the 1987 season and the tag numbers reported. A tagged female with cubs added to our age of first reproduction data. ~/ [.i / /' ;: Prepared by ~~~ __,_~~>/?_~~_~~~ __v.~y_·~,/_:,~_/,'_~c_~~.~~·· _ Thomas D. I. Beck Wildlife Researcher ��Wildlife Research Report July 1987 253 JOB PROGRESS REPORT State of Colorado Project No. 01-03-048 (FW 26 p) --~----------~------~- Mammals 2 Research Work Plan No. 6 Mountain Lion Investigations Job No. 1 Mountain Lion Population Dynamics Period Covered: July 1, 1986 - June 30, 1987 Author: A. E. Anderson ABSTRACT Sixteen puma (Felis concolor) were captured for the first time and 6 puma recaptured and recollared. Since 1981, 52 puma have been captured and 44 of those radiocollared. Aerial telemetric locations of 29 puma ranged from 1 to 44 per puma and totaled 714. Twenty-two puma were being radiotracked at the end of the fiscal year. The death of 7 puma this fiscal year reflects the continuing trend of high mortality; 22 confirmed deaths among 52 captured puma. Sport hunting outside the study area and our capture activity accounted for 7 and 9 deaths, respectively; and both factors combined accounted for about 73% of the total known mortality among 15 radio-collared puma. The mean, SD, minimum-maximum number of days elapsed from capture to death was 308.3 ~ 246.7, 5-829, and the median number days elapsed was 243. There is some evidence that the method of capture may affect sex ratios among age groups. ��255 MOUNTAIN LION POPULATION DYNA}1ICS Allen E. Anderson P. N. OBJECTIVE To assess the effects of sport hunting on mountain lion populations. SEGMENT OBJECTIVES 1. Capture and mark up to 12 mountain lions. 2. Monitor mountain lion movements. ACKNOWLEDGMENTS I thank D. Allen, D. Bowden, P. Burke, G. Cheney, C. Chipman, D. Coven, F. Fields, M. Gallagher, B. Gill, J. Gray, L. Green, M. Gruebel, G. Hanson, M. Hershcopf, J. Herwick, J. Humphry, B. Kattner, D. Kattner, E. Kattner, D. Masden, D. McCauley, K. Miller, J. Olterman, P. Pope, M. Potter, D. Rowley, S. Steinert, M. Stevens, and M. Stone for their assistance. D. M. Kattner, project puma hunter, whose efforts once again were consistently above and beyond reasonable performance, deserves special recognition. I am also grateful to D. Miller, Vice President, Research and Education Department, for facilitating financial assistance from the National Wildlife Federation and to the laboratory staff of Montrose Memorial Hospital, especially C. Cruz for providing facilities and technical services for the "Collaborative Wild Cougar Project." Finally, we thank Dr. M. Roelke for saving the life of a pumainjured hound. METHODS AND MATERIALS Methods are described in Anderson (1982). One-hundred-sixteen days were spent hunting from November 17, 1986, to May 21, 1987. Puma were located with aerial telemetry at approximate weekly intervals. Using criteria previously described (Anderson 1983); only those locations subjectively judged as being within 1.5 mm of the actual location are presented in this report. Estimated telemetry locations of each puma were plotted on USGS topographic county maps (scale 1:50,000; contour interval 80 ft) or USGS topographic guadrangles (scale 1:24,000; contour interval 20 ft) if locations were densely aggregated. Hunting effort was allocated proportional to surface area among 4 previously established strata (Fig. 1 and Table 1) so that each puma within the study area might have a similar probability of capture. �RESULTS AND DISCUSSION Allocation of Hunting Effort Proportional allocation of hunting effort (Table 1) was, again, not achieved. Hunting conditions and accessibility tend to differ greatly among strata, and some strata are rather consistently better, or worse, than others. So far, these factors have precluded a reasonable proportional allocation of hunting effort. Puma Capture and Telemetry During 1986-87, 6.9 days were required to radiocollar 1 puma for the first time (Table 2); our best capture rate thus far. Since 1981, 42 puma have been radiocollared for the first time in 560 days of hunting with hounds for an overall capture rate of 13.3 days per radiocollared puma (Table 2). Sixteen puma were captured and radiocollared for the first time during this fiscal year (Table 3). In addition, 6 puma were recaptured and their radiocollars replaced. A summary of the single and mUltiple captures of 52 puma since the study began is given in Table 3A. Recap~ures offer the opportunity to obtain data on puma growth (Table 4). Aerial telemetric locations of 29 puma radiotracked during this fiscal year ranged from 1 to 44 per puma and totaled 714 (Table 5). Of the 29 puma, there were only 18 whose locations were all within the study area (GMU 62). Individual locations for each of the 29 puma were listed in Appendix Tables A through Z3. Twenty-two puma were being radiotracked as the fiscal year ended. Mortality Note first that 7 puma died during this fiscal year (Table 5). This value exceeds the maximum legal kill of puma recorded during each of 5 hunting seasons on this study area (1977-82) (Game Management Files, Colo. Div. Wildl.). Since the study began, there have been 22 confirmed deaths among 52 captured puma (Table 6). Among 8 sources of mortality generalized (Table 6), sport hunting and capture related accounted for 7 and 9 deaths, respectively, and together comprised about 73% of the known total of 22 deaths. Among 15 puma whose date of death was exact or could be closely (± 3 days) approximated, the mean + SD, minimum-maximum number of days elapsed from capture to death were 308.3-± 246.7, 5-829 and the median number of days was 243. While there was a preponderance of males less than 24 months of (Table 6), the sex ratios of dead puma in this age group do not singificantly (p < 0.05) from equality. So far, the numbers of both sexes less than 24 months of age exceed those of 24 months age dying differ dead puma of and older. There is some indication that the sex ratios of small samples of living puma at first capture may be influenced by the method of capture. Thus, among 52 puma captured with hounds and leghold traps (Table 7), there were no significant (P < 0.05) differences from equality of the sex ratios regardless of age group. But among 44 radiocollared puma (Table 8) captured exclusively with hounds, males were in preponderance and differed significantly (P < 0.05) from equality among puma less than 24 months of age, whereas females were in �257 preponderance and differed significantly (p 24 months of age and older. < 0.10) from equality among puma Size of Home Range Analyses of data for 1 adult male and 5 adult female puma are essentially complete, and a first draft of a manuscript is in progress. Completion and submission of this manuscript for publication will be a priority item during 1987-88. Cervids Killed by Puma Eleven mule deer and 1 elk killed by puma included 8 heavily scavenged mule deer carcasses whose sex could not be determined (Table 9). Since 1981, 60 mule deer and 4 elk have been recorded as being killed by puma (Table 10). Often, carcasses were found while trailing with hounds. Only 42 mule deer carcasses could be identified by both sex and age class. Among 18 mule deer, 217 mos of age, the sex ratio (50:100) did not differ significantly (p < 0.05) from equality. Among 24 mule deer >18 months of age, however, the sex ratio (20:100) differed significantly (p < 0.01) from equality. Among 4 elk killed by puma, the 3 whose sex could be determined were all females (Table 10). "Collaborative Wild Cougar Project" The goal of this study, staffed by Melody Roelke, DVM--Florida Panther Project Veterinarian, Florida Game and Fresh Water Fish Commission, Gainsville; Steve J. O'Brien, PhD--Laboratory of Viral Carcinogenetics, National Cancer Institute, Frederick, Maryland; and D. E. Wildt, PhD; E. Jacobson, DVM, PhD, and J. G. Howard, DVM--Dept. of Animal Health, National Zoological Park, Washington, D.C., is to examine genetic diversity among subspecies of Felis concolor and how the endangered Florida panther compares to other presumably heterogeneic species. The protocol for these studies is described in Roelke et ale (n.d.). Since several puma required recapture and recollaring this spring, we were able to assist the Cougar Project by 3 recaptures; Stacey Hudelson and Brett Parker, freshman and senior veterinary medicine students, respectively, at Coloardo State University, obtained samples from puma #7 on 4-25-87, and Melody Roelke obtained samples from puma #5 on 5-9-87 and puma #21 on 5-12-87. But on May 2, 3, 1987, we were unsuccessful in capturing puma so Howard Martin, DVM, Colorado State University and Dave Kinny, DVM--Denver Public Zoo, returned home without the samples. Blood and skin biopsy samples were otained from each puma and, in addition, semen was obtained by electroejaculation from puma #5. I anticipate this cooperative study will continue during the next fiscal year and possibly beyond. LITERATURE CITED Anderson, A. E. 1983. Program Narrative Proj. 45-01-503-15050, Work Plan 6, Job 1, Mountain lion population dynamics 7pp + 3 tables and Appendix A. Colo. Div. Wildl., Fort Collins. �Roelke, M. E., S. J. O'Brien, D. E. Wildt, J. G. Howard, E. Jacobson. n.d. Collaborative wild cougar project; genetic, reproductive, and biomedical studies protocol. 4005 S. Main St., Gainsville, Florida, 32601. l5pp. Prepared by ~~.~ Allen E. Anderson Wildlife Researcher �Table 1. Chronology of puma hunting effort by strata, 1986-87 Uncompahgre Plateau (GMU 62). represents 1 day__of_hun_ting effort. Each date Strata and dates ----------------------------------------------------------------------Out of 1 E 2 E 3 L 4 E study area Month Nov 21,24,25 26,27,28 6 o 17,20 2 18,19 Grand total 2 10 o 22 2 22 Dec 3,4,5 3 8,18,19,23, 24,29,30,31 8 1,2,9,22, 26 5 7,10,11,12, 15,16- 6 Jan 8,17,19,21,22, 26,29 7 10,12,15,16, 18,20,27,28 8 2 1 3,9,23,30 4 Feb 2,12,17,19, 22 5 27,28 2 6,10,11,15 27 5 5 1 o 13 March 31 1 2,9,10,12 13,17,20 7 16,23 2 4,18,19,24, 25,28,30 7 o 17 April 7,9 2 8,14,18,20, 24,25,26 7 10,23 2 13,15,16,21, 22,27,30 7 o 18 May 5,21 2 1,2,3,9,10, 11,12 ,13,14, 15 7 1 6 1 o 14 10 §_,]_ Total 20 48 16 28 4 116 Target allocation 27 38 23 22 o 110 a19 days (underlined) were spent in activity other than capturing and radioco11aring puma for the first time; 6 days during which 6 puma were recaptured and reco11ared, 10 days in unsuccessful attempts to recapture and reco11ar 5 puma, 1 day locating a dead puma, and 2 days retrieving hounds. N (,Jl 1.0 �260 Table 2. Rate of radioco11aring puma using hounds, Uncompahgre Plateau, GMU 62,4-9-81 to 5-21-87. Period From 4- 9-81 12-14-81 1- 2-83 11-19-83 11-19-84 11-18-85 11-17-86 No. radioco11ared No. days hunted 4-29-81 2-21-82 5- 6-83 5-17-84 6-28-85 5-17-86 5- 7-87 1 3 7 5 7 5 l4b 16 32 91 111 77 136 97 16.0 10.7 13.0 22.2 11.0 27.2 6.9 Total 42 560 13.3 To Radioco11aring ratea aNo. days to radiocollar 1 puma; does not include replacing or refitting radiocol1ars or puma captured by methods other than pursuit by hounds and drug immobilization. bTwo additional puma, each 5 mos in estimated age, were caught inadvertently in legho1d traps by a recreational trapper and ear-tattooed 48 and 49, respectively. �261 Table 3. Details On the capture and body measurements Plateau (GMU 62). of 16 puma handled during FY 1986-87, Uncompahgre Ear tattoo number 37 Sex Date of capture Estimated age (months) Legal descr., capture site 1/4, S T Ii: U.T.M., capture site Elevation (m) capture site Radiocollar serial no. Transmitter freq. (MHz) Drug (ketamine:rompum,180: 90 mg/ml) Dosage (cc injected) Induction time (mins) Body wt (kg) Measurements (cm) Total body length Tail length Head-body length Chest girth Neck circumference Height at shoulder Head length Zygomatic breadth Ear length Hind foot Hind paw length Heel pad, max. diameters: Left front, length width Left rear, length width 38 39 40 41 42 43 M F 44 M M 12-12-86 36 M 12-19-86 7 M M 12-9-86 6 1-3-87 32 1-8-87 14 NEll 48N 121/ 748-4258 2,347 12909-01 149.9600 NII13 47N 101/ 244-4246 2,225 6583-01 148.5605 SE30 5IN 131/ 729-4281 1,768 6570-01 148.7195 SI/ 8 46N 81/ 257-4236 2,196 8391 149.6105 NE12 3 6 20 3 4 3.5 3 32.2 70.8 8.8 10 53.5 49.8 39.9 5 5 59.9 196 80 116 61 38 61 19.5 12.7 7.0 26.6 9.0 232 92 140 83 55 80 21.8 16.2 9.2 31.0 10.3 211 81 130 78 42 75 21.9 14.8 8.5 29.5 9.3 216 81 135 74 42 72 22.5 14.4 8.8 27.0 9.2 208 76 132 74 39 69 18.0 14.2 7.7 26.7 8.8 229 87 142 79 46 80 23.4 14.9 9.2 31.5 11. 7 4.0 6.0 4.0 5.0 4.1 6.8 5.2 6.1 5.3 6.6 4.7 5.8 4.8 7.2 4.6 5.6 3.8 5.4 3.5 4.8 5.6 6.9 5.0 6.5 F 6 27.8 SON 151/ 718-4276 2,164 17198-01 149.9410 1-10-87 12 S\l34 SON 13'W 734-4270 2,073 3958-01 148.1310 1-16-87 24 1-17-87 36 NE13 50N 131/ 738-4275 1,920 12903-01 149.900 N1117 ISS 100W 708-4291 2,377 3968-01 148.2100 Ear tattoo number 45 Sex Date of capture Estimated age (months) Legal descr., capture site 1/4, S T R U.T.M., capture site Elevation (m) capture site Radiocol1ar serial no. Transmitter freq. (MHz) Drug (ketamine:rompum,180: 90 mg/ml) Dosage (cc injected) Induction time (mins) Body wt (kg) Measurements (cm) Total body length Tail length Head-body length Chest girth Neck Circumference Height at shoulder Head length Zygomatic breadth Esr length Hind foot length Hind paw length Heel pad, max. diameters: Left front, length width Left rear, length width F 47 46 F M 48 F 49 50 51 52 F M M M 4-30-87 30 1-29-87 9 2-4-87 5 2-4-87 5 2-17-87 13 3-16-87 99. 717-4289 2,440 3962-01 148.0900 5.15 155 10011 712-4291 2,256 7009.01 148.3090 SII19 145 100. 716-4299 2,012 SII19 145 10011 716-4299 2,012 51117 155 10011 708-4291 2,347 26307-01 148.5300 NE9 48N 11\1 754-4257 1,891 12903-01 149.9000 NEl9 47N 9\1 246-4244 2,256 12900-01 149.B700 3 5 39.0 2 2.5 3 17.2 4 46.3 4 5 46.3 4 6 51.8 2.7 5 29.1 4 2 56.3 219 84 135 73 43 74 20.0 13.9 8.3 29.5 9.8 210 76 135 79 41 75 21.3 14.0 8.4 27.7 8.8 205 77 127 68 39 72 21.0 13.3 8.4 27.1 B.9 165 66 99 57 33 56 16.8 11.6 7.7 25.2 7.7 170 69 101 5B 30 59 16.3 11.5 25.0 7.3 211 79 132 78 43 74 20.7 14.1 B.7 29.0 9.9 197 74 123 66 39 6B 19.B 12.7 8.7 29.7 10.3 222 B3 139 78 47 78 23.5 15.1 9.5 31.1 12.1 5.4 6.5 4.6 5.6 4.4 5.7 4.3 4.9 5.2 6.5 4.3 5.7 3.6 4.B 3.3 5.2 3.9 5.B 3.7 4.B 5.2 6.0 3.8 5.5 5.2 6.8 4.2 6.0 5.1 6.4 4.9 6.0 1-20-87 30 1-22-87 60 SE20 S.'19 SON ISS 1211 740-4273 2,044 8769-02 149.6300 4 19.5 B.O 7 �262 Table 3A. Fifty-two puma captured 4-16-81 to 4-30-87, Uncompahgre Plateau (GMU 62). Radiocollar Ear tattoo number Date captured Number Frequency Sex Est. age (months) Body wt (kg) Status as of June, 1986 1 4-16-81 8389 149.5500 F 20 33.1 Disappeared GMU 40 8-82 2 1- 5-82 8772 149.7010 F 60 43.5 Illegal kill 1-10-82 3 1- 8-82 8775 149.8005 F 26 38.5 Disappeared GMU 62 4-83 4 1-21-82 8773 149.7215 F 24 42.5 4 4-14-84 12898 149.8400 51 Radiocollar replaced 4 11-17-86 12911-01 149.9800 84 Alive radiocollar replaced 5 2- 7-83 8774 149.7815 5 5-17-85 8772-01 5 5- 9-87 6 60 64.0 149.7010 87 68.1 3965-01 148.1700 112 2-10-83 12904 149.9090 6 8-24-85 12897-01 149.8310 7 3- 4-83 12909 149.9605 7 5-14-85 12901-01 149.8810 74 7 4-25-87 8390-01 149.5020 97 8 2-20-83 12901 149.8810 8 8- 4-83 10 3-23-83 M F F M 149.8705 M Litter of 2 on 12-14-83 and 4-14-84, litter of at least 1 on 1-24-85, and litter of at least 1 on 6-19-86, 10-9-86 Radiocollar replaced Radiocollar replaced; alive but with possible broken right rear leg, left paw healing but 2nd outer toe miSSing Blood and tissue biopsy samples obtained for Nat1. Cancer lnst. Microscopic count, on Site, of electroejacu1ated semen indicated very low sperm count. Looked thin. Badly worn lower canines, left upper canine broken near tip. Probably 2 young winter, 1984-85 and 2 young (D29, #30) during winter, 1985-86. Possibly 2 young May, 1986. 34.0 Radiocollar replaced, alive 60 149.8810 12900 30 Breeding status and comments 48 2 cubs of capture probable litter of 2 spring 1984, litter of unknoWIl size, spring 1985. 46.0 Radioco11ar replaced No sign of pregnancy Radiocollar replaced, alive No sign of pregnancy Blood and skin biopsy sample obtained for Natl. Cancer lnst. 6 25.9 12 32.9 Died during recapture (to adjust radiocollar) 6 26.3 Found dead of unknown causes 6-2-83. �263 Table 3A. continued 12 3-25-83 12911 149.9800 12 4-30-85 8773-01 149.7210 F 60 Mot her 0: 1113 and 1 other cub. 44.0 85 Radiocollar Lact at ing replaced, alive 13 4-13-83 12896 149.8200 13 10-11-83 12896 149.8200 M 9 36.3 15 52.0 Offspring Unknown, disappeared GMU 40 summer 1984. of g12. Recaptured to adjust radiocollar 14 12-12-83 12906 149.9310 M 84 15 12-15-83 12902 149.8900 F 60 15 3-21-86 16 1-15-84 12897 149.8310 M 30 55.0 Killed by puma, found 4-2-84 17 4-14-84 12910 149.9710 M 8 36.2 Killed by hunter 140.5 killfrom capture site in GMU 42 on 1-11-85 18 4-16-84 12903 149.9010 M 36 64.3 Killed by hunter in GMU 65 on 3-18-86 19 7-18-84 12908 149.9510 F 28.0 Killed by ADC Caught in 1eghold usn;s hunter tr ap for killing sheep in GMU 70 on 1-15-85 20 11-26-84 12899 149.8600 M 10 44.5 Proba b1e offsping of ii15. 20 1-12-87 6582-01 148.5500 36 62.7 21 12- 7-84 12905 149.9200 12 36.8 21 12- 8-86 12905 149.9200 36 21 5-12-87 3963-01 148.1400 41 49.9 22 1- 5-85 8771 149.6710 15 56.0 22 1- 7-87 3957-01 148.0800 39 69.8 Radioco11ar replaced, alive 23 1- 9-85 8390 149.5020 M 12 44.0 Killed by hunter in GMU 40 on 12-12-85 24 1-23-85 8773 149.7210 M 18 Found dead 6-22-84, possible predacide victim. Not lactating Radioco11ar replaced, found dead of unknown causes 3-28-86 87 F M 2 kittens 5-12 days old at capture. Mother of #20 48.0 Radioco11ar replaced, alive .Probable offspring of 117. Treed but not handled Nipples greatly enlarged. Radloco11ar replaced, alive Probably pregnant blood and skin biopsy samples obtained for Nat1. Cancer lnst. Died at cap- Severely ~ounded by ture another Site, probably on 1-23-85 puma a few days prior to capture. �264 Table 3A. continued 25 1-24-85 26 3-20-85 12906 149.9305 26 2-18-87 12906 149.9305 27 4-24-85 8769-01 149.6310 F 14 36.3 Killed by hunter in GHU 61 on 12-9-85 28 11-20-85 12910-01 149.9700 F 24 43.1 Killed by hunter in GHU 40 on 11-22-86 None 12-14-85 F 3.0 14.5 Killed by our hounds 29 12-17-85 F 2.3 5.9 30 12-17-85 F 2.3 6.4 30 5- 3-86 7.3 21.8 F 4 14.0 M 10 44.0 35 Unknown Too small to radiocollar. Caught in a leghold trap. Offsping of 112 left her at about 12 1I0S of age. Killed by hunter in Utah on 2-18-87 Unknown No evidence of lactation Caught in leghold trap Probable offspring of #6. As above. Left front paw missing probably as a result of trap injury. Unknown, alive on 5-3-86 12.9 Died in vet. hospital on 1-7-86 Leghold trap injury. 30 40.0 Alive Lactating; a litter of 1-3 cubs in Vicinity. 7 23.6 16 40.0 Killed by hunter in GHU 65 on 11-6-86 F 7 28.1 Alive M 7 30.9 31 12-30-85 32 1- 8-86 8770-01 149.6515 F 33 2- 4-86 12904-01 149.9090 F 33 11- 6-86 12904-01 149.9090 34 2-12-86 12908-01 149.9500 35 2-13-86 8774-01 149.7815 35 11-19-86 8774-01 149~7815 F 3.0 Verified by Steve Boyle DWM. Not verified is the legal description of the kill site: 528,T325, R22E, Indian Ck., north side of Blue Mtn., Utah. (West of Monticello). 16 Caught in 1eghold trap. 3 toes amputated trap Littermate Died of unknown causes of #35, #36. Last "alive" signal on 10-10-86, Roc Ck., Utah in GHU 40. Carcass dried. F 7 25.4 Alive M 8 33.1 Dead 149.9600 F 6 27.8 Alive 6583-01 148.5605 M 36 6570-01 148.7195 M 7 32.2 Alive? 1- 3-87 8391 149.6105 M 32 70.8 Alive 1- 8-87 17198-01 149.9410 M 14 53.5 Died at capture site, probably on 1-8-87 3960-01 36 2-13-86 None 4- 3-86 37 12- 9-86 12909-01 38 12-12-86 39 12-19-86 40 41 148.1105 Killed by our hounds. Alive Last signal received 4-6-87 Possible fatal injury in fall from tree �265 Table 3A. 42 continued 1-10-87 3958-01 148.1310 12 ~1 ':'9.0 Liist ~iO!:1J~ :-pce~"'c'j :n ';'1[; 63 .';11 V('? 4-24-87 43 1-16-87 12903-01 149.9000 39.~ 24 F Died at Cause 0: de a t h unknovn caoture site, pr o ba b Ly on 1-16-87 44 1-17-87 3968-01 148.2100 M 36 59.9 Alive 45 1-20-87 8769-02 149.6300 F 30 46.3 Alive 46 1-22-87 3962-01 148.0900 F 60 46.3 Alive 47 1-29-87 7009-01 148.3090 M 9 39.0 Alive In 1eghold t rap vhe n dogs caught up with it. Pulled "ut of trap leaving l digit and dis1ocat~ng ldt hind foot 48 2- 4-87 F 5 17.2 !JnknoW"'Q Caught in Le ghc Ld trap, litteroate or #49. 49 2- 4-87 F 5 19.5 unknown Caught in 1eghold trap. 50 2-17-87 26307-01 148.5300 tl 13 51.8 Alive 51 3-16-87 12903-01 149.9000 M 7 29.1 Died at capture Full stocach. site, probably ]-16-87 52 4-30-87 12900-01 149.8700 Table 4. Body weights (kg) and measurements (G~!L'62). Sex, ear tattoo no. Dates of capture Est. age (mos) Body lOt Total body length Tail length Head-body length Chest girth Neck circumference Height at shoulder Head length Zygomatic breadth Ear length Hind foot length Hind Eaw 1ensth M 11-26-84 10 44.5 206 81 125 70 39 69 14.4 8.0 27.0 8.1 30 M Alive (cm) of puma at first and second capture, Uncompahgre 20 1-12-87 36 62.1 212 89 123 78 44 74 22.9 16.2 8.6 29.2 8.8 56.3 16.3 12.1 8.1 24.5 8.3 5-12-87 41 49.9 206 83 123 72 37 73 20.7 13.9 8.4 28.0 8.5 F 33 F 30 F 21 12-7-84 12 36.8 193 77 116 63 35 12-17-85 2 6.4 110 39 71 33 21 40 13.5 8.3 8.3 17.8 6.1 5-3-86 5 21.8 167 65 102 57 30 57 16.8 13.7 7.9 26.0 7.4 Plateau 2-4-86 7 23.6 176 74 103 58 31 62 19.0 12.1 8.3 27.0 7.6 11-6-86 16 40.0 192 77 114 61 36 67 21. 6 14.9 8.9 27.9 8.9 �266 Table 5. Number of aerial telemetric locations of 29 puma radiotracked during FY 1986-87; 22 were bein~ tracked at end of the fiscal year. Ear tatoo no. 4 5 6 7 12 20 21 22 26 28 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 50 51 52 Capture date 1-21-82 2- 7-83 2-10-83 3- 4-83 3-25-83 11-26-84 12- 7-84 1- 5-85 3-20-85 11-20-85 1- 8-86 2- 4-86 2-13-86 11-19-86 2-13-86 12- 9-86 12-12-86 12-19-86 1- 3-87 1- 8-87 1-10-87 1-16-87 1-17-87 1-20-87 1-22-87 1-29-87 2-17-87 3-16-87 4-30-87 Sex F M F F F M F M M F F F F M F F M H M M M F M F F M M M M Est. age <months) on 6-87a GMU locations No. locations 1986-87 89 113 82 99 112 32 41 44 41 15 15 12 42 13 38 18 42 36 66 14 17 31 Total 44 44 44 43 43 38 44 42 17 10 44 11 42 9 32 27 26 13 23 2 10 2 20 20 22 17 16 1 8 714 a"Dashes" indicate puma died before 6-87. ----------------Majorit:t: 62 62 62 62 62 62 62 65 62 40 62 65 62 62 65 62 62 62 62 62 62 62 62 62 62 62 62 62 62 Minor 0 0 0 0 0 0 0 62,64 40,61 0 0 62 65 61,40 62 0 0 0 0 0 53,63,64 0 40 0 0 40 65 0 0 �267 Table 6. 1981-87. Mortality among puma captured on the Uncompahgre Plateau, GHU 62, Ages estimated at the time of death. Less than 24 months Male Female 24 months and older Male Female Total Radiocol1ared Unconfirmeda Illegal kill b Unknown cause(s)C Sport huntingd Suspected predacidee Intraspecific strifef Predator manae~ntg . Capture-related Subtotal 1 1 1 1 2 2 1 1 1 2 2 1 4 -2 8 4 1 2 9 6 5 5 3 1 2 7 1 1 1 6 22 Not Radiocollared Capture re1atedi Total 3 5 5 25 apumas #1, #3, #13 bPuma #2 died 5 days after capture. cPumas #10 and #35 died about 69 and 243 days from capture, respectively. dPumas #17, #18, #23, #26, #27, #28, #33 died 272, 701, 337, 700, 229, 367, 275 days, respectively, after capture. epuma #14 died about 186 days after capture. fPuma #16 died about 73 days after capture. gPuma #19 died 181 days after capture. hPumas #24, #41, #43, #51; #8 died 165 days after capture, #15 died about 829 days after capture. iPuma #31, 1 unnumbered male, 1 unnumbered female. �263 Table 7. Age and sex classes of 52 puma at fist capture on the Uncompahgre Plateau, GMU 62, 1981-87. Includes all Euma handled. Year 1981-82 1982-83 1983-84 1984-85 1985-86 1986-87 Total Less than 24 months of age 24 months and older Male Male Female 4 2 3 8 12 52 1 0 3 1 5 2 0 1 3 6 17 15 Total 4 7 6 9 11 16 0 1 4 7 3 Female 0 0 3 3 1 0 Table 8. Age and sex classes of 44 puma radiocollared at first capture, Uncompahgre Plateau, GMU 62, 1981-87. Excludes 8 puma handled but not radiocollared. Less than 24 months of age Year 1981-82 1982-83 1983-84 1984-85 1985-86 1986-87 Total Male Female 24 months and older Male Total 4 2 3 4 7 6 7 6 14 6 14 44 0 3 1 0 0 1 5 1 7 1 2 1 1 17 7 3 0 Female 0 0 3 3 3 0 aThe 8 puma excluded are; 5 caught in leghold traps by commercial trappers and too small (~ 27.2 kg) to radioco1lar, 1 caught in a 1eghold trap and died of trap injury in Vet. Hospital, and 2 cubs killed by hounds. �269 Table 9. Eleven mule deer and 1 elk that were killed bZ Euma, 1986-87, Uncom2ah~re Plateau (GHU 62). Legal description ----------- Date found SEecies Sex 7- 2-86 Dlule deer F 9-17-86 Est. ase (mos.) S T R 18+ SW 20 47 10 oak, PJ Third neck vertebrae Dlu1e deer <6 NE 18 47 10 PJ Inside cave 9-17-86 Dlu1e deer 18+ NE 18 47 10 PJ Inside cave 9-17-86 mule deer 18+ NE 18 47 10 PJ Inside cave 10- 8-86 mule deer 18+ NE 18 47 10 PJ First 5 entries from Spring Ck 10- 9-86 Dlule deer <6 NIl 8 47 10 PJ 8 NE 33 47 8 9 NE 9 48 11 NIl 4 47 9 F 1/4 Prob. killed by #22, GMU 65, Billy Ck Thoracic organs and portions of lover neck and 1 thigh eaten PJ Thoroughly oak, PJ Partially leaves 13 PJ Thoroughly eaten 50 13 PJ Thoroughly eaten 6 48 11 riparian Partially needles 34 48 11 oak Not covered, elk 3-16-87 mule deer 3-25-87 mule deer 4- 8-87 mule deer NIl 26 51 4- 8-87 mule deer NIl 6 4-22-87 mule deer F 46 NIl 6- 2-87 mule deer F 20 SE 108+ broken Edge of irrigated meadow 1-7-87 F Comments Habitat eaten eaten covered with covered with pine thoroughly �270 Table 10. Age classes and sex of cervids killed by puma, Uncompahgre Plateau (GMU 62), 1981-87.a mos 13-17 mos ------- --------- 92 Fiscal ear 18+ mos ------- SEecies M F M F M F Sex unk Age unk Sex and age unk 1980-81 mule deer elk 0 0 1 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 3 0 1981-82 mule deer elk 1 0 0 0 0 0 0 0 2 0 6 0 0 0 0 0 0 0 9 0 1982-83 mule deer elk 0 0 1 0 1 0 2 0 0 0 5 0 0 0 0 0 0 0 9 0 mule deer elk 3 4 0 0 0 1 1 0 0 2 0 1 1 0 1 0 0 0 12 2 mule deer elk 1 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 4 0 7 0 mule deer elk 0 0 1 0 0 0 -1 0 1 0 2 0 4 1 0 0 0 0 9 1 mule deer elk 0 0 0 1 0 0 0 0 0 0 3 0 6 0 0 0 2 0 11 1 Total mule deer 5 8 1 4 4 20 11 1 6 60 Total elk 0 1 0 1 0 1 1 0 0 4 x 1983-84 1984-85 1985-86 1986-87 "i elk (1986-87) from GMU 65 (Billy Ck) probably killed by puma 1122. Total �271 APPENDICES �272 AEEendix Date 8-18-86 8-22-86 8-29-86 9- 5-86 9-12-86 9-19-86 9-25-86 10- 5-86 10-10-86 10-17-86 10-24-86 10-31-86 11- 6-86 11-14-86 11-21-86 11-25-86 12- 5-86 12-12-86 12-19-86 1- 3-87 1- 9-87 1-16-87 1-24-87 1-30-87 2- 6-87 2-15-87 2-20-87 2-28-87 3- 6-87 3-13-87 3-21-87 3-28-87 4- 6-87 4-10-87 4-17-87 4-24-87 5- 1-87 5- 8-87 5-15-87 5-22-87 5-30-87 6- 5-87 6-12-87" 6-20-87 Table A. Aerial telemetry locations Legal description U.T.M. ------------------ ----------- of adult female Euma 114. 1/4 5 T R X Y Approx. elev (m) Nil 17 18 7 18 18 18 18 8 8 33 34 17 18 8 8 28 5 34 4 33 28 28 9 33 5 33 33 28 32 8 8 9 8 8 17 8 9 33 33 30 19 26 26 4 47 47 47 47 47 47 47 47 47 48 48 47 47 47 47 48 47 48 47 48 48 48 47 48 47 48 48 48 48 47 47 47 47 47 47 47 47 48 48 47 47 47 47 47 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 LO 10 10 10 10 10 10 10 10 10 10 10 9 10 10 10 761 761 761 761 760 761 761 762 238 240 241 762 761 762 762 240 239 241 240 240 239 240 240 239 762 240 239 240 762 762 762 239 762 762 762 761 239 240 240 760 246 243 243 240 4246 4246 4246 4245 4246 4246 4246 4247 4247 4250 4250 4245 4246 4248 4247 4251 4248 4250 4248 4251 4252 4251 4247 4251 4248 4251 4251 4252 4251 4247 4248 4248 4248 4247 4245 4246 4247 4251 4251 4242 4243 4242 4241 4248 2,316 2,408 2,408 2,377 2,438 2,438 2,347 2,286 2,225 2,134 2,164 2,347 2,438 2,347 2,347 2,134 2,196 2,164 2,225 2,164 2,134 2,134 2,256 2,164 2,316 2,134 2,196 2,103 2,196 2,347 2,316 2,316 2,347 2,316 2,438 2,316 2,347 2,042 2,103 2,560 2,256 2,408 2,438 2,286 NE SE 51< NE NE NE SI< NE SE SI< SI< NE NW NW SE SE SI< SE N" )',"\01 S\'" NE Nh 51.' NE l,1< Sf )',"\; )',1: Nl> m; ]I•••• m, SE S\'" SI< NE H SI-' S\; SIo: S\i SE Distance (km) between locations 4.3 0.6 0.2 1.4 1.3 2.2 0.6 1.1 1.8 2.9 0.9 5.9 1.6 2.3 0.5 4.5 3.6' 3.2 2.0 2.9 1.4 1.3 3.9 2.0 3.0 3.5 2.6 1.9 2.8 3.7 0.4 1.2 1.4 0.3 2.5 1.7 1.7 5.7 0.6 9.9 10.3 2.9 0.7 7.5 Rating Major draina!le Middle Fk Spring Spring Spring West Fk Spring Spring Spring Spring Spring Spring Spring Spring East Fk Spring Spring Spring Spring Spring Spring Spring Spring Devinny Lindsay Devinny Spring Spring Spring Spring Devinny Devinny Lindsay Spring Spring Spring Spring Spring Spring Spring Spring Spring Spring Middle Fk Spring Dolores Happy Happy Spring -----------------Signal Location G F G G G F G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G F G G G G G G G G G G G G G G G G G G G G F G G F G G F G G G G F F F G G F G F G G G F G G G �273 AEl~endix Table B. Aerial telemetry locations of adult male Euma liS. Legal description -----------------Date 8-18-86 8-22-86 8-29-86 9- 5-86 9-12-86 9-19-86 9-25-86 10- 5-86 10-10-86 10-17-86 10-24-86 10-31-86 11- 6-86 11-14-86 11-21-86 11-25-86 12- 5-86 12-12-86 12-19-86 1- 3-87 1- 9-87 1-16-87 1-24-87 1-30-87 2- 6-87 2-15-87 2-20-87 2-28-87 3- 6-87 3-13-87 3-21-87 3-28-87 4- 6-87 4-10-87 4-17-87 4-24-87 5- 1-87 5- 8-87 5-15-87 5-22-87 5-30-87 6- 5-87 6-12-87 6-20-87 1/4 NE NW NW SE NI sv SI SE ffi' SW SE SE SE NE SE NW Ni.' SE 1\..••. SE NE N10i l'.,\; S\\ SE NE I'<,\; NE SE SW NE SE SW SW NE ffi' NE SW I'<,\; NE SE SE SE SW U. T.M. ----------- S T R 35 3 28 27 3 15 7 6 24 27 29 29 20 11 11 11 35 1 24 14 18 16 4 6 18 15 15 12 13 30 31 26 5 6 31 9 6 19 17 4 2 17 2 30 49 48 49 49 48 49 48 48 49 49 49 49 49 48 48 48 49 49 49 49 47 49 48 49 49 49 49 49 49 50 50 50 49 49 50 49 48 49 49 49 48 49 48 49 13 13 12 13 13 13 12 12 13 13 12 12 12 12 12 12 12 12 13 13 12 11 12 11 12 12 12 12 13 12 12 13 12 12 12 13 12 12 12 12 13 13 13 12 X 736 736 742 735 737 734 741 741 737 734 74.1 741 741 748 748 747 745 742 737 736 740 751 744 748 739 744 743 748 739 738 739 736 740 740 740 732 742 739 740 742 738 732 738 739 Y Approx. e1ev (m) 4261 4259 4263 4262 4259 4265 4256 4258 4264 4262 4262 4262 4262 4257 4257 4257 4261 4268 4264 4265 4265 4266 4259 4269 4265 4266 4266 4268 4265 4272 4270 4271 4268 4268 4271 4267 4259 4264 4266 4269 4258 4265 4258 4262 2,499 2,560 2,134 2,499 2,621 2,438 2,530 2,408 2,438 2,499 2,256 2,134 2,256 2,377 2,377 2,438 2,316 2,012 2,377 2,225 2,134 1,920 2,256 1,951 2,164 2,134 1,859 2,042 2,316 2,042 2,164 2,103 2,225 2,134 2,134 2,286 2,469 2,073 2,073 2,012 2,560 2,469 2,560 2,286 Distance (km) between locations 4.3 2.3 7.5 7.3 3.0 6.4 11.4 2.9 6.9 3.4 7.4 0.4 0.4 8.3 0.5 0.7 5.0 7.6 10.7 1.7 3.7 11.5 11.1 10.7 9.5 5.0 1.0 5.1 10.0 6.8 1.6 3.0 4.6 1.7 2.6 8.3 12.4 5.2 3.0 3.3 11.8 9.3 9.4 3.8 Rating Major drainage -----------------Signal Criswell Criswell Roubideau Potter Moore Monitor Beach Terrible Potter Potter Traver Traver Traver Cushman Cushman Cushman Roubideau Middle Fk Roatcap Criswell Potter Criswell Coal bank Roubideau Roatcap Gulch Criswell Roubideau Roubideau East Fk Roatcap Gulch Criswell Monitor Potter Monitor Potter Potter Potter Cottonwood Wright Criswell Criswell Criswell Traver Little Monitor Traver Moore G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G Location F F G F F G G G G F G G G G G G G F F G F G G F G G F F F G F F G F F F F F G F G F G G �274 Appendix Table C. Aerial telemetry locations of adult female Euma 116. Legal description -----------------Date 8-18-86 8-22-86 8-29-86 9- 5-86 9-12-86 9-19-86 9-25-86 10- 5-86 10-10-86 10-17-86 10-24-86 10-31-86 11- 6-86 11-14-86 11-21-86 11-25-86 12- 5-86 12-12-86 12-19-86 1- 3-87 1- 9-87 1-16-87 1-24-87 1-30-87 2- 6-87 2-15-87 2-20-87 2-28-87 3- 6-87 3-13-87 3-21-87 3-28-87 4- 6-87 4-10-87 4-17-87 4-24-87 5- 1-87 5- 8-87 5-15-87 5-22-87 5-30-87 6- 5-87 6-12-87 6-20-87 U.T.M. ----------- S T R X y 25 34 3 mo' 3 S"_' 12 NE 8 f-,'\, 16 NE 7 SE 6 f-,'j.; 30 S"_' 9 NE 16 SE 6 SE 29 Nj.; 18 NW 30 N\; 20 S"_' 19 NE 8 NE 30 NE 30 Sj.; 14 NE 30 I'<'W 28 NO; 21 1'<1\ 28 S"_' 21 n 28 S"_' 17 }.1\ 17 )','j.; 8 N\; 4 Sh 7 S"_' 25 Sh 18 }.1\ 24 SE 29 SE 24 )',"1,,' 31 NE 35 Sh 31 Sh 30 SE 36 mo' 31 49 49 48 48 48 48 48 48 48 49 48 48 48 49 49 49 49 49 49 49 49 49 49 49 49 49 49 49 49 49 49 49 49 49 49 49 49 49 49 49 49 49 49 49 13 13 13 13 13 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 13 12 12 12 12 12 12 11 12 12 12 12 13 12 13 12 13 12 13 12 12 13 12 738 735 736 736 739 743 744 742 741 740 744 744 741 741 739 739 740 740 741 740 740 736 740 742 742 742 742 743 750 740 740 744 739 737 739 738 742 738 739 737 739 739 738 739 ",262 4260 4258 4259 4256 4257 4256 4257 4258 4262 4256 4255 4258 4262 4265 4262 4264 4263 4267 4263 4262 4265 4262 4262 4264 4263 4264 4263 4265 4265 4268 4269 4266 4262 4265 4264 4262 4263 4261 4261 4260 4262 4260 4261 1/4 SO; SE Sj.; Approx. elev (m) Distance (km) between locations 2,377 2,621 2,591 2,621 2,621 2,377 2,499 2,438 2,347 2,316 2,377 2,377 2,408 2,256 2,286 2,377 2,164 2,134 2,012 2,286 2,134 2,256 2,408 2,196 2,164 2,073 2,073 1,920 2,073 2,164 2,196 1,920 2,256 2,347 2,256 2,316 2,196 2,256 2,438 2,530 2,377 6.7 2.8 2.0 1.8 4.4 4.3 1.9 2.6 1.9 4.5 7.8 1.0 4.0 4.6 4.3 3.3 2.1 0.9 4.7 4.7 0.4 4.3 4.5 2.8 1.9 1.7 1.1 1.3 7.2 9.5 2.4 2.5 4.0 4.9 3.4 1.8 4.4 3.3 2.4 2.4 2.6 2,499 2,408 1.1 1.6 Rating Major draina!le Criswell Criswell Criswell Criswell Wright Bull Ck Roubideau Beach Terrible Moore Roubideau Roubideau Terrible Traver Criswell Moore Moore Criswell Criswell Moore Moore Potter Moore Traver Roubideau Roubideau Roubideau Roubideau Coalbank Criswell Potter Criswell Criswell Criswell Criswell Criswell Traver Criswell Moore Criswell Traver Criswell Moore Moore -----------------Si!lnal Location G G F F F F F G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G F F F G G G G G G G G G F F G F F F G G G F G G F F F F F G G G G G G G G F G P P G F F G �275 Appendix Table D. Aerial telemetr~ locations of adult female Euma 117. Legal description -----------------Date 8-18-86 8-22-86 8-29-86 9- 5-86 9-12-86 9-19-86 9-25-86 10- 5-86 10-10-86 10-17-86 10-24-86 10-31-86 11- 6-86 11-14-86 11-21-86 11-25-86 12- 5-86 12-12-86 12-19-86 1- 3-87 1- 9-87 1-16-87 1-24-87 1-30-87 2- 6-87 2-15-87 2-20-87 2-28-87 3- 6-87 3-21-87 3-28-87 4- 6-87 4-10-87 4-17-87 4-24-87 5- 1-87 5- 8-87 5-15-87 5-22-87 5-30-87 6- 5-87 6-12-87 6-20-87 1/4 S SW 20 NE 5 S1ft 22 SE 9 m; 10 SE 33 NE 3 SE 36 SW 1 NW 12 SE 36 NE 36 SE 8 SW 31 NE 36 NW 6 NE 1 NW 21 m; 20 SE 29 NE 30 SW 29 NE 5 NE 31 NW 21 NW 6 SE 20 NW 29 5E 25 SE 28 SW 26 NE 30 SE 32 NW 9 SW 4 SW 29 }''\; 12 NW 23 N\" 23 SE 31 SE 27 51< 25 51< 2 U.T.M. --------- T R X Y Approx. elev (m) SO 13 13 14 14 14 14 14 14 14 14 14 14 13 13 14 13 14 13 13 13 13 13 13 13 13 13 13 13 14 13 13 13 13 13 13 13 14 14 14 13 14 14 14 730 730 724 723 724 723 725 729 727 728 728 728 731 729 728 729 728 732 732 731 730 731 731 730 732 729 731 731 728 733 735 730 731 732 732 730 728 726 727 730 725 727 726 4273 4268 4273 4266 4267 4270 4269 4269 4268 4267 4270 4270 4276 4270 4271 4268 4269 4273 4274 4271 4272 4271 4269 4270 4274 4269 4273 4272 4271 4271 4271 4272 4270 4267 4268 4271 4267 4264 4264 4269 4261 4262 4268 2,012 2,408 2,164 2,196 2,347 2,408 2,164 2,103 2,286 2,256 2,256 2,256 2,196 2,225 2,256 2,347 2,196 2,134 2,073 2,256 2,012 2,377 2,347 2,377 2,134 2,316 2,134 2,288 2,316 2,225 1,890 2,225 2,347 2,286 2,134 2,316 2,347 2,469 2,530 2,377 2,621 2,530 2,408 49 SO 49 49 SO 49 SO 49 49 SO SO SO SO SO 49 49 SO 50 SO SO 50 49 SO SO 49 SO SO 50 SO SO SO SO 49 49 SO 49 49 49 SO 49 49 49 Distance (km) between locations 14.0 4.8 8~0 6.7 1.2 3.1 1.8 3.5 1.7 1.7 2.4 0.8 6.2 6.6 1.0 0.8 0.4 6.2 1.0 2.8 1.6 1.6 2.1 2.0 4.1 6.0 6.0 1.4 2.8 5.1 2.0 5.1 2.3 2.5 1.2 3.4 5.0 3.3 0.9 7.1 9.3 2.4 4.7 Rating Major draina8e Dry Fk Escalante Cottonwood Middle Fk Escalante East Fk Escalante East Fk Escalante Middle Fk Escalante East Fk Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Grade Gulch Dry Fk Escalante Grade Gulch Cottonwood Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Mud Springs Gulch Cottonwood Dry Fk Escalante Mud Springs Gulch Cottonwood Cottonwood Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Dry Fk Escalante Cottonwood Dry Fk Escalante Cottonwood East Fk Escalante -----------------S18na1 G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G Location F F F F F F G G G F F G F G F G F G F G F G F F F F F F G G G F F F F F F G F F F F F �276 Appendix Table E. Aerial telemetry locations of adult female Euma 1112. Legal description -----------------Date 8-18-86 8-22-86 8-29-86 9- 5-86 9-12-86 9-19-86 9-25-86 10- 5-86 10-10-86 10-24-86 10-31-86 ll- 6-86 ll-14-86 ll-21-86 11-25-86 12- 5-86 12-12-86 12-19-86 1- 3-87 1- 9-87 1-16-87 1-24-87 1-30-87 2- 6-87 2-15-87 2-20-87 2-28-87 3- 6-87 3-13-87 3-21-87 3-28-87 4- 6-87 4-10-87 4-17-87 4-24-87 5- 1-87 5- 8-87 5-15-87 5-22-87 5-30-87 6- 5-87 6-12-87 6-20-87 1/4 N'h 5W SE 5 13 20 30 Nii 30 SE 7 SE 21 SW 19 NE 17 SE 18 NE 29 SW 21 NE 30 NE 30 SW 21 SW 13 SW 10 SW 10 NE 3 SE 10 NE 35 N\..' 13 NE 3 N\..' 22 SW 32 SE 32 NW 35 NW 25 SE 30 SI>" 10 NW 15 SIo: 32 NE 15 NE 16 NE 17 m,36 NW 21 S\,' 15 51> 27 SE 26 SE 6 SE 12 NE 19 SE 20 T 50 50 50 50 50 50 50 50 50 50 50 50 50 50 51 50 50 50 50 51 51 50 51 15S SIN 51 51 51 50 50 51 50 50 50 51 50 50 50 50 49 49 50 50 U.T.M. ----------R 15 15 14 14 14 15 14 15 14 15 14 14 14 14 31 14 14 14 14 14 13 14 13 97W 13\;1 14 14 14 14 14 13 14 14 14 14 14 14 15 15 14 15 14 14 X Y Approx. e1ev (m) 717 714 720 720 720 714 719 712 720 712 722 720 720 723 729 724 724 724 725 726 737 724 734 737 731 726 727 720 724 724 730 725 723 722 727 722 724 715 717 720 719 720 721 4275 4274 4271 4272 4275 4272 4272 4274 4274 4271 4273 4271 4272 4273 4279 4276 4276 4278 4275 4280 4285 4278 4283 4286 4280 4280 4281 4281 4276 4276 4280 4225 4274 4275 4280 4273 4274 4271 4271 4267 4266 4273 4273 2,377 2,560 2,469 2,408 2,341 2,530 2,408 2,499 2,377 2,438 2,256 2,469 2,438 2,347 2,164 2,196 2,134 2,073 2,196 2,196 1,647 2,073 2,103 1,676 2,134 2,164 2,012 2,438 2,164 2,225 2,073 2,196 2,316 2,196 1,920 2,377 2,347 2,591 2,682 2,438 2,438 2,134 2,438 Distance (km) between locations 7.3 3.2 6.4 1.2 3.9 6.8 5.8 7.8 7.8 8.1 10.1 2.4 0.4 3.2 7.4 2.3 0.4 2.4 1.0 4.8 ll.5 rr.o rr.s 5.0 9.5 5.0 2.1 7.1 6.5 0.5 7.5 7.0 1.5 1.8 7.2 8.0 1.5 9.4 2.0 4.7 1.5 6.3 1.4 Rating Major drainalle North Fk Escalante North Fk Escalante Middle Fk Escalante Kelso North Fk Escalante Kelso Kelso North Fk Escalante Kelso Kelso Middle Fk Escalante Kelso Kelso Middle Fk Escalante Escalante Kelso Kelso North Fk Escalante Escalante Escalante Dry Fk Escalante North Fk Escalante Escalante Dry Fk Escalante Tatum Draw North Fk Escalante Escalante North Fk Escalante Escalante Escalante Escalante Escalante Kelso Kelso Escalante Kelso Escalante Kelso Kelso Middle Fk Escalante Middle Fk Escalante Kelso Kelso -----------------Sisna1 G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G G Location F F F F F F G F F G G G G F G F G G F G F G F F F G G G F G G F G F F F F F G F F F G �277 Appendix Table F. Aerial telemetrv locations of male Euma #20. Legal description -----------------Date 8-18-86 8-22-86 8-29-86 9- 5-86 9-12-86 9-19-86 9-26-86 10- 5-86 10-10-86 10-17-86 10-24-86 10-31-86 11-14-86 11-21-86 11-25-86 12- 5-86 12-12-86 12-19-86 1- 9-87 1-24-87 1-30-87 2- 6-87 2-15-87 2-20-87 2-28-87 3- 6-87 3-13-87 3-21-87 3-28-87 4- 6-87 4-10-87 4-17-87 4-24-87 5- 1-87 5- 8-87 5-15-87 5-22-87 5-30-87 6- 5-87 6-12-87 6-20-87 1/4 S T U.T.M. ----------R X Y Approx. elev (m) Distance (km) between locations 49 11.1 22 13 735 4264 2,316 26 49 3.2 13 4262 2,469 737 30 49 2.2 12 2,408 739 4261 49 13 13 4.6 738 4265 2,347 49 13 13 0.8 738 4265 2,377 50 14 1 728 4278 17.1 2,073 22 51 14 4284 6.0 725 2,256 NW 6 50 13 4278 728 2,164 6.0 N.••. ; 3 50 734 4278 13 1,951 5.5 m; 2 49 4269 12 745 2,012 14.7 SII' 19 50 12 739 4273 7.7 1,890 SE 6 49 4268 12 739 2,134 4.9 NW 33 50 743 4270 4.5 12 1,981 NE 25 49 4263 13 738 2,316 9.5 NE 49 16 12 742 4266 6.0 1,951 No location, faint momentary Signal over lower Monitor Mesa SW 10 12 743 50 4276 1,920 10.2 SW 4 49 12 742 4268 8.3 1,951 NE 4270 36 50 13 738 4.4 2,196 NE 4 49 742 4269 12 2,042 5.0 NE 4278 8 50 12 741 1,920 9.0 ]'.'W 2 49 12 745 4269 1,951 9.3 Nil' 35 50 4270 13 735 2,196 9.7 SW 32 50 12 740 4270 1,920 4.5 NE 32 50 12 741 4271 2,012 1.5 N\.,' 14 50 12 745 4276 1,829 6.0 NE 32 50 12 741 4270 2,012 6.3 SE 29 50 12 741 4271 1,859 1.0 m; 4273 26 50 12 745 1,737 3.7 SW 49 4267 12 745 11 2,134 5.8 NE 4276 18 50 12 739 2,073 10.2 NE 32 50 12 741 4271 1,890 5.0 SE 12 742 4272 28 50 2,073 2.0 NE 743 4273 28 50 12 2,042 0.8 NE 12 4274 19 50 739 1,981 4.3 S\\ 20 4273 50 12 740 2.2 2,073 NE 4276 14 50 12 745 1,707 5.7 )';0 location, faint signals over Winter and Monitor Mesas Nl, 50 12 740 4279 5 1,798 6.4 No location, faint Signals over Lower Criswell Ck NE 49 4265 17 12 741 2,164 13.5 NE SE SW SE SE NE NE Rating Major drainage Potter Criswell Moore Criswell Criswell Escalante Palmer Gulch Escalante Dry Fk Escalante Roubideau Monitor Potter Criswell Criswell Roubideau -----------------Signal Location G G F F F G G F G G G G G F F G F F G G G G G G G G G Roubideau Criswell Monitor Criswell Monitor Roubideau Monitor Potter Potter Roubideau Potter Potter Criswell Roubideau Monitor Potter Roubideau Potter Monitor Potter Roubideau G G G G G G F F G G G G G G G G F G G G F F F F F F F G F G G G G G G F F G F F F F G G Cottonwood G F Moore G G G �278 Appendix Table G. Aerial telemetr~ Legal description -----------------Date 8-18-86 8-22-86 8-29-86 9- 5-86 9-12-86 9-19-86 9-25-86 10- 5-86 10-10-86 10-17-86 10-24-86 10-31-86 11- 6-86 11-14-86 11-21-86 11-25-86 12- 5-86 12-12-86 12-19-86 1- 3-87 1- 9-87 1-16-87 1-24-87 1-30-87 2- 6-87 2-15-87 2-20-87 2-28-87 3- 6-87 3-13-87 3-21-87 3-28-87 4- 6-87 4-10-87 4-17-87 4-24-87 5- 1-87 5- 8-87 5-15-87 5-22-87 5-30-87 6- 5-87 6-12-87 6-20-87 1/4 S 51,' 36 S\,' 36 NE 1 NE l6 Sl-.' 26 NE 12 SE 7 Nl.' 2 S\,' 30 SE 3 Sl-.' 2 m; 26 S\\ 34 SW 26 NE 20 SE 31 m; 21 SW 24 NE 20 NE 6 :Nl-.' 4 SE 29 Nl.' 32 SW 13 ]','\" 23 S\,' 16 NE 8 m; 33 NE 31 SW 13 1\..••. 9 m; 32 S\,' 9 S\,' 20 51,' 12 S\,' 34 NE 4 NE 14 N'w 29 NW 6 SE 3 Sl-.' 17 SE 3 NE 10 locations of adult female Euma 112l. !j.T.H. Y Approx. elev (m) Distance (i<m) between locations 4269 4269 4268 4265 4271 4267 4266 4269 4271 4267 4268 4272 4269 4271 4264 4279 4274 4273 4273 4278 4278 4281 4280 4275 4284 4284 4277 4280 4280 4274 4277 4280 4276 4272 4276 4289 4278 4275 4272 4268 4268 4265 4267 4267 2,377 2,377 2,469 2,621 2,652 2,347 2,621 2,377 2,560 2,499 2,377 2,103 2,347 2,256 2,499 2,164 2,134 2,164 2,103 2,225 2,103 2,042 2,103 2,347 1,951 2,012 2,134 2,042 1,890 2,316 2,164 2,073 2,103 2,073 2,256 2,134 2,073 2,012 2,196 2,377 2,225 2,530 2,438 2,408 5.1 0.5 1.1 4.5 5.8 4.8 1.5 4.4 4.8 1.7 0.7 10.5 3.2 2.1 8.0 12.8 5.8 4.6 5.4 5.2 2.3 2.3 0.6 6.5 12.0 2.8 3.0 3.1 2.8 5.8 4.4 3.6 5.2 3.8 4.0 7.0 1.1 7.5 4.5 12.0 6.1 4.8 5.2 0.6 ----------- T R X 50 50 49 49 50 49 49 49 50 49 49 50 50 50 49 51 50 50 50 50 50 51 51 50 51 51 50 51 51 50 50 51 50 50 50 51 50 50 50 49 49 49 49 49 15 15 15 15 15 15 14 15 14 15 15 14 14 14 14 13 13 13 13 13 13 13 13 14 13 13 13 13 13 14 13 13 13 13 14 13 13 14 13 14 14 14 14 14 718 718 719 714 716 719 720 717 719 715 716 725 724 725 722 730 732 737 732 729 732 731 731 727 735 732 731 732 729 728 732 731 732 731 727 733 733 726 730 719 725 721 726 725 Rating Major drainase Middle Fi< Escalante Middle Fi< Escalante Middle Fi< Escalante Middle Fi< Escalante Kelso Middle Fi< Escalante Middle Fi< Escalante Middle Fi< Escalante Middle Fi< Escalante Middle Fi< Escalante Middle Fi< Escalante East Fi< Escalante East Fi< Escalante East Fi< Escalante East Fi< Escalante Escalante Dry Fi< Escalante Monitor Dry Fi< Escalante Tatum D Dry Fi< Escalante Escalante Tatum Escalante Tatum Escalante Dry Fi< Escalante Tatum Ci< Escalante Dry Fk Escalante Dry Fi< Escalante Escalante Dry Fi< Escalante Dry Fi< Escalante Escalante Dry Fi< Escalante Dry Fi< Escalante Escalante Dry Fi< Escalante Middle Fi< Escalante East Fi< Escalante East Fk Escalante East Fk Escalante East Fk Escalante -----------------Sisnal G G G Location G G F G G G G G G G G G G G G F F G G G G G G G G G G G G G G G G G F F G G G G G G G G G G G F F F F F F G F F G F G G F F F F F F F F F F F F F G G G G G G G G G G G G �279 AEEendix Table H. Aerial telemetr~ Legal description -----------------Date 8-18-86 8-22-86 8-29-86 9- 5-86 9-12-86 9-19-86 9-25-86 10- 5-86 10-10-86 10-17-86 10-24-86 10-30-86 11- 6-86 11-14-86 11-21-86 11-25-86 12- 5-86 12-12-86 12-19-86 1- 3-87 1- 9-87 1-16-87 1-24-87 1-30-87 2- 6-87 2-15-87 2-20-87 2-28-87 3- 6-87 3-13-87 3-21-87 3-28-87 4- 6-87 4-10-87 4-17-87 4-24-87 5- 1-87 5- 8-87 5-15-87 5-22-87 5-30-87 6- 5-87 6-12-87 6-20-87 1/4 SE SI< NW NE S T locations of adult male Euma 1122. U.T.M. ----------R X Y Approx. elev (m) Distance (km) between locations Rating Major draina!:e -----------------Sisnal Location 46 5 8 257 4.9 4239 2,134 Uncompahgre R G 13 46 8 264 4235 7.4 2,316 G Deer 24 46 8 263 4234 0.8 G 2,225 Deer 30 46 7 266 4233 2.0 G 2,408 Martin m,' 6 46 7 265 4239 G 6.6 2,591 Burro NW 27 47 8 260 4242 G 2,286 4.9 Uncompahgre NE 46 29 258 4232 8 G 2,134 10.2 Uncompahgre SIo.' 24 48 8 264 4252 G 2,469 Dry Cedar Ck 21.5 m,' 14 47 262 4246 G 8 2,164 6.8 Beaton NE 46 5 8 258 4239 2,012 G 15.0 Uncompahgre SI< 22 48 260 4252 G 8 2,316 11.5 Dry Cedar N'W 30 46 4332 7 265 G 2,225 22.8 Martin Ck NW 33 46 259 4231 G 8 2,256 6.2 Uncompahgre SO; 19 46 265 4233 7 2,377 G 6.1 Tommy SE 4238 1 46 8 264 2,347 4.5 G Burro NW 46 19 7 265 4233 2,438 3.5 G Tommy SE 28 48 4251 8 259 2,196 7.1 G Uncompahgre R SW 48 22 8 261 4252 2,316 2.2 Dry Cedar G NW 46 4234 19 7 265 2,438 19.2 Tommy G m; 25 47 8 263 4242 2,377 8.9 Brook G SE 4 46 8 260 4238 2,225 5.2 Chaffee G SE 30 47 4242 7 265 2,621 7.1 Billy G 24 SE 47 4242 265 2,560 1.7 F 8 Billy SE 4236 7 46 266 2,560 7 6.8 Deer G NE 25 47 264 4242 8 2,438 6.0 G Billy SE 12 46 4236 8 264 2,560 5.5 G Burro NW 25 47 264 4242 2,499 6.2 8 Billy G SW 4254 15 48 8 260 2,103 12.5 Dry Cedar Ck G 51< 16 48 4254 8 259 2,012 1.9 Dry Cedar Ck G N'W 12 46 264 4237 8 2,377 Burro G 17.0 NE 4235 G 15 46 8 261 2,134 3.1 Burro NE 23 48 4254 8 263 2,316 20.0 Dry Cedar Ck G NIo.' 25 47 263 4242 G 8 2,377 11.5 Billy Ck No location, fair signals heard over Montrose, 1 mi. S of Jet. Hwys. 50 & 347, Lower Spring Ck NE 7 49 265 4267 2,652 G 7 25.0 Cedar Ck 4244 NE 23 47 8 263 2,377 20.0 Onion Ck G 51< 25 4272 G 50 8 262 2,469 28.5 Gunnison SI~ 4 46 4238 G 8 259 2,073 Uncompahgre R 17.1 No location, faint signals north end of home range SI< 27 4242 G 47 8 260 2,256 4.0 Billy 4234 SE 46 266 2,530 G 19 7 10.2 Martin NE 4242 28 4i 8 260 2,286 G 10.5 Uncompahgre G SE 34 48 261 4249 8.0 8 2,196 Beaton 51< 21 268 4233 46 7 2,682 G 14.5 Taylor G G G F F F G F G G F G G G G G G G F G G F F G F F F G F F F F F F F F G F F F F F �280 AEpendix Table I. Aerial telemetry locations of adult male Euma 1126. Legal description U.T.M. -----------------Date 8-22-86 8-29-86 9- 5-86 9-12-86 9-19-86 9-26-86 10- 5-86 10-10-86 10-17-86 10-24-86 10-31-86 ll- 6-86 ll-14-86 11-21-86 11-25-86 12- 5-86 12-12-86 1- 9-87 1-24-87 2- 6-87 2-15-87 2-18-87 1/4 T S ----------- R X Rating Major draina~e -----------------1/4 -----------------Si~nal Location F G F F G F G F F G G F G F F G F F F F G P F G F F G G G G G F F G Aerial telemetry locations of adult female Euma 1128. Legal description 8-29-86 9- 5-86 9-12-86 9-19-86 9-26-86 10- 5-86 10-10-86 10-17-86 10-24-86 ll- 6-86 11-14-86 ll-21-86 ll-25-86 Distance (km) between locations 5E 24 ISS 99\1 725 1,707 38.0 4290 Little Dominquez NE 30 155 100\1 708 2,560 17.4 4288 Big Dominquez Good momentary signal, Upper Escalante Good momentary signal, wend Winter Mesa NW 15S 98••.• 731 23.5 Gunnison Gulch 15 4292 1,798 511 35 5lN 131.' 735 4279 1,829 13.0 Dry Fk Escalante NW 15 SON 1411 724 4275 2,256 12.5 Escalante No location, good on-frequency signal east of Gateway NW 24 SON 1511 718 4273 Kelso 2,256 6.7 511 30 50 14 4271 720 2,499 Middle Fk Escalante 2.8 5E 15 50 14 724 4274 2,256 5.3 Escalante NW 29 49 12 4262 741 2,316 7.9 Traver NW 18 4282 63.4 19 51 681 2,134 Larsen 4292 9 15 103 680 2,196 9.4 North Fk West Ck 51' 14 NE 4285 51 2,103 46.1 Palmer Gulch 141' 726 NE 4 1411 723 4268 16.4 49 2,499 East Fk Escalante 5E 4272 2,560 13.6 19 50 15 710 North Fk Escalante 4271 50 13 734 2,256 21.0 Mud Spring Gulch 51' 27 34 4251 2,042 NE 48 10 241 37.0 Spring 744 4265 2,134 NE 22 49 12 12.5 Roubideau No location, strong momentary signal heard over foothills about 1 mi. II. of Colona Reportedly killed in 528, T32S, R22E, N. side Blue Mtn, on Indian Ck, Utah AEEendix Table J. Date Y Approx. elev (m) S T R U.T.H. ----------X Y Approx. elev (m) Distance (km) between locations Rating Major draina~e Intermittent, off-frequency signals, south rim Western Unaweep Canyon 4296 32.0 NW 2,408 Granite Ck 30 145 104,1 676 NE 4307 2,560 9.9 McKenzie 25 135 104.••. 677 4308 SE 678 2,225 1.8 Briar Canyon 19 135 104" 4313 2,196 7.0 Coates Ck 3 13S 104;;' 672 5" NW 4312 2,196 1.0 Coates Ck 10 135 1041<1 672 4308 2,103 4.8 Spring Ck SE 20 135 104\01 670 Nil' 29 4307 2,256 1.1 Spring Ck 13S 104W 669 4309 2,134 1.8 Spring Ck NE 20 135 104\01 670 No location, strong off-frequency signal over upper Dominquez drainages 4307 2,377 3.6 Hill NE 27 13S 1041' 673 135 104\01 671 4310 2,286 3.8 Cook NE 16 Mortality signal at CDW, NW Regional Office, killed 11-22-86 by J. Brent ---------------5i~na1 G G G G G Location F F F F G G F G G F F G F G G �231 AEEendix Table K. Aerial telemetry Legal description -----------------Date 8-18-86 8-22-86 8-29-86 9- 5-86 9-12-86 9-19-86 9-25-86 10- 5-86 10-10-86 10-17-86 10-24-86 10-31-86 11- 6-86 11-14-86 11-21-86 11-25-86 12- 5-86 12-12-86 12-19-86 1- 3-87 1- 9-87 1-16-87 1-24-87 1-30-87 2- 6-87 2-15-87 2-20-87 2-28-87 3- 6-87 3-13-87 3-21-87 3-28-87 4- 6-87 4-10-87 4-17-87 4-24-87 5- 1-87 5- 8-87 5-15-87 5-22-87 5-3(i-87 6- 5-87 6-12-87 6-20-87 locations of adult female Euma 1132 •• U.T.M. ----------- S T R X Y Approx. elev (m) 17 8 18 17 S•• 1 NE 8 NE 8 NE 9 NW 10 NE 10 SI< 1 SW 21 NW 17 NW 29 SE 30 NW 19 NW 20 NE 18 NE 20 SE 30 SE 19 SE 30 NE 29 S," 17 SW 17 S\.I 20 m,' 30 S;;' 17 SE 17 NW 20 NE 20 NW 30 NE 29 SW 5 Nli 5 NE 7 SW 28 NE 5 SE 30 S;;' 17 NW 32 SI< 17 NW 27 SW 24 45 45 45 45 45 45 45 45 45 45 45 46 46 46 46 46 46 46 46 46 46 46 46 46 46 46 46 46 46 46 46 46 46 45 45 46 46 45 46 46 46 46 46 46 9 9 9 9 9 9 9 9 9 9 9 8 8 8 8 8 8 8 8 8 8 10 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 9 9 247 247 246 247 253 247 247 249 250 251 253 258 257 257 256 255 257 256 258 256 256 256 258 257 257 257 255 257 257 257 258 256 258 257 257 256 259 258 256 257 257 257 250 254 4226 4227 4226 4227 4229 4228 4228 4228 4228 4228 4229 4234 4235 4232 4232 4234 4234 4236 4234 4232 4232 4232 4232 4235 4235 4233 4232 4235 4235 4234 4234 4232 4232 4229 4229 4238 4232 4229 4233 4235 4230 4235 4233 4234 2,469 2,438 2,438 2,438 2,499 2,621 2,652 2,469 2,591 2,499 2,408 2,196 2,134 2,286 2,347 2,286 2,256 2,225 2,134 2,286 2,347 2,316 2,042 2,196 2,196 2,256 2,377 2,196 2,196 2,225 2,164 2,347 2,196 2,316 2,408 2,225 2,225 2,316 2,286 2,196 2,347 2,042 2,499 2,377 1/4 ,,-.; NE NE NE Distance (kIn) between locations Major drainaEe 13.4 1.5 1.7 0.4 6.7 6.9 0.2 1.7 1.4 0.8 2.6 7.0 2.4 3.2 0.9 1.9 1.3 1.7 2.0 3.3 1.8 1.6 1.5 2.2 0.5 2.0 2.1 3.3 0.5 0.5 1.3 3.0 1.5 3.9 0.5 8.6 6.3 2.7 4.4 1.7 5.3 4.7 7.2 3.5 Pleasant Valley Pleasant Valley Pleasant Valley Pleasant Valley Uncompahgre R Pleasant Valley Pleasant Valley Pleasant Valley Pleasant Valley Pleasant Valley Pleasant Valley Uncompahgre R Fisher Uncompahgre Uncompahgre Fisher Fisher Fisher Uncompahgre Uncompahgre Uncompahgre Uncompahgre Uncompahgre Fisher Fisher Uncompahgre Uncompahgre Fisher Uncompahgre Fisher Uncompahgre Fisher Uncompahgre Dallas Ck Dallas Ck McKenzie Uncompahgre Dallas Ck Fisher Fisher Uncompahgre Fisher McKenzie Fisher Rating -----------------Signal Ck Ck Ck Ck G G G G G G G G G G G G G G G G G G G G Location G G G G G F F G G G G G G G F F F G G G F F G G F G G F G G G G G G G G G G G G G G G G G G G F G F G G G G G G G F F F F G F F F F G F �202 Aj2j2endix Table L. Aerial telemetr~ locations u.r.u. Legal description -----------------Date 8-29-86 9- 5-86 9-12-86 9-19-86 9-25-86 10- 5-86 10-10-86 10-17-86 10-24-86 10-31-86 11- 6-86 1/4 T S NE liE N\" 24 30 28 N\\ 29 SIo: 20 Nil 29 NE 6 SE 29 N',; 32 SIo: 29 NE 9 Killed by Aj2j2endix Table H. 46 46 46 46 46 46 47 46 46 46 46 hunter ----------R 8 7 7 7 7 7 7 7 7 7 8 Aerial telemetr~ 8-18-86 8-22-86 8-29-86 9- 5-86 9-12-86 9-19-86 9-25-86 10- 5-86 10-10-86 10-17-86 10-24-86 10-31-86 11- 6-86 11-14-86 11-21-86 11-25-86 12- 5-86 12-12-86 12-19-86 1- 3-87 1- 9-87 1-24-87 1-30-87 2- 6-87 2-15-87 2-20-87 2-28-87 3- 6-87 3-13-87 3-21-87 3-28-87 4- 6-87 4-10-87 4-17-87 4-24-87 5- 1-87 5- 8-87 5-15-87 5-22-87 5-30-87 6- 5-87 6-12-87 6-20-87 S T Approx. elev (m) Distance (km) between locations 264 4234 2,256 .,4232 2,347 265 2,530 268 4232 266 4232 2,591 266 4233 2,560 264 4234 2,256 266 2,499 4249 267 4231 2,438 267 4231 2,286 266 4230 2,347 4237 259 2,134 11-6-86 in GMU 65 on Uncompahgre locations U.T.H. -------- -----------------1/4 Y X Legal description Date of subadult female Euma (133. R 46 SE 19 9 NW 46 5 9 46 N\" 17 9 Faint, intermittent 47 NE 35 10 t-.'1,.,' 47 16 11 47 SE 25 10 S;; 28 46 9 N',; 34 46 9 NE 47 22 9 47 SE 12 10 47 10 S\" 24 S',; 19 47 9 NE 47 13 10 S\,' 19 47 9 S;; 19 47 9 4 SE 47 10 NE 24 47 10 24 NE 47 10 SI; 4 47 9 NE 24 47 10 51: 17 47 9 24 NE 47 10 Nh' 30 47 9 47 5E 13 10 Nh' 27 47 9 47 9 5\" 19 47 SW 7 9 SW 7 47 9 NE 24 47 10 Nj,' 7 47 9 Nj,' 13 47 10 5E 12 47 10 SW 13 47 10 Nil 23 47 10 NE 23 47 10 47 NE 19 9 47 10 SE 13 NE 47 13 10 47 9 S'" 19 NE 26 47 10 NW 47 10 35 47 5E 26 10 X 246 247 247 signal, 243 754 245 248 250 251 245 244 246 245 245 245 240 245 245 249 244 247 245 246 245 250 245 245 245 245 245 244 245 244 242 243 246 245 245 246 243 242 243 Y 21. 7 2.2 4.3 1.5 0.4 2.6 16.5 17.2 1.1 0.6 9.3 River Rating Major draina8e Tommy Martin Lou Taylor Martin Tommy Beaton Lou Lou Lou Uncompahgre -----------------Si8nal Location G G G G F G G G G F G G G F F F G G G G G G of subadult female Euma #34. Approx. elev (m) Distance (km) between locations Rating Major draina8e 4234 2,621 2.7 East Fk Horsefly 4239 2,377 4.2 Horsefly 2,499 3.6 Cottonwood 4236 East Horsefly and N. of Upper Spring Ck 2,438 4241 6.3 Dolores 4245 2,591 14.7 East Fk Dry Ck 4242 2,377 16.1 Dolores 4232 2,591 10.1 McKenzie 4231 2,560 2.0 Fisher 4244 2,073 13.6 Horsefly Happy 4246 2,164 5.1 4243 2,256 3.5 Happy 4244 2,256 1.5 Dolores 4246 2,164 2.6 Happy 4237 2,256 2.5 Dolores 4243 2,256 0.1 Dolores 4248 2,225 2.1 Spring 4244 2,256 Happy 2.7 4244 2,286 Happy 0.2 4248 2,012 6.5 Dolores 4244 2,225 Happy 6.5 4245 2,164 3.2 Dolores Happy 4245 2,225 2.5 4243 2,316 1.8 Dolores 4245 Happy 2,196 2.8 4242 2,196 6.7 Horsefly 4243 2,256 5.2 Dolores 4247 2,134 3.8 Happy Happy 4247 2,103 0.2 Happy 4244 2,256 2.5 3.0 Happy 4247 2,103 Happy 4246 2,286 2.0 1.0 Happy 4247 2,164 2.0 Happy 4245 2,256 Happy 4244 2,408 2.0 Happy 4244 2,286 0.9 Dolores 3.3 4243 2,196 Happy 4246 2,225 2.0 Happy 0.2 4245 2,196 2.6 Dolores 4243 2,316 2.4 Happy 4243 2,377 Happy 4241 2,469 1.8 4242 2,469 0.3 Happy -----------------Sisnal G G G Location F F F G G G G G G G F G F G G G G G G G G G G G G G G G G G G G G G F G G G F G F F G G G G G G G G G G G G G G G G G F G G G G F F F G F G G G G G F G F G G G G �203 AEEendix Table N. Aerial telemetrl locations of Juvenile male Euma #35. Legal description Date 9- 5-86 9-12-86 9-19-86 9-26-86 10- 5-86 10-10-86 10-17-86 10-24-86 11- 6-86 11-14-86 11-19-86 1/4 T 5 R y Distance (km) between locations Rating Major drainage -----------------Signal Location Aerial telemetr~ locations of subadult female Euma #36. Legal description -----------------8-18-86 8-22-86 8-29-86 9- 5-86 'l-12-86 9-19-86 9-25-86 L,- 5-86 10-10-86 10-17-86 10-24-86 10-31-86 11- 6-86 11-14-86 11-21-86 11-25-86 1- 3-87 1- 9-87_ 3- 6-87 3-13-87 3-21-87 3-28-87 4- 6-87 4-10-87 4-17-87 4-24-87 5- 1-87 5- 8-87 5-15-87 5-22-87 5-30-87 6- 5-87 6-12-87 6-20-87 X Approx. elev (m) G 5W 49 G 32 19 673 4259 2,256 10.5 Roc Ck G 5E 49 4259 31 19 673 G 2,225 0.3 Roc Ck G SE 49 31 672 4259 0.2 G 19 2,225 Roc Ck G SW 49 32 673 4259 G 19 0.2 Roc Ck 2,196 G G SW 32 49 19 673 4259 0.5 2,012 Roc Ck NW G 19 671 4261 G 2,316 2.7 ~O 49 Roc Ck No location, mortality signals vicinity of Sinbad Ridge and Pace Lake, Utah G SW 20 145 102W 689 4298 mortality, G 2,714 38.5 West Ck F m< 31 145 102W mortality, G 688 4295 2,134 2.9 West Ck NE G 29 normal, G 145 10211 689 4297 West Ck 2,682 2.4 NE 29 4298 Animal dead. Recovered 689 2,745 0.5 West Ck 145 102" Cause death radiocollar. unknown but dried carcass suggested 1+ mos. from death to discovery. AEEendix Table O. Date U.T.M. -------- ------------------ 1/4 S T R U.T.M. -------X Y Approx. elev (m) Distance (km) between locations NW 48 16 259 4255 8 2,103 17.0 NW 22 48 4253 8 261 2,256 2.8 NW 247 17 46 4236 9 2,499 22.0 NW 46 17 247 4236 9 2,499 0.3 NW 34 47 4241 9 250 2,256 5.4 NE 48 261 4253 15 8 2,286 17.5 NE 48 4255 15 8 261 2,256 0.4 NW 47 263 4246 13 8 2,316 9.6 47 SW 12 8 263 4246 2,408 0.5 NE 48 4254 19 7 266 2,499 8.8 NE 48 15 8 261 4235 2,256 5.0 NW 48 4253 22 8 260 2,164 2.1 NW 47 260 4248 3 8 2,134 5.2 NW 48 262 26 8 4251 2,438 3.5 NE 4252 28 48 8 260 2,103 2.1 NW 47 261 4241 2,134 35 8 11.4 No location; faint, sporadic signal Lower Cimaroon Drainage No location; faint, sporadic signal Lower Cimaroon Drainage SW 48 260 4254 15 8 2,134 14.0 NE 48 28 8 260 4252 2,196 2.4 SW 47 262 4243 23 8 2,286 9.0 NE 48 4255 15 8 261 2,286 12.0 NW 4 46 261 8 4237 2,377 18.0 SE 47 261 4241 27 8 2,196 6.3 SW 48 262 4256 11 8 2,286 13.5 NE 266 2,745 48 7 4252 5.6 30 NW 47 8 261 4243 2,377 10.0 23 NE 47 7.2 8 261 4249 2,256 3 SE 48 263 4254 6.0 14 8 2,438 NW 48 262 4250 2,316 4.2 35 8 NW 262 4250 48 2,256 0.4 35 8 NW 260 4242 2,286 47 8 8.5 27 NW 260 4242 47 8 2,316 0.7 27 261 4244 2,408 1.2 SE 47 8 22 Rating Major drainage -----------------Signal Dry Cedar Dry Cedar Cottonwood Cottonwood Horsefly Dry Cedar Dry Cedar Beaton Beaton Dry Cedar Dry Cedar Dry Cedar Beaton Beaton Uncompahgre Billy Ck G G Dry Cedar Ck Dry Cedar Ck Onion Dry Cedar Ck Cow Ck Billy Ck Dry Cedar Ck Hairpin Ck Brook Beaton Dry Cedar Ck Beaton Beaton Uncompahgre Billy Ck Onion G G G G G G G G Location G F F F F G F F G G G F F F G G G G G G G G G G F F G G G G G G F F F F F G G G G G G G F G G G G G G F F F �234 AEEendix Table P. Aerial te1emetr~ locations Legal description -----------------Date 12- 9-86 12-12-86 12-19-86 1- 3-87 1- 9-87 1-16-87 1-24-87 1-30-87 2- 6-87 2-15-87 2-20-87 2-28-87 3- 6-87 3-13-87 3-21-87 3-28-87 4- 6-87 4-10-87 4-17-87 4-24-87 5- 1-87 5- 8-87 5-15-87 5-22-87 5-30-87 6- 5-87 6-12-87 6-20-87 1/4 NE NE NW SI< NW NE SE SI< NE SE SW NE NE SW NW SW SW SW SE NE SE NW NE No; SE NE NW NE Date 12-12-86 12-19-86 1- 3-87 1- 9-87 1-16-87 1-24-87 1-30-87 2- 6-8-Z 2-15-87 2-20-87 2-28-87 3- 6-87 3-13-87 3-21-87 3-28-87 4- 6-87 4-10-87 4-17-87 4-24-87 5- 1-87 5- 8-87 5-15-87 5-22-87 5-30-87 6- 5-87 6-12-87 6-20-87 ----------- S T R X 11 11 12 33 5 32 .7 3-3 5 33 33 11 5 33 17 33 34 34 13 9 33 18 35 36 26 24 24 24 48N 48 48 49 48 49 48 49 48 49 49 48 48 49 48 49 49 49 48 48 49 48 48 48 48 48 48 48 121< 12 12 11 11 11 11 11 11 11 11 12 11 11 11 11 11 11 12 11 12 11 12 12 12 12 12 12 748 748 749 752 752 751 751 752 753 453 753 748 752 752 752 751 753 753 750 754 743 750 748 749 748 749 749 749 A1212endix Table Q. Aerial te1emetr~ Y 4258 4258 4257 4260 4259 4262 4256 4261 4260 4260 4260 4257 4260 4261 4256 4260 4260 4260 4255 4257 4260 4255 4251 4251 4251 4254 4254 4254 locations Legal description U.T.M. ------------------ ---------- 1/4 S N" 13 3 4 24 28 9 34 21 21 13 33 17 34 22 9 35 18 13 27 7 22 34 34 34 3 34 3 NW SI; SE NI.' SE NW NW SE NW NE SE sw NE sw SW NE SW NW SE NE SW SE NE NE SE NE T 47 48 46 47 49 47 47 49 49 47 49 49 50 48 48 49 48 48 48 47 49 48 48 48 48 48 47 . of subadult U.T.M. R X 10 12 9 10 11 10 9 11 11 10 11 11 12 11 11 12 11 12 11 10 11 11 11 11 11 11 11 244 745 248 244 751 240 250 752 752 244 753 750 743 756 754 745 751 749 755 761 754 755 756 756 756 756 756 Y 4247 4259 4239 4244 4263 4247 4241 4265 4264 4246 4261 4265 4270 4254 4257 4260 4256 4254 4252 4248 4265 4250 4250 4251 4250 4250 4250 female Euma {i37. Approx. e1ev (m) Distance (Iem) between locations 2,347 2,347 2,256 2,196 2,225 2,225 2,347 2,196 2,196 2,164 2,134 2,286 2,196 2,103 2,196 2,196 2,164 2,256 2,469 1,920 2,196 2,408 2,621 2,560 2,591 2,499 2,438 2,560 capture 0.0 0.9 4.0 0.8 2.4 5.2 4.2 1.7 0.8 0.4 6.0 5.5 0.8 4.9 4.6 1.8 0.4 6.3 5.3 11.3 8.5 5.1 0.5 0.7 3.3 0.6 0.3 Rating ------------------ Major drainage Cushman Cushman Piney Piney Piney Cushman West Fk Piney Dry Ck Piney Piney Cushman Piney Piney West Fk Piney Dry Ck Dry Ck Piney East Fk Terrible West Fk West Fk West Fk West Fk West Fk West Fk West Fk Sisnal Location G G G G G G Dry Ck G F F F G G G G G F G G G G Dry Ck G G G G G G G G G G G G G Dry Ck G G Dry Dry Dry Dry Dry Dry Dry G G Ck Ck Ck Ck Ck Ck Ck F F F F F F F F G G G F G G G G G of adult male 12uma 1138. Approx. elev (m) 2,225 2,042 2,377 2,225 2,134 2,286 2,164 2,012 1,981 2,225 2,042 2,042 1,981 2,286 2,042 2,347 2,256 2,499 2,316 2,438 1,981 2,438 2,377 2,347 2,408 2,377 2,438 Distance (kill) between locations capture 25.5 20.9 6.6 28.5 21.0 7.7 16.0 0.8 12.0 22.1 4.8 8.3 19.8 3.1 9.4 7.4 2.6 6.8 8.4 17.2 14.3 0.8 0.5 1.1 0.7 1.3 Rating ------------------ Major drainase Happy Roubideau Horsefly Happy Cushman Spring Horsefly Cushman Cushman Happy Piney Coal bank Roubideau Coal Ck East Fk Dry Roubideau West Fk Dry Piney East Fk Dry Spring Dry Ck East Fk Dry East Fk Dry East Fk Dry East Fk Dry East Fk Dry East Fk Dry Sisna1 G G G G G G F G G G G G G G G G Ck Ck Ck Ck Ck Ck Ck Ck Ck Location G G G G G G G G G G G G G G F G G F F G F G F F G F F F F F G G G F G G �285 AEEendix Table R. Aerial telemetr~ locations of subadult male Euma 1139. Legal description Date 12-19-86 1- 3-87 1- 9-87 1-16-87 1-24-87 1-30-87 2- 6-87 2-15-87 2-20-87 2-28-87 3- 6-87 3-13-87 3-21-87 4- 6-87 1/4 S T SE 30 16 NE 35 Slol 13 NE 3 SE 16 SW 32 Slol 16 tm 35 SE 26 SE 30 10 S" NE 15 NO< 35 51 50 51 51 50 51 l5S 51 51 51 51 50 50 51 NO< AEEendix Table S. ----------R 13 14 14 13 14 13 97101 13 14 14 14 14 14 14 X Y 729 723 426 737 725 732 737 732 726 726 720 724 725 725 4281 4275 4280 4285 4278 4284 4286 4284 4280 4281 4280 4276 4275 4280 -----------------1- 3-87 1- 9-87 1-16-87 1-24-87 1-30-87 2- 6-87 2-15-87 2-20-87 2-28-87 3- 6-87 3-13-87 3-21-87 3-28-87 4- 6-87 4-10-87 4-17-87 4-24-87 5- 1-87 5- 8-87 5-15-87 5-22-87 5-30-87 6- 5-87 6-12-87 1/4 T R X 8 29 NE 16 NE 5 SE 20 SE 2 NO< 33 SW 1 51.' 5 5E 5 NW 11 5E 30 5E 5 5E 6 NE 9 5E 1 5E 30 5E 16 NE 27 24 5E 5E 4 NE 5 SW 2 NO< 31 46 46 46 45 46 45 46 45 46 45 45 46 45 45 45 45 46 46 46 46 45 45 45 46 8 8 9 8 8 9 8 9 8 8 9 8 8 8 9 9 8 9 9 9 8 8 9 8 257 257 249 257 258 252 258 253 257 258 252 256 257 256 249 254 256 250 251 254 257 258 252 255 SW AEEendix Table T. Date 1- 8-87 1- 9-87 1-16-87 1,768 2,196 2,134 1,676 2,073 1,951 1,676 2,012 2,164 2,134 2,438 2,164 2,012 2,196 capture 9.0 6.5 11.5 14.0 8.0 5.3 5.9 7.5 1.2 6.8 6.5 1.0 4.5 U.T.M. Rating Major drainage Escalante Kelso Escalante Dry Fk Escalante N. Fk Escalante Escalante Dry Fk Esclante Escalante North Fk Escalante Escalante North Fk Escalante Kelso Escalante North Fk Escalante Y Approx. elev (m) Distance (km) between locations 4236 4233 4236 4229 4233 4228 4231 4229 4238 4229 4228 4231 4229 4229 4228 4228 4232 4235 4233 4234 4229 4229 4228 4231 2,196 2,286 2,438 2,438 2,196 2,499 2,256 2,438 2,134 2,316 2,469 2,377 2,256 2,316 2,469 2,316 2,316 2,499 2,499 2,256 2,286 2,316 2,499 2,408 capture 3.9 8.0 10.0 4.0 7.4 6.5 5.9 10.3 9.3 6.1 5.6 3.1 1.2 7.1 4.5 4.0 7.5 2.4 8.5 6.0 0.9 6.3 4.5 ---------- S t;'1I Distance (km) between locations -----------------Location Signal G G G G G G G G G G G G G G G F G F G F G F G G F G Aerial telemetry locations of adult male Euma 1/40. Legal description Date Approx. elev (m) U.T.M. ------------------ Rating Major draina!le Uncompahgre Uncompahgre Busted Boiler Draw Dallas Ck Uncompahgre Pleasant Valley Ck Uncompahgre Pleasant Valley McKenzie Dallas Pleasant Valley Uncompahgre Pleasant Valley Dallas Pleasant Valley Ck Pleasant Valley Ck Uncompahgre R Busted Boiler Draw McKenzie Fisher Dallas Ck Dallas Ck Pleasant Valley Ck Uncompahgre R --------------Si!lnal Location F G G G G G F G G F G G G G G G G G G G G G G G G G G G G G G G F G G F F F G G G G G G F Aerial telemetry locations of subadult male Euma 1/41. Legal description -----------1/4 R S T U.T.M. x y Approx. elev (m) Distance (km) between locations capture NE 12 50 15 718 4276 2,164 0.0 NE 12 50 15 718 4276 2,164 NE 12 0.0 50 15 718 4276 2,164 ·.Cause of death unknown; carcass intact where we left it. skull on 2-9-87 Rating Major drainage North North North Recovered Signal Location Fk Escalante Fk Escalante mortality Fk Escalante mortality radiocollar and removed �286 Appendix Table U. Aerial telemetr~ locations Legal description ---------- -----------------Date 1-10-87 1-16-87 1-30-87 2- 6-87 2-15-87 3- 6-87 3-13-87 3-21-87 3-28-87 4- 6-87 4-10-87 4-17-87 4-24-87 AEEendix Date 1-16-87 1-24-87 1-30-87 2- 2-87 1/4 T 5 R v. Aerial te1emetr~ ------------------ ----------- 5 T R y X 13101 737 NE 13 50 13\,' 735 SW 26 50 NE 13 50 13 737 Radioco11ar recovered Aerial te1emetr~ Legal description -----------------1-17-87 1-24-87 1-30-87 2- 6-87 2-15-87 2-20-87 2-28-87 3-13-87 3-21-87 3-28-87 4- 6-87 4-10-87 4-17-87 4-24-87 5- 1-87 5- 8-87 5-15-87 5-22-87 5-30-87 6- 5-87 6-12-87 6-20-87 locations U.T.M. AEEendix Table W. Date 4270 4268 4276 4259 4256 4238 4243 4244 4245 4255 4273 4287 signal Legal description 1/4 1/4 5 T NW NE 17 9 34 16 16 10 35 31 19 35 17 35 21 32 7 8 30 20 24 21 33 16 IS SO 15 51 51 51 ISS ISS ISS 145 155 ISS 51 155 51 51 155 51 15 51 51 51 NW NE NW SW SE SW SW NW NE NW SE NE SE NW SE SW NE NE SE NE 4276 4272 4276 locations 2,073 2,256 X 708 16 742 12 711 100 713 15 712 15 714 15 100W 714 99l.' 716 99101 716 99W 722 100W 709 100W 713 713 15 719 99 710 15 711 15 100W 707 701 16 706 101 694 17 704 16 704 16 y 4291 4277 4287 4284 4284 4285 4286 4286 4289 4296 4291 4287 4282 4286 4285 4285 4288 4282 4289 4282 4278 4284 Distance (lem) between locations capture 2.7 2,042 2,073 2,164 2,408 2,377 2,225 2,042 1,737 2,196 5W of Crawford 27.0 3.8 28.2 6.2 1.8 1.3 12.0 18.0 19.5 Rating ------------------ Major draina~e Location Si~nal Cottonwood Cottonwood Potter Coal Ck Coal Ck McKenzie Brook Onion Onion Dry Cedar Gulch Gunnison R Smith Fk G P G G G G G G G G G G P F G G G F F F F G of adult female Euma 1143. Approx. e1ev (m) Distance (lem) between locations 1,920 2,042 1,920 capture 4.3 4.4 Rating ------------------ Major drainage Cottonwood Cottonwood Cottonwood Signal mortality, mortality, Location G G F F of adult male Euma 1144. U.T.M. ----------R Approx. elev (m) y X 50 734 51.' 34 13 4 49 Sw 13 732 SW 9 50 12 742 48 SE 2 11 757 NW 48 760 18 10 46 SW 5 257 8 SE 22 47 261 8 SW 14 47 262 8 SE 47 15 261 8 SE 48 16 259 8 NE 27 50 259 8 NW 31 155 91W 273 No location; weak, momentary Table of subadult male Euma #42. U.T.M. Approx. elev (m) Distance (km) between locations 2,377 capture 2,438 2,438 2,530 2,377 2,499 2,256 2,438 1,981 2,286 2,499 2,499 2,012 2,499 2,652 2,591 2,438 2,499 2,560 2,652 2,499 5.2 3.0 0.7 2.1 1.1 2.5 3.1 9.4 14.3 6.2 5.3 13.8 8.6 0.3 4.0 8.5 8.8 13.6 10.3 6.2 Rating Major draina~e Big Dominquez Potter Big Dominquez Little Dominquez Little Dominquez Little Dominquez Little Dominquez Little Dominquez Rough Draw Big Dominquez Rocky Pitch Gulch Big Dominquez Little Dominquez Little Dominquez Keith Big Dominquez Big Dominquez Smith Ck Gill Ck Cow Ck Big Dominquez Smith Ck -----------------Si~nal P G G G G G G G G G G G G G G G G G G G G Location P F F F F F G G F G F G F F G G G F F F G �287 A:eEendix Table X. Date 1-20-87 1-24-87 1-30-87 2-15-87 2-20-87 2-28-87 3- 6-87 3-13-87 3-28-87 4- 6-87 4-10-87 4-17-87 4-24-87 5~ 1-87 5- 8-87 5-15-87 5-22-87 5-30-87 6- 5-87 6-12-87 6-20-87 Aerial te1emet!1 locations of adult female Euma 1145. Legal description --------------1/4 S T R SE SE S;; SE m; NE NE NE ~.",. m,' NE NE SE N\< N;; m; SW NE NE SIo: J',w A:e:eendixTable 1. 20 17 21 9 22 32 32 32 26 2 32 33 33 5 5 32 17 19 6 21 20 50 50 50 50 50 50 50 50 50 49 50 50 50 49 49 50 49 49 48 49 49 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 U.T.H. ------Y X Legal description 1-22-87 1-24-87 1-30-87 2- 6-87 2-15-87 2-20-87 2-28-87 3- 6-87 3-13-87 3-21-87 3-28-87 4- 6-87 4-10-87 4-17-87 4-24-87 5- 1-87 5- 8-87 5-15-87 5-22-87 5-30-87 6- 5-87 6-12-87 6-20-87 1/4 SW S;; Distance (km) between locations 2,044 1,981 1,951 1,768 2,134 2,073 2,042 2,012 1,707 1,981 2,042 1,890 1,981 2,073 2,103 2,042 2,225 2,196 2,438 2,196 2,225 capture 2.0 1.6 3.0 2.6 3.9 0.2 0.2 4.5 3.6 4.5 1.6 0.5 3.0 0.8 1.8 6.0 1.0 5.2 4.0 1.8 Rating Major draina!1!e Potter Potter Potter Potter Roubideau Potter Potter Potter Criswell Roubideau Potter Criswell Criswell Potter Criswell Potter Criswell Criswell Wright Roubideau Hoore -----------------Sisna1 Location G G G G G G G G G G G G G G G G G G G G F F F F F G G F G F F F F F F G G G F F Aerial te1emetrr locations of adult female Euma 1146. -----------------Date 4273 4275 4273 4276 4274 4271 4270 4271 4273 4269 4271 4271 4270 4269 4269 4271 4265 4264 4259 4264 4264 740 741 742 743 744 741 741 741 745 745 741 743 743 740 741 740 740 739 742 742 740 Approx. elev (m) 5 19 1~ 25 N'" KE 29 510: 26 5;; 1.3 SE 16 NI-: 11 NE 20 SE 26 SO: 29 SE S 5;; 28 No: 14 NE 21 SI:: 16 No: 19 m; 30 NE 30 SW 27 SW 34 NE 28 NW 20 T 15S 155 155 15S 15S 155 15S 51 51 51 155 51 ISS SlN 5lN lSS 51 51 51 51 ISS 155 ISS U.T.H. Y Approx. e1ev (m) Distance (km) between locations 4289 4289 4289 4288 4287 4291 4291 4286 4284 4280 4288 4285 4288 4284 4282 4291 4283 4281 4281 4280 4285 4288 4290 2,440 2,438 2,499 2,316 2,256 2,073 1,890 2,196 2,042 2,164 2,012 2,164 2,316 2,196 2,408 2,164 2,347 2,438 2,438 2,499 2,621 2,377 2,408 capture 0.0 1.9 3.9 0.9 5.5 5.5 7.5 6.5 6.5 8.5 6.0 3.9 5.3 3.3 11.0 8.3 1.1 1.3 5.7 6.2 2.6 3.0 --------R 99W 9910: 100W 99W 99W 99\1 98\01 14 13 14 98 14 99W lSW lSIi 99\01 14 14 14 15 100W 100W 100W X 717 717 715 719 720 724 730 725 731 726 728 722 720 716 714 721 719 719 720 714 711 711 709 Rating Major drainage Little Dominquez Little Dominquez Little Dominquez Little Dominquez Little Dominquez Little Dominquez Little Dominquez West Palmer Gulch Escalante Escalante West Palmer Gulch Little Dominquez Little Dominquez Little Dominquez Little Dominquez Wildhorse Rose Rose Open Draw Red Ck Big Dominquez Big Dominquez Big Dominquez -----------------Signal G G G G G G G G G G G G G G G G G G G G G G Location F F F F F F F F G F F F F F F F G F G G F F �2dd AEl:endix Table 2. Aerial telemetr~ locations Legal description -----------------Date 1-29-87 1-30-87 2- 6-87 2-15-87 2-20-87 2-28-87 3- 6-87 3-13-87 3-21-87 3-28-87 4- 6-87 4-24-87 5- 1-87 5- 8-87 5;"'22-87 5-30-87 6- 5-87 6-12-87 1/4 5 51.' 15 SW 15 SW 15 m; 35 SE 14 NW 13 SW 14 5101 14 SW 14 SE 14 SE 14 SW 30 NE 34 SE 21 SE 11 NE 7 SE 15 SE 11 ----------- T R X 155 155 155 155 155 51 155 15S 155 15S 155 50N 5lN 155 145 14S 145 14S 100101 100101 1001.' 100101 99\1 13 97 97 97 97 97 14 17 102101 1041.' 103W 103101 103101 712 712 712 714 724 736 742 742 742 743 743 720 696 691 675 678 683 684 Aerial telemetr~ A;e;eendix Table 21- Legal description 2-17-87 2-20-87 2-28-87 3- 6-87 3-13-87 3-21-87 3-28-87 4- 6-87 4-10-87 4-24-87 5- 1-87 5- 8-87 5-15-87 5-22-87 5-30-87 6- 5-87 6-12-87 6-20-87 1/4 S\~ SE SE 5 T Y 4291 4291 ,4291 4286 4291 4285 4291 4291 4291 4291 4291 4271 4279 4288 4301 4302 4300 4302 locations Approx. elev (m) Distance (kIn) between locations 2,256 2,250 2,256 2,560 2,073 1,829 1,525 1,525 1,525 1,488 1,494 2,499 2,408 2,775 2,316 2,652 2,682 2,560 capture 0.1 0.1 5.3 11.4 14.0 8.3 0 0 0.6 0.3 31.0 13.0 9.9 20.6 2.9 5.4 2.5 ----------R 17 155 1001.' 13 155 99W 14 15 51 NW 25 155 98•• Not located, strong SE 5 47 8 NW 30 49 11 SE 18 49 11 N,," 4 47 8 NE 48 6 11 5,,' 32 48 10 SE 27 11 48 N\,' 24 47 11 SE 47 17 10 NW 26 47 11 Nw 47 35 10 NE 35 47 11 SW .10 47 11 X Y Rating Major drainalle Big Dominquez Big Dominquez Big Dominquez Big Dominquez Little Dominquez Dry Fk Escalante Gunnison Gunnison Gunnison Gunnison Gunnison Middle Fk Escalante Indian Yel10wjacket Canyon Granite Ck Granite Ck North Fk West Ck North Fk West Ck -----------------Sillnal Location G G G G G G G G G G G G G G G G G G F F F F G G G G G F F F F G F F of subadult ·male Euma 1150. U.T-M. -----------------Date of subadult male ;euma 1147. U.T.M. Approx. elev (m) Distance (kIn) between locations Rating Major drainalle 708 4291 2,349 capture Big Dominquez 4287 717 2,316 9.6 Little Dominquez 4284 2,256 8.4 725 Palmer Gulch 734 4289 1,890 11.5 Escalante on-frequency signal from Dry Cedar Ck area 258 4248 2,012 60.2 Beaton 749 4263 2,164 36.0 Coal bank 749 4265 2,103 2.3 Sandy Wash 259 4248 2,073 36.0 Beaton 4259 2,225 34.2 751 Piney 238 4250 2,286 14.0 Devinny 4251 2,316 756 10.0 East Fk Dry Ck 4244 2,469 4.6 West Fk Spring 759 4245 238 2,438 4.3 East Fk Spring 4243 757 2,652 5.9 west Fk Spring 242 4241 2,408 9.3 Happy 9.4 4241 2,745 Middle Fk Spring Ck 758 4247 2,469 6.0 East Fk Dry Ck 755 -----------------Sillnal Location G G G F F F G G G G G G G G G G G G G G F F F F F F F F G G F F �239 AEEendix Table 22. Aerial telemetrl locations Legal description -----------------Date 3-16-87 3-21-87 3-21-87 AEEendix 1/4 5 T ----------R Tal.l~ 23. Y Aerial telemetrl locations U.T.M. ------- -----------4-30-87 5- 1-87 5- 8-87 5-15-87 5-22-87 5-30-87 6- 5-87 6-12-87 6-20-87 X Approx. elev (m) Distance (kIn) between locations 48 NE 9 11 754 4257 1,891 capture 9 NE 48 11 754 4257 1,891 Cause of death unknown; recovered skull and.radiocol1ar. Carcass undisturbed--died in position we left it in. Legal description Date c~ juvenile male Euma U51. U.T.H. 1/4 5 T R X Nt SW St 19 27 24 16 19 19 19 8 33 47 47 46 47 47 47 47 46 47 9 9 9 9 9 9 9 9 9 246 250 254 249 246 245 245 248 249 5101 Nt SW SW St 51< 4244 4241 4234 4245 4244 4243 4243 4237 4240 East Fk Dry Ck East Fk Dry Ck -----------------Location Signal G mortality of adult male Ellt:.:: 1152. Dd s t at.r.e Y Rating Major drainage Rating (Ian) Approx. e1ev (m) locations 2,256 2,225 2,256 2,256 2,225 2,256 2,286 2,408 2,256 capture 4.5 8.5 12.2 2.3 1.6 0.7 6.1 3.0 be tveen Major drainage Dolores Horsefly Fisher Duckett D Dolores Dolores Dolores Horsefly Horsefly -----------------Signal G G G G G G G G Location G G F F G G F G �'-:lU o 10 I 20 I KM Figure 1. Puma study area GMU 62, Uncompahgre subunits. Plateau, showing �VV.VLQUV "~~""'''''''6.j. ~J- """_ •• _ Wildlife Research Report July 1987 . JOB PROGRESS REPOR1 State of Colorado Project No. Mammals 01-03-048 (FW 26 p) ~~--------~------~-- 2 Research Work Plan No. 7 Mountain Goat and Mountain Sheep Investigations Job No. 1 Improve Matching Harvest Regulations with Available Mountain Goat and Sheep Populations and Improve Methods for Obtaining Better Population Estimates Period Covered: July 1, 1986 - June 30, 1987 Author: D. F. Reed ABSTRACT First during this segment, 4 program narratives were tentatively developed in accordance with 'the overall objectives of improving mountain goat and mountain sheep census data and of evaluating census methodology. These 4 program narratives were Work Plan 7, Jobs 1-4 as follows: Job 1 - Improve matching harvest regulations with available mountain goat populations through better counts and better analyses of count data (Appendix A), Job 2 - Improve methods for obtaining better population estimates of mountain goats through mark-recapture data (Appendix B), Job 3 - Improve matching harvest regulations with available mountain sheep populations through better counts and better analyses of count data (Appendix C), and Job 4 - Improve methods for obtaining better population estimates of mvuntain sheep by combining mark-recapture and telemetry data (Appendix D). Secondly, after review of possible microcomputer simulations (resighting probabilities and aggregation or clumping scenarios) and budgetary constraints, Jobs 1-3 were dropped, and Job 4 modified. Job 4 was redesigned as Work Plan LA, Job 5 and titled: �Experimental Tests of the Mark-Recapture Method to Estimate Mountain Sheep Numbers (Appendix E). Selection of an appropriate study area for Job 5 is judged to be critical to its successful completion. Such selection will not be made until after the end of this segment. ) Wildlie Researcher �293 APPENDIX A PROGRAM NARRATIVE State Colorado --~~~~--------------------- Project No. Work Plan No. 01-03-048 - 11700 7 ------~--------------- Job No. A. 1 Mammals 2 Research Mountain Goat Investigations Improve matching harvest regulations with available mountain goat populations through better counts and better analyses of count data NEED Problem Mountain goats (Oreamnos americanus) are not indigenous to Colorado. They were first introduced into the Collegiate Range in 1948 and subsequently into other areas including Mt. Evans, San Juan Mountains, Gore Range, and Marcellina Mountain (Denney 1977). These areas were often within the historical, if not present, range of native mountain sheep (Ovis canadensis canadensis). The problem is that as mountain goat populations increase, they may expand their range into areas managed specifically for mountain sheep. The Division's policy in regard to areas managed specifically for mountain sheep and areas where mountain goat populations will be permitted or considered for translocations was addressed at a recent Biologists' Coordination Meeting (28 August 1986). The concern has been and continues to be that at some threshold density of one or both of these species, "competition" may occur resulting in a competitive advantage for mountain goats (Bassow 1985). Consequently, it is imperative that the distribution and abundance of mountain goats be determined, updated periodically, and published in a format that is readily available for review. Literature Review Although there is not a substantial volume of literature concerning techniques of censusing mountain goats, some workers have reported on ground (Fox 1984, Nichols 1980) and aerial counts (Fox 1977, 1984; Smith and Bovee 1984). The results of such censuses have yielded variable results. Hence, development of mountain goat inventory techniques is a high priority research need (Eastman 1977). Nichols (1980) indicated that counts made under good conditions, in early to midsummer, included about 90% of the goats found from the ground or helicopter. Although he suggested that the 2 methods, aerial and ground counts, were of equal accuracy, he had only limited samples for comparison. Conversely, Fox (1984) found that aerial counts averaged 32% lower than those of ground counts. Houston et al. (1986) estimated the efficiency of aerial census at 66% by making aerial counts before and after a reduction in the �mountain goat population. This lower efficiency in aerial counts may be due in part to the reported propensity of mountain goats to seek escape terrain (cliffs and/or rock crevices, overhangs, etc.) when alarmed by low-flying aircraft. Most of the counts of mountain goats reviewed here were purported as total censuses where there is no question of variance or confidence limits in the samplng sense (Overton and Davis 1969). Sample counts, in addition to being described by variance and confident limits, have the advantages of being less labor-intensive and time-consuming (Gill et al. 1983), reducing counting some individuals more than once and missing others, being conducted over longer periods of time, and of disturbing the population less (Caughley 1977:26). Since wild ungulates tend to be clumped, i.e. their densities vary from place to place within given population ranges, some investigators have found it advantageous to use stratified sampling vs. simple random sampling. Siniff and Skoog (1964) in censusing caribou (Rangifer tarandus) doubled the precision of their census estimates by use of stratified sampling (Caughley 1977:27). They also used equal-area sampling units since the line-transect method was found impractical in rugged terrain. Houston et ale (1986) used census strata and a design involving block count sampling because of variable mountain goat densities and precipitous terrain, respectively. B. OBJECTIVES The overall objective of this study is: To improve the Division's ability to match harvest regulations with available mountain goat populations through better counts and better analyses of count data. Specific subobjectives are: C. 1. To improve estimates of the distribution of mountain goats in the populations or subpopulations statewide. 2. To improve estimates of the number of mountain goats in the populations or subpopulations statewide. 3. To prepare a publication delineating the brief history and current distribution and abundance of mountain goats statewide. EXPECTED RESULTS OR BENEFITS Better counts and better analyses of count data should improve the Division's ability to match harvest regulations with available mountain goat populations. �295 D. APPROACH 1. Methods: Reconnaissance - During the first year of study, ground reconnaissance of each of the populations or subpopulations will be conducted during July when mountain goat aggregations have become relatively stable and the animals are conspicious against the low, green vegetation of alpine areas. These efforts are necessary in order to establish their distribution and estimates of various _ densities for purposes of stratification during the sampling phase of the study (2nd.and 3rd year). Reconnaissance will be conducted in each of the areas where mountain goats are known to occur. However, it is anticipated that the major effort will be concentrated within 2-3 areas that will be selected for the sampling phase where densities range from high to low and access is least difficult. The following areas are likely candidates: a. Mt. Evans (Central Region) - Population estimates have ranged from 118-168 and a highway provides access to a major portion of the alpine area. Annual ground counts have been conducted in the area since 1978 and numerous ground counts have been made throughout the year in the east-central portion of the area from 1980-85. Preliminary tasks include reconnaissance of peripheral areas and determination of densities per unit area. b. Gore Range (Northwest Region) - Population estimates have ranged from 100-111 (Thompson 1980) and access into the area is limited to trails and nonmotorized travel because of wilderness designation. Both aerial and ground counts were conducted from 1977-79. Summer distribution was reported as covering 75 km2 in the alpine areas of Otter Creek, Cataract Creek, Piney River, Black Creek, Slate Creek, and Boulder Creek with the greatest portion of the population inhabiting Dora Mtn. (Thompson 1981). Preliminary tasks include reconnaissance of entire range in order to update both distribution and density. c. Mt. Antero-Mt. Shavano (Southeast Region) - A minimum population value for the area including Mt. Antero, Mt. Mamma, Mt. Shavano, and Mt. Aetna was 110 in 1978 (West 1978). The access into the area is provided by the road from Maysville to North Fork Reservoir campground. Preliminary tasks include reconnaissance of the entire range in order to update both distribution and density. The other areas in which ground reconnaissance will be conducted are listed by Division Region in which they occur: �296 Central Grey and Torres Southeast Northwest Mt. Princeton Jones-Sheep Mtn.l Mt. HarvardMt. Yale Holy Cross Ten-mile Southwest Needles Tincup-Monarch Raggeds Marcellina Mtn. The candidate areas mayor may not stand, depending upon 1) the results of the reconnaissance of all areas and 2) the monetary and time constraints of this study. Sampled counts - During the sampling phase, a block count sampling method similar to that of Houston et al. (1986) will be used except that aerial counts will be replaced with ground counts. Census blocks will be delineated on 1:62,000 scale contour maps. Blocks will be designed to include as many slopes and aspects as possible. Conspicuous terrain features (ridges, rock outcrops, forested areas) will be used as block boundaries. Further details of the design including the number and location of census routes and the number of censuses to be conducted will no~ be determined until after reconnaissance. 2. Analysis: the population totals will be calculated using Jolly's (1969) method for unequal sized sample units. Schedule Activity Conduct ground reconnaissance, select areas for sampled counts, and delineate census blocks. Fiscal Year 1987-88 Conduct sampled counts. 1988-89 Same as 1988-89. 1989-90 Data analysis and publication 1990-91 Personnel D. F. Reed Principal Investigator No. positions undetermined Volunteers lThis area is considered to be covered by on-going CSU graduate studies. �297 Estimated Cost Person days $24,461.00 (01) Personal Services (21) Operating Supplies and Services 2,539.00 (28) Travel Expenses 1,000.00 (31) Capital Expenditures (Equipment) TOTAL E. 140 Amount $28,000.00 GEOGRAPHIC LOCATION Various areas have been mentioned in the Approach section and are subject to change. F. RELATED FEDERAL PROJECT Colorado Federal Aid Project FW 26P. LITERATURE CITED Bassow, S. 1985. Potential competition between the Mt. Evans populations of Rocky Mountain bighorn sheep (Ovis canadensis) and mountain goats (Oreamnos americanus). B.A. Thesis, Princeton Univ., Princeton, NJ. 77pp. Caugh1ey, G. 1977. Analysis of vertebrate populations. Sons, NY. 234pp. J. Wiley and Denney, R. N. 1977. Status and management of mountain goats in Colorado. Pages 29-36 in W. Samuel and W. G. MacGregor, eds. Proc. First Int'l. Mtn. Goat Symp. 243pp. Eastman, D. S. 1977. Research needs for mountain goat management. Pages 160-168 in W. Samuel and W. G. MacGregor, eds. Proc. First Int'l. Mtn. Goat Sym~ 243pp. Fox, J. L. 1977. Summer mountain goat activity and habitat preference in coastal Alaska as a basis for the assessment of survey techniques. Pages 190-199 in W. Samuel and W. G. MacGregor, eds. Proc. First Int'l. Mtn. Goat Symp. 243pp. Fox, J. L. 1984. Population density of mountain goats in southeast Alaska. Pages 51-60 in M. Hoefs, ed. Proc. Fourth Biennial Symp. N. Wild Sheep and Goat Counc.--513pp. �293 Gill, R. B., L. H. Carpenter, and D. C. Bowden. 1983. Monitoring large animal populations: the Colorado experience. Trans. N. Amer. Wi1d1. Conf. 48:330-341. Houston, D. B., B. B. Moorhead, and R. W. Olson. 1986. An aerial census of mountain goats in the Olympic mountain range Washington. NW Sci. 60(2):131-136. Jolly, G. M. 1969. Sampling methods for aerial censuses of wildlife populations. E. Afr. Agric. For. J. 34(Specia1Issue):49-49. Nichols, L. 1980. Aerial census and classification of mountain goats in Alaska. Pages 523-589 in W. O. Hickey, ed. Proc. Biennial Symp. N. Wild Sheep and Goat Counc. 405pp. Overton, W. S., and D. E. Davis. 1969. Estimating the numbers of animals in wildlife populations. Pages 403-455 in R. H. Giles, ed., Wildlife Management Techniques, 3rd Ed. The Wildlife Society, Washington, DC. 623pp. Siniff, D. B., and R. O. Skoog. stratified random sampling. 1964. Aerial censusing of caribou using J. Wi1dl. Manage. 28:391-401. Smith, C. A., and K. T. Bovee. 1984. A mark-recapture census and density estimate for a coastal mountain goat population. Pages 487-498 in M. Hoefs, ed. Proc. Fourth Biennial Symp. N. Wild Sheep and Goat Counc. 5l3pp. Thompson, R. W. 1980. Population dynamics, habitat utilization, recreational impacts and trapping of introduced Rocky Mountain goats in the Eagles Nest Wilderness Area, Colorado. Pages 459-462 in W. O. Hickey, ed. Proc. Biennial Symp. N. Wild Sheep and Goat Counc. 405pp. 1981. Ecology of Rocky Mountain goats introduced in the Eagles Nest wilderness, Colorado and some factors of geographic variation in the lambing season of bighorn sheep. M.S. Thesis, Univ. of Wyoming, Laramie. 359pp. West, R. L. 1978. The Mt. Shavano mountain goat study. l8pp. Mimeogr. Colo. State Univ. �299 APPENDIX B PROGP~ State Colorado ------------------------------- Project No. Work Plan No. 01-03-048 - 11700 7 ----------------------- Job No. A. NAP~TIVE 2 Mammals 2 Research Mountain Goat Investigations Improve methods for obtaining better population estimates of mountain goats through mark-recapture data. NEED Problem Mountain goats (Oreamnos americanus) are not indigenous to Colorado. They were first introduced into the Collegiate Range in 1948 and subsequently into other areas including Mt. Evans, San Juan Mountains, Gore Range, and Marcellina Mountain (Denney 1977). These areas were often within the historical, if not present, range of native mountain sheep (Ovis canadensis canadensis). The problem is that as mountain goat populations increase, they may expand their range into areas managed specifically for mountain sheep. The Division's policy in regard to areas managed specifically for mountain sheep and areas where mountain goat populations will be permitted or considered for translocations was addressed at a recent Biologists' Coordination Meeting (28 August 1986). The concern has been and continues to be that at some threshold density of one or both of these species, "competition" may occur resulting in a competitive advantage for mountain goats (Bassow 1985). Consequently, it is important that methods for obtaining better population estimates of mountain goats be determined. Literature Review There is a substantial volume of literature concerning mark-recapture theory, models, and various statistical treatments (see reviews by Caughley 1977; Otis et al. 1978; Overton and Davis 1969; Seber 1973, 1982; White et al. 1982). The amount of lite=ature is greatly reduced when conSidering only the "size of the popUlation" versus 8 other properties that can be investigated by some form of the mark-recapture technique (Caugh1ey 1977:133) and when limiting the application to wild ungulates (Strandgaard 1967, Woolf 1973, Rice and Harder 1977, Bartmann et al. 1987). Aspects of mark-recapture techniques most applicable to a study of free-ranging wild ungulates include the importance of experimental design, of equal catchability of all individuals, marked and unmarked, where truth of the assumption can be tested in a pilot experiment or have such a test built into the experimental design (Caughley 1977:134), and of �3UO adequate sample sizes including the proportion of marked animals in the population. Concerning the latter aspect, large proportions (>67% Strandgaard 1967, >45% - Bartmann et ale 1987) of small populations need to be marked to generate reliable population estimates (White et ale 1982). Approaches to the study of population size in free-ranging wild ungulates might include 1) the Petersen estimate (Petersen 1896, Lincoln 1930) or the simplest mark-recapture procedure which calls for marking on one occasion and recording proportion of marked animals captured (or "sighted" since actual capture is not necessary if marks can be recognized [Eberhardt 1969]) on a second occasion (Caughley 1977:141), 2) Schumacher's method which calls for marking on several occasions and is estimated from the rate at which the proportion of marked individuals rise as progressively more are marked (Caughley 1977:145), 3) methods involving the assumption of "closure" - the simplest appropriate model of 8 possibilities (models Mo' Mt, Mb, Mh, Mtb, Mth, Mbh, and Mtbh) (Otis et ale 1978, White et ale 1982),4) methods involving the assumption of "open" populations where the process of birth, death, and migration are allowed to operate--essential elements are incorporated in the classic Jolly-Seber model (Seber 1978:196-232, Seber 1982:196-232), and 5) procedures involving maximum likelihood estimates (MLE) used in combining repeated Lincoln-Petersen estimates (Bartmann et ale 1987). B. OBJECTIVES The overall objective of this study is: To improve the Division's methods for obtaining better population estimates of mountain goats through mark-recapture data. Specific subobjectives can be stated as hypotheses to be tested: 1. 2. (to be determined - see Analysis section) 3. C. EXPECTED RESULTS OR BENEFITS Methods for obtaining better population estimates should improve the Division's ability to match harvest regulations with the number of available mountain goats. D. APPROACH 1. Methods: Counts - During each of 3 years of study, 5 ground counts will be conducted in the Mt. Evans area during July and August. One of the counts will be the previously established annual ground count usually conducted during the last week of July (Yamashita per. comm.). A schedule of the counts for 3 years is as follows: �301 1987 1988 1989 9 Jul 6 Jul 6 Jul 16 Jul 13 Jul 13 Jul 23 Jul 20 Jul 20 Jul 30 Jul 27 Jul 27 Jul 6 Aug 3 Aug 3 Aug It is estimated that a minimum of 26 observers, minimum of 2 per route, will be required to cover 13 routes (previously established by Ya~ashita 1985) during each of the count days. It is further es~imated that at least 2 different groups of 26 observers will be needed for the 5 count days, i.e. the first group will conduct the first 3 counts and the last count, and the second group (used in previous years and organized by 5. Yamashita) will conduct the 4th count (30 Jul, 27 Jul, and 27 Jul for 1987, 1988, and 1989, respectively). For the first gFOUp, successful G4 mountain goat and 53 and 53A mountain sheep permittees for last year and the new permittees drawn for the current year, selected Division, U5F5, U5F&W, USNPS, and BLM employees, selected area biology teachers, selected members of the Colorado Chapter of the Wildlife Society, the Colorado Wildlife Federation, Volunteers for Outdoor Colorado, Mountain Goat Society, and Bighorn Sheep Society, and selected individuals will be invited to participate. Responses from those invited will be used in scheduling and in selecting team leaders. Team leaders will be responsible for their group, for assuring that marked animals are identified, and for reporting the number and location of marked and unmarked animals sighted. Marking - Mountain goats will be trapped and marked with individually identifiable (unique) collars at 3 locations. They will be handled by previously established methods (Reed 1981, 1982). The trapping and marking will be done during a relatively short period, between the time adult females return from kidding areas with their neonates and the date of the first count (approximately 2 weeks), after which no new marks will be added to the population for the year. In addition to these animals, previously marked animals (Reed 1986) will be utilized in the mark-recapture procedure. This can be done only if the old marks can be reliably placed in the population at the time of the counts. Hence, a complete survey will be conducted in all areas to identify old marks still in the population. This survey will be conducted during the same 2 weeks that marking is in process and must also be complete before the first count. An attempt will be made to mark approximately 50% of an estimated 140 animals (30 Jul 1986 ground count of 123 [Yamashit~ per. comm.] plus at least 17 missed) or 70 animals in 1987. It is estimated that about 20 old marks will be in the population and hence, about 50 new marks will need to be entered into the population. During the subsequent years of 1988 and 1989, an attempt will be made to mark approximately 60 and 70~, respectively. �302 2. Analysis: Mark-recapture analyses for purposes of estimating population size will follow models whose assumptions can best be met. Models assuming demographic closure (closed populations) and models allowing the demographic closure assumption to be relaxed (open population) (White et al. 1982:48-77, 180-186) are considered in relation to some estimated practical limitations of the study. If closure can be assumed based on the relatively short period between marking and recapturing (sighting), model Mt (Otis et al. 1978:24-28, White et al. 1982:51-55) could possibly be used. However, it is estimated that model Mt is not applicable for this study for the same reason that it was not applicable for a study of mark-recapture estimates of confined mule deer (Bartmann et al. 1987). At this juncture in the formation of this study, it is suggested that several procedures and/or several proportions of marked animals be tested with analyses following those of Bartmann et al. (1987) and White and Garrott (1987). Tentatively, several hypotheses for testing procedures and proportions of marked animals are: Those involving procedures to combine repeated Lincoln-Petersen estimates: 1. simple arthmetic mean and median yield different efficiencies (confidence intervals) 2. median and joint hypergeometric MLE yield different efficiencies (confidence intervals) 3. simple arthmetic mean and joint hypergeometric MLE yield different efficiencies (confidence intervals) those involving proportions of marked animals in a joint hypergeometric MLE procedure to combine repeated Lincoln-Petersen estimates: 1. the proportion of marked animals of 50% and 60% yield different efficiencies (confidence intervals) 2. the proportion of marked animals of 60% and 70% yield different efficiencies (confidence intervals). Replication of counts (n = 5) and years (n = 3) provide data points for comparing these procedures and proportions of marked animals. Schedule Activity Fiscal Year Trap and mark, conduct ground counts 1987-88 Same as 1987-88 1988-89 Same as 1988-89 1989-90 Data analysis and publication 1990-91 �303 Personnel D. F. Reed Principal Investigator No. positions undetermined Volunteers Estimated Cost Person Days (01) Personal Services (02) Operating Supplies and Services (28) Travel Expenses (31) Capital Expenditures (Equipment) 70 $12,230.50 1,269.50 500.00 $14,000.00 TOTAL E. Amount GEOGRAPHIC LOCATION Part of the Mt. Evans area has been described previously (Reed 1981). In addition, Yamashita (1985) has briefly described contiguous areas to be covered on the various census routes. F. RELATED FEDERAL PROJECT Colorado Federal Aid Project FW26P. LITERATURE CITED Bartmann, R. M., G. C. White, L. N. Carpenter, and R. A. Garrott. 1987. Aerial mark-recapture estimates of confined mule deer in pinyon-juniper woodland. J. Wildl. Manage. 51:41-46. Bassow, S. 1985. Potential competition between the Mt. Evans populations of Rocky Mountain bighorn sheep (Ovis canadensis) and mountain goats (Oreamnos americanus). B.A. Thesis, Princeton Univ., Princeton, NJ. 77pp. Caughley, G. 1977. NY. 234pp. Analysis of vertebrate populations. J. Wiley and Sons, Denney, R. N. 1977. Status and management of mountain goats in Colorado. Pages 29-36 in W. Samuel and W. G. MacGregor, eds. Proc. First Int'l. Mtn. Goat Symp. 243pp. Eberhardt, L. L. 1969. Population estimates from recapture frequencies. Wildl. Manage. 33:28-39. J. �304 Lincoln, F. C. 1930. Calculating waterfowl abundances on the basis of banding returns. U.S. Dep. Agric. Circ. 118. 4pp. Otis, D. L., K. P. Burnham, G. C. White, and D. R. Anderson. 1978. Statistical inference from capture data on closed animal populations. Wildl. Monogr. 62:1-135. Overton, W. S., and D. E. Davis. 1969. Estimating the numbers of animals in wildlife populations. Pages 403-455 in R. H. Giles, ed., Wildlife Management Techniques, 3rd Ed. The Wildl. Soc., Washington, DC. 623pp. Petersen, C. G. J. 1896. The yearly immigration of plaice into the Limfjord from the German Sea. Rep. Dan. BioI. Stn. 1895 6:1-77. Reed, D. F. 1981. Rocky mountain goat ecology study. Res. Rep. July, Part 2:209-222. Colo. Div. Wild1. Game. 1982. Rocky Mountain goat-bighorn sheep competition study. Div. of Wildl. Game Res. Rep. July:79-l28. Colo. 1986. Seasonal habitat selection and activity of sympatric mountain goat and bighorn sheep populations. Colo. Div. of Wildl. Game Res. Rep. July Part 2:205-216. Rice, W. R., and J. D. Harder. 1977. Application of multiple aerial sampling to a mark-recapture census of white-tailed deer. J. Wildl. Manage. 41:197-206. Seber, G. A. F. parameters. 1973. The estimation of animal abundance and related Hafner Press, NY. 506pp. 1982. The estimation of animal abundance and related parameters. ed. MacMillan Publishing Co., NY. 654pp. 2nd Strandgaard, H. 1967. Reliability of the Petersen method tested on a roedeer population. J. Wi1dl. Manage. 31:643-651. White, G. C., D. R. Ander~on, K. P. Burnham, and D. L. Otis. 1982. Capturerecapture and removal methods for sampling closed populations. Los Alamos National Laboratory. LA-8787-NERP. Los Alamos, NM. 235pp. , and R. A. Garrott. 1987. Analysis of biotelemetry data - a primer. ----~Co10. State Univ., Unpubl. Draft. 225pp. Woolf, A. 1973. Population dynamics and remote censusing of a large, captive white-tailed deer herd. Ph.D. Thesis, Cornell Univ., Ithaca, NY. l68pp. Yamashita, S. 9pp. 1985. Mt. Evans goat, sheep and elk survey 1985. Mimeogr. �305 APPENDIX C PROGRAM NARRATIVE State Colorado ----~~----------------------- Project No. Work Plan No. 01-03-048 - 11700 7 ------~---------------- Job No. A. 3 Mammals 2 Research Mountain Goat Investigations Improve matching harvest regulations with available mountain sheep populations through better counts and better analyses of count data NEED Problem Mountain sheep (Ovis candensis canadensis) are indigenous to Colorado. They reached North America and probably Colorado during the late Pleistocene. As with essentially all wildlife ungulates with fragmentary fossil records, little is known about their early population levels. Historical accounts of their abundance were not recorded until around 1922 when Seton (1929) estimated that there were about 8,000 mountain sheep in Colorado. The Division's Comprehensive Management Plan (Division of Wildlife 1977) indicated a statewide population of about 3,000 for the base year of 1983. More important, however, is that the plan called for increasing the population to over twice this number in the next 15 years. The projected population curve continues a relatively steep increase through 1993. Concomitant with this planned increase is the consistent highest ranking by each Region of the number one problem: need for· additional inventory. If results of the management program for increasing distribution and abundance of mountain sheep by trapping and transplanting is to known, better counts and analysis of count data will be necessary. Literature Review Traditional procedures used in censusing mountain sheep have been ground (Geist 1971:64-66, Hudson 1982, Shannon et a1. 1975) and aerial counts (Hudson 1982) where the latter may be the most effective method (Simmons and Hansen 1980). Denney (1976) indicated that aerial mountain sheep counts had all but been abandoned in Colorado, due to inconsistent data, difficulties of observation and classification, hazardous flying conditions in rough terrain, vagaries of weather, and budgetary constraints. Irby et a1. (1987) used ground counts and found that none of the 7 indices tested performed well on a single count basis. They further indicated that if intensive management is desired, winter ranges should be observed for 3-5 years prior to initiation of a sampling scheme. Census units most typically have been delineated by terrain characteristics. Hudson (1982) suggested that December is the optimal time to census mountain sheep in �306 most parts of their range. This coincides with the rut and when snow cover improves visibility of the animals. Total counts rather than sample counts are most commonly used. When total counts are used, there is no question of variance or confidence limits in the sampling sense (Overton and Davis 1969). Sample counts in addition to being described by variance and confidence limits, have the advantages of being less labor-intensive and time-consuming (Gill et al. 1983), reducing counting some individuals more than once and missing others, being conducted over longer periods of time, and of distrubing the population less (Caughley 1977:26). Since wild ungulates tend to be clumped, i.e. their densities vary from place to place within given population ranges, some investigators have found it advantageous to use stratified sampling vs. simple random sampling. Siniff and Skoog (1964) in censusing caribou (Rangifer tarandus) doubled the precision of their census estimates by use of stratified sampling (Caughley 1977:27). They also used equal-area sampling units since the line-transect method was found impractical in rugged terrain. Houston et al. (1986) used census strata and a design involving block count sampling because of variable mountain goat densities and precipitous terrain, respectively. B. OBJECTIVES The overall objective of this study is: To improve the Division's ability to match harvest regulations with available mountain sheep populations through better counts and better analyses of count data. Specific subojectives are: C. 1. To improve estimates of the distribution of mountain sheep in the populations or subpopulations statewide. 2. To improve estimates of the number of mountain sheep in the populations or subpopulations statewide. 3. To prepare a publication delineating history and current distribution and abundance of mountain sheep statewide. EXPECTED RESULTS OR BENEFITS Better counts and better analyses of count data should improve the Division's ability to match harvest regulations with available mountain sheep populations. D. APPROACH �307 1. Methods: Reconnaissance - During the first year of the study, ground and aerial reconnaissance of selected populations or subpopulations will be conducted during August-September and November-December, respectively. The purpose of ground reconnaissance during August-September is primarily for area familiarization. Aerial (helicopter) reconnaissance during November-December will utilize animal's aggregations on rutting ranges and snow cover for improving their visibility. These efforts are necessary in order to establish distribution and estimates of various densities for purposes of stratification during the sampling phase of the study (2nd and 3rd year). Aerial or ground reconnaissance will not be conducted in each of the areas where mountain sheep are known to occur. Time and manpower constraints simply will not allow. It is anticipated that the major effort will be concentrated within 2-3 areas that will be selected for the sampling phase where densities range from high to low and aerial counts are least hazardous. The following areas are possible candidates: a. Mt. Evans (Central Region) - Population estimates have ranged from 103-126 and a highway provides access to a major portion of the alpine area. Annual ground counts have been conducted in the area since 1978, and numerous ground counts have been made throughout the year in the e~~t-central portion of the area from 1980-85. Preliminary tasks include ground reconnaissance of peripheral areas and determination of densities per unit area. b. Sangre de Cristo range (Southeast Region) - The population has been approximated at 300-400 (Wakelyn 1984). Access is provided to Crestone campground and by adjacent trails (N. end range). The range includes extremely rugged, precipitous, and rocky alpine and subalpine forest. Winter ranges and migration routes are apparently not well known. Preliminary tasks include reconnaissance of entire range in order to determine both distribution and density. c. Trickle Mtn. (Southwest Region) - A minimum population count for the area was 122 with estimates ranging from 165-175 for 1974 (Shepherd 1975). The access to the area is provided by Highway 114. Preliminary tasks include reconnaissance of the entire range in order to update both distribution and density. Other areas in which aerial and/or ground reconnaissance may be conducted are listed by Division Region in which they occur: �303 Central Northwest Georgetown Clear Creek South P1atte4 Maroon Be11sl Gore2 Never Summers3 Redstone Colorado NM5 Battlement Mesa Cline top Mesa Snowmass Southeast Buffalo Peaksl Arkansas River2 Beaver Creek3 Pike's Peak Cottonwood Chalk Cliffs Tarrya11s Kenosha Mtns. Rampart Mt. Silverhee1s Southwest San Juansl La Garitas2 Cebolla Creek Ouray Taylor River Lake City Black Canyon Pole Mtn. Sheep Mtn. Marshall Pass Hovenweep NM Northeast Poudre Canyon RMNP4 Rawahs Big Thompson Lone Pine Button Rock Cow Creek The candidate areas mayor may not stand depending on 1) the results of the reconnaissance of other areas and 2) the monetary and time constraints of this study. Sampled counts - During the sampling phase, a block count sampling method similar to that of Houston et a1. (1986) will be used. Census blocks will be delineated on 1:62,000 scale contour maps. Blocks will be designed to include as many slopes and aspects as possible. Conspicuous terrain features (ridges, rock outcrops, forrested areas) will be used as block boundaries. Further details of the design including the number and location of census routes and the number of censuses to be conducted will not be determined until after reconnaissance. 2. Analysis: the population totals will be calculated using Jolly's (1969) method for unequal sized sample units. Schedule Activity Fiscal Year Conduct ground and aerial reconnaissance, select areas. for sampled counts, and delineate census blocks. 1987-88 Conduct sampled counts. 1988-89 Same as 1988-89. 1989-90 Data analysis and publication 1990-91 1-3Ranked by region in order of census need. 4Areas considered to be covered by on-going CSU graduate studies. 5NM denotes National Monument. �309 Personnel D. F. Reed Principal Investigator No. positions undetermined Volunteers Estimated Cost Person days (01) Personal Services (21) Operating Supplies and Services 2,539.00 (28) Travel Expenses 1,000.00 (31) Capital Expenditures (Equipment) 140 TOTAL E. Amount $24,461.00 $28,000.00 GEOGRAPHIC LOCATION Various areas have been mentioned in the Approach section and are subject to change. F. RELATED FEDERAL PROJECT Colorado Federal Aid Project FW 26P. LITERATURE CITED Caugh1ey, G. 1977. Analysis of vertebrate populations. Sons, NY. 234pp. J. Wiley and Denney, R. N. 1976. The status and management of bighorn sheep in Colorado. Desert Bighorn Counc. Trans. 20:5-10. Division of Wildlife. 1977. Div. Wildl. 96pp. Today's strategy •••tomorrow's wildlife. Colo. Geist, V. 1971. Mountain sheep: a study in behavior and evolution. Chicago Press, Chicago and London. 383pp. Univ. Gill, R. B., L. H. Carpenter, and D. C. Bowden. 1983. Monitoring large animal populations: the Colorado experience. Trans. N. Amer. Wildl. Conf. 48:330-341. Houston, D. B., B. B. Moorhead, and R. W. Olson. 1986. An aerial census of mountain goats in the Olympic mountain range Washington. NW Sci. 60(2):131-136. �310 Hudson, R. J. 1982. Bighorn sheep. Page 271 in CRC Handbook of Census Methods for Terrestrial Vertebrates. D.E. Davis, ed. 397pp. Irby, L. R., J. E. Swenson, and S. T. Stewart. 1987. Techniques for assessing population trends and age/sex structure in mountain sheep. J. Wildl. Manage. (In process) Jolly, G. M. 1969. Sampling methods for aerial censuses of wildlife populations. E. Afr. Agric. For. J. 34 (Special Issue):49-49. Overton, W. S., and D. E. Davis. 1969. Estimating the numbers of animals in wildlife populations. Pages 403-455 in R. H. Giles, ed., Wildlife Management Techniques, 3rd Ed. The Wildlife Society, Washington, DC. 623pp. Seton, E. T. 1929. Lives of game animals. Garden City, NY. 4l2pp. Siniff, D. B., and R. O. Skoog. stratified random sampling. Vol. III. Doubleday and Doran Co. 1964. Aerial censusing of caribou using J. Wild1. Manage. 28:391-401. Shannon, N. H., R. J. Hudson, V. C. Brink, and W. D. Kitts. 1975. Determinants of spatial distribution of Rocky Mountain bighorn sheep. Wildl. Manage. 39:387-401. J. Shepherd, H. R. 1975. Vegetation of two dissimilar bighorn sheep ranges in Colorado. Div. Rep. 4. Colo. Div. Wildl. 223pp. Simmons, N. M., and C. G. Hansen. 1980. Population survey methods. Pages 260-272 in The Desert Bighorn: Its Life History, Ecology, and Management. G. Monson and L. Summer, eds. Univ. Ariz. Press, Tucson. 370pp. Wakelyn, L. A. 1984. Analysis and comparison of existing and historic bighorn sheep ranges in Colorado. M.S. Thesis, Colo. State Univ., Fort Collins. 274pp. �311 APPENDIX D PROGRAM NARRATIVE State Colorado Project No. Work Plan No. 01-03-048 - 11700 A. 7 Mountain Sheep Investigations 4 Improve methods for obtaining better population estimates of mountain sheep by combining mark-recapture and telemetry data. ----------------------- Job No. Mammals 2 Research NEED Problem Mountain sheep (Ovis canadensis canadensis) are indigenous to Colorado. They reached North America and probably Colorado during the late Pleistocene. As with essentially all wildlife ungulates with fragmentary fossil records, little is known about their early population levels. Historical accounts of their abundance were not recorded until around 1922 when Seton (1929) estimated that there were about 8,000 mountain sheep in Colorado. The Division's Comprehensive Management Plan (Division of Wildlife 1977) indicated a statewide population of about 3,000 for the base year of 1983. More important, however, is that the plan called for increasing the population to over twice this number in the next 15 years. The projected population curve continues a relatively steep increase through 1993. Concomitant with this planned increase is the consistent highest ranking by each Region of the number one problem: need for additional inventory. If results of the management program for increasing distribution and abundance of mountain sheep by trapping and transplanting is to be known, methods for obtaining better population estimates will be necessary. Literature Review There is a substantial volume of literature concerning mark-recapture theory, models, and various statistical treatments (see reviews by Caughley 1977; Otis et ale 1978; Overton and Davis 1969; Seber 1973, 1982; White et ale 1982). The amount of literature is greatly reduced when considering only the "size of the population" versus 8 other properties that can be investigated by some form of the mark-recapture technique (Caughley 1977:133) and when limiting the application to wild ungulates (Strandgaard 1967, Woolf 1973, Rice and Harder 1977, Bartmann et ale 1987). Aspects of mark-recapture techniques most applicable to a study of .f r ee-rrang Lng wild ungulates include the importance of experimental design, of equal catchability of all individuals, marked and unmarked, where truth of the assumption can be tested in a pilot experiment or have such a test built into the experimental design (Caughley 1977:134), and of adequate sample sizes including the proportion of marked animals in the population. Concerning the latter aspect, large proportions (>67% - Strandgaard �312 1967, >45% - Bartmann et ale 1987) of small populations need to be marked to generate reliable population estimates (White et ale 1982). Approaches to the study of population size in free-ranging wild ungulates might include 1) the Petersen estimate (Petersen 1896, Lincoln 1930) or the simplest mark-recapture procedure which calls for marking on one occasion and recording proportion of marked animals captured (or "sighted" since actual capture is not absolutely necessary if marks can be recognized [Eberhardt 1969]) on a second occasion (Caughley 1977:141), 2) Schumacher's method which calls for marking on several occasions and is estimated from the rate at which the proportion of marked individuals rise as progressively more are marked (Caughley 1977:145), 3) methods involving the assumption of "closure" - the simplest appropriate model of 8 possibilities (models Mo' Mt, Mb, Mh' Mtb' Mth' Mbh' and Mtbh) (Otis et al. 1978, White et al. 1982), 4) methods involving the assumption of "open" populations where the process of birth, death, and migration are allowed to operate--essential elements are incorporated in the classic Jolly-Seber model (Seber 1978:196-232, Seber 1982:196-232), 5) procedures involving maximum likelihood estimates (MLE) used in combining repeated LincolnPetersen estimates (Bartmann et ale 1987), and 6) a procedure of combining mark-recapture and telemetry data (Kufeld et ale 1987). B. OBJECTIVES The overall objective of this study is: To improve the Division's methods for obtaining better population estimates of mountain sheep by combining mark-recapture and telemetry data. Specific subobjectives can be stated as hypotheses to be tested: 1. C. (To be determined - see Analysis section.) EXPECTED RESULTS OR BENEFITS Methods for obtaining better population estimates should improve the Division's ability to match harvest regulations with available mountain sheep. D. APPROACH 1. Methods: Counts - During each of the 2 years of study, 5 aerial counts will be conducted in the study area during November and December to coincide with the rut and when snow conditions improve visibility of the animals. A schedule of counts for the 2 years will depend on these conditions and the favorability of wind and cloud cover. Flights will �313 be conducted with either helicopters or ultralight aircraft (Markowski 1982, Knight et al. 1986) which have adequate performance characteristics for elevation and other conditions of the area. During the counts, the number of telemetered, uniquely marked, and unmarked animals will be noted. Attempts will be made to identify all marks, the ease of which will depend largely on how well the collars are color-coded and how large the numbers are on the collars. Flight patterns will be standardized prior to commencement of counts of which details must await selection of the study area. Study area selection will depend on information collected during the reconnaissance phase of Work Plan 7, Job 3 (Reed 1987). Marking - About 50 mountain sheep will be trapped with a drop net and marked with equal numbers of telemetry collars and individually identifiable collars (unique marks) during the first year (1988). Whether trapping and marking will need to be done during the second year (1989) will depend on how many collars are "out" of the population (as delineated by a defined geographical area) during counts of the first year. An attempt will be made ~o maintain a minimum of 20 active telemetry collars in the population during both years. Either for one or both years, the trapping and marking will be done during JanuaryMarch, a period when mountain sheep are most likely to respond to baiting. Marking will necessarily take place some months before the counts are to be initiated. In order to establish the number of marks in the population just prior to the counts, telemetered animals will be located and placed "in" or "out" of the population to be counted. Any telemetered animals that cannot be located (transmitter failure, animals killed and removed from area, etc.) will be treated the same as those placed out of the population. Hence, the telemetry collars will also need to have unique marks so that if transmitter failures occur and they are subsequently sighted in any of the counts, they can be subtracted from the number of marks counted. 2. Analysis: Each count is expected to yield recaptures (sightings) from each of the 2 subgroups: 1) telemetered marks, and 2) unique marks. These different subgroups of marks will be related in that whatever number of telemetered marks are determined to be "out", the same number of unique marks will be ascribed to the same status. This assumption will, in effect, double the sample size of "known" marks in the population. At this juncture in the formation of this study, it is suggested that 2 or more procedures be tested for their efficiencies (confidence intervals). Tentatively, a hypothesis for testing the procedures involving joint hypergeometric MLE used in combining repeated Lincoln-Petersen estimates (Bartmann et al. 1987) and an extended mark-recapture method (Kufeld et al. 1987) is: joint hypergeometric MLE and extended mark-recapture yield different efficiencies (confidence intervals) in estimating population sizes of mountain sheep. �314 Analyses will follow those of Bartmann et ale (1987) and Kufeld et ale (1987) • Schedule Activity Fiscal Year Conduct ground and aerial reconnaissance, select area. 1987-88 Conduct sampled counts. 1988-89 Same as 1988-89. 1989-90 Data analysis and publication. 1990-91 Personnel D. F. Reed Principal Investigator No. positions undetermined Volunteers Estimated Cost Person Days (01) Personal Services (02) Operating Supplies and Services (28) Travel Expenses (31) Capital Expenditures 70 $12,230.50 1,269.50 500.00 (Equipment) TOTAL E. Amount $14,000.00 GEOGRAPHIC LOCATION The area is yet to be determined. F. RELATED FEDERAL PROJECT Colorado Federal Aid Project FW26P. LITERATURE CITED Bartmann, R. M., G. C. White, L. N. Carpenter, and R. A. Garrott. 1987. Aerial mark-recapture estimates of confined mule deer in pinyon-juniper woodland. J. Wildl. Manage. 51:41-46. �315 Caughley, G. 1977. NY. 234pp. Analysis of vertebrate populations. Division of Wildlife. 1977. Div. Wildl. 96pp. J. Wiley and Sons, Today's strategy •••tomorrow's wildlife. Colo. Eberhardt, L. L. 1969. Population estimates from recapture frequencies. Wildl. Manage. 33:28-39. J. Knight, J. E., C. L. Foster, V. W. Howard, and J. G. Schickedanz. 1986. A pilot test of ultralight aircraft for control of coyotes. Wildl. Soc. Bull. 14:174-177. Kufeld, R. C., D. C. Bowden, and D. L. Schrupp. 1987. Estimating mule deer density by combining mark-recapture and telemetry data. J. Mammal. (In Press) Lincoln, F. C. 1930. Calculating waterfowl abundances on the basis of banding returns. u.S. Dep. Agric. Circ. 118. 4pp. Markowski, M. A. 1982. Ultralight airplanes. Scientific Am. 247:62-68. Otis, D. L., K. P. Burnham, G. C. White, and D. R. Anderson. 1978. Statistical inference from capture data on closed animal populations. Wildl. Monogr. 62:1-135. Overton, W. S., and D. E. Davis. 1969. Estimating the numbers of animals in wildlife populations. Pages 403-455 in R. H. Giles, ed., Wildlife Management Techniques, 3rd Ed. The \~ildl. Soc., Washington, DC. 623pp. Petersen, C. G. J. 1896. The yearly immigration of plaice into the Limfjord from the German Sea. Rep. Dan. BioI. Stn. 1895 6:1-77. Reed, D. F. 1987. Improve matching harvest regulations with available mountain sheep populations through better counts and better analyses of count data. Colo. Div. of Wildl., Program Narrative, Work Plan 7, Job 3. 6pp. Mimeogr. Rice, W. R., and J. D. Harder. 1977. Application of multiple aerial sampling to a mark-recapture census of white-tailed deer. J. Wildl. Manage. 41:197-206. Seber, G. A. F. parameters. 1973. The estimation of animal abundance and related Hafner Press, NY. 506pp. 1982. The estimation of animal abundance and related parameters. ed. MacMillan Publishing Co., NY. 654pp. Seton, E. T. 1919. Lives of game animals. Co., Garden City, NY. 4l2pp. Vol. III. Doubleday and Doran Strandgaard, H. 1967. Reliability of the Petersen method tested on a roedeer population. J. Wildl. Manage. 31:643-651. 2nd �White, .G. C., D. R. Anderson, K. P. Burnham, and D. L. Otis. 1982. Capturerecapture and removal methods for sampling closed populations. Los Alamos National Laboratory. LA-8787-NERP. Los Alamos, NM. 235pp. Woolf, A. 1973. Population dynamics and remote censusing of a large, captive white-tailed deer herd. Ph.D. Thesis, Cornell Univ., Ithaca, NY. l68pp. �317 APPENDIX E PROGRAM NARRATIVE State of Colorado Project No. Mammals Research Work Plan No. 2A Mountain Sheep Investigations Job No. 5 Experimental Tests of the MarkRecapture Method to Estimate Mountain Sheep Numbers A. NEED Colorado has an aggressive mountain sheep translocation program. The objectives of this program are to expand the distribution of mountain sheep back into historically occupied habitats which are currently vacant or are occupied by small, stagnant, remnant populations, and to increase increase mountain sheep numbers throughout the state. Currently, objective progress towards those objectives is hindered because we have no demonstrably reliable methods to estimate mountain sheep numbers. In recent years, markrecapture techniques have become the method of choice to estimate mountain sheep numbers (Furlow et al. 1981, Douglas and Leslie, Jr. 1986, Leslie, Jr. and Douglas 1986). Leslie, Jr., and Douglas (1986) summarized the state of the art for mountain sheep as follows: "Despite intensive field investigations in the river Mountains, we must admit that our ability to estimate population size is limited. Confidence intervals around population estimates derived from fall helicopter surveys conducted by the NDOW are large (Fig. 4). Furthermore, because of the number of marked animals in the population was unknown, confidence intervals from 1979 assume that efficiency of fall helicopter surveys has remained unchanged." There is a substantial volume of literature concerning mark-recapture theory, models, and various statistical treatments (see reviews by Overton and Davis 1969; Seber 1973, 1982, 1986; Caughley 1977; Jensen 1981; Otis et ale 1978; White et al. 1982). The amount of literature is greatly reduced when considering only the "size of the population" versus 8 other properties that can be investigated by some form of the mark-recapture technique (Caughley 1977:133) and when limiting the application to wild ungulates (Strandgaard 1967, Woolf 1973, Rice and Harder 1977, Biggens and Jackson 1982, Bartmann et al. 1987). Aspects of mark-recapture techniques most applicable to a study of free-ranging mountain sheep include the importance of experimental design, equal probabilities of catchability within sex-and-age categories, equal probabilities of recapturing (resighting) of both marked and unmarked individuals, and of adequate sample sizes both of marked animals in the population and recapture or resighting samples. According to the results of Strandgaard (1967) and Bartmann et al. (1987), large proportions ( 45%) of small populations need to be marked to generate reliable population estimates (White et ale 1982). But this requirement may be mitigated to some extent by combining large numbers of small resighting samples (Cox and Solomon 1986, Bartmann et ale 1987. �31d This study intends to examine the magnitude and the consequences of violating the fundamental assumptions of the mark-recapture method or Lincoln Index when applied to small aggregated populations of mountain sheep. B. OBJECTIVE Develop standard operating procedures for estimating mountain sheep numbers with the Lincoln Index or comparable mark-recapture procedures. C. EXPECTED RESULTS OR BENEFITS Methods for obtaining better population estimates should improve the Division's ability to evaluate the success of mountain sheep drug treatment and translocation efforts and to evaluate the efficacy of mountain sheep hunting strategies. D. APPROACH 1. Up to 40 mountain sheep will be marked with radiocollars to determine the probability of sighting each radio-collared animal during each resighting sample. Resighting samples will be replicated up to 5 times in preliminary tests to determine resighting sample size requirements. Data from the sighting probabilities of individual radio-collared animals will be used to generate the probability density function of the probability of sighting distribution. Radio-collared animals are necessary to be sure they are on the study and are alive at the time of each resighting sample. 2. Degree of association or aggregation among radio-collared animals will be computed to quantify the extent to which the assumption of independent observations is violated. 3. Methods of resighting (helicopters vs. ground surveys) will be compared to evaluate their performance relative to their costs. Approaches that improve the probability of sighting and still meet the assumptions of the technique within acceptable tolerance limits while holding survey costs should be developed. 4. Field surveys will be repeated for 2 years to determine repeatability of the results. Microcomputer Laboratory Activities 1. A range of heterogeneities among the population of resighting probabilities will be simulated to document their effects upon various estimators of mountain sheep population parameters. 2. A range of aggregation or clumping scenarios will be simulated (i.e. degrees of nonindependence of sighting probabilities) to document their effects upon various estimators of mountain sheep population parameters. �319 3. From these combinations of field and simulation activities, guidelines will be developed for standard operating protocols in the use of markrecapture techniques to estimate mountain sheep numbers and population dynamics. Schedule Fiscal Year Activity Period 1987-88 Select Study Area and Design Specific Field Protocol October-May 1988-89 Capture Mountain Sheep and Conduct Resighting Samples December-February Conduct Population Estimation Simulations March-June 1989-90 Same as FY 1988-89 1990-91 Data Analysis, Publication, Development of Mountain Sheep Inventory Standard Operating Procedures Year-round Personnel Co-Principal Investigator Co-Principal Investigator Co-Principal Investigator Graduate Research Assistant Dale F. Reed Dr. Gary C. White R. Bruce Gill Andrea Neal Estimated Annual Costs Costs FTE Requirements PFTE = 1.08 TFTE = 0.00 TOTAL = 1.08 E. (01) Personal Services (21) Operating Supplies and Services (21) Utilities (28) Travel Expenses (31) Capital Outlay $63,000 18,000 o 1,500 500 $83,000 LOCATION Tentatively, the Poudre Canyon northwest of Fort Collins, Colorado, to reduce travel expenses. F. RELATED FEDERAL PROJECTS None �320 LITERATURE CITED Bartmann, R. M., G. C. White, L. C. Carpenter, and R. A. Garrott. 1987. Aerial mark-recapture estimation of confined mule deer in pinyon-juniper woodland. J. Wi1d1. Manage. 51:41-46. Biggens, D. E., and M. E. Jackson. 1982. Biases in aerial surveys of mule deer. Pages 60-65 in R. D. Conner et a1. (eds.). Issues and technology in the management of impacted western wildlife. Thorne Ecological Institute. Boulder, CO. Caugh1ey, C. 1977. York, NY. Analysis of vertebrate populations. J. Wiley & Sons. New Cox, D. R., and P. J. Solomon. 1986. Analysis of variability with large numbers of small samples. Biometrika 73(3):543-554. Douglas, C. L., and D. M. Leslie, Jr. 1986. Influence of weather and density on lamb survival of desert mountain sheep. J. Wi1d1. Manage. 50(1):153-156. Eberhardt, L. L. 1969. Population estimates from recapture frequencies. Wi1d1. Manage. 33:28-39. J. Furlow, R. C., M. Hader1ie, and R. Van den Berge. 1981. Estimating a bighorn population by mark-recapture. Desert Bighorn Council, Trans. 1981:31-33. Jensen, A. L. 1981. Sample sizes for single mark and single recapture experiments. Amer. Fish. Soc., Trans. 110:445-458. Leslie, Jr., D. M., and C. L. Douglas. 1986. Modeling demographics of bighorn sheep: current abilities and missing links. N. Amer. Wildl. Natur. Res. Conf., Trans. 51:62-73. Lincoln, F. C. 1930. Calculating waterfowl abundances on the basis of banding returns. USDA Circ. 118. 4pp. Otis, D. L., K. P. Burnham, G. C. White, and D. R. Anderson. 1978. Statistical inference from capture data on closed animal populations. Wi1d1. Monogr. 62:1-135. Overton, W. S., and D. E. Davis. 1969. Estimating the numbers of animals in wildlife populations. Pages 403-455 in R. H. Giles (ed.). Wildlife management techniques, 3rd ed. The Wi1d1. Soc. Washington, DC. Petersen, C. G. J. 1896. The yearly immigration of plaice into the Limfjord from the German Sea. Rep. Dan. BioI. Stn. 1895 6:1-77. Rice, W. R., and J. D. Harder. 1977. Application of mUltiple aerial sampling to a mark-recapture census of white-tailed deer. J. Wi1d1. Manage. 41:197-206. Seber, G. A. F. parameters. 1973. The estimation of animal abundance and related Hafner Press. New York, NY. �321 Seber, G. A. F. parameters. 1986. Strandgaard, H. population. 1982. The estimation of animal abundance and related 2nd ed. MacMillan Publ. Co. New York, NY. A review of estimating and abundance. Biometrics 42(2):267-292. 1967. Reliability of the Petersen method tested on a roe-deer J. Wildl. Manage. 31:643-651. White, G. C., D. R. Anderson, K. P. Burhnam, and D. L. Otis. 1982. CaptureLos Alamos recapture and removal methods for sampling closed populations. National Laboratory. LA-8787-NREP. Los Alamos, NM. Woolf, A. 1973. Population dynamics and remote censusing of a large captive. white-tailed deer herd. Ph.D. Thesis, Cornell Univ. Ithaca, NY. ��Wildlife Research Report July 1987 JOB PROGRESS REPORT State of Colorado Project No. 01-03-048 (FW 26 P) Mammals --~--~~--~~----~-- 2 Research Work Plan No. 8 Small Carnivorous Mammals Investigations Job No. 1 Experimental Introduction of River Otter Period Covered: July 1, 1986 - June 30, 1987 Author: T. D. I. Beck ABSTRACT A detailed study plan was developed with management and research objectives. An Environmental Analysis Report was submitted to the BLM. A reliable source for river otters was located, and a tentative order was placed for March, 1988, delivery. ��325 SMALL CARNIVOROUS MAMMALS INVESTIGATIONS Thomas D. I. Beck P. N. OBJECTIVE Develop procedures for river otter reintroductions in Colorado and establish a self-sustaining population of river otters from which to collect river otters for future trans1ocations. SEGMENT OBJECTIVE Prepare a study plan detailing programmed activities to experimentally introduce river otter into the Dolores River. ACKNOWLEDGMENTS Assistance with preliminary field studies was provided by the following CDOW personnel: S. Boyle, K. Kehmeir, D. Langlois. RESULTS AND DISCUSSION An extensive literature review on mustelids was conducted preparatory to writing a study plan. A study plan was prepared with both management and research objectives. A draft Environmental Assessment was prepared and submitted to Bureau of Land Management officials. Considerable time was spent trying to develop sources for river otters for transplant. The best source (cost, health of animals, reliability, desired sex) appears to be LeeRoy Sevin of Louisiana. River otters from this source have been successfully transplanted to other northern states (Pennsylvania, Iowa, Missouri). Preliminary habitat surveys were begun in late June, 1987, with the intent being to identify the areas within the 160-km study area for release sites next March. The river is boatable for approximately 9 mos. each year. Ice on the long pools in the lower canyon prohibits winter boating, but most of the area is then accessible by foot. Inquiries have been made regarding video documentation of the program, both in Colorado and California. Discussions with private groups wishing to raise funds for purchase of river otter were conducted. No firm proposals were developed on either. Prepared by JL~n.~ ;!)~;j4 ~T~h-o~m~a~s~D~.~I~.~B-e-c7k------~-Wildlife Researcher � Dublin Core The Dublin Core metadata element set is common to all Omeka records, including items, files, and collections. For more information see, http://dublincore.org/documents/dces/. Title A name given to the resource Federal Aid Wildlife Quarterly/Annual Reports Description An account of the resource In 1937 the U.S. Congress enacted, and President Franklin D. Roosevelt signed, the Federal Aid to Wildlife Restoration Act (50S-917). This milestone in wildlife financing is popularly known as the Pittman-Robertson act. The Colorado Federal Aid to Wildlife Restoration program began in 1937, shortly after enabling legislation by the Congress in 1936. Reports of progress, as required by the Federal Aid Act, first appeared in 1938. Federal Aid reports have been prepared continuously since. Beginning in 1947 and continuing through 2000, Colorado Federal Aid Reports were issued quarterly, in January, April, July and October. <strong>Starting in 2001</strong>, reports were issued annually.&nbsp;<br /><br />Current reports cover both federal aid and non-federal aid research and summarize (≤6 pages each with tables and figures) preliminary results of wildlife research projects and support services updates conducted by the Mammals<br />Research Team.&nbsp;&nbsp;<br /><br />The title of the reports has changed over the years. Reports from 1938-1945 were issued under specific job title.&nbsp; For a listing of individual projects see <a href="https://cpw.cvlcollections.org/exhibits/show/mammals-research/progress-reports">Mammals Research: Progress Reports (1939-current)</a>.<br /><br />Quarterly Progress Report (1946-1956)<br /><a href="https://cpw.cvlcollections.org/items/show/674">1946-1956</a><br /><br />Quarterly Report (1957-1962)<br /><a href="https://cpw.cvlcollections.org/items/show/673">1957-1962</a><br /><br />Game Research Report (1963-1979)<br /><a href="https://cpw.cvlcollections.org/items/show/454">1963-1969</a><br /><a href="https://cpw.cvlcollections.org/items/show/455">1970-1979</a> <p>Wildlife Research Report (1980-2000)<br /><a href="https://cpw.cvlcollections.org/items/show/451">1980-1987</a><br /><a href="https://cpw.cvlcollections.org/items/show/452">1988-1994</a><br /><a href="https://cpw.cvlcollections.org/items/show/453">1995-2000</a></p> <p>Wildlife Research Report. Mammals (2001-2023)<br /><a href="https://cpw.cvlcollections.org/items/show/611" class="permalink">2001-2023</a><br /><br />Mammals Research Summary Report (2024-&nbsp; )<br /><a href="https://cpw.cvlcollections.org/items/show/612">2024-present</a></p> Dublin Core The Dublin Core metadata element set is common to all Omeka records, including items, files, and collections. For more information see, http://dublincore.org/documents/dces/. Title A name given to the resource Wildlife Research Report (1980-1987) Description An account of the resource Published under title: <em>Wildlife Research Report</em>&nbsp;from 1980-2000. This set contains <strong>1980-1987</strong>.<br /><a href="https://cpw.cvlcollections.org/items/show/452">Wildlife Research Report (1988-1994)</a><br /><a href="https://cpw.cvlcollections.org/items/show/453">Wildlife Research Report (1995-2000)</a><br /><br />Research topics issued in specific months. January 1983-1986: Non-game investigations; January 1987-1993: Raptor investigations. April 1980-1982: Game bird survey; April 1983: Small game investigations; April 1984: Game bird survey; April 1986-2000: Avian research. July 1980-1984, July 1985- Mammals. September 1983-1984: Law Enforcement investigations; September 1986: Wildlife law enforcement research. October 1980-1984: Migratory bird investigations; October 1999: Avian research, migratory birds; October 1989-1993: Migratory game bird research. <p>Progress reports for mammals and avian Federal Aid research. See&nbsp;<a href="https://cpw.cvlcollections.org/exhibits/show/mammals-research/progress-reports">Mammals Research: Progress Reports (1939-current)</a> for a listing of collated reports for each mammals project (avian to be added in the future).<br /><br />Some quarters contain multiple volumes.</p> <br />Continues: Game Research Report (1963-1969)<br />&nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp;&nbsp;<a href="https://cpw.cvlcollections.org/items/show/454">1963-1969</a>&nbsp; |&nbsp;&nbsp;<a href="https://cpw.cvlcollections.org/items/show/455">1971-1979</a><br /><br />Continued by: <a href="https://cpw.cvlcollections.org/items/show/611">Wildlife Research Report. Mammals (2001-current)</a>. Avian research did not publish quarterly/annual report from 2001-2009.<br /><br />Print copy: Federal Aid binders Rights Information about rights held in and over the resource <a href="http://rightsstatements.org/vocab/InC-NC/1.0/">IN COPYRIGHT - NON-COMMERCIAL USE PERMITTED</a> Type The nature or genre of the resource Text Format The file format, physical medium, or dimensions of the resource application/pdf Language A language of the resource English Creator An entity primarily responsible for making the resource Colorado Division of Wildlife